."'.'■,:'

(. .

ILLINOIS BIOLOGICAL
MONOGRAPHS

PUBLISHED QUARTERLY
UNDER THE AUSPICES OF THE GRADUATE SCHOOL

BY THE UNIVERSITi' OF ILLINOIS \

VOLUME VII

Urbana, Illinois
1922

Editorial Committee

Stephen Alfred Forbes William Trelease

Henry Baldwin Ward

TABLE OF CONTENTS
VOLUME VII

NUMBERS PAGES

1. Studies on Gregarines, II. By Minnie Watson Kamm. With 4 plates 1-104

2. The Molluscan Fauna of the Big Vermilion River, Illinois. By F. C. Baker.

With 15 plates 105-224

3. North American Monostomes. By E. C. Harrah. With 9 plates 225-324

4. A Classification of the Larvae of the Tenthredinoidea. By H. Yuasa.

With 14 plates 325-490

/ ? 7^

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII January, 1922 No. i

Editorial Committee

Stephen Alfred Forbes William Trelease

Henry Baldwin Ward

Published under the

Auspices of the Graduate School by

THE University of Illinois Press

COPVKIGHT, 1922 BY THE UNIVERSITY OF IlUMQIS

Distributed June 30, 1922

STUDIES ON GREGARINES II

SYNOPSIS OF THE POLYCYSTID GREGARINES

OF THE WORLD, EXCLUDING THOSE

FROM THE MYRIAPODA,

ORTHOPTERA, AND

COLEOPTERA

WITH FOUR PLATES

BY

MINNIE WATSON KAMM

Contributions from the

Zoological Laboratory of the University of lUinois

under the direction of Henry B. Ward, No. 195

TABLE OF CONTENTS

Introduction 7

Classification of the Tribe Cephalina with the Type Species 10

Group of Tables Showing the Phylogenetic Relationships of Gregarines 16

Table 1. Showing the Intermediate Position of the Two Families, Lecudinidae and

Polyrhabdinidae 16

Table 2. Showing Relationships of the Families of the Tribe Cephalina 16

Table 3. Showing Relationships Between Genera in the Family Gregarinidae 17

Table 4. Showing Relationships Between Genera in the Family Actinocephalidae . ... 17

Table 5. Showing Relationships Between Genera in the Family Acanthosporidae . ... 19

Table 6. Showing Relationships Between Genera in the Family Stylocephalidae 19

Table 7. Showing Relationships Between Genera in the Family Dactylophoridae . ... 20

Synopses of the Gregarines from Various Groups of the Animal Kingdom 21

List and Descriptions of Species Found in the Phylum Coelhelminthes 21

Family Lecudinidae Kamm (Nov. fam.) 21

Discussion of the New Family Lecudinidae 22

Family Polyrhabdinidae Kamm 1922 26

Discussion of the New Family Polyrhabdinidae 26

List and Descriptions of Species Found in the Phylum Mollusca 30

List and Descriptions of Species Found in the Class Crustacea of the Phylum Arthro-

poda 31

Family Didymophyidae Leger 32

Family Gregarinidae Labbe 1899 34

Family Cephaloidophoridae Kamm 1922 36

List and Descriptions of Species Found in the Class Acerata 49

Family Actinocephahdae Leger 1892 49

Family Acanthosporidae Leger 1892 53

List and Descriptions of Species Found in the Class Malacopoda 54

List and Descriptions of Species Found in the Order Thysanura of the Class He.xapoda 54

Family Gregarinidae Labbe 1899 54

List and Descriptions of Species Found in the Order Isoptera 56

Family Gregarinidae Labbe 1899 56

List and Descriptions of Species Found in the Order Hemiptera 57

Family Actinocephalidae Leger 1892 57

List and Descriptions of Species Found in the Order Neuroptera 59

Family Gregarinidae Labbe 1899 60

Family Actinocephalidae Leger 1892 62

Family Acanthosporidae Leger 1892 70

Family Menosporidae L^ger 1892 71

6 ILLINOIS BIOLOGICAL MONOGRAPHS [6

List and Descriptions of Species Found in the Order Lepidoptera 72

Family Gregarinidae Labbe 1899 72

List and Descriptions of Species Found in the Order Diptera 73

Family Gregarinidae Labbe 1899 73

Family Actinocephalidae Leger 1892 75

List and Descriptions of Species Found in the Order Siphonaptera 79

Family Actinocephalidae Leger 1892 79

List and Descriptions of Species Found in the Subphylum Tunicata 80

List of Hosts with Their Gregarine Parasites 82

Bibliography 86

Explanation of Plates 92

Index 101

71 STUDIES ON GREGARINES—KAMM

INTRODUCTION

This monograph constitutes the second half of a problem in which it is
endeavored to bring together in small compass the more important facts
ooncerning the systematic position of polycystid gregarines. The first
part of the work (Watson, 1916a) was published in 1916 and included the
polycystid gregarines recorded from the Coleoptera, Orthoptera, and
Myriapoda of the world. The present paper comprises those from the
remainder of the animal kingdom. It is designed chiefly for the use of
workers on gregarines who do not have access to the vast literature which
has developed upon the subject and with which the University library is
so richly endowed.

I wish to thank Professor Henry Baldwin Ward for kindly encouraging
the publication of this paper, for his advice and judgment concerning some
of the knotty systematic questions involved, and for the use of the labora-
tory and library facilities in his Department.

When the first paper was published Dr. C. W. Stiles, of Washington,
informed me concerning the desirability of designating the type species
of the various genera, and this has been done in the present paper. He
also gave his opinion concerning the type species of the genus Cephaloido-
phora, for which I wish to express my thanks.

The larger groupings used in the present paper are made from the
standpoint of the host-material rather than the parasite itself in order to
facilitate work in the field. From the descriptions it can be at once
determined whether or not a particular gregarine has ever been found in
identical or similar material. If it has not been described from hosts in
the same group and its position is definitely established from some one or
more of the generic characters, one may turn to the index of this and of the
preceding paper, where all the known species in each genus are ranged in
alphabetical order under the generic name. From a comparison of all
the known species with the species in hand it may be determined whether
or not the latter has been described from some other group. It is true,
however, that except in rare instances a species is confined to a single
host or to nearly-related hosts, not being found in insects as far removed
from each other as the Coleoptera and the Orthoptera.

In this paper I have not tabulated a complete alphabetic list of the
gregarine genera with their hosts or of the hosts with their parasites for this
reason: Two such lists were given in the former paper covering the species

8 ILLINOIS BIOLOGICAL MONOGRAPHS [8

up to 1915, and those described since then are listed in the same manner
at the end of the present paper.

The previous monograph covers the species known from the Coleoptera,
Orthoptera, and Myriapoda up to the year 1915. The gregarines parasitic
in the remainder of the animal kingdom up to the beginning of 1920 are
described in the present work. The arrangement of hosts in orders and
classes of the Hexapoda follows that given by Comstock (1912).

Because more and more species are being described in such great genera
as Gregarina, Actinocephalus, etc., which are very similar in all their charac-
teristics but yet are distinct species, I believe accurate and detailed
measurements are necessary for the sporonts, trophozoites, cysts, and
spores, or as many of these characters as may be determined. Measure-
ments should be made of quiescent but not water-swollen animals while
alive on the slide. These measurements must needs be made very quickly
after the host is opened for a water medium is highly destructive to the
ectosarc of the gregarine, at first causing it to swell out of all normal pro-
portions and then to break, with the collapse of the animal. When the
parasites do not die within a short time (e.g., the Stenophoridae) they at
least lose their original proportions and measurements become valueless.
Those taken on preserved material are far from accurate.

Ratios used in this paper indicate length of protomerite to total length
(LP:TL) and width of protomerite to width of deutomerite (WP:WD) as
indicated in the author's drawings even though no measurements may
have been made.

One would like to make a clean-cut and sweeping statement that
polycystid gregarines are parasites of the arthropods exclusively and
unless an exhaustive study be made this may seem to be true.

But just as higher animals cannot be placed under such definite groups,
so it is with these simply organized forms. There is a gradual transition
from simple to complex — from the unilocular to the septate forms, and
from the simpler to the higher septate organisms. As shown in the group
of tables in this paper, I believe there is an interesting and not entirely
imaginary parallelism in the character of the hosts which harbor these
developing parasites. The transitional forms between non-septate and
septate gregarines are found exclusively in the polychaetes. The two
lowest (and very similar) families of septate gregarines are found in lowly
organized arthropods — the Crustacea and Diplopoda. These gregarines
possess no epimerites or very rudimentary ones.

The next higher family, in which the epimerite is a very simple struc-
ture, is abundantly represented in the Orthoptera, but also in the Coleop-
tera (which is not one of the lower orders), and is fairly general throughout
the Hexapoda. Just as the line of evolution in higher animals may take
a single track however aberrent it may prove to be, this is apparently

9] STUDIES ON GREGARINES—KAMM 9

true in gregarines, for many instances may be mentioned in which genera
are found exclusively in a single order of insects; the genus most frequently
present in the Acerata (Sciadophora) is found nowhere else and is highly
specialized and unique along a particular line.

A gregarine has even been found in Peripatus; and the Tunicates
contain true septate forms. It would be interesting to know what becomes
of these parasites above the Tunicates — whether they lose their gregarinoid
and take on higher characteristics, continuing to develop in vertebrates, or
whether the alimentary tract of the latter contain enzymes not suited to their
continued existence and they are thereby extinguished.

A suggested problem is that of the possibility of the existence of greg-
arines in the insects found in Baltic amber; the insects being transparent
their presence or absence should be easily determined, and possibly some
of the generic characters.

10 ILLINOIS BIOLOGICAL MONOGRAPHS [10

CLASSIFICATION OF THE TRIBE CEPHALINA

WITH THE TYPE SPECIES

Order 1. Gregarinoidea Minchin 1912.

Suborder 1. Eugregarinae L6ger 1900.

Tribe 1. Cephalina Delage and Herouard 1896 (Intestinal parasites almost ex-
clusively).

Family 1. Lecudinidae Kamm 1922. Non-septate gregarines inhabiting
digestive tract of polychaetes. Epimerite a simple knob.
Genus 1. Lecudina Mingazzini 1891:469. Non-septate, epimerite simple,
spores ovoidal with thickening at one pole.
Tj'pe: Lecudina pellucida (Kolliker) Mingazzini.
Family 2. Polyrhabdinidae Kamm 1922. Septate gregarines inhabiting the
digestive tract of polychaetes. Epimerites varied.
Genus 2. Polyrhabdina Mingazzini 1891 :229. Septate, sporonts flattened,
ovoidal, epimerte a corona of hooks. Intestine of poly-
chaetes of the family Spionidae.
Type species: Polyrhabdina spionis (Koll.) Ming.
Genus 3. Sycia L6ger 1892:52. Epimerite knobbed, bordered by a thick
ring. Protomerite subspherical, deutomerite conical with
many inclosures in adult.
Type: Sycia inopinata Leger.
Genus 4. Ulivina Mingazzini 1891: 235. Elongate-ellipsoidal, protomerite
the more dense.
Type: Ulivina elliptica Ming. ''

Family 3. Cephaloidophoridae Kamm 1922. Sporonts associated in twos,
no epimerite. Development intmcellular; cyst dehiscence
by simple rupture, spores ovoidal with equatorial line.
Parasites of Crustacea.
Genus 5. Cephaloidophora Mawrodiadi 1908:101. Characters of the
family.
Type: Cephaloidophora communis Mawrodiadi.
Family 4. Stenophoridae Leger and Duboscq 1904:361. Sporonts solitary,
epimerite absent or rudimentary. Development intracellu-
lar; cyst dehiscence by simple rupture, spores ovoidal with
equatorial line. Confined to Diplopods.
Genus 6. Stenophora Labbe 1899: 15. Characters of the family.

Type: Stenophora juli (Frantz.) Schn.
Genus 7, Fonsecaia Pinto 1918 : ? Like type genus except spores elongate-
ellipsoidal, no endospore.
Type: Fonsecaia polymorpha Pinto.
Family 5. Gregarinidae Labbe 1899:9. Epimerite symmetrical, simple.
Sporonts solitary or in association. Cysts with or without
spore-ducts, spores symmetrical. (Confined to insects with
two exceptions, development extracellular.)
Genus 8. Leidyana Watson 1915:35. Sporonts solitary, epimerite a
simple sessile knob. Dehiscence by spore-ducts. Spores
barrel-shaped, in chains.
Type: Leidyana erratica (Crawl.) Wats.

Remarks in pareathesis are added by the writer.

11]

STUDIES ON GREGARINES—KAMM

11

Genus 9. Gregarina Dufour 1828:366. Sporonts associated in twos,
epimerite a simple globular or cylindrical papilla. Cysts
with spore-ducts, spores barrel-shaped, extruded in chains.

Type: Gregarina ovata Duf.
Genus 10. Protomagalhaensia Pinto 1918: ? Sporonts in associations
of several individuals, often attached laterally. Myonemes.
prominent. Sporonts attenuate. (Cysts and epimerites
unknown.) Spores barrel-shaped, spine at each corner.

Type: Prolomagalhaensia serpentula (Magal.) Pinto.

Genus 11. Gamocystis Schneider 1875:587. Protomerite only in tro-
phozoite, sporonts in twos; sporulation partial, with spore-
ducts, spores cylindrical.

Type: Gamocystis tenax Schn.

Genus 12. Hyalospora Schneider 1875:583. Sporonts in associations of
two, endoplasm yellow-orange. Epimerite a simple
globular knob. Cyst dehiscence by simple rupture, spores
ellipsoidal.

Type: Hyalospora roscoviana Schn.
Genus 13. Hirmocystis Labb^ 1899:12. Sporonts in associations of from
two to twelve (or more). Epimerite a small cylindrical
papilla. Cysts spherical, dehiscence by simple rupture,
spores ovoidal.

Type: Hirmocystis venlricosa (L6ger) Labb6.
Genus 14. Euspora Schneider 1875:582. Sporonts in twos. (Epimerite
unknown.) Cysts spherical, dehiscence by simple rupture,
spores prismatic.
Type: Euspora fallax Schn.

Genus 15. Sphaerocystis L^ger 1892:115. Protomerite only in trophic
stages, sporonts solitary, subspherical. Cysts spherical,
dehiscence by simple rupture, spores ovoidal.

Type: Sphaerocystis simplex L6ger.
Genus 16. Cnemidospora Schneider 1882 :446. Sporonts solitary, anterior
half of protomerite gray, posterior half yellow-green. (Epi-
merite not known.) Cyst dehiscence by simple rupture,
spores ellipsoidal.

Type: Cnemidospora lutea Schn.

Genus 17. Uradiophora Mercier 1912 :198. Sporonts in twos, deutomerite
with small appendix. Epimerite an elongate papilla. Cysts
ovoidal, dehiscence by simple rupture, spores spherical or
subspherical, extruded in chains. Development extra-
cellular.

Type: Uradiophora cuenoti (Merc.) Merc.

Genus 18. Pyxinoides Tr6goubofif 1921: liii. Sporonts in associations of
two, epimerite a slightly stalked globular papilla with
sixteen longitudinal furrows, small cone at apex. Extra-
cellular development.

Type: Pyxinoides bcdani (Koll.) Treg.

12 ILLINOIS BIOLOGICAL MONOGRAPHS [12

Family 6. Didymophyidae L6ger 1892:105. Sporonts in associations of two or
three. No septum in satellites.

Genus 19. Did>Tnophyes Stein 1848:186. Epimerite a small pointed
papilla. Cysts spherical, dehiscence by simple rupture,
spores ellipsoidal.

Type : Didymophyes gigantea Stein.

Family 7. Actinocephalidae Leger 1892:166. Sporonts solitary, epimerites
complex and varied. Cyst dehiscence by simple rupture,
spores irregular, biconical, or cylindro-biconical. (Practi-
cally confined to insects.)

Genus 20. Amphoroides Labbe 1899:20. Protorperite cup-shaped,
epimerite a globular sessile papilla. Spores curved.

Type: Amphoroides polydesmi (Leger) Labbe.

Genus 21. Pileocephalus Schneider 1875:591. Epimerite a lance-shaped
cone on a short neck. Spores biconical.

Type: Pileocephalus chinensis Schn.

Genus 22. Stylocystis Leger 1899:526. Epimerite a sharply recurved
cone. Spores biconical.
Type: Stylocystis praecox Leger.

Genus 23. Discorhjmchus Labb6 1899:20. Epimerite a large globular
papilla with a thin collar at the base, neck short. Spores
biconical, slightly curved.
Type: Discorhynchus truncalus (Leger) Labb6.

Genus 24. Steinina Leger and Duboscq 1904:352. Epimerite a short
mobile digitiform process changing into a flattened disc.
Spores biconical.
Type: Steinina ovalis (Stein) L & D.

Genus 25. Anthorhynchus Labbe 1899:19. Epimerite a large fluted
flattened disc. Spores biconical, extruded in chains
laterally.
Type: Anthorhynchus sophiae (Schn.) Labb^.

Genus 26. Sciadophora Labbe 1899:18 Epimerite a large sessile disc
with crenulate periphery. Protomerite with numerous
backwardly-directed vertical laminations. Spores biconical.
Type: Sciadophora phalangii (Ldger) Labb6.

Genus 27. Amphorocephalus Ellis 1913:462. Epimerite a sessile peri-
pherally fluted disc set upon a short dilated neck. Proto-
merite constricted superficially across middle. (Spores
unknown.)
Type: Amphorocephalus amphorcllus Ellis.
Genus 28. Pyxinia Hammerschmidt 1838:357. Epimerite a flat crenulate
crateriform disc from the center of which rises a long or
short style. Spores biconical.
Tjrpe: Pyxinia rubecula Hamm.

Genus 29. Schneideria L^ger 1892:153. EpiAerite sessile, a thick hori-
zontal disc with milled border, from center of which pro-
jects up a short style. No protomerite in adults. Spores
biconical.

13J STUDIES ON GREGARINES— KAMA! U

Type: Schneider ia mucronata Leger.
Genus 30. Asterophora L^ger 1892 : 129. Epimerite like Schneideria,
with longer style. Spores cylindro-biconical.
Type: Asterophora mucronata L6ger.
Genus 31. Beloides Labbe 1899:27. Epimerite a short stout neck with
spiny globule at af>ex. Spores biconical.
Ty-pe: Beloides firmus (L€ger) Labbe.

Genus 32. Actinocephalus Stein 1848:196. Epimerite sessile or with
short neck, at apex of which are eight to ten short sharp
or simple digitiform processes. Spores biconical.
Type: Actinocephalus conicus (Duf.) Frantz.

Genus ii. Taeniocystis Leger 1906:307. Epimerite small sessile sphere
set with six or eight recurved hooks. Deutomerite divided
by septa into many linear segments. Spores biconical.
Tjrpe: Taeniocystis mira L^ger.
Genus 34. Stictospora L^ger 1893:129. Epimerite with a short neck, a
spherical crateriform ball with twelve backwardly-directed
laminations set close to neck. Spores biconical, slightly
curved.
Type: Stictospora provincialis L6ger.
Genus 35. Bothriopsis Schneider 1875:596. Epimerite sessile, small,
ovoidal, set with six or more long slender filaments directed
upward. Spores biconical. Protomerite wider than
deutomerite, septum convex upward.
Type: Bothriopsis hislrio Schn.
Genus 36. Coleorh^nchus Labbe 1899:23. Protomerite a shallow disc
with scalloped cape down over upper part of deutomerite.
Spores biconical.
Type: Coleorhynchus heros (Schn.) Labbe.

Genus 37. Legeria Labbe 1899 :24. Protomerite broader than deutomerite,
septum convex upward, spores cylindro-biconical. (Epi-
merite unknown.)
Type: Legeria agilis (Schn.) Labbe.
Genus 38. Geneiorhynchus Schneider 1875:594. Epimerite a tuft of
short bristles at apex of a long slender neck, spores cylindro-
biconical.
Type: Geneiorhynchus monnieri Schn.

Genus 39. Hoplorhynchus Carus 1863:570. Epimerite a flat disc with
eight to ten short digitiform processes at apex of a long
neck. Spores biconical.

Type: Hoplorhynchus oligacanthus (von Sieb.) Schn.
Genus 40. Phialoides Labbe 1899:24. Epimerite a cushion set peripher-
ally with stout teeth around the base a collar larger than
the cushion, upon a long slender neck. Spores biconical.

Type: Phialoides ornata (Leger) Labb6.

Family 8. Acanthosporidae Leger 1892:167. Sporonts solitar>-, epimerites
complex. Cyst dehiscence by simple rupture. Spores with
equatorial and polar spines.

14 ILLINOIS BIOLOGICAL MONOGRAPHS [14

Genus 41. Acanthospora L^ger 1892:145. Epimerite a simple conical
knob. Spores biconical with a row of equatorial and one
of polar spines.
Type: Acanthospora pileata L^ger.
Genus 42. Corycella L^ger 1892:144. Epimerite globular, with eight
large recurved hooks. Spores biconical with one row of
polar spines.
Type : Corycella armata L6ger.
Genus 43. Prismatospora Ellis 1914:215. Epimerite subglobular with
eight lateral recurved hooks. One row of spines at each
pole. Spores hexagonal.
Type: Prismatospora evansi Ellis.
Genus 44. Ancyrophora L^ger 1892:146. Epimerite globular with
five to ten backwardly-directed digitiform processes.
Spores biconical with one row of equatorial and one of
polar spines.
Type: Ancyrophora gracilis lAgei.
Genus 45. Cometoides Labb6 1899:29. Epimerite a globe with six to
eight long slender filaments directed upward. Spores
cylindro-biconical with one row of polar and two of equa-
torial spines.
Type: Cometoides crinitus (L^ger) Labb6.
Family 9. Menosporidae L6ger 1892:168. Sporonts solitary. Epimerite a
large cup bordered with hooks and placed on a long slender
neck. Cyst dehiscence by simple rupture. Spores cres-
centic, smooth.
Genus 46. Menospora L6ger 1892:151, 168. Characters of the family.
, Type: Menospora polyacantha L^ger.

Family 10. Stylocephalidae Ellis 1912:25. Sporonts solitary, epimerites varied.
Nucleus ovoidal. Dehiscence by pseudocyst. Spores hat-
shaped, in chains.
Genus 47. Stylocephalus Ellis 1912:25. Dilated papilla at end of a long
neck. Spores hat-shaped. Cysts covered with minute
papillae.
Type: Stylocephalus oUongatus (Hamm.) Wats.
Genus 48. Bulbocephalus Watson 1916b :66. Epimerite a dilated papilla
in middle of a long slender neck.
Type: Bulbocephalus wardi Wats.
Genus 49. Sphaerorhynchus Labb6 1899:32. Epimerite a small sphere
at the end of a long slender neck.
Type: S phaerocystis ophioides (Schn.) Labb^.
Genus 50. Cystocephalus Schneider 1886:99. Epimerite a large lance-
shaped papilla at end of a short neck. Spores hat-shaped.
Type: Cystocephalus algerianus Schn.
Genus 51. Oocephalus Schneider 1886:101. Epimerite a sphere at apex
of a short stout neck.
Type : Oocephalus kispanus Schn.
Genus 52. Lophocephalus Labb^ 1899:31. Sessile crateriform disc with
crenulate periphery, set at the base with many short
upwardly-directed digits.
Type: Lophocephalus insignis (Schn.) Labb^.

15] STUDIES ON GREGARINES—KAMM IS

Family 11. Dactylophoridae L^ger 1892:165. Sporonts solitary, epimerites
highly complex, asymmetric, digitate. Cyst dehiscence
by pseudocyst, spores eIongate-C}'lindrical. Parasites in
Chilopods.
Genus 53. Echinomera Labb6 1899:16. Epimerite an eccentric cone with
eight or more short upwardly-directed digitiform processes,
persistent. Cyst dehiscence by simple rupture, spores
cylindrical, extruded in chains.
Type: Echinomera hispida (Schn.) Labb€.
Genus 54. Rhopalonia L^ger 1893 :1285. Epimerite a subspherical cushion
with ten or more short thick outwardly directed digits.
Pseudocyst. Spores cylindrical.
Type: Rhopalonia geophili L^ger.
Genus 55. Trichorhynchus Schneider 1882:438. Protomerite with long
slender neck dilated at end in an eccentric cone. Cyst-
dehiscence by pseudocyst, spores cylindrical to ellipsoidal.
Type: Trichorhynchus pulcher Schn.
Genus 56. Seticephalus Kamm 1922. Epimerite a dense tuft of short
upwardly-directed brush-like bristles on top of the broad
fiat protomerite.
Type: Seticephalus elegans (Pinto) Kamm.
Genus 57. Dactylophorus Balbiani 1889:41. Protomerite dilated laterally
at the top and set with peripheral digitiform processes.
Cyst dehiscence by pseudocyst, spores cylindrical.
Type: Dactylophorus robustus Leger.
Genus 58. Nina Grebnecki 1873: ? Protomerite two long lobes fused at
one end, set peripherally with teeth and long filaments.
Cyst dehiscence by pseudocyst. Spores long-ovoidal, in
chains obliquely.
Type: Nina gracilis Grebnecki.
Genus 59. Acutispora Crawley 1903:632. Epimerite not described.
Spores long, ellipsoidal with a lengthwise-directed rod at
each end. Cyst dehiscence by pseudocyst.
Type: Acutispora macrocephala Crawl.

GENERA OF UNCERTAIN POSITION

Genus 60. Metamera Duke 1910:261. Epimerite an eccentric cone set
peripherally with numerous branched digitiform processes.
Cyst dehiscence by simple rupture. Spores obese biconical.
Type: Metamera schubergi Duke.
Genus 61. Agrippina Strickland 1912:108. Sporonts solitary, epimerite
a circular disc armed with peripheral digitiform processes,
on a short neck. Spores long ovoidal.
Type: Agrippina bona Strick.
Genus 62. Ganymedes Huxley 1910:155. Sporonts associative. A
motile stalked sphere at anterior end, cup at posterior end.
No septum, no true epimerite. Cysts spherical. Ali-
mentary tract of Crustacea.
Type: Ganymedes anaspidis Hux.
Genus 63. Nematoides Mingazzini 1891:233. Dicystid, no septum in
sporonts. Epimerite forked, at apex of a long neck.
T3rpe: Nematoides Jusijormis Ming.

16

ILUNOIS BIOLOGICAL MONOGRAPHS

[16

A GROUP OF TABLES SHOWING THE PHYLOGENETIC RELA-
TIONSHIPS OF GREGARINES

Table I. Showing tht InUrmediate Position of Two Families, LECUDINIDAE AND POLYRHABDINIDAE

Septum

Epimerite, character

Spores

Host

Habitat

Tribe Acephalina

Absent

Absent

Biconical or with
dissimilar poles

Ecbinoderms, ma-
rine and terres-
trial Annelids, a
few ArthropKxls

Generally coel-
omic

Family Lecudi-

nidae
Family Poly-

rhabdinidae
Tribe Cephalina

Absent
Present
Present

Present, simple
Present, varied
Present, varied

Dissimilar poles

?
All poles alike

Marine Annelida
Marine Annelida
Arthropoda

Digestive tract
Digestive tract
Digestive tract

Table 2. Showing Relalionships of the Families of the Tribe Cephalina

Family Sporont Epimerite Develop- Cyst de- Spores Host

ment hiscence

Remarks

1. Lecudinidae

2. Polyrhabdinidae

See Table I

3. Cephaloidophori-

Twos

Absent

Intracellular

Simple rup-

Ovoidal

Crustacea

dae

ture

with equa-
torial line

4. Stenophoridae

Solitary

Absent or ru-
dimentary

Intracellular

Simple rup-
ture

Ovoidal
with equa-
torial line

Diplopods

Solitary or

Simple, sym-

Extra-cellu-

Spore-ducts

Ellipsoidal

Insects

S. Gregarinidae

in associa-
tions up
to 12

metrical

lar

or simple
rupture

chiefly

6. Didymophyidae

Associations

Small, simple

(C

Simple rup-

Ellipsoidal

Crustacea

No septum

of 2 or 3

papilla

ture

and Cole-
optera

in satellites

7. Actinocephalidae

Solitary

Complex

<i

Simple rup-
ture

Irregular or
biconical or
cylindro-
biconical

Insects and
Chilopods

8. Acanthoeporidae

Solitary

Complex

It

Simple rup-
ture

With equa-
torial and
polar spines

Insects

9. Menosporidae

Solitary

Long neck,
cup with
spines at top

l€

Simple rup-
ture

Crescentic

Insects

10. Stylocephalidae

Solitary

Long neck
(except one
genus) with
simple apex

t(

Pseudocyst

Irregular
brown or
black

Insects

11. Dactylophoridae

Solitary

Highly com-
plex asym-
metric digi-
tate

((

Pseudocyst

Elongate-
cylindrical

Chilopods

17]

STUDIES ON GREGARINES—KAMM

Table 3. Shoving Relationships Between Genera in the Family Gregarinidae

17

Genus

Sporont

Epimerite

Cyst dehiscence

Spores

Remarks

1. Leidyana

Solitary

Sessile globe

Spore-ducts

Barrel-shaped, in
chains

2. Gregarina

In twos

Simple globular or cy-
lindrical papilla

Spore-ducts

Barrel-shaped, in
chains

3. Protomagalhaen-

sia

Several

?

?

Barrel-shaped,
with spine at
each comer

Sporonts attenu-
ate, attachment
often lateral

4. Gamocystis

In twos

?

Spore-ducts, spor-
ulation partial

Cylindrical

Protomerite only
in trophic stages

5. Hyalospora

In twos

Simple globular knob

Simple rupture

Ellipsoidal

Endoplasm
yellow-orange

6. Hirmocystis

Two to 12

Small cylindrical pa-
pilla

Simple rupture

Ovoidal

7. Euspora

In twos

?

Simple rupture

Prismatic

8. Sphaerocystis

Solitary

?

Simple rupture

Ovoidal

Protomerite
only in trophic
stages

9. Cnemidospora

Solitary

?

Simple rupture

Ellipsoidal

Protomerite
one-half gray,
other yellow-
green

10. Uradiophora

In twos

Elongate papilla

Simple rupture

Spherical or
subspherical, in
chains

Deutomerite
with small ap-
pendix, Crus-
tacea

11. Pyxinoides

In twos

Slightly stalked globu-
lar papilla with 16
longitudinal furrows
and a small cone at
apex

?

Crustacea

Table 4, Showing Relationships Between Genera in the Family Actinocephalidae

Genus

Epimerite

Spores

Remarks

1. Amphoroides

2. Pileocephalus

3. Stylocystis

4. Discorhynchus

5. Steinina

6. Anthorhynchus

7. Sciadophora

8. Amphorocepha-
lus

Sessile globular papilla

Short dilated neck, lance-shaped

cone at apex
Recurved sharply pointed cone
Short neck, large globe with thin

collar around base
A short mobile digitiform process

changing into a flat crenulate

disc
Sessile large fluted flattened disc

A large sessile peripherally crenu-
late disc

Concave peripherally crenulate
fluted disc set upon a short di-
lated neck

Curved biconi-
cal
Biconical

Biconical
Biconical,
slightly curved
Biconical

Biconical
Biconical

Extruded in
chains, laterally

Protomerite with
numerous back-
wardly-directed
vertical lamina-
tions

18

ILLINOIS BIOLOGICAL MONOGRAPHS

[18

Table 4. (Continued)

Genus

Epimerite

Spores

Remarks

9. P3ndnia

Flat crenulate crateriform disc
from center of which rises a long
or short style

Biconical

10. Schneideria

Sessile, a thick horizontal disc

Biconical

No protomerite in

with milled border, short style

adults; a degener-

projecting up from center

ate rather than

rudimentary

character

11. Asterophora

Like last, with central style longer

Cylindro-bicon-
ical

12. Beloides

Short stout neck with spiny glob-
ule at apex

Biconical

13. Actinocephalus

Sessile or with short neck, at apex
8 to 10 short sharp spines or sim-
ple processes

Biconical

14. Taeniocystis

Small sessile sphere set with 6 or

Biconical

Deutomerite di-

8 recurved hooks

vided by septa
into many linear
segments

15. Stictospora

Short neck, spherical crateriform

Biconical,

ball with 12 backwardly-directed

slightly curved

laminate filaments set close to

neck

16. Bothriopsis

Sessile, very small, ovoidal, with

Biconical

Protomerite wider

6 or more long slender filaments

than deutomer-

directed upward

ite, septum con-
vex upward

17. ColeorhjTichus

?

Biconical

Septum convex
upward. Prot. a
shallow disc with
scalloped cape
over deutomerite

18. Legeria

?

Cylindro-bicon-

Protomerite

ical

broader than
deutomerite,
septum convex
upward

19. Geneiorhynchus

Long slender neck, tuft of short

Cylindro-bi-

bristles at apex

conical

20. Hoplorhynchus

Long neck, flat disc with 8 to 10
digitiform processes at apex

Biconical

21. Phialoides

Long slender neck, at apex a
broad cushion with peripheral
teeth surrounded at base with a
collar larger than the cushion

Biconical

19]

STUDIES ON GREGARINES—KAMM

19

The epimerites of this family are surprisingly well correlated. The genera with short
necks form a regular succession; the simple globe (1) becomes slightly modified (2), (3); then
a disc develops at the base (4), (5), (6), (7), (8); with a style rising in the center (9), (10).
This seems to be the end of one line of development.

The simple sessile or nearly sessile globule becomes spiny (12), (13); the spines become
simple digitiform processes or recurved hooks (14); the recurved hooks become attached to
the neck along their whole length (15). The processes change into long slender filaments (16)
— and probably (17) and (18). The neck becomes long and slender, and at the apex there
develops a tuft of short bristles (19), a flat disc with 8 to 10 peripheral digits (20), or stout
teeth (21).

Table 5. S homing the Relationships Between the Genera in the Family Acanthosporidae

Genus

Epimerite

Spores

1. Acanthospora

2. Corycella

3. , Prismatospora

4. Ancyrophora

5. Cometoides

Simple conical knob

Globular, with 8 large recurved
hooks

Subglobular, with 8 lateral re-
curved hooks

Globular, with 5 to 10 backward-
ly directed digitiform processes

Globe, with 6 to 8 long slender
filaments directed upward

Biconical, row of equatorial and one
of polar spines
Biconical, one row of polar spines

One row of spines at each pole.
Hexagonal

Biconical, with one row of equato-
rial and one of polar spines.

Cylindro-biconical, with one row of
polar and two of equatorial
spines

Table 6. Showing Relationships Between the Genera in the Family Stylocephaiidae

Genus

Epimerite

Spores

Remarks

1. Stylocephalus

Long neck, dilated papilla at apex

Hat-shap>ed

Cysts covered with
small papillae

2. Bulbocephalus

Long neck, dilated papilla mid-
way

?

3. Sphaerorhynchus

Long neck, small sphere at apex

?

4. Cystocephalus

Short stout neck, large lance-
shaped papilla at apex

Hat-shaped

5. Oocephalus

Short neck with sphere at apex

?

6. Lophocephalus

Sessile crateriform disc with cren-
ulate periphery, set at base with
many short upwardly-directed

Hat-shaped

digits

I can see very little relationship between the genera of this family as a whole. (1), (2),
and (3) are related, as are (4) and (5), but the three groups seem to bear no epimeritic rela-
tionship whatever. The hat-shaped spores, however, connect the seemingly aberrant genus
(6), with the type.

20

ILLINOIS BIOLOGICAL MONOGRAPHS

[20

Table 7. Showing Relationships Between the Genera in the Family Dactylophoridae

Genus

Character of protomerite or
epimerite

Cyst dehis-
cence

Spores

Remarks

1. Echinomera

Epimerite an eccentric cone

Simple rup-

Cylindrical in

with eight or more short up-

ture

chains

wardly-directed digitiform

processes, persistent

2. Rhopalonia

Epimerite a subspherical cush-
ion with ten or more short
thick outwardly-directed dig-
its

Protomerite with long slender

Pseudocyst

Cylindrical

No proto-
merite in
adult

3. Trichorhyn-

Pseudocyst

Cylindrical to

chus

neck dilated at end in an ec-
centric cone

ellipsoidal

4. Seticephalus

A dense tuft of short upward-
ly-directed brush-like bristles
on top of a broad flat proto-
merite

?

?

5. Dactylophorus

Protomerite dilated laterally
with peripheral digitiform
processes

Pseudocyst

Cylindrical

6. Nina

Protomerite two long lobes
fused at one end, set peri-
pherally with teeth and long
filaments

?

Long-ovoidal

Spores in
chains
obliquely

7. Acutispora

?

Pseudocyst

Long-ellipsoidal
with a length-
wise-directed
rod at each end

21]

STUDIES ON GREGARINES—KAMM

21

SYNOPSES OF THE GREGARINES FROM VARIOUS GROUPS OF

THE ANIMAL KINGDOM

LIST OF SPECIES FOUND IN THE
Parasite

LECUDINIDAE

Lecudina pdlucida (Kolliker) Mingazzini

Type species
Lecudina leuckarii Mingazzini
Lecudina aphrodiiae (Lankester) Kamm
Lecudina elongata (Mingazzini) Kamm
Lecudina hekrocephala (Mingazzini) Kamm
Lecudina polydorae (L^ger) Kamm

Lecudina sp. Saint- Joseph

Lecudina legeri (Brasil) Kamm
Lecudina sp. Faria, Cunha, and Fonseca

PHYLUM COELHELMINTHES*
Host

Nereis cullrifera

N. beaucourdrayi Aud.

Sagitta sp.

Aphrodite aculeata

Lumbriconereis impatiens Clap.

Nephthys scolopendroides delle Chiaje

Poly dor a agassizi Clap.

P. ciliata

Polymnia nehulosa M.

Notomastus exserlilis N. S.

Petaloproctus terricola Qfg.

Glycera convolula Kef.

Polydora socialis Schm.

POLYRHABDINrDAE

Polyrhabdina spionis (Kolliker) Mingazzini

Tj^e species
Polyrhabdina brasili Caullery and Mesnil
Polyrhabdina pygospionis Caullery and

Mesnil
Sycia inopinata lAger

Type species
Ulivina elUptica Mingazzini

Tj^e species

Ulivina rhynchoboli (Crawley) Kamm

Scolohpsis fuliginosa

S. ciliala

Spio martinensis

Pygospionis seticornis
Audouinia Lamarkii

Audouinia filigera (d. Chiaje)
A . tenlaculata Mont.
Petaloproctus terricola Qfg.
Nicolea venustula Mont.
Polymnia nebulosa Mont.
Rhynchobolus americanus Ver.

GENUS OP UNCERTAIN POSITION

Metamera schubergi Duke

Type species
Metamera sp. Ellis

Clossosiphonia complanata
Hemichpsis marginat<i
Clepsine elongata

SPECIES OF UNCERTAIN POSITION

? Taeniocystis ligeri Cognetti de Martiis

Kynotus Pittarelli

Family LECUDINIDAE Kamm (nov. fam.)
Epimerite symmetrical, simple. Body non-septate. Spores ovoidal,
asymmetrical, thickened at one pole. Intestine of marine annelids.

*Tbe parasites are arranged in chronological order under each genus in this and all similar succeeding lists.

22 ILLINOIS BIOLOGICAL MONOGRAPHS [22

DISCUSSION OF THE NEW FAMILY LECUDINIDAE

The type genus of this family, Lecundina was named by Mingazzini
in 1891. Two years later Leger, working independently, designated a
new genus Doliocystis for the species described by Mingazzini. The
earlier work has been overlooked by subsequent workers while the name
used by Leger has come into frequent usage, the genus being raised to
family rank (Doliocystidae) by Labbe, in 1899.

In Table I of Chapter II, is shown the intermediate position of the family
Lecudinidae and the somewhat related family, the Polyrhabdinidae. The
Lecudinidae are related to the Tribe Acephalina, for they are non-septate,
there being but one division in the body at all stages of development.

All the members of the Tribe Cephalina, on the other hand, are char-
acterized by the presence of a septum, which divides the body into a
protomerite and deutomerite, if not in the adult, at least in the trophic
stages of development. When the septum is absent from the adult, it is
clearly a degenerative rather than a rudimentary character, all other
generic features conforming to the type.

(Cf. Schneideria, Sphaerocystis, Rhopalonia, Gamocystis.)

The Lecudinidae, however, possess only the epimeritic demarkatioQ
from the rest of the body, and when this structure disappears, the body is
unilocular. Leger (1893) remarks:

... la gr6garine pr^sente tou jours deux segments: le segment intra-cellulaire ou
ipimirite, et le segment extra-cellulaire dans lequel s'est port6 le noyau. C'est done seulement
i ce moment que la gr^garine se montr6 comme une veritable dicystidee. . . . les jeunes
individus abandonnent leur ipimirite et deviennent libres dans I'intestin, prSsentant a lore
toutes les apparences de veritable Monocystis ....

In the type species, Lecudina pellucida (Koll.) Ming., there is a differ-
entiation in the protoplasm of the anterior end (what would be the pro-
tomerite in polycystids) from that of the remainder of the body. This
was illustrated by Kolliker (my fig. 126) but not mentioned by Leger in
1893. Brasil (1909) illustrates this differentiation clearly in both tropho-
zoite and sporont (my figs. 134 and 135).

If normally present, and I have no doubt that it is but often not
mentioned or discovered, this character is an important one in assigning
the family in question to an intermediate position between the monocystids
and the polycystids.

In its cyst-formation and spore type, the family follows the polycystid
type.

After the above had been written, this statement from Minchin (1903)
was found:

The possession of an epimerite is a feature is used for classifying which the Gregarines,
and the legion Eugregarinae is separated into the two sub-orders Cephalina and Acephalina,
according to the presence or absence of this appendage. As a general rule the forms which

23] STUDIES ON GREGARINES—KAMM 23

possess an epimerite have the body behind it divided into protomerite and deutomerite by a
septum, and have hence been termed Polycystida . . . , while those without an epimerite
are also without a septum; hence Monocystida . . . But in one family, Doliocystidae, Labb€,
an epimerite is present, and may attain a considerable size . . . without any septum dividing
the rest of the body ... It is purely a matter of definition whether those fonns be con-
sidered as Cephalina without a septum, or as Monocystis with an epimerite. The Cephalina
in which the body is non-septate are sometimes distinguished as Dicystida from those in
which there is a distinct protomerite and deutomerite (Tricystida) . These terms are to be
understood, however, in a purely descriptive sense, and cannot be used for classificatory
purposes, as there is no doubt that many dicystid species are derived from tricy^tid forms
secondarily, by obUteration of the protomerite .... On the other hand, such forms as the
Doliocystidae .... appear to be truly and primitively dicystid, and are to be regarded as
intermediate forms transitional from Acephalina to Cephalina.

Genus LECUDINA Mingazzini 1891: 469
Body non-septate, epimerite regularly simple. Spores ovoidal, thick-
ened at one pole.

LECUDINA PELLUCIDA (Kolliker) Mingazzini Type species

1848 Gregarina pelliicida

1851 Gregarina pellucida

1859 Gregarina pellucida

1872 Monocystis nereidis

1891 Lecudina pellucida

1893 Lecudina pellucida

1893 Doliocystis pellucida

1899 Doliocystis pellucida

1903 Doliocystis pellucida

1909 Doliocystis pellucida

1913 Doliocystis pellucida

Sporonts solitary, ellipsoidal or "bottle-ehaped," rectangular or rounded
at anterior end and broadly rounded at posterior. Nucleus spherical.
Epimerite a simple small papilla.

Cysts small, dehiscence by simple rupture, spores ovoidal, 7 by 5^1, with
thickening at one pole.

Taken at Triest, Naples, and in the Gulf of Marseilles.

Intestine of Nereis cultrifera Grube and N. beaucourdrayi Aud. Kol-
liker's figure indicates a differentiation in the protoplasm of the anterior
end of the body from that of the remainder.

Mingazzini describes the body as nearly transparent, especially at the
anterior end, and both mentions and illustrates the retractility of the
anterior end within the body.

[Figure 126]

Kolliker

1848: 35

Diesing

1851: 17

Diesing

1859: 739

Lankester

1872:343

Mingazzini

1891:469

Mingazzini

1893: 51

Leger

1893: 204

Labbe

1899:33

Minchin

1903: 202, 327

Brasil

1909: 119

Ellis

1913: 287

24

ILLINOIS BIOLOGICAL MONOGRAPHS

[24

LECUDINA LEUCKARTI Mingazzini

1860 No name Leuckart 1860: 263

1891 Lecudina Leuckarti Mingazzini 1891: 469

1893 Lecudina Leuckarti Mingazzini 1893: 51

Similar to the type species, the anterior end not constricted bottle-like.
Intestine of Sagitta sp.

LECUDINA APHRODITAE (Lankester) Kamm

Figure 125]

1863

Monocystis aphroditae

Lankester

1863: 90, 94

1871

Unnamed

Stuart

1871:498

1899

Doliocystis aphroditae

Labbe

1899: 33

1903

Doliocystis aphroditae

Minchin

1903: 176, 202, 326

1909

Doliocystis aphroditae

Brasil

1909: 120

1922

Lecudina aphroditae

Kamm

1922 (this paper)

One-half inch in length. "This is the only unilocular form of Gre-
garina which at present has been found provided with a proboscis."
Lankester.

Intestine of Aphrodite aculeata.

LECUDINA ELONGATA (Mingazzini) Kamm

[Fig

ures 134 and 135

1891

Ophioidina elongata

Mingazzini

1891:471

1893

Ophioidina elongata

Mingazzini

1893: 53

1899

Doliocystis elongata

Labbe

1899: 33

1903

Doliocystis elongata

Minchin

1903: 327

1908

Doliocystis elongata

Brasil

1908:355

1908

Doliocystis elongata

Brasil

1908a: 425

1909

Doliocystis elongata

Brasil

1909: 112

1922

Lecudina elongata

Kamm

1922 (this paper)

Sporonts solitary, elongate-cylindrical, attaining 500/i by 40/i. Pro-
toplasm of anterior end highly specialized from that of remainder. Nu-
cleus large, spherical, with several karyosomes.

An intracellular stage was encountered by Brasil, who thinks it may be
schizogonic in character. It is possible, however, that it may have been a
portion of the life-history of another parasite,

Mingazzini describes the epimerite as a "piccolo bottone sferico";
Brasil finds a long thread-like filament which penetrates to the base of the
cell, which is certainly atypical.

Intestine of Lumbriconercis impatiens Claparede.

Taken at Naples.

25] STUDIES ON GREGARINES—KAMM 25

In three of Mingazzini's twelve illustrations of the species, there is
some indication of a septum present, the protoplasm being quite different
in the two portions.

I can see no differentiation between the two genera described by Min-
gazzini, Lecudina and Ophioidina, from the meagre data known con-
cerning each. The intracellular development and the spores may reveal
differences, however. From the existing data, I have united the two under
the name of the first-named genus, Lecudina.

LECUDINA HETEROCEPHALA (Mingazzini) Kamm
1891 Ophioidina heterocephala Mingazzini 1891:473

1893 Ophioidina heterocephala Mingazzini 1893: 54

1899 Doliocystis heterocephala Labbe 1899: 34

1903 Doliocystis heterocephala Minchin 1903: 327

1922 Lecudina heterocephala Kamm 1922 (this paper)

Sporonts very elongate, vermiform. Epimerite "a sort of papilla"
at anterior end. Protoplasm of anterior end differentiated from that of
body proper.

Nucleus spherical, with one or two karyosomes.

Intestine of Nephthys scolopendr aides delle Chiaje.

LECUDINA POLYDORAE (Leger) Kamm

1893 Doliocystis poly dor ae

1899 Doliocystis polydorae

1903 Doliocystis polydorae

1909 Doliocystis polydorae

1914 Polyrhabdina polydorae Caullery and Mesnil 1914: 520

1922 Lecudina polydorae Kamm 1922 (this paper)

Epimerite similar to that described for the type species, but larger.
"Epimerite a la forme d'un tronc de cone a petite base inferieure et il se
continue directement avec I'extremite anterieure du second segment,
allongee en forme de col." Leger.

Intestine of Polydora agassizi Clap.; P. ciliata.

Taken in the Gulf of Marseilles, France.

LECUDINA sp. Saint- Joseph

1907 Doliocystis sp. Saint- Joseph 1907: 145, 172, 173

1911 Doliocystis sp. Sokolow 1911:287

1922 Lecudina sp. Kamm 1922 (this paper)

Measurements are given of sporonts from various polychaetes, as
follows: 84m by 29/x; 470/1 by 63m; 300m by 48m. It is quite possible that
several species are involved, but no description is given of any one.

L6ger

1893: 205

Labbe

1899: 33

Minchin

1903: 327

Brasil

1909: 119

26 ILLINOIS BIOLOGICAL MONOGRAPHS [26

Intestine of Polymnia nebulosa M.; Notomastus exsertilis N.S.; and
Petaloproctus terricola Qfg.

LECUDINA LEGERI (Brasil) Kamm

1909 Doliocystis legeri Brasil 1909: 71, 123

1911 Doliocystis legeri Sokolow 1911: 284

1922 Lecudina legeri Kamm 1922 (this paper)

Sporonts cylindrical, lOOiu by 25/x. Nucleus ovoidal, with one karyo-
some. Epimerite a rhizoid filament which penetrates to the base of the
host-cell (Brasil).

Cysts small, 45/x in diameter, spores typical, releasing eight sporozoites.

Intestine of Glycera convoluta Kef.

Taken on the Mediterranean Coast.

LECUDINA sp. Faria, Cunha, and Fonseca

1918 Doliocystis sp. Faria, Cunha, and

Fonseca 1918: 17

1922 Lecudina sp. Kamm 1922 (this paper)

Sporonts spindle-shaped, nucleus spherical.

Host: Polydora socialis Schmerda.

Taken near Rio de Janeiro, Brazil.

Not enough data exists to definitely fix the position of this species.

Family POLYRHABDINIDAE Kamm 1922 (this paper)
Polycystid (septate) gregarines inhabiting the digestive tract of poly-
chaetes. Epimerites varied.

DISCUSSION OF THE NEW FAMILY POLYRHABDINIDAE

The true septate intestinal gregarines of marine annelids fall under
three genera, which are placed in various positions by diflferent workers on
the subject:

(a) Two genera were described very meagerly by Mingazzini in 1891
under one name, Polyrhabdina, two very different parasites being found in
the same host material. The one has been removed to the Schizogregarines
(Selenidium), while the other represents the genus as named, and is a
pyriform septate species which possesses an epimerite in the shape of a
corona of hooks.^

(b) The genus Sycia Leger 1892 is characterized by its septate sporonts
and unique epimerite in the form of a large rounded papilla with a thick

• Mingazzini's description follows: . . . specie dimorfe. Individue a forme di nematode
e pirifonni. I primi hauno il corpo allungato fusiforme e la cuticula striata longitudinalmente
du rialzi numerosi finissimi.

271 STUDIES ON GREGARINES—KAMM 27

ring or collar around the base. The deutomerite of the adult sporont,
only, contains numerous elongate crescentic or ellipsoidal inclusions
probably "reserve protoplasm."

(c) The third genus, Ulivina Mingazzini 1891, has little to characterize
it. In fact so little is known concerning its development that it cannot be
placed with either of the two named genera and hence must stand distinct
until its position shall have been proven. The epimerite, is simple so far
as known and cyst and spores are unknown.

The cysts and spores are still unknown for all three genera.

Since a septum is present without exception in all three, they must be
included in the suborder Cephalina, but stand near the borderline with the
Acephalina because of their presence only in polychaetes.

Genus POLYRHABDINA Mingazzini 1891:229
(Polyrhabdina Labbe 1899:48)
Septate, sporonts flattened, ovoidal, epimerite a corona of hooks. In-
testine of polychaetes of the family Spionidae. (Cyst and spores unknown.)

POLYRHABDINA SPIONIS (KoUiker) Mingazzini Type Species

[Figures 128, 129]

1848 Gregarina Spionis KoUiker 1848: 4

1851 Gregarina Spionis Diesing 1851:18

1891 Polyrhabdina Spionis Mingazzini 1891:229

1893 Polyrhabdina Spionis Mingazzini 1893:56

1903 Doliocystis sp. Minchin 1903: 327

1914 Polyrhabdina spionis Caullery and Mesnil 1914: 516

Septum present. Sporonts 100 by ZSn, ellipsoidal, nucleus spherical,
one large karyosome.

Epimerite a flattened apical corona of 8 to 10 ameboid digitiform
processes frequently bifurcate.

Development extracellular. Cyst and spores unknown. •

Intestine of Scololepsis fuliginosa; S. ciliata (?).

Taken at Naples.

Minchin mentions a species from the same host, but places it in the ge-
nus Doliocystis. Since no data is given and the hosts are identical, it is
placed here.

POLYRHABDINA BRASILI Caullery and MesnU
1914 Polyrhabdina brasili Caullery and Mesnil 1914: 518

Sporonts ovoidal, 200)u in length. Epimerite characteristic, but spines
shorter than type.

Cyst and spores unknown.

Host: Spio martinensis.

28 ILLINOIS BIOLOGICAL MONOGRAPHS [28

POLYRHABDINA PYGOSPIONIS Caullery and Mesnil
1914 Polyrhahdina pygospionis Caullery and Mesnil 1914: 520

No description given.
Host: Pygospionis seticornis.

Genus SYCIA Leger 1892: 52

Polycystid intestinal parasite of marine annelids. Epimerite knobbed,
bordered by a thick ring. Protomerite subspherical, deutomerite conical,
with numerous inclosures.

SYCIA INOPINATA

Leger Type Species

Figures

118 and 119]

1892

Sycia inopinata

Leger

1892:

52,90

1899

Sycia inopinata

Labbe

1899:

34

1903

Sycia inopinata

Minchin

1903:

203, 326

1909

Sycia inopinata

Brasil

1909:

121

Septate gregarine, ellipsoidal. No dimensions given. Ratio LP : TL
:: 1:6; WP : WD :: 1 : 1.5. Protomerite conoidal, broadest at septum.
Deutomerite ellipsoidal, tapering rapidly to a sharp point. Nucleus large,
spherical, one karyosome.

Epimerite an elongate papilla upon a short neck around which is a
collar a little larger than the papilla itself.

Intestine of Audouinia Lamarkii {A. tentaculata — Brasil).

Taken at Belle-Isle-sur-Mer, France.

Caullery and Mesnil (1914) give the host name as the latter above, and
note

. . . c'est une espece voisine, sinon identique, que Mingazzini (1891) a d6nomm6e
Ulivina n. g. elliptica

but since the cyst and spores of both species are unknown, this cannot at
present be determined.

This species possesses characteristic elongate crescentic or biconical
inclusions in the deutomerite only of adult sporonts. (Fig. 119). L6ger
offers the hypothesis that they may be condensations of protoplasm in re-
serve masses.

Brasil found this species with its epimerite and the inclusions, thus
verifying Leger's work of sixteen years previous.

Genus ULIVINA Mingazzini 1891: 235

Body elongate-ellipsoidal, "external membrane continuous around
animal." Protomerite the more dense. Epimerite simple, spores unknown.
Intestine of marine polychaetes.

29] STUDIES ON GREGARINES—KAMM 29

ULIVINA ELLIPTICA Mingazzini Type Species

Figure 127

1891

Ulivina elliptica

Mingazzini

1891:235

1899

Ulivina elliptica

Labbe

1899: 34

1903

Ulivina elliptica

Minchin

1903: 203, 326

1907

Ulivina elliptica

Saint-Joseph

1907: 164, 174

Sporonts solitary, ellipsoidal, tapering to a sharply rounded posterior
end. Nucleus ovoidal, two or three karyosomes. Dense and opaque.
No dimensions given by the original author. Ratio LP:TL::1:4
to 7; WP: WD:: 1.1:1.

Saint- Joseph mentions dimensions as 150 by 90)u; 75 by 33;u. He illus-
trates (his fig. 106) a trophozoite with a simple, small, papillate epimerite.

Taken at Naples and off the coast of France.

Intestine of Audouinia filigera (delle Chiaje); A. tentaculata Mont.,
Petaloproctus terricola Qfg., Nicolea venustula Mont., and Polymnia nebu-
losa Mont.

ULIVINA RHYNCHOBOLI ((

[Crawley)

Kamm

1897

Unnamed

Porter

1897a: 8

1903

Doliocystis rhyncoholi

Crawley

1903: 56

1913

Doliocystis rhyncoholi

Ellis

1913:287

1922

Ulivina rhynchoboli

Kamm

1922 (this paper)

Sporonts attain 700;u in length. A distinct septum, with protoplasm
in protomerite the more dense. Epimerite a small pointed papilla with a
long slender filament at the end. Nucleus small, spherical.

Intestine of Rhynchoholus americanus Verrill.

While the epimerite is described for this and for the preceding species,
I am not ready to state that it is the type for the genus.

Porter notes a peculiar attachment of the parasite while free in a watch-
glass to the glass, being attached by the anterior end of the protomerite
with such force that the animal cannot be withdrawn without destroying
it.

Genus of Uncertain Position
METAMERA Duke 1910:261

Sporonts solitary, epimerite subconical, apex eccentric with corona of
numerous branched digitiform appendages. Cyst dehiscence by simple
rupture. Spores biconical.

30 ILLINOIS BIOLOGICAL MONOGRAPHS [30

METAMERA SCHUBERGI Duke Type Species
[Figures 131, 132, and 133]

1910 Metatnera schubergi Duke 1910: 261
1913 Metatnera schubergi Ellis 1913: 285

Sporonts 150 by 45/1. Deutomerite with one to three septa posterior
to the nucleus. Cysts spherical, spores ovoidal 9 by 7/i.

Epimerite subconical, apex eccentric with corona of numerous branched
digitiform processes.

Taken at Heidelberg and Cambridge, Eng,

Intestine of Glossosiphonia complanata and Hemidepsis marginata.

I am not ready to assign this species to a position in the family Dactylo-
phoridae, as placed by the author. The method of cyst-dehiscence is differ-
ent from the type for that family and the spores are not elongate cylindrical.
The host is so far removed from the Chilopoda (to which the named family
is confined) that I feel certain the species belongs in a group as yet unde-
scribed. When more similar species shall have been described its position
can the better be determined.

METAMERA sp. Ellis

1913 Metatnera sp.? Ellis 1913:285,287

Host: Clepsitie elotigata.

Species of Uncertain Position

TAENIOCYSTIS LEGERI Cognetti de Martiis
[Figure 130]

1911 Taeniocystis legeri Cognetti de Martiis 1911:247

Polycystid gregarine, solitary, protomerite and deutomerite segmented
by 16 to 19 septa. 700 to I6OO/1 in length.

Epimerite, cyst, and spores unknown.

Taken at Moramanga, Madagascar.

Host: Kytiotus Pittarelli (oligochaete). Coelomic.

The "protomerite" is divided into three segments, which is unique.
The parasite is coelomic rather than intestinal, as are all other polycystid
gregarines. The epimerite is unknown. For these reasons it is not
placed with the type for the genus. The single nucleus seems to place it
with the Protozoa; otherwise it might be a haplosporidian.

LIST OF SPECIES FOUND IN THE PHYLUM MOLLUSCA

Species of Uncertain Position
Gregaritia pterotracheae (Stuart) Labbe Pterotrachea sp.

31] STUDIES ON GREGARINES—KAMM 31

GREGARINA PTEROTRACHEAE (Stuart) Labbe

[Figure 136]

1871 Zygocystis Pteroiracheae Stuart 1871:498

1891 Zygocystis Pteroiracheae Mingazzini 1891: 235

1899 Gregarina pteroiracheae Labbe 1899: 37

1903 " " Minchin 1903: 338

Contour of body ovoidal, typically that of the genus Gregarina, biasso-
ciative. Dimensions not given. Ratio LP :TL :: 1 :4; WP : WD ::
1 : 1.5. Protomerite dome-shaped with slight constriction in middle.
Deutomerite ovoidal, well-rounded posteriorly. Epicyte unusually thick.
Nucleus large, spherical, one to four karyosomes.

Cyst formed of one individual seen, embedded in muscular tissue of host.

Coelom of Pierotrachea sp.

Taken at Odessa.

This species is regarded as of uncertain position because it is the first
species of the genus Gregarina to be found in the body-cavity of a host.
It is the only species described from the molluscs, where one would not
expect to find a polycystid and biassociative gregarine. This phylum cer-
tainly offers splendid research opportunities, if not to substantiate to repu-
diate the above work.

Stuart describes in detail the motion of the species, and because I
believe it is the first good record of the movements of gregarines, it is
reproduced here, as follows:

Die Bewegungen des Thieres sind schlangelnder Art, aber neben den Hauptcontractionen,
als deren Resultat die Vorwartsschiebung der Gregarine erscheint, bemerkt man eine Reihe
partieller Contractionen, welche die ausseren Contouren des Korpers wellenformig umandern.
Bei diesen Contractionen wird die weiche innere Kornermasse hin und hergeschoben und
nimmt die durch die partiellen Gestaltanderungen des contractilen Schlauches gebildeten
Innenraume ein. Der Nucleus folgt diesen Bewegungen in beschranktem Masse mit.

LIST OF SPECIES FOUND IN THE CLASS CRUSTACEA OF THE PHYLUM

ARTHROPODA

Parasite Host

didymophyidae

Didytnophyes longissima (von Siebold) Frant- Gammarus pulex von Sieb.

zius Orchestia littorea

GREGARINIDAE

Uradiophora cuenoti (Mercier) Mercier Alyaephyra Desmaresli Millet

Type species
Pyxinoides balani (Kolliker) Tr^goubofF Balanus pusillus Ecker

Type species B. tintinnabulum L.

B. amphitrite Darw.
B. eburneus Gould
Pyxinoides cthamaii (L6ger and Duboscq)

Trdgouboff Cthamalus sieUalus Ranz.

32

ILLINOIS BIOLOGICAL MONOGRAPHS

[32

CEPHALOIDOPHORIDAE

Cephaloidophora conformis (Diesing) L6ger

and Duboscq
Cephaloidophora communis Mawrodiadi

Type species

Cephaloidophora fossor (L6ger and Duboscq)

Tr^gouboff
Cephaloidophora ocellata (Ldger and Duboscq)

Kamm
Cephaloidophora maculata L6ger and Duboscq
Cephaloidophora talilri Mercier
Cephaloidophora olivia (Watson) Kamm
Cephaloidophora nigrofusca (Watson) Kamm
Cephaloidophora delphinia (Watson) Kamm
Cephaloidophora ampelisca (Nowlin and

Smith) Kamm

GENEEA OF UNCERTAIN POSITION

Nematoides fusiformis Mingazzini

Type species
Ganymedes anaspidis Huxley

Type species

UNNAMED SPECIES OF UNCERTAIN POSITION

[? gammari von Siebold]
[sp. Mawrodiadi]

[? valettei Nussbaum]
[? praemorsa Diesing)
[? clausii Frenzel]

[? nicaeae Frenzel]
[sp. Minchin]

Pachygraspus marmoratus Fabr.
Balanus improvisiis Darw.
B. tinlinnabulum communis L.
B. improvisus gryphica Miin.
B. perforans Brug.
B. amphitrile Darw.
B. eburneus Gould

Pinnotheres pisum Penn.

Eupagurus Prideauxi Leach
Gammarus marinus Leach
Talitrus saltator Mont.
Libinia dubia
Uca pugnax, U. pugilalor
Talorchestis longicornis Say.

Ampelisca spinipes

Balanus perforalus
Pollicipes cornucopia
Anaspides tasmaniae (Thomp.)

Gammarus pulex

Balanus amphilrite pallidus Darw.

B. amphitrile Darw.

B. eburneus Gould

Pollicipes polymerus Say

Cancer pagurus

Phronima sp.

Phronimella sp.

Hyale pontica Rathke

Nebalia serrata

Family DIDYMOPHYIDAE Leger 1892:105
Sporonts in associations of two or three. No septa in satellites.

Genus DIDYMOPHYES Stein 1848:186
Characters of family, Epimerite a small pointed papilla, cyst dehis-
cence by simple rupture, spores ellipsoidal.

von Siebold

1839: 58

Frantzius

1848: 196

KolUker

1848: 34

Diesing

1851: 15

Diesing

1859: 735

Lankester

1863: 95

Plate

1886: 235

Pfeiffer

1895: 60

Labbe

1899: 8

Minchin

1903: 330, 331

Wellmer

1911: 119

33J STUDIES ON GREG A RINES—KA MM 33

DIDYMOPHYES LONGISSIMA (von Siebold) Frantzius

1839 Gregarina longissima

1848 Didymophyes longissima

1848 Gregarina longissima

1851 Gregarina longissima

1859 Gregarina longissima

1863 Gregarina longissima

1886 Gregarina longissima

1895 Gregarina sp.

1899 Didymophyes longissima

1903 Didymophyes longissima

1911 Gregarina longissima

Sporonts very elongate, cylindrical, vermiform in movements. Ratio
LP : TL : : 1 : 30 ; WP : WD : : 1 : 1 . Primite longer than satellite. Dimen-
sions not given (Kolliker). Nucleus small, spherical, one large karyosome.

Intestine of Gammarus pulex von Siebold, Orchestia littorea.

Taken in Germany.

The species was first described by von Siebold, who also found another
species in Gammarus pulex, typically like the Cephaloidophoridae in outline
but regarded by him as another form of the same species.

Kolliker studied fresh material adding the data that associations exist
of two or three elongate slender individuals arranged in single file and of
two satellites attached to the posterior end of the primite. Kolliker recog-
nized the possibility of a new species being involved in von Siebold' s
material and illustrates this as well as the two modifications mentioned in
the type species.

The exclusion of the doubted form was also recognized by Diesing
(1859).

Lankester (1863) regards all the following as synonymons:

Gregarina longissima Siebold

? Gr. diffluens Diesing

Gr. miliaria Diesing (Actinocephalus Stein)

Gr. putanea Leuckart

Gr. Gammari Siebold (Didymophyes Stein)
His basis was the identity of the hosts only.

Plate found in the intestine and coelom of Gammarus pulex "band-
formig" gregarines 255jLi in max. length, three or four attached together
and regards them as belonging to the species in question.

Minchin (1903) questions the inclusion of the last named host as
authentic.

34 ILLINOIS BIOLOGICAL MONOGRAPHS [34

Leger and Duboscq (1911) have described a new species, Cephaloido-
phora maculata, from Gammarus marinus which is identical with the aber-
rant species referred to above.

Family GREGARINIDAE Labbe 1899:9

Epimerite symmetrical, simple. Sporonts solitary or in association.
Cysts with or without spore-ducts, spores symmetrical.

Genus URADIOPHORA Mercier 1912a

Intestinal parasites. Development extracellular. Epimerite an
elongate papilla. Cysts without sporeducts. Spores spherical or sub-
spherical, not united in chains, faint equatorial line. End portion of deuto-
merite of satellite greatly constricted. Associations of two sporonts.

URADIOPHORA CUENOTI (Mercier) Mercier Type Species

[Figure 63]

1911 Cephaloidophora Cuenoti

1911 Cephaloidophora Cuenoti

1912 Uradiophora Cuenoti
1912 Uradiophora Cuenoti

1912 Uradiophora Cuenoti

1913 Uradiophora Cuenoti

Characters of the genus. Syzygies of two sporonts, infrequently of
three. Sporonts very elongate, cylindrical. Ratio LP:TL::1:20
(primite) 1 : 28 (satellite without appendage). WP: WD:: 1:1.9 to
1 : 2. Sporonts 700^ in max. length. Nucleus sub-spherical. Epimer-
ite an elongate papilla, persistent in sporonts. Deutomerite with small
atrophied appendix. Nucleus in adults spherical, with one large karyo-
some; chromidial bodies frequently found in both protomerite and deu-
tomerite. Cysts ovoidal, 38 to 44/i in diam. Spores 4)u in diam.

Intestine of Atyaephyra Desmaresti Millet.

Taken at Nancy, France.

Genus PYXINOIDES Tregouboff 1912

Intestinal parasites, extracellular in development. Epimerite a short
stype dilated in middle to form a globe with sixteen longitudinal grooves,
the free upper end of the style being about 2n in length. Septum with
small disc-shaped horizontal swelling in middle. Sporont ovoidal, nucleus
large, one karyosome. Syzygies of two sporonts, the primite always
larger. Cyst and spores unknown.

Mercier

1911: 51

Sokolow

1911: 286

Mercier

1912a: xli

Mercier

1912b; 177

Tregouboff

1912: Ivi

Ellis

1913: 264

35] STUDIES ON GREGARINES—KAMM 35

PYXINOIDES BALANI (Kolliker) Tregouboff Type Species
[Figures 65, 82, 88]
1848 Gregarina halani Kollilcer 1848: 35

1848 Stylorhynchus Balani Frantzius 1848: 195

1851 Gregarina Balani Diesing 1851: 10

1859 Gregarina Balani Diesing- 1859: 728

1863 Gregarina Balani Lanlcester 1863: 95

1899 Gregarina halani Labbe 1899: 36

1903 ''Gregarina'' halani Mincliin 1903: 329

1912 Pyxinoides balani Tregouboff 1912: liii

Sporonts associated in pairs, ovoidal to cylindrical. Max. length seen
by Kollilier 63/1, by Tregouboff 130 fi primite, 60/i satellite. Ratio LP :
TL::1:1.3 to 1:3; WP : WD :: 1:1 to 1:1.4. Protomerite dome-
shaped, slightly dilated below middle, terminating in a long cone upon
which is surmounted an epimerite in the form of a short stylet dilated cen-
trally into a bulb (Kollilier). Deep constriction at septum, deutomerite
widest in middle. Nucleus larger, spherical with one karysome.

Intestine of Balanus pusillusEckeT; B.tintinnahulum'L. (Koll.), B. am-
phitrite Darw., B. ehurneus Gould (Treg.).

Taken at Triest, Cette, France.

Kolliker first described this species, although he credits "Professor
Ecker" with having seen it before, evidently without placing it upon
record.

Frantzius reasonably considered it a member of the genus Stylorhyn-
chus from the character of the epimerite, which resembles that of a new
genus Bulbocephalus Watson (1916) of the family Stylocephalidae.

The species was not described again until the time of Tregouboff, who
contributes many new observations, assigning to it a new name, evidently
ignorant of the earlier work, and creating for it a new genus, Pyxinoides.

From the beautiful drawing of Kolliker, of a cephalont, there can be
no doubt that one and the same species is involved. He even shows in the
tiny figure the longitudinal grooves in the bulb of the epimerite.

Members of the genus Balanus also harbor Nematoides fusiformis,
very similar in some respects but yet regarded as a separate species. See
discussion under that species.

PYXINOIDES CTHAMALI (Leger and Duboscq) Tregouboff

[Figures 78 and 811
1909 Frenzelina chtamali Leger and Duboscq 1909a: 112, 114, 115

1911 Frenzelina cthamali Sokolow 1911: 281

1912 Pyxinoides cthamali Tregouboff 1912: Iviii
Sporonts regularly cylindrical, primite 230/4 in max. length, satellite

170m. Ratio LP : TL :: 1 : 4.5; WP : WD :: 1. 1. Protomerite wid-

36 ILLINOIS BIOLOGICAL MONOGRAPHS [36

est below mid-region, with characteristic specialized crescentic zone at
apex, with what appears to be a canal leading from apex a short distance
into protomerite. Deutomerite cylindrical, blunt at posterior end. No
constriction at septum. Nucleus ovoidal with one large karyosome.
Development extracellular. Epimerite atypical for that of the genus,
but still Pyxinia-like, a long style reaching in large trophozoites to base
of the cell and having a large crenulated cup-like enlargement at its mid-
point, the rim of the cup being turned away from the body of thegregarine.

Cysts spherical, 85 to lOOju in diameter. Spores unknown.

Intestine of Cthamalus stellatus Ranzani.

Taken at Cette, France.

In the first mention of the species, Leger and Duboscq 1909:112, the
spelling is given as chtamali and that of the host genus chtamalus, while
on pp. 114 and 115 the spelling is chthamali and chthamalus. Tregouboff
gives it for both parasite and host as cthamali (us).

Family CEPHALOIDOPHORIDAE Kamm 1922

Intestinal parasites of Crustacea, development intracellular, early
syzygies of two sporonts. Cysts without sporeducts, spores ovoidal with
equatorial line. Entire life cycle passed within a single host.

DISCUSSION OF THE NEW FAMILY CEPHALOIDOPHORIDAE

The genus Cephaloidophora was created in 1908 by Mawrodiadi for
intestinal parasites of the Crustacea. Leger and Duboscq in 1907 created
a genus in all respects identical with the former genus naming it Frenze-
lina; but in 1911 they discovered that the name Frenzelina was preoccu-
pied, so the name Cephaloidophora becomes the accepted genus designa-
tion.

The genus Frenzelina had been placed with the Gregarinidae because
of the precocious association of sporonts in pairs; but there the simi-
larities stop. Cephaloidophora was placed in the family Stenophoridae
(Sokolow 1911:286) and it must be acknowledged that there are many
characters in common with that family, including (a) intracellular devel-
opment, (b) rudimentary epimerite, (c) spores ovoidal with equatorial line,
(d) cysts without spore-ducts, (e) spores not united in chains. (A) is
confined to these two groups alone, in both of which there is cell-destruc-
tion, all other known forms possessing extracellular trophozoitic stages
without or with but little injury to the host. The only important variance
in the two groups lies in the fact that the Cephaloidophoridae invariably
form early syzygies of two individuals while the Stenophoridae as invari-
ably do not. The shape of the body, usual shape and character of the
nucleus and texture of the protoplasm are different in the two groups.
For complications given below, the gregarines of the Crustacea should be
considered apart from all others, at least until many of the knotty points

371 STUDIES ON GREGARINES—KAMM 37

are cleared up by exhaustive researches. For these reasons I have raised
the genus Cephaloidophora to a new family, Cephaloidophoridae.

Unlike other great groups of gregarines, the complete life-history of a
crustacean parasite must be known before it can be assigned to a particular
genus with a reasonable assurance of permanence; generally two out of
three or four characters will determine the location of a gregarine. Unless
the evolution of a sporozoite from the spore (Cephaloidophoridae) or from
the cyst direct (Aggregatidae) is known and the intracellular development
has been observed or proven to be absent, a crustacean parasite cannot
be accurately placed. Leger and Duboscq, the closest students of the
gregarines of the Crustacea, have removed many species hitherto classed as
gregarines to the family Aggregatidae, now placed in the order Coccidia,
because the sporozoites develop in the cyst without the intervention of
spores. The sporogonic cycle of the Aggregatidae is passed in another
host, a Cephalopod. Thus an alternation of hosts corresponds to an
alternation of generations, the crab being eaten by the Cephalopod.

Another family of gregarines inhabiting Crustacea is the POROSPORI-
DAE, parasitic in Decapods only. These animals are very large, usually
but not always solitary in the adult and are capable of forming cysts from
a single individual. This raises the question that such cysts are schizo-
gonic cysts and that the sporogonic stage, which possibly has not been seen,
is passed in another host.

Leger and Duboscq (1911 :lix) offer this hypothesis:

.... si Cephaloidophora et Porospora ne tomberont pas en synonymie, Porospora
representant la schizogonie et Cephaloidophora la gamogonie d'un meme cycle.

Tregouboff (1912) says, however:

Je suis convaincu que les Cephaloidophora effectuent leur cycle dans un seul h6te et presentent
un seul ty\)t des germes.

Minchin (1912:340) says:

A character such as the power of multiplication by schizogony is clearly one of great adaptive
importance in the life-history of a parasitic organism, and therefore not likely to be of class-
ificatory value. The classification of the future will probably be one which divides all grega-
rines into Cephalina and Acephaljna, and distributes the Schizogregarines (into which
suborder the Porosporidae are now placed) amongst these two divisions.

I think the last statement holds the clue to the classification of these
puzzling forms. Leger and Duboscq first placed two of their early forms
Cephaloidophora fossor and C. ocellata among the Aggregatidae, finding
only coelomic cysts. After more exhaustive researches they found a life-
history typically that of the genus Cephaloidophora, and removed the
species to this genus, considering that the coelomic cysts they had pre-
viously found belonged to different gregarines, of which nothing else was
known. I feel certain the latter really belonged in the same life-cycle
and that the power of producing coelomic cysts and their concommitant

38 ILUNOIS BIOLOGICAL MONOGRAPHS [38

sporozoites within the original host is one means the parasite has of
"holding its own" in Nature and reproducing at an enormous rate upon
certain occasions. This migration of the young gregarine through the
intestinal epithelium and production of a cyst from a single individual
resulting in schizogonic sporozoites may be due to a change of food in the
host animal unadapted to the parasite's requirements, to sickness of the
host, to a scarcity of food, to the time of the year, or to a natural phe-
nomenon occurring at stated intervals in the economy of the organism
itself. Or it may be due to some extraneous cause not mentioned.

I am convinced that all the gregarines in the Crustacea will again
come under one family head and that the Aggregatidae, the oldest known
group, will be restored to its position among the gregarines. The Poro-
sporidae and Cephaloidophoridae will of necessity be dropped. But in
the meantime, until much more work on the crustacean parasites shall
have been done and complete life-histories become the rule rather than the
exception, the present group headings must remain.

Type Genus CEPHALOIDOPHORA Mawrodiadi 1908: 101
Intestinal parasites of Crustacea. Characters of the family.

CEPHALOIDOPHORA CONFORMIS (Diesing) Leger and Duboscq

[Figures 66 and 75]

1787 Unnamed Cavolini 1787: 169

1810 ? Distome Rudolphi 1810: 287

1819 ? Distome Rudolphi 1819: 197

1851 Gregarina conformis Diesing 1851: 15

1859 Gregarina conformis Diesing 1859: 735

1863 Gregarina conformis Lankester 1863: 95

1885 Gregarina conformis Frenzel 1885: 579

1899 Aggregata conformis Labbe 1899: 6

1903 Aggregata conformis Minchin 1903: 331

1907 Frenzelina conformis Leger and Duboscq 1907: 773

1908 Frenzelina conformis Leger and Duboscq 1908: 99

1909 Frenzelina conformis Leger and Duboscq 1909: 733
1909 Frenzelina conformis Leger and Duboscq 1909a: 113
1911 Frenzelina conformis Sokolow 1911: 280
1911 Cephaloidophora conformis Leger and Duboscq 1911: lix
1913 Aggregata conformis Ellis 1913: 264

(Frenzel) Sporonts associated in pairs, not more than two, irregularly
cylindrical, tapering slightly, well-rounded posteriorly. Often a slight
shoulder exists. Protomerite broadly dome-shaped, slightly constricted

39] STUDIES ON GREGARINES—KAMM 39

at septum. 400 to SOOju in length. Ratio LP:TL::1:6 (primite)
1 :8 (satellite). WP:WD::1:5. Protomerite clear with large chro-
midial inclusions and crescentic apical area. Deutomerite dense, nucleus
not seen in vivo, spherical, one or more karyosomes.

Cysts ISO/i in diameter. Spores ellipsoidal with faint equatorial line,

6.4 by 5;u.

Intestine and gastric caeca of Pachygraspus marmoratus Fabr. {Cancer

depressus).

Taken at Cavaliere, France, Naples, etc.

This is recognized as the earliest known species of Gregarines. Redi
in 1708 found a parasite in the crab Cancer pagurus but Leger and Duboscq
(1908) say:

La courte description et les images que Redi (1708) donne des vers du Cancer pagurus ne peu-
vent s'appliquer aux Gregarines des Crustaces, dont la decouverte revient k Cavolini, quo-
iqu'en aient pens^ Diesing (1851) et Labbe (1899).

Cavolini in 1787 discovered in "Cancer depressi" a parasite which he
did not name and which Rudolphi (1810 and 1819) placed among his
"vermes generis dubii," suggesting that it may be a Distome.

Diesing (1851) named the parasite Gregarina conformis. In the same
paper he named another species seen by the same earlier workers Gre-
garina praemqrsa. Hence either species might with equal right be named
the type species. Diesing (1859) remarks only "Individua solitaria
(Cavolini)."

Frenzel studied the species at length and gives several figures. The
above description is taken mostly from his work.

Leger and Duboscq studied it again and find in their specimens that the
satellite is the longer, while the primite is short and relatively stouter.
The nucleus in their specimens is subspherical to ovoidal. Dimensions of
the two descriptions compare favorably, an association given by the later
workers as 1050ai in length, 400/i for the primite, the former being 80m
wide and the latter 40^. Ratios of drawing of the latter's work are as
follows: LP:TL::1 : 9 (primite) 1 : 21 (satellite); WP : WD :: 1
: 1.5 to 1 : 1.2. Thus their specimens conform to the type.

In 1907 these workers report schizogonic stages encountered in the
coelom. They also found sporogonic encystment in the intestine with
cysts 150m' in diameter found in the excrement and two sizes of spores,
6.4 by 5/i and microspores 5/i by 4.7jLt.

40

ILLINOIS BIOLOGICAL MONOGRAPHS

[40

CEPHALOIDOPHORA COMMUNIS Mawrodiadi Type Species

[Figure 77]

Solger

Labbe

Minchin

Mawrodiadi

Sokolow

Mercier

1890-1: 233
1899: 36
1903: 329
1908: 101
1911: 286
1912a: xli
1912: liii
and deutomerite rounded.
Intracellular trophozoites

1890-1 No name

1899 Gregarina sp.

1903 Gregarina sp.

1908 Cephaloidophora communis

1911 Cephaloidophora communis

1912 Cephaloidophora communis
1912 Cephaloidophora communis Tregouboff

Sporonts 65-80^1 in length. Protomerite
Associations in pairs, the primite the larger,
found in intestinal epithelium and hepatic cells. A thickened hyalin
crescentic area at apex of protomerite corresponding to a rudimentary
epimerite. Ratio LP : TL :: 1 : 4; WP : WD : 1 : 1.5. Cysts spheri-
cal. Spores with eight sporozoites, ovoidal, 4.5 by 4/i, with faint equator-
ial line. Sporozoites subspherical

Hosts: Balanus improvisus Darw.; B. improvisus gryphica Miinter;
B. tintinnahulum communis L.; B. perforans Brug.; B. amphitrite Darw.; B.
eburneus Gould.

Habitat: Intestine and caeca.

Taken at Odessa, Cette, France, etc.

The original description is in Russian and is not available. The above
data is taken from Sokolow's translation and from Tregouboff's new
observations upon live material.

Buddington (1910:470) gives a brief description of an unnamed species
found in Balanus eburneus Gould, presumably at Woods Hole, Mass.,
merely stating the animal to be a polycystid gregarine of rapid movement,
with a nucleus having five karyosomes and chromidial bodies in the
protomerite. It is quite probable that he found the above species.

CEPHALOIDOPHORA FOSSOR (Leger and Duboscq) TregoubofiF

{Figure 79]
Leger

Leger and Duboscq
Leger and Duboscq
Leger and Duboscq
Leger and Duboscq
Sokolow

1901
1903
1907
1908
1909
1911
1912
1912

Aggregata coelomica

Aggregata coelomica

Frenzelina fossor

Frenzelina fossor

Frenzelina fossor

Frenzelina fossor

Cephaloidophora fossor Tregouboff

Cephaloidophora fossor Mercier

ISOjj. in length. Ratio

LP:TL

1901
1903:
1907:
1908:

1909a
1911
1912
1912

:1 : 3

: 1343
: cxlvii
: 774

99
: 112,114,115
: 280
: Iv
: xliv
to 1 :4; WP;

Sporonts

WD:: 1:1.1. Sporonts obese, primite larger than satellite. Proto-
merite of primite broadly dome-shaped, twice as broad as high, with cres-

41] STUDIES ON GREGARJNES—KAMM 41

cent at anterior end and an apparent pore at the apex. Very slight con-
striction at the septum. Deutomerite ovoidal, broadly rounded at the
distal end. Nucleus apparently ovoidal, situated near septum. Satellite
more nearly cylindrical than primite.

Intracellular stage noted.

Intestine, coelom, and "sub-epithelium" of Pinnotheres pisum Penn.

In the first paper above, Leger reports finding coelomic schizogonic
cysts 150/i in diameter which produce sporozoites directly.

Leger and Duboscq (1907) conclude that the two types of cysts encoun-
tered respectively in the coelom and alimentary tract represent different
species of parasites, only one type of sporont having been seen, however.
They have found no other stages of the coelomic gregarine except the
cysts and contained sporozoites and have found all the stages in the life-
history of the intestinal parasite. I am convinced that but one species is
involved producing two kinds of cysts depending on conditions not yet
determined. I am not ready, however, to remove any of the species now
placed to another position. Much more work is needed upon crustacean
gregarines and parasites placed elsewhere before any species can be assigned
positions with a fair degree of permanence.

CEPHALOIDOPHORA OCELLATA (Leger and Duboscq) Kamm

[Figure 80

1903 Aggregata vagans Leger and Duboscq 1903: cxlvii

1907 Frenzelina ocellata Leger and Duboscq 1907: 774

1908 Frenzelina ocellata Leger and Duboscq 1908: 99

1909 Frenzelina ocellata Leger and Duboscq 1909a: 112,114,115
1911 Frenzelina ocellata Sokolow 1911: 280-1

1922 Cephaloidophora ocellata Kamm 1922 (this paper)

Sporonts in pairs, elongate cylindrical, 200/i in maximum length.
Primite the larger. Ratio LP :TL :: 1 :6 to 1 :9; WP : WD :: 1:
0.9. Protomerite widest anterior to the mid-region with two large cor-
puscles resembling eyes in both primite and satellite. A flattened cone
projects at the anterior end. Slight constriction at septum. Deutomerite
widest at shoulder, tapering from thence gradually and terminating in a
broadly rounded end. Nucleus ovoidal to rectangular, situated at or above
mid-region of deutomerite.

Intestine of Eupagurus Prideauxi Leach.

Taken at Banyuls, France.

This species was first described as Aggregata vagans and shown to exist
during the sporont stage in the intestine but to be able to pass while
comparatively large in size through the epithelium into the coelom. A
single individual here swells, its protoplasm changes in character, and it
finally becomes a schizogonic cyst 150/i in diameter in which are produced
sporozoites directly, as in the family Aggregatidae.

42 ILLINOIS BIOLOGICAL MONOGRAPHS [42

Typical sporont cysts are also noted in the excrement from the alimen-
tary tract.

The authors think that the two kinds of sporozoites represent different
species of parasites, the sporogonic representing the new species, Frenz-
lina occllata. I am of the opinion that but one species is involved. See
discussion under Cephaloidophora fossor and also Discussion of the New
Family Cephaloidophoridae, at the beginning of Crustacean Parasites.

CEPHALOIDOPHORA MACULATA Leger and Duboscq

[Figure 68]

1911 Cephaloidophora maculata Leger and Duboscq 1911 : lix

1912 Cephaloidophora maculata Tregouboff 1912 : liv
1912 Cephaloidophora maculata Mercier 1912a: xliii

Sporonts small,, ovoidal, maximum length 80/x. Ratio LP :TL :: 1
: 4 to 5; WP : WD :: 1 : 1.5 to 2. Cytoplasm yellowish in deutomerite,
clear in protomerite. Spherical chromidial bodies in protomerite. Nu-
cleus small, spherical, one karyosome. Characteristic crescentic apical
area in protomerite. Cysts spherical, lOO^i in diameter. Spores spherical
to subspherical, 4ju in diameter, with equatorial line. Intracellular stage
observed.

Intestine of Gammarus marinus Leach.

Taken at Roscoff, France.

CEPHALOIDOPHORA TALITRI Mercier
[Figure 86]
1912 Cephaloidophora talitri Mercier 1912:38

1912 Cephaloidophora talitri Mercier 1912a: xliv

1912 Cephaloidophora talitri Tregouboff 1912: Iv

Sporonts in pairs, primite the larger, ovoidal, average length 40^.
Ratio LP : TL :: trophozoites 1 : 4.5 to 6; WP : WD :: 1 : 1.2 to 2.
Protomerite dome-shaped with rudimentary epimeritic area at apex.
Slight constriction at septum. Deutomerite broadly ovoidal, nucleus
spherical with one large karyosome. Intracellular stage noted.

Cyst and spores unknown.

Intestine of Talitrus saltator Mont.

Taken at Roscoff, France.

CEPHALOIDOPHORA OLIVIA (Watson) Kamm

[Figure 83]
1912 Frenzelina olivia Watson 1916: 133

1922 Cephaloidophora olivia «Kamm 1922 (this paper)

Sporonts in pairs, max. length association 218m, ellipsoidal, largest
sporont 118 by 36^. Ratio LP:TL::1 : 5. WP : WD :: I : 1.3.

43] STUDIES ON GREGARINES— RAM M 43

Protomerite dome-shaped, constriction at septum slight, papillate area at
apex of protomerite orange in color. Deutomerite long-ovoidal. Nucleus
spherical, one karyosome. Cysts spherical, 60/u. Spores unknown.

Intestine of Libinia dubia.

Taken at Cold Spring Harbor, Long Island.

CEPHALOIDOPHORA NIGROFUSCA (Watson) Kamm
* [Figure 84]

1912 Frenzelina nigrofusca Watson 1916: 134

1922 Cephaloidophora nigrofusca Kamm 1922 (this paper)

Sporonts stout-bodied, ovoidal to rectangular, maximum size 125iLi by
75/x- LP :TL:: 1 :4; WP :WD ::1 :1.5. Protomerite broadly
dome-shaped with characteristic papillate apical area. Slight constric-
tion at septum. Deutomerite regularly cylindrical, broadly-rounded at
posterior end. Protoplasm very dense. Nucleus small, spherical with
one or two karyosomes.

Cysts and spores unknown.

Hosts: Uca pugnax, Uca pugilator. Intestine.

Taken at Cold Spring Harbor, Long Island.

CEPHALOIDOPHORA DELPHINIA (Watson) Kamm

[Figure 85]

1916 Frenzelina delphinia Watson 1916: 29

1922 Cephaloidophora delphinia Kamm 1922 (this paper)

Sporonts ovoidal, maximum length association 215^i, largest sporont
115 by 64m. Ratio LP :TL :: 1 : 4; WP : WD :: 1 : 1.5. Proto-
merite irregularly dome-shaped, dilated medianly. Papillated apex. Very
slight constriction at septum. Deutomerite ovoidal. Nucleus spherical,
one karyosome.

Cysts spherical, 80^. Spores unknown.

Intracellular stage noted.

Intestine of Talorchestia longicornis Say.

Taken at Cold Spring Harbor, Long Island.

CEPHALOIDOPHORA AMPELISCA (Nowlin and Smith) Kamm

[Figure 71]

1917 Frenzelina ampelisca Nowlin and Smith 1917: 83

1920 Cephaloidophora ampelisca Kamm 1922 (this paper)

Sporonts elongate cylindrical, 62/x by 15/1 in maximum dimensions.
Sporonts in pairs. Ratio LP :TL ::1 :6 to 1 :9; WP : WD :: 1 :1. Proto-
merite broadly dome-shaped, dilated in middle, more or less con-
stricted at septum and possessing a rudimentary epimerite or broad

44 ILLINOIS BIOLOGICAL MONOGRAPHS [44

apical papilla. Several chromidial bodies in protomerite. Deutomerite
cylindrical, blunt at posterior end. Nucleus spherical with one or more
karyosomes.

Intracellular development noted.

Intestine and hepatic caeca of Ampelisca spinipes.

Taken at Woods Hole, Mass.

Genera of Uncertain Position
NEMATOIDES Mingazzini 1891 2nd sem.: 233

Dicystid, no septum in sporonts. Epimerite forked, at apex of a long
neck. "Corpo allungato fusiforme, aguzzo ad entrambi gli apici. Cuticola
liscia."

NEMATOIDES FUSIFORMIS Mingazzini Type Species

1891 Nematoides fusiformis Mingazzini 1891: 2nd sem. 233

1899 Nematoides fusiformis Labbe 1899: 34

1903 Nematoides fusiformis Minchin 1903:203,329,331

Characters of the genus. "Trophozoite vermiform, without septum.
Epimerite in form of a fork or pair of pincers, borne on an elongated neck."
Minchin.

Intestine of Balanus perforatus, Pollicipes cornucopia.

Balanus sp. also harbor Cephaloidophora communis Mawr. and Pyxin-
oides halani (Kolliker) Tregouboff .

Mingazzini says the new species above is the Gregarina halani of
Kolliker. But the cephalont depicted by Kolliker corresponds in every
particular with that of an early cephalont of Pyxinoides halani. See dis-
cussion under that species.

If, however, the fully developed epimerites of the two species, Pyxin-
oides halani and Nematoides fusiformis are considered, they are suspiciously
similar. Labbe emends the original description of the latter thus:
Epim. en forme de fourche ou de pince,
separe par un col allonge du reste du corps.

The only character, then, in which the two species differ is this: The
latter possesses no septum in the sporont and is in appearance a Monocys-
tid, while the former has a complete septum in all stages except the very
earliest. Exhaustive researches may contribute something upon this
point and prove the two species identical.

Below is the original description of the species:

Cercando nell'intestino del Balanus perforatus la Gregarina balani der Kolliker che t una
policistidea, ho trovato invece una monocistidea appartenente a questo gruppo di gregarine
venniformi. £ pittosto lunga, ha la membrana intieramente liscia, un nucleo ovale al centre
con un nucleolo. Sembra assai rara. L'apice anteriore, troncato, termina con una specie di
ventosa, il posteriore invece e affatto puntuto. Nell'apice anteriore vi e un po'di metaplasma,
nel resto vi ba I'endoplasDQa.

45] STUDIES ON GREGARINES—KAMM 45

GANYMEDES Huxley 1910:55

A possible fixation organ at interior end, consisting of a motile stalked
sphere. Cup at posterior end which fits into ball of a satellite. No sep-
tum, no true epimerite. Typical gregarinoid encystment, cysts spherical.
Alimentary tract of Crustacea.

Huxley offers the hypothesis that this gregarine lies between the
polycystids and the monocystids and creates for it a new family, Gany-
medidae. Since his studies were made only upon fixed material and all
characters of this very unusual parasite are unknown, I think the rank of
a new family is hardly justifiable and prefer to consider it among the
Genera of Uncertain Position.

GANYMEDES ANASPIDIS Huxley Type Species ♦

[Figure 89]

1910 Ganymedes anaspidis Huxley 1910: 155

1913 Ganymedes anaspidis Ellis 1913: 264

Characters of the genus. Sporonts in pairs, elongate cylindrical,
maximum length 200/i, width 120/z. Average size 250^ to 300/x by IT/z
to 20/i. Nucleus large, ellipsoidal, one large karyosome. Cysts spherical,
100/x in diameter.

Anterior end of protomerite a "distinct stalked sphere," 8 to 10m across.
Protoplasm of anterior end highly specialized, posterior end a socket into
which fits the "ball" of another individual, during the associative period.

Intestine and pyloric caeca of Anaspides tasmaniae (Thompson).

Taken on the Island of Tasmania.

Species of Uncertain Position

[? gammari von Siebold]

1839 Gregarina gammari von Siebold 1839: ?

1848 Gregarina Gammari? Frantzius 1848: 196

1848 ? Gregarina longissima Koiliker 1848: 35

1859 Gregarina Gammari Diesing 1859:735

1863 Gregarina longissima Lankester 1863: 95

1886 Gregarina Gammari Plate 1886: 236

1895 Gregarina sp. Pfeiffer 1895:60

1899 Gregarina sp. Labbe 1899: 36

1903 Gregarina sp. Minchin 1903:330

Epimerite present. Sporonts 425/u in length, long-ovoidal.

Intestine and ? coelom of Gammarus pulex.

von Siebold described Didymophyes longissima from the same host,

considering it a second form of the same species. In the same paper,

V.

46 ILLINOIS BIOLOGICAL MONOGRAPHS [46

however, he described the above species from the same host, overlapping
his first observations with two names.

Diesing (1859) refers to this species thus:
Receptaculum globosum. Corpus ovoideum receptaculo duplo longius. Longit ....
crassit ....

He regards it as synonymous with the specimens seen by von Siebold and
described by Kolliker as follows:

Neben dieser Gregarine fand v. Siebold in Gammarus pulex noch eine andere Form (Fig. 29c),
von der es zweifelhaft bleibt, ob sie als Entwicklungsform zu der Gr. longissima zu rechnen
ist oder nicht.

KoUiker's figure obviously represents an entirely different species which
seems to correspond to one mentioned later in this discussion.

Lankester (1863) regards these as synonymous:

Gregarina longissima Sieb., Gr. diffluens Dies., Gr. miliaria Dies., Gr.
putanea Leuckart, Gr. Gammari Sieb.

Plate (1886) found in the intestine of Gammarus pulex two polycystid
gregarines. One is small, elongate cylindrical, 255 in length, the other
long-ovoidal, 425/i in length. The one seems to correspond with Didy-
mophyes longissima and the other with Gregarina gammari.

This species agrees in form and proportions with Cephaloidophora
maculata from Gammarus marinus but sporonts of the present species
attain 425/x in length while those of the former reach only 80jli.

Two species described by Diesing (1851:7) both from Gammarus pulex
are sufficiently alike to be considered synonymous. No detailed descrip-
tion of the species (? Gammari von Sieb.) is extant and so they cannot be
compared in minutae with the present species, but since all three are
found in the same host they should obviously be considered together under
the oldest name. These species are mentioned below

1851 Gregarina miliaria Diesing 1851: 7
1859 Gregarina miliaria Diesing 1859:731

Proboscis cylindrica gracilis, apice obtuse conica, uncinulorum seriebus 10-20. Receptaculum
subglobosum papillosum. Corpus ellipticum turgidum receptaculo tripio longius, laete
auranticum. Longit. 3^' ' ', crassit. . . .
Hosts: Gammarus pulex, Astacus fluviatilis. Intestine.

1851 Gregarina diffluens Diesing 1851: 7

1859 Gregarina diffluens Diesing 1859: 731

Proboscis cylindrica brevis gracilis, apice longe conica obtusa, uncinulorum seriebus 10-20.
Receptaculum subglobosum papillosum. Corpus oblongum per intervalla coarctatum postice
rotundatum, saturate auricatum, facillime difBuens. Longit. 1 ' ' '; crassit. . . .

Host: Gammarus pulex.

[? sp. Mawrodiadi]

1908 No name Mawrodiadi 1908: 101

1911 Gregarina sp. Sokolow 1911: 287

1912 No name Tregouboff 1912: Iviii

47] STUDIES ON GREGARINES—K AM M 47

Sporont 130-140/x in length, same general form as Pyxinoides halani
(Kolliker) Tr^gouboflf, but with a long epimerite which penetrates to the
muscular layer.

... la gr^garine poss^de un long 6pim6rite, qui traverse tout le tissu 6pith61ial, s'accolle i
la galne musculaire sous-jacente et la suce. (Quoted by Tr^goubofiF from Mawrodiadi.)

Tregoubofif adds:
D est impossible de juger mdme de la parents entre ces deux Gr^garines 6tant donne la d^fec-
tuosit6 de la description qui d'ailleurs tient toute dans les quelques mots cit6s plus haut.

Intestine of Balanus amphitrite pallidus Darw., B. amphitrite Darw., B.
eburneus Gould.

Taken in the Black Sea.

[? valettei Nussbaum]
1890 Gregarina valettei Nussbaum 1890: 156

1899 Gregarina valettei Labbe 1899: 36

1903 ''Gregarina'' valettei Minchin 1903: 331

1912 Gregarina valettei Tregouboff 1912: Ivii

Epimerite a simple stylet. Sporont S^n in length.

Intestine of Pollicipes polymerus Sow.

Taken in California.

Tregoubofif concludes
. . . cette Gregarine, . . . n'est certainement pas une Gregarina sensu stricto.

[? praemorsa Diesing]

1684 ?

1729 ?

1810 Distqme or Monostome?

1819 Distome or Monostome?

1851 Gregarina praemorsa

1859 Gregarina praemorsa

1863 Gregarina praemorsa

1899 Aggregata praemorsa

1903 Aggregata praemorsa

1908 Frenzelina praemorsa

1909 Frenzelina praemorsa
1911 Frenzelina praemorsa
1922 Cephaloidophora (?) praemorsaKannm 1922 (this paper)

Intestine and "ovarian appendage" of Cancer pagurus L. {Platycarcinus
(Lankester)).

The first reference in literature to what may possibly have been a
gregarine was that of Redi in 1684, who found in the "vesicular ovariorum"
of Cancer paguri a parasite which Rudolphi (1810, 1819) refers to as a
possible Distome or Monostome, and places in a list of "Vermes generis
dubii."

Redi

1684: 183

Redi

1729: 270

Rudolphi

1810: 287

Rudolphi

1819: 197

Diesing

1851: 287

Diesing

1859: 735

Lankester

1863: 95

Labbe

1899: 6

Minchin

1903: 329

Leger and

Duboscq

1908: 99

Leger and

Duboscq

1909a: 112

Sokolow

1911: 280

48 ILLINOIS BIOLOGICAL MONOGRAPHS [48

Diesing (1851) named the species Gregarina praemorsa from the
originally observed material and contributes no new observations. In
1859 he merely reports "Individua solitaria (Redi)."

As stated under Cephaloidophora conformis, Leger and Duboscq (1908)
do not credit Redi's observations as referring to authentic gregarines but
give to Calvolini (1787) the honor of having first seen and recorded un-
doubted specimens of this group. Rudolphi does not mention Cavolini
in connection with the present species, however.

Leger and Duboscq (1908 and 1909a) merely mention the species by
name in a list with this heading:

Le genre Frenzelina comprend ainsi actuellement les especes suivantes:

Sokolow lists it with the authentic species of the genus Frenzelina, not
with a group of uncertain species.

Thus there is no record of positive data concerning the actual existence
of this parasite as a gregarine. No description or drawing is extant.
It is remarkable that the mere reference has held its place in literature
for over two hundred years.

?

clausii

Frenzel

Figure 69

1879 ?

Claus

1879: 78

1885 Gregarina Clausii

Frenzel

«

1885: 575

1899 Gregarina clausi

Labbe

V

1899: 37

1903 "Gregarina" clausi

Minchin

1903: 331

Sporonts solitary, ovoidal

, 100m

in maximum

length.

Ratio LP :

TL::1:5.5; WP : WD :

:1:]

L.5.

Epimerite

unknown.

Protomerite

domeshaped, striated longitudinally. Deutomerite widest at shoulder,
tapering to a rounded point. No constriction at septum. Chromidial
bodies in protomerite. Nucleus spherical.

Solitary encystment within two thick hyalin cyst-walls. Cysts and
spores not described.

Intestine of Phronima sp., Phronimella sp.

Taken at Naples.

The position of this species cannot be determined from the known
data. Inclusion within the Cephaloidophoridae is doubtful from the
solitary encystment.

[? nicaeae Frenzel]

[Figure 70]

1885 Gregarina Nicaeae Frenzel 1885: 578

1899 Aggregata nicaeae Labbe 1899: 6

1903 Aggregata nicaeae Minchin 1903: 330

49] STUDIES ON GREGARINES—KAMM 49

Sporonts associated in pairs. Length 60^1. Ratio LP : TL :: 1 :3 to
4; WP:WD::1:L Protomerite dome-shaped, deutomerite tapering
to a blunt point. No constriction at septum. Nucleus large, spherical,
15/i in diameter.

Cyst and spores unknown.

Intestine of Hyale pontica Rathke (Nicaea Nilsonii).

Taken at Naples.

There is an equal possibility of this species belonging with the Cephaloi-
dophoridae and with the Aggregatidae from the characters recorded.
The exact position can only be determined after cyst and spores (or their
absence) are demonstrated.

[? sp. Minchin]

1903 Septate Gregarine Minchin 1903: 330

"Original observation."
Intestine of Nebalia serrata.
No comment whatever is offered concerning this parasite.

LIST OF SPECIES FOUND IN THE CLASS ACERATA

Parasite Host

actinocephalidae

Anthorhynchus sophiae (Schneider) Labb6 Phalangitis opilio

Type species

Sciadophora phalangii (L6ger) Labb6 Phdangium crassum

Type species P. cornutum

Opilio grossipes Herbst.

Sciadophora fissidens (Rossler) Labbe Phalangidae sp.

Sciadophora caudatus (Rossler) Kamm Phalangidae sp.

Sciadophora goronoivitschi (Johansen) Labb6 Phalangium opilio

ACANTHOSPORIDAE

Acanthospora repelini L6ger Phdangium cornutum

P. opilio
UNCERTAIN SPECIES

[? Wellmer] Oribata geniculata (L.)

Not named Sokolow] Scorpio indicus

Family ACTINOCEPHALIDAE Leger 1892: 166

Sporonts solitary, epimerites complex and varied. Cyst dehiscence by
simple rupture, spores irregular, biconical, or cylindro-biconical.

Genus ANTHORHYNCHUS Schneider 1887: 69, emend Labbe 1899: 19
Epimerite a large flattened and fluted button, spores ovoidal, knobbed
at sides, united in chains laterally.

Schneider

1887: 69

Leger

1897: 11

Labbe

1899: 19

Minchin

1903: 199, 338

Ellis

1913: 279

50 ILLINOIS BIOLOGICAL MONOGRAPHS [50

ANTHORHYNCHUS SOPHIAE (Schneider) Labb6 Type Species

[Figures 58, 108]

1887 Anthocephalus Sophiae

1897 Anthocephalus Sophiae

1899 Anthorhynchus Sophiae

1903 Anthorhynchus Sophiae

1913 Anthorhynchus Sophiae

Sporonts solitary, obese. Maximum length 2 mm. Width protomerite
330ai, deutomerite 600/x. Ratio LP : TL :: 1 : 11 (without epimerite);
WP:WD::1.2. Protomerite small, flattened, three times as wide as
high, no constriction at septum. Deutomerite conoidal, widest just
below septum, tapering thence and ending in a broad blunt extremity.
Epimerite a series of short blunt digitiform processes united laterally
and curved inward to form a broad flattened corona. Epimerite 200^ high.
Nucleus not described.

Spores 6.9 by 4.6/x, broad, biconical, extruded in chains laterally
attached.

Intestine of Phalangium opilio.

Taken at Poitiers, France.

Genus SCIADOPHORA Leger 1897: 36, emend Labbe 1899: 18

Epimerite a large flattened centrally indented papilla with a crenulate
periphery. Protomerite with numerous backwardly directed leaf-like
processes arranged vertically, each sharply pointed at its posterior extrem-
ity.

SCIADOPHORA PHALANGII (Leger) Labbe Type Species

[Figures 59, 60, 62, and 106]

1897 Lycosella Phalangii Leger 1897:12,36

1899 Sciadophora phalangii Labbe 1899: 18

1903 Sciadophora phalangii Minchin. 1903:199,338

1911 Sciadophora phalangii Wellmer 1911: 127

1913 Sciadophora phalangii Ellis 1913:280

Sporonts solitary, very elongate, up to 2.5 mm., one of the largest
known species. Width not given. Ratio LP:TL::1: 12; WP:WD::
1.3:1. Protomerite broadly conical at apex with 15 or 16 vertical
lamillar plates starting below the apical cone recurving backward and
terminating in sharp hooks. The whole bears a resemblance to an um-
brella. Deep constriction at septum. Deutomerite widest at shoulder,
tapering to a very long slender acuminate extremity. Epimerite a large

51] STUDIES ON GREGARJNES—KAMM 51

nearly sessile papilla indented in middle and crenulate on periphery.
Nucleus ovoidal, nearly spherical, with many karyosomes.

Cysts spherical, 3^ mm., dehiscence by simple rupture, spores biconical
but rounded off at poles. Unique. 9 x 5/x.

Intestine and caeca of Phalangium crassum, P. cornutum, Opilio grossi-
pes Herbst.

Taken in Provence, France and East Prussia.

Two species have been described previous to this one and considered

by Leger as synonymous; I have, however, not considered them so. They

are incompletely described but quite different from the type species and

because of the regularity of the digitiform bifurcate processes may not

belong in the genus. This is what Leger says concerning the species:

.... sans rien prejuger des especes observ6es par M. Johansen et M. Rossler, je donne k
celle que j'ai 6tudiee le nom specifique de Lycosella phalangii qui indique son origine. II ne
parait pas douteux cependant que, d'apres leurs caracteres morphologiques, les deux gr6-
garines signal^es par ces auteurs appartiennent au genre Lycosella et soient des expeces tres
voisines de celle que je d^cris dans ce travail, c'est ce que 1 etude complete de leur cycle
viendra nous confirme un jour on I'autre.

SCIADOPHORA FISSIDENS (Rossler) Labbe
[Figures 110 and 111]
1882 Actinocephalus fissidens Rossler 1882: 700

1897 Lycosella Phalangii Leger 1897: 12,36

1899 Sciadophor a fissidens Labbe 1899: 18

1903 Sciadophora fissidens Minchin 1903: 338

Sporonts solitary, 2 to 3 mm, in length. Ratio LP : TL :: 1 : 8;
WP : WD :: 1 : 1.5. The protomerite is broadly dome-shaped, at the
mid-region there is developed a corona of processes directed gently back-
ward in two rows, the upper consisting of twelve long sharp spines, the
lower of twelve broad plates widest at the middle and bifurcate at the
ends, superficially resembling a lobster's claw. These are arranged
alternately with the spines of the upper row. (Fig. 111). Deutomerite
widest at shoulder, tapering gently backward, the posterior third, however,
being a very much narrowed cylinder. Nucleus small, spherical.

Intestine and caeca of Phalangidae sp.

Taken at Freiburg, Germany.

This peculiar parasite in appearance is very unlike the type species,
(Cf. Figs. 62 and 110) but because the crenulations are a part of the
protomerite and not an epimerite, must be placed in this genus, or in a yet
undescribed genus very similar. The complete life-history remaining
unsolved, the species is placed here tentatively.

Rossler describes the peculiar protuberances thus:

.... zeigt am Kopf zwolf gespaltene Hakenpaare und zwischen je zweien dieser Paare einen
einfachen, stachelformigen Dorn.

52 ILLINOIS BIOLOGICAL MONOGRAPHS [52

SCIADOPHORA CAUDATUS (Rossler) Kamm
[Figures 112 and 113]

1882 Stylorhynchus caudatus Rossler 1882: 700

1899 Stylorhynchus caudatus Labbe 1899: 33

1903 Stylorhynchus caudatus Minchin 1903: 338

1913 Stylocephalus caudatus Ellis 1913: 338

1922 Sciadophora caudatus Kamm . 1922 (this paper)

Sporonts solitary, 2 to 2.5 mm. with a long "tail-process" 2 to 3 mm.
in addition. This makes the species the longest known gregarine. Body
similar in shape to S. fissidens except for the tail. The protomerite is
situated upon a short neck with a dome-shaped top and a corona of twelve
digitiform processes at the mid-region. Nucleus spherical.

Intestine and caeca of Phalangidae sp.

Taken at Freiburg, Germany.

This species and the preceding one are differentiated by the character of
the protomeritic corona. Cf. Figs. Ill and 113. It is evident that they
belong to the same genus, for the corona of each is a part of the proto-
merite rather than of an epimerite. Rossler considers the fact that this
species possesses a long neck upon which is superimposed the protomerite
sufl&cient evidence to place it in the genus Stylocephalus, but the coronae
are so similar and the host identical — the two found in the same intestines
— leads one to suspect that the two forms may represent parts of a single
life-history. Further investigations are, however, necessary to establish
this statement or refute it.

Rossler says concerning the species:

.... besitzt einen gestielten Kopf , der mit zwolf Erhebungen oder Leisten verstehen ist, die
liber den Rand desselben hinausragen und sich theilen. Diese Form ist ausserdem mit einem
diinnen, schwanzartigen Anhang versehen der durch keine Scheidewand von dejn eigentlichen
Korper getrennt ist, jedoch auch keine einspringenden Konturen zeigt, die auf einen ver-
stiimmelten Zustand schliessen liessen.

SCIADOPHORA GORONOWITSCHI (Johansen) Labbe
[Figures 104 and 105]

1894 Actinocephalus Goronowitschi Johansen 1894: 140

1897 Lycosella Phalangii Leger 1897: 11, 36

1899 Sciadophora goronowitschi Labbe 1899: 18

1903 Sciadophora goronowitschi Minchin 1903: 338

No dimensions or complete figure are given. Johansen describes the

peculiar parasite thus:

. . . naher zum Deutomerit gelegenen Aquator eine Reihe von Hackchen und Dornchen zur
Befestigung des Parasiten an dem Gewehen der Phalangide dient.

His meagre illustration. Fig. 105, indicates a bifurcation of the processes,
as in the species found by Rossler.

53] STUDIES ON GREGARINES—KAMM 53

Maximum length 5 mm. This length is reached only by the species
just preceding, the two being the longest known species of gregarines.
Intestine of Phalangium opilio.
Taken at Tomsk, Russia.

Leger also found Acanthrospora repelini in his material.

Family ACANTHOSPORID AE Leger 1892 : 167

Sporonts solitary. Epimerite simple or appendicular. Dehiscence by
simple rupture. Spores with equatorial and polar spines.

Genus ACANTHOSPORA Leger 1892: 167

Epimerite a simple large conical papilla on a short neck. Spores biconi-
cal or ovoidal with a row of equatorial spines and a tuft of four spines at
each pole.

ACANTHOSPORA REPELINI Leger

[Figures 57 and 107]

1897 Acanthospor a Repelini Leger 1897:13,42

1899 Acanthospor a repelini Labbe 1899: 28

1903 Acantkospora repelini Minchin 1903: 338

1911 Acantkospora repelini Wellmer 1911: 139

Sporonts solitary, obese. Maximum length 1 mm., width 250/i.
Ratio LP : TL :: 1 : 4 (without epimerite); WP : WD :: 1 : 1.2. Proto-
merite dome-shaped, deeply constricted at septum. Deutomerite widest
in anterior third, tapering irregularly to a sharp point. Epimerite a
broad flattened papilla indented in the middle and crenulate along the
margin. Nucleus spherical, one karyosome.

Cysts spherical, 500/z, dehiscence by simple rupture. Spores 13 by 4.8/i,
biconical, spines at apices and equator.

Intestine of Opilio grossipes Herbst, Phalangium cornutum, and P.
opilio.

Taken in Tourraine, France, and in East Prussia.

This species was found in association with Sciadophora phalangii by
Leger. There is no possibility of confusion, however, in the two species.

The epimerite of this species does not conform closely to that of the
type, ^./;r7e(z/a (Leger, 1892: 115).

Uncertain Species

[? Wellmer:

1911 Gregarina sp. Wellmer 1911:148

Intestine of Orihata geniculata (L.). Taken in East Prussia.
There are no data whatever concerning this gregarine form.

54 ILLINOIS BIOLOGICAL MONOGRAPHS [54

[? Sokolow]
[Figure 109]
1908 Not named Sokolow 1908:500

1911 Not named Sokolow 1911:295

SSfj, in length. Nucleus spherical, 3/x. Seen in copulation.
Host: Scorpio indicus.
No other data are given concerning this gregarine form.

SPECIES FOUND IN THE CLASS MALACOPODA
SPECIES OF DOUBTFUL POSITION

[? Moseley] Peripatus capensis Grube

[? Moseley]

1874 "Gregarinae" Moseley 1874: 762

1899 Gregarina sp. Labbe 1899: 37

1903 Gregarina sp. Minchin 1903: 331

The only data which exist concerning this parasite are the following:

Some very small encysted Gregarinae were found in the stomachs of all the specimens
examined.

Host: Peripatus capensis Grube.

Taken in Equatorial Africa.

One might wish that more data existed concerning the type of gregarine
which inhabit this intermediate host. Theoretically, it should be a form
intermediate between the family Polyrhabdinidae and the true Cephaline
Gregarines.

LIST OF SPECIES FOUND IN THE ORDER THYSANURA OF THE CLASS

HEXAPODA

Parasite Host

gregarinidae

Gregarina lagenoides (Leger) Labb6 Lepisma saccharina L.

SPECIES OF DOUBTFUL POSITION

? Gregarina podurae (L^ger) Labb6 Podura villosa

Gregarina sp. Wellmer Smiiithurus fuscus L.

Family GREGARINIDAE Labbe 1899: 9

Genus GREGARINA Dufour 1828: 366

Sporonts in pairs, epimerite a small sessile knob or cone. Spores barrel-
shaped to cylindrical. Cysts with spore ducts.

GREGARINA LAGENOIDES (Leger) Labbe

Figure 52

1892

Clepsidrina Lagenoides

Leger

1892:

118

1899

Gregarina lagenoides

Labbe

1899:

11

1903

Gregarina lagenoides

Minchin

1903:

335

1911

Gregarina lagenoides

Wellmer

1911:

117

55] STUDIES ON GREGARINES—KAMM 55

Sporonts in pairs, elongate. Length 150/i, width not given. Ratio
LP : TL :: 1 : 3.5. WP : WD :: 1 : 1 .3. Protomerite of primite cylin-
drical, with conical apex, the conical portion nearly equalling the cylindri-
cal in length. Slight constriction at septum. Deutomerite constricted
at end of anterior third, rapidly swelling to an almost perfect globe in
posterior two-thirds. Satellite of practically same shape but less accen-
tuated. Epimerite a simple spherical or ovoidal papilla.

Cysts spherical, dehiscing by spore-ducts, spores in chains. Spores
biconical, blunt at poles.

Intestine of Lepisma saccharina L.

Taken in the Valley of the Creuse, France, and in East Prussia.

Species of Doubtful Position

? GREGARINA PODURAE (Leger) Labbe

[Figures 53 and 54]

1892 Clepsidrina Podurae Leger 1892:119

1899 Gregarina podurae Labbe 1899: 12

1903 Gregarina podurae Minchin 1903: 336

Sporonts in pairs or threes. Maximum length 90/x. Protomerite either
separated from deutomerite by septum or lacking the septum. Ratio
LP : TL :: 1 : 5 . 5; WP : WD :: 1 : 1 . 8. In normal individuals proto-
merite dome-shaped, flattened apically. No constriction at septum.
Deutomerite cylindrical, broadly truncate at base.

In individuals which lack the septum the whole animal is ovoidal with
or without a very small papillate epimerite. Normal epimerite much
larger, a slightly stalked papilla. The individuals which form associations
of more than two are of the abnormal ovoidal type.

The protoplasm is gray-green with large orange granules. The nucleus
is spherical, with one karyosome.

Cysts spherical, 80/z, one long spore-duct, spores in chains, 6.8 by 3^-

Intestine of Podura villosa and Orchesella sp.

Taken in the Valley of the Loire, France.

This species seems to be an aberrent one representing a transitory
species to another genus or an intestine in which for some reason the
gregarines became abnormal. The possibility of more than one species
being present is excluded from the same type of protoplasm of peculiar
color and texture in both kinds of individuals. The author suggests that
the species may be intermediate between the genus Gamocystis, in which
a septum is always absent, and Gregarina in which it is always present.

56 ILLINOIS BIOLOGICAL MONOGRAPHS [56

Gregarina sp. Wellmer

1911 Gregarina sp. Wellmer 1911: 146

There are no data whatever concerning this form.
Intestine of Sminthurus fuscus L.
Taken in East Prussia.

LIST OF SPECIES FOUND IN THE ORDER ISOPTERA OF
THE CLASS HEXAPODA

SPECIES OF DOUBTFUL POSITION

? Gregarina termitis Leidy Termes flavipes; T, lucifugus Rossi.

Family GREGARINIDAE Labb^ 1899: 9
Genus GREGARINA Dufour 1828: 366
? GREGARINA TERMITIS Leidy
[Figures 51 and 56]

1881 Gregarina termitis Leidy 1881: 425

1897 Gregarina termitis Porter 1897: 65

1899 Gregarina termitis Labbe 1899: 36

1903 Gregarina termitis Crawley 1903: 44

1903 ''Gregarina" termitis Minchin 1903: 337

1913 Gregarina termitis Ellis 1913: 289

Sporonts solitary, obese ovoidal. Length 60/i, width 36/i. Ratio
LP : TL :: 1 : 3.3 WP : WD :: 1 : 1.2. Protomerite dome-shaped, twice as
wide as high, deep constriction at septum. Deutomerite ovoidal, tapering
to a rather sharp point. Nucleus spherical, one karyosome.

Epimerite and cysts unknown.

Intestine of Termes flavipes; T. lucifugus Rossi.

Taken at Philadelphia, Pa., Cambridge, Mass., and Boulder, Colo.

Leidy found but one gregarine in his host, but Porter reports the
parasites "very common in some hosts," in the anterior part of the small
intestine only. Porter illustrates a section of the intestine in which are
"great numbers of cysts." The sections, however, are undoubtedly those
of adult sporonts cut crosswise and obliquely, with dark-staining nuclei
often sectioned, as often not. A figure (Fig. 74, PI. VI, of Porter) showing
an adult solitary sporont in which the protoplasmic granules are fairly
large and regularly spherical is described as "that of one filled with sporo-
cysts." Porter's figure of a sporont is almost identical with that of Leidy
(Plate 52, fig. 27).

Crawley opened "perhaps a dozen termites" in search for the parasite
but did not encounter it.

I have examined many termites at difiFerent seasons. They were
abundantly parasitised with Infusoria (Leidy 1881, and Kofoid and Swezey
1919) but no gregarines were recovered.

57] STUDIES ON GREGARINES—KAMM 57

Ellis described the species from western termites. His drawing
(my figure 56) compares favorably with that of Leidy except that the
posterior end of his is broadly rounded instead of bluntly pointed. The
newer specimens are also much larger — 570/x long. Dimensions given by
Ellis are 'T. 25/x x 170m, D- 30m x 400m." They correspond with his figure
(PI. XVII, fig. 6) if changed to "P. 250m by 170m, D- 300m by 400m.

This species is considered doubtful because three important characters
are lacking, viz. epimerite, cyst and spores. None of the workers has
encountered associations.

LIST OF SPECIES FOUND IN THE ORDER HEMIPTERA OF
THE CLASS HEXAPODA

Parasite Host

ACnNOCEPHALIDAE

Coleorhynchus heros (Schneider) Labb6 Nepa cinerea L.

UNCERTAIN SPECIES

[? reduvii Ramdohrl Reduvius personalus (L.)

MISCELLANEOUS

[no name] Phymafa crassipes

Family ACTINOCEPHALIDAE Leger 1892: 166

Genus COLEORHYNCHUS Schneider 1885: 94, emend.

Labbe 1899: 23

Epimerite unknown. Protomerite a rounded shallow disc depressed
centrally with a cape overlapping the deutomerite. Septum convex up-
ward. Simple rupture of cysts, spores biconical.

COLEORHYNCHUS HEROS (Schneider) Labbe Type Species

[Figures 48 and 49]

1885 Cokophora heros Schneider 1885:95

1899 Coleorhynchus heros Labbe 1899: 23

1903 Coleorhynchus heros Minchin 1903:200,335

1913 Coleorhynchus heros Ellis 1913: 276

Sporonts solitary, length 2 to 3 mm. Width not given. Ratio LP :
TL :: 1 : 4; WP : WD :: 1 : 1. Protomerite a round flattened disc cen-
trally depressed, border slightly irregular with a cape extending down over
the deutomerite. Septum convex upward. Deutomerite ovoidal, blunt
at posterior end. Epimerite and nucleus not known.

Cyst dehiscence by simple rupture, spores sharply biconical.

Taken at Poitiers, France.

Intestine of Nepa cinerea L.

58 ILLINOIS BIOLOGICAL MONOGRAPHS [58

Uncertain Species

? reduvii Ramdohr

[Figure 50]

Ramdohr

1811:

194

Stein

1848:

213, 223

Frantzius

1848:

195

Diesing

1859:

734

Lankester

1863:

94

Labbe

1899:

14

Minchin

1903:

336

1811 Vibrio Reduvii
1848 Sporadina Reduvii
1848 Sporadina Reduvii
1859 Gregarina Reduvii
1863 Gregarina Reduvii
1899 Hyalospora reduvii
1903 Hyalospora reduvii

Taken at Berlin.

Intestine of Reduvius personatus (L.).

Ramdohr placed the species with the Infusoria, in the genus Vibrio.

Stein gives these dimensions:
Sie zeigten sehr verschiedene Grosse, die grossten waren etwa /^ ' ' ' lang und 1/38 ' ' ' breit

He also observed cysts in which two sporonts were united. Nine spore-
ducts were seen, with spore extrusion in chains. The spores were
. , kung spindelformig oder fast eiformig , 1/250'" by 1/570'".

His illustrations show them to be well-rounded at the ends.

The genus Hyalospora, in which the specimens recorded were placed
by Labbe, is characterized in part by (a) sporonts in associations of two,
(b) cysts with simple rupture, (c) spores fusiform, sharply pointed at the
ends.

The species in question does not belong in this genus because (a) among
the great numbers of parasites seen by Stein none is mentioned as found
in association and his figure indicates a solitary individual, (b) cyst dehis-
cence is accomplished through spore-ducts (nine in the figure of Stein),
and (c) with spores extruded in chains. The spores are ellipsoidal, rounded
at the ends.

It is evident that the species belongs in the family Gregarinidae from
the method of cyst dehiscence. I have placed it with the Uncertain
Species because of insufiicient data. It seems to most closely resemble
the genus Leidyana Watson.

Miscellaneous

No Name

1828

Gregarina Phymatae

crassipedis

Dufour

1828:

206

1837

Gregarina soror

Dufour

1837:

12

1846

Gregarina soror

Frantzius

1846:

11, 28

1851

Gregarina soror

Diesing

1851:

11

1863

Gregarina soror

Lankester

1863:

94

1903

Gregarina soror

Minchin

1903:

336

59]

STUDIES ON GREGARINES—KAMM

S^

Dufour says concerning this animal life:
Subsphericum alba, cephalothorace abdominis dimidiam partem adaequante.

From the figure and description of Dufour it is obvious that what he
observed were not sporonts but cysts. There is no record of sporonts being
found, and the error was a very natural one at the time.

LIST OF SPECIES FOUND IN THE ORDER NEUROPTERA OF
THE CLASS HEXAPODA

Parasite
gregarinidae

Gregarina clavata Kolliker

Gregarina mys'acidarum Frantzius

Gregarina marteli L€ger

Gamocys is ephemerae (Frantzius) Labb6

ACTINOCEPHAI.IDAE

Actinocephalus sieboldii (Kolliker) Frantzius
Actinocephalus oclacanthus Frantzius
Actinocephalus brachydactylus EUis
Geneiorhynchus monnicri Schneider

Type species
Genriorhynchus aeschnae Crawley
Bothriopsis claviforviis Pinto
Asterophora mucronata Leger

Tj'pe species
Asterophora elegans Leger

Discorhynchus truncalus (Leger) Labbe

Type species
Pileocephalus chinensis Schneider

Type species
Pileocephalus heerii (Kolliker) Schneider
Hoplorhynchus oligacantkus (von Siebold)
Schneider

Type species

ACANTHOSPORIDAE

Ancyrophora imcinata L^ger

Prismatospora evansi EUis
Type species

MENOSPORIDAE

Menospora polyacantha Leger

SPECIES OF DOUBTFUL POSITION

[? psocorum von Siebold]

Host

Ephemera vulgara larv.
Mystacides sp., larv.
Embia Solieri Ramb., larv.
Ephemera vulgata, larv.
Ephemera sp., larv.

Agrion sp., larv.
Phryganea sp.
Aeschna sp.
Libellules sp.

Aeschna constricta Say.
Aeschnida sp., larv.
Rhyacophila sp.

Phryganea grandis, larv.
Sericostoma sp., larv.
Sericostoma sp. larv.

Mystacides sp., larv.

Phryganea varia (Fab.) larv.
Calopteryx virgo L., larv.
C. splendens Harb. larv.

Sericostoma sp., larv.
Phryganea rhiimbica, larv.
Dytiscus sp., larv.
Colymbetes sp., larv.
Limnophilus rhombicus, larv.
Tramea lacerata Hagen
Sympetrum rubicundulum Say

Type species Agrion puella, larv.

Psocus sp.

60 ILLINOIS BIOLOGICAL MONOGRAPHS [60

Family GREGARINIDAE Labbe 1899: 9
Genus GREGARINA Dufour 1828: 366
GREGARINA CLAVATA Kolliker
[Figures 22 and 89]

1848 Gregarina clavata Kolliker 1848: 10

1848 Gregarina clavata Frantzius 1848: 194

1848 Sporadina clavata Frantzius 1848: 195

1851 Gregarina clavata Diesing 1851: 14

1859 Gregarina clavata Diesing 1859: 734

1863 Gregarina clavata Lankester 1863: 94

1863 Zygocystis clavata Lankester 1863: 94

1887 Ckpsidrina granulosa Schneider 1887:74,77,78

1899 Gregarina granulosa Labbe 1899: 11

1903 Gregarina granulosa Minchin 1903: 334

1911 Gregarina granulosa Wellmer 1911: 119

Of the original observation by Kolliker only a single immature specimen
is illustrated which furnishes a clue to the identity of the parasite. The
sporont is short and stout, 30 by 16/i, the ratio of LP : TL :: 1 : 2.5;
WP : WD :: 1 : 1. The protomerite is dome-shaped with a small apical
cone and a deep constriction at the septum. The deutomerite is ovoidal
and the nucleus spherical with one karyosome.

Intestine of Ephemera vulgata, larva.

Taken at Zurich, Poitiers, France, and in East Prussia

Kolliker records that Frantzius had found this species (1845) but
Frantzius' drawings (1848, PI. VII, figs. VII, 1, 2, 3, and 4) are very unlike
the figure of Kolliker (Fig. 89, this paper) and represent clearly the species
Gamocystis ephemerae.

Diesing (1859) records:

Receptaculum subhemisphaericum apiculo terminali conico. Corpus longe ovatum, lacteum.
Longit. 1/26 " ' ; crassit. 1/69 " ' .

Schneider (p. 74) reports finding Gamocystis ephemerae and also another
gregarine:

Le Gregarine qui domine qu'on trouve toujours dans ces larves, est la curieux Gamocystis
Francisci. Beaucoup plus rare et toujours en petit nombre est au contraire la Gregarine ac-
tuelle, qui se fait en revanche remarquer par une belle taille (518u le primite).

It is characterized by the very large granules in the endocyte of the
protomerite. Chromidial inclusions in the deutomerite. Ratio LP< TL
:: 1 : 5 (Primite) to 1 : 9 (Satellite). WP : WD :: 1 : 1.5. The nucleus
is spherical, 5\yi in diameter with one karyosome 26/i in diam. The
sporont is cylindrical, the protomerite dome-shaped with a deep constric-
tion at the septum. The deutomerite is cylindrical with a shoulder and
truncate at the free end.

Frantzius

1848: 193, 196

Stein

1848: 188

Diesing

1851: 12

Diesing

1859: 734

Lankester

1863: 94

Schneider

1875: 592

Labbe

1899: 11

Minchin

1903: 335

Wellmer

1911: 118

cal, Ratio

LP :TL ::1 :6 to IC

61] STUDIES ON GREGARINES—KAMM 61

While the observations of the two authors are not comparable, the
first from an immature specimen only, yet there is a probability of their
being identical, the hosts being the same. In order therefore to reduce
the number of older incompletely described species they should be absorbed
wherever possible. In the present species there are at least no conflicting
characters.

GREGARINA MYSTACIDARUM Frantzius

[Figure 76]
1848 Gregarina Mystacidarum
i848 Gregarina Mystacidarum
1851 Gregarina Mystacidarum
1859 Gregarina Mystacidarum
1863 Gregarina Mystacidarum
1875 Clepsidrina mystacidarum
1899 Gregarina mystacidarum

1903 Gregarina mystacidarum
1911 Gregarina mystacidarum

Sporonts in pairs, long-cylindrical,
WP : WD :: 1 : 1. Protomerite cylindrical, truncate at free end, slightly
constricted at septum. Deutomerite cylindrical, broadly rounded pos-
teriorly. Nucleus spherical. Epimerite not linown. Cysts spherical.

Taken at Berlin and in East Prussia. Host Mystacides sp., larva.

This host is also parasitized by Pileopcehalus chinensis Schn.

GREGARINA MARTELI Leger
[Figures 36 and 37]

1904 Gregarina Marteli Leger 1904: 358
1911 Gregarina Marteli Sokolow -1911:279

Sporonts in pairs, elongate, slender. Length 100 to 150/t. Width
not given. Ratio LP : TL :: 1 : 10; WP : WD :: 1 : 1. Protomerite of
primite conoidal, blunt at apex and dilated in posterior two-thirds. Pro-
tomerite, of satellite deeply cupped in anterior portion. Constriction at
septum. Deutomerite elongate-cylindrical, of same width throughout,
ending in a broad blunt extremity. Epimerite a simple globular papilla.
Nucleus spherical. Cyst and spores unknown.

Host: Embia Solieri Ramb., larv. Intestine.

Taken at Cavaliere, France.

This host is also parasitized with Diplocystis clerci Leger (Monocystid
Gregarine) and Adela transita Leger (Coccidian).

Genus GAMOCYSTIS Schneider 1875: 586; 1882: 443
Sporonts biassociative, united in apposition, ovoidal, stout-bodied,
resembling a Monocystid. Protomerite only in trophozoite. Sporulation
partial. Cyst without spore-ducts. Spores elongate-cylindrical.

62 ILLINOIS BIOLOGICAL MONOGRAPHS [62

GAMOCYSTIS EPHEMERAE (Frantzius) Labbe
[Figures 38 and 90]

1843

Zygocystis Ephemerae

Frantzius

1843:

194

1848

Gregarina clavata

Kolliker

1848:

10

1848

Zygocystis Ephemerae

Frantzius

1848:

194

1851

Gregarina clavata

Diesing

1851:

553

1859

Gregarina Ephemerae

Diesing

1859:

736

1882

Gamocystis Francisci

Schneider

1882:

444

1887

Gamocystis Francisci

Schneider

1887:

74

1899

Gamocystis ephemerae

Labbe

1899:

12

1903

Gamocystis ephemerae

Minchin

1903:

197, 334

Sporonts in pairs, in apposition. No protomerite. Body broadly
ovoidal, truncate at apices, flattened at place of union. Length of primite
110/i, of satellite 130/z. Width of former 80^, of latter same. Sarcocyte
thickened at the two apices. Endocyte dense, with large irregular gran-
ules. Nucleus spherical, 30/1, one karyosome. Cyst and spores not known.

Intestine of Ephemera sp., larva. Ephemera vulgata, larva.

Taken at Zurich and in Indre-et-Loire, France.

There seems to be no doubt that the species seen by Frantzius and by
Schneider are identical from a comparison of the figures given by each.
The ratio of length to width in the figures of Frantzius is about two to
one, in the figure of Schneider one and one-half to one.

Labbe considers Kolliker's Gregarina clavata identical with the above
species, probably because of a similarity of the host. The species are,
however, quite different.

Minchin considers the present species identical with Gamocystis fran-
cisci Schn.

Schneider (1882: 445) offers this phylogenetic hypothesis: The genus
Ephemera has been considered a pseudo-neuropteran closely allied to the
Orthoptera, in which group the only other species of the genus Gamocystis
has been found.

Family ACTINOCEPHALIDAE Leger 1892: 166
Genus ACTINOCEPHALUS Stein 1848: 196

Epimerite small, sessile or upon a short neck, with 8 to 10 short sharp
spines or simple bifurcate digitiform processes. Spores biconical.
ACTINOCEPHALUS SIEBOLDII (Kolliker) Frantzius

[Figure 12]

1839

?

von Siebold

1839:

67

1848

Gregarina Sieholdii

Kolliker

1848:

10

1848

Actinocephalus Sieholdii

Frantzius

1848:

195

1851

Gregarina Sieholdii

Diesing

1851:

7

1859

Gregarina Sieholdii

Diesing

1859:

731

1863

Gregarina oligacantha

Lankester

1863:

94

1899

Hoplorhynchus oligacanthus

Labbe

1899:

30

63] STUDIES ON GREGARINES—KAMM 63

Sporonts solitary, rather obese. Total length '0.162 to 0.188"" (KoUi-
ker), Width 0.045'". Ratio LP : TL :: 1 : 4 to 5; WP : WD :: 1.3 : 1.
Protomerite rounded, slightly wider than high, constriction at septum.
Deutomerite narrower than protomerite, tapering to a blunt posterior
extremity. Epimerite composed of 4 to 6 sharp recurved hooks set upon
a long cylindrical neck. Nucleus spherical with one or two large karyo-
somes.

Host: Larva of several species of Agrion. Intestine.

Taken at Danzig, Wurtzburg.

Kolliker says this species was first taken by von Siebold but not
described. Kolliker described it in detail and assigned to it a name.

Frantzius placed it in the genus Actinocephalus because of the charac-
ter of the epimerite.

Labbe placed the species with Eoplorhynchus oligacanthus Schn. because
of a close similarity with this genus. While the epimerites of the two
species are similar, Kolliker's species having 4 to 6 recurved hooks and
Schneider's 6 to 8, the hooks on the former species are sharply pointed and
decidedly recurved while those on the latter number eight as a maximum
and are but slightly recurved and blunt at the tips. (Cf. Figs. 13 and 14
{Eoplorhynchus oligacanthiis) with Fig. 12, the present species).

The writer has differentiated the two species on this point and a
dissimilarity of the nuclei. Schneider's new genus, Hoplorhynchus, is
characterized by an elongate-ovoidal nucleus while the nucleus in Kolliker's
species is spherical.

ACTINOCEPHALUS OCTACANTHUS Frantzius

[Figure 74]

194, 195

1848

Gregarina Heerii

Frantzius

1848:

193

1848

Actinocephalus octacanthus

Frantzius

1848:

192

1851

Gregarina Heerii

Diesing

1851:

552

1851

Gregarina Frantziusiana

Diesing

1851:

8

1863

Gregarina Heerii

Lankester

1863:

94

Host: Phryganea sp. Intestine.

Frantzius recognized that this species was different from the Gregarina
Heerii of Kolliker from the same host, and illustrates epimerites of the two
species, but his drawings are confusing for in another place he calls the
same drawing now one name, now the other.

Diesing regarded the two species as synonymous on one page (552) and
still separates them on another, creating a new species for the present
species, Gregarina frantziusiana. He differentiates the two thus: *

64 ILLINOIS BIOLOGICAL MONOGRAPHS [64

Gregarina frantziusiana Diesing :

Proboscis globosa, uncinulorum, octo coronula tenninali simplici (octolobia Frantzius).
Corpus subellipticum turgidum ....

Gregarina heerii Kolliker.

Proboscis antrorsum obovato-lanceolata, granulis exasperata, retrorsum subelliptica laevis,
inermis? .... Corpus longe ovatum ....

Hab. Phryganea grandis larva, intestine, in both.

From then on the present species is lost in the species Pileocephalus
heerii. It is obvious from a comparison of the epimerites that two very-
different species have been confused and that the original designation of
Frantzius still holds. ^

ACTINOCEPHALUS BRACHYDACTYLUS Ellis
[Figures 44 and 45]

1913 AcHnocephalus brackydactylus Ellis 1913: 279, 289

Sporonts solitary, elongate-ovoidal. Length 501/i, Ratio LP: :1 :3 to 4;
WP:WD: :1:1. Protomerite well-rounded, with same width as height,
slight constriction at septum. Deutomerite slightly wider at shoulder,
tapering gently and ending in a blunt point. Epimerite very short, a
corona of eight short digitiform processes.

Cyst and spores unknown.

Habitat not given. Host: Nymphs of Aeschna sp.

Taken at Douglas Lake, Michigan.

Genus GENEIORHYNCHUS Schneider 1875: 594

Epimerite a tuft of short bristles set at the apex of a long slender neck.
Spores cylindro-biconical.

GENRIORHYNCHUS MONNIERI Schneider Type species

[Figures 19, 20, and 21]

1875 Geneiorhynchus Monnieri Schneider 1875: 595

1899 Geneiorhynchus monnieri Labbe 1899: 25

1903 Geneiorhynchus monnieri Minchin 1903: 200, 335

1913 Geneiorhynchus monnieri Ellis 1913: 274

Sporonts solitary, obese, ovoidal. No dimensions given. Ratio
LP:TL: :1:4; WP:WD: :1:1.3. Protomerite very broadly rounded,
acuminate at apex. Slight constriction at septum. Deutomerite widest
at mid-portion, ending in an acute point. Epimerite a tuft of short sharp
bristles set at apex of a long slender neck. Nucleus spherical with several
karyosomes. Cyst dehiscence by simple rupture, spores sub-navicular.

Intestine of Libellules sp.

Taken near Paris.

65J STUDIES ON GREGARINES—KAMM 65

GENEIORHYNCHUS AESCHNAE Crawley
[Figure 40]
1907 Geneiorhynchus aeshnae Crawley 1907: 227

1911 Geneiorhynchus aeshnae Sokolow 1911: 282

1913 Generorhynchus aeshnae Ellis 1913: 275, 289

Sporonts solitary, cylindrical. Length of trophozoite 420jU. Ratio
(without epimerite) LP:TL: :1:3. WP:WD: :1.1:1 Protomerite widest
at base, curving gently anteriorly, ending in a truncate cone. No con-
striction at septum. Deutomerite narrower than protomerite at septum,
tapering thence and ending in a broad truncated extremity, Epimerite a
large globular papilla borne on a short stout neck from the protomerite and
"liberally provided with short spines directed backward." (Crawley).
Nucleus spherical with several karyosomes. Sarcocyte in apex of pro-
tomerite very thick. Endocyte not dense, nucleus visible in vivo. Cyst
and spores not known.

Intestine? of Aeschna coustricta Say. {Aeschna c.)

Taken in Southeastern Pennsylvania.

Genus BOTHRIOPSIS Schneider 1875: 596
Epimerite an ovoidal structure with six or more long slender filaments.
Protomerite very large, septum convex upward. Spores biconical.

BOTHRIOPSIS CLAVIFORMIS Pinto
[Figure 61]

1918 Bothriopsis claviformis Pinto 1918: ?

1919 Bothriopsis claviformis Pinto 1919: 87
Sporonts qlongate-triangular, widest at anterior end, bluntly acuminate

posteriorly. Dimensions: 100 to 200At in length, 70 in width (of protomer-
ite). Ratio LP:TL: :1:7; WP:WD: :1.4 : 1. Nucleus spherical to irregular
in shape.

Intestine of Aeschnida sp.

Taken at Manguinhos, Brazil.

Genus ASTEROPHORA Leger 1892: 129
Epimerite a thick long disc with a milled border and a long stout style
projecting upward from the center. Spores cylindro-conical.

ASTEROPHORA MUCRONATA Leger Type species
[Figures 25 and 26]
1892 Asterophora mucronata Leger 1892: 130

1899 Asterophora mucronata Labbe 1899: 22

1903 Asterophora mucronata Minchin 1903: 200, 336

Sporonts solitary, elongate-ovoidal. Length 350m, width not given.
Ratio LP:TL: :1:2.2. WP:WD: :1:1.1. Protomerite elongate (twice as

66 ILLINOIS BIOLOGICAL MONOGRAPHS [66

long as wide), tapering upward from septum to a blunt conoidal anterior
end. Deep constriction at septum. Deutomerite of same shape as proto-
merite and but little longer. Epimerite a flat disc with milled border and
long blunt central style set upon a fairly stout neck. Nucleus ovoidal with
several karyosomes. Cysts spherical, 150-200/i. Dehiscence by simple
rupture. Spores cylindro-biconical, 8 by 3.5^.

Host: Larva of Rhyacophila sp. Intestine.

Taken in the Valley of the Loire, France.

ASTEROPHORA ELEGANS Leger

[Figures 23, 24, and 43]

1892 Asteropkora elegans Leger 1892: 131

1899 Asteropkora elegans Labbe 1899: 22

1903 Asteropkora elegans Minchin 1903: 200,336

Sporonts solitary, elongate ovoidal. Length 450^- Width not given.
Ratio LP:TL: :1:3; WP:WD: :1:1.1. Protomerite very elongate (almost
twice as long as wide), tapering regularly upward from septum to a long
blunt anterior end. Deep constriction at septum. Deutomerite of same
shape as protomerite but twice as long. Epimerite a flat disc with a milled
border and a short central style set upon a fairly stout neck which is of the
same length as the protomerite. Nucleus slightly ovoidal with several
karyosomes.

Cysts spherical, 300 to 400/i. Spores biconical, much larger than in
Asteropkora mucronata, 12.7 by 4.25^4.

Intestine of Pkryganea grandis, larva, Sericostoma sp., larva.

The former host is also parasitized with Pileocephalus heerii, the two
parasites generally found together. Sericostoma is also parasitized with
Discorkynckus truncatus.

Genus DISCORHYNCHUS Leger 1892: 134; Labbe 1899: 20

Epimerite a short-stalked, large flattened globular structure with a still
larger horizontal collar around its base. Spores biconical.

DISCORHYNCHUS TRUNCATUS (Leger) Labbe Type species

[Figures 28 and 29]

1892 Discorkynckus truncatus Leger 1892: 134

1899 Discorkynckus truncatus Labbe 1899: 20

1903 Discorkynckus truncatus Minchin 1903: 199, 337

1913 Discorkynckus truncatus Ellis 1913:275

67] STUDIES ON GREGARINES—KAMM 67

Sporonts solitary, almost rectangular. Length 300/i, width not given.
Ratio LP:TL: :1:4; WP:WD: :1.2 : 1. Protomerite irregularly rounded,
bluntly conoidal at apex, widest at mid-portion. Constriction at septum.
Protomerite wider than deutomerite. Deutomerite rectangular, in optical
section, flat at base. Epimerite a short stout neck with a large flattened
papilla at apex. Around the base of the papilla fits a much larger flattened
disc like a collar. Nucleus spherical, with several karyosomes.

Cysts spherical, 140/li. Spores biconical, slightly crescentic.

Taken in Poitou, France.

Intestine of a neuropteran doubtfully identified as the larva of Seri-
costoma sp.

Genus PILEOCEPHALUS Schneider 1875:591

Epimerite a small lance-shaped or simple conoidal papilla placed un-
stalked upon the protomerite. Spores biconical.

PILEOCEPHALUS CHINENSIS Schneider Type species
[Figures 17, 18, 93, and 94]

1875 Pileocephalus chinensis Schneider 1875: 592

1885 Pileocephalus chinensis Schneider 1885: 10

1899 Pileocephalus chinensis Labb6 1899: 19

1903 Pileocephalus chinensis Minchin 1903: 199, 335

1911 Pileocephalus chinensis Wellmer 1911: 128

1913 Pileocephalus chinensis Ellis 1913: 274

Sporonts solitary, stout bodied, ovoidal. Measurements not given;
"... tou jours de taille assez petite ..." Ratio LP:TL: :1:5; WP:
WD: :1:1. Protomerite broadly rounded, apex pointed, slightly wider
than high. Constriction at septum. Deutomerite widest at shoulder,
tapering from thence and ending in a broad blunt extremity. Epimerite
a small unstalked papilla, conoidal at apex. Nucleus spherical.

Cysts spherical, spores roughly triangular with long slender ends.
(Figs. 93, 94.)

Taken near Paris.

Intestine of Mystacides sp., larva.

This species received its name from the character of the epimerite
concerning which Schneider says :
Epimerite triangulaire ou cordifonne en maniere de bonnet chinois ....

The insect is also the host of Gregarina mystacidarum Frantzius, and
Schneider and Wellmer found associations of the parasite in the host
with the above described species.

68 ILLINOIS BIOLOGICAL MONOGRAPHS [68

PILEOCEPHALUS HEERII (Kolliker) Schneider
[Figures 15, 16, 46, 72 and 73]

1845

Gregarina Heerii

KoUiker

1845:

100

1848

Gregarina Heerii

Kolliker

1848:

6

1848

Gregarina Heerii

Frantzius

1848:

193

1848

Stylorhynchus Heerii

Frantzius

1848:

195

1851

Gregarina Heerii

Diesing

1851:

552

1851

Gregarina Frantziusiana

Diesing

1851:

8

1859

Gregarina Heerii

Diesing

1859:

728

1859

Gregarina Frantziusiana

Diesing

1859:

730

1863

Stylorhynchus Heerii

Carus

1863:

570

1863

Gregarina Heerii

Lankester

1863:

95

1887

Pileocephalus Heerii

Schneider

1887:

199

1899

Pileocephalus heerii

Labbe

1899:

19

1903

Pileocephalus heerii

Minchin

1903:

199, 336

Sporonts solitary, capable of great contractility due to unusually well-
developed myonemes. Trophozoite when quiescent long and arrow-shaped,
widest in middle and tapering in both directions, sharply acuminate at
extremities (Fig. 15). In sporonts the same general shape but much
contracted longitudinally and proportionally broader. (Fig. 16). Di-
mensions not given. Ratio LP:TL: : (quiet trophozoite without epimerite)
1:3 (contracted sporont 1:2). WP:WD: :1:1. Protomerite widest just
above septum, conical, apex truncate. Deep constriction at septum.
Deutomerite same shape, ending acuminately. Nucleus ellipsoidal
with many small karyosomes. ' Epimerite highly specialized. In young
trophozoites an elongate papilla sharply acuminate and situated upon a
short neck. In order specimens there devlops a spade-shaped or lanceolate
(in optical section) holdfast, at the end of a bulbous neck (the former
papilla). The neck and lance are of equal length and surmount the
truncate protomerite. The trophozoite in this form is a very beautiful
animal (Fig. 15). *

Spores from cysts in feces biconical, which is the accepted form for the
type, as mentioned in the genus synopsis.

A possible schizogony exists, for in this species cysts were encountered
in the coelom which developed triangular spores. Schneider attributes
these latter spores to the species in question but he suggests that they may
belong to another parasite. If the discovery is authentic, the fact adds
one link in the hypothesis of Minchin that the Cephaline Gregarines are
capable of schizogony as well as sporogony. If this should be proven an
impossibility, then the present species must be removed from the Eugre-
garinae and placed with the Schizogregarinae.

Schneider found only triangular spores in the type species P. chinensis,
but does not state whether they emanated from coelomic or fecal cysts,
mentioning only the latter type of cysts.

69] STUDIES ON GREGARINES—KAMM €9

Habitat: Intestine of Phryganea varia (Fabricius), larva.

Taken at Wurzburg, and Poitiers (France).

Kolliker gives as hosts Phryganea grandis 'and other Phryganeidae
larvae.' The only point of serious conflict between the two sets of obser-
vations lies in the shape of the nucleus. We are led to believe this one of
the fixed characters of a species, but Kolliker finds the nucleus to be
spherical, while Schneider says it is ovoidal. 'In P. chinensis, however, it
is spherical, so this is not a genus character.

Schneider does not doubt the identity of the two species.

La figure donnee par Kolliker ne laisse aucun doute sur la synonymic, ....

Frantzius gives a beautiful figure of a trophozoite which agrees in every
particular with the observations of others except that the nucleus here too
is spherical (Fig. 72, this paper). I am inclined to think Schneider's obser-
vations were in error or mis-interpreted in the rapidly moving animals and
that the correct shape in all three sets of data is spherical.

Genus HOPLORHYNCHUS Carus 1863:570

Sporonts slender with elongate ovoidal nuclei. Epimerite a very long
neck with six to eight slightly recurved hooks. Cyst dehiscence by simple
rupture. Spores ellipsoidal.

HOPLORHYNCHUS OLIGACANTHUS (von Siebold) Schneider

Type species
[Figures 13 and 14]
1839 Gregarina oligacanthus
1845 Gregarina oligacantha
1845 Gregarina oligacantha
1848 Gregarina oligacantha
1848 Stylorhynchus oligacanthus
1848 Stylorhynchus oligacanthus
1851 Gregarina oligacantha
1859 Gregarina oligacantha
1863 Gregarina oligacantha
1875 Hoplorhynchus oligacanthus
1899 Hoplorhynchus oligacanthus
1903 Hoplorhynchus oligacanthus
1911 Hoplorhynchus oligacanthus
1913 Hoplorhynchus oligacanthus

Sporonts solitary, slender. Dimensions
1:4; WP:WD: :1:1. Protomerite irregularly rounded, slighlty wider than
deutomerite. Widest at mid-region, constricted at septum. Deutomerite
irregularly cylindrical, tapering posteriorly and ending in a blunt point.
Epimerite a corona of six to eight slightly recurved hooks set upon a long

von Siebold

1839:

67

Dujardin
Kolliker

1845:
1845:

638

?

KoUiker

1848:

10

Stein

1848:

195,

222

Frantzius

1848:

195

Diesing
Diesing
Lankester

1851:
1859:
1863:

7

730-

95

1

Schneider

1875:

591

Labbe

1899:

30

Minchin

1903:

201,

Z?,3

Wellmer

1911:

139

Ellis

1913:

275

not given.

Ratio

LP:TL:

70 ILLINOIS BIOLOGICAL MONOGRAPHS [70

slender cylindrical neck. Nucleus elongate-cylindrical with many small
karyosomes.

Cyst and spores as in genus characters.

Intestine of Cahpteryx {Callopteryx, Schneider) virgo L., larva, and
C. splendes Harb. larva.

Taken at Danzig, Berlin and Paris.

With all the authors who mention the species, only one gives adequate
figures, Schneider, from whom the data and drawings is taken.

Diesing (1859) reports:

Proboscis longissima gracilis medio ventricosa, apice subglobosa, spinularum (circa 9) corona
simplici. Receptaculum subglobosum compressum. Corpus conicum lacteum receptaculo
quinquies longius.

Lankester considers Actinocephalus sieholdii and the present species
synonymous.

Family ACANTHOSPORIDAE, Leger 1892: 167
Sporonts solitary, epimerite simple or appendicular. Dehiscence by
simple rupture. Spores with equatorial and polar spines.

Genus ANCYROPHORA Leger 1892: 146
Epimerite a globe with a corona of not more than 12 backwardly-
directed blunt hooks. Spores biconical with equatorial and polar spines.

ANCYROPHORA UNCINATA Leger
[Figures 27, 41 and 42]

1848 Gregarina Dytiscorum Frantzius 1848: 194

1892 Ancyrophora uncinata Leger 1892: 147

1899 Ancyrophora uncinata Labbe 1899: 28

1903 Ancyrophora uncinata Minchin 1903: 201,335

1916 Ancyrophora uncinata Watson 1916a: 164

Sporonts solitary, elongate. Length 150-200/x. Protomerite well
rounded, deeply constricted at septum. Deutomerite widest at shoulder,
tapering from thence to a long slender acuminate posterior extremity.
Epimerite a corona of twelve or less short blunt recurved hooks, superim-
posed upon a distinct but rather short stout neck. The hooks are arranged
in two levels, the upper group being larger and stouter. Nucleus spherical
with several karyosomes.

Cysts spherical, 200;u, spores biconical, hexagonal in optical section,
with four spines at each pole and six on equator. Spores 8 by 6/Le.

Intestine of larvae of Sericostoma sp., Phryganea rhumbica, and of two
beetles, Dytiscus sp. and Colymbetes sp. Minchin adds Limnophilus
rhombicus, probably a synonym of the second.

Taken in France.

711 STUDIES ON GREGARINES—KAMM 71

The fact that this species is found in such widely varying hosts is
unusual but not unique in the history of gregarines.

Genus PRISMATOSPORA EUis 1914: 215

Spores hexagonal, truncate at ends with one row of long spines at each
pole. Epimerite subglobose with lateral recurved hooks.

PRISMATOSPORA EVANSI Ellis Type species
[Figures 95, 96, 97, and 99]

1914 Prismatospora evansi Ellis 1914: 215

Sporonts broadly conical 400/i in average length. Ratio LP:TL: :1:3;
WP: WD: :1:1. Protomerite broad, blunt, deutomerite tapering. Nucleus
small, spherical.

Cysts subspherical, 370/1 in diameter. Dehiscence by simple rupture.
Spores as in genus, six long spines encircling each pole a short distance
from the end, 11 by 5.8/i.

Intestine of Tramea lacerata Hagen. and Sympetrum ruhicundulum
Say.

Taken at Douglas Lake, Michigan.

The sporonts of this species are peculiar in that the protomerite is drawn
out into digitiform processes at the apex in vivo, which may aid the
animal in retaining its hold upon the intestine in so active a host where ab-
dominal contractions might thrust it from the canal. These processes disap-
pear when the animal is placed in water.

Family MENOSPORIDAE Leger 1892:168

Sporonts solitary. Epimerite a large cup bordered with hooks and
placed on a long slender collar. Cyst dehiscence by simple rupture.
Spores crescentic, smooth.

Genus MENOSPORA Leger 1892:151, 168
Characters of the family.

MENOSPORA POLYACANTHA Leger Type species
[Figures 30, 31 and 32]

1892 Menospora polyacantha Leger 1892: 151

1899 Menospora polyacantha Labbe 1899: 30

1903 Menospora polyacantha Minchin 1903: 201, 332

1913 Menospora polyacantha Ellis 1913: 275

Sporonts ovoidal, lanceolate, 600-700/i in length. Width not given.
Ratio LP:TL: : (trophozoite, without epimerite) 1:5; WP:WD: :1:1.1.
Protomerite well rounded, deeply constricted at septum. Deutomerite
widest just below septum, tapering gradually to a long sharply pointed

•72 ILLINOIS BIOLOGICAL MONOGRAPHS [72

extremity. Epimerite a deep urn bordered with many short recurved
hooks, superimposed upon a long slender neck two-thirds as long as the
whole animal. Nucleus elongate-ovoidal, with one dumb-bell shaped
karyosome.

Cysts spherical, 200^*, spores smooth crescents 15 by 4n.

Taken in Poitou, France.

Intestine of Agrion puella, larva.

Species of Uncertain Position

[? psocorum von Siebold]
1839 Gregarina Psocorum von Siebold 1839: 67

1846 Gregarina Psocorum Frantzius 1846: 25

1848 Gregarina Psocorum Frantzius 1848: 195

1851 Gregarina ovata Diesing 1851: 10

1863 Gregarina Psocorum Lankester 1863: 94

1911 Hyalospora psocorum Wellmer 1911: 124

The only data outside the original reference, which is not available,
are those of Diesing (1851):

Proboscis .... Receptaculum compressum. Corpus ovalum utrinque obtusum, recepta-
culo triple longius. Longit. crassit . . .

Host: Psocus sp.

LIST OF SPECIES FOUND IN THE ORDER LEPIDOPTERA OF
THE CLASS HEXAPODA

Pakasite Host

gkegarinedae

Leidyana tinei Keilin Endrosis feneslrella Stain., larv.

UNNAMED SPECIES Keilin Oecophora pseudopretella Stain., larv.

Tinea pallescenldla Stain., larv.

Family GREGARINIDAE Labbe 1899: 9
Genus LEIDYANA Watson 1915: 35

Sporonts solitary, epimerite a small sessile knob, dehiscence by
spore-ducts. Spores in chains, barrel-shaped.

LEIDYANA TINEI Keilin
[Figures 115 and 116]

1918 Leidyana tinei Keilin 1918: 406

Sporonts solitary, long-ellipsoidal. Ratio LP:TL: :1: 7.5; WP:WD: :
1 : 1.7. Maximum length 300/x, width 85^. Protomerite hemispherical,
deutomerite regularly ellipsoidal, tapering gently to a blunt posterior
end.

Cysts spherical 110/i in diameter. Spores barrel-shaled, 7^ long.

Intestine of Endrosis fenestrella Stain.

Taken at Cambridge, England.

73]

STUDIES ON GREGARJNES—KAMM

73

Unnamed Species Keilin

1918 Unnamed species Keilin 1918: 406

Intestine of Oecophora pseudospretella Stain, and Tinea pallescentella
Stain.

Taken at Cambridge, England.

"Very similar to species here described," above, but a complete life-
cycle was not procured, and the specimens left unnamed.

LIST OF SPECIES FOUND IN THE ORDER DIPTERA OF THE CLASS

HEXAPODA
Parasite Host

GREGASINIDAE

Gregarina longa (L6ger) Labb6
Gregarina ctenocephalus Ross
Hirmocystis veniricosa (L^ger) Labb6

Type species
Eirmocystis polytnorpha (L^ger) Labb6

ACTINOCEPHALIDAE

Actinocephalus tipulae (Hammerschmidt)

L€ger
Actinocephalus sp. L^ger
Schneideria caudata (von Siebold) Ldger
Schneideria mucronata L^ger Type species
Schneideria sp.

Slylocystis praecox L^ger Type species
Taeniocystis mira L^ger Type species
Pikocephaliis striatus L^ger and Duboscq

Tipula sp., larva
Ctenocephalus serraticeps
Tipula oleracea, larv.

Limnobia sp., larv
Syslenocerus caraboides L., larv.

Tipuia oleracea L., larv.

Pachyrhina pratensis L., larv.

Ctenophora sp., larv.

Sciara nitidicollis Meig. larv.

Bibio marci (L.), larv,

Chironomus sp., larv.

T any pus sp., larv.

Ceratopogon solstitialis Winn., larv.

Ptychoptcra contaminala, larv.

Family GREGARINIDAE Labbe 1899: 9
Genus GREGARINA Dufour 1828
GREGARINA LONGA (Leger) Labbe
[Figure 7]
1892 Clepsidrina longa Leger 1892: 117

1899 Gregarina longa Labbe 1899: 11

1903 Gregarina longa Minchin 1903: 337

1911 Gregarina longa ^ Wellmer 1911: 114

Sporonts in pairs, very slender and elongate. Maximum length of
Bporont 500m. Width not given. Ratio LP:TL: :1: 10 to 12; WP:WD: :
1:1.2. Another form is found in which this ratio exists: LP:TL: :1:7,
body less slender. Protomerite of primite well-rounded, deeply con-
stricted at septum. Protomerite of satellite flattened at top, constricted
in middle. Deutomerite regularly cylindrical, of approximately same
width throughout in primite and tapering very slightly in satellite, ending
in a broad blunt extremity. Epimerite a simple spherical papilla. Nucleus
large, spherical, with one large karyosome.

/

74 ILLINOIS BIOLOGICAL MONOGRAPHS [74

Cysts spherical, 140|u. Dehiscence by 7 or 8 short spore-ducts. Spores
extruded in chains, dolioform, 6 by 5/i.

Intestine of Tipula sp., larva.

Taken in the Valleys of the Vienne and Loire, France and in East
Prussia.

GREGARINA CTENOCEPHALUS Ross

1909 Gregarina ctenocephalus canis Ross 1909: 359

1922 Gregarina ctenocephalus Kamm 1922 (this paper)

Sporonts spherical, no dimensions given. Epimerite pyriform, spores
barrel-shaped. No dimensions given.

Intestin^of Ctenocephalus serraticeps.

Taken at Port Said, Egypt.

Genus HIRMOCYSTIS Leger 1892: 110, emend. Labbe 1899: 12
Associations of from two to twelve sporonts. Epimerite small, cylin-
drical papilla. Cyst dehiscence by simple rupture. Spores ovoidal.

HIRMOCYSTIS VENTRICOSA (Leger) Labbe Type species

[Figures 8 and 9]

1892 Eirmocystis ventricosa Leger 1892: 111

1899 Eirmocystis ventricosa Labbe 1899: 13

1900 Eirmocystis ventricosa Magalhaes 1900: 39
1903 Eirmocystis ventricosa Minchin 1903: 336, 337
1911 Eirmocystis ventricosa Wellmer 1911: 123
1913 Eirmocystis ventricosa Ellis 1913: 264

Sporonts bi- or rarely tri- associative, obese. Length 180ai, width not
g^iven. Ratio LP:TL: :1:3.5. WP:WD: :1:1.5 Protomerite broadly
rounded in front, widest in the anterior half, much constricted and cylin-
drical in second half. No constriction at septum. Deutomerite broadens
appreciably just below septum, widest in anterior third, tapering from
thence, ending in a square extremity. Epimerite a small cylindro-conical
papilla. Nucleus spherical, one karyosome.

Cysts spherical, lOOju, dehiscence by simple rupture, spores ovoidal,
not fusiform, 9n by 6/x.

Intestine of Tipula oleracea and Pachyrhina pratensis, larvae.

Taken in the Valley of the Loire, France and in East Prussia.

Tipula sp. is also parasitized by Gregarina longa and Actinocephalus
tipulae.

HIRMOCYSTIS POLYMORPHA (Leger) Labbe
[Figures 10 and 11]
1892 Eirmocystis polymorpha Leger 1892: 113

1899 Eirmocystis polymorpha Labbe 1899: 13

1903 Eirmocystis polymorpha Minchin 1903: 197,335

1911 Eirmocystis polymorpha Wellmer 1911: 123

751 STUDIES ON GRECARINES—KAMM 75

Sporonts in associations of from two to twelve, arranged linearly or
bi- or tri-furcate, largest individuals always at anterior end of chain.
Length sporonts 100 to 150/i, whole series attaining 13^ mm. Ratio LP:
TL: :1:6 to 8; WP:WD: :1:1. First protomerite rounded at apex, wider
than high, constriction at septum. Deutomerite elongate cylindrical,
truncate at extremity. Epimerite a simple cylindrical knob. Nucleus
spherical, one karyosome.

Cysts spherical, lOO/x. Dehiscence by rupture, spores ovoidal, 11 by S/ci.

Intestine of Limnobia sp., larva; Systenocerus carahoides L. larva.

Taken in Poitou, France and in East Prussia.

Family ACTINOCEPHALIDAE Leger 1892: 166

Genus ACTINOCEPHALUS Stein 1848: 196

ACTINOCEPHALUS TIPULAE (Hammerschmidt) Leger

[Figure 6]

1838 BulluUna Tipulae Hammerschmidt 1838: 357

1846 Gregarina Tipulae

1851 Gregarina Tipulae

1859 Gregarina Tipulae

1863 Gregarina Tipulae

1892 Actinocephalus tipulae

1899 Actinocephalus tipulae

1899 Actinocephalus tipulae

1903 Actinocephalus tipulae

1911 Actinocephalus tipulae

Sporonts solitary, obese. Length nearly a millimeter. Width not given
Ratio LP:TL: :1:3.5 to 5; WP:WD: :1:1. Protomerite sub-spherical,
widest in middle, deeply constricted at septum. Deutomerite widest at
shoulder and tapers from thence to a rather short sharply pointed ex-
tremity. Epimerite not seen when alive and penetrating a cell, hence its
supposed digitiform appendages not delineated. Portion seen a large
round apically pointed papilla superimposed upon a short thick neck.
Nucleus large, spherical. Several karyosomes.

Cysts spherical, 250 to 300^ spores fusiform, 9 by 4^.

Intestine of Tipula oleracea L., larva, Pachyrhina pratensis, larva.

Taken in the Valley of the Vienne, France, and in East Prussia.

Diesing credits Hammerschmidt with having first seen and named
this species. Since no data whatever exists concerning the species until
Leger's time, it is doubtful that the two species are identical, especially
since Tipula is parasitized by several gregarines, but the old record stands.
Diesing gives as host Ctenophora pectinicomis , larva, in body cavity.

Frantzius

1846:

28

Diesing

1851:

16

Diesing

1859:

739

Lankester

1863:

94

Leger

1892:

141

Labbe

1899:

26

Leger

1899:

532

Minchin

1903:

337

Wellmer

1911:

130

76 ILLINOIS BIOLOGICAL MONOGRAPJIS [76

Leger places the species positively in the genus stated:

Ces caract^res, (the epimerite as described above) joints k ceux fournis par la forme du spora-
din du kyste et des spores, font renter, sans aucun doute, cette Gr6garine dans le genre
Actinocephalus.

ACTINOCEPHALUS sp. Leger

1899 Actinocephalus sp. Leger 1899: 532

1903 7 Actinocephalus sp. Minchin 1903: 333

Host: Ctenophora sp., larva.

This species is not described, but merely mentioned in a list of other
species found by Leger.

Genus SCHNEIDERIA Leger 1892: 153

Sporonts solitary. Protomerite separated from deutomerite by
constriction but no septum (pseudomonocystid). Epimerite a flat disc
with milled border, with or without central spine. Cyst dehiscence by
simple rupture. Spores biconical, smooth. (It might be possible that a
septum exists in the young trophozoites although this is not suggested
by Leger.)

SCHNEIDERIA CAUDATA (von Siebold) Leger
[Figures 4 and 5]

1839

Gregarina caudata

von Siebold

1839:

51

1845

Gregarina caudata

Dujardin

1845:

638

1846

Gregarina caudata

Frantzius

1846:

23

1848

Actinocephalus caudatus

Frantzius

1848:

195

1851

Gregarina caudata

Diesing

1851:

8

1859

Gregarina caudata

Diesing

1859:

729

1863

Gregarina caudata

Lankester

1863:

94

1875

Gregarina caudata

Schneider

1875:

591

1892

Schneideria coronata

Leger

1892:

155

1899

Schneideria caudata

Labbe

1899:

22

1903

Schneideria caudata

Minchin

1903:

337, 199

Sporonts solitary, elongate. Maximum length 1 mm. Ratio LP:TL: :
1:9; WP:WD: :1.1 : 1. Protomerite flattened at apex, wider than high,
widest in middle, constricted considerably to meet deutomerite but no
septum present. Deutomerite long, slender, tapering from anterior end,
a very long conical extremity. Epimerite a flattened disc with a milled
border, no style in center, as in 5. mucronata. Nucleus slightly ellipsoidal,
one large karyosome.

Cysts not seen. Spores 20 by 10/x.

Intestinal caeca of Sciara nitidicoUis Meig., larva.

Taken in Poitou, France.

77] STUDIES ON GREGARINES—KAMM 77

This parasite is one-fifth the length of its host, and because of the
transparency of the host can easily be seen from without when both are
alive.

No clue to the actual identity of this parasite exists in the earlier work
except this by Diesing (1851):

Proboscis brevissime cylindrica apice subglobosa truncata crenata, uncinis retractus?
which seems to apply to the epimerite. It seems strange, if this species is
meant, tliat no one mentions the absence of a septum until Leger described
anew the species with a new name. He does not refer to the earlier workers.

Labbe, however, unites the species under the older name. The hosts
are identical.

SCHNEIDERIA MUCRONATA Leger Type species
[Figures 1, 2 and 3]

1892 Schneideria mucronata Leger 1892: 153

1899 Schneideria mucronata Labbe 1899: 22

1903 Schneideria mucronata Minchin 1903: 199, 333

Sporonts solitary, elongate. Length 700-800/x. Width not given.
Ratio LP:TL: :1:7; WP:WD: :1:1. Protomerite irregularly triangular,
pointed, widest two-thirds its length from the apex. Deep constriction
between protomerite and deutomerite. No septum. Deutomerite slender,
gradually tapering from anterior end to a long slender point. Epimerite
a wide flattened disc with a milled border and a short style in the center.
Nucleus large, spherical, one karyosome. Very agile in movement, also
polymorphic.

Cysts 270 by 190/i, fusiform spores 15 by 9/x.

Intestinal caeca of Bibio marci (L.), larva.

Taken in Tourraine and Poitou, France.

SCHNEIDERIA sp. Leger
1899 Schneideria sp. Leger 1899: 526, 532

1903 Schneideria sp. Minchin 1903:333

Very briefly described as unilocular, less than 250/i in length, greatly
dilated in the anterior portion and gradually tapering posteriorly. Epi-
merite not seen. Judging by context from similar parasites in other
fly larvae, the author considers it a Polycystid rather than a Monocystid,
which it resembles.

Host: Chironomus sp., larva.

Taken in Poitou, France

Genus STYLOCYSTIS Leger 1899: 529
Dicystid gregarine with a simple epimerite in form of a sharply pointed
recurved spine. Spores biconical.

78 ILLINOIS BIOLOGICAL MONOGRAPHS [78

STYLOCYSTIS PRAECOX Leger Type species

[Figure 33]

1899 Stylocystis praecox Leger 1899: 529

1903 Stylocystis praecox Minchin 1903: 199, 337

1911 Stylocystis praecox Sokolow 1911: 283

Sporonts solitary, elongate, maximum length 500/x, width not given.
Ratio LP:TL: :1:10. WP:WD: :1:1. Body cylindrical, somewhat dilated
in anterior end. Tapering slightly at posterior end, end blunt. Nucleus
spherical with one karyosome. Epimerite a slender recurved pointed
spine.

Cysts elongate-ovoidal, 200/i in long axis. Spores biconical, 8 by 5/x.

Intestine of Tanypus sp., larva.

Taken at Grenoble, France.

Genus TAENIOCYSTIS Leger 1906: 307

Deutomerite of sporont divided by septa into numerous segments one
behind the other. Epimerite a small sphere set with six or eight recurved
hooks. Spores biconical.

TAENIOCYSTIS MIRA Leger Type species
[Figures 34 and 35]

1905' Leger 1905: ?

1906 Taeniocystis mira Leger 1906: 307

1908 Taeniocystis mira Leger and Duboscq 1908a: 105
1911 Taeniocystis mira Sokolow 1911: 284
1913 Taeniocystis mira Ellis 1913: 280

Sporonts solitary, elongate cylindrical, deutomerite segmented, super-
ficially resembling a tapeworm. Length 400-500/x. Width not given.
Ratio LP:TL: :1:11. WP:WD: :1:1. Epimerite a relatively very small
bulb with a corona of 6 to 8 recurved hooks set upon a short neck.

Cysts spherical, 130/Lt, spores 7 by 3/:.

Host: Ceratopogon solstitialis Winn., larva.

Taken at Cavaliere, France.

Genus PILEOCEPHALUS Schneider 1875: 591
Epimerite a small lance-shaped or simple conoidal papilla placed
unstalked upon the protomerite. Spores biconical.

PILEOCEPHALUS STRIATUS Leger and Duboscq

[Figure 100]

1909 Pileocephalus striatus Leger and Duboscq 1909b: 887
Sporonts solitary, length 150/i. Protomerite more or less hemispherical,

deutomerite broad, obtusely pointed. Epimerite a long striated bulb
upon a short stout neck. Nucleus large, spherical, often found in the

79] STUDIES ON GREGARINES—K AM M 79

protomerite (a phenomenon which has been nowhere else except in the
species Pileocephalus chinensis by Schneider in 1886 and in Acanthospora
polymorpha by Leger in 1897).

Cysts spherical.

Development extracellular, no cell destruction.

Intestine of Ptychoptera contaminata, larva.

Taken in Belgium?

LIST OF SPECIES FOUND IN THE ORDER SIPHONAPTERA OF
THE CLASS HEXAPODA
Parasite Host

actinocephalidae

Actinocephalus parvus Wellmer Ceratopsyllus JringiUae (Wlk.)

C. gallinae (Schrank), larvae
Steinina rotunda Ashworth and Rettie Ceratopsyllus styx Roths.

GENTJS OF UNCERTAIN POSITION

Agrippina bona Strick. Ceratopkyllus fasciatus Bosc. larv.

FamUy ACTINOCEPHALIDAE Leger 1892: 166 .
Genus ACTINOCEPHALUS Stein 1848: 196
ACTINOCEPHALUS PARVUS Wellmer
[Figures 114 and 120]

1911 Actinocephalus parvus Wellmer 1911:131
Sporonts ovoidal, maximum dimensions 140ju by 75m- Ratio LP:TL: :

1:5; WP:WD: :1:1.3. Nucleus ovoidal, several karyosomes. Epimerite
a corona of eight digitiform processes situated upon a short neck.
Cyst and spores unknown.

Intestine of Ceratopsyllus JringiUae (Wlk.), larva {Ceratopkyllus /.);
and C. gallinae (Schrank), larva.
Taken at Konigsberg.

Genus STEININA Leger and Duboscq 1904: 352
Epimerite a short mobile digitiform process changing into a flat
button. Spores biconical.

STEININA ROTUND ATA Ashworth and Rettie
[Figures 121, 122, 123 and 124]

1912 Steinina rotundata Ashworth and Rettie 1912: 31
Sporonts solitary, obese to pyriform. No constriction at septum.

Maximum dimensions 180/1 by 80^. Nucleus spherical, one or two large
karyosomes.

Cysts spherical, 110/i to 185^ in diameter. Dehiscence by simple
rupture. Spores ovoidal, \2n by 7/i. Extracellular development. Intes-
tine of Ceratopsyllus styx Roths. (Ceratopkyllus s.)

Taken near Edinburgh.

80 ILLINOIS BIOLOGICAL MONOGRAPHS [80

• Genus of Uncertain Position

AGRIPPINA Strickland 1912: 108

Sporonts solitary, epimerite a circular disc armed with digitiform
processes on periphery, a short neck. Spores ellipsoidal.

AGRIPPINA BONA Strickland 1912: 108 Type species
[Figures 101, 102, and 103]

1912 Agrippina bona Strickland 1912:108

Sporonts elongate conoidal, average length 175At. Nucleus ellipsoidal
Protomerite dome-shaped, deutomerite tapering to a long pointed extremity
from the shoulder. Epimerite as in genus.

Cysts spherical, dehiscing by rupture. Spores smooth, ellipsoidal,
6.6 by 7/Li.

Intestine of Ceratophyllus fasciatus Bosc, larva.

Taken at Cambridge, England.

LIST OF SPECIES FOUND IN THE SUBPHYLUM TUNICATA

Parasite
[? Kolliker]
[? Ritter]
[? Frenzel]
[? Roboz]
[? Bargoni]

Host

Phallusiae mamillaris Ecker

Perophora annectens

Salpa maxima

Salpa confoederata,

S. vagina

Salpa aeruginosa

[PKoUiker]

Figure 55

Kolliker

1848: 35

Frantzius

1848: 195

Diesing

1851: 9

Diesing

1859: 728

Lankester

1863: 95

Labbe

1899: 49

Minchin

1903: 339

1848 Gregarina phallusiae

1848 Stylorhynchus Phallusiae

1851 Gregarina Phallusiae

1859 Gregarina Phallusiae

1863 Gregarina Phallusiae

1899 Pleurozyga phallusiae

1903 Pleurozyga phallusiae

The figure given by Kolliker indicates the presence of a septum. Animal j

solitary, length 30 to 40/i. Ratio LP:TL: :1:4.5. WP:WD: :1:1. Proto- j

merite dome-shaped, constricted at septum. Deutomerite cylindrical, '

tapering slightly and ending in a broad, rounded extremity. Nucleus ,

spherical. Epimerite a short bulbous style.
Cyst and spores unknown.

Taken at Triest. i

Intestine of Phallusiae mamillaris Ecker. '

The position of the gregarines in Tunicates is still uncertain. Little |

work has been done on them for thirty years and they offer an interesting '

81] STUDIES ON GREGARINES—KAMM 81

field from a phylogenetic point of view — both as regards the hosts and the
parasites themselves. The Tiinicates represent the highest type of hosts
recorded for gregarines. Whether or not the gregarines are also the most
complex is a problem of the future.

[? Ritter]
[Figure 98]

1893 Gregarina sp. Ritter 1893: 69
1899 Gregarina sp. Labbe 1899: 37
1903 Gregarina sp. Minchin 1903: 339
1913 Gregarina sp. Ellis 1913: 271, 290

40/x in diameter. The parasites were found in the digestive tract,
boring through the walls, and in the coelom. Three bands characterize
all but the smallest, made up of "many elongated granules possibly
cuticular in structure though they seem to take the stain."

Host: Perophora annedens.

The specimens described have little the character of polycystid gre-
garines (Fig. 98). There is a superficial resemblance to the genus Taenio-
cystis, but they possess no protomerite, and in one instance two nuclei are
present, which leads to the hypothesis that they may belong to the genus
Haplozoon.

[? Frenzel]
[Figures 39 and 117]

1885 Gregarina Salpae Frenzel 1885: 565
1899 Gregarina salpae Labbe 1899: 37
1903 "Gregarina" salpae Minchin 1903:339

Intestine of Salpa maxima.

[? Roboz]

1886 Gregarina flava Roboz 1886: 1, 46
1899 Gregarina flava Labbe 1899: 37
1903 "Gregarina'' flava Minchin 1903: 339

Intestine of Salpa confoederata, S. vagina.

[? Bargoni]

1894 Gregarina ensij or mis Bargoni 1894: 44
1899 Gregarina ensiformis Labbe 1899: 37
1903 "Gregarina" ensiformis Minchin 1903: 339

Intestine of Salpa aeruginosa.

The following data is given, by Bargoni, concerning this species:

Sospettai tratarsi di qualche parassita e particolarmente di una Gregarina avendone gii altra,
di forma per6 semilunare, ma molto simile pel contenuto giallognolo, avuta in esame dal Prof.
Kleinenberg, che la trov6 frequentemente parassita nella Cydippe e nella Alciopa. Di questa
nuova forma, che non va confusa colla Gregarina flava del Roboz da me potuta osservare una
volta sola in Salpa aeruginosa, mi occuper6 altra volta, accontentandomi pel momento di
distingueria col nome di Gregarina ensiformis n. sp.

82

ILLINOIS BIOLOGICAL MONOGRAPHS

[82

LIST OF HOSTS WITH THEIR GREGARINE PARASITES

Host

Platyhelminthes
Geoplana backi
G. amagensis
Planaria sp.
Polyporus sulphttreus
Sorocoelis sp.

Annelida: Polychaeta
Capitella capitala
Clymenella torquata
Glycera siphonostotna

Ophelia neglecta
Pareudrilus pallidus

Polydora ciliala
Polydora socialis

Pygospionis selicornis
Rhinodrilus incertus
Scolekpsis fuliginosa
Spio martinensis
Annelida: Oligochaeta
Kynotus Pitlarellii
Lumbricus ierrestris

Lumbricus variegatus
Glossoscolex wiengreeni

Annelida: Hirudin ea

Glossophonia complanala
Hemiclepsis marginata

Rotifera

Euchlanis dilalala
Salpina mucronata

Echinodermata

Echinocardiiim cordalum

it n

sp.
Spatangus sp.
Synapta purpureus
Synapta galliennei
Synapta digitala
Mollusca

Cerithiutn vulgalum

Parasite

Rhynochocyslis geoplanae Fuhrman
Rhynchocystis geoplanae Fuhrman
Lankesleria sp. Swarczewskj-
Gregarine form, Wellmer
Lankesleria sp. Swarczewsky

Ancora lutzi Hasselmann
Microlaeniella clymenellae Calkins
Gonospora glycerae Pixell-Goodrich
Gonospora inleslinalis Pixell-Goodrich
Three unnamed parasites Pixell-Goodrich
Rhytidocyslis henneguyi deBeauchamp
Monocystis pareudrili Cognetti de Martiis
Rhynchocystis hessei Cognetti de Martiis
Polyrhabdina polydorae Caullery and Mesnil
Doliocystis sp. Faria, Cunha and Fonseca
Selendium cruzi Faria, Cunha and Fonseca
Polyrhabdina pygospionis Caullery and Mesnil
Monocystis thamnodrili Cogn. de Martiis
Polyrhabdina spiotiis Caullery and Mesnil
Polyrhabdina brasili Caullery and Mesnil

Taeniocystis legeri Cogn. de Martiis
Monocystis roslrata Muslow
Monocystis catenata Muslow
Spirocyslis nidula Leger and Duboscq
Monocystis perforans Pinto

Metamera schubergi Duke

Monocystis minima Konsuloff

Lithocystis foliacea Pixell-Goodrich
Urospora neapolitana Pixell-Goodrich
Ursopora echinocardii Pixell-Goodrich
Urospora echinocardii Pixell-Goodrich
Lithocystis microspora Pixell-Goodrich
Urospora synaptae Cuenot
Gonospora mercieri Cuenot

Gonospora testiculi Trebougoff

83)

STUDIES ON GREGARINES—KAMM

83

Crustacea

Ampelisca spinipes
Anaspides tasmaniae
Atycphyra desmaresti
Balanus amphitrite
Balanus ehurneus

Eriphia spinifrons
Gammarus marinus
Libinia dubia
Nephrops norbegkus
Portuniis depuralor
Pisa gihosii
Talitrus saUator
Talorchestia longicornis
Uca pugnax
Uca pugilator

Chilopoda

Scolopendra heros

" subs pin Ipes

sp.

sp.

sp.

Scolopendrella sp.

DlPLOPODA

Callipus ladarius
Euryurus erythropygus
Fontaneria coarclata
Orthomorpha coarclata
" gracilis

sp.

sp.
Parajidus impressus
" venuslus
" sp.
Rhinocricus pugio

sp.

sp.

sp.

sp.

sp.

Thysanura

Sminthurus fuscus

Orthoptera

Ceuthophilus lalens

" maculalus

" neglectus
" stygius
" valgus

Ccphaloidophora ampelisca Kamm
Ganymedes anaspidis Huxley
Uradiophora cuenoli Mercicr
Pyxinoides bal'ani Tregouboff
Pyxinoides balani Tregouboff
Unnamed parasite, Buddington
Porospora legeri deBeauchamp
Cephaloidophora macidata Leger and Duboscq
Ccphaloidophora olivia Kamm
Porospora nephropsis Leger and Duboscq
Porospora maraisi Leger and Duboscq
Porospora pisae L^ger and Duboscq
Cephaloidophora talilri Mercier
Cephaloidophora dclphinia Kamm
Cephaloidophora nigrofusca Kamm

Amphoroccphalus amphorellus Ellis
Nina indicia Merton
Echinomera magalhaesi Kamm
Seticephalus elegans Kamm
Gregarina brasiliensis Pinto
Gregarine form, Wellmer

Stenophora lactaria Watson
Stcnophora diplocorpa Watson
Slcnophora catidata Watson
Stenophora elongala Ellis
Stenophora robtisla Ellis

Fonsecaia polymorpha Pinto
Stenophora impressa Watson
Stenophora robusla Ellis
Stenophora cockerellae Ellis
Stenophora cunhai Pinto
Stenophora lutzi Pinto
Stenophora cruzi Pinto
Stenophora viannai Pinto
Stenophora umbilicala Pinto
Stenophora tenuicollis Pinto

Gregarine form, Wellmer

Gregarina longiducta Ellis
Gregarina longiducta Ellis
Gregarina neglecta Watson
Gregarina stygia Watson
Gregarina consobrina Ellis

*

84

ILLINOIS BIOLOGICAL MONOGRAPHS

[84

Conocephalus frater
Encoptolophus sordidus
Forficidaria auricularia
Gryllus abbreviatus
Ischnoptera pcnnsylvanicus
Mclanoplus diferenlialis

" femur-rubrum
Udeopsyllae nigra

Hemiptera
Spiniger sp.

Neuroptera

Aeschnidae Iv.
Aeschna sp.
Phryganca grandis
Sympclrum rubicundulum
Tramea lacerata

DiPTERA

Anopheles bifurcatus Iv.
Aphiochaela rufipes Iv.
Ceratophyllus fasciatus
Ceratopsyllus farreni

" fringillae Iv.

" gallinae Iv.

" ad.

" Styx

Ficalbia doflcini Iv.
Simulium braclealiim Iv.
Stegomyia fasciala Iv.

COLEOPTERA

Alobates pennsylvanicus
Amara angtislata
As id a opaca
Asida sp.
Broscus cephaloks
Carahus sp.
Clerid Iv.
Coccinella sp.
'' sp.
Coccinella novemnolata
Coptotomti-s inlerrogatus

Cryplicus quisquilius
Cucujus Iv.
Cychrus rostratus
Dermestes lardarius
Diabrolica viltata
Elateridae Iv.
Eleodes sp.
Eusatlus sp.

Gregarina chagasi Pinto
Gregarina nigra Watson
Gregarine form, Pantel
Gregarina galliverl Watson
Gregarina illinensis Watson
Gregarina nigra Watson

Gregarina udeopsyllae Watson

Schisocystis spiniger Machado

Bolhriopsis claviformis Pinto
Actinoccphalus brachydactylus Ellis
Diplocystis phryganeae Berg-von-Emme
Prismatospora evansi Ellis

Caulleryella anopkelis Hesse

" aphiochactae Keilin

Agrippina bona Strickland
Steinina rotundala Ashworth and Rettie
Actinoccphalus parvus Wellmer
Actinoccphalus parvus Wellmer
Steinina rotundata Ashworth and Rettie

Unnamed par. Guenther

Unnamed par. Strickland

Lankesteria culicis Stevenson and Wenyon

Actinoccphalus zophus Ellis
Steinina rotunda Watson
Stylocephalus gigantcus Ellis
Stylocephalus gigantcus Ellis
Gregarina erecla Wellmer
Cotnctoides sp. Wellmer
Bidbocephalus wardi Watson
Gregarina fragilis Watson
Gregarina katherina Watson

Gregarina globosa Watson
Gregarina coptotomi Watson
Gregarina ovoidea Wellmer
Bidbocephalus clongalus Watson
Gregarine form, Wellmer
Pyxinia bulbifera Watson
Gregarina diabrotica Kamm
Gregarina gracilis Watson
Stylocephalus gigantcus Ellis

85]

STUDIES ON GREGARINES—KAMM

85

Harpalus aeneus
Harpalus pennsylvanicus

Harpalus ruficornis
Heledona agricola
Helophorus aqiiaticus
Hydrophilus aterrimus Iv

sp.
bylobius abielis
Ips typographus
Lagria hirta
Leptochirus edax

a tt

Ninus inlerstitialis

Nydotheres barbarata

Omoplala normalis

Platydema excavatum

Platymis ruficollis

Procrustes coriaceiis

Pterostichus niger
« «

Pterostichus stygicus
(( ((

" vulgaris

Systena sp.

Tenebrio castaneus

Tenebrionidae Iv.

Tribolium ferrugineum
« ((

(C ((

« <c

Tritoma quadripuslulata

Gregarina polyaulia Wellmer
Aclinocephalus gimbeli Watson
erythropus Hirmocystis harpali Watson
longior Steinina harpali Watson

Gregarina polyaulia Wellmer
Gregarine form, Wellmer
Monocystis sp. Wellmer
Cometoides-like form, Wellmer
Botkriopsis terpsichorella Ellis
Gregarina hylobii Kamm
Gregarina typographi Fuchs
Gregarina rostrata Wellmer
Aclinocephalus craMW5 Ellis
Stylocystis ensiferns Ellis
Gregarina guatetnalensis Ellis
Aclinocephalus zophiis Ellis
Gregarina watsoni Pinto
Gregarina platydema Kamm
Gregarina platyni Watson
Aclinocephalus permagnus Wellmer
Gregarina exiguus Kamm
Actinocepkabis echinatus Wellmer
Gregarina monarchia Watson
Gregarina inteslinalis Watson
Aclinocephalus echinatus Wellmer
Gregarina aragaoi Pinto
Gregarina grisea Ellis
Gregarina tenebrionella Watson
Gregarina mintita Ishii
Gregarina crassa Watson
Didymophyes minuta Kamm
Steinina obconica Ishii
Gregarine form, Wellmer

Lepidoptera

Endrosis fenestrella Iv.
Oecophora pseudospretella
Tinea pallescentella

ARACHNffiA

Ctenocaphelus serraticeps
Oribata geniculata

TUNICATA

Stolonica socialis

Enteropneusta

Glossobalanus minutus

Leidyana tinei Keilin
Unnamed Gregarine
Unnamed Gregarine

Gregarina clenocephalus Ross
Gregarina sp. Wellmer

Selysina perforans Duboscq

Selenidium metchnikovi Leger and Duboscq

86 ILLINOIS BIOLOGICAL MONOGRAPHS [86

BIBLIOGRAPHY

ASHWORTH, J. H., AND THEODORE ReTTIE

1912. On a Gregarine Steinina rolundata nov. sp. Present in the Mid-Gut of Bird-Fleas
of the Genus Ceratophyllus. Proc. Roy. Soc. Lond., B86:31-38; 1 pi.
Balbiani, G.

1889. Sur trois Entophytes nouveaux du tube digestif des Myriapodes. Jour. anat.
physiol., 25: 5-45; 1 pi.
Bakgoni, E.

1894. Di un foraminifero parassita nelle salpe e considerazioni sui corpuscoli amilace i
dei protozoi superiori. Ric. Lab. Anat. Normale Univ. Roma, 4: 43-64; 2 pi.
Brasil, Loms

1908. La croissance de Doliocystis dongata dans I'intestin de Lumbriconereis impaiiens
Clap. C. R. soc. biol., 64:355-6.

1908a. Le genre Doliocystis L6ger. C. R. acad. sci. Paris, 146: 425-7.

1909. Documents sur quelques Sporozoaires d'Annelides. Arch. Protist. 16: 107-42,
4 pi.

BUDDINGTON, R. A.

1910. The Behavior and Structure of a New Species of Gregarine. Science n s., 31:470.
Cakus, J. V.

1863. Protozoa, in Handbuch der Zoologie by W. Ch. Peters, J. V. Cams and C. E. A.
Gerstaecker, 2, 1-421.
CaULLERY, M., AND F. Mesnil

1914. Sur resistance de Grdgarines dicystid^es chez les Annelides polychetes. C. R.
soc. biol., 77: 516-20, 10 figs.
Cavolini, F.

1787. Memoria sulla generazione dei Pisci e dei Granchi. Napoli. (Quoted from
Rudolphi, 1808: 130.)

Cavolini, Prni-ipp

1792. Abhandlung iiber die Erzeugung der Fische und der Krebse. (Tr. into German
by E. A, W. Zimmermann, Berlin) (Quoted from Rudolphi, 1810: 130.)

Cecconi, J.

1905. Sur V Anchorina sagittaia Leuck., parasite de la Capilela capitata O. Fabr. Arch.
Protist., 6: 230-244; 2 pi.

Claus, C.

1879. Der Organismus der Phronimiden. Wien. (Quoted from Frenzel, 1885: 556.)

COGNETTI DE MaRTDS, LuIGI

1911. Des crizione d'una nuova Gregarine Policistidea parasita d'un Oligochaete. Arch.
Protist., 23:247-52; 1 pi.

CoMSTOCK, J. H. and a. B.

1912. A Manual of the Study of Insects. Ithaca. 10th. Ed. 701 pp.

Crawley, H.

1903. List of Polycystid Gregarines of the United States. Proc. Acad. Nat. Sci. Phila.,

55: 41-58; 3 pi.
1907. The Polycystid Gregarines of the United States (Third Contribution). Proc.
Acad. Nat. Sc. Phila., 59: 220-8; 1 pi.
Cunningham, J. T.

1907. On Kalpidorhynchus aretiicolae a new Gregarine parasitic in Arenicola ecaudata.
Arch. Protist., 10: 199-215; 2 pi.

87] STUDIES ON GREGARINES—KAMM 87

DiESING, K. M

1851. Systema helminthium, vol. 2. 591 pp.

1859. Revision der Rhjmgodeen. Stizb. Akad. Wiss. Wien math.-natw. Kl., 37:
719-82; 3 pi.

DUFOUK, L.

1828. Note sur la gr^garine nouveau genre de ver qui vit en tropeau dans les intestins
de divers insectes. Ann. sci. nat., (1) 13: 366-8; 1 pi.

1837. Recherches sur quelques entozoaires et larves parasites des insectes OrthoptSres
et Hymenopteres. Ann. sci. nat., (2) 7: 5-20; 1 pi.

DUJARDIN, F.

1845. Histoire naturelle des helminthes ou vers intestinaux. Paris. (Quoted from
Labb6.)

Duke, H. Lyndhtjrst

1910. Some Observations on a New Gregarine (Metamera schuhergi nov. gen., nov.
spec). Quar. Jour. Micr. Sci., 55: 261-86; 2 pi.
Elus, M. M.

1912. A New Species of Polycystid Gregarine from the United States. Zool. Anz.,
39: 25-7.

1913. A Descriptive List of the Cephaline Gregarines of the New World. Trans. Amer.
Micr. Soc, 32: 259-96, 4 pi. •

1914. An Acanthosporid Gregarine from North American Dragonfly Nymphs. Trans.
Amer. Micr. See, 33: 215-22, 7 figs.

Fasia, Gomez de, Marques de CtWHA, and O. da Fonseca

1918. Protozoarios parazitos de "Polydora Socialis." Mem. Inst. Osw-Cruz 10:
17-19; 1 pi.
Feantzius, a. von

1846. Observationes quaedam de Gregarinis. Berolini. (Quoted from Frantzius, 1848.)
1848. Einige nachtragliche Bemerkungen iiber Gregarinen. Arch. Naturg., 14: 188-96;

Ipl.
Fbenzel, J.

1885. Ueber einige in Seethieren lebende Gregarinen. Arch. mikr. Anat., 24: 545-88;
2 pi.
Grebnecki, ?

1873. Mem. Soc. Nat. Nouvelle-Russie. Odessa. (Quoted from Sokolow, 1911.)
Hammerschmidt, K. E.

1838. Hehninthologische Beytrage. Isis (Oken), (5), 351-8; 4 pi. (Quoted from Stein,
1848.)

Huxley, J. S.

1910. On Ganymedes anaspidis (nov. gen., nov. sp.), a Gregarine from the digestive
tract of Anaspides tasmaniae (Thompson). Quart. Jour. Micr. Sci., 55: 155-75;
Ipl.

JOHANSEN, H.

1894. Aclinocephalus goronowilschi, eine anscheinend neue Gregarineform. ZooL Anz.,
17: 140-5; 4 figs.

Kfh.tn, D,

1918. On the Occurrence of a Cephaline Gregarine, Leidyana tinei n. sp., in Lepidopter-
ous Larvae. Parasit., 10:406-10; 1 pi.

Kororo, C. A. and Olive Swezey

1919. Studies on the Parasites of the Termites. Univ. Calif. Publ. Zool., 20: 1-116;
14 pi .

88 ILLINOIS BIOLOGICAL MONOGRAPHS [88

KOLLIKER, A.

1845. Die Lehre von der thierischen Zelle und den einfachen thierischen Formelementea
nach den neuesten Fortschritten. Zeit. wiss. Bot., 1; 97.

1848. Beitrage zur Kenntniss niederer Thiere. Zeit. wiss. ZooL, 1: 1-37; 3 pi.
Labb£, a.

1899. Sporozoa. Das Tierreich, pt. 5; 180 pp.
Lankester, E. R.

1863. On Our Present Knowledge of the Gregarinidae. Quar. Jour. Micr. Sci., 3: 83-96.
L£ger, Louis

1892. Recherches sur les gr^garines. Tabl. zool., 3: 1-182; 22 pi.

1893. L'evolution des gr^garines intestinales des vers marins. C. A. acad. sci., Par.,
116: 204-6.

1897. Nouvelles recherches sur les polycystidees parasites des arthropodes terrestres.
Ann. fac. sci. Marseille, 6: 54 pp., 2 pi.

1899. Sur les Gr^garines des Dipt^res et description d'une esp^ce nouvelle. Ann. See
Ent. France, 68: 526-33; 4 pi. ,

1901. Sur une nouvelle Gr^garine parasite des Pinnotheres des mouies, C. R. acad.
sci., Par., 132:1343-6;! fig.

1904. Sporozoaires parasites de I'Embia Solieri Rambur. Arch. Protist., 3: 358-66; 7
figs

1905. Un nouveau type cellulaire de Gr6garine h, cytoplasme m6tam6ris6. C. R. acad.
sci., Par., 140: 524-6.

1906. fitude sur Taeniocystis mira L€ger, Gr6garine m6tam6rique. Arch. Protist.,
7: 307-29; 2 pi.

L£ger, Lotns AND O. Duboscq

1902. Les Gregarines et I'^pith^lium intestinal chez des Trach6ates. Arch, parasit.,
6: 377-473; 5 pi.

1903. Aggregata vagans n. sp, Gr6garine gymnosporee parasite des Pagures. Arch,
zool. expfir., (4) 1: cxlvii-cli; 1 fig,

1904. Nouvelles recherches sur les Gregarines et I'^pith^lium intestinal des Trach^ates.
Arch. Protist., 4: 335-83; 2 pi.

1907. L'evolution des Frenzelina (n.g.) Gregarines intestinales des Crustac^s decapodes.
C. R. acad. sci.. Par. 145: 773-4.

1908. L'6volution schizogonique de \ Aggregata (Eucoccidium) eberthi Labbfi. Arch.
Protist., 12: 44-108; 3 pi.

1909. Sur une micros poridie parasite d'une Gr^garine. C. R. acad. sci.. Par., 148: 733-4.

1909a. fitudes sur la sexuality chez les Gregarines. Arch. Protist., 17: 19-134; 5 pi.

1909b. Protistes parasites de I'intestin d'une larve de Ptychoptera et leur action sur
I'hdte. Bull. Acad. Roy. Belgique, CI. sc, 1909: 885-902; 4 pl.

1911. Deux Gregarines de Crustaces. Porospora porlunidarum Frenz. et Cephaloido'

phora maculata n. sp. Arch. zool. exper., (5) 6: lix-lxx; 6 figs.
1911a. Deux nouvelles especes de Gregarines appartenant au genre Porospora. Amu
Univ. Grenoble, 23: ?. (Quoted from Mercier, 1912a.)
Leuckart, R.

1860. Bericht uber die Leistungen. Arch. Naturg., 26 (2): 263.
Magalhaes, P. S. de

1900. Notes d'helminthelogie bresilinne. Arch, paras t., 3: 34-69.

89] STUDIES ON GREGARINES—KAMM 89

Mawrodiadi. p.

1908. Les Balanes de la Mer Noire et les Gr^garines leurs parasites. Note pr^liminaire
(in Russian). Mem. Soc. Natural. Nouvelle-Russie, Odessa, 32:101-33. (Quoted
from L^ger and Duboscq 1911.)
Mercter, Louis

1911. Cephdoidophora cuenoti n. sp. Gr^garine parasite du tube digestif de la Caridine.
C. R. soc. biol., 71:51-3.

1912. Cephaloidophora talitri n. sp., Gr^garine parasite du Talitre. C. R. soc. biol.,
72:38-9, 1 fig.

1912a. N^cessit^ de retirer la Gr^garine de la Caridine {Cephaloidophora Cuenoti Mercier)
du genre Cephaloidophora. Arch. zool. exp6r., (5) 9: xli-xlvi.

1912b. Monographic d'Uradiophora Cuenoti. Gr6garine parasite du tube digestif de la
Caridine. Arch. zool. exp6r., (5) 10: 177-202; 2 pi.
MiNCBIN, E. A.

1903. Introduction and Protozoa. Pt. I, Fasc. 2, A Treatise on Zoology. Ed. by E. R.
Lankester. London 451 pp..

1912. An Introduction to the Study of the Protozoa. London. 520 pp. 194 fig.

MiNGAZZINI, P.

1891. SuUa distribuzione delle gregarine policistidee, Atti Ace. Lincei, Rend., Roma,

(4) 7, 1, 234-7, 2 figs.
1891a. Gregarine monocistidee, nuove o poco conosciute, del Golfo di Napoli. Atti
^ Ace. Lincei. Rend., Roma, (4) 7, 2: 229-35.

1891b. Gregarine monocistidee, nuove o poco conosciute del Golfo di Napoli. Atti Ace.

Lincei Rend., Roma, (4) 7, 1: 467:74.
1893. Contributo alia conoscenza degli Sporozoi. Ric. Lab. Anat. Normale Univ. Roma,

3: 31-85; 2 pi.

MOSELEY, H. N.

1874. On the Structure and Development of Peripatus capensis. Phil. Trans. Roy. Soc.
Lond., 164: 757-82; 3 pi.
NowLiN, Nadine and Inez Smith

1917. The Intracellular Development of a Gregarine, Frcnzelina ampdisca n. sp. Jour.
Parasit., 4: 83-88; 1 pi.

NUSSEAUM, ?

1890. Anatomische Studien an califomischen Cirripedien. Bonn, 97 pp., 12 pi. (Quoted
from Tr^gouboff, 1912).
Pfeiffer, L.

1895. Die Protozoen als Krankheitserreger, Nachtrage. Jena; 60 pp.
Pinto, Cesar F.

1918. Sobre as eugregarines parasitas dos artropodes brasileiros. Brazil Medico, 7, 8,
9, 12, 13, 15, 26, 30, 35, 38, 39, 41.

1919. Contribuigao ao estudo das Gregarinos. Rio de Janeiro. 113 pp., 6 pi.
Plate, Lpdwig

1886. Untersuchungen einiger an den Kiemenblattem des Gammarus pulex lebenden
Ektoparasiten. Zeit. wiss. Zool., 43: 175-241; 2 pi.

POKTES, J. F.

1897. Trichonympha, and Other Parasites of Termes fiavipes. Bull. Mus. Harvard,

31: 45-48; 6 pi.
1897a. Two New Gregarinida. Jour. Morph., 14: 1-20; 3 pi.
Ramdohr, ?

1811. Abhandlung ijber die Verdauungswerkzeuge der Insekten. 194 pp., 22 pi

(Quoted from Stein, 1848.)

90 ILLINOIS BIOLOGICAL MONOGRAPHS [90

Redi, Francesco

1684. Intorni agli animali viventi, che si trovano negli animali viventi. Firenze. 232

pp., 26. (Quoted from Rudolphi.)
1708. De animalculis vivis qui in Corporibus Animalium vivorum reperiuntur, observa-

tiones. Amstelodami.
1729. Opusculorum Pars tertian sive de animalculis vivis, quae in corporibus animalium

vivorum reperiuntur, observationes. Ex Etruscis Latinas fecit Petrus Coste.

342 pp., 26 pi. (Quoted from L^ger and Duboscq, 1908).
RlTTER, W. E.

1893. Tunicata of the Pacific Coast of North America. I. Perophora annectens n. sp.

Proc. Calif. Acad. Sci., (2) 4: 37-85; 3 pi.

ROBOZ, ?

1886. Math, naturw. Ber. Ungarn, 4: 146. (Quoted from Labb6).

Ross, E. H.

1909. A Gregarine Parasitic in the Dog-Flea Clenocepkalus serralicsps. Ann. Trop.
Med. Par., 2:359-63.

ROSSLER, R.

1882. Beitrage zur Anatomic der Phalangiden. Zeit. wiss. Zool., 36: 671-702; 2 pi.

Rudolphi, C. A.

1808-10. Entozoorum, sine Vermium Intestinalium Historia Naturalis. Amstelaedami.

1 (1808): 527 pp., 6 pi.; 2, pt. 1 (1809): 458 pp., 6 pi.; 2, pt 2 (1810):

386 pp.
1819. Entozoorum Synopsis. Berolini, 811 pp., 3 pi.

Saint- Joseph, M. le Baron de

1907. Les Ann^lides Polych^tes de cotes de France. Ann. sci. nat., (9) 3: 145-260; 5 pi.

Schneider, Aime

1875. Contributions 4 I'histoire des gregarines des invertebres de Paris et de Roscoff.

Arch. zool. exp6r., 4: 493-604; 7 pi.
1882. Seconds contribution i I'etude des gregarines. Arch. zool. exp^r., 10: 423-50; 1 pi.
1885. Etudes sur les d^veloppement des gregarines. Tabl. zool., 1: 10-24; 6 pi.

1887. Grdgarines nouvelles ou peu connues. Tabl. zool., 2: 67-85; 1 pi.

Scudder, S. H.

1882. Nomenclator Zoologicus. Bull. U. S. Nat. Mus., 19; 716 pp.

SlEBOLD, C. Th. von

1839. Beitrage zur Naturgeschichte der wirbellosen Thiere IV. Neuest. Schrift. Naturf.
Gesell., Danzig, 3: 56-71; 1 pi. (Quoted from Diesing, 1851.)
SOKOLOW, B.

1911. Liste des Gregarines d^crites depuis 1899. Zool. Anz., 38: 277-295.

1911a. Liste des Gregarines d^crites depuis 1899a. Zool. Anz., 38: 304-14.
SoKOLOW, Ivan

1908. Zur Kenntniss der phagocytSren Organe von Scorpio indicus. Zool. Anz., 33:
497-503; 8 figs.

SOLGER, BeRNH.

1890-1. Notiz iiber eine im Darmkanal von Balanus improvisus Darw. (var. gryphica
Miinter) lebende Gregarine. Mitt. Naturw. Vereine Neuvorpommern und Riigea
im Greifswald, Berlin, 32: 99-102; 1 fig. (Quoted from Tr^gouboff, 1912).
Stein, F. von

1848. Uber die Natur der Gregarnen. Arch. Anat. Physiol. Med., 1848: 182-223; 1 pi.

911 STUDIES ON GREGARINES—KAMM 91 |

Strickland, C.

1912. Agrippina bona n.g., et n. sp., representing a New Family of Gregarines. Parasit., ,

5:97-108; 1 pi. I

Stuart, Alex.

1871. Uber den Bau der Gregarinen. Bull. Acad. Imper. Sci. St. Petersbourg, 15: i

498-502; 1 pi. 1

Tr£gouboff, G.

1912. Sur les Gr6garines des Balanes. Arch. zool. exp^r., (5) 10: liii-lxi, 3 figs. |

Watson, M. E. i

1915. Some New Gregarines from Arthropoda. Jour. Parasit., 2: 27-36; 2 pi.

1916. Three New Gregarines from Marine Crustacea. Jour. Parasit., 2: 129-36; 1 pi.
1916a. Studies on Gregarines. 111. Biol. Monogr., 2: 211-468; 15 pi.

1916b. Observations on Polycystid Gregarines from Arthropoda. Jour. Parasit., 3:

65-75; 1 pi. j

Wellmer, Leo '

1911. Sporozoen ostprussischer Arthropoden. Schr. Physik-Okon. GeseUsch. Konigsb., •

52: 103-64; 1 pi. ;

92 ILLINOIS BIOLOGICAL MONOGRAPHS [92

EXPLANATION OF PLATES

Drawings were either traced directly from the author's plates or
enlarged by means of the camera lucida. Magnifications are not given
for they did not appear in most of the original sources. The text will
furnish the exact size of each species so far as data were available in the
original publication or in the later articles.

93]

STUDIES ON GREGA JUNES— K AM M 93

PLATE I

94 ILLINOIS BIOLOGICAL MONOGRAPHS [94

EXPLANATION OF PLATE I

Fig. 1 — Schneideria mucronata L^ger. After L6ger 1892, Plate II, fig. 9.

Fig. 2 — Schneideria mucronata L€ger. After L^ger 1892, Plate II, fig. 8. i

Fig. 3 — Schneideria mucronaia Leger. After Leger 1892, Plate II, fig. 10. }

Fig. 4 — Schneideria caudata L6ger. After Leger 1892, Plate II, fig. 4. 1

Fig. 5 — Schneideria caudata Leger. After Leger 1892, Plate II, fig. 3. \

Fig. 6—Actinocephalus tipulae Leger. After Leger 1892, Plate XIV, fig. 8. |

Fig. 7—Gregarina longa Labb6. After Leger 1892, Plate VIII, fig. 7. j

Fig. 8 — Hirmocystis ventricosa Labbe. After Leger 1892, Plate VI, fig. 2. I

Fig. 9 — Hirmocystis ventricosa Labbe. After L^ger 1892, Plate VI, fig. 1. '

Fig. 10— Hirmocystis polymorpha Labb6. After Leger 1892, Plate III, fig. 1. '

Fig. 1 1 — Hirmocystis polymorpha Labbe. After L6ger 1892, Plate III, fig. 5.

Fig. 12 — Actinocephalus sieboldii Frantzius. After Kolliker 1848, Plate II, fig. 16.

Fig. 13 — Hoplorhynchus oligacanthus Schneider. After Schneider 1875, Plate XVI, fig. 25.

Fig. 14 — Hoplorhynchus oligacanthus Schneider. After Plate XVI, fig. 30. '.

Fig. 15 — Pileocephalus heerii Schneider. After Schneider 1877, Plate XXXI, fig. 4. '

Fig. 16— Pileocephalus heerii Schneider. After Schneider 1887, Plate XXXI, fig. 5.

Fig. 17 — Pileocephalus chinensis Schneider. After Schneider 1875, Plate XVI, fig. 23.

Fig. 18 — Pileocephalus chinensis Schneider. After Schneider 1875, Plate XVT, fig. 21.

Fig. 19 — Geneiorhynchus monnieri Schneider. After Schneider 1875, Plate XX, fig. 23. /

Fig. 20 — Geneiorhynchus monnieri Schneider. After Schneider 1875, Plate XX, fig. 25. ' I

Fig. 21 — Geneiorhynchus monnieri Schneider. After Schneider 1875, Plate XX, fig. 26. j

Fig. 22 — Gregarina clavata Kolliker. After Schneider 1887, Plate X bis, fig. 3. j

Fig. 23—Asterophora elegans L6ger. After Leger 1892, Plate XII, fig. 9. !

Fig. 24r—Asterophora elegans Leger. After L6ger 1892, Plate XII, fig. 12.

Fig. 25 — Asterophora mucronata Leger. After Leger 1892, Plate XV, fig. 7. I

Fig. 26— Asterophora mucronata L6ger. After Leger 1892, Plate XV, fig. 8. j

Fig. 27 — Ancyrophora uncinata Leger. After Leger 1892, Plate XIX, fig. 8.

Fig. 2S—Discorhynchus truncalus Labbe. After L6ger 1892, Plate XV, fig. 10. i

Fig. 29— Discorhynchus truncalus Labbe. After Leger 1892, Plate XV, fig. 11. i

Fig. 30—Menospora polyacantha Leger. After L^ger 1892, Plate XIX, fig. 5. j

Fig. 31 — Menospora polyacantha I>eger. After Leger 1892, Plate XIX, fig. 3.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

STUDIES ON GREGARINES II

PLATE I

95] STUDIES ON CRECARINES—KAMM 95

PLATE II

96 ILLINOIS BIOLOGICAL MONOGRAPHS [96

EXPLANATION OF PLATE II

Fig. 32— Menospora polyacantha L^ger. After L6ger 1892, Plate XIX, fig. 2.

Fig. 33—Slylocystis praecox Leger. After L^ger 1899, fig. la.

Fig. 34 — Taeniocystis mira Leger. After L^ger 1906, Plate XII, fig. 8.

Fig. 35 — Taeniocystis mira L^ger. After L6ger 1906, Plate XII, fig. 7.

Fig. 36 — Gregarina marteli L4ger. After Leger 1904, fig. 2.

Fig. 37 — Gregarina marteli L^ger. After L^ger 1904, fig. 1.

Fig. 38 — Gamocystis ephemerae Labb6. After Schneider 1882, Plate XIII, fig. 31.

Fig. 39_ ? Frenzel. After Frenzel 1885, Plate XXVI, fig. 35. !

Fig. 40 — Geneiorhynchus aeschnae Crawley, After Crawley 1907, Plate XVIII, fig. 4. i

Fig. 41 — Ancyrophora uncinate L6ger. After Leger 1892, Plate XIX, fig. 10.

Fig. 42 — Ancyrophora uncinala L^ger. After Leger 1892, Plate XIX, fig. 9, I

Fig. 43 — Asterophora elegans L6ger. After L6ger 1892, Plate XII, fig. 14.

Fig. 44 — Actinocephalus brachydactylus Ellis. After Ellis 1913, Plate XVIII, fig. 33.

Fig. iS— Actinocephalus brachydactylus Ellis. After Ellis 1913, Plate XVIII, fig. 31.

Fig. 46 — Pileocephalus heerii Schneider. After Kolliker 1848, Plate I, fig. 11.

Fig. 47 — Pileocephalus heerii Schneider. After Schneider 1887, Plate III, fig. 1.

Fig. 48 — Coleorhynchus heros Labb6. After Labbe 1899, fig. 52.

Fig. 49 — Coleorhynchus heros Labbe. After Labb^ 1899, fig. 53.

Fig. 50— [? reduvii Ramdohr]. After Stein 1848, Plate IX, fig. 35. |

Fig. 51 — ?Gregarina termilis Leidy, After Leidy 1881, Plate LII, fig. 27. ;

Fig. 52— Gregarina lagenoides Labb6. After L^ger 1892, Plate VI, fig. 7. I

Fig. 53 — ?Gregarina podurae L6ger. After L^ger 1892, Plate X, fig. 2. I

Fig. 54—?Gregarina podurae L6ger. After L^ger 1892, Plate X, fig. 4. i

Fig. 55— [? KoUiker]. After Kolliker 1848, Plate III, fig. 33. j

Fig. 56— ?Gregarina termitis Leidy. After Ellis 1913, Plate XVII, fig. 6. j

Fig. 57— Acanthospora repelini L^ger. After L6ger 1897, Plate III, fig. 16. i

Fig. 58 — Anthorhynchus sophiae Labb6. After Schneider 1887, Plate X, fig. 11. ';

Fig. 59— Sciadophora phalangii Labb6. After L^ger 1897, Plate III, fig. 2. ]

Fig. ^y— Sciadophora phalangii Labb6. After L6ger 1897, Plate III, fig. 3. j

Fig. 61— Bothriopsis claviformis Pinto. After Pinto 1919, Plate IV, fig. 65. |

The following two figures are also included in this plate: \

Fig. 90— Gamocystis ephemerae Labb6. After Frantzius 1848, Plate VII, fig. VII, 1. j

Fig. 99 — Prismatospora evansi Ellis. After Ellis 1914, fig. 6. j

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

KAMM

STUDIES ON GREGARINES II

PLATE II

971 STUDIES ON GREGARINES—K AM M 97

PLATE III

98 ILLINOIS BIOLOGICAL MONOGRAPHS [98

EXPLANATION OF PLATE III )

r
(

Fig. 62— Sciadophora phalangii Labbe. After L6ger 1897, Plate III, fig. 4. !

Fig. 63 — Uradiophora cuenoti Mercier. After Mercier 1912 b, PI. XIII, fig. 1. I

Fig. 64 — Didymophyes longissima Frantzius. After Kolliker 1848, PI. Ill, fig. 29, a. j

Fig. 65— Pyxinoides balani Trdgouboff. After Frenzel 1885, PI. Ill, fig. 34. li

Fig. 66 — Cephaloidophora conformis L6ger and Duboscq. After Frenzel 1885, PI. XXVI, i

fig. 65. ' ;

Fig. 67— [? gammari von Siebold]. After Kolliker 1848, PI. Ill, fig. 29, c. ,

Fig. 68 — Cephaloidophora tnaculata L^ger and Duboscq. After L^ger and Duboscq 1911, |

fig. 4. I

Fig. 69— [? clausii Frenzel]. After Frenzel 1885, PI. XXVI, fig. 57.
Fig. 70— [? nicaeae Frenzel]. After Frenzel 1885, PI. XXVI, fig. 62.

Fig. 71 — Cephaloidophora ampelisca Kamm. After Nowlin and Smith 1917, PI. I, fig. 11.
Fig. 72, 73 — Pileocephalus heerii Schneider. After Frantzius 1848, PI. VII, fig. 1, 4, 6.
Fig. 74 — Aciinocephalus octacanthus Frantzius. After Frantzius 1848, PI. VIT, fig. VI, 2. |

Fig. 75 — Cephaloidophora conformis L^ger and Duboscq. After L6ger and Duboscq 1909 a, ;

fig. 31. I

Fig. 76 — Gregarina mystacidarum Frantzius. After Frantzius 1848, PI. VII, fig. VI, 2. j

Fig. 77 — Cephaloidophora communis Mawrodiadi. After TrdgoubofF 1912, fig. lb. j

Fig. 78 — Pyxinoides cthamali Tr^gouboff. After Tregouboff 1912, fig. 3, c. ]

Fig. 79 — Cephaloidophora fossor Tregouboff. After L^ger and Duboscq 1909 a, fig. 29.
Fig. 80 — Cephaloidophora ocellata Kamm. After L6ger and Duboscq 1909a, fig. 27. i

Fig. 81 — Pyxinoides cthamali Tr6gouboff. After Tregouboff 1912, fig. 3, e. I

Fig. 82 — Pyxinoides balani Tregouboff. After Tregouboff 1912, fig. 2, a. ,

Fig. 83 — Cephaloidophora olivia Kamm. After Watson 1916, PI. I, fig. 8.
Fig. 84 — Cephaloidophora nigrofusca Kamm. After Watson 1916, PI. I, fig. 11.
Fig. 85 — Cephaloidophora delphinia Kamm. After Watson, 1916, PI. I, fig. 1.

Fig. 86 — Cephaloidophora talitri Mercier. After Mercier 1912, fig. 1. i

Fig. 87 — Ganymedes anaspidis Huxley. After Huxley 1910, fig. 1.

Fig. 88 — Pyxinoides balani Tr6gouboff. Tregouboff 1912, fig. 2, b. |

Fig. 89— Gregarina clavata Kolhker. After Kolliker 1848, Plate I, fig. 10. ;

Fig. 90 — See explanation of Plate II for figure 90.
Fig. 91 — Coelomic spore of Pileocephalus heerii Schneider. After Schneider 1887, Plate

XXXII, fig. 2. .

Fig. 92 — Coelomic spore of Pileocephalus heerii Schneider. After Schneider 1887, Plate j

XXXII, fig. 3.
Figs. 93, 94 — Two forms of intestinal spores of Pileocephalus chinensis Schneider. After |

Schneider 1875, Plate XVI, fig. 24. \

Fig. 95 — Prismatospora evansi EUis. After Ellis 1914, fig. 3. '

Fig. 96 — Prismatospora evansi EUis. After Ellis 1914, fig. 4.

Fig. 97 — Prismatospora evansi EUis. After EUis 1914, fig. 5. j

Fig. 98— [sp. Ritter.] After Ritter 1893, Plate III, fig. 22.
Fig. 99 — See explanation of Plate II for figure 99. ;

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

KAMM

STUDIES ON GREGARINES II

PLATE III

991 STUDIES ON GREG ARINES— RAM M 99

PLATE IV

100 ILLINOIS BIOLOGICAL MONOGRAPHS [100 :

>

EXPLANATION OF PLATE IV

Fig. 100 — Pileocephaltis striatus L6ger and Duboscq. After L6ger and Duboscq 1909 b,
Plate II, fig. 1.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.

Fig.
Fig.

Fig-
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.

Fig.

Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

01 — Agrippina bona Strickland. After Strickland 1912, fig. 5.

02 — Agrippina bona Strickland. After Strickland 1912, fig. 9.

03 — Agrippina bona Strickland. After Strickland 1912, fig. 31.

04 — Sciadophora goronowilschi Labbe. After Johansen 1904, fig. 1.

05 — Sciadophora goronowilschi Labb6. After Johansen 1904, fig. 3.

06 — Sciadophora phdlangii Labb6. After Leger 1896, Plate III, fig. 7.

07 — Acanthospora repelini L6ger. After L6ger 1896, Plate III, fig. 17.

08 — Anthorhynchus sophiae Labb6. After Schneider 1887, Plate X, fig. 16.

09— [Not named] Sokolow. After Sokolow 1908, fig. 8a.

10— Sciadophora fissidens Labb6. After Rossler 1882, Plate XLII, fig. 21a.

11 — Sciadophora fissidens Labbe. Protomerite from above. After Rossler 1882,

Plate XLII, fig. 21b.
12 — Sciadophora catidatus Kamm. After Rossler 1882, Plate XLII, fig. 22a.
13 — Sciadophora caudatus Kamm. Protomerite from above. After Rossler 1882,

Plate XLII, fig. 22b.
14 — Actinocephalus parvus Wellmer. After Wellmer 1911, text-figure 9a.
15 — Leidyana tinei Keilin. After Keilin 1918, Plate EX, fig. 5.
16 — Leidyana tinei Keilin. After Keilin 1918, Plate IX, fig. 6.
17— [? Frenzel]. After Frenzel 1885, Plate XXVI, fig. 39.

18 — Sycia inopinata Leger. After L6ger 1892, Plate V, fig. 12.
19 — Sycia inopinata L6ger. After Leger 1892, Plate V, fig. 14.
20 — Actinocephalus parvus Wellmer. After Wellmer 1911, fig. 9b.
21 — Steinina rolundata Ashworth and Rettie. After Ashworth and Rettie 1912, Plate

I, fig. 3.
22 — Steinina rotundata Ashworth and Rettie. After Ashworth and Rettie 1912,

Plate I, fig. 5.
23 — Steinina rotundata Ashworth and Rettie. After Ashworth and Rettie 1912, Plate

I, fig. 8.
24 — Steinina rolundata Ashworth and Rettie. After Ashworth and Rettie 1912, Plate

I, fig. 11.
25 — Lecudina aphroditae Labb6. After Minchin 1903, fig. 19, p. 177.
26 — Lecudina pellucida Leger. After Kolliker 1848, Plate I, fig. 31a.
27 — Ulivina elHptica Mingazzini. After Mingazzini 1891, 1st sem. fig. 2, p. 236.
28 — Polyrhabdina spionis Mingazzini. After Kolliker 1848, Plate I, fig. 7.
29 — Polyrhabdina spionis Mingazzini. After Caullery and Mesnil 1914, fig. 9.
30—Taemocystis legeri Cognetti de Martiis. After Cognetti 1911, Plate I, fig. 3.
31 — Metamera schubergi Duke. After Duke 1910, Plate XV, fig. 5.
32 — Metamera schubergi Duke. After Duke 1910, Plate XV, fig. 1.
33— Metamera schubergi Duke. After Duke 1910, Plate XVI, fig. 35.
34 — Lecudina elongala Labbe. After Brasil 1909, Plate IX, fig. 6.
35 — Lecudina elongata Labbe. After Brasil 1909, Plate IX, fig. 13.
36 — Gregarina pterotracheae Labbe. After Stuart 1871, Plate I, fig. 4.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

126 ^loe

KAMM

STUDIES ON GREGARINES II

PLATE IV

101]

STUDIES ON GREGARINES—KAMM

lOi

INDEX

Acanthospora pileata, 53

polymorpha, 53

repelini, 53
Actinocephalus brachydactylus, 64

caudatus, 76

fissidens, 51

goronowitschi, 52

octacanthus, 63

parvus, 79

sieboldii, 62, 70

sp., 75, 76

tipujae,- 74, 75
Acerata, List of Gregarines in, 49
Aeschna constricta, 65

sp., 64
Aeschnida sp., 65
Aggregata coelomica, 40

conformis, 38

nicaeae, 48

praemorsa, 47

vagans, 41
Agrion puella, 72

sp., 63
Agrippina bona, 80
Ampelisca spinipes, 44
Anaspides tasmaniae, 45
Ancyrophora uncinata, 70
Anthocephalus sophiae, 50
Anthorhynchus sophiae, 50
Aphrodite aculeata, 24
Astacus fluviatilis, 46
Asterophora elegans, 66

mucronata, 65, 66
Atyaephyra desmaresti, 34
Audouinia filigera, 29

Lamarkii, 28

tentaculata, 28, 29

Balanus amphitrite, 35, 40, 47
amphitrite pallidus, 47
eburneus, 35, 40, 47
improvisus, 40
improvisus gryphica, 40

perforans, 40

perforatus, 44

pusillus, 35

sp., 44

tintinnabulum, 35

tintinnabulum communis, 40
Bibio marci, 77
Bothriopsis claviformis, 65
Bullulina tipulae, 75

Calopteryx splendes, 70

virgo, 70
Cancer depressus, 39

pagurus, 39, 47
Cephaloidophora ampelisca, 43

communis, 40, 44

conformis, 38, 48

cuenoti, 34

delphinia, 43

fossor, 40, 42

maculata, 34, 42

nigrofusca, 43

ocellata, 41

oHvia, 42

praemorsa, 47

talitri, 42
Ceratophyllus fasciatus, 80
Ceratopogon solstitialis, 78
Ceratopsyllus fringillae, 79

gallinae, 79

Styx, 79
Chironomus sp., 77
Clepsidrina granulosa, 60

lagenoides, 54

longa, 73

nystacidarum, 61

podurae, 55
Clepsine elongata, 30
Coelhelminthes, List of Gregarines found in,

21
Coleophora heros, 57
Coleorhynchus heros, 57
Colymbetes sp., 70

102

ILLINOIS BIOLOGICAL MONOGRAPHS

[102

Crustacea, List of Gregarines found in, 31
Ctenocephalus serraticeps, 74
Ctenophora pcctinicornis, 75

sp., 76
Cthamalus stellatus, 36

Didymophyes longissima, i2, 45, 46
Diptera, List of Gregarines found in, 73
Discorhynchus truncatus, 66
Doliocystis aphroditae, 24

clongata, 24

heterocephala, 25

legeri, 26

pellucida, 23

polydorae, 25

rhyncoboli, 29

sp., 25, 27
Dytiscus sp., 70

Eirmocystis polymorpha, 74

ventricosa, 74
Embia solieri, 61
Endrosis fenestrella, 72
Ephemera vulgata, 60, 62
Eupagurus prideauxi, 41
Frenzelina ampelisca, 43

conformis, 38

cthamali, 35

delphinia, 43

fossor, 40

nigrofusca, 43

ocellata, 41, 42

olivia, 42

praemorsa, 47
Gammarus marinus, 34, 42

pulex, 32), 45, 46
Gamocystis ephemerae, 60, 62

francisci, 60, 62
Ganymedes anaspidis, 45
Geneiorhynchus aeshnae, 65

monnieri, 64
Glossophonia complanata, 30
Glycera convoluta, 26
Gregarina balani, 35, 44

caudata, 76

clausi, 48

clavata, 60, 62

conformis, 38, 39

ctenocephalus, 74

ctenocephalus canis, 74

difBuens, ii, 46

dytiscorum, 70 ^

ensiformis, 81

ephemerae, 62

flava, 81

frantziusiana, 63, 68

gammari, Zi, 45, 46

granulosa, 60

heerii, 63, 68

lagenoides, 54

longa, 73, 74

longissima, Z2>, 46

marteli, 61

miliaria, 33, 46

mystacidarum, 61, 67

nicaeae, 48

oligacantha, 62, 69

ovata, 72

pellucida, 23

phallusiae, 80

phymatae crassipedis, 58

podurae, 55

praemorsa, 39, 47, 48

psocorum, 72

pterotracheae, 30, 31

putanea, ii, 46

reduvii, 58

salpae, 81

sieboldii, 62

soror, 58

sp., 3i, 40, 45, 46, 53, 54, 56, 81

spionis, 27

termitis, 56

tipulae, 75

valettei, 47
Hemiclepsis marginata, 30
Hemiptera, List of Gregarines found in, 57
Hirmocystis polymorpha, 74

ventricosa, 74
Hoplorhynchus oligacanthus, 62, 63, 69
Hyale ppntica, 49
Hyalospora psocorum, 72

reduvii, 58

Isoptera, List of Gregarines found in, 56

Kynotus pittarelli, 30

Lecudina aphroditae, 24
elongata, 24
heterocephala, 25
legeri, 26
leukarti, 24
pellucida, 22, 23

103]

STUDIES ON GREGARINES—KAMM

103

polydorae, 25

sp., 25, 26
Leidyana tinei, 72

Lepidoptera, List of Gregarines found in, 72
Lepisma saccharina, 55
Libellules sp., 64
Libinia dubia, 43
Limnobia sp., 75
Limnophilus rhombicus, 70
Lumbriconereis inipatiens, 24
Lycosella phalangii, 50, 51, 52

Malacopoda, List of Gregarines found in, 54
Menospora polyacantha, 71
Metamera schubergi, 30

sp., 30
Monocystis aphroditae, 24

nereidis, 23
Mystacides sp., 61, 67

Nebalia serrata, 49
Nematoides fusiformis, 35, 44
Nepa cinerea, 57
Nephthys scolopendroides, 25
Nereis beaucourdrayi, 23

cultrifera, 23
Neuroptera, List of Gregarines found in, 59
Nicolea venustula, 29
Notomastus exsertilis, 26

Oecophora pseudopretella, 73
Ophioidina elongata, 24

heterocephala, 25
Opilio grossipes, 51, 53
Orchesella sp. 55
Orchestia littorea, 33
Oribata geniculata, 53

Pachygraspus marmoratus, 39
Pachyrhina pratensis, 74, 75
Peripatus capensis, 54
Perophora annectens, 81
Petaloproctus terricola, 26, 29
Phalangidae sp., 51, 52
Phalangium cornutum, 51, 53

crassum, 51

opilio, 50, 53
Phallusiae mamillaris, 80
Phronima sp., 48
Phronimella sp., 48

Phryganea grandis, 64, 66, 69

rhumbica, 70

sp., 63

varia, 69
Pileocephalus chinensis, 61, 67, 79

heerii, 64, 66, 68

striatus, 78
Pinnotheres pisum, 41
Pleurozyga phallusiae, 80
Podura villosa, 55
Pollicipes cornucopia, 44

polymerus, 47
Polydora agassizi, 25

ciliata, 25

socilis, 26
Polymnia nebulosa, 26, 29
Polyhabdina brasili, 27

polydorae, 25

pygospionis, 28

spionis, 27
Prisma tospora evansi, 71
Psocus sp., 72
Pterotrachea sp., 30, 31
Ptychoptera contaminata, 79
Pygospionis seticornis, 28
Pyxinoides balani, 35, 44, 47

cthamali, 35

Reduvius personatus, 57, 58
Rhyacophila sp., 66
Rhynchobolus americanus, 29

Sagitta sp., 24
Salpa aeruginosa, 81

confoederata, 81

maxima, 81

vagina, 81
Schneideria caudata, 76

coronata, 76

mucronata, 77

sp., 77
Sciadophora caudatus, 52

fissidens, 51

goronowitschi, 52

phalangii, 50, 53
Sciara nitidicoUis, 76
Scololepsis ciliata, 27

fuliginosa, 27
Scorpio indicus, 54
Sericostoma sp., 66, 67, 70
Siphonaptera, List of Gregarines found in, 79

104

ILLINOIS BIOLOGICAL MONOGRAPHS

[104

Sminthurus fuscus, 56
Spio martinensis, 27
Sporadina clavata, 60

reduvii, 58
Steinina rotundata, 79
Stylocephalus caudatus, 52
Stylocystis praecox, 78
Stylorhynchus balani, 35

caudatus, 52

heerii, 68

oligacanthus, 69

phallusiae, 80
Sycia inopinata, 28
Sympetrum rubicundulum, 71
Systenocerus caraboides, 75

Taeniocystis legeri, 30

mira, 78
Talitrus saltator, 42
Talorchestia longicornis, 43

Tanypus sp., 78
Termes flavipes, 56

lucifugus, 56
Thysanura, List of Gregarines found in, 54
Tinea pallescentella, 73
Tipula oleracea, 74, 75

sp., 74
Tramea lacerata, 71
Tunicata, List of Gregarines found in, 80

Uca pugilator, 43

pugnax, 43
Ulivina elliptica, 29

rhynchoboli, 29
Uradiophora cuenoti, 34

Vibrio reduvii, 58

Zygocystis clavata, 60
ephemerae, 62
pterotracheae, 31

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII April, 1922 No. 2

Editorial Committee

Stephen Alfred Forbes William Trelease

Henry Baldwin Ward

Published under the

Auspices of the Graduate School by

THE University of Illinois Press

COPYMGHT, 1922 BY THE UNIVERSITY OF lUJNOIS

Distributed August 5, 1922

THE MOLLUSCAN FAUNA OF THE

BIG VERMILION RIVER,

ILLINOIS

With Special Reference to its Modification as the Result of
Pollution by Sewage and Manufacturing Wastes

WITH FIFTEEN PLATES AND ELEVEN TABLES

BY

FRANK COLLINS BAKER

Curator of the Museum of Natural History
University of Illinois

Contributions from the

Museum of Natural History of the University of IIliDois

No. 22

TABLE OF CONTENTS

Introduction 7

Acknowledgments 8

Physical Features of the Big Vermilion Basin 10

General Biology of the Big Vermilion River 15

Plants 15

Worms 15

Crustacea 16

Aquatic Insects 16

Higher Vertebrates 18

Fish Fauna of the Big Vermilion River 18

General Distribution of the Mollusk Fauna 24

Ecological Variation 25

Comparisons with Other River Systems 28

Systematic Discussion of the Mollusca 31

Unionidae 31

Sphaeriidae 58

Pleuroceridae 59

Amnicolidae 60

Viviparidae 61

Ancylidae ... 63

Physidae 64

Planorbidae 64

Lymnaeidae 65

Pollution of the Salt Fork by Sewage and Manufacturing Wastes 67

General Nature of Stream Pollution 67

Sewage Pollution in the Salt Fork 72

Recent Examination of the Polluted Portion of Salt Fork 80

Summary of Salt Fork Conditions 88

Summary and Conclusions 90

Unfavorable Influences 90

Parasites and Pathologic Agencies 91

Economic Considerations 91

Bibliography 94

Explanation of Plates 97

105 ! FA UNA OF BIG VERMILION RI VER—BA KER

INTRODUCTION

The present paper embodies the result of investigations of the mollusk
fauna of one of the smaller rivers of the State of Illinois, the Big Vermilion,
carried on during the years 1918 to 1920. To this are added notes on
another river, the Sangamon, for comparison. While all groups of mol-
lusks were considered, special emphasis has been placed on the Unionidae
or pearly fresh water mussels (Naiades), on account of their abundance,
their significance in matters of geographic distribution, and their impor-
tance from an economic standpoint, as raw material from which pearl
buttons are made.

For a number of years, the United States Bureau of Fisheries has been
engaged in conducting a series of investigations of several rivers of Illinois
and Indiana, for the purpose of ascertaining the mussel resources of these
streams. In view of the rapid depletion of the supply in the larger rivers
(Mississippi, Ohio, Illinois) it becomes necessary to search the smaller
streams to replenish the older beds. The Maumee and the Kankakee
(Wilson and Clark, 1912), as well as the Illinois (Danglade, 1914), have
recently been rather carefully surveyed with interesting and important
results.

It was thought that a somewhat similar investigation of the Big Ver-
milion River would be of value. The scope of the investigation included
everything that appeared to affect the moUuscan life of the river, physical
characters, pollution, general relation and number of species. It was
believed, also, that the Big Vermilion, as well as other smaller streams
in the State, might provide good breeding stock with which to carry on
artificial glochidial infection of fish, and the results seem to warrant the
assumption. It will be necessary to make more or less detailed surveys of
all of the smaller rivers and their tributaries, and the present paper may
be considered a contribution toward this end, covering fully the Big Ver-
milion from its upper waters to the vicinity of Danville, a distance of about
forty-five miles by stream. A portion of the Sangamon is considered, and
this river may also prove a valuable source of mussel material.

Little has been done by field naturalists in the study of the distribution
of the molluscan fauna of a stream from the headwaters to the larger
portions of these rivers. Perhaps the most thorough and notable study
of this character was carried on by Adams (1900, 1915), on the genus lo
in which the Tennessee River and its tributaries (Powell, Clinch, Holston,
French Broad, Nolichucky, etc.) were studied from sources to Chattanooga.

8 ILLINOIS BIOLOGICAL MONOGRAPHS [106

Ortmann has made a study of the distribution of the Naiades in the
streams of Tennessee from this standpoint, using material collected by
Adams (Ortmann, 1918); and Wilson and Clark (1912) have added greatly
to our knowledge of the comparative distribution of the mussel faunas
of the Kankakee and Maumee rivers.

Studies of this kind bring out thfe fact that certain species are charac-
teristic of the upper reaches of a stream while others are found only in the
lower part. Barriers, such as falls and polluted water, are also seen to have
a large influence on the distribution of mollusks, especially river mussels.
The same species may also vary in size or shape in dififerent parts of the
stream, as noted by Ortmann (1920). In fact, a true picture of the charac-
teristics of the fauna of a river or smaller stream can be gained only by this
method, which the present paper clearly shows.

The Salt Fork of the Big Vermilion is a striking example of the ill
effect of sewage and other pollution on the fauna of a stream. It was
carefully studied from this standpoint in order to determine accurately
the distance that the polluted stream must flow before a normal fauna can
establish itself, and to ascertain the relative effect of pollution on different
species of mussels as well as on other aquatic animals. Mussels and cray-
fishes are considered good indicators of the measure of pollution in a stream
and the absence of both of these groups of animals from the upper part of
the Salt Fork is ample evidence of the septic condition of this stream.

The work was largely carried on during the months of August, Septem-
ber, and October, when the water was low. Collecting was done by wading
in the stream as deep as hip rubber boots would permit. In the shallow
water (up to two feet in depth) the entire bottom was gone over with the
hands, and the mussels and other mollusks thus picked from the bottom
of mud, gravel, and sand. Samples of all mollusks, as well as associated
a;nimals of the other groups, have been preserved in the Natural History
Museum of the University of Illinois.

The different areas of the region studied are covered by three maps of
the United States Geological Survey, the Mahomet, Champaign, and Dan-
ville FoUos, and on these the stations recorded in this paper may easily
be located.

ACKNOWLEDGMENTS

The writer is greatly indebted to Professor Frank Smith, not only for
invaluable assistance in collecting much of the material upon which the
paper is based, but also for many notes on the distribution of the Naiades
and other mollusks in the two river systems. Professor Smith has studied
the fauna of the Salt Fork and the Sangamon streams for many years,
collecting from them at different times of the year. His classes in zoology
have visited Homer Park yearly for a long period and thus the moUusk

1

107] FAUNA OF BIG VERMILION RJVER— BAKER 9 [

fauna of this locality is better known than that of any other part of the
stream.

To Professor Smith the writer is also indebted for the identification of
the worms and crayfish, and the determination of the animal life in the
samples of bottom sludge from the polluted part of Salt Fork. The sincere
thanks of the writer are due to the persons noted below; the group of ani-
mals or plants which they have identified is indicated: Dr. Charles P.
Alexander, Coleoptera and other insects; Dr. Edward Bartow; Dr. Stephen
A. Forbes; Mr. Calvin Goodrich, Goniobasis; Dr. George M. Higgins; Mr.
John Malloch, Diptera; Dr. J. Percy Moore, Hirudinea; Professor James E.
Smith; Dr. E. N. Transeau, Algae; Dr. Victor Sterki, Sphaeriidae; Dr.
Harley J. Van Cleave; Dr. Bryant Walker, Ancylidae and other moUusks.

10 ILLINOIS BIOLOGICAL MONOGRAPHS [108

PHYSICAL FEATURES OF THE BIG VERMILION BASIN

The Big Vermilion River drains about 1,500 square miles in Champaign,
Ford, and Vermilion counties in Illinois and a small portion of Warren and
Fountain counties, Indiana. The North Fork also drains from a small
territory in the southeastern part of Iroquois County, Illinois. The main
stream, known as Middle Fork, rises in the southern part of Ford County
near the town of Melvin, in the Bloomington morainic system, at a height
of 800 feet above sea level. Its course is southeastward, between the hills
of the moraine known as the Roberts and Melvin ridges, passing through
the latter and uniting with a tributary known as the West Branch of the
Middle Fork, which also rises at an elevation of 800 feet in the Roberts
ridge. At Potomac, the stream turns southward, cuts through the outer
ridge of the Bloomington moraine and crosses the plain of the Champlain
till sheet, uniting with the Salt Fork about six miles west of Danville.

The largest western tributary, known as the Salt Fork, rises in the till
plain in the north-central part of Champaign County, near Thomasboro,
at an elevation of about 740 feet above the sea. It drains the till plain
lying between the Bloomington moraine on the north and the Champaign
moraine on the south. It flows in a south and east direction for about
55 miles^ and unites with the Middle Fork as described above. A large
tributary of Salt Fork, known as Spoon River^ rises in the northeastern
part of Champaign County, in two branches, not far from the outer ridge
of the Bloomington moraine. Its general course is southward for a dis-
tance of about ten miles, where it unites with the Salt Fork near St. Joseph.

Another large tributary is known as the North Fork, which rises in the
southeafetern corner of Iroquois County in the inner ridge of the Blooming-
ton moraine. It flows southward, cutting through the middle and outer
ridges of the moraine, crosses a part of the Champaign till plain and unites
with the Big Vermilion at Danville. This tributary has a length of about
40 miles. From Danville the larger stream flows southeastward for about
20 miles, crossing a part of Vermilion County, Indiana, and empties into
the Wabash River 10 miles from the Illinois State line.

The basin of the Big Vermilion River lies in or is surrounded by glacial
moraines of the Early Wisconsin glaciation, the Bloomington moraine on

1 Length of rivers designates total length including all meanders.

* Not to be confounded with Spoon River entering the Illinois River near Havana, Mason
County.

1091 FAUNA OF BIG VERMILION RIVER— BAKER 11

the north and the Champaign moraine on the south. In its course it cuts
through the Bloomington moraine in several places. The territory drained
includes a small part of the Bloomington till plain and a considerable part
of the Champaign till plain (Leverett, Illinois Glacial Lobe, plate VI.)
As there are no outcroppings of rock in this area the streams have cut well
defined, though meandering, channels. The upper, small, creek-like tribu-
taries have sunk their beds but a few feet below the general level of the
country, but lower down, the stream, after receiving several large tributa-
ries, has cut its bed to a depth of ten or fifteen feet. In Middle Fork and
North Fork, and in the Big Vermilion from above Middle Fork to the
Wabash, the river has cut deep canyons upwards of 200 feet in depth,
which produce some of the most picturesque scenery in the State of Illinois.
This river valley varies from half a mile to a mile in width and in one place,
about four miles below Danville, it widens to form a large amphitheater
two miles wide and a mile long, with cliffs and hills rising on all sides to a
height of over 150 feet.

Outside of the stream valleys the country is a flat till plain, largely
devoted to crop purposes. Ridges, made up of the Champaign and Bloom-
ington moraines and their branches, occur and have been largely instru-
mental in directing the course of some of the stream drainage. The Salt
Fork turns to the north after leaving Urbana, and passes around Yankee
ridge (a branch of the Champaign moraine) near the Brownfield woods,
and spurs from the Champaign moraine occur in several places on the west
and south banks of the Salt Fork, which are relatively high and the stream
skirts their bases.

The bottom of the main stream and its tributaries varies greatly.
The small tributaries mostly have mud bottoms. In Spoon River the
bottom is of mud in most places and the water is of considerable depth,
even in summer (three to four feet maximum in August). In places
there are riffles where the water is very shallow (a foot or less) and the bot-
tom here is of sand and fine gravel. The Salt Fork below St. Joseph is
made up of stretches of stream where riffles with sand and gravel bottom
alternate with deeper back water with mud bottom. Near Muncie there
are outcrops of rock, a small tributary. Stony Creek, flowing over a stony
bed. In the Middle Fork, shale rock outcrops in several places, notably
below the interurban bridge, where the whole bottom is composed of a
sheet of rock with a thin coating of sediment in spots. The same conditions
are found in patts of the North Fork and in the Big Vermilion. These
varying conditions provide the most favorable environment for the growth
of river mussels, a fact made evident by the large naiad fauna found in
the stream despite the unfavorable effects of sewage and waste pollution.

Natural dams occur in a few places in the Salt Fork, caused by the
accumulation of debris which has lodged against the trunk of a tree that

12 ILLINOIS BIOLOGICAL MONOGRAPHS [110

has fallen across the stream. At Homer Park, an artificial dam about five
feet in height causes slack water for several miles up the stream. This dam
markedly affects the mollusk fauna, the tumbling of the water over the
dam mixing air with the polluted water and providing the dissolved oxygen
so necessary to naiad life. It is probable that the large number of species
of mussels found at some stations above this dam is due largely to the
presence of sections of the stream where rififles provide the oxygenating
agent. Mussels were usually found in or near such habitats. The mussel
fauna below the Homer Park dam numbers 28 species while above the dam,
as far up stream as Sidney, only 17 species occur, 10 species not passing
the barrier, although the environment does not differ essentially. The
current in the river is rapid over the riffles but rather sluggish in the deeper
places. The difference between high and low water (spring and fall) is
about six feet. The streams usually vary in width from ten to thirty feet.

In the late summer and fall the small tributary streams (creeks and
rivulets) flowing into Salt Fork and other branches of the Big Vermilion
are usually either dry or contain scattered pools of water throughout their
length. They do not contribute any water, therefore, to the larger stream
at this time of the year. The mollusks living in these tributaries bury
themselves in the mud during this period of dry bottom and hibernate.
Many die at this time.

The banks of the stream valleys, exclusive of the small tributaries, are
for the most part high and well wooded especially where the valley floor
is wide enough to permit meandering, in which cases the flat floodplains
are abundantly wooded. These flat areas vary from a few hundred feet
to a half mile in width. About two miles above Sidney an island has been
formed by the forking of the stream, the area embraced being about 650
by 1200 feet. At this place the right bank is 20 feet high and the left bank
quite low. The presence of fossil shells indicates that the island was prob-
ably the result of silt accumulation during a long period of time. The
wooded banks of the stream alternate with farm lands, some in pasture
and others in crops. Many of the crop lands have a fringe of timber border-
ing the stream. That the stream is high and powerful during the spring
when it is in flood is evidenced by the tangled mass of logs and other woody
debris which thickly cover the flood plain areas along the valley. Such
conditions were especially noted between Sidney and Homer Park.

The current varies somewhat, being relatively sluggish in the backwater
above dams and riffles, but quite swift over the shallow places. During the
flood periods of spring and early summer the current is quite swift and in
places becomes torrential. This condition is indicated by the large number
of trees which have been thrown on the flood plains far above the margin
of low water. As measured during the month of October the current in
the Salt Fork at Urbana and a few miles down the stream had a velocity

Ill] FAUNA OF BIG VERMILION RIVER— BAKER 13

of from half a mile to a mile an hour. The water was very low when these
measurements were taken by the State Water Survey. In its course of 90
miles to the Wabash River, the Big Vermilion River falls about 320 feet or
ZV2 feet per mile. It thus has a greater fall than either the Kankakee,
which is 300 miles long and has a fall of 250 feet, or less than a foot per mile,
or the Maumee River, which has a length of 150 miles and a fall of 154 feet,
or about a foot per mile (Clark and Wilson, 1912).

The water in the normal parts of the stream is usually clear at depths of
one to two and a half feet, especially on the riffles. This condition was
noted in Salt Fork east of Sidney, and in the Middle Fork. Above Sidney,
except where the water is very shallow, the stream is murky and laden with
fine silt. During times of high water the stream is in this condition in all
parts of the Big Vermilion. The upper part of Salt Fork, from Urbana to a
point six or eight miles down stream is always more or less brownish in
color from the large amount of sewage, equalling as much as a third of the
total volume, and putrient matter as well as an oily scum is usually to be
seen on the surface. The shore of the stream is rendered very unsightly by
the mass of filth that is despoiled above the usual level by high water. At
St. Joseph, ten miles from Urbana, much the same condition is found.

The upper part of the Salt Fork has been greatly modified by ditching
and dredging. North of Urbana for the distance of a mile and a quarter
above Crystal Lake Park a large ditch carries the surface drainage in a
straight line to the park thus cutting off the tortuous windings of the
original stream bed, which have been left as long, narrow, shallow ponds,
reminding one of the 'ox-bows' so common in the valley of the Mississippi
River. The bed of this ditch is about six feet below the general level of
the surface. For several miles above this ditch the original stream has been
deepened by dredging and the bed is now pretty generally five or six feet
lower than that of the original stream.

From Crystal Lake Park, Urbana, to a point near St. Joseph, the stream
has been ditched to straighten the bed, leaving numerous 'cut-offs' of the
old stream bed. This canal permits a better flow of water for the disposal
of the sewage. Where not ditched the stream bed has been deepened.
The ditching has greatly modified the original stream bed, providing a new
and different kind of environment for the mussels and other aquatic life.
It is probable that all of the old fauna was exterminated during the ditching
operations and the sewage pollution provides an unfavorable environment,
which the aquatic bottom life does not seem able or inclined to enter. The
effect of sewage pollution may be seen all the way down the stream from the
source of contamination. At St. Joseph, where the stream bed has not been
modified, conditions are very bad, the mud in the bottom being filled with
gas forming bacteria which are constantly causing bubbles of gas to break
at the surface of the water. Were it not for the sewage pollution, the stream

14 ILLINOIS BIOLOGICAL MONOGRAPHS [112

at this place would be quite normal for the life of mussels and other bottom
animals. (See the chapter on sewage pollution.)

Several other river systems have their beginning near Urbana and
Champaign, on the southwest side of the Champaign moraine. These are
the Embarras, which has tributaries rising south of Urbana; the Kaskaskia,
which has its inception northwest of Champaign near the village of Rising;
the Little Vermilion, which rises in the southeastern part of Champaign
County; and the Sangamon, which rises in the extreme northwestern part
of Champaign County not far from Gibson. Only the last stream has been
examined for its mussel fauna for the purpose of making comparisons with
the fauna of the Big Vermilion River.

The Sangamon River has a length of about 180 miles and a drainage
basin of some 5670 square miles. It rises on the south side of the Blooming-
ton moraine in McLean County, "at an elevation of 850 feet above sea
level, and cuts through two moraines in its course, the Champaign and the
Shelbyville. The river channel is tortuous and meandering and the flood
plain in many places very wide. The banks of the stream are low for the
most part and wooded in spots. The stream has been examined carefully
at only two points, Mahomet and west of White Heath, the first locality
receiving the most attention. Near Mahomet the river is notably mean-
dering, the banks are high, the Champaign moraine rising 90 feet above
the water level near the village. The river bank is usually five or six feet
above the stream, which has cut vertical clifif-like banks in many places.
Below Mahomet the woodlands are abundant, extending well back from
the river in some places. The river varies in width from 40 to 50 feet.
The bottom alternates between riffles with sand and gravel bottom and
deeper back water stretches with mud bottom. The former habitats are
a foot or so in depth and the latter habitats two to four feet in depth in the
summer. In the spring the river is ten to twelve feet deep and very swift,
at times becoming torrential. The average fall of the stream is 2.3 feet
per mile. Many logs and other debris thrown up on the flood plains attest
the power of the river during spring floods.

West of White Heath, for a mile above the I. C. railroad bridge to
two miles below, the river has been examined, though no systematic
collecting comparable to that carried on at Mahomet has been done. The
stream in the portion of the river valley examined is similar to that near
Mahomet in its general physiographic features. The banks of the stream
are on the whole lower than farther up the river near Mahomet. Studies
of this stream similar to those carried on in the Big Vermilion and its
tributaries would doubtless yield interesting and valuable results. The
Sangamon is a characteristic mussel stream and should contain a much
larger mussel fauna than at present known and listed.

113] FAUNA OF BIG VERMILION RIVER— BAKER IS

GENERAL BIOLOGY OF THE BIG VERMILION RIVER

Although the chief purpose of the study of this stream was to ascertain
the general conditions and distribution of the fresh water Mollusca,
especially the Naiades or river mussels, such attention was given to other
groups as came easily under observation. At least one of these groups
bears an intimate relation to the Naiades in the matter of distribution
as restricted by pollution, e.g., the crayfishes, and another, the fish, is
closely related to the breeding habits of the mussels. Others, as some of
the oligochaete worms, are especially characteristic of polluted waters.
A few notes are given under each group.

The Protozoa and other microscopic groups are omitted because given
little or no attention, except in the polluted part of Salt Fork. These
are mentioned in the section on sewage pollution. The Big Vermilion and
its tributaries present a wide field for the study of other kinds of aquatic
life that are especially characteristic of the upper reaches of a river system

PLANTS

The larger aquatic plants were notably scarce in most parts of the
Salt Fork. In several places in Salt Fork, especially near Sidney, between
the cement and railroad bridges, the shores are lined with the spatterdock,
Nymphaea advena Ait (see Fig. 11). The cat-tail, Typha latifolia, Linn.,
is common in various stretches of the stream, bordering the shore. The
arrow-head, Sagittaria latifolia Willd., and the larger blue flag. Iris versi-
color, Linn., were observed along the shore in many places. The water
willow, Dianthera americana Linn., is abundant in the shallows in many
parts of the Salt Fork below Homer Park. Elodea canadensis Michx.,
was abundant at Homer Park.

Filamentous algae occurs in many places, attached to submerged
objects. Among these Cladophora and Spirogyra were noted. Septic
algae growing in the polluted portion of Salt Fork are noted under the
section on pollution of the stream.

WORMS

Nematoda. Gordius rohustus Leidy, was collected in several places in
Salt Fork between the first bridge below St. Joseph and the bridge above
Sidney. It was found in mud bordering the shore, in shallow water. A
number of minute nematodes were observed in the sludges of the bottom
in Salt Fork between Urbana and St. Joseph. These were not identified.

16 ILLINOIS BIOLOGICAL MONOGRAPHS [114

Oligochaeta. These aquatic earthworms were abundant in places
along the shore of Salt Fork, Sparganophilus eiseni Smith occurred in
abundance in the mud on the margin of Salt Fork at Homer Park, and
immature worm of this genus, as well as cocoons, were collected from near
the bridge below St. Joseph to the bridge above Sidney. A single specimen
of Helodrilus chloroticus (Savigny) was collected in the Salt Fork near St.
Joseph. This species has not heretofore been recorded from Illinois and
its occurrence at this locality adds another species of earthworm to the
State list (see Smith, 1915:557). Limnodrilus was abundant in bottom
sludges from the Salt Fork, from Urbana to near St. Joseph. Tubifex was
also found, but not as abundantly.

Hirudinea. Two species of leeches occurred in several places in Salt
Fork, below St. Joseph. They were nowhere abundant. The two species
are: Erpobdella punctata (Leidy) Moore and Placobdella rugosa (Verrill)
Moore.

CRUSTACEA

Three species of crayfish (Cambarus) were determined by Professor
Smith from the material collected in Salt Fork and other parts of the Big
Vermilion River. The most abundant was Cambarus propinquus Girard,
which was found commonly from bench mark 655 to the Big Vermilion
River below Middle Fork. It is especially abundant in the latter place
where an individual may be found under every piece of stone. The same
species was abundant in Stony Creek and in the Salt Fork near Muncie.

Cambarus blandingi acutus Girard, both young and adult, occurred in
the upper parts of Salt Fork, but preferred a mud bottom rather than a
rock bottom. Cambarus immunis Hagen was collected only at bench mark
655 where it was rare.

It will be noted that Cambarus is first met with about two miles below
St. Joseph and twelve miles from Urbana. The only ones seen here were
dead. Living crayfish begin to appear in abundance about six miles below
St. Joseph or sixteen miles below Urbana. Crayfish and mussels both be-
come common or abundant at about the same time, viz., fourteen and
sixteen miles below the source of sewage pollution at Urbana. This agree-
ment in distribution indicates the close relationship between these other-
wise diverse groups of animals, as regards resistence to septic conditions,
neither being able to thrive under pollutional conditions of the bottom.

AQUATIC INSECTS

Aquatic insects are doubtless abundant during spring and early summer
in the lower parts of Salt Fork and in the Big Vermilion River. A few
species were collected incidentally at some of the molluscan stations.
These are listed in Table I, in the order of their occurrence in the stream.

115]

FAUNA OF BIG VERMILION RIVER— BAKER

17

It will be noted that the nymph and larval forms of most species appear
coincident with the degree of sewage pollution. Chironomus and Libellula
are, seemingly, able to accommodate themselves to the contaminated or
polluted condition of the water in the neighborhood of St. Joseph. The
adult beetles, breathers of free air, are not aflFected directly by these un-
favorable conditions and occur in great abundance, even in the heavUy
polluted parts of Salt Fork above St. Joseph. The Ephemerids occur for
the most part far down the stream where the water is at most only con-
taminated. The Neuroptera and Plecoptera are clean water forms and
were found only in the Middle Fork which does not carry sewage. These
are also to be found, probably, in the Big Vermilion below Middle Fork,
but no attempt was made to discover these animals when that part of the
stream was examined.

Table I. Distribution of Insects

•

T3

lU

M
>

o

D

o
>—>

Bridge below
St. Joseph

IS

a

x:
<j
c

Mile above iron bridge
near Sidney

4»
.£3

a
o

1— (

-a

Diptera
Chironomus decorus Joh., larva

X
X

X

X

X
X
X
X
X

X

X

Odonata

Libellula pulchella Drury, nymph

Coleoptera

Gyrinus analis Say, adult

Dineules assimilis Aube, adult

Cnemidotus 12-punctatus Say, adult

X

Laccophilus maculosus Say, adult

Copelatus glyphicus Say, adult

Hemiptera

Corixa, nymph

X
X

X

Diptera

Cricolopus Irifasciatus ¥., larva (?)

Ephemerida

Hexagenia bilineata Say, nymph

X

Heptagenia, nymph

X

Trichoptera, larva

X

Odonata

Ophiogomphus, nymph

X

Neuroptera

Corydalis, larva

X

Plecoptera

Perla, nymph

X

18 ILLINOIS BIOLOGICAL MONOGRAPHS [116

HIGHER VERTEBRATES

Vertebrates characteristic of aquatic environments were fairly common
in most parts of the Big Vermilion River valley. Three species of turtles
were observed: the western painted terrapin (Chrysemys cinerea Bonna-
terre), the snapping turtle (Chelydra serpentina Linn.) and the soft shelled
turtle (Platypeltis spinifera LeSusur). The garter snake (Thamnophis
sirtalis Linn.) was observed swimming across the stream in several places.
The fox snake {Elaphe vulpinus Baird and Girard) was seen on several
occasions near the margin of Salt Fork above St. Joseph. Frogs, among
which the pickerel frog {Rana palustris LeConte) was noted, were abun-
dant in many places, and tadpoles of all ages were abundant in both the
Big Vermilion and Sangamon rivers.

Aquatic birds were occasionally seen in both river valleys. The little
green heron {Ardea virescens Linn.) and the great blue heron {Ardea hero-
dias Linn.), as well as the American bittern {Botaurus lentiginosus Mon-
tague) were seen repeatedly, especially in the old river cut-offs between
Urbana and St. Joseph. Where high banks occur the kingfisher {Ceryle
alcyon Linn.) made the woods resound with its rattle-like notes. These
birds feed largely on young fish and in this way affect the mussel distribu-
tion by reducing the number of fish that may bear glochidia. Shore birds,
among which were noted Wilson's snipe {Gallinago delicata Ord.), solitary
sandpiper {Helodromus soUtarius Wilson), yellowlegs (Totanus), sora rail
(Forzana Carolina Linn.), and kill deer (Oxyechus vocijerus Linn.). The
pied-billed grebe (Podilymbus podiceps Linn.) is seen frequentJy during
spring and autumn on Crystal Lake and in the more pond-like reaches of
the Salt Fork. Many of the shore birds feed on mollusks and insects which
they find along the shores of the Big Vermilion River system. During
migrations, the river valley and its tributaries are fairly alive with birds of
all kinds, and at such times the region is well adapted for bird study.

The presence of the muskrat {Ondatra zibethica Linn.) is attested by
the number of piles of opened mussel shells, the animals of which have
provided this mammal with many a meal. In these muskrat piles have
been found the shells of many species that are rare or difl&cult to find alive
in the streams.

The abundance of all groups of animal life in the Big Vermilion system
indicates that it is a favorable environment for an optimum biota. With
the exception of the upper twenty miles more or less seriously affected by
sewage pollution, the stream is one of the best collecting grounds in the
State, a condition indicated by the very large mussel fauna of thirty-eight
species and races, recorded in the following pages.

FISH FAUNA OF THE BIG VERMILION RIVER

The close relationship between the mussel fauna and the fish fauna,
through the breeding habits of the former, render a knowledge of the fish

117] FAUNA OF BIG VERMILION RIVER— BAKER 19

fauna of the Big Vermilion necessary. As a large mussel fauna lives in the
stream it follows as a corollary that there must also be a fish fauna of com-
parable size. Fortunately, Dr. S. A. Forbes, caused extensive collections
to be made in 1901, and these are listed on the maps accompanying the
work on the Fishes of Illinois, by Forbes and Richardson (1908). Fifty
species are recorded from the Big Vermilion River and its tributaries (see
Table II). It will be seen that at this date 28 species were found below
Urbana, in that portion of the stream now heavily polluted.

Between the years 1908 and 1912, the Salt Fork between Urbana and
St. Joseph was deepened and straightened and the bottom fauna was com-
pletely destroyed. In the canal thus formed, for the purpose of carrying
ofif the sewage of the Twin-Cities, no living clams, crayfish, or other clean
water animals have been found, and but one school of young fish (bull-
heads) was observed during several examinations of this part of the stream.
It is possible that during periods of high water in the spring, some hardy
fish may venture into this heavily polluted area. Below St. Joseph it is
quite probable that some fish are found during high water periods. A
few minnows were observed near the station called bench mark 655, about
fifteen miles below Urbana. A farm boy reported that bullheads could be
caught at high water on set lines.

Below bench mark 655, and for some distance above it, fish must resort
in some numbers because of the presence of a fair sized mussel fauna (see
Table III). Young mussels, however, were not seen in any number above
the Homer Park dam, and it is possible that the upper stream is now little
visited by fish suitable for glochidial infection. Below the dam, young
mussels are plentiful at all points examined.

Information concerning the species of fish that carry glochidia of the
river mussels is still of a fragmentary character. Suber (1912), Howard
(1914), and other workers of the U. S. Bureau of Fisheries have published
considerable data on this subject, but much more is needed before one
can fully understand the relation between mussels and fish.

Three species of mussels living in Salt Fork and other parts of the
Big Vermilion River are known to have glochidia encysted on five species
of fish, all of which have been reported from the Big Vermilion, viz.:

Mussels Fish

Lampsilis anodontoides Lepomis humilis

" anodontoides Pomoxis sparoides

" anodontoides " annularis

Quadrula metanevra Lepomis pallidus

" pusiulosa Pomoxis annularis

" pustulosa Ictalurus punctatus

20

ILLINOIS BIOLOGICAL MONOGRAPHS

[118

The abundance of desirable species of mussels in the Big Vermilion
indicates that it may serve as a reservoir for button material, the species
being easily transported to laboratories anywhere in the state for artificial
infection of fish.

Table II. Distribution of the Fish Fauna in the Big Vermilion

Car piodes diffiformis Cope. Blunt-nosed silver carp

Carpiodcs veli/er (Raf .). Quillback; Silver carp

Erimyzon sttcetta oblongus {MitchiH) . Chub-sucker

Minytretna melanops (Raf.). Spotted Sucker

Catostomus commersonii (Lac). Common Sucker

Catoslomus nigricans (Le Sueur). Hog sucker

Moxostoma aureolum (LeSueur). Common red-horse

M 0X0 stoma breviceps {Copt). Short-headed red-horse . . . .

Campostoma anomalum (Raf.). Stone-roller

Hybognathm nucalis Agassiz. Silvery minnow

Pimephales notatus (Raf.). Blunt- nosed minnow

Semotilus atromactdatus (Mitchill). Horned-dace

Abramis crysoleucas (Mitchill). Golden shiner

Cliola vigilax (B. & G.). Bullhead minnow

Notropis cayuga Meek. Cayuga minnow

Notropis blennius (Girard). Straw-colored minnow

Notropis Ulecebrosus (Girard). Minnow

Notropis whipplii (Girard). Steel-colored minnow

Notropis cornulus (Mitchill). Common shiner

Notropis atherinoides Raf. Shiner

Notropis umbratilis atripes (Jordan) . Blackfin

Ericymba buccala Cope. Silver-mouthed minnow

Phenacobius mirabilis (Girard). Sucker-mouthed minnow

Hybopsis amblops (Raf.). Big-eyed chub

Hybopsis storerianus (Kirtland). Storer'schub

Hybopsis kentuckiensis (Raf.). River chub

lotalurus punctatus (Raf.). Channel-cat

Ameiurus natalis (LeSueur). Yellow bullhead

Ameiurus melas (Raf.). Black bullhead

Noturus flavus Raf. Stonecat

Schilbeodes gyrinus (Mitchill). Tadpole cat

Schilbeodes murus (Jordan). Brindled stonecat

Esox vermicularis Le Sueur. Little pickerel

Fundtdus notatus (Raf.). Top minnow

Labidesthes sicculus (Cope). Brook silversides

>

a
o
o
a
in

>

2

a
o
o
a

0)

a

.'S
'in

■a

o

1^

o

119]

FAUNA OF BIG VERMILION RIVER— BAKER

21

Table II — (continued)

C

-e
>

o

<

a

m

4)
>

a
o
o

a.

>

5

a

8

a-

1)

pq

c

nJ

§

u
C

3

O

o

a
Q

o
o

Pomoxis annularis Raf . White crappie

X
X
X
X

Pomoxis sparoides (Lacepede). Black crappie

X
X

X

X
X
X
X

X
X
X

X

X

X

X

X
X

X

Lepomis megalotis (Raf.). Long-eared sunfish

X

X

Lepomis humilis (Girard). Orange-spotted sunfish

X

Lepomis pallidus (MitchUl). Blue gill

Micropterus dolomieu Lacepede. Small-mouthed black
bass

X

X

X

Micropiertis salmoides (Lac). Large-mouthed black bass.
Percina caprodes (Raf.). Log-perch

X

X
X
X

X

X

X
X

X

X
X

X

Hadropterus pkoxocephalus (Nelson)

Hadropterus aspro (C. & J.). Black-sided darter

X

X
X
X

X

X
X
X

X
X
X

X

X
X

X
X

X
X

X

Diplesion blennoides (Raf.). Green-sided darter

X

Boleosoma nigrum (Raf.). Johnny darter

X

X

Elheosloma jessiae (Jordan & Bray ton)

X

Etkeosloma coeruleum Storer. Rainbow darter

X
X

X
X

X

X
X

X

X
X

27

X

Etheostomaflabellare Raf. Fan-taUed darter

X

Total SDecies from each station

,19

34

42

28

30

27

21

?6

>

z
o

g

PK

H
>

O

o

g

o

S5
O

&

0

n

ailtAn^d JB8N "^ : :

Q

►i

00 •

O boO

j3Aiy noiiinija^ atg n m->

Q

08— • C8 d 00 t.

-

.J

e.

c

: *

&

-

a

c

C

c

b

c

0
CM

05 61)<M
3(J0,^ 3|PP!IAf =^"-

C

o

o> u OS d u

CN

o— *

08

cs

&

cs

V

' ^"

d 08 —

«-H wm

<o
<o

OO bOO

noijonnr jjjo j •Ji«s « «i -h

Q

CS OS e) 09 C8>-

Cs

• »-«
d^^

• 09

*"

^ ^^

C-j

U

^^

CO
CM

t- go

5[38J3 XnOIC! =~^ CO -

C

«

■-- : ; :

•-

«

-

0

CO

to txio
apanjY jo q^nog cm m —

Q

6 •

«■- . d 08 t

-

d e.

' OS

c

-

c

-

-

• t

•-

o<

0

CO

0

CM

CO

m«p jaraojj Mopg '^ " " | ^q

u

d d oj OS 03 d

-

-

d— • t

09

•-

^^ t>"-

0
1^

mcp jsroou Mopg cm o cm

O
h3°

OS

oj OS OS e« 08 d'-

e> o

d d d 08 u

-

es

—

J..

-

— ; ti-'—

0

00
CM

1 c^

msp janiojj aAoqc saiini z c-i -^

•-

es o 08— rtr ■

• i-<

w- *-^ d

CO

a

•-

d d os-i 08'-

.-;.-;

o— « o e>

0

;i?=M

CO
CM

mcp jamoH SAoqB 63|ini f cm «

Q

d • 08— • d •

;""

.-H .-« • ■

CD

t*r-c^

CM

CM

JO -g '93puq pBOJiiB^ ^ '-

*"*

•CM • o ■ —
.« .a OS ■

^

eq

CM CM

CM
CO

OCM
CM

'fa"P!S ogo
JO -g '33puq ^naraag cm a =.,

3«^

d

— ••- • 08-H--

0

at

;i?c.

iCanpig JO 'N 'aSpijq aojj -^ '-'

•-i d d 09

t-H

^H

>o

CO bflOO

83puq nojt jo qijon ajij^ -h m -h

Q :

■•-

•CM • • •

d • oSrt' 03 —

0<-tCM •

• • ■ 08

00

2"

.«

• cq . . . .

■'

(N

^

00
to

»o

to ao
piraq 3iq uo|3g — « cm

Q
►J

IN

CM • -coco ■

•"■

10

1-1

;?;«

^

pnaq Sig '-« o _

Q

CM

^

! *"^ '.'^

I—

'"'

to

::?«

■*

2 92

a
►J

• -eo •cc

CM

•^ ^

CO a o
qdasop 'jg M0[3q aSpug — Qci

Q

o^^ ueo ed

to

CMCO

qdasof •■jg — n -<

Q

00

— ' aoo
83itm 2 J8Aiy noodg -h o «

Q

-

• '^N *-« ■
•-« . • ot

CM

Or).

o aoo
ajira I J3At'}j noodg -< n «

Q

-

U

d— i d« d —

"■

wm

^^

OCO

05 bCCM

6 aapijq ':(joj ■Ji^g

Q

00 to

00 M<M

Q

CO

f aapijq -^jjioj 5IBS '^ "-

1 :

■*>o

Z 93prjq '3IJ0J ^iBg '° "2

Q

CM

T aaptjq *3iJ0j ij^g "^ S'SJ

Q

►J

a

u

OQ

og

•^

CO

CO

CO fl to

peq niBajjs pjo ° "

■^

CO

0

„ cfl a CM

2

o

«

0

3(jog }|Bg J9ddfi '^ a «

Q

►J"

u

c»

CO

•0

Symbols
a, abundant
c, common
i, infrequent
1, 2, 3, specimens found
L, living
D, dead
(/, gravel
m, mud
«, sand

Stati on number

i

a

c
03

03

J
1

c

X

c
Q

S

C3

P.

"a

K
3

00

3

1

OO

3

5

3

R
O

1
1

1

c

_«o

■3

1

1

.1

Si

a

c

■§

■a

2

00

3
S

■3

S

00

E

s

LampHlia luteola

Eurynia lienosa

Lasmigona complanata

Fusconaia rubiginoaa

Amblema undulata

Quadrula vuatuloaa

09

c

■1

3

5

1
i

3

1

1

09

_3

e

3

1
1

1

5

§

ft:

8

o

s

.1

d

a

s

:§
.3

5

3

R
■§

8
0

S
§
a.

8

R
_ao

s

0

^00

1

•0

s

■§

0

i

E
8

g

■1

i

00

1

1
§

_00

E
0

"S

3
g

00
E

,0{

S

i

§
s

R
0
R

■■s

0

.5

E

i

s

St

■*«»

1
1

e
a
"a

0

3

tj

K

e
3

1

s

c
"3

03
3

0

3
g

3

03

c

■§

R
3

1

1

1

.0
S

1

.0
J

i

■2

<u

C

•n
3

1

0

e:

s

-i

R

1
1

•2

R
3)

1

t3
.§

2

1

1
s

R
3

w

6-1

■§

a,
to

OS

a

1

1

a

1
"i

a
a

a:

S

1

t»

0

a
>
'So

1

>

z
o

M

>-)

a

»
>

a
pq

H

Z

M
CO

I-)
•«:

a

Z
<

Q
M

<

O
O

C.T

m

•<

Q
Z
<

t/T

Q

o
0.
o

0$
H
en
<

a

<

a

OS

u
<

o
o

t-H

H
S

n
1-^

M
H
(/}

n

aO!itmjB;\ aig ^ ^^ q : :

. . 0 .

. y • .

;•■■■•■• OS

■* :!;;:!:;;

:::::::""::::

jjjo^ 3[PP!IM

OS &0(M

1 Q : :
1 iJ : -■

lid'.!!*.

0 Oi-« '.

uopoant 3(ioj jjBg

00 bCO

Q :

jpajQAuo^s ^ ^o

Q :

J '■

: :- :

-;

:::::::*:::;

'■ '• <i '■

aiounjv JO qjnog j^a mco

1 CM 01

1

1 Q : :
1 hj : :

;;;;;■•• os'

CM

: 1 : ! 1 ! ! a : : : :

:(JEJ jaraog

•0 ea 00

Q :

::::*::::

»-«

(4iopq) ra^p jamajj

•0 bflOO

a :

-- i

.'.'.'.'.'.'.'. a

'.'.'.'.'. a

•^

: ta 1 : : : : : : 1 ; I

niBp
jamou BAoqc 8a|im z

^-'^

Q :

rasp im M=M
jaraog aAoqB saftm z/jg ]« «> -^

Q :
h-) : :

• • ■ -«....

«-t

jaraojj BAoqc saifoi ^ ]" '^

Q :

J :

. . . ! ; 0 ■ - '

-■

aSpuq psoijiBjj

?;"2

! ^^

;;;;«••••

CM

'. d '.','.'.'.'. '.

:::!:"'•'*''!:::

aSpuq ■juamaQ

gas 2 :^

.'.'.'.'. ^ '.'. '.

. .^^ '. '. '.

CO

'.'.'.'.'.'.'. 6 - '■ 6^

aSpijq uojj

2 "■SS

Q :
►J :

• -co • (M

*"^ c8 ca ca

] '■ J — ' OS « \

0

CM

0

'. '. I '. \ '. '. \ '. . 0 '.

a3puq uoat q:)jon a[tai j

00 gO

2;=^

: : :-^

. . .

: : :---' : :'

"::;::

' 0

Or^ '• '• '• '•■^ii 0 ■« •

509 jjj^m qauag

^ a2

Q : :

: : ':■■'

- 0 0

• -o • • • •

CM

. . . .OUC^^H*-^ • • •

puaqSig U q : :

1-^- ^ : :

. • -co • --H •

::::*:::

CM

mep |Bjn}B(^ Mojag

2 as

Q :

mep [BjnjB^^

2: a2

Q : :
►J : :

• * 0 e>

: : i""

. . -ro • • • •

'. a '■ '■ '■~i '• '• d

'. '.^^ 0 0 '•'•'•'•'• '•

qdasof jg Mopq aSpug

-H Sim

^\-

_» . . .

. -c^

■ ■ -St ■ ■ ■ ■

CM

CO

'. a ■'•'.'■'.'• 0

.^ '.....','. l 0 '

qdasof -^g

-a« ^ .^

• -CM ;

• ■ • -co • • •

CO

■ rt ^ C8 C8 -^ *** •-» ■

•

• -^

sajim z '■•SAia uoodg

:2 a2

Q :

:::•-::::

-h"

8|ira I 'iaAiy uoodg

2 as

Q :
J :

.-;.- oj' .

. . .CM ....

1-
|CO

'.'.'.'. ^ '.'.'. '.

■ '■'■ si '.'■'■ a '■'■'■ '■

6 33P!Jq '^-"'d ?|SS

05 e^CM

Q : :

'.'.'.'. ci '.'.'. '.

L 33puq 'JJJOJ }iBg ^ ^^

'.'.'.'. ei '.'.'. '.

{. agpijq -^ijoj ?[Bg

t» boo

CO ^H

Q : ;

'.'.'.'. a) '■'■'■ '■

Z a3p!iq 'jjjoj ■jiBg

<o bOO

00 ^H

Q : :

I ^SPl-iq '^JOJ 11^3

»o bflC^

Q : :

:

a '.'.'.'.'.'.'. '.

ajjBi [b^sAjq

'82 1

Q :
►J : '■

! : oi 'a

t» 0 «"^

O • M

00'

paq raB9ns p|o

n g|0

Q :

i a ■ 0 '■

. . 0 .

«••■•;;

lo

• t>

qa^ip aSBuiBjQ

01 gCM

Q : :

• cj— ■■- •

OS ■ ■ ■

>o

V°d llBg iaddQ

-as

a : ;

-■

Symbols
a, abundant
c, common
(', infrequent
1,2, 3, specimens found
L, living
D, dead
g, gravel
m, mud
», sand

is
■i

a^

!^

o
-a

Mollusca

Goiia caperata

Phyaa gyrina

Muaculium truncatum

Phyaa crandalli

Planorbia trivolina

Galba humilia modicella

Sphaerium atriatinum, var

Sphaerium atamineum

Sphaerium aolidulum

Musculiuvi partumeium

Muaculium tranaversum

Ferriasia tardua

Gundlachia meekiana

Campeloma rufum

Ferriaaia rivularia

Piaidium compreaaum

Pleuroccra elevatum

Gotiiobaaia livescena

60/60 par 10

Piaidium kirklandi

Piaidium aplendidulum

Sphaerium species

"o

a

£
8

Q.

00

■s

: ■ : : :3*i • :

-^ :«- -■si-ss^-l-
oS?o,33«„.gi;5

2-3^ 3.;5 a 0 oC c
CtlO>JQ3C>0-JOOQ.

i.-? i : : ig : : : ;^

:-s-...ca|-§,ri -.^ii

liiriiiHiil

3 Cob N-g <5 0-0.0 H;!i.«.=

il|-i III iiSl-Si

Perla nymph

Corydalia larva

Benacua species

24 JLUNOIS BIOLOGICAL MONOGRAPHS [122

GENERAL DISTRIBUTION OF THE MOLLUSK FAUNA

It will be noted in Table III that the mussels were found in greatest
abundance in a bottom composed of sand or gravel, or both, and were
fewer in number of species, as well as in individuals, on a mud bottom.
As a rule the mussels were found in abundance on the shallow riffles and
were often absent from the deeper places in which the bottom was composed
of soft mud. A notable exception to this rule occurs at Homer Park where
the largest mussel fauna is found in a mud bottom, and where this kind of a
bottom produced eleven more species (28) than did the sand and gravel
riffles a short distance below (17). This station has been used by Professor
Frank Smith for many years as a field habitat for his zoology classes and
the mussel fauna is, therefore, better known than that of any other locality
on the stream. Although visited several times a year for nearly a score of
years, there seems no diminution of the fauna in either species or individ-
uals. A day spent at this station, during which two collectors examined the
stream, yielded 24 of the 28 species. This indicates the great abundance of
the fauna, which may be due in large measure to the aerating influence
of the dam situated just above the collecting grounds.

The stream below the dam at Homer Park is an excellent place in which
to study the ecological conditions governing the distribution of the mollusk
fauna in a small stream. There is first a very shallow stream below the
dam (Fig. 14) flowing over gravel and boulders, in which a few gastropods
and small bivalves (Sphaerium) live in considerable abundance (Fig. 15).
Mussels are rare. Then follows a comparatively deep area of the stream
(2-3 feet) flowing over a bed of fine sand or mud in which mussels are
abundant and gastropods rare (Fig. 13). This is followed by a moderately
shallow stretch of the stream (1-2 feet) which flows over a bottom of coarse
sand and gravel in which mussels are found in abundance, although not as
numerous in species, as in the mud habitat. A few gastropods live here.
These conditions are exceptional in Salt Fork and are not duplicated in
any other part of the stream above Danville.

Some species of mussels, as Anodonta, Anodontoides, and Uniomerus,
prefer a mud bottom and thrive only in such situations, but the great
majority of the naiades prefer a sand or gravel bottom in water with
considerable current, and this is the reason they are so abundant in the
riffles of all streams. Among the smaller bivalves (Sphaerium and Pisi-
dium), the majority of species prefer a mud or fine sand habitat. Of the
snails or gastropods, Pleurocera and Goniobasis are usually found in a

123] FAUNA OF BJG VERMILION RIVER— BAKER 25

rocky (gravel) habitat while Campeloma prefers a mud or fine sand bot-
tom. Physa lives in both mud and on rocks and Ancylus on vegetation or
in empty shells of mussels. The ecological preferences of these species, as
well as the associated animals, are shown in Table IV.

ECOLOGICAL VARIATION

The 35 species and varieties of Unionidae found in the Big Vermilion
River west of Danville show an interesting distribution. Table III clearly
indicates that there is a more or less gradual increase in the number of
species as the stream increases in size. Taking into consideration both dead
and living naiads and ignoring for the time the effect of sewage pollution
on the distribution, the increase in species correlated with the increase in
distance in miles from Urbana may be expressed in the following table:

Table Number V. Increase in Species with Distance

Station No. of Species Distance from Urbana

Big bend 6 14.25 miles

One mile north iron bridge 12 16.50 "

Railroad bridge 15 20.00 "

Two miles above Homer dam 14 24. 75 "

Homer Park.. 28 27 .00 "

South of Muncie 23 36.00 "

Salt Fork junction 15 44.00 "

Middle Fork 22 45.00 "

Big Vermilion 21 46.00 "

The sudden rise in number of species at Homer Park is noteworthy and
is due to the exceptionally favorable environment, good depth of water,
favorable bottom, plenty of food, and a fully normal supply of dissolved
oxygen provided by the dam just above the Park. The dam appears to be
an effective barrier to the migration of mussels, and it would also seem
difl&cult for fish to pass the dam, except at very high water, and thus
migration in the glochidial stage is rendered difficult or impossible. In
the table it may be noted that 17 species occur at five stations, two to
eight miles above the dam and 28 species occur below the dam. Seventeen
species are common to both areas and 12 species are found below but not
above the dam. This distribution is shown in Table VI.

A striking feature of naiad distribution, noted repeatedly in several
species, is the conspicuous change in the shape of the shell as the distance
from the headwaters of the stream increases. Species that normally have
swollen or globose shells in the larger rivers, occur as flat or compressed
forms in the headwaters of these streams. An increase in length of shell
as correlated with decreased obesity is also noted, and also, a decrease in
tuberosity. Ortmann (1920) has recently ably discussed this matter and
shows that the rule holds good for many species in widely separated areas;
the writer cannot fully agree with Ortmann in reducing so many species

26 ILLINOIS BIOLOGIC A L MONOGRA PUS [ 1 24

of Naiades to varieties on the basis of compression in the headwaters of
streams. The same species varies in obesity, but it is the same species,
whether thin or fat. Other characters are usually present which separate
the allied species.

Table VI. Species of Unionidae Found Above and Below Homer Park Dam

Both Above and Below Dam Below Dam Only

parea anodontoides

fcrussacianus dlipsiformis

grandis ligamenlina

imbecillis muUiradiata

edenhdus lackrymosa

pavonius melanevra

luteola wardii

lienosa R- tubercvlata

complanala T. htherculata

rubiginosa circulus

undulata clava

ptislulosa glans
costata
marginata
cocsineum
veniricosa
compressa

In the Big Vermilian this variation in compression is marked in several
species. Thus Rotundaria tuberculata, Pleurobema coccineum, and Amblema
are more compressed than are individuals from the Wabash River below
the junction of the Big Vermilion with that river. Quadrula pustulosa is
smaller than the same species lower down in the Salt Fork, and the same
may be said of Alasmidonta marginata and Strophitus edentulus. That the
rule does not always hold good is shown by the variation of Fusconia rubi-
ginosa which is abundant in most parts of the Big Vermilion and its tribu-
taries. Measurements are given in Table VII, showing the length and
breadth of several species in different parts of the Salt Fork from below
Urbana to the Big Vermilion. The percentage of width to length is also
shown.

It will be seen that the average index for the first lot is 42 per cent and
for the last lot, 46 miles down stream, is almost the same, 45 per cent.
These averages compare well with some of those given by Ortmann (1920:
283). It was observed, however, that in the Salt Fork and Big Vermilion
the obese individuals occurred with tlie compressed specimens the former
increasing in ratios as the distance down stream increased. The variety
wardii of Quadrula metanevra occurred in two places in Salt Fork but always
in company with the typical form. In the cases cited above ecological
features cannot be called into account in locating the cause of the com-

125] FAUNA OF BIG VERMILION RIVER— BAKER 27

pression of the shell, for all live in the same section of the stream under
identical conditions. The rule cited by Ortmann, however, seems to be
applicable in most cases.

Several species increase in size toward the lower part of the river. This
is especially true of Anodonta grandis, Strophitus edentulus, Alasmidonta
marginata, Eurynia lienosa, Lampsilis luteola, Amblema undulata, and
Lampsilis ventricosa. A few others show some increase at different stations.
One species, Uniomerus tetralasmus, is apparentl)'' confined to the upper,
smaller tributary streams of Salt Fork. It was common in the ditch north
of Urbana; in Crystal Lake, Urbana, in Spoon River; and at Muncie in a
small tributary. Two broken valves were found at the station called the
natural dam, but these are believed to have been washed into this stream
from a nearby tributary which was dry when this part of the Salt Fork
was examined (September 25). Tetralasmus probably also occurs in the

Table VII. Variation of Fusconaia Rubiginosa

Length Width Per cent Station No. Distance from

Urbana

58 23 39 17 15^ mUes

62 26 42 ..

61 31 SO ..

68 27 39

79 33 42

86 33 38 24 243^ miles

89 41 46

90 31 34 25 27 miles

82 35 42 ..

89 47 52 ..

71 30 42 26 36 miles

67 30 44

102 41 40 30 46 miles

65 30 46 ..

86 43 50 ..

upper part of Middle Fork and in North Fork, but the upper parts of these
streams have not been examined. The majority of the other species occur
at several stations along the Salt Fork and its tributaries and no particular
variation in distribution was observed except as already noted.

Three mussel species and varieties were found in Crystal Lake that
are exotic as far as the Big Vermilion River is concerned. These are Ano-
donta grandis gigantea, Anodonta corpulenta, and Uniomerus tetralasmus
sayii. These species were artificially introduced into this body of water
about 1908 by a member of the zoological department of the University of
Illinois. They originally came from a stream in western Indiana. That
these mussels found a favorable environment and have thrived during these
years is evidenced by the number of fine specimens recently collected when
the lake was partly drained. Only the Uniomerus was rare, but one speci-
men being found. As Crystal Lake is not connected with the Salt Fork

28

ILUNOIS BIOLOGICAL MONOGRAPHS

1126

stream, these species have not been able to enter the Big Vermilion drain-
age.

As has been noted in the Illinois River (Forbes and Richardson, 1919),
the mussel fauna gradually increases as the distance from the source of
sewage pollution becomes greater. In the Salt Fork the fauna becomes
normal at about 20 miles from the source of pollution at Urbana. In the
Illinois River a normal fauna is not found within 80 miles (Hennepin) of
the source of pollution indicating that the quantity of sewage is so great
that the river must flow this distance before purifying itself suflSciently for
the residence of normal aquatic life. That a normal fauna should be
found within 20 miles of the source of pollution in the Salt Fork, though a
much smaller stream carrying a smaller amount of sewage, is quite sur-
prising when it is remembered that no large tributaries enter the stream
above Spoon River, and indicates that self purification is active. The
shallowness of the water (less than a foot on the average in fall and winter)
probably provides a larger quantity of dissolved oxygen than woiTld be
possible in waters of a deeper stream. It was especially noted that Am-
blema undulaia and Lasmigona complanata, of the larger species, withstood
the absence of water better than any of the other comparable species.
These mussels also resisted polluted conditions better than others and this
fact is important in connection with mussel propagation for button shells.

COMPARISONS WITH OTHER RIVER SYSTEMS

It is of interest and value to compare the mussel fauna of the Big
Vermilion River with that of some other rivers of comparable size and
development. The United States Bureau of Fisheries has conducted
mussel investigations of several of the rivers of Illinois and adjacent states
and one of these, the Kankakee (Wilson and Clark, 1912), may well be
compared with the Big Vermilion. The mussel fauna of the Sangamon
River is also included, the data given being gathered from several sources,
but principally from personal collections and from collections in the
Museum of Natural History of the University of Illinois. Some species
not listed by Wilson and Clark are included from Baker's Catalog of
Illinois Mollusca (1906). These are indicated by an asterisk.

Table VIII. Distribution of Unionedae in Three River Systems

Length of river in miles

Quadrula cylindrica

" metanevra

" metanevra wardii. . .

" puslulosa

" lachrymosa

" ebena

Trilogonia tuber ctUala

Amblema unduleta

" peruviana (plicata).

Vermilion
90

X
X
X
X
X

X
X

Kankakee
300

X
X

X

*

X

Sangamon
150

X
X
X
X
X
X
X

127]

FAUNA OF BIG VERMILION RIVER— BAKER

29

Table VTII — (continued)

Fusconaia rubiginosa

" trigona

" solida

Rolundaria tuhercidata

Pleurobema clava

" coccineum

Plethobasus aesopus

Elliptio gibbosus

Uniomerus tetralasmus

" tetralasmus say it

Slrophittis edentulus

" edentulus pavonius

Anodonta grandis

" grandis gigantea

" corpulenta

" imbecillis

Anodontoides fertissacianus

" /. buchanensis

Arcidens confragosus

Lasmigona compressa

" costata

" complanata

Alasmidonta marginala

" calceola

Ptychobranchus phaseolus

Obliquaria reflexa

Plagiola securis

Amygdalonaias elegans

" donaciformis

Propter a alata

Paraptera gracilis

Obovaria circulus

" ellipsis

A ctinonaias ligamenlina

" ligamenlina nigrescens.

" ellipsiformis

Caruncidina parva

" glans

Eurynia recta

" subrostrata

trts.

" fabalis

" lienosa

Lampsilis fallaciosa

" anodontoides

" ventricosa

" multiradiata

" capax

" luteola

" higginsi

Truncilla sulcata

" perplexa rangiana .

Total species in each river .

Vermilion

X

X
X
X

X

X
X
X
X
X
X
X
X
X

X
X
X
X

X
X

X

X

X
X

38

Kankakee

X
X

*

X
X
X
X
X

X
X

X
X
X
X

*
X

*

X
X
X
X
X

*

X
X
X
X
X
X

X
X

X

Sangamon

X

X
X
X

X
X
X
X

X
X

X
X

48

25

30 ILLINOIS BIOLOGICAL MONOGRAPHS [128

It is noteworthy that with a length of 300 miles and with two tributa-
ries of large size (Iroquois River, 100 miles in length, Yellow River, 65 miles
long) the Kankakee River has a mussel fauna only 21 per cent greater than
the Big Vermilion River with a length of 90 miles and no very long tribu-
taries. Other species will proably be found in the Big Vermilion below
Danville, which was not examined during this survey, and these may
bring the total nearer to that of the Kankakee River. The Sangamon
River undoubtedly contains many more species than listed in the table,
and these will be found when additional collecting is carried on. The table
shows that the mussel fauna of the Big Vermilion River is of large size as
compared with other streams of similar character.

129] FAUNA OF BIG VERMILIOIi RIVER— DAKER 31

SYSTEMATIC DISCUSSION OF THE MOLLUSCA

In this chapter the species of mollusks, both Pelecypoda (mussels,
clams) and Gastropoda (snails), are discussed in relation to their distribu-
tion in the Big Vermilion River, special emphasis being given the Unionidae
or river mussels on account of their economic importance. The influence of
sewage pollution of the stream on the mollusk fauna is also referred to.
The species collected in the Sangamon River at Mahomet and elsewhere are
included for purposes of comparison.

The classification followed for the Unionidae is that proposed by Simp-
son (1900, 1914) and extended by Ortmann (1912, 1918). The sequence of
groups is that set forth in Walker's Synopsis recently published (1918).
It will be noted that the newer classification necessitates the adoption of
several new names, both generic and specific, but these seem, on the
whole, justified by the rules of nomenclature and are a natural result of
the advancement of knowledge on the subject.

For the purpose of providing reliable data on the particular character-
istics of the mussels and other mollusks in this stream for comparison with
similar features of this group of animals in other streams, a feature almost
totally lacking in the literature, considerable space is devoted to descrip-
tions of the minor variations and pathological conditions of each species
in dififerent environments. This has been done, more or less extensively,
in several reports on the mussel faunas of three or four of our Illinois,
Indiana, and other streams (Wilson and Clark, Danglade). It will be
noted that there are certain features characteristic of the species in one
stream not shared by the same species in other streams, as, for example,
Lampsilis ventricosa which differs markedly in coloration and even in
shape in the two river systems herein considered. Similar data on our other
rivers would provide a body of facts of considerable importance.

FAMILY UNIONIDAE

1. Ouadrula (Quadrala) cylindrica (Say). Rabbits-Foot.

This species was not found in the Salt Fork above a point about a mile
west of its junction with the Middle Fork, 44 miles from Urbana. As it is
not listed from the neighborhood of Muncie its westward extension in
the stream lies somewhere between Muncie and Middle Fork. Even in
this part of the river it is rare and the specimens obtained are small, of
dark color and resemble the form called strigillatus by Wright. Cylindrica
is not a widely distributed species in Illinois, if one may judge by the
records at hand. Danglade (1914) did not find it in the Illinois River

32 ILLINOIS BIOLOGICAL MONOGRAPHS [130

nor is it listed by other students. It has been reported by several concholo-
gists from the Wabash River (Baker, 1906:79) and the Ohio River, in which
streams it is common and of large size and fine color. The species probably
would not thrive in polluted water. It was not found in the Sangamon
River at the places visited. Owing to its peculiar shape it is not adapted
for the cutting of button blanks and is considered worthless by the mussel
fishermen.

2. Quadrula (Quadrula) metanevra Rafinesque. Monkey-Face.

This naiad is apparently a rare species in Salt Fork occurring sparingly
from Homer Park to Middle Fork. It begins to increase in number of
individuals near the Middle Fork, where the specimens are also larger
and more brilliantly colored. All but one of the specimens collected are
typical in form and coloring. Individuals from the Big Vermilion below
Middle Fork are larger than those collected above this point. The species
is also more abundant. It is found on both a mud and a gravel-sand bot-
tom. Specimens from Homer Park are darker and less conspicuously rayed
than those from Middle Fork, and are also less pustulose. Metanevra is
rare in the Sangamon River, living on both a sand and gravel bottom.

2a. Quadrula (Quadrula) metanevra wardii (Lea).

Two specimens referable to this variety have been collected from the
Big Vermilion; one near Muncie, in the Salt Fork, and one in the Middle
Fork, above its entrance into the Vermilion River. These individuals
are more elongated and compressed than the typical form and the tubercles
are not as heavy, in fact are reduced to large pustules. Professor Smith
has found the variety more common in the Sangamon River than the
typical form, and until these two specimens were found in the Big Vermilion
drainage, wardii was supposed to be the predominant form in the Sanga-
mon while the typical form was believed to be the only form of this species
found in the Big Vermilion, at least above Danville. Wardii i^, as far as
present material indicates, very rare in the Salt Fork and other tributaries
of the Big Vermilion.

3. Quadrula (Theliderma) pustulosa (Lea). Warty-^&ack; Pimple-Back.

This is the most abundant Quadrula in both the Salt Fork and the San-
gamon River, rivalling in number any other mussel species in the lower
part of the Salt Fork. It does not occur in any abundance above the
Homer Park dam, but below this point it is common, of large size, fine
color, and good nacre. The sewage pollution has evidently affected this
species as others and, with rare exceptions, only dead shells could be found
above the Homer Park dam. Below the dam it occurs commonly and the
increase in number of individuals is largely due to the aerating effect of
the flow of water over the dam which provides the dissolved oxygen so

131] FAUNA OF BIG VERMILION RIVER— BAKER 33

necessary to these animals. The young shells (20mm. in length) are almost
smooth with a broad dark green ray or stripe extending from the umbones
to the ventral margin of the valve. No young shells were found in the
stream above Homer Park dam and the species may not be breeding in
this part of the stream at the present time.

The pustulosa from Salt Fork exhibit some variation in the pustulosity
of the surface, but all are referable to typical pustulosa. A few specimens
from Homer Park are more quadrate than the average but are otherwise
typical. The individuals from the Sangamon River, however, show con-
siderable variation in both form and pustulosity, ranging from nearly
circular to quadrate and from almost smooth to quite pustulose. On an
average, however, they are less pustulose than the species occurs in Salt
Fork. Individuals might be picked out that could be referred to both
dorfeuillianus Lea and schoolcraftensis Lea, but the range of variation is so
great that they seem better referred to pustulosa. In the Sangamon River
pustulosa occurs on a sand and gravel bottom but in the Big Vermilion
River it is found most abundantly on a mud bottom. The individuals
from Homer Park and the Sangamon River, especially the later, are of
good size, 70 to 80mm. in length, and the thickness of the shell combined
with the clear pearly luster would seem to make them good shells for the
button trade. No evidences of parasitism were observed in the shells
examined.

4. Quadrula (Theliderma) lachrymosa (Lea). Maple-Leaf.

This handsome shell is very rare in the Salt Fork and was not found
at any of the localities in the Sangamon River. In the Salt Fork it has
been found only at Homer Park below the dam, and only occasional speci-
mens have been collected here. These are quite typical of the species.

5. Tritogonia tuberculata (Barnes). Buck-Horn; Pistol-Grip.

The buck-horn first makes its appearance in Salt Fork at Homer Park
below the dam where it is of large size (female 145, male 115 mm.) and fine
quality. The shells are densely covered with tear-like pustules which in a
few individuals cover the entire surface, though usually confined to the
middle and anterior end behind the posterior elevated ridge. Of the speci-
mens collected 40 percent are males. Young specimens 46 mm. in length
were found at Homer Park. Individuals from Homer Park are larger than
those collected in the Sangamon River, the largest specimens being found
on a mud bottom, although it also lives on a sand and gravel bottom. Both
the Salt Fork and Sangamon specimens are of good quality from the button
makers standpoint.

Abnormalities and pearly growths due to injuries or parasitism are
rare in the specimens of this species examined. A few individuals from
Homer Park had scattered pin-head pearls and a small patch of discolored

34 ILLINOIS BIOLOGICAL MONOGRAPHS [132

blister formation near the anterior and posterior end, one in each end of
two specimens.

6. Amblema undulata (Barnes). Blue-Point; Three-Ridge.

This characteristic mussel is the most abundant species in the Vermilion
River, greatly exceeding (with the possible exception of Lasmigona com-
planata) in number of individuals all other species. It is also able to resist
much of the ill effects of sewage pollution and is the first shell met with in
the polluted waters of the Salt Fork. Living specimens, however, were
not seen above the station called bench mark 655, a distance of over 15
miles from the source of pollution. Empty shells and odd valves occur
more or less abundantly from St. Joseph, 10 miles below Urbana, to the
station mentioned. As the species lives in fair abundance in the tributary
known as Spoon River, for a distance of over two miles up stream from near
the mouth of the stream, it is evident that at one time its distribution was
equally continuous in the Salt Fork below the junction of Spoon River
with Salt Fork, where now there is a break of nearly six miles. This break
in the distribution is in all probability due to the sewage pollution, for the
stream is admirably adapted by nature as a habitat for this species and
has not been disturbed by dredging.

There is great variation in the form of the shell. Many specimens from
the upper part of the stream, both Spoon River and Salt Fork as far down
as the natural dam, are almost round with a broad 'wing' above the undula-
tions, which may be reduced in number and form (Fig. 34). These
shells may be inflated or rather compressed. Other shells are more quad-
rate and in occasional individuals the umbones are elevated simulating
Amblema peruviana {plicata of authors). These shells have a black or
dark brown epidermis in the adult condition. In Spoon River young
shells 25 mm. long were common, but few young specimens were found in
the Salt Fork above the Homer Park dam. Shells from the lower part of
Salt Fork, below Sidney, are as a rule cleaner, the epidermis is of a brighter,
lighter brown and are more uniformly quadrate than those from above
Sidney. The largest specimen collected measured 140 mm. in length and
this seems to be the maximum size for the undulata in this stream. Many
of this size were seen.

In the Spoon River, and in the upper part of Salt Fork, injured shells
are common. The injuries consist of breaks in the shells and subsequent
repairs. In one specimen from the upper part of Spoon River, an injury
had been received when the mussel was small which resulted in a deep
channel across the right valve (Fig. 29) and a ridge, also slightly chan-
nelled, on the left valve (Fig. 30). Another shell had nearly a hundred
blister pearls on the edge of the posterior margin of the left valve (Fig. 28)
and a large blister pearl about midway of the pallial line in the right valve

133J FAUNA OF BIG VERMILION RIVER—BAKER 35

(Fig. 27). Still another shell had covered a quantity of mud, which had
gotten in between the mantel of the animal and the shell, with a thin layer
of pearl, forming a large pad-like blister covering the greater part of the
interior of the left valve (Fig. 26). The right valve was normal (Fig. 25).
These blister pearls, as well as the more valuable free pearls, are believed
to be caused by parasites, perhaps distomids. These injured shells are
eagerly sought by the pearl hunters and mussel fishermen in the belief
that they may contain pearls of value.

The undulata from the Sangamon River also exhibit a wide range of
variation in the shape of the shell, but not to the degree seen in the material
from the Spoon River and Salt Fork of the Big Vermilion River. In the
Sangamon, undulata is very abundant on a gravel and sand bottom.
In the branches of the Big Vermilion it occurs on both a mud and a sand-
gravel bottom. As this species is very successful in resisting adverse
conditions it is a valuable mussel for propagation in the streams subject
to pollution. It is probably not much affected by a moderate amount of
sewage in its environment. The button manufacturers consider it a good
shell when the undulations are not too heavy to render the cutting of
blanks difi&cult.

7. Fuscunaia rubiginosa (Lea). Wabash Pig-Toe.

This mussel is abundant in the lower part of the Salt Fork. It was not
found in any abundance above the dam at Homer Park, where it is abun-
dant, and the large number of dead, empty shells observed attest the
presence of an unfavorable environment. As it is rare in Spoon River,
where some other species are abundant, it is probable that this species
requires fairly deep water and a large stream bed to attain good size and
abundance in individuals. This characteristic of distribution was also
noted by Wilson and Clark (1912:43) in the Kankakee River where ruhigi-
nosa was found to be more common in the lower part of the river.

There is considerable variation in the form of the shell; some examples
are compressed, others quite inflated. Nearly all are distinctly quadrate,
but in some examples the ventral margin is convex; in others it is some-
what concave; while in a few it is straight. The individuals from the
upper part of the stream, above Homer Park dam, are usually dark brown
with a satiny sheen to the epidermis and are almost rayless. Those from
the lower part of the river, especially from Middle Fork, are light yellowish
brown, quite distinctly rayed. The nacre varies from white to pink or
salmon, but is white in the great majority of specimens collected. Young
shells 28 mm. in length were common below the Homer Park dam, but were
apparently rare above the dam. This may indicate adverse conditions
due to sewage pollution and the species may not now be breeding freely,
possibly for lack of suitable fish for the glochidia. Young specimens were

36 ILLINOIS BIOLOGICAL MONOGRAPHS ■ [134

also collected in the Big Vermilion River. One of the largest adult individ-
uals found, below Homer Park dam, measured 95 mm. in length; another
from the Big Vermilion River measured 103 mm. in length. The species
occurs about equally on a mud or sand-gravel bottom.

A single example from Homer Park is worthy of special note. It is
large, inflated, almost twice as wide as the average shell of the same size,
and is elongate-quadrate in outline. When viewed from within, the
valves are basin-shaped. All of the muscle scars are very heavily impressed
and the pseudocardinal teeth are much modified and heavier than in normal
rubiginosa. The lateral teeth are very high, wide and massive. The
shell was dead when picked up and badly discolored and notes on the ani-
mal, which would have been very desirable, could not be made. The
measurements of this shell, together with that of a normal shell of the
species, from the same habitat, are given below:

Length, 90; height, 59; width, 50 mm. Z11163 A, Variety.
" 82 " 58 " 36 mm. Z11163 B, Normal.

This species also occurs in the Sangamon River, but does not, appar-
ently, attain the dimensions of the Salt Fork specimens, nor does it occur
as abunantly. There is but slight variation in form in the Sangamon
shells. No pathological specimens were observed in individuals from either
river. Rubiginosa is not much esteemed by either pearlers or mussel fisher-
men.

Rubiginosa frequently closely resembles Pleurobema coccineum in the
form of the shell and specimens occur which seem difiicult to place satisfac-
torily. The animals diflfer in that in rubiginosa all four of the gills are used
as marsupia while in coccineum only the outer gills are so used. As far as
the Salt Fork and Sangamon River shells are concerned there has been no
difficulty in placing any individual. In this material rubiginosa is always
quadrate with the umbones large and full, the posterior end of shell is
almost sharply truncated and there is a more or less distinct ridge extend-
ing from the umbones to the posterior angle of the shell. In coccineum
the outline is rather ovate, or rounded, there is no posterior ridge and the
position of the umbones gives to the shell an oblique appearance which is
very characteristic and is absent in rubiginosa. The ventral margin in
coccineum is almost always convex and seldom straight or concave as in
rubiginosa. The young shell in coccineum is also usually more distinctly
rayed. The surface of the two species is also different, that of coccineum
not being 'satiny' as is that of rubiginosa. The interior, and even the
exterior, of coccineum is usually pinkish or salmon colored, although
individuals occur with white nacre.

135] FAUNA OF BIG VERMILION RIVER— BAKER 37

8. Pleurobema clava (Lamarck). Club-Shell.

This species is rare in most parts of the Big Vermilion examined, and
was not found in the Sangamon. It occurs sparingly at Homer Park,
abundantly in the Salt Fork near Muncie, and sparingly in the Middle
Fork. At Muncie the largest specimen measured 90 mm. in length. The
specimens from the Big Vermilion are beautifully marked with broad green
rays on young individuals and on the umbonal half of older specimens.
Large individuals are almost rayless.

This species has been previously known only from the Wabash River in
Illinois (Baker, 1906:77) and the present records, although in the same
drainage basin, extend the range of its distribution.

9. Pleurobema coccineum (Conrad). Thin Niggerhead.

The shell known as coccineum attains large size in certain parts of the
Salt Fork. It is common, however, only at one place, below the dam at
Homer Park. No shells of this species were observed above the station
called bench mark 655, which is about 15 miles below Urbana. The shells
from habitats above the dam at Homer Park exhibit evidence of an un-
favorable environment, the shells having heavy lines of growth which on
some specimens are raised to form ridges. These are especially marked at
the rest periods (seasonal). The individuals from the upper part of the
stream are also more or less pathologic, 50 per cent of the shells being
abnormal in form or with pearly growths on the inside of the valves.
Coccineum is found on both a mud and a sand-gravel bottom.

There is considerable variation in the outline and general shape of the
shells from Salt Fork. The outline varies from quadrate to roundly ovate
and the ventral margin from nearly straight to strongly convex. Young
and half-grown shells seem more uniform than large adult shells. The
quadrate individuals may easily be confounded with Fusconaia ruhiginosa.
In the last species, however, the shell is more inflated {coccineum is com-
pressed), the umbones are directed upward and not backward, as in
coccineum, and the posterior portion of the shell has a depressed area and
a strong ridge which are absent in coccineum. The nacre of coccineum is
pink of various shades, only two specimens from the Salt Fork having
white nacre. In this respect the coccineum of the Big Vermilion drainage
differ almost constantly from rubiginosa which has white nacre. The
epidermis in specimens from the upper part of the stream (Homer Park and
above) is usually very dark brown with faint evidences of rays. In some
specimens, especially from the station three and a half miles above Homer
Park dam, the whole shell, inside and outside, is of a delicate pink shade.
Individuals from the Middle Fork and the Big Vermilion, where the species
is rare, are lighter in color. The two largest specimens collected from the
Big Vermilion drainage measure as follows:

Length, 96; height, 75 mm. Quadrate form above Homer dam, Z11114 A.
" 96 " 80 mm. Rounded form, Homer Park, Zl 1164 A.

38 ILLINOIS BIOLOGICAL MONOGRAPHS [136

In the Sangamon River coccineum is a most abundant and variable
mussel, having a smooth, polished shell on which the rays are many and
distinct. It also attains a large size, though not as large as specimens
from the Big Vermilion drainage. The shape of the shell is more oblique
and eliptical than is the species as it occurs in the Big Vermilion and the
shell is a trifle more inflated on the average. So marked is the difference
that it is comparatively easy to say from the shape of the shell and the
surface markings what drainage a particular individual may have come
from. The twelve sets of coccineum in this collection indicate in a marked
degree the fact that species may differ conspicuously in both sculpture
and form in different river systems. The nacre of the Sangamon River
coccineum is more often white than in the Big Vermilion shells. Pearly
secretions or pathologic malformations have not been observed in the
specimens from the Sangamon River, indicating, without doubt, a more
favorable environment than is provided by the waters of the Salt Fork.
Young specimens (25-30 mm. long) are more abundant in the Sangamon
River than in the Big Vermilion River, and these individuals are beautifully
marked with dark green rays on a yellowish or light brown background.
Occasional shells are pinkish. The beak markings on the umbones are
especially well preserved in these young specimens.

A large right valve from Mahomet is very peculiar. In outline it is
ovate, a trifle oblique. The posteroir portion of the valve is much elongated
the hinge line is long and straight, and the posterior margin is sharply,
obliquely truncated. The umbonal region is near the anterior margin of
the valve. The lateral tooth is longer and straighter than in normal
coccineum. The shell recalls Pleurohema clava but is much larger and
differently shaped. The valve measures as follows: length 92, height
70 mm.

Ortmann (1918:549) considers coccineum a variety or race of obliquum
(Conrad), together with solidus{ = catillus Conrad), which is also rated as
a variety of obliquum. To this disposition the writer cannot agree, the
forms here listed as varieties being quite as much entitled to specific rank
as are many other forms recognized as distinct species which have marked
variation and a similar facies. The whole group of obliquum- solidum-
coccineum are closely related, but I have seen no good reason after
examining a large series in the Hinkley and other collections in the
Museum collections, for lumping these species as varieties of obliquum.
As far as Illinois specimens of obliquum and coccineum are concerned, the
two species seem sufficiently distinct for recognition.

10. Rotundaria tuberculata Rafinesque. Purple Warty-Back.

This species was found at but two places in Salt Fork, at Homer Park
and South of Muncie, and in the Big Vermilion below Middle Fork. At

1371 FAUNA OF BIG VERMILION RIVER— BAKER 39

the first two places it is very rare, only a few individuals having been
found by Professor Smith in a number of years. In a days search at
Homer Park by two experienced collectors, only two living specimens and
odd valves of two others were found. The largest specimen from Homer
Park measures 72 mm. in length and 63 mm. in height. This species is one
of the most abundant of shells in the Big Vermilion below Middle Fork,
where specimens measuring 100 mm. in length are common. The distri-
bution of this species is a good example of the progressive development of a
species in the downward course of a stream, for in the course of about
twenty miles the size nearly doubles. Beginning as a rare form at Homer
Park it becomes one of the most common forms in the Big Vermilion,
twenty miles downstream.

All of the Big Vermilion tuherculata are of the compressed type, and
the shell is covered posteriorly and ventrally with large tear-like pustules.
The anterior third of the valve is free from pustulation. The nacre of all
shells seen is rich purple, which renders the species valueless for the button
makers. Tuberculala does not occur in the portions of the Sangamon River
examined.

11. Elliptic gibbosus (Barnes). Lady-Finger; Spike.

This mussel does not occur in Salt Fork, nor in any tributaries of the
Big Vermilion above Danville that have been examined. It is fairly com-
mon in the Sangamon River at Mahomet on a sand and gravel bottom.
Young and immature shells are distinctly rayed. The nacre of all speci-
mens examined has been purple, no white-nacred individuals being seen.
In the Kankakee River white-nacred specimens occur and become the
dominant form in the lower part of the stream (Wilson and Clark, 1912 :45).
In the Illinois River beds of shells occur which have either a white or a pur-
ple interior. (Danglade, 1914:42). This familiar shell will probably have
to be known as dilatatus (Rafinesque) if the original description is definite
enough to identify it as the gibbosus of Barnes. Dilatatus was described
in 1820. It is a pity that these names of Rafinesque could not have been
applied earlier to these shells and thus saved the confusion which is now
resulting from the changes of the old familiar names which zoologists in
our universities have used for years in connection with their classes in
systematic zoology.

12. Uniomerus tetralasmus (Say).

This species has been found living only in the upper waters of Salt
Fork and in Stony Brook near Muncie. It occurs in fair numbers in the
stream above Urbana and in Spoon River. Two broken valves were found
in Salt Fork at the station called natural dam about 12 miles below Urbana.
No living mussels could be found in the stream at this point and it is
believed that the odd valves were washed into Salt Fork from a small

40 ILLINOIS BIOLOGICAL MONOGRAPHS [138

tributary nearby which was dry at the time of our examination. Tetralas-
mus is a species of the small, mud-bottom tributaries and seems not to
occur in the larger part of the stream with the larger and heavier mussels.
The largest specimen collected measures 55 mm. in length. The colors of
the shells are yellow, black, and greenish, the latter in indistinct ray-form.

12a. Uniomerus tetralasmus sayi (Ward).

Among the Anodontas collected in Crystal Lake is a specimen of the
shell known as variety sayi. It is large for the species but seems otherwise
typical. The dimensions are: length, 123; height, 58; breadth, 38 mm.
(No. Z 11369). As only tetralasmus is found in the Salt Fork and its
tributaries it is probable that this form was introduced with the Anodontas
described on a subsequent page.

13. Strophitus edentulus (Say). Squaw-Foot.

This mussel once occurred in nearly all parts of the Big Vermilion River
but it is now found in any number only below the dam at Homer Park.
No living specimens were found above a point four miles above Homer
Park dam or 22 miles below Urbana. It occurs, rarely, living, in Spoon
River and its absence in a living state for a distance of 12 miles between
this tributary and the first habitat in which it was found alive in Salt
Fork is striking and suggestive of the harmful effect of sewage pollution.
This species reaches its greatest perfection below the dam at Homer Park
on a gravel bottom. Edentulus also occurs in the Sangamon River, but
the individuals from that stream are not as large and are more compressed
than the specimens from Salt Fork, which are as a rule quite corpulent.
It is also not as abundant in the Sangamon as in the Big Vermilion. Meas-
urements are given below of the largest specimens from the Salt Fork and
the Sangamon.

Length, 90; height, 54; width, 41 mm. Salt Fork, Z11174.
" 89 " 57 " 32 mm. Sangamon, Z11227 A.

There is great variation among the shells referred to this species.
Typical edentulus is rhomboid in form, rather inflated, with prominent,
inflated umbones; the posterior margin of the shell is usually sharply,
obliquely truncated, and the ventral margin is straight or even slightly
convex. The color is usually black without rays. From this t3^e the shell
varies to an ovate or elliptical outline, a more or less compressed form, with
a rounded, convex ventral margin and with an almost total absence of the
strong posterior ridge so characteristic of the usual form. The variation is,
as would be expected, toward the variety known as pavonius. The shells
from Salt Fork are very thick and solid, much more so than in specimens
from the Sangamon River. The nacre of the majority of specimens is
yellowish in color.

139] FAUNA OF BIG VERMILION RJVER— BAKER 41

13a. Strophitus edentulus pavonius (Lea).

This variety is described by Simpson (1914:348) as "Shell generally long
elliptical; epidermis yellowish-green, more or less covered with green or
brownish-green rays." In its typical form pavonius is easily separable
from edentulus. The variety is almost as common in Salt Fork as is the
typical form and there are many intermediate individuals. It is possible,
however, to separate all of the edentulus from the different stations, 17 lots,
into two groups; one with rhomboid or long-ovate outline and with black
or brownish, rayless surface; and the other with long-elliptical outline,
brownish surface, and many distinct rays. In pavonius the height is less
as compared with the length than in edentulus. These two forms of
Strophitus were almost always associated together, indicating their close
relationship. On the whole pavonius is much less variable than typical
edentulus. Specimens from the Sangamon River at Mahomet are brilliantly
rayed with patches of bright green on the ventral margin of the shell.

Wilson and Clark (1912:48) state that the "question of rays appears
to be closely related to clearness of water; in turbid streams mussels are
usually dull colored, while in clear streams they are usually brightly
rayed." This has been our observation in many cases, but the rayed
pavonius in Salt Fork occurs with the rayless edentulus in quiet water
on a mud bottom; both also occur in riffles on a sand-gravel bottom.

Pavonius is credited by Simpson to Ohio and Indiana; it is probably
widely distributed in Illinois, but has most likely been listed under edentu-
lus in most cases. It is known from^ Cook County, Will County, and the
Wabash River (Baker, 1906:72). Its presence in the Sangamon River
indicates that it is also an inhabitant of the Mississippi River drainage,
as well as the Wabash and Ohio drainages. It is quite probable that the
distribution of the variety is coincident with that of edentulus. The nacre
of both edentulus and pavonius is usually yellowish or salmon colored al-
though white-nacred specimens occur. Pearly growths are not as common
among the shells of this species as found in the region under consideration
as among the same species from other places. A few individuals had blister
and pin-head pearls. A specimen each of the type and the variety had a
peculiar pearl formation on the pallial line at or near the posterior end of
the shell. These are somewhat dome-shaped, about 5 mm. in diameter
and 4 mm. in height and evidently were caused by an effort on the part of
the mollusk to cover some irritating object, possibly a nematode worm
(Figs. 31, 32). As both shells were without the animal {pavonius had
been alive very recently) this point could not be determined. The
edentulus was from the railroad bridge east of Sidney (No. Z11098) and
the pavonius from below Homer Park dam (No. Z11144 A).

42 ILLINOIS BIOLOGICAL MONOGRAPHS [140

14. Anodonta grandis Say. Floater.

The floater or paper-shell is more or less abundant in Salt Fork and
other parts of the Big Vermilion drainage. In Spoon River it is common,
livijig in the lower part of the stream. From this station to the station
called bench mark 655, over five miles below, not a living Anodonta could
be found, and the species does not become abundant until the cement
bridge east of Sidney is reached, nine miles below Spoon River. This
distribution is again indicative of the harmful influence of sewage on the
bottom inhabiting animals. From the cement bridge to the Homer Park
dam grandis is fairly common. It was very rare below the station at Homer
Park, at which place it is common. The best habitat observed appears
to be between the cement and railroad bridges east of Sidney, where the
water is fairly deep in summer (three-four feet) and where there is a soft
mud bottom and not much of a current in the stream. The species is
t3q)ically a pond-inhabiting mussel. Gravid individuals were collected on
September 13, 1918.

At Mahomet, on the Sangamon River, grandis is abundant and of large
size, and occurs on a fine sand bottom. The Sangamon specimens are
on the whole more cylindrical in form than those from the Big Vermilion
and have a brown or brownish-green epidermis. The Big Vermilion
specimens are mostly grass-green in color and are more elongate-ovate
in form, the ventral margin being almost universally rounded while in the
Sangamon shells this margin is nearly straight. The Sangamon River
grandis are on the whole more solid than the same species from Salt Fork.

The nacre of the great majority of the Salt Fork specimens is bluish-
white, while that of the Sangamon specimens is salmon-colored for the
most part. A few individuals from both streams have salmon-colored
patches and small pearl growths indicating that the animals had suffered
from the attack of distomid worms, possibly the distomid of Osborn, which
is known to infest this species in other places (Wilson and Clark, 1912).
These shells, however, were rare and infection from this source seem
uncommon among the grandis of these streams. No Unionicola {Atax)
or other water-mites were observed in this species. These parasites are
common in grattdis irihabiting other streams (Wilson and Clark, 1912:
61-71).

An empty shell from the big bend in the Salt Fork showed evidences
of distomid infection in the form of elongated blisters on the ventral
margin of the valves, near the pallial line. In the right valve, near the
anterior adductor muscle scar, there is a large blister, 8 by 12 mm. which
evidently covered a distomid. The left valve of this specimen had suf-
fered an injury when the animal was about two-thirds grown, which has
caused a part of the antero-ventral margin to become folded inward, a part
of the folded portion having the epidermis well preserved. The animal

141] FAUNA OF BIG VERMILION RIVER— BAKER 43

continued its shell formation so perfectly that from the outside no evidence
of an injury is visible. This shell indicates plainly a case of a hard struggle
for existence against both mechanical injury and heavy parasitism. The
interior of the shell is spotted with grayish patches and salmon-colored
streaks (No. Z11029-A). (Figs. 22,23.)

14a. Anodonta grandis gigantea Lea. Floater.

Specimens of an Anodonta from Crystal Lake, Urbana, are apparently
referable to Lea's gigantea. Simpson (1914:420) diagnoses this variety
as "Shell large, ovate or sabrhomboid, a little higher in proportion to the
length than the type; beaks full and high." The specimens from Crystal
Lake agree with this diagnosis. The largest individual measures 152 mm.
in length and 92 mm. in height. The umbonal region is more corpulent
than in the grandis from the other parts of the Salt Fork. The color
is brownish or greenish, the two colors frequently in alternating zones
on the same specimen. Evidences of distomid infection are common in the
form of salmon or pink discolorations and ridges. One individual has many
long, thin, curved ridges on the interior of the shell, principally in the left
valve. One of these ridges measures 93 mm. in length and 1 . 50 mm. in
height (Fig. 24). Another individual has a round pearl attached to
the posterior end of the shell, measuring 5 mm. in diameter. This variety
has not been observed in any collections from the Big Vermilion or Sanga-
mon rivers. Marsh has recorded gigantea from the Big Vermilion (Baker,
1906:73) but the exact location is not known, and must have been below
the points examined by the writer.

15. Anodonta corpulenta Cooper. Floater,

The large Anodontas from Crystal Lake are divisible into two groups;
one is the variety of grandis described above; the other seems to be the
corpulenta of Cooper, although the shells are smaller than examples of
this species from other rivers. The shells referred to corpulenta are sub-
rhomboid, somewhat elongated in a few of the individuals. The umbonal
swelling is very pronounced, extending well downward on the shell. The
anterior end is broadly rounded and the posterior end is distinctly plow-
shaped and rather strongly biangulate. The epidermis is olive or brownish.
The surface is very rough, the growth lines in some specimens being
elevated into longitudinal ridges. As in gigantea, the inner surface is
ridged and salmon-colored in many specimens due to the presence of
distomid worms. No specimens of this species were seen which did not in
some degree show evidences of the work of this parasite. Characteristic
measurements of this shell ate given below (Z11368):

Leegth, 127; height, 82; breadth, 53 mm.
" 119 " 71 " 58 mm.

" 128 " 70 " 50 mm.

" 111 " 64 " 50 mm.

44 ILLINOIS BIOLOGICAL MONOGRAPHS [142

The large Anodontas in Crystal Lake are apparently not members of
the original Salt Fork fauna. Neither grandis gigantea or corpulenia are
found anywhere in the Big Vermilion drainage, at least above Middle Fork,

45 miles below Urbana. Since these shells were planted in the lake (see
p. 27) they have evidently thrived and multiplied. Anodonta grandis foot-
iana is parasitic in the glochidial stage on the Johnny Darters {Boleosoma
nigrum, Hankinson, 1908:235) and as this fish also inhabits Crystal Lake
it may have been the medium for the propagation of the alien fauna. That
this fauna should have been so easily detected as alien is due to the method
of examining a stream from its source to its mouth and the distinguishing
of the foreign population is a striking recommendation of this mode of
stream study.

16. Anodonta imbecilMs Say. Paper-Shell.

This beautiful paper-shell occurs abundantly in but one place in the
Salt Fork — near the cement bridge east of Sidney. Here it is of good size,
grass-green in color, the rest periods showing as black longittidinal bands.
The shell is easily known from all others in this State by the very flat
umbonal region which is flush with the upper or dorsal margin of the shell.
The largest specimen in the collection measures 75 mm. in length. Imbe-
cillis was not collected or observed above the cement bridge, 19 miles down
stream from Urbana. It was, also, not seen below the bed at Homer Park
and it appears to inhabit only that portion of the stream between these
points, a distance of about 8 miles. This mussel thrives best on a mud bot-
tom in quiet water and it is not found, normally, on a sand or gravel bottom.
It did not occur in our Sangamon River collections.

All of the individuals from Salt Fork bear evidences of distomid infec-
tion. In nearly all of the valves there are many small pearl-like blisters
about the size of a pin head which are in all cases confined to the posterior
two-thirds of the shell. None were noted near the anterior end.

The species is peculiar and almost unique among naiades in being
hermaphroditic and in carrying the glochidia within the gills until they are
ready for independent life, there being no parasitic stage encysted on fish
as in the case of most Unionidae (Howard, 1914:353). It has an almost
continuous breeding season, glochidia or embryos having been found in
the gills during almost every month of the year. The Salt Fork specimens
were gravid on August 26 and contained well formed glochidia. In this
mode of reproduction imbecillis is parallelled by Strophitus edentulus,
which also passes through its metamorphosis without parasitism.

17. Anodontoides ferussacianus (Lea). Paper-Shell.

This small naiad was found abundantly in but two places — the Middle
Fork and Stony Creek near Muncie. It occurred infrequenty at all other
stations. In the upper Salt Fork, north of Urbana, it was common at one

143] FAUNA OF BIG VERMILION RIVER—BAKER 45

time, near Lincoln Avenue, and in Crystal Lake. Two summers collecting
failed to find it common at the present time at these places. Living speci-
mens were not found in the Salt Fork betwen Spoon River and the big
bend below the natural dam, a distance of over four miles, and it did not
occur even infrequently above Homer Park dam, a distance of 17 miles.

All of the specimens collected are fresh, bright colored shells, greenish
or olive with distinct grass-green rays. All but one specimen were normal in
form and coloration. An individual from the cement bridge station was
thicker than usual, had a short truncated posterior end and somewhat
resembled small specimens of Strophitus edentulus. The beak sculpture was
characteristic of Anodontoides. Pearly growths and abnormalities are
rare in the shells collected. Gravid females were found September 26 and
October 8. It seems to be rare in the Sangamon River, only a stray valve
being found in this river near White Heath.

17a. Anodontoides ferussacianus buchanensis (Lea).

Specimens from the Salt Fork near Muncie and from the Big Vermilion
are referable to this variety, long known under the name subcylindracea
of Lea. The variety in the Big Vermilion drainage is more elongate, more
cylindrical, and has a less height in comparison with the length than in the
typical form. It is also decidedly biangulate behind, a characteristic lack-
ing in the typical form. At the two localities it is associated with, ferussacia-
nus, but at Muncie it is the prevailing form.

18. Arcidens confragosus (Say). Rock-Shell.

This species is a rare inhabitant of the Sangamon River and is not found
in the Big Vermilion River. It was reported from the Sangamon River at
White Heath and Monticello by Mr. James Zetek, about ten years ago.
Professor Smith has not found it at Mahomet during many years of col-
lecting. Recently (September 1920) a single specimen, dead, was picked
up by the writer in the Sangamon River at a point about four miles above
Mahomet, thus establishing its presence above White Heath. It probably
lives sparingly in the river and may inhabit water too deep for examination.
It has been reported from the Sangamon at Springfield (Baker, 1906:74).

19. Lasmigona (Platynaias) compressa (Lea).

This characteristic species is rare in the Big Vermilion River. Several
fine specimens were collected from the station three and a half miles above
Homer Park in rifiles on a sand-gravel bottom. One of these shells is
pathologic, the umbones being almost in the center of the shell, the anterior
end having a strong depression in front of the umbones. The posterior
end is much shorter than usual and is rounded instead of broadly truncate.
The interior shows distomid parasitism near the posterior end with a large
elongated blister near the postero-ventral margin. The pseudocardinal in

46 ILLINOIS BIOLOGICAL MONOGRAPHS [144

the right valve is elongated and thinner than in normal individuals and
the lateral teeth in both valves are scarcely visible. In the left valve there
is an abnormally high, long and narrow tooth under the beak. There is
a pronounced lunule in front of the umbones which is absent in typical
compressa. This species occurs infrequently in mud at Homer Park. In
the Middle Fork a single dead and broken shell was found. No represen-
tatives of this species were found in the Sangamon River.

20. Lasmigona (Lasmigona) costata Rafinesque. Fluted Shell.

This characteristic mussel is fairly common at most stations visited
from bench mark 655 (fifteen miles below Urbana) down the stream to
Middle Fork. It probably inhabits the lower Big Vermilion to the Wabash
River. Living specimens, however, were not seen above the station four
miles above Homer Park dam, nearly 23 miles from Urbana. From Homer
Park down stream it is a common mussel. The individuals are for the most
part fine, large, heavy shells with good clean lustre. The shells from Homer
Park have an olive epidermis beautifully marked with green rays. The
largest individual collected measured 145 mm. in length and was found at
the station three and a half miles above Homer Park dam (Z11116A).

The majority of the specimens of this species are colored light salmon
on the interior of the shell. Pathological individuals are rare in the collec-
tions. One specimen from Salt Fork near Middle Fork, found on a sand
bottom, has a large pearl blister on the posterior margin. A shell from
Homer Park, taken from a gravel bottom, has an injury in the form of a
crack in the shell on the outside which had been repaired on the inside by
the addition of pearly matter forming a long, raised blister, 45 mm. long
and 2 to 5 mm. wide (Fig. Z2)). This nodulous blister reaches almost
to the center of the shell (Z11192 A). Gravid individuals were collected on
October 8 and 13, 1920.

The costata from the Sangamon River, where the species is common,
are somewhat heavier than those from the Big Vermilion River. The
shell is also less high in comparison with its length. Young individuals of
this species from either river di'ainage are very rare, judging by our collec-
tions.

21. Lasmigona (Pterosygna) complanata (Barnes). White Heel-Splitter.
This large, roundish, flat mussel is the most abundant species in the

Big Vermilion River, occurring commonly or abundantly in all parts of the
stream, excepting a small stretch of about five miles near St. Joseph, from
Spoon River to the neighborhood of Danville. It also probably occurs in
equal abundance below Danville. The abundance of this species in Spoon
River and below the station bench mark 655, with the break of five miles
between the beds of living mussels, is strong evidence of the effect of sewage
pollution. In this barren area only empty shells and odd valves could be

145] FAUNA OF BIG VERMILION RIVER— BAKER 47

found after careful search; in one place, below the first bridge south of St.
Joseph, dead, empty shells were abundant, but a careful search, conducted
on two days, failed to discover a single living specimen. The largest and
finest shells occur at Homer Park on a mud bottom in water from two to
three feet deep. The largest shell from this station, a female, measured:
length, 185; height, 130 mm. The species was observed to be gravid on
the 6th of November, in 1918, and on October 8, in 1920.

The shells of complanata are very uniform in general shape, nacre, and
condition. Pearly growths or pathological forms are rare. A few specimens
contained small pin-head pearls and an occasional individual had suffered
slight injury to the posterior part of the shell. The presence of many young
and immature specimens indicates that the species is now breeding well
and that the glochidia are finding suitable fish hosts. Young shells from the
Sangamon River, where the species is abundant, are more of an olive color
and not as green as those from the Salt Fork and other parts of the Big
Vermilion River. Old shells from both drainages are dark brown or black.

22. Alasmidonta (Pressodonta) calceola (Lea).

This species has been recorded by Mr. Zetek from west of White Heath
in the Sangamon River. No specimens were observed during the recent
survey. As far as known it does not occur in the Big Vermilion River.

23. Alasmidonta (Rugifera) marginata Say. Elk-Toe.

In the big Vermilion River, this species is found, infrequently, at most
stations from bench mark 655 as far down as the stream has been examined.
It is not abundant anywhere and common at but three stations — below
the dam at Homer Park, south of Muncie, and in the Big Vermilion. The
first station where living specimens were found is four miles above the
Homer Park dam. Individuals, both above and below the dam, are of
good size and fine color, the characteristic green rays being very brilliant.
Specimens from Middle Fork have many black spots on the shell. The
largest specimen collected from the station two miles above the Homer dam,
measured 80 mm. in length and 45 mm. in height; one from the Big Vermil-
ion measured 96 mm. in length and 48 mm. in height.

Marginata exhibits little evidence of parasitism or abnormalities. One
specimen collected from two miles above the Homer dam had several
pearly growths and blisters indicating distomid infection. Gravid females
were collected on October 8 and 13, 1920.

This species is rare in the Sangamon River, only one specimen being
found at Mahomet during a days search. Marginata is a species more com-
mon in the upper waters of rivers and streams and is not, as a rule, found
in any number in the larger rivers.

48 ILLINOIS BIO LOGIC A L MONOGRA PHS [1 46

24. Amygdalonaias elegans (Lea). Deer-Toe.

But one specimen of this peculiar species was found. This individual
was collected by Professor Smith at the big bend below the natural dam.
The specimen was an empty shell, iron stained outside and inside, and
apparently had not been living for a considerable time. It is typical in
form but not as large as individuals from the larger rivers. The measure-
ments of this specimen are: length, 57; height, 47; width, 27 mm. That
but one specimen of this species should have been collected in this river
drainage seems quite surprising, indicating, probably, that the species
has not been able to become established. It is not found in the Sangamon
River as far as known.

25. Obovaria circulus (Lea).

This nearly circular mussel was collected at but five stations in the
Big Vermilion River; Homer Park below the dam, near Muncie, Salt Fork
near Middle Fork, in Middle Fork, and in the Big Vermilion below Middle
Fork. It is infrequent at the first place and abundant only in Middle Fork,
near its junction with Salt Fork, on a gravel and sand bottom, in fairly
shallow water (September) and in the Big Vermilion (October). In the
Big Vermilion this species is very uniform in shape and size, the ratios of
height to length ranging between 70 and 90 per cent. The shells are usually
nearly circular and have a distinct light yellowish-brown zone at the pos-
terior margin. The nacre is pearly and there are no evidences of discolora-
tion from injury or parasitism. It has not been found in the Sangamon
River.

As in the case of so many of our mussels whose names have become
familiar, this species may have to be changed to that of siibrotunda Rafin-
esque (1820).

26. Actinonaias ligamentina (Lamarck). Mucket.

This mussel was not found above the Homer Park dam. It occurred
infrequently (almost rarely) below the dam on both a mud and a gravel
bottom. In the Salt Fork near its junction with Middle Fork but one dead
shell was found in half a days search for a distance of nearly a mile up
stream. In Middle Fork, between the interurban bridge and the mouth
of the stream, it was fairly common on a gravel and fine sand bottom in
shallow water near riffies. In the Big Vermilion below Middle Fork it is
the most abundant mussel, attaining a length of over 135 mm.

The individuals of this species are all normal specimens; the young and
half-grown shells are greenish with many dark green rays of various width.
Older shells are yellowish with few rays. In an old shell measuring 120 mm.
in length and 75 mm. in height the rays showed but faintly (Z 11216).
Evidences of distomid enfection were rare in this species in the Big Ver-

147] FAUNA OF BIG VERMILION RIVER— BAKER 49

milion specimens. A long narrow pearly blister on the posterior margin of
one specimen might have been due to the presence of the marginal distomid
described by Kelly. Clark and Wilson (1912:62) found this distomid com-
mon as cysts in ligamentina from the Maumee River. A specimen from
the Big Vermilion had a large pearl-like blister at the lower edge of the
posterior adductor muscle scar. (Z 11482 A). An abnormal shell without
the animal, was found in Middle Fork. The posterior end is sharply trun-
cated and bent inward in the right valve and bent outward in the left valve.
The hinge teeth are much more elevated and heavier than in normal speci-
mens. This abnormality was due to an injury received when the shell was
about three years old. It must have lived upward of two years after
receiving the injury, judging by the rest marks on the shell. This individual
measures length, 84, height, 52 mm. (Z11216A).

The Middle Fork and Big Vermilion shells are thick and of good, clear,
peerly- white lustre. The mucket is one of the most valuable shells for the
cutting of button blanks and the Middle Fork and Big Vermilion speci-
mens appear to be of excellent quality for this purpose. The species from
this stream could be used for glochidial infection of fish. Gravid females
were collected on September 26. Ligamentina was not found in the Sanga-
mon River.

27. Actinonaias ellipsiformis (Conrad),

This small naiad occurs rarely at three places in the Big Vermilion
River; below the dam at Homer Park, in the Salt Fork south of Muncie,
and in the Big Vermilion below Middle Fork. Whether this rarity is due
to a real scarcity of the mussel or to the inability of the collectors to find it,
is not known. Diligent search was made in the various places visited,
and the fact that but few specimens have been found by Professor Smith
after years of constant collecting at Homer Park, leads to the conclusion
that the species is rare in this stream.

Ellipsiformis is common in the Sangamon River at Mahomet where it
occurs on a sand and gravel bottom, more commonly on rifiles, where the
current is rather swift. The shells from this station are not large (maximum
size, length, 63, height, 31 mm.) but are very thick, with pearly- white
nacre. The shape is normal, the posterior end being sharply pointed with a
narrow truncation. The color is yellowish or greenish with many dark
green rays, often wavy. Rest periods show as elevated longitudinal ridges,
especially near the ventral mai"gin of the shell. A half-grown specimen,
length 38, height 23 mm., resembles in outline young shells of Actinonaias
ligamentina and if found with that species would probably be so identified.
Ligamentina, however, is not found at Mahomet, and this specimen is
doubtless referable to ellipsiformis. Utterbach's reference of the species
to the genus Nephronaias ( = Actinonaias) is strengthened by the similarity
of these shell characters, which this author has also noted (1916: 142).

50 ILLINOIS BIOLOGICAL MONOGRAPHS [148

In the largest specimen of this species, from the Sangamon River, the
right valve has a large blister parallel with the postero-ventral border,
measuring 17 by 7 mm., and anterior to this blister a group of twenty of
more 'pin-head' pearls. There are several of these small pearls in the center
of the valve. The left valve has a number of 'pin-head' pearls bordering the
pallial line and one larger (1 mm.) black pearl in the center of the valve.
(Z11230 A). These may have been caused by distomid parasites, although
none were observed in the animals of this species. Only one shell in a dozen
were thus afifected.

28. Carunculina parva (Barnes).

This diminutive mussel is characteristic of the smaller tributaries of
the Big Vermilion River. It occurs commonly in the ditch above Urbana
and also in Spoon River. No living specimens were found between the
Spoon River tributary and the station one mile above iron bridge north of
Sidney, a distance of over six miles. From this station to the railroad
bridge east of Sidney, a distance of three miles, this species was abundant
or common. It is infrequent at Homer Park and was not found in the
Salt Fork below Muncie or in Middle Fork. Its normal habitat in the Salt
Fork is on a mud bottom in quiet water. At Homer Park, however, it
occurs sparingly on a gravel bottom in very shallow water.

The species as found in Salt Fork is normal in form but not of large
size, the largest specimen observed, from the railroad bridge east of Sidney,
measuring 30 mm. in length and 18 mm. in height (Z11096). The surface
in the Salt Fork specimens is rayless, the color being brown from the um-
bones to the center of the valve, the balance of the shell being black.
Specimens from Spoon River were smaller than those from farther down
the stream.

Parva occurs infrequently in the Sangamon River at Mahomet on a
mud or sand bottom.

29. Carunculina glans (Lea).

This small mussel, which is much higher in proportion to its length
than parva, is very rare in Salt Fork, where it was found at but three
stations, living individuals being collected only below the dam at Homer
Park. Like parva, it is a species of the smaller tributaries and its norm.al
habitat is on a mud bottom. It was not found in the Sangamon River at
any station examined.

Frierson (1914: 7) has identified Lea's glans with the shell called
Unio (Toxolasma) lividus of Rafinesque, changing the generic, name,
accordingly, to Toxolasma Rafinesque. In this he is followed by Ortmann
(1918:572, 573). Should these names be plainly identifiable from the
original descriptions, these small shells will bear the names Toxolasma
parvum (Barnes) and T. lividum Rafinesque. The writer wonders whether

149] FAUNA OF BIG VERMILION RIVER— BAKER 51

some of these Rafinesquean names have not been adopted too hastily and
whether the fact of the identified specimens in the Poulson collection has
not unduly influenced the references, which, without these identifications,
are not as clear. The test must be, it would seem, a clear case of identifi-
cation from the writings of Rafinesque alone.

30. Eurynia (Micromya) lienosa (Conrad).

The distribution of this small mussel seems to be coincident with that of
Lampsilis luteola in the Big Vermilion and its tributaries. It occurred
infrequently living in the Spoon River. It was not again collected in a
living condition above the iron bridge north of Sidney, seven miles below
Spoon River. At this place only one living specimen was found, and as
far down as the cement bridge it is only infrequent. Between this point
and the station three and a half miles above Homer Park dam no living
specimens were found after diligent search in favorable habitats. It does
not occur in any abundance above Homer Park dam; but below the dam
the species is abundant and of large size. The largest specimens from
Homer Park measure: length, 65; height, 31; breadth, 28 mm. (male);
length, 55; height, 34; breadth, 25 mm. (female). In Salt Fork near its
junction with Middle Fork it is abundant and in the Middle Fork it is
common.

The species varies somewhat in coloration. From Homer Park up-
stream the color is black or dark brown with rarely faint indications of
rays. Specimens from Middle Fork and Salt Fork near Middle Fork are
more of a chestnut color, slightly reddish, often with quite distinct rays.
By arranging the different lots by consecutive stations down the river it
may be at once observed that there is a marked and striking increase in
size, the increase being in some cases as much as 50 percent between Spoon
River and Homer Park. The Homer Park shells are on the average some-
what larger than those from farther down the stream and from Middle
Fork.

The shells of many individuals of this species contain pearly growths
in the form of blisters and 'pin-head' pearls. These occur near the margin
of the valve, more frequently near the posterior end near the siphonal
region. Gravid females were collected at Homer Park on July 30.

Lienosa was not collected from the Sangamon River and no records
have been seen of its occurrence in that stream.

31. Eurynia (Micromya) iris (Lea).

A broken valve of this species was found in Middle Fork below the
interurban bridge. The rays are distinct and the shell seems typical.
It measures, length 37, height 21 mm. It appears to be a very rare shell
in the parts of the Big Vermilion examined but may be more abundant

52 ILLINOIS BIOLOGICAL MONOGRAPHS [150

below the stations examined where the river is larger. Iris has not been
recorded from the Sangamon River.

32. Lampsilis luteola (Lamarck). Fat Mucket.

The mussel known among fishermen as the fat mucket is common or
abundant almost everywhere in the Big Vermilion and Sangamon rivers.
It has been killed by the sewage of the Salt Fork from St. Joseph to bench
mark 655, a distance of five miles down the stream, but the number of dead
and empty shells found almost everywhere between these points indicates
that at one time, not very remote, it was common continuously from Spoon
River, where it now lives in some abundance, to the Wabash River. Below
the dam at Homer Park it is very common and of large size, and this abun-
dance continues down the stream and was also noted in the tributary Mid-
dle Fork. At Mahomet on the Sangamon River it is also abundant.

There is great variation both in form and coloration among the shells
of this species in all of the habitats examined. The male shells are usually
pointed at the posterior end and are elongated and somewhat compressed.
From this form they vary by being quadrate in outline with a distinctly
plow-shaped posterior end, corpulent and almost cylindrical, or flattened
and oval, in this form greatly resembling Actinonaias ligamentina, from
which they may be distinguished by the numerous double-looped ridges
on the umbones. The female shells do not differ so greatly in shape, the
post-basal swelling for the accommodation of the enlarged branchial marsu-
pium giving more uniformity to the shell, the variation being principally
in the width o f the shell, which in old specimens is very pronounced. Male
shells greatly predominate in the collections. In color there is every
gradation between a bright yellow shell with distinct, narrow dark green
rays, to a shell that is dark yellowish or brownish without rays or with the
rays only faintly developed. A few specimens are dark brown or even
pinkish with narrow, greenish rays. Young shells are very brightly rayed,
the rays being dark grass-green on a light yellowish background, forming a
beautiful surface ornamentation. The rays on the adult shells may be
narrow or broad, or the broad rays may be made up of many fine rays,
which may also be a trifle wavy. The nacre in all specimens examined from
the two rivers here considered is pearly white, unmarked by color of any
kind. The largest specimens seen occur at Homer Park; measurements of
these are given below:

Length, 110; height, 59; breadth, 35 mm. Male

" 116 " 68 " 40 mm. Male

" 116 " 69 " 46 mm. Female

" 100 " 65 " 48 mm. Female

Pearly growths were observed in many of the specimens collected.
Occasionally a few pin-head pearls occur in a valve but the greatest number

151] FAUNA OF BIG VERMILION RIVER— BAKER 53

of abnormalities consist of cyst-like pearly growths near the posterior
end, in or near the adductor muscle and pallial line. These may occur
in one or both valves. Occasionally the anterior adductor scar is almost
wholly changed in character by these abnormal growths. In several shells
the space between the pallial line and the margin of the valve is stained
purple or brown, accompanied by few or many blisters of various sized.
The posterior end of the shell may also be affected so that it ceases to grow,
forming a blunt posterior end marked by blisters and discolorations.
Whether this condition is due to distomid infection as mentioned by
Wilson and Clark (1912 63) as occurring in Actinonaias ligamentina from
the Maum^ee River is not known definitely, but the inference is strong that
it is. It is noteworthy that the largest number of individuals affected by
parasitism or other injurious agencies occurred in the upper part of the
Salt Fork above Homer Park dam. Specimens collected from stations
below the dam, including Middle Fork, were as a whole remarkably free
from pearly growths or discolorations. Occasional individuals from Maho-
met, on the Sangamon River, have round, pin-head pearls in the shell.
Though no distomids were observed in the animals of luteola it is quite
probable that these pearls and blisters, which were observed for the most
part in empty shells, were caused by distomid larvae, possibly the mar-
ginal cyst described by Wilson and Clark (1912:62). Many young shells
of luteola were collected and many more observed, indicating that the
species is breeding freely from the Homer Park section of the stream down-
ward. No young shells were seen above the Homer Park dam. Gravid
females were observed at several places in September and October.

A specimen collected at Mahomet (Z11223 A) somewhat resembles
Actinonaias ligamentina in the form of the shell, absence of strong umbonal
markings and heavy hinge teeth; it seems to be one of those individuals
which has led many students to say that ^^ligamentina runs into luteola.'^
The shell is free of all abnormalities and has a clear pearly-white nacre.
It measures, length 87, height 54, breadth 30 mm.
33. Lampsilis ventricosa (Barnes). Pocket-Book.

This large fine species was not collected in Salt Fork above the station

about two miles north of Sidney, 16 miles below Urbana. As it does not

occur in Spoon River (as far as known from our collections) it is probably

a species that does not inhabit the smaller tributary streams. From the

station mentioned as far down stream as the river has been examined,

including Middle Fork, ventricosa is common or abundant in most places.

Below Homer Park dam it is abundant and of large size, and more or less

ponderous. The largest female shell was found at Homer Park and the

largest male shell in Salt Fork near Middle Fork. These shells measure as

follows:

Length, 138; height, 60; breadth. 56 mm. Male
" 117 " 60 " 60 mm. Female

54 ILLINOIS BIOLOGICAL MONOGRAPHS [152

Shells from the Sangamon River are equally large.

The shape of the shell is fairly constant and there is little variation
except in the females, which are rounder posteriorly and have a large post-
basal swelling for the enlarged marsupia. In color the Big Vermilion shells
are all yellowish with dark green rays on the posterior slope. No specimen
was seen in this river that was rayed all over. Very old specimens are
entirely rayless. At Mahomet, on the Sangamon River, the yellow shell
occurs and also another form in which the shell is yellowish-green with
bright, grass-green rays, often of considerable width. One specimen is
in outline like Actinonaias ligamentina and the surface is densely covered
with dark green rays. Two other specimens have green rays on a pink
background, have pink hinge teeth, and the whole interior of the shell is
pinkish. These bright colored shells are the form called occidens by Lea.

These pink shells and the specimens with the numerous green rays are
so strikingly different from the ventricosa as found in the Big Vermilion
and also from the other shells found in the Sangamon, that the name
occidens might be retained for these shells for ecological purposes. The
color is not an age stage, for young yellow ventricosa were found associated
with these distinctly rayed forms, and the rayed forms were collected
at Mahomet and were not found at White Heath in the Sangamon. The
river below Mahomet has not been carefully searched, however, and the
occidens form may occur in some of this unexplored territory. It is also
to be noted that, as far as the material from these two rivers is concerned,
the beak sculpture of the occidens type of shell is very much larger and
coarser than in the ventricosa iyp^ {ZWlll). This form falls under the
group of individuals called mutations by DeVries and others.

Ortmann (1918:583) makes ventricosa a variety or race of ovata (Say).
As far as the authors' experience goes, this seems unwarranted, the two
species being as easily separable as many other closely allied species.
Individual specimens from Illinois localities approach ovata in that the
posterior ridge is somewhat accentuated, but no specimens have been seen
that could not be placed readily in one species or the other. Ovata is
reported from the Ohio River in Illinois by Marsh but this species is more
southern in its distribution, reaching its maximum development in Ala-
bama and Tennessee. Ventricosa is a more northern species attaining its
maximum development in the rivers of Illinois and Indiana.

Ventricosa is subject to the attack of distomid worms as well as to
parasitism by mites (Unionicola) and the material of this species from the
Big Vermilion have suffered more or less from this cause. Blisters, pin-
head pearls and various abnormalities occur in many individuals. Among
the shells collected at the station one mile above iron bridge north of Sid-
ney there are several individuals of this sort. One has a large blister ex-
tending nearly the whole length of the ventral margin and covering the

153] FAUNA OF BIG VERMILION RIVER— BAKER 55

space between the margin and the pallial line. This was evidently formed
by distomid parasites (Z11058 A). Another specime i from the same
lot (Z11058 B) has a large group of pearls (over 100) filling the space
between the ventral margin and the pallial line over an area of about two
inches. Other specimens have an abnormally thickened ventral margin
in addition to blisters, indicating the presence of some irritating material
between the shell and the mantle lobes, probably mud or sand (Z11149
A). A specimen from Mahomet had suflFered a curious injury which had
caused a large blister extending from the upper third of the posterior end
to the center of the ventral margin, a distance of about three inches. This
was plainly due to the presence of a quantity of mud getting in between
the shell and the mantle, a small amount of this material being retained
under the cylindrical blister on the ventral margin. The outside of the
shell was so perfectly repaired that there was no evidence of the pathologi-
cal condition within the shell (Z11202 A). Fig. 35. This injury was only
in the left valve, the right valve being normal.

Nearly all of these pearly growths, blisters, and abnormalities have
been noted to occur almost universally without the pallial line, between
this muscle attachment and the ventral, anterior, and posterior margins
of the shell. This limited area is easily understood when it is remembered
that the adductor muscles and the pallial line form a barrier to the entrance
of foreign material because the muscles at these points are firmly attached
to the shell and parasites or foreign material cannot readily gain entrance
to the interior of the animal between the mantle and the shell. This is
true of all the shells of other species examined from these two river systems.
It was noted that pearly growths and abnormalities were rarer in the
shells from the Middle Fork and from Salt Fork near the Middle Fork
than from Homer Park and up the stream from this habitat. The shells
from the Sangamon River at Mahomet were much less subject to parasitism
or abnormalities than those from the Big Vermilion River.

Young shells of ventricosa were common only in Middle Fork, in Salt
Fork near Middle Fork, and in the Sangamon River at Mahomet. No
young shells were collected from the stream above the dam at Homer Park.

Gravid females were observed on Septem.ber 26 in the Salt Fork near
Middle Fork in the act of spawning. The shells were buried in the sand,
only the tips of the siphons showing above the general level of the bottom
of the stream. The bright yellow siphon fringes, which are enlarged during
the spawning period, were observed to wave about with graceful undula-
tory motion. The movements were more or less intermittent, a waving
period being followed by a resting stage. These mussels had well formed
glochidia and were evidently discharging the embryos from the marsupia.
These glochidia are of the bookless type and develop in the mouth of fish
and the undulatory motions possibly attract these fish which may be thus

56 ILLINOIS BIOLOGICAL MONOGRAPHS [154

more easily infected with the glochidia, as suggested by Wilson and Clark
(1912:13, 14). This undulatory motion of the siphon fringes is so striking
that it at once attracts the attention of an observer,even if this person is
not interested in the study of these creatures. Its habit of burying itself
so deeply in the mud or sand of the bottom (four to seven inches) renders
this species difficult to collect, especially if the sand be packed rather hard
and the water is eighteen or more inches in depth, as frequently occurs.
This condition was found in several of the Salt Fork habitats. Gravid
females were also collected from other localities in October.

34. Lampsilis multiradiata (Lea).

This handsome mussel occurs rarely at but three stations, Homer Park,
near Muncie, and in Middle Fork. It is common in the Big Vermilion
below Middle Fork. The specimens from the Salt Fork Stations are all
small for the species, being not over half the size of individuals from
White River, near Muncie, Indiana, and the hinge teeth are much weaker
than those from Indiana streams. This is especially true of the pseudo-
cardinal teeth which are narrower and more elongated than the speci-
mens examined from other streams. Individuals from Homer Park
are brilliantly rayed, grass green on a yellowish ground. Middle Fork
specimens are not as brilliantly rayed, the general color being yellowish
with scanty rays. The largest specimens collected measure as follows,
a specimen from Muncie, Indiana also being included for comparison:

Length, 34; height, 22 mm. Homer Park.

" 43 " 31 mm. Middle Fork. Male

" 69 " 55 mm. Big Vermilion. Female

" 85 " 64 mm. Muncie, Ind. Female

This species is rare in Illinois waters, judging by the few records
available. These indicate, however, a wide range over the State, from
Cook County to southern Illinois (Baker, 1906:64). Ortmann (1918:584,
1920:309) adopts the na.Tne fasciola of Rafinesque (1820) for this species,
stating that the original description of Rafinesque is definite enough to
indicate without much question a shell of the multiradiata type. If
this be so, then Lea's very appropriate name must give place to the
one used at an earlier date by Rafinesque. The species has not been
recorded from the Sangamon River.

35. Lampsilis anodontoides (Lea). Yellow Sand Shell.

This fine shell, which is usually abundant throughout Illinois waters,
was collected at only three stations in the Big Vermilion River, all of
which were below the dam at Homer Park. The largest and finest in-
dividuals occur at Homer Park and the smallest were collected in
Middle Fork. Specimens from the first named station have a yel-

155J FAUNA OF BIG VERMILION RIVER— BAKER 57

lowish shell with little or no indication of rays. Middle Fork specimens
include yellowish shells without rays, as well as, more rarely, individ-
uals with rather bright, wide rays, approaching in this respect the
related species fallaciosa of Simpson. The largest specimens from the
two localities measure as follows:

Length, 124; height 57; breadth 47 mm. Male, Homer Park.

90 " 41 " 33 mm. Female, Middle Fork.

The nacre is tinted with pinkish or salmon color. Pearly growths
are not uncommon in specimens from the Big Vermilion. These are
in the form of blisters and pin-head pearls, which are usually confined
to the margin of the shell between the pallial line and the external margin
of the valve. One individual from Homer Park had the entire area between
the pallial line and the ventral margin of the shell abnormally enlarged
and thickened, due possibly to the presence of distomid larvae and to some
extent to the intrusion of small amounts of soil between the mantle and
the animal (Z11147A). Specimens from Middle Fork are, as a rule,
free from pearls and abnormal growths. Gravid females were found on
September 26 in Middle Fork, and on July 30 in Salt Fork at Homer
Park. Anodontoides has not been recorded from the Sangamon River at
Mahomet or in the other places examined.

The early writers. Say, Conrad, and others, have identified this species
with the Elliptio teres of Rafinesque (1820) and if the shell is clearly identi-
fiable from the description of Rafinesque the familiar name of Lea must
become a synonym.

36. Truncilla (Pilea) perplexa rangiana (Lea).

Four specimens of this race of perplexa are in the naiad collection
of the Museum of Natural History, University of Illinois, collected
by Mr. A. A. Hinkley, in the Big Vermilion River at Danville. Three
are females and one is a male. They are much smaller than specimens
from Florence, Alabama, where the species attains its greatest develop-
ment. The miale and largest female shell measure as follows, corre-
sponding measurements being also given for the Alabama shells.

Length, 41; height, 29; breadth, 22 mm. Danville, male, Z3770.

" 44 " 34 " 20 mm. Danville, female, Z3770.

" 55 " 43 " 32 mm. Alabama male, Z3947.

" 73 " 53 " 35 mm. Alabama female, Z3947.

As no specimens of Truncilla were found in the Big Vermilion or its
tributaries as far down as Middle Fork, which is but a few miles west
of Danville, this species evidently does not inhabit the stream above the
locality from which Hinkley collected his shells. Truncilla perplexa
as well as its variety rangiana is known in Illinois only from the Ohio

58 ILLINOIS BIOLOGICAL MONOGRAPHS [156

and Wabash rivers. The small size of the Danville specimens may be
due to the fact of their inhabiting the upper part of their distributional
range, in a rather small river. Rafinesque's name torulosa has been
used by Conrad, Agassiz, Reeve, and Ortmann for Truncilla perplexa
and if the original description is clear enough to reasonably fix the shell
as Lea's perplexa this form will stand as Truncilla torulosa rangiana
(Lea). Neither perplexa or rangiana are known from the Sangamon River.

FAMILY SPHAERIIDAE

37. Sphaerium striatinum (Lamarck). Variety.

This finger-nail clam occurs abundantly only in the ditch north of
Urbana. It also occurred infrequently in Spoon River, and near the
iron bridge in Salt Fork north of Sidney. Dead specimens were
found at bench mark 655 and below the Homer Park dam.

Sterki remarks of this form "possibly a form of striatinum Lam;
apparently the same as a Sphaerium from Iowa and other places which for
many years has been left unnamed, except in manuscript. It still seems
impossible to say where the limits of striatinum are." The writer fully
agrees with Dr. Sterki regarding the variation in this common species.
This variety appears quite distinct from the ordinary specimens of the
species and it is to be hoped that Dr. Sterki will give it a name. It is
noteworthy that the form occurs commonly only in the drainage ditch above
Urbana and was not found in any numbers below this place. A Sphaerium
identified as striatinum byZetek some years ago, occurring in Crystal Lake,
was doubtless this form.

38. Sphaerium stamineum (Conrad).

This small mussel is abundant at several places in Salt Fork above
Sidney. It occurs only infrequently at two other places. Reported by
Zetek from Crystal Lake. Some of the shells are typical but others are
similar to straitinum and it is difficult to separate some specimens from that
species. At the station called bench mark 655 a form of this species
occurs that is somewhat like the variety forbesi Baker, described from
Mason County, Illinois.

39. Sphaerium solidulum (Prime).

An abundant species in the Spoon River and in the Salt Fork above
Sidney. Occurs rarely at other places. Reported by Zetek from Crystal
Lake. On the whole this species is quite uniform in characters,

40. Sphaerium species.

A Sphaerium occurs abundantly in Salt Fork below the Homer Park
dam which is thought by Sterki to be possibly a new species. It is related to
solidulum but is larger and more robust (mus. no. Z11383). It lives in

157] FAUNA OF BIG VERMILION RIVER— BAKER 59

shallow water on a more or less rocky bottom a short distance below
the dam.

41. Musculium transversum (Say).

This fragile finger-nail clam is apparently rare in Salt Fork and its
tributaries, occurring only in Salt Fork near Sidney, It has been reported
from Crystal Lake by Zetek. The specimens are typical.

42. Musculium truncatum Linsley.

This very small clam was found only in the old river bed near the
cemetery north of Urbana. Here it is common and typical.

43. Musculium partumeium (Say).

Reported by Zetek from Crystal Lake. It has not been found recently.

44. Pisidium compressum Prime.

This small clam occurs in abundance only at the iron bridge north of
Sidney. A few specimens were found below the natural dam in Salt Fork.
The individuals are quite typical.

45. Pisidium kirklandi Sterki.

A single specimen of this species was foumd at the iron bridge north
of Sidney. It is apparently very rare.

46. Pisidium splendidulum Sterki.

A few specimens of this tiny species were found in the Salt Fork near
Muncie. The individuals are young and not quite typical.

The distribution of the Sphaeriidae in Salt Fork is interesting and
significant in connection with the sewage pollution of the stream. It
will be noted that no Sphaeriidae were found in the Salt Fork between the
Urbana ditch and the station called natural dam, 14 miles below Urbana.
These mussels are characteristic mud dwellers and their absence from the
intervening territory in the stream is striking evidence of the unfavorable
conditions on the bottom. They were not found in abundance above
bench mark 655. It will also be noted that Sphaerium appears, living,
before Pisidium and is also found in abundance higher up the stream.
This may mean that Sphaerium is better able to withstand these adverse
conditions than Pisidium, which is more of a mud dweller. Pisidium is
much less common everywhere in the stream and but one species occurs
abundantly at one place (Table IV).

FAMILY PLEUROCERIDAE

47. Pleurocera elevatum (Say).

This long-spired snail is rare or wanting in most parts of the Big
Vermilion River. Two white, weathered specimens of this species were
found below the natural dam in Salt Fork. They may have been washed

60 ILLINOIS BIOLOGICAL MONOGRAPHS [158

from a post-glacial fossil deposit. None were found either alive or dead
in any other part of the Big Vermilion examined, excepting the two speci-
mens mentioned above.

In the Sangamon River, at Mahomet, elevatum is very abundant on a
sandy bottom in water a foot in depth (low water period). The speci-
mens are large (maximum length 35 mm.) and vary in color from light yel-
lowish to black or dark chestnut with a light zone below the suture. The
characteristic peripheral keel is developed on the majority of specimens
and ascends the spire just above the suture. In a few individuals this keel
is absent and the body whorl is rounded. Many intermediate forms occur
connecting these rounded shells with the more distinctly keeled forms.
Usually there are from one to six spiral ridges on the base of the shell extend-
ing longitudinally parallel with the peripheral keel. The strongly carinate
whorls of the young shell are six in number and these lose their sharpness
more or less abruptly on the seventh whorl. No individuals were seen in
which the protoconch or nucleus of the first whorl was preserved. Young
and half-grown specimens have two brown bands on the spire whorls and
four bands on the body whorl and base of the shell. This species also occurs
in the Sangamon at White Heath, and it has been found in several Pleisto-
cene deposits near Mahomet, the white shells of the fossils being washed
out of the bank by periods of high water.

48. GonJobasis livescens Menke.

This river snail occurred in but four places in Salt Fork, It was
infrequent at bench mark 655 and abundant just below the dam at Homer
Park and near Muncie. It is abundant in the big Vermilion below Middle
Fork. It has been identified by local naturalists as Goniohasis pulchella
Anthony and has been reported as this species by other students as from the
Big Vermilion River (Baker, 1906:98). Specimens were sent to Mr. Calvin
Goodrich, who is making a study of this family, and were identified as
livescens. I quite agree with Mr. Goodrich that they are livescens rather
than pulchella. Young and immature specimens are banded like Goniobasis
depygis Say.

Goniohasis semicarinaia Say is reported from the Big Vermilion River
by Marsh (Baker, 1906:98) but no specimens have been seen from this
stream. The species may live in the river below Danville or the citation
may be based on long-spired livescens.

FAMILY AMNICOLIDAE

49. Pomatiopsis lapidaria (Say).

This tiny snail is abundant in small, cold streams flowing into the
Sangamon River. At one locality, about three-fourths of a mile below
Mahomet, it was extremely abundant in a small brook about two feet

159] FAUNA OF BIG VERMILION RIVER— BAKER 61

wide and not exceeding six inches in depth. The bottom was of mud and
fine sand. The largest specimen measures 7 mm. in length. This spe-
cies was not collected from streams flowing into the Big Vermilion River.

50. Amnicola limosa (Say).

This common Amnicola occurred at three stations, only two of these
yielding living specimens. At the station above the iron bridge north of
Sidney, the shells were secured by sweeping the vegetation bordering the
shore with the Walker dredge. The lifnosa from the Salt Fork are rather
corpulent but are not as globular as the variety known as porata (Say). It
is noteworthy that this species (and in fact the genus) should be found only
in a stretch of the stream three miles in length, from 16 to 19 miles below
Urbana. None were seen below the dam. Individuals were rare and
difficult to find. Near the cement bridge east of Sidney, dead shells
occurred among water plants {Nymphaea advena) but no living specimens
could be found. Limosa was not collected at the stations in the San-
gamon River.

51. Amnicola (Cincinnatia) clncinnatiensis (Anthony).

This Amnicola occurred rarely, but living, at but two stations in the
Salt Fork, 16 and 17 miles below Urbana. The largest and only adult
individual collected measures 6 mm. in length. Several half-grown shells
were found. About two years ago Mr. James Zetek found cincinnatiensis
near St. Joseph. A careful search of this region, both in the Salt Fork
and in the small streams flowing into Salt Fork, failed to produce a single
specimen of this species. During the intervening ten years the species
appears to have died out, either from the effect of sewage pollution or
from some other unknown cause.

FAMILY VIVIPARIDAE

52. Campeloma rufum (Haldeman).

The Campelomas of the Big Vermilion all appear to be referable to
Haldeman's rufum. The shell is more or less pinkish, especially on
the spire. One specimen from the iron bridge north of Sidney resembles
Haldeman's figure 1 on plate 3 of the Monograph, which is the type
of rufum. Specimens from Homer Park, below the dam, are strongly
suggestive of integrum (Say), many of the individual shells being like
Binney's figure 96 which represents Say's integrum. With these are
short-spired shells recalling the obesum of Lewis as figured by Binney
(figure 95). These are not quite like the figures of Lewis (1875, pi. 23,
figs. 4-5) which are rather broader. Variation in rufum seems to parallel
that of integrum in the length of the spire. Typical integrum has a white
aperture and a bright green shell devoid of the peculiar pink tint of rufum.

62 ILLINOIS BIOLOGICAL MONOGRAPHS [160

The rujum from Homer Park have the spire whorls more or less gibbous,
strongly shouldered, the first three whorls seeming to be telescoped into
the later whorls. In this respect they resemble integrum ohesum as sug-
gested above. Measurements of a few of the Homer Park specimens are
given below (Z11168):

Length, 37; breadth, 24; aperture length, 21; breadth, 14 mm.
" 33 " 22 " " 19 " 13 mm.

" 30 " 21 " " 18 " 12 mm.

To this and other lots of Campelomae from the Big Vermilion River the
statement of Lewis may be well applied: "These and many other forms in
my collection, all part of a series, go far to show that it is unsafe to attempt
to decide the limits of species from a few individuals" (1875:337).

The distribution of this species in Salt Fork is interesting and sug-
gestive. Living specimens, small and few in number, were found over two
miles upstream in Spoon River. Dead, mostly old and bleached shells,
were collected at nearly all stations in the Salt Fork, but living shells of
rufum were not seen above the station two miles north of Sidney. Here
only one living specimen could be found. A mile farther down the stream
another living specimen was collected. The presence of so many dead
shells with so few living individuals above the Homer Park dam indicates
clearly an unfavorable environmicnt. There are many normal and favor-
able habitats for this moUusk in this stretch of nearly twenty miles in Salt
Fork and the unfavorable agencies must be wholly those contributed by
Man — the disposal of sewage and other wastes by means of this stream.
Below Homer Park dam the species is abundant and as fine as can be
found anywhere. Rufum is rare on a sand and gravel bottom and abundant
on a mud bottom.

The Campelomae from the Sangamon River at Mahomet are also refer-
able to rufum. The spire is longer and the shell narrower, however, than in
the Salt Fork specimens, and there is no tendency to vary toward the
obesum form of shell. The interior of the aperture is slightly pinkish. One
specimen from Mahomet has a very heavy shell recalling the subsoUdum
of Anthony, a common species in most parts of Illinois but absent from
either of the rivers under consideration. Reversed individuals are rare,
only one specimen being found in the Sangamon River, a mile below
Mahomet. This is a young individual.

The air-breathing snails, belonging to the genera Physa, Ferrissia,
Planorbis, and Galba, are better able to withstand the ill effects of sew-
age and other stream pollution than are their relatives, the snails and
clams that take their oxygen directly from the water (dissolved oxygen).
They were therefore found in Salt Fork in places where the water breathers
were entirely wanting, as at St. Joseph and the first stations below. It has

161] FAUNA OF BIG VERMILION RIVER— BAKER 63

been observed in other places, notably in the Genesee River, at Rochester,
N. Y., where sewage pollution was at one time very severe, that these
pulmonate water snails were the last to succumb to the toxic influences
of pollution and they have been known to live in water that was filled
with putrescent matter and also in water strongly impregnated with arsenic.
When these snails begin to disappear, conditions must indeed be deplorable.

FAMILY ANCYLIDAE

53. Ferrissia rivularis (Say). River Limpet.

The tiny limpet-like shell known as Ancylus {Ferrissia) rivularis was
very abundant in parts of Salt Fork, its usual habitat being the inside of
empty valves of the naiades. The individuals are large (6.4 mm. in length)
and fine and apparently normal in form. This species was found alive at
St. Joseph where pollutional conditions are bad, and was also abundant
at the stations lower down the stream where living mussels or pectinibran-
chiate snails were very rare or absent. Ancyli were not observed in Spoon
River, in Middle Fork, or below the dam at Homer Park. Found by Mr.
Zetek in the Sangamon River at White Heath.

54. Ferrissia tardus Say. River Limpet.

This Ancylus is in the author's collection from White Heath and
Monticello, Sangamon River, and from the Salt Fork near Urbana, col-
lected by Mr. Zetek and identified by Dr. Bryant Walker. No tardus
were collected during the present survey.

55. Gundlachia meekiana Stimpson.

Specimens of this characteristic mollusk are in the author's collection
from Crystal Lake, Urbana, collected by Mr. Zetek in August 1904, and
identified by Dr. Walker. None were seen during the present survey.

In the catalogue of the Mollusca of Illinois (Baker, 1906: 101, 102)
Ancylus {Ferrissia) shimekii Pilsbry is recorded from Salt Fork, Urbana,
and Ancylus {Laevapex) kirklandi Walker from Crystal Lake, Urbana.
These species were contained in the collection of the State Laboratory
of Natural History. A recent examination of the material upon which
these records are based indicates that there has been an error in the habitat
given. None are from the Salt Fork or Crystal Lake. Ancylus kirklandi,
identified by Walker, is in the laboratory collection from Havana, Illinois
River (Nos. 13792, 13811, 24123) and Elizabethtown, Illinois (No. 24527).
Ancylus shimekii (No. 24541) is in a bottle with Ancylus rivularis, both
identified by Walker. The bottle is without locality and no record was
found in the laboratory catalogs of the specimens bearing this number.
It is evident, therefore, that these species of Ancylus must be eliminated
from the list of the fauna of Salt Fork.

64 ILLINOIS BIOLOGICAL MONOGRAPHS [162

FAMILY PHYSIDAE

56. Physa gyrina Say. Tadpole Snail.

This snail is usually abundant wherever found. In the old cut-offs
of the Salt Fork above Urbatia, as well as in the ditch north of Urbana,
it is abundant and quite typical with long, slender shell and spire, the
immature individuals with a short, dome-shaped spire. Below St. Joseph,
where it occurs sparingly, the shell is broader and even in adult ''specimens
the spire is more or less dome-shaped. Gyrina is more common above than
below the dam at Homer Park. The species is more characteristic of slow-
moving, pond-like bodies of water than of larger streams. It occurs also
in Stony Creek near Muncie, in a small pond near Middle Fork, and in
the Big Vermilion below Middle Fork.

57. Physa crandalli Baker.

Specimens of a Physa with a shouldered whorls, a wide body whorl,
the shell thick and heavy for the genus are referred to crandalli. This
mollusk is abundant in the drainage ditch above Urbana associated with
gyrina. None were found in the cut offs of the old stream and the species
probably does not inhabit the pond-like habitats in which gyrina is usually
found, preferring running water. The specimens referred to sayii Tappan,
from Urbana (BakeT, 1906:99) are also this species. It is noteworthy that
living specimens of this species were collected at St. Joseph where polluted
conditions are bad. None were found below the iron bridge one mile
north of Sidney, or in any part of the Salt Fork below this point. Charac-
teristic specimens were collected in the Big Vermilion below Middle Fork,
on a stony bottom in riffles.

FAMILY PLANORBIDAE

58. Planorbis (Helisoma) trivolvis Say. Wheel Snail.

This species of wheel snail is apparently not common in Salt Fork, only
scattering specimens being found along the stream. It occurred more
abundantly in a small stream, dry in summer, which runs through low.
swampy ground on the east bank of Salt Fork south of the interurban
bridge at St. Joseph. Living trivolvis were not found in the stream above
the first bridge below St. Joseph, twelve miles below Urbana. Even this
air-breathing snail seems to be unable to live in any abundance in the
polluted water of South Fork.

59. Planorbis (Helisoma) pseudotrivolvis Baker.

This recently described wheel snail (Baker, 1920:123) occurs abun-
dantly in the old stream bed (cut-offs) of the Salt Fork near the Woodlawn
cemetery, Urbana, and it is here the predominating species of the genus,
true trivolvis being rare. The differences between this species and trivolvis

163] FAUNA OF BIG VERMILION RIVER— BAKER 65

have been clearly pointed out in the paper referred to above. Two speci-
mens, one living, referable to this species were found in Salt Fork about
two miles below St. Joseph, associated with typical trivolvis.

60. Planorbis (Helisoma) antrosus Conrad.

A single, small, bleached shell of this species was found near the cement
bridge northeast of Sidney. It may have been washed from a Pleistocene
fossil deposit near by. None were collected living.

61. Planorbis (Gyraulua) parvus Say.

This small species was found in limited number associated with Planor-
bis trivolvis and Galba parva in the low ground subject to spring overflow
south of the interurban bridge at St. Joseph. It was typical in form as
compared with authentic specimens collected near Philadelphia by Dr.
H. A. Pilsbry. None were found in Salt Fork or in the Sangamon River.

FAMILY LYMNAEIDAE

62. Galba parva (Lea).

This tiny pond snail was found at but one place in the Salt Fork. A
dead shell was obtained at the iron bridge, a mile north of Sidney. In
the low ground south of the interurban bridge at St. Joseph before referred
to, parva occurs abundantly in the bed of a small stream which has water
in it only in spring and early summer. On the Sangamon River, this species
was noted in abundance on wet mud flats bordering the margin of the
stream. This locality was about three-fourths of a mile below Mahomet.
The polluted water at St. Joseph appears to have little effect on this species
or the other pulmoniferous mollusks associated with it. It is probable
that at the times of high water so much oxygen is mixed with the upper
layer which overflows these low places that the ill effect of sewage pollution,
from decomposition of organic matter, is so reduced in quantity and quality
as to be little noticed by these air-breathers.

63. Galba humilis modicella (Say).

This is usually a very common species where it occurs at all. It is rare,
however, in the Big Vermilion, scattered specimens, mostly dead shells,
being found at four stations. It was collected living in the drainage ditch
above Urbana, in Crystal Lake (Zetek), and a large typical individual was
found alive on the mud bordering the stream a short distance below the
mouth of Spoon River in Salt Fork west of St. Joseph. The species occurs
sparingly in the Sangamon River associated with Galba parva at the locality
mentioned under that species. In a small pond in the gravel pit north of
the interurban tracks west of the Middle Fork, modicella occurs living
among cat-tails {Typha) associated with Physa gyrina.

66 ILLINOIS BIOLOGICAL MONOGRAPHS [164

64. Galba obrussa (Say),

Obrussa occurs in Stony Creek near Muncie, collected by Mr. John R.
Malloch, May 29, 1919. The specimens are typical though small.

65. Galba caperata (Say).

This species occurs abundantly in swampy woodlands bordering the
Salt Fork about three miles north of Urbana, the ground in dry weather
being almost paved with the dead shells of this snail. Some of the snails
escape the dry period by crawling into cracks and holes and there hiber-
nating. This species has not been found in or near Salt Fork or the other
tributaries of the Big Vermilion River. Because of its preference for small,
summer-dry ponds and pools this Galba will not be found, probably, in any
part of the streams herein considered.

165] FAUNA OF BIG VERMILION RIVER— BAKER 67

POLLUTION OF SALT FORK BY SEWAGE AND
MANUFACTURING WASTES

GENERAL NATURE OF STREAM POLLUTION

Stream pollution may be broadly divided into two main divisions:
contamination by organic sewage from cities and towns and by chemical
wastes from factories and mines. Both are inimical to life but the latter
is especially fatal to animal life, causing wide stretches of otherwise
fertile streams to become veritable deserts. Organic sewage, in a crude
or highly concentrated form, is also very injurious, effectually eliminating
most forms of life from the polluted body of water.

The importance and seriousness of the problem of stream pollution in
its effect on the life of the rivers and streams into which the contaminating
material is discharged has not until very recently been given the attention
the subject demands. The diminishing fish supply, and in many places
the very objectionable physical character of the polluted waters, have
caused the authorities of several states to pass laws governing the discharge
of these wastes into streams and the establishment of penalties for dis-
regarding these laws. New York and Massachusetts have led in the fram-
ing of these laws and other states are following the good example set by
these two older commonwealths, where the conditions seem to have reached
a maximum of harmfulness (see Ward, 1918, 1919).

During recent years stream pollution has enormously increased and the
problems arising from this condition have been investigated by many
biologists and sanitary engineers. The former have studied the problem
from the viewpoint of its effect on the useful animal life, especially fishes
and river mussels, and this phase probably bears as close a relation to
human welfare as any other. Of course, from the standpoint of health, the
pollution problem is of paramount importance because of its bearing on
such diseases as typhoid fever which may be caused by a polluted water
supply.

Perhaps the worst effect of chemical pollution is to be found in the
streams of western Pennsylvania, where water heavily loaded with oil or
acid water from coal mines is permitted to flow into the rivers and streams
of this part of the state. Studies by Ortmann (1909) show that whole
stretches of the Allegheny, Ohio, and Monongahela rivers have been made
into deserts, as far as the animal life is concerned, by the large amount of
poisonous substances discharged into these streams by the mines, oil indus-

68 ILLINOIS BIOLOGICAL MONOGRAPHS [166

tries, and chemical and other factories that border these rivers. In the
Susquehanna River the same condition prevails in many places (Leighton,
1904). Such pollution causes a complete extermination of the fauna
(and largely of the flora) and leaves the streams in such condition that
restocking by either natural or artificial means is practically impossible.

Pollution by sewage, when the polluting material is of small percentage
as compared with the pure water of the stream (as 200 to 1), causes little
inconvenience to the animal life and is doubtless of some benefit because of
the additional food material that is added (Forbes and Richardson, 1919:
146). But the streams seldom remain long in this innoxious condition, the
sewage becoming more and more concentrated until the whole stream
may be supersaturated with noxious substances, the amount of oxygen in
saturation reduced, and the biota finally driven out or killed.

The Illinois River is one of the most striking examples of the effect of
sewage pollution on the life of a stream. Under the direction of Dr. S. A.
Forbes, studies of this river have been carried on for more than forty-two
years (since 1877) and a mass of reliable data has been gathered. The open-
ing of the Chicago Drainage Canal in 1890 produced most revolutionary
changes in the life of the Illinois river, by the discharge into it of the
sewage of Chicago as well as commercial wastes from this city and other
places along the river (Forbes and Richardson, 1913, 1919). The effect
of this sewage pollution has been to cause the animal life to be almost
excluded from the upper parts of the river. That the polluted condition
is creeping down stream is shown by comparisons of collections made in
1911 with those made in 1918. In the earlier years a foul-water fungus
disappeared from the river near Starved Rock; in 1918 it v/as found at
Henry and Lacon, 35 and 41 miles farther down the river (Forbes and
Richardson, 1919:145). At the present time (1919) optimum conditions
and a normal river fauna are not encountered until Peoria is reached, a
distance of about 120 miles from the chief source of pollution at Lockport.
Sewage from the towns and cities along the river also contribute to the
general septic condition.

A striking example of the deadly effect of sewage pollution on the
mussel life of a stream is given by Wilson and Clark (1912:34) in their
study of the Kankakee River mussel fauna. "The DesPlaines River,
which joins the Kankakee to form the Illinois River, is simply an immense
sewer bringing down the Chicago sewage. Both rivers, but especially the
DesPlaines, are full of the characteristic algae and other vegetation whicli
grow in such waters^ and the combination of a copious vegetation with the
sewage has effectually killed off all the mussels in the vicinity. Not a
single living specimen could be found in either river; but there were hun-
dreds of dead shells along the banks, most of these old and well bleached,
but still capable of identification." This statement, of course, applies only

167] FAUNA OF BIG VERMIUON RJVER— BAKER 69

to the lower part of the Kankakee River where the influence of the polluted
DesPlaines has worked upstream for some distance. The Kankakee River
for the most part is a highly productive stream with a high rate of dissolved
oxygen, in fact, the water is supersaturated with this life-giving element.

In the Maumee River (Wilson and Clark:1912, 26, 28) shell beds were
found which had probably been killed by the refuse from gas works near
the junction of the St. Mary's and St. Joseph's rivers. "Spots of tar were
found on dead mussels some distance below this point. The water was
covered with an oily scum in places and a tarry odor was perceptible for
several miles down the river." Lower down the river the mussels were
showing the effect of increased pollution of the river by sewage.

The pollution is worst and usually the most deadly to animal life during
periods of low water and in winter when the amount of water in the stream
is small and the decomposing organic material has less water to deprive
of its dissolved oxygen. During times of floods the putrescent material
is also carried down the stream for many miles and contaminates areas not
previously affected.

While all clean-water forms of animal life are more or less affected
by sewage pollution, the decomposition of organic matter abstracting
dissolved oxygen from the water and rendering it unsuitable for aquatic
life, the fish, river mussels and crayfish are particularly affected, most
fish being especially sensitive to contaminated water. Some fish (as the
brook silversides, Labidesthes sicculus) are notably sensitive, while others
(as the black bullhead, Ameiurus mclas) will endure water that is badly
polluted (Shelford, 1918:27; Wells, 1918:562-567). The young fish are
relatively more sensitive than the adult fish. It is noteworthy that the more
resistant species of fish are inhabitants of sluggish bodies of water, as ponds
and shallow lakes, while the least resistant species live in running streams.
It seems to be a question of the amount of oxygen necessary for the well
being of the fish.

The ill effect of sewage pollution is most marked on the bottom of
bodies of water, where a sludge is formed, often of great thickness (as
much as ten feet in some instances), consisting of a mass of soft, black,
sediment with a high content of organic matter, in which only a few
organisms, normally inhabitants of polluted streams, can live (e.g., septic
Protozoa and Rotifera, foul- water algae, and slime worms, Tubificidae).
This effect on the bottom is perhaps the most serious phase of stream
pollution because the septic condition of this area continues in operation
long after the original source of contamination ceases to operate. This
sludge formation renders the bottom unfit for clean-water life upon which
many fish depend for food.

The effect of sewage pollution on the fish population of the upper Illi-
nois River has been marked, many species, such as catfishes, red-horse,

70 ILLINOIS BIOLOGICAL MONOGRAPHS [168

buffalo, and sheepshead, which were formerly very common and taken in
quantity by the fishermen several years ago, are now either wanting, or
greatly reduced in numbers. Other fish, not bottom feeders, such as
sunfishes, crappies, and the basses, are reported to be decreasing in recent
years as polluted conditions are creeping down the river (Forbes and
Richardson, 1913:544). It has been observed that fish entering a polluted
stream from a clean-water tributary soon die if unable to return to clean
water. The fauna of a polluted stream also becomes gradually of greater
size as the distance from the source of pollution increases. This has been
observed by Forbes and Richardson in the Illinois River, by Ortmann in
the Allegheny River, and by the author in the Big Vermilion River.
The time necessary for the recovery of the normal biota of such a stream
will in most cases be of long duration and in the case of a stream polluted
by wastes from mines and chemical manufacturies, there may never be a
return to the original condition.

In New York State, the Genesee River, at Rochester, has afforded a
striking example of stream pollution, of the effect of this pollution on cer-
tain animal life in the river, and of the return of this life when the amount
of pollution has been largely reduced. This stream has been under observa-
tion by the writer for a period of twenty-seven years (1892 to 1919) and
collections of the molluscan life have been made from time to time, both
before the period of maximum pollution and since that time. The portion
of the river studied lies below the lower falls north of the city, and about a
quarter of a mile below the outfall of several trunk sewers, the sewage
being discharged into the river in a crude condition. Refuse and other
waste matter, both liquid and solid, also enter the stream from gas works,
tanneries, and manufacturing plants above the lower falls.

Collections made in 1892, before pollution became notably apparent,
included nine species of gastropod moUusks, three being water breathers
and six air breathers. These species included:

Musculium iransversum Physa sayid

Musculium partumeium Physa oneida

Bythinia tentaculala Galha calascopium

Planorhis Irivolvis Galba caperata
Physa gyrina

Individuals were notably abundant, thickly covering the rocks and
the shore. In 1897, it was observed that the sewage was increasing in
volume and pollution was becoming more noticable, the water appearing
like very heavy, greasy dish water. The river was visited and examined
at short intervals from 1898 to 1919. Each year it was noted that pollu-
tion was rapidly increasing. In 1907, the water-breathing moUusks,
Musculium and Bythinia, had succumbed and none could be found.
The air-breathers, Galba, Planorbis, and Physa, still held out, though

169] FAUNA OF BIG VERMILION RIVER— BAKER 71

reduced in number of individuals. An examination made in 1910 failed
to discover a single living moUusk of any species. Apparently the water
had reached such a state of concentrated pollution that even the air-breath-
ing mollusks, which normally come to the surface to take free air, could
not adapt themselves to this unfavorable environment and were either
killed or compelled to migrate down the river to a point where pollution
was less deadly. During the following years, 1910 to 1913, the river was
visited but no mollusks were found.

During the summer of 1912, G. C. Whipple, made a study of the effect
of the sewage pollution on certain animal and vegetal life in the Genesee
River (Fisher, 1913:179-200). This study was made when pollution was
at its maximum and during the period when molluscan life had disappeared
from the lower part of the river. The dissolved oxygen in the lower river,
below the trunk line sewer, in July and August, when the temperature
was high and the water low, varied from 5 to 41 per cent of saturation.
The water at the bottom of the river almost always contained less oxygen
than that at the surface. On one day in August, the percentage of satu-
ration in a distance of three miles did not exceed 5 per cent from the
surface to the bottom of the stream, which has a depth of a.bout twenty-
six feet. The number of bacteria per cc for this period was 1,650,000 near
the source of pollution and but 67,000 per cc near the mouth of the river
where the influence of the pure water from Lake Ontario increased the
amount of dissolved oxygen.

In 1917, a large part of the city sewage was diverted to a disposal
plant situated near the shore of Lake Ontario. Here an average of 32
million gallons of sewage are treated daily and the treated sewage is dis-
charged into Lake Ontario in deep water at some distance from shore.
It is at once apparent that when this large amount of sewage was discharged
into the Genesee River in a crude condition, it could not but render the
water totally unfit for animal life and a menace even to the inhabitants
who visited the beautiful parks bordering both sides of the lower Genesee
River.

The result of the diminution in the amount and character of the sewage
discharged into the river has been that the molluscan fauna, as well as
other forms of aquatic animal life, have returned and are rapidly taking
possession of the favorable environments which were in use previous
to the maximum period of pollution. Collections made in September,
1919, contained six species, two being water-breathers and four air-
breathers.

Musculium transversum Planorbis trivolvis

Bylhinia tentaculata Physa Integra

Galba catascopium Physa oneida

72 ILLINOIS BIOLOGICAL MONOGRAPHS [170

It will be noted that of the returned species, one is different {Physa integra),
while four are missing, Galba caperata, Physa gyrina, Physa sayii, and
Musculium partumeium. It frequently happens that when a fauna returns
to a habitat from which it has been driven by unfavorable conditions, it
is made up of a different aggregation of species (see Ortmann, 1909, for
additional notes on this subject).

The Genesee River is a striking example of the history of a polluted
stream and its effect on one group of animal life. Previous to the stage
of greatest pollution there is a varied fauna of mollusks very numerous in
individuals. In the course of eleven years the gill-bearing species are
forced out and after a lapse of fourteen years all molluscan life ceases to
live in this part of the river. Seven years later the greater amount of
sewage is diverted to another outlet. Two years after this change the mol-
lusks have returned in as great numbers as before the maximum stage of
pollution. The significance of all this lies in the fact of the early return of
this life and strikingly indicates that streams may become restocked with
life in a short period after pollution has ceased to be of an unfavorable
character, provided, of course, the bottom of the stream has not been
made permanently untenable by the deposition of poisonous substances
that cannot be washed away by ordinary river currents. It is quite prob-
able that the large fall of water, some 60 feet in height, immediately above
the sewage outlet, has had a marked effect in the return of these favorable
conditions.

No additional data are at hand indicating the changes in a polluted
stream after septic conditions have ceased or become greatly modified.
It is probable that similar beneficial results would be obtained in other
streams if the sewage was diverted or treated to remove the large amount
of organ,ic matter. In the case of a stream like the Salt Fork, the septic
condition of which will be discussed in the following pages, it would
probably not require a very long time to reduce the septic conditions if
the sewage from Urbana and Champaign were properly treated. While
the putrescible matter at present covers everything, in some places to a
considerable depth, the high water during the spring would in several
seasons remove a large part of this material, and i£ no additional matter
was permitted to flow into the stream, the lapse of a few years v.^ould
enable nature to bring the stream back to a normal, healthy condition,
and make it a place to seek for recreation instead of a place to avoid on
account of its filth, as at present.

SEWAGE POLLUTION IN THE SALT FORK

The sewage and other wastes of the Twin Cities of Urbana and Cham-
paign are discharged into the waters of the Salt Fork by separate systems,
that of Urbana emptying into the Boneyard near the Big Four shops, while

171] FAUNA OF BIG VERMILION RIVER— BAKER 73

the S€wer outlet of the Champaign system is situated on the Salt Fork
about a mile below th,e Urbana outlet. There are two systems for each
city, one for dome3tic wastes and the other for the care of storm water,
the sanitary sewage. Septic tanks were installed with the systems, about
the year 1894, to reduce the amount of putrescible matter, but at the pres-
ent time the sewage receives little treatment and practically enters the
Salt Fork in a crude condition.* The population in 1914 was estimated to
be 13,750 for Champaign, and 9,252 for Urbana, or a total population of
about 23,000 for the Twin Cities. At the present time, 1920, six years later,
the increase has probably brought the total up to nearly 30,000. The
sewage system, therefore, provides disposal for this population, and is all
discharged into the waters of the Salt Fork, It is estimated by G. C. Haber-
meyer, that the total flow of sewage from the Urbana plant is about 500,000
gallons per day and from the Champaign plant about 1,000,000 gallons
per day.**

The flow of the Salt Fork below the Champaign sewage disposal plant
is 3,000,000 gallons per day. These figures indicate that the sewage forms
one-half of the total water flowing down the Salt Fork. These data were
taken in October, when the stream was low, and may be a trifle too high
for those periods when there is a rise of water following a period of rainy
weather. During a greater part of the year, however, the water is low and
these figures will be approximately correct. The fresh water added to
the sewage is derived from the stream north of Urbana which contributes
250,000 gallons per day, and the Boneyard, which adds 1,500,000 gallons
per day, about two-thirds being clear water. "In October, 1917, the
flow in the Boneyard below the Urbana tank was about one-third sewage
and probably contained considerable other waste and sewage discharged
above the Urbana sewage outlet. The flow in Salt Fork below the Cham-
paign sewer outlet was probably one-half sewage."

H. E. Babbitt,*** thus describes the condition at the Champaign dispo-
sal plant at this date. "The appearance of the efiQuent from the Champaign
septic tank is that of fresh sewage, having the typical color of sewage, and
carrying fecal matter and paper. The appearance of the Salt Fork at the
point of entrance of the sewage from the tank is good. It is about twenty

* New septic tanks have been installed at the Champaign sewage disposal plant on Salt
Fork and a portion of the sewage is well treated before it enters the canal.

** Data for the sewage conditions, stream flow, chemical analyses, etc., of the Salt Fork are
taken from an unpublished report of G. C. Habermeyer (assisted by S. D. Kirkpatrick, assis-
tant chemist, and J. F. Schellbach, engineer) made for the State Water Survey Division of
the Department of Registration and Education, of Illinois, and here used by permission of the
late Chief of the Division, Dr. Edward Bartow.

*** From unpublished Report on the Champaign-Urbana Water Works System, prepared
June 23, 1914. Extracts here published by permission of Edward Bartow.

74 ILLINOIS BIOLOGICAL MONOGRAPHS [172

feet wide, fifteen to eighteen inches deep, clear, colorless, and odorless.
The stage of water at the time of inspection was low. No septic action
was present except in the open ditch through which the sewage flows from
the tank into the stream. There is a large sludge bank in the stream
immediately below the outlet and a most obnoxious odor." Effluent from
the Urbana tank appeared as typical fresh sewage, but was not representa-
tive of the ordinary effluent as the tank had been cleaned only the day
previous to the visit (page 9).

The stream known under the name of the ''Boneyard" carries both
waste and sewage. This stream is about three and a half miles in length and
rises about half a mile beyond the northern limits of the City of Cham-
paign. The stream flows southward to near Third and Green streets,
where it abruptly turns eastward, emptying into the Salt Fork near the
Big Four shops in Urbana. This stream has a drainage area of about eight
and a half square miles. Sewage enters the Boneyard near Goodwin
Avenue and at the old high school on Stoughten Street, Urbana. Water
bearing wastes, presumably from business houses near by, enters the stream
north of Main Street. The banks close to the water line near Main Street
culvert are slimy and green (data from G. C. Habermeyer, 1918).

The Boneyard is subject to great fluctuations of water level, due to the
severe storms of spring and summer when heavy rains occur. At such times
a rise of three or four feet in a few hours is not unusual and the waters
overflow all adjacent low land. At one time (reported very bad in 1915)
chemical wastes in the form of oil and tar were discharged into the stream
from the gas works of the Champaign Street Railway, Gas and Electric
Company, situated at the corner of Fifth and Hill streets, Champaign,
east of the Illinois Central tracks. The oil at one period extended the
entire length of the Boneyard, covering lawns, when the water was high,
with an unsightly layer of heavy oil. The shores and bottom of the stream
in many places were covered with tar, which has not subsequently been
removed and also cannot be removed by the natural flow of water, but must
be artificially taken from the stream bed.*

The oil discharged from the gas works, as well as from some other
points along the Boneyard, is absolutely inimical to any life in the stream.
The tar, if carried down stream, finally settles to the bottom and unless
artificially removed, will remain there and render the stream unfit for
bottom inhabiting life of any kind. Fortunately, such pollution is confined
to the upper part of the Boneyard. Waters charged with sewage may
become purified in a year or two after pollution ceases, but they will sel-
dom or never recover from chemical pollution such as is produced by tar

* From Ralph Hilscher, Report on Contamination of the Boneyard in Champaign by
Gas House Wastes August 28, 1915. Here published by permission of Edward Bartow.

1731 FAUNA OF BIG VERMILION RIVER— BAKER 75

and similar wastes. Ortmann's work on the rivers of western Pennsylvania
clearly indicate the baneful results of such pollution (Ortmann, 1909).

The Boneyard is apparently now barren of all clean water life. Fre-
quent examinations made during 1918 failed to supply any life of this
character. Fresh water pulmonate mollusks, and Cambarus and other
smaller Crustacea were once abundant, but appear to be wanting at the
present time. A large snapping turtle was observed in the Boneyard near
Lincoln Avenue, in 1918. The ditched portion of Salt Fork above Urbana,
is a clean water stream, filled with aquatic life, which abruptly terminates
at the junction of this stream with the Boneyard, with its load of sewage
from Urbana. From here to a point fourteen miles below Urbana not a
living mussel was found, and no air breathing snails were observed, except
in one instance, believed to have been introduced from a near-by portion
of the old river bed, which still retains clean water life, above St. Joseph,
a distance of ten miles below Urbana. The large number of empty valves
and paired shells of the river mussels below St. Joseph indicate an environ-
ment that has become, more or less recently, inimical to these creatures.
Crayfish were also absent from the same area.

Habermeyer's report of Salt Fork, October 1, 1917, describes condi-
tions as follows: "The stream below the Champaign outlet to the north
line of section 10, one and a half miles below the outlet, was in very foul
condition. At tb^ east line of section 11, four miles below the Champaign
sewer outlet, the stream appeared to be quite clear and there was no
offensive odor noticeable. At the outlet of the west branch of Salt Fork,
the water was clearer than that in the north branch (Spoon River). A
resident near the junction of the two branches stated that at times when
the creek flow increased, foul matter was washed down from above and
there was considerable odor in the vicinity for a day or two."

The dredging of a new channel has been responsible for the water
being clearer at times in Salt Fork than in Spoon River, providing a sand
bottom which is hard and resistant and has not yet silted up to any degree.
Spoon River has a mud bottom and the waters are turbid a large part of
the time. In a photograph of the Salt Fork taken some eight miles below
Urbana (two miles northwest of St. Joseph), the water was so clear that
the ripple marks on the sandy bottom may be distinctly seen in the picture
(Fig. 45) yet the stream at this point was totally barren of clean water
life and the water was laden with masses of decomposing matter, made up
of foul water algae and Protozoa, and the bottom was filled with slime
worms (Limnodrilus). Below the junction of these two branches, bottom
conditions are still very bad, and clean water life does not appear for a
distance of several miles, as has already been shovvn.

The people mentioned in the report were interviewed in 1918. They
reported that no trouble was experienced during periods of low water, but

76 ILLINOIS BIOLOGICAL MONOGRAPHS [174

that after rains, when the stream rapidly rose, putrescible matter was
washed down stream, cast on shore, and the odor was then very bad. This
lasted until the water subsided. In the fall of 1920, the same people were
again interviewed and conditions were reported to be much worse than
previously, the unusually low stage of the water causing the putrescible
matter to decay on the exposed sand bars in the river, from which some
odor was noticed.

Stream measurements of the velocity of the current were made by the
Water Survey at several points along the Salt Fork. These were made by
Habermeyer on October 1, 1917, when the water was low in the stream.
All data given in feet.

Table IX. Mouth of the Boneyard

Distance from east bank 1 2 3

Depth of water 0.6 0.7 0.4

Flow per hour 4608 4500 4320

Salt Fork, Four Miles below Champaign Sewer Outlet

Distance from bank 2.9 2.4 1.8 2.2

Flow per hour 3888 4320 4212 2412

These measurements indicate a very slow current, from about one half
to nearly a mile an hour. At high water the rate of flow is undoubtedly
several times as great. Tliis slow rate of flow would cause much of the
heavy matter in the sewage to be precipitated to the bottom and form
sludge banks, and this has occurred at many places below both the Urbana
and Champaign outlets. High water and more rapid current probably
moves some of this sludge farther down stream, also carrying the fresh
sewage farther down at such times. A recent examination made at very
low water indicates that this has been the case, for hundreds of bars were
observed out of water, each covered with a sludge formation of greater
or less size.

Analyses of water from various places in the Salt Fork and tributary
streams have been made by the Water Survey. The two tables that fol-
low, taken from Habermeyer's report, indicate some of the conditions of
the upper waters of Salt Fork (Tables X and XI).

In Table X the low percentage of dissolved oxygen (saturation) in
the ditched stream above Lincoln Avenue is noteworthy as compared with
the larger amount at Cunningham Avenue. At both localities, however,
the water is relatively pure, there being little or no pollution. The sample
from a mile and a quarter below the champaign outlet is striking because
of the total absence of oxygen. The sudden rise in saturation at four
and eight miles is also notable, and indicates that at these points on this
date little decomposition was taking place. The sample from Salt Fork
near Spoon River shows a marked fall in the amount of dissolved oxygen,

1751

FAUNA OF BIG VERMILION RIVER

77

Pi
o

H

<

Q
<
a
<

Z
O
P5

a

o
«

tn
H
»-l

<

o

><
X

o

Q

w
>

o

t/J

o
z
<

<

H
CO

H
>

t— I
H
<

i-l
H

O

H
<

g

<:

X
H

p-i
<:
u
I— )

o
o

1-4

o
w

H
O
<

1-1
<

i

3

s

1

a

(

a

a
o

3

T3

o

<

a

o
<J

c

12

Cunningham Avec
\^ miles below Cha

u

S o

J3

u

<U O

sewer
of Spo

on River
, sewage o

of Boneyai

hJ

■t-l

a S

OO

■g o

River
Up Spo
Urbana

-1

»-H

2

[1^

a
S

I-l

00 -H

o

r~~

^ •^

3

lU

\0 t--

t-~

O

\0 vO

■*-»

ni

^

-*-»

u

>%

C

_>

-Ij

<u

.^^

^

Tj-

•* --1

^

1<

Tj<

«— ( v-^

*-(

rt

3

OO

00 CN

ca

00

00

(N4 CN

r-t

"33

a

a

0^

4->

in

_g

o

c
.2

a

"1

UI O

r-

r^

CN

*

t^

X

o

3

lO

Tf o

<3

Ov

"*

00 to

d

Id

lO

00 o

O

CO

LO

\0 CS

^-<

^3

I)

■if

c/)

g

t-»

to o

O

LO

r--

O in

o

»— t

t^ o

T-<

o

O

CO T*

o

Q

d

lO

r- O

vO

OO

lO

O r^i

*— <

rH

o

^

o

o

o

o

»-l

c

o

"^

CO

Q

s

^

o

C

rt

£?

T— t

o

o

o
o

♦

*

M

o

'5

.,— (

c

O

o

o

«

»

*

«-(-<

o

tn

o

*-H

o
c

*

*

♦

*

*-H

o

*

*

*

*

^

1— i

• * *

*

*

jf

*

*

d

^ r ^

c

o c

J ^

) c

c

O C

> o

u

c:

CN C

) c

) c

c

O C

> o

\C

CN C

> c

I c

c

o c

> o

o ^

aS;

rj

CN C

r c

r CN

<::

o" c

J LO

l/~

) ""

r<-

CN C

> OS

§-

o

•^

LT

— . 1^

.s

rt •£:

.4_)

C

o c

) c

> c

C

o c

) O

■^ ■"

oc

) o c

) c

) c

> c

o c

> O

O -1-1

Cv

' "*. ^

>_ c

)_ c

) t-

o c

I o_

H ^

'-T c

r c

r cv

CS

r o" c

3 *~4

-Q

a
O

) t-

c<-

)

-*§

> OO

1

Ui

c

) -^ "■

> v:

5 t^

oc

) CN C

> »-<

U4

O

h

^

t~

- <^ r-

I--

r-

t--

»^ oc

3 00

i-i

s

^—

i •.-1 —

^ ^M

4 «-

< ^-1 w-

1 »-H

^

O

«

3 00 OC

3 a

3 S

) oc

) 00 oc

3 00

oi

•4-*

3

r<-

5 ro r<-

5 r'

> C

) c

) ro f

3 ro

1-

-J

(3

1

o

OO

00

I-l

<u

3
C

>

o

XI

<U
u

a

tn

IH
O

a

u3

_>

ji

IH

d

C

03
fcfl
>>

o

tn

(U

78

ILLINOIS BIOLOGICAL MONOGRAPHS

[176

o

M
l-l
1^

W

CM
H

<

p<

O
P-,

H

i-l
<

o

<

Q
H
<

W

O

m
m

H

a

o

1-1

s
<

Pm
O
[«
M

<

<

<
u

H
K
U

X!

•-;
<

'

1 • 1

I

1

>-l

■>->

t 1

p

coin Ave
Ave.
ow Cham

E
U

6

ta
U

oon Rive

ver

ge outlet

C

o

60

lU

e close to Ur
wage tank.

5

D

.S S ID

-Ft tn

In O

paign sewer
Mouth of Sp
Up Spoon Ri
Urbana sewa

C
bO

a

West of
Cunning
Hi mile:

c -Si

c ^
&

00

o
o

ir-l

'3
ex

S
u

manhol
bana se

>>

■4-J

"

— 1

M

fN o o

00

OO

\0 00 •<*
lO -^ VO

o

CN

r^

lO •^ ■rt

^*

"0

I^

OO

a

kA

fN CN -^

t^

<^

r^ CN •«*

CO

■<*

■*<

"<

C

^ u

D

lO o o

o

o

O CN O

o

o

Jh o

r^i !^3 O

lO

o

lo o o

o

o

o o o

CN

rO

CN O O

o

o

2-1

trogen

itrate

trogen

r-1 •^ O

CO

O

O O OO

NO

NO

J^ \0 tN

'^

O

rq -^ ■*

rn

lO

r-l

Csl

i^ ^ '^

73

o o o

o

Q)

o o o

o

o

."2

T

<u

■* Tf O

00

T^

OO vo OO

^

o

cn

_>

Pvl !M O

\o

O

rt< fo O

rfl

NO

mino
rogen

p

tn

CN

CN

tH

y—^

^

o ^ o
^ ■* o

o

r^

o o o

o

O

3 *->

c3

tN

in

CN O O

o

o

a c

*J

CN »-^ IN

t^

1--

f- CN -^

TT'

-*

O

<:

H

ro

•^

CN

NO

■g 1

■H

rsi O) O

o

O

o o o

o

o

u

^ r^ O

o

O

O CO o

o

o

11

->

■^ O O

OO

o

O O CN

'^

o

CO

Tt<

t^

ro C\

NO

CN

tM

^H

(N »-i

CO

< <

^

,

T3

vO o o

o

(VI

CN O O

(N

CN

S

TJH Tf Tt*

CN t^ O
CN

CO

A c

"rt

\0 o o

CN

00

O 00 00

•<*

CO

O o

(J

o

-* ^ o

*-H

<vj

cr> t^ 00

NO

00

H

*— (

''"'

T— (

-H VO

CO

CO

.s

)-H <>

5 1

Tt< rO CN

«-H

vO

On O \0

CO

lO

^ •-

-H VO

lO

iO

in lo

NO

00

-fl

s

u

CJ

. T

3

c c S

J 1>

o o i

3-5

O O O

o

lO

lO o o

lO

o

tn *^ C

t^ 0\ 00

lO

•*

"* O cs

NO

NO

t/3 •!— « Ci

■-H -r-l tN

CM

CN

CN 1-H ro

CN

CO

i-J .W)^

5 ^

1

-O

lO lO lO

o

o

lO o «o

o

«iO

t/3

1>

'o

en

vo r-- •*

o

Ov

CO 00 1^

00

(M

(5

ro lO \0

o

lO

lO rr> >0

>o

00

Residi
evapoi

rj

ir> o in

lO

ID

O O O

IT)

o

■4-J

lO OO Tji

o

ro O CO

■*

NO

O

■* lO t^

o

vO

NO "* OO

NO

i

-1 ,

c

3_§

3 E

<^ ■>* lO

vO

t^

00 OS O

»-l

(N

■4.

J^ t^ 1^

r^

J^

1^ t^ 00

OO

OO

a

^-H ^— ( vH

T-l

^-H

<^^ ■^^ -r^

*-4

y—i

00 OO 00

00

CO

00 OO 00

00

00

^ 3

«^ <n fO

rO

<^J

CO ro ro

CO

CO

c

j a

)-

177] FAUNA OF BIG VERMILION RIVER— BAKER 79

which coincides with the absence of mollusks and crayfish from this part of
the stream. The rise of the oxygen content 200 feet upstream in Spoon
River shows the effect of the cleaner water, and this also coincides with
the presence of clean water life in this part of Spoon River. The lower
temperatures were probably responsible for the rise in oxygen content at
the stations four and eight miles below Urbana. In the summer months,
during low water and high temperature conditions, the percentage would
probably be much lower. This difference was noted by Forbes and
Richardson in their study of the Illinois River. The percentages of dis-
solved oxygen are relatively higher in these two stations of the Salt Fork,
four and eight miles from the source of pollution, than in the Illinois River
at Starved Rock, Hennepin, and Chillicothe, where samples taken in Octo-
ber showed smaller percentages, though many miles from the chief source
of pollution at Lockport (Forbes and Richardson, 1913:565).

The rise and fall in the number of bacteria and the presence of many
gas-forming organisms in the Salt Fork is well shown in the table. The
sudden rise from Cunningham Avenue to the point one and a quarter
miles below the Champaign sewer outlet is very striking and indicates
in a graphic manner the difference between clean water and that heavily
polluted by sewage wastes. The high number 200 feet up from the mouth
of Spoon River indicates that there is some pollution in this stream, al-
though not far above this point living mollusks (Naiades) were found in
abundance.

In Table XI, the relative high amounts of albuminoid nitrogen, as well
as ammonia nitrogen, supplied by the sewage material, in contrast with the
small amounts in the purer water of the stream, are strikingly shown.
All of these agencies are inimical to clean water life, especially fish, mussels,
and crayfish, which by their relative abundance or absence, indicate in a
most satisfactory manner the quality of the environment, and its fitness
for the well-being of its inhabitants.

The sewage in the Salt Fork has greatly increased in recent years, fol-
lowing the increase in population, and, as in the case of the sewage from
Chicago in the Illinois River, the polluted condition is gradually creeping
down the stream below St. Joseph, changing the pure water inhabitants
to those that can live under septic conditions. As no collections were made
during previous years from the stream below St. Joseph it is not now pos-
sible to make comparisons of the present with previous conditions. Such
lists as have been available have not indicated precisely the location from
which they were collected (as in the neighborhood of St. Joseph, which
might mean in the Salt Fork or in Spoon River) and they cannot be used
for this reason. The value of exact local lists is emphasized in studies of this
kind, showing that strictly technical (or pure science) information is often
of the greatest value in the study of economic probems.

80

ILLINOIS BIOLOGICAL MONOGRAPHS

[178

RECENT EXAMINATION OF THE POLLUTED PORTION OF SALT FORK

Figures 21, 36 to 45

During the fall, winter, and spring of 1919 and 1920 Salt Fork was
carefully examined from Urbana to St. Joseph. All parts of the stream
were searched for macroscopic life and samples of the bottom and of the
water were taken for microscopic study, to determine the character and
abundance of the foul-water organisms present. Examinations were made
during the months of May, September, October, November, and December.

During the spring months the water is so high in the stream that collec-
tions can not be made and examination of the water is difficult. The
great volume of water, laden with both sewage and silt, is of a dark lead
color and polluted conditions are not apparent, although some putrescent
matter was observed on several occasions. In the fall months conditions
are much more favorable for critical examination, the water being so low
that one may wade over any part of the stream. It is at this time, which
extends from July through the fall and winter to the rainy period in March
or April, that the polluted conditions are very apparent and the most
satisfactory studies can be made. The general conditions of the stream at
this period are summarized below.

The water in the Boneyard near the Big Four shops, below the Urbana
sewage outlet is usually shallow, from a foot or less to two feet in places.
The bottom is a mass of sludge with putrescent material covering all
objects in the water and floating down the stream, which has an estimated
flow of about half a mile an hour. Several large sludge banks are exposed
and the odor is usually nauseating (Fig. 39). At the junction of the
Boneyard with the Salt Fork the water varies from six inches to a foot
in depth, there are several sludge banks and the putrescent matter covers
all objects and lines the shores to a height of several feet, indicating former
high water marks (Fig. 36). Samples of the sludge and green putrescent
material from one of these sludge banks were examined by Professors
Smith and Transeau and the following algae, Protozoa and other animals
observed:

Blue-green algae

Animals

Pediaslrum simplex, common.
Senedesmus abundans, rare.
Phormidium inundatum, abundant.

Diatoms
Navicula cryptocephala, common.
Synedea pulchella, not common.

Paramoecium, not common.
Euglena geniculata, very abundant.
Limnodrilus (sludge worm), very abun-
dant.

Tubifex (sludge worm), one specimen.
Nematode worms, several.

Ciliate Protozoa were numerous, including Colpodium and a few
hypotrichous and peritrichous species.

179J FAUNA OF BIG VERMILION RIVER— BAKER 81

At the junction of the Boneyard with the Salt Fork the waters of the
two streams usually formed two distinct bands, the clear water of the latter
on the left or north bank and the murky, sewage-laden water of the Bone-
yard on the right or south bank, the line in the center of the stream dividing
the two waters being clearly marked. (Fig. 21). The waters of these
two streams do not fully mingle until they have flowed a distance of
several hundred yards. All the way down the Salt Fork, however, as far
as the first bridge, the right side is more discolored than the left side,
indicating that the sewage-laden waters of the Boneyard and Champaign
outlet flow mostly on this side of the stream.

At the Champaign sewage outlet, about half a mile below the Bone-
yard, there is usually a good flow of water from the discharge pipe into an
open ditch, which empties into the Salt Fork a short distance away. At
the point of entrance of the open ditch, the chocolate-colored water of the
Champaign sewage is clearly marked as a dark band extending around the
upstream end of a large sludge bank, similar to the one described in Haber-
meyer's report in 1917 (Fig. 41). Some fecal matter is generally pre-
sent. The water in the Salt Fork at this point is usually less than a foot
deep and the bottom is made up of a soft sludge which covers everything in
the water. The odor is very bad, almost nauseating.

Below the Champaign outlet the stream is in very foul condition, as
noted in Habermeyer's report. From the outlet to the first bridge, more
than a mile down stream, the water is less than a foot deep, in most places
but a few inches, and the bottom has much sludge, and putrescent matter
covers every object in the water, as well as the shores and all objects on the
shores to a height of several feet, indicating former stream levels. Fecal
matter, in dark brown masses, as well as partly decomposed organic matter
colored green by the presence of blue-green algae and the protozoan
Euglena, are usually floating down the stream. The bottom substratum
is of sand and gravel, and over this sludge banks have been formed of
greater or less thickness. Bars of sand and gravel occur at irregular inter-
vals and are covered with masses of putrescent matter forming long, alter-
nating streaks of black and green. An oily scum resembling petroleum
covers the surface in many places, and the bottom, when disturbed, emits
an oily substance which spreads over the surface as an oily scum. This may
represent some gas house wastes as well as heavy oil from other places
along the Boneyard. The odor in this section of the stream is almost over-
powering in many places, being distinctly a privy smell. In one place the
observer was compelled to leave the vicinity of the stream, the odor was
so strongly nauseating. In places, bubbles of gas may be seen to break at
the surface from submerged sludge banks.

Samples of the bottom sludge and putrescent matter floating in the
water were collected just above the bridge, about a mile and a quarter

82

ILLINOIS BIOLOGICAL MONOGRAPHS

[ISO

below the Champaign outlet. Algae and animal life were noted, but not as
abundantly as in the sludge bank at the junction of the Salt Fork and Bone-
yard (Fig. 21).

Animals
Euglena geniculata, very few.
Paramoecium, one specimen.
Vorticella, very few.

Blue-green algae

Phormidium inundatum, abundant.

Diatoms
Navicula salinarum, abundant.

The scarcity of microscopic life and the total absence of clean-water
life is paralleled by the chemical condition of the water at this station,
where the examination of samples showed a total absence of dissolved
oxygen and the presence of multitudes of bacteria and gas-forming organ-
isms (see Tables X, XI).

From the first iron bridge to the Brownfield woods bridge, a distance
of about a mile and a quarter, the stream is very shallow, less than a foot
in depth on the average, the water grayish with a sloppy appearance, and
the odor foul in places, though not as nauseating as belov/ the first bridge.
Putrescent masses of soft, grayish-black or greenish matter, ranging in size
from a penny to a platter, may be seen floating down stream, held together
by algal strands. Brown masses observed on shore resembled human ex-
crement. The oily appearance of the surface of the water continues and
oily matter ascends when the bottom is disturbed.

From the Brownfield bridge to the third (farmer's) bridge, a distance
of about three-quarters of a mile, the conditions are the same as above this
bridge (Fig. 38). Between the third bridge and the Cottonwoods road
bridge, a distance of about a mile, the water is very shallow, scarcely
exceeding six-inches in depth, with gravel or sand bottom. Sludge bars,
of sand mixed with organic matter, are frequent, in many places occupying
more than half the width of the stream, the channel, a few inches to a foot
in depth, meandering over the bottom between these bars (Fig. 40).

These bars presented a striking appearance, the stones and sand being
black from the decomposing organic matter, and the foul water algae
being arranged in long streaks, presenting in combination a striped green
and black design. The green algae is here very abundant, floating in the
stream or covering the shores. In places the encrusting material on the
shore margins is bright yellow. Everywhere along the stream the exposed
surface of the bottom is black from the decomposing organic matter,
which covers all objects and has been baked hard by the hot summer's
sun. The vegetation bordering the shore also shows the effect of sewage
action, being either black in color or having the dried pieces of polluted
material attached to the lower part of the plants. The same conditions
prevail from the Cottonwoods road to the Mayview road, a distance of
about two miles.

181]

FAUNA OF BIG VERMILION RIVER— BAKER

83

Collections of materials made just below the third farmer's bridge, about
three and a half miles below the Champaign outlet, contained the following
organisms:

Blue-green algae Animals

Phormidium inundatum, abundant. Flagellate Protozoa, very minute.

Diatoms Euglena geniculala, very abundant.

Navicula salinarum, abundant. Many in stage of encystment.

Dineutes assimilis, very abundant.

On the surface of the water.

Collections at the Cottonwoods bridge (Fig. 42) contained a larger
variety of animal life, which was rather meagerly represented above
this bridge. This place is on the east line of section 11, about four miles
below the Champaign outlet. The bottom here is of fine sand and mud.

Blue-green algae

Phormidium inundatum, abundant.

Diatoms
Navicula salinarum, abundant.

Animals

Euglena geniculala, very abundant.
Rotifer, illoricate, one specimen.
Limnodrilus, two specimens.
Dineutes assimilis, a few examples.

Mussel shells or other mollusks were entirely absent in a living state
and their shells were notably rare. About three quarters of a mile below
the first bridge east of Urbana, a half valve of Lampsilis luteola was found
on a sand bar (Fig. 37). Near the Brownfield woods bridge many broken
pieces of mussel shells, as well as a few mutilated half valves, were ob-
served. At a farmer's bridge half a mile below this bridge several broken
valves of Lampsilis luteola and Anodonta grandis were collected, (Fig.
38) and from this point down stream to the Cottonwoods road bridge
detached valves or broken pieces of shell wer€ more or less common.
From observations of this and other parts of the stream it seems evident
that these mutilated shells were washed from the spoil banks on either side
or from the bed of the old stream channel where it crosses the canal. At the
junction of the Boneyard with the Salt Fork a layer of these shells was
observed in the bank, about eighteen inches above the water line (the water
being low), in a position that indicated the old bed of the Salt Fork before
the canal was excavated. High water would wash this material way and
provide the odd valves of mussels observed in different parts of this stream.

Below Mayview road bridge the conditions are much the same as in
the neighborhood of the Cottonwoods bridge. The bottom is of sand and
gravel, with some mud bordering the shore. The water is from a few
inches to a foot in depth, the channel meandering among a continuous
series of sand bars. The sand is ripple-marked in places and streaked
with bands of dark green algae, with yellowish algae in spots. The surface
of the flowing portion of the stream is thickly covered with patches of dark

84

ILLINOIS BIOLOGICAL MONOGRAPHS

[182

green putrescent matter, measuring in size from a peanut to a dish pan.
Some of these masses are brownish, where the algae and Protozoa have not
completely taken possession of them.

A large amount of oily scum may be observed on the surface and
when disturbed the bottom emits quantities of oily matter, as is the case
higher up the stream. On exposed bars and along shore the algae and
putrescent matter have dried and caked, forming a pavement-like layer.
The water is clearer here than in the portion of the stream previously
examined, but no clean water life could be found; mussels, crayfish, and
insects were entirely absent.

Samples of the bottom from the stream about 300 feet east of the
Mayview road bridge, about six miles below the Champaign outlet were
examined. The following life was present:

Blue-green algae

Pediastrum simplex, rare.
Phormidium inundatum, common.

Diatoms
Navicula salinarutn, abundant.
Fragilaria capucina, abundant.

Animals

Ciliata, minute, abundant.

Colpodium, several.

Euglena genictdata, very abundant.

Limnodrilus, common.

Nematode worms, minute, abundant.

About a mile and a half below this locality additional samples were
taken for examination. Conditions are similar but the water is not as
clear, holding more sediment in suspension.

Blue-green algae

Pediastrum simplex, rare.
Phormidium inundatum, abundant.

Diatoms
Fragilaria capucina, abundant.

Animals

Ciliata, minute, abundant.
Paramoecium, several.
Euglena geniculata, abundant.
Limnodrilus, about a dozen.
Nematode worms, minute, many.

At the last north and south farmer's bridge, the canal makes a wide
sweep, in a southeasterly direction (Fig. 43) leaving the old stream bed
to the west of the new channel, in the form of an 'ox-bow' almost half
a mile in length, which, during the greater part of the year, forms a large
elongated pond, that drains into Salt Fork canal by means of a small outlet
at the south end which turns abruptly northeastward as it empties into the
canal. At the time examined, the bed of this old stream was almost dry,
following a period of very dry weather, and the fauna had retired to several
small, shallow, muddy pools which remained in the deeper parts of the
stream bed. An examination of these pools disclosed a number of bull-
heads (Ameiurus melas), many dragonfly larvae {Libellula piilchella),
and a few moUusks {Planorbis trivolvis, Physa gyrina, and MuscuUum
transversum).

183] FAUNA OF BIG VERMILION RIVER— BAKER 85

It seems evident that this portion of the old stream forms a reservoir
from which certain species of mollusks, as well as fish, are carried, or volun-
tarily migrate, through the outlet into Salt Fork. By this means only
can the presence of these animals in the polluted water be accounted for,
because none have been seen either above or far below the drain from the
old stream channel. The specimens of Planorhis trivolvis, that have been
observed in the Salt Fork canal below the east and west road bridge, were
probably derived from this source.

Specimens of Planorbis trivolvis have been observed in the Salt Fork
canal which probably came from this source. Near this locality a school
of about fifty fingerling bullheads was observed on May 29, in a small
part of the stream where a rather deep pool had developed. They were
making frantic efforts to get out of the pool but the surrounding water
^as too shallow. The low water and general polluted condition of the
stream evidently provided a very unfavorable environment. The source
from which these fish came was quite likely the old cut off portion of the
original Salt Fork from which place they had been carried or had volun-
tarily migrated, when the water was higher from the April rains.

From the east and west road bridge (Fig. 45) to the first farmer's bridge,
a distance of about three-quarters of a mile, the conditions are the same
as in the preceding portion of the canal. The water is clear but no living
mollusks or other animals could be found. A half valve of Anodonta
grandis, badly weathered, and a few bleached valves of Sphaerium solidu-
lum, were the only evidences of molluscan life. These had evidently been
washed from the old stream bed at some point where it was exposed by
erosion in the canal walls. The same algae as previously observed was
floating down the stream in large green patches. Samples of these green
particles were examined by Professor Smith and found to contain such
animials as Euglena geniculata, Paramoecium, and numerous ciliate Pro-
tozoa, mostly inhabitants of polluted water.

From the next farmer's bridge to the eastward turn of the stream the
bottom conditions were also similar, except that the surface of the water
was covered with an oily scum. Groups of greater or less size of gyrinid
beetles {Dineutes assimilis) were seen at different places down stream for
a mile or more. A single empty shell of Planorbis trivolvis was observed
on the shore of the west bank below this bridge. No living clams were
seen, nor any mutilated valves.

From the eastward turn of the stream to the mouth of Spoon River,
the canal is in bad condition, the water being dirty and oily, with numer-
ous bubbles of gas rising constantly from the bottom, which also gave off
quantities of oily scum when disturbed (Fig. 44). The water is shallow
and conditions as already described for the stream higher up. Green

86

ILLINOIS BIOLOGICAL MONOGRAPHS

[184

putrescent matter is not quite as abundant as higher up in the canal
and the odor is not as bad.

Samples were collected from a point about a mile above the junction
of Salt Fork with the Spoon River. Organisms were fewer in both species
and individuals.

Blue-green algae

Phormidium inundatum, common.

Diatoms
Synedea pulchella, abundant.
Fragilaria capucina, abundant.

Animals

CUiata, abundant.
Euglena geniculata.
Limnodrilus, a few individuals.

Below the mouth of Spoon River conditions are also bad. Where the
current is strong, sand bars have been formed which cover the black mud
beneath. Everywhere, except in the channel where it is sandy, the bottom
is of soft mud, eight to fifteen inches deep, filled with ill-smelling gasses.
Sewage conditions here are bad, the water having a foul smell. When
wading in the water, the disturbed mud, which is black and oozy, con-
stantly gives off bubbles of gasses that rise to the surface and break. This
condition is uniform for the entire stretch of stream bed examined, about
a third of a mile, from near the mouth of Spoon River to the middle of the
big bend above St. Joseph. The surface of the water is usually covered
with a film of oil resembling petroleum.

No living mussels could be found in this area. A single valve of Am-
blema undulata was seen near the mouth of Spoon River which was prob-
ably brought to its resting place from Spoon River during a period of
flood. It is noteworthy that while no mussels or gill-bearing snails were
observed in this part of Salt Fork, several species of fresh water pul-
monates were seen in considerable numbers, indicating that here, as
elsewhere, the pulmonates are able to live in water which is totally
unfit for mussels and gill-bearing moUusks.

Examples of the bottom sediments and green masses floating in the
water were taken from the stream about a hundred feet below the mouth
of Spoon River. Algae was plentiful and included Phormidium inundatum,
a blue-green species, in abundance. Tv/o species of diatoms, Navicula
salinarum and Fragilaria capucina, were also present. Of animals, Limno-
drilus was well represented, but Euglena geniculata was not common, and
ciliate Protozoa were much less numerous than in samples taken from
stations higher up in the stream. Salt Fork here shows the influence of
the added cleaner water from Spoon River.

At the mouth of Spoon River a bar has been formed, by natural or
artificial agencies, which, except for a space about four feet wide, holds
back the waters of that stream (Fig. 16). An examination was made
of Spoon River about 400 feet above this bar. Here five species of mussels

185] FAUNA OF BIG VERMILION RIVER— BAKER 87

were found, in more or less abundance, and insects were also abundant.
The water is low, two or three feet deep in the channel in the fall and the
bottom is very muddy. A very small amount of green algae was noted
on the surface, evidently brought up from Salt Fork. The mussels col-
lected included:

Lasmigona complanata Lampsilis luteola

Amblema undulata Eurynia lienosa

Carunculina parva

A water boatman (Corixa, near burmeisteri) was very abundant,
especially in the nymph stage, as were also numerous gyrinid beetles
(Dineutes assimilis). Though the water was low, men were observed
fishing for bullheads in this part of Spoon River. The contrast between
this stream, with its abundance of clean water life, and the adjacent parts
of Salt Fork, which is utterly devoid of clean water life, strikingly indicates
the harmful efifects of sewage pollution on the fauna of our streams.

Samples of the green matter floating on the water of the Spoon River
were examined and found to contain Euglena geniculata, common, ciliate
Protozoa, many. These masses seem to be made up almost entirely of
these animals and low plants, of which three species were noted: Phormi-
dium inundatum alga; Navicula salinarum and Fragilaria capucina, dia-
toms. A single pupa of Chironomus decorus was found at this place.

For a distance of several miles below St. Joseph unfavorable conditions
seem to prevail, no living mussels being found for a distance of over four
miles, and here they occur rarely. Living mussels are not found in any
number for a distance of over five miles below St. Joseph. These animals
are not abundant in species and individuals until a distance of twenty miles
has been traversed below Urbana. Conditions for a distance of two miles
below St. Joseph are similar to those described for the area just below
Spoon River, the bottom consisting of black mud from which bubbles of
ill-smelling gas rise when the bottom is disturbed. Below this point condi-
tions begin to improve, though very gradually.

That conditions along the polluted portion of Salt Fork are often, if not
always, highly objectionable was evidenced from conversations with far-
mers living near the stream. One farmer, who had built a small house
within a few hundred feet of the stream, stated that the "stench was at
times almost unbearable" and that people living half a mile away were
strongly conscious of the odor. This was about five miles down the
stream from Urbana. A gentleman driving along the road which parallels
the Salt Fork east of Cottonwoods road, stated that the odor on Septem-
ber 10 was very obnoxious. People living a mile south and north of the
stream do not suffer from these odors.

88 ILLINOIS BIOLOGICAL MONOGRAPHS [186

SUMMARY OF SALT FORK CONDITIONS

It has been shown in the previous pages that the sewage and other
wastes that drain into the Salt Fork from the Twin Cities have driven out
or killed all clean water life from the junction of the Boneyard with Salt
Fork to a point about four miles below St. Joseph, or fourteen miles below
Urbana. At this point a few living mussels are found and also a few cray-
fish. One must pass down the stream for a distance of twenty miles before
encountering a normal river fauna, comparable to that found in Spoon
River at a point less than a mile above the junction of that stream with
Salt Fork. The abundance of clean-water life in Spoon River is in marked
contrast with the total absence of this kind of life in Salt Fork, which
normally would have, in suitable habitats, a similar fauna in the barren
stretch of ten miles between the two localities compared. No better
example is known of the total annihilation of a fauna from so great a dis-
tance as the result of polluted conditions.

Foul water algae and Protozoa, as well as some other animal life (slime
worms) characteristic of polluted Vv'ater, are abundant in that portion of
the' stream devoid of clean water life. The same relative conditions were
observed by Forbes and Richardson in their study of the Illinois River.

Fish, especially young fish, have been made an index to the degree of
pollution of streams. It would seem from observations made during the
course of the present study, as well as from other occasions and in other
places, that bottom-inhabiting animals, such as river mussels and cray-
fish, provide a better index for this purpose. Fish are able to migrate
easily and swiftly from an unfavorable to a more favorable environment,
but these more sedentary animals, especially the mussels, cannot change
their environment so easily and must either adapt themselves to the more
unfavorable conditions or perish. For example, young bullheads were
observed in Salt Fork about three miles above St. Joseph in the spring
when the water was comparatively high. But no mussels or crayfish
have been seen within five miles of this point. This indicates clearly
that fish are more flexible in this matter than the mussels and crayfish,
which are not as mobile. Ortmann (1909:93-94) believes that crayfish are
slightly more resistant than mussels to polluted conditions, and as scaven-
gers (they have been observed eating dead mussels) they could naturally
withstand a limited degree of unfavorable environment. Observations
made on the Salt Fork, however, indicate that the two groups appear at
about the same time.

Forbes and Richardson (1913:498) distinguish three stages of impurity
of streams, which may apply equally well to either the stream itself or to
the organisms living in the stream. These terms are "given in the order
of diminishing impurity, namely, (1) septic or saprobic, (2) polluted or

187] FAUNA OF BIG VERMIUON RIVER— BAKER 89

pollutional, and (3) contaminated or contaminate; and to these we will
add 'clean-water' to indicate the conditions and organisms substantially
equivalent to those of the natural, uncontaminated stream."

Judging Salt Fork by these standards of impurity, we would say that
from the Urbana sewage outlet to the first bridge below the Champaign
sewer outlet, a distance of about two miles, the stream is in a septic condi-
tion. From this point to about two miles below St. Joseph it is polluted.
From this region to Homer Park, below the dam, it is contaminated.
Below the dam there is probably some contamination at times, but the
fauna is a clean water one and the stream would be classed as a clean water
stream, being unmodified by sewage conditions. The lower part of Salt
Fork near Middle Fork, and Middle Fork and the Big Vermilion as far as
Danville are clean-water streams with a large and varied fauna of mussels,
crayfish, and insect larvae. Below Danville there is sewage pollution, and
conditions are again unfavorable.

90 ILLINOIS BIO LOGIC A L MONOGRA PHS 1 1 88

SUMMARY AND CONCLUSIONS

The mussel fauna of the Big Vermilion River consists of at least 35
species and varieties. Ten species of small pelecypods and 15 species
of gastropods also occur, making a total molluscan fauna of 60 species
and varieties occurring in a distance of upwards of 50 miles. The mussel
fauna of this stream compares well with that of two other Illinois streams
of comparable size, the Sangamon, 150 miles long, with 25 species, and the
Kankakee, 300 miles long, with 48 species. The total length of the Big
Vermilion River is 90 miles, with 35 species. Seventeen species of Naiades
occur in the Big Vermilion River that have not been found in the Sangamon
River, while five species have been collected from the Sangamon that have
not yet been detected in the Big Vermilion. The former stream belongs
to the Mississippi drainage while the latter is a part of the Wabash drain-
age.

In the Big Vermilion it was noted that there was a progressive increase
in number of species as the distance down stream increased, the head-
waters containing but few species, which are usually smaller than the
same species from lower down in the stream. It was also observed that
the headwater inhabitants, as well as many naiads farther down the stream,
were more compressed and of greater comparative length than the same
species as it occurred in the larger part of the river.

The dam at Homer Park, 27 miles below Urbana, appears to form a
barrier between the fauna above and below this obstruction. Above the
dam, 19 species occur, while below the dam, 32> species have been observed.
It is noteworthy that immediately above the dam the largest number of
species found at one habitat was 14, while below the dam, at Homer
Park, 28 species have been collected. It is probable that the fall of water
over the dam aerates the stream and provides an especially favorable
environment for the mussels.

UNFAVORABLE INFLUENCES

Sewage pollution has killed all clean water life for a distance of fourteen
miles below Urbana and has made the stream an unfavorable environ-
ment for a distance of twenty miles. Below this point the fauna is normal
and is not affected by sewage conditions. In the desert area between St.
Joseph and Urbana, slime worms and septic Protozoa were observed
among the animals and foul water algae among the plants. A few beetles,
breathing free air, were observed in the stream near St. Joseph and for
some distance above this locality. Many of the old pond-like bodies of

189] FAUNA OF BIG VERMILION RIVER— BAKER 91

water left on either side of the new drainage ditch are inhabited by
clean water animals that occasionally get into the ditch during periods
of high water. Fish, mollusks, and insects have been observed which
doubtless came from this source.

PARASITES AND PATHOLOGIC AGENCIES

As a rule, parasites, either distomids or Unionicola (Atax) were rare
in the naiads collected in the Big Vermilion and Sangamon rivers. Some
of the Anodontas had marginal cists of distomids and many shells of this
genus were discolored from this cause. Clark and Wilson (1912:61), in
their study of the Maumee River fauna, observed distomids in various
mussels which were believed to be the distomids described by Osborn and
Kelly. The affected mussels were Aciinonaias ligamentina, Obovaria
cir cuius, Elliptio gibbosus, Lampsilis ventricosa, Amblema undulata, Las-
migona costata (thought to be the distomid of Kelly), and Anodonta
grandis (thought to be the distomid of Osborn). It is probable that these
flat-worms also infest many of the mussels of the rivers investigated, but
they were not observed in the specimens collected. Cotylaspis insignis
and forms of Unionicola were also found by Clark and Wilson, but these
parasites were not seen in the mussels examined from the area under con-
sideration.

Pearls, usually of small size, were frequently seen attached to the
shells of mussels, and many pearly growths cause by injuries were also
observed. A large round pearl was found in a shell of Anodonta grandis
gigantea from Crystal Lake, which measured almost a quarter of an inch
in diameter (5 mm.). It has been suggested that many of these pearls and
pearly growths attached to the mussels may have been caused by parasites,
such as the distomids before mentioned. The same is also true of the round
pearls found in the animal tissues of the mussels.

Crippled shells, those individuals have abnormal valves, were not com-
mon in either of the rivers examined. Mud was found to cause trouble in
many cases, getting in between the mantle and the shell below the pallial
line and causing large blisters.

ECONOMIC CONSIDERATIONS

The shells known as river mussels or Naiades are used in the manufac-
ture of pearl buttons. As this industry, the making of pearl buttons, has
reached large proportions it is imperative that the raw material be con-
served for the maintenance of the industry. The continued fishing of the
mussel beds in the larger rivers has greatly depleted the amount of avail-
able raw material — the mussel beds — , and the whole industry, shell collect-
ing and button making, is threatened with disaster if means are not found to
restock the depleted beds (see Coker, 1919:44). The United States Bureau

92 ILLINOIS BIOLOGICAL MONOGRAPHS [190

of Fisheries has conducted many interesting experiments on the propaga-
tion of mussels by the artificial infection of fish with mussel glochidia and
the means and methods for restocking these cleaned-out areas are at hand.
It only remains for proper laws to be passed and enforced, by the states or
federal government, or both, regulating the time and place in which shelling
operations may be carried on. Reasonable time must be given, at least
three years, for the recovery of a depleted mussel bed.

In this connection it would seem that the mussel fauna of such a stream
as the Big Vermilion River might form a reservoir from which the depleted
beds farther down the stream might be restocked by fish which had been
infected with glochidia from the commercial species living in the smaller
stream. The Big Vermilion contains eleven species that are used for
cutting button blanks and are considered valuable for this purpose by the
button manufacturers. These are:

Amblema undulata Blue point
Lampsilis luteola Fat mucket
Lampsilis anodontoides Yellow sand-shell
Lampsilis ventricosa Pocket-book
Trilogonia iuberculata Buckhom
Quadrula pustulosa Warty-back
Quadrula lachrymosa Maple-leaf
Actinonaias ligamentina Mucket
Fusconaia rubiginoas Wabash pig-^;oe
Lasmigona coniplanata White heel-splitter
Lasmigona costata Fluted shell

Several of the smaller shells are also used when shells are scarce, as
Lampsilis compressa, Quadrula metanevra, Obovaria circulus, and Strophitus
edentulus. In the Sangamon River about the same number of species suit-
able for the button industry occur and these are usually of fine quality.

In their survey of the mussel fauna of the Kankakee basin, Wilson and
Clark (1912:35) recognize the value of these smaller streams, with a fauna
too small in individuals to be used by the shell fishermen, but containing
many of the essential species from which good button blanks may be cut.
These authors say: "The most valuable species are all good breeders
throughout the basin. This, taken in connection with the excellent quality
of the shells they produce and the good railroad facilities everywhere
available, makes this basin one of the best yet examined for the supply
of glochidia to be used in artificial mussel propagation." This statement
might apply with almost equal force to the Big Vermilion, which may
sometime be needed for a reservoir from which to propagate the mussels
in the larger rivers.

Whether all of the fishes which have proved the most satisfactory hosts
for glochidia are abundant here is not known, but as young of nearly all the

191] FAUNA OF BIG VERMILION RIVER— BAKER 93

commercial shells were observed, it follows that the fishes carrying the
glochidia must also be present.

The Big Vermilion River is seen, therefore, to be a valuable asset to the
State, containing a fauna of both biologic and economic importance,
which should be conserved for possible use as a restocking reservoir for
the larger rivers. Pollution should be reduced to a minimum and the
timber along the banks, especially of the headwaters, should be conserved
in order that the water may be held in the ground and gradually flow into
the streams, instead of running off in floods, causing excessive high water
in the spring and extremely low water in the fall and winter. It would be
possible to reforest many parts of the upper branches of the streams,
where they are low or irregular places unfit for farming operations.

94 ILLINOIS BIOLOGICAL MONOGRAPHS 1192

BIBLIOGRAPHY

Adams, C. C.

1900. Variation in lo. Proc. Amcr. Assoc. Adv. Sci., 49:208-225.
1915. The Variations and Ecological Distribution of the Snails of the Genus lo. Mem-
oir Nat. Acad. Sci., 12, 184, pp. 61 pi.
Baker, F. C.

1906. A Catalogue of the MoUusca of Illinois. Bull. State Lab. Nat. Hist., 7:53-136.
1920. Animal Life and Sewage in the Genesee River, New York. Amer. Nat., 54:

152-161.
1920. A New Planorbis from Illinois. Nautilus, 33 :123-125.
BiNNEY, W. G.

1865. Land and Fresh Water Shells of North America. Part 11, 161 pp., 261 figs.;
part III, 120 pp., 232 figs,
Clark, H., and Wilson, C. B.

1912. The Mussel Fauna of the Maumee River. Bureau Fish., Doc. 757, 72 pp., 2 pi.
CoKER, Robert E.

1914. The Protection of Fresh-water Mussels. Bureau Fish., Doc. 793, 23 pages, 2 pi.

1919. Fresh-water Mussels and Mussel Industries of the United States. Bull. Bur.
Fish., 36:11-89, 46 pi.
Danglade, Ernest.

1914. The Mussel Resources of the Illinois River. Bureau Fish., Doc. 804, 48 pp., 5 pi.
Eldridge, John A.

1914. The Mussel Fishery of the Fox River, op. cit., 8 pp.
Forbes, S. A., and Richardson, R. E.

1908. The Fishes of Illinois. Nat. Hist. Surv. 111., Vol. Ill, Ichthyology. 357 pp.
Many plates and an Atlas of maps.

1913. Studies on the Biology of the Upper Illinois River. Bull. 111. State Lab. Nat. Hist.

9:481-574, 21 pi.
1919. Some Recent Changes in Illinois Biology. Bull. 111. State Lab. Nat. Hist., 13:
137-156.
Haldeman, S. S.

1842-1844. A Monograph of the Fresh-water Univalve Mollusca of the United States.
Philadelphia. Many colored plates.
Howard, Arthur D.

1914. Experiments in the Propagation of Fresh-water Mussels of the Quadrula group.

Bur. Fish., Doc. 801. 52 pp., 6 pi.
1914. A Second Case of Metamorphosis without Parasitism in the Unionidae. Science,

40:353-355.
Iseley, Frederick B.

1911. Preliminary Note on the Ecolog>- of the Early Juvenile Life of the Unionidae.

Biol. Bull., 20:79-115.
1914. Experimental Study of the Growth and Migration of Fresh-water Mussels.

Bur. Fish., Doc. 792, 24 pp., 3 pi.
LeFevre, Geo., and Curtis, Winterton C.

1908. Experiments in the Artificial Propagation of Fresh-water Mussels. BuU. Bur.

Fish., 28:615-626.

193] FAUNA OF BIG VERMILION RIVER— BAKER 95

1910. Reproduction and Parasitism in the Unionidae. Journ. Exper. Zool., 9:79-115,
pi. 1-5.
Leighton, Marshall O.

1904. Quality of Water in the Susquehanna River Drainage Basin. Water Supply and
Irrigation Paper No. 108, U. S. Geol. Surv., 76 pp.
Leverett, Frank.

1899. The Illinois Glacial Lobe. U. S. Geol. Surv. Mon. 38.
Marsh, M. C.

1908. Notes on the Dissolved Content of Water in its Effect upon Fishes. Bull. Bur.

Fish., 28:891-906.
Ortmann, a. E.

1909. The Destruction of the Fresh-water Fauna in Western Pennsylvania. Proc.

Amer. Phil. Soc, 48:90-110, 1 pi.
1912. Notes on the Families and Genera of the Najades. Ann. Carnegie Mus., 8:222-

365.
1918. The Nayades (Fresh-water Mussels) of the Upper Tennessee Drainage, with Notes

on Synonymy and Distribution. Proc. Amer. Phil. Soc, 57:521-626.
1920. Correlation of Shape and Station in Fresh-water Mussels (Naiades). Proc. Am.

Phil. Soc, 59: pp. 267-312.
1920a. A Monograph of the Naiades of Pennsylvania. Part III. Systematic Account of

the Genera and Species. Mem. Carnegie Mus., 8:1-384, 21 pi.
Pennsylvania.

1918. Industrial W'aste Survey of the Clarion River Basin. Ann. Rept. Commis.

Health, Penn., 10:1279-1316.
Shelford, V. E.

1917. An Experimental Study of the Effects of Gas Waste upon Fishes, with Especial

Reference to Stream Pollution. Bull. State Lab. Nat. Hist., 11:378-425.

1918. Ways and Means of Measuring the Dangers of Pollution to Fisheries. Bull. 111.

Nat. Hist. Surv., 13:23-42.
SmpsoN, C. T.

1914. A Descriptive Catalogue of the Naiades, or Pearly Fresh-Water Mussels. Bryant

Walker, Detroit, Mich. 1540 pp.
Smith, Frank.

1915. Two New Varieties of Earthworms with a Key to Described Species in Illinois.

Bull. State Lab. Nat. Hist. 10:551-559, 1 pi.

SUREER, ThADDEXJS.

1912. Identification of the Glochidia of Fresh-water Mussels. Bur. Fish. Doc, 771,

10 pp. 3, pi.

1913. Notes on the Natural Hosts of Fresh-water Mussels. Bull. Bur. Fish., 32:

101-116, 3 pi.
.1915. Identification of the Glochidia of Fresh-water Mussels. Bur. Fish. Doc, 813,9
pp. 1 pi.
Utterbach, W. I.

1916. The Naiades of Missouri. Amer. Midland Nat., 4, nos. 1-10, 200 pp., 29 pi.
1916. Breeding Record of Missouri Mussels. Privately printed, 8 pp.

Walker, Bryant.

1918. A Synopsis of the Classification of the Fresh-water MoUusca of North America
north of Mexico, and a Catalogue of the More Recently Described Species,
with Notes. Univ. Mich., Mus. Zool., Misc. Publ., no. 6. 213 pp., many cuts.
Ward, H. B.

1918. The Elimination of Stream Pollution in New York State. Trans. Amer. Fish.
Soc, 48:25 pp.

96 ILLINOIS BIOLOGICAL MONOGRAPHS [194

19 19. Stream Pollution. Jour. Amer. Water Works Assoc. , 6 :823-83 1 .

1919. Stream Pollution in New York State. A Preliminary Investigation of the Prob-

lem from the Standpoint of the Biologist. New York Conserv. Comm., 79 pp.,

3 pi.
Wells, M.

1918. The Reactions and Resistance of Fishes to Carbon Dioxide and Carbon Monoxide.

BuU. lU. State Lab. Nat. Hist., 11:557-571.
Wells, W. F.

1920. Conservation of the Quality of Water of New York State as a Natural Resource.

Science, 52:279-284.
Wilson, C. B. and Clark, H. W.

1912. The Mussel Fauna of the Kankakee Basin. Bur. Fish. Doc, 758, 52 pp., pi. and

map.
1914. The Mussels of the Cumberland River and its Tributaries. Bur. Fish. Doc, 781,

63 pp., 1 pi.

195] FAUNA OF BIG VERMILION RIVER— BAKER 97

PLATE I

98 ILUNOIS BIOLOGICAL MONOGRAPHS [196

EXPLANATION OF PLATE

Fig. 1. Drainage ditch above Urbana, east of cemetery. Station 2.

Fig. 2. Salt Fork south of Muncie. Note water willow in center of stream bed; also
numerous bare portions of stream bed, the river appearing as a series of pools. October, 1919.
Station 26.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER

PLATE I

197] FAUNA OF BIG VERMIUON RIVER— BAKER 99

PLATE II

100 ILLINOIS BIOLOGICAL MONOGRAPHS [198

EXPLANATION OF PLATE

Fig. 3. Old stream bed of Salt Fork in Busey woods, north of Crystal Lake, Urbana.
Original habitat for Planorbis pseiidolrivolvis. Station 3.

Fig. 4. Junction of Middle Fork with Salt Fork to form the Big Vermilion River. Look-
ing south from road bridge crossing Middle Fork. September 26, 1920.

Fig. 5. Middle Fork a fourth of a mile above junction with Salt Fork. Bed of river
exposed in foreground. Station 29. September 26, 1920.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

.MS^^

BAKER

FAUNA OF BIG VERMILION RIVER PLATE II

199] FAUNA OF BIG VERMILION RIVER— BAKER 101

PLATE III

102 ILLINOIS BIOLOGICAL MONOGRAPHS [200

EXPLANATION OF PLATE

Fkj. 6. Spoon River, seven-tenths of a mile above Salt Fork. Station 10. September
28, 1918.

Fig. 7. Spoon River, riffles below bridge, same locality as Fig. 6.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE III

201]

FA UNA OF BIG VERMILION RI VER—BA KER 103

PLATE IV

104 ILLINOIS BIOLOGICAL MONOGRAPHS [202

EXPLANATION OF PLATE

Fig. 8. Bench mark 655, Salt Fork, above road bridge. Station 17. September 13, 1918.
Fig. 9. Salt Fork two and a half miles north of Sidney, looking north, the Champaign
moraine on the left, a flood plain on the right. Station 18. September 13, 1918.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

.5. -im

' ' ' - - ■ -^

8

■
I

^'^^"' -• ■*^

BAKER

FAUNA OF BIG VERMILION RIVER PLATE IV

203] FAUNA OF BIG VERMILION RIVER— BAKER 105

PLATE V

106 ILLINOIS BIOLOGICAL MONOGRAPHS [204

EXPLANATION OF PLATE

Fig. 10. Iron bridge one mile north of Sidney. Station 19. August 26, 1918.
Fig. 11. Cement bridge northeast of Sidney. Note large area of water lily, Nym-
phaea advena. Station 20, August 26, 1918.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER

PLATE V

2051 FAUNA OF BIG VERMILION RIVER— BAKER 107

PLATE VI

108 ILLINOIS BIOLOGICAL MONOGRAPHS [206

EXPLANATION OF PLATE

Fig. 12. Salt Fork three and a half miles above Homer Park, Station 23. October 4,
1918.

Fig. 13. Salt Fork. Deep pool below dam and rapids. Professor Smith's field laboratory
during a period of twenty years. Station 25. November 4, 1920.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE VI

207] FAUNA OF BIG VERMILION RIVER— BAKER 109

PLATE VII

no ILUNOIS BIOLOGICAL MONOGRAPHS (208

EXPLANATION OF PLATE

Fig. 14. Daxn in Salt Fork at Homer Park. The dam is just below the interurban
bridge and is five feet high. November 4, 1920.

Fig. 15. Salt Fork below dam at Homer Park. Shallow water and rocky bottom. No-
vember 4, 1920. Part of Station 25.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE VII

209] FA UNA OF BIG VERMIUON RI VER— BAKER 1 1 1

PLATE VIII

112 ILLINOIS BIOLOGICAL MONOGRAPHS [210

EXPLANATION OF PLATE

Fig. 16. Salt Fork and mouth of Spoon River. Rowboat marks outlet of Spoon River
through bar. September 24, 1920.

Fig. 17. Valley of Big VermUion River from crest of bank at Gray's Siding. October 8,
1920.

Fig. 18. Cutting through shale rock covered with glacial deposits. Big Vermilion River
one mile below mouth of Middle Fork. Station 30. October 8, 1920.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE VIII

211) FAUNA OF BIG VERMILION RIVER— BAKER} 113

PLATE IX

114 ILLINOIS BIOLOGICAL MONOGRAPHS [212

EXPLANATION OF PLATE

Fig. 19. Rock bed of Big Vermilion River one mile below Middle Fork. Station 30.
October 8, 1920.

Fig. 20. Sandbar in center of riverbed, several hundred feet below fig. 1. Station 30.
October 8, 1920.

Fig. 21. Junction of Boneyard with Salt Fork. Boneyard on right above ice. December
20, 1919.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE IX

213] FA UNA OF BIG VERMILION RI VER— BAKER 1 15

PLATE X

116

ILLINOIS BIOLOGICAL MONOGRAPHS

[214

EXPLANATION OF PLATE

Fig. 22. Anodonta grandis.
Fig. 23. Anodonta grandis.
Fig. 24. Anodonta grandis.
Fig. 25. Amblemaundulata.
Fig. 26. Amblema undulata.
Fig. 27. Amblemaundulata.
Fig. 28. Amblema undulata.
margin of shell.

Right valve with pearly growths at anterior end.

Left valve with injured portion folded inward.

Pearly formation in left valve.

Right valve uninjured.

Left valve with large blister inside pallial line.

Left valve with abnormal pallial line.

Left valve with pin-head pearls between pallial line and

Pathologic mussels from Salt Fork.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

24

£6

BAKER

FAUNA OF BIG VERMILION RIVER PLATE X

215J FA UNA OF BIG VERMILION RI VER-BAKER

PLATE XI

117 '

118 ILLINOIS BIOLOGICAL MONOGRAPHS [216

EXPLANATION OF PLATE

Fig. 29. Amblema undulata. Left valve with channel due to injury.

Fig. 30. Amblema undulata. Right valve with ridge due to same injury.

Fig. 31. Strophitus edentulus. Right valve with pearl near posterior end.

Fig. 32. Strophitus edentulus. Left valve with distomid discoloration.

Fig. a. Lasmigona costata. Right valve with repaired injur}' near posterior end.

Fig. 34. Amblema undulata. Spoon River form with rounded shell.

Fig. 35. Lampsilis venlricosa. Left valve with injured postero-ventral margin.

Pathologic mussels from Salt Fork.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

29

94

BAKER

FAUNA OF BIG VERMILION RIVER PLATE XI

217] FAUNA OF BIG VERMILION RIVER— BAKER 119

PLATE XII

120 ILLINOIS BIOLOGICAL MONOGRAPHS [218

EXPLANATION OF PLATE

Fio. 36. Junction of Boneyard with Salt Fork canal, looking west. Note high bank on
left where mussel shells were found, indicating bottom of bed of Salt Fork stream before the
canal was dug. Sludge bank in foreground. September 14, 1920.

Fig. 37. Salt Fork canal three-fourths of a mile below first iron bridge east of Urbana.
December 31, 1919.

Fig. 38. Farmer's bridge across Salt Fork canal, one-half nule below Brownfield Woods
bridge. December 31, 1919.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE XII

219] FAUNA OF BIG VERMILION RIVER— BAKER 121

PLATE XllI

122 ILLINOIS BIOLOGICAL MONOGRAPHS [220

EXPLANATION OF PLATE

Fig. 39. Boneyard just below Urbana septic tank outlet. September 14, 1920.

Fig. 40. Salt Fork canal west of Cottonwood's road bridge. Note bare sand bars with
stream meandering between. Bars are covered with green putrescent matter dried by the
sun. September 14, 1920.

Fig. 41. Salt Fork canal at entrance of small ditch from Champaign outfall pipe. Note
sludge bank in foreground and sewage indicated by dark color of water in center of picture.
September 14, 1920.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

BAKER

FAUNA OF BIG VERMILION RIVER PLATE XIII

2211 FAUNA OF BIG VERMILION RIVER— BAKER 123

PLATE XIV

124 ILUNOIS BIOLOGICAL MONOGRAPHS [222

EXPLANATION OF PLATE

Fig. 42. Salt Fork canal looking east from Cottonwood's road bridge. Note bare patches
of bottom. September 14, 1920.

Fig. 43. Salt Fork canal looking southeast from last north and south road bridge.
September 24, 1920.

Fig. 44. Salt Fork canal looking west from last farmer's bridge above St. Joseph. Sep)-
tembcr 24, 1920.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

L.
BAKER

1

-^I^^^^^^H^I^

%

''T^^^^^^^^^BI^HH

FAUNA OF BIG VERMILION RIVER PLATE XIV

223] FA UNA OF BIG VERMILION RI VER—BA KER

125

PLATE XV

126 ILLINOIS BIOLOGICAL MONOGRAPHS [224

EXPLANATION OF PLATE

Fig. 45. Salt Fork canal looking south from last east and west road bridge west of
St. Joseph. September 28, 1918.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

—

W^^M

w

*

'?*

1

If \

M^^H

^^^HhhhL-jbhH^ ^ k

^Bh

. f^ 4^H

i

BAKER

FAUNA OF BIG VERMILION RIVER PLATE XV

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII July, 1922 No. 3

Editorial Committee

Stephen Alfred Forbes William Trelease

Henry Baldwin Ward

Published under the

Auspices of the Graduate School by

THE University of Illinois Press

Copyright, 1922 by the University of Illinois
Distributed December 31, 1922

NORTH AMERICAN MONOSTOMES

PRIMARILY FROM FRESH WATER HOSTS

WITH NINE PLATES AND TWO TABLES

BY
E. C. HARRAH

Contributions from the

Zoological Laboratory of the University of Illinois

under the direction of Henry B. Ward No. 208

THESIS

SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE

DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY IN THE GRADUATE

SCHOOL OF THE UNIVERSITY OF ILLINOIS

1922

TABLE OF CONTENTS

Introduction 7

Methods of investigation 8

Material 8

Historical data 10

Species previously described from North America 11

New species described in this paper 12

Families of Monostomidae 13

Key to families 13

Description of families 14

Cyclocoelidae validity of the older name 14

Diagnosis 14

Key to subfamilies and genera 16

Cyclocoelum 15

Historical 16

Structure of the genus 20

Key to species of cyclocoelum 34

Description of species 35

Cyclocoelum leidyi nov. spec 35

Cyclocoelum pseudomicrostomum nov. spec 37

Cyclocoelum halli nov. spec 38

Cyclocoelum wilsoni nov. spec 40

Cyclocoelum cuneatum nov. spec 41

Cyclocoelum obscurum (Leidy) 42

Cyclocoelum macrorchis nov . spec 44

Cyclocoelum triangularmn nov. spec 46

Cyclocoelum vicarium (Arnsdorff) 47

Notocotylidae 48

Diagnosis of family 49

Key to subfamilies and genera 49

Notocotylinae 50

Notocotylus 50

Notocotylus urbanensis (Cort) 51

Stages in the life history 52

Notocotylm quinqueserialis (Barker and Laughlin) 53

Catatropis 54

Catatropis filamentis Barker 54

Paramonostomum 55

Paramonostomum echinum nov. spec 55

Nudocotylinae 56

Nudocotyle novicia Barker 56

Heronimidae 57

Heronimns chelydrae MacCallum 57

CoUjTiclidae 60

Collyriclum colei Ward 60

Diagnosis 60

Remarks on the life history 64

Species inquirendae 66

Clinostomum (?) incotnmodum Leidy 60

Monostomum ornatum Leidy 66

Monosiomum spatulalum Leidy 68

Monostomum affine Leidy 68

Monostomum asperum Vaillant 69

Monostomum amiuri Stafford 70

The polyphyletic origin of the monostomes 72

Introduction 72

Interrelationship of the monostome famihes 74

Apparent relation of these families to other groups 75

Summary 80

Bibliography 82

Description of plates 88

225] NORTH AMERICAN MONOSTOMES

INTRODUCTION

The study of the Monostomata was undertaken four years ago following
the suggestion of Professor Henry B. Ward, who expressed the need for a
thorough study of the group. A preliminary survey of the available mate-
rial served to convince me that a more complete and comparative study
of the group was desirable and that such a study would prove of value to
helminthologists. This study seemed more desirable since but two com-
parative studies of the group had been made in the last twenty-five years.
The first of these was that of Stossich (1902) which is based wholly on
European material; the second that of Kossack (1911) is based largely on
the same material. The fact that very Httle had been done on this group
in America and that no comparative study of the American fauna had been
undertaken furnished further incentive for this work.

The writer undertook an investigation of the group which should be
comprehensive in scope and at the same time comparative. For this pur-
pose he had free and unUmited use of the large personal collections of
Professor Ward secured thru extensive field work as well as obtained by
exchange. These had been augmented, with a view to getting a complete
series, by loans of material from European and American investigators.
Unfortunately specimens of certain originals could not be secured either
because they had been lost or because they could not be removed from the
collections.

Thanks are due to the following investigators who at the request of
Professor Ward kindly sent material from valuable collections for study
and comparison; Professor Anton Collin, University of Berlin; Professor
Theodor Pintner, University of Vienna; the curator of the Museum,
University of Gottingen; Professor Fr. Zschokke, University of Basel;
Dr. C. W. Stiles, Hygienic Laboratory, Washington, D. C; Dr. B. H.
Ransom, Bureau of Animal Industry. Dr. John C. Johnson has also
very kindly loaned the writer material for study. For the study of Cyclo-
coelum halli and Cydocoelum wilsoni, use has been made of the field notes
of W. C. Hall filed with the records of that series.

Especially to Professor Henry B. Ward under whose supervision the
work has been done, the writer desires to express most sincere thanks for
the use of his extensive collections and of his library, as well as for his
efforts to secure rare material and for the continued deep interest and
hearty cooperation which have given the inspiration to finish this work.

ILLINOIS BIOLOGICAL MONOGRAPHS

[226

METHODS OF INVESTIGATION

The methods employed in this study are those ordinarily used with
similar material. No special technique has been found necessary. In
general sections were stained in toto with Erlich's acid hematoxylin and
counterstained in section with alcoholic eosin. Sections were cut 10 to
20 micra in thickness. Frontal, sagittal, and transverse sections were
used. In order to obtain a clearer perception of more minute and obscured
parts wax reconstructions were made. These were employed particularly
to find the relationship and constancy of form of the female genital organs.

MATERIAL

The material on which this study is based is composed of various collec-
tions from different regions of the United States. It is limited to that
obtained from North American fresh water hosts and in each instance is
listed with the species description. However for comparison with the
Amerian material the following specimens were secured from the sources
indicated.

EUROPEAN MATERIAL
From the Zoological Museum, Berlin

Species
Cyclocoelum problematicum
Cyclocoelum problematicum
Cyclocoelum problem alicmn
Cyclocoelum ovopunctatum
Cyclocoelum braziiianum
Cyclocoelum tringae
Haematolrephus similis
Haemalolrephus similis
Monostomum vanelli

From the Zoological Museum, University of Gottingen

Host

Cat. No.

Collector

Totanus glottis

2449

Ehrenberg

To tonus glottis

2455

Ehrenberg

Canis vulpecula

2454
1342

Ehrenberg

Scolapax ftavicepa

2494

V. Olfers

2495

Ehrenberg

Himantop. atropterus

2486

Ehrenberg

2309

Ehrenberg

Vanellus cristalus

1326

Rudolphi

Cyclocoelum (Monosiomum)

mutabile (Zed.)
Monostomum verrucosum
Monostomum attenuatum
Monostomum attenuatum

284

Gallimila chloropus

Taken at Claustal
Anas bernicla
Mergus merganser
Anas fuligula

From the Zoological JSIuseum, University of Vienna

Monostomum mutabile Totanus flaviceps IX 552

Taken in Brazil

234
238
220

Mehlis

Mehlis
Mehlis
Mehlis

From the Collection of Henry B. Ward, University of Illinois

Collyriclum faba

Passer domesticus 21 . 764

Taken in Switzerland

Zschokke

Cyclocoelum leidyi

{Monostomum mutabile
Zeder of Leidy)

NORTH AMERICAN MATERIAL
From the Leidy Collection

GalUnago wilsoni 160

Leidy

227]

NORTH AMERICAN MONOSTOMES

Species Host

Cyclocoelum pseudoniicrostomum Fulica americana
(Monostomum miUabilc
Zeder of LeidjO

Cat. No. Collector
186 Leidy

From the collection of the Army Medical Museum
Cyclocoelum ohscuriim Stereolepsis sp? 1035 Leidy

From the collection of the Bureau of Animal Industry

Notocotylus urbanensis Dafila acuta 5772 Stiles & Hassall

(Monostoma sp. Stiles & Hassall)

Notocotylus urbanensis Fiber zibethicus

(Monostoma sp. Stiles & Hassall)

5769

5770 Stiles & Hassall

Notocotylus urbanensis Aix spotisa

(Monostoma sp. Stiles & Hassall)

Notocotylus quinqueserialis Arvicola riparius

(Monostoma sp. Stiles & Hassall)

5771 Stiles & Hassall

5773 Stiles & Hassall

From the Collection of Henry B. Ward, University of Illinois

Cyclocoelum pseudomicrostomum
Cyclocoelum cuneatum'
Cyclocoelum obscurum
Cyclocoelum obscurum
Cyclocoelum niacr orchis
Notocotylus quinqueserialis
Paramonostomum echinum
Heronimus chelydrae

Collyriclum colei

Cyclocoelum halli

Cyclocoeluvi wilsoni
Cyclocoelum triangularum

Wild duck
Gallinago deli^ata
Symphaemia seviipalmata
(unknown)

Straight billed curlew
Fiber zibethicus
Fiber zibethicus
Snapping turtle
Emys blatidingi
Chrysemys marginata
Kinosternon odoratus
Cinosternum pennsyhanicum
Graptemys geographicus
Passer domesticus

Totavus melanoleucus
Totanus solitarius
Gallinago delicata
Tringa maculata

10.41

08.172

08.179

08.183

08.180

15.120

21.91

08.176

12.161, 162, 163

N.B. 15,21.760,22.169

16.425

22.40,41

22.166

11.11 L.J.Cole

11.12,21.961,762

21.90 W.C.Hall

21.763 W.C.Hall

21.89 W.C.Hall

21.88 W.C.Hall

10 ILLINOIS BIOLOGICAL MONOGRAPHS [228

HISTORICAL DATA

More than a century ago the first of the Monostomata were described.
Since that time many helminthologists have contributed to the study of
this group. Most notable of the older authors are Zeder, Rudolphi, von
Siebold, Van Beneden and Diesing. In more recent times Brandes, Stos-
sich, Looss, Liihe, Monticelli, Kossack and Odhner have contributed
materially to our knowledge of the European forms.

Apart from the work of Goeze (1782) who described two species which
he believed to have only one sucker and that of Schrank (1788) which was
a mere catalogue of species, Zeder (1800, 1803) was the first to establish
this group. In 1800 he created the genus Monostoma based on five species;
Monostoma ocreatum, and Monostoma bombynae which have since been
removed to the distomes; Monostoma verriicosum a Notocotylid; and
Monostoma prismaticum and Monostoma mutabile which are now trans-
ferred to the genus Cyclocoelum. Rudolphi in his Synopsis Entozoorum
served in the early organization of the group. While a number of species
were imperfectly known at this time it was not until the work of von
Siebold (1835) was published that the anatomy of these worms was clearly
understood. In this work the author gave a good description of Monostoma
mutabile Zeder along with the early stages of the life history as shown in
the development of the egg before it is discharged from the uterus. Diesing
(1850) reorganized the genus including all species described up to this
time. Following this Van Beneden (1861) reviewed the anatomy of
Monostomum mutabile and added a description of the anatomy of Mono-
stomum verrticosum Frdlich with a study and description of a cercaria which
he believed to be the larval form of this species. The next important
contributions to the knowledge of this group were those of Monticelli
(1892) in which he gave a complete account of the genus Notocotyle Dies-
ing and in a second paper in the same year a similar account of Monosto-
mum cymhium Diesing. Closely following these was the Revision of Brandes
(1892) in which he proposed the new genus Cyclocoelum to include Mono-
stomum mutabile, M.flavum, M. arcuatum, M. tringae and M. elUpticiim.

In more recent time the work of Stossich (1902), Odhner (1905, 1907)
and Kossack (1911) stands out as important contributions to the knowl-
edge of this group. Looss and Liihe have likewise had a share in the
organization of the group as a unit. The most recent European work on
members of this group is that of Jegen (1917) in which he presents data on
the life history and relationship of Collyriclum faba Bremser.

2291 NORTH AMERICAN MONOSTOMES 11

In North America little has been done on this group of trematodes and
aside from the systematic arrangement of species by Pratt (1902) and the
descriptive key of Ward (1918) only isolated descriptions of species have
appeared. The earliest report of species which have been assigned to this
group are those of Joseph Leidy (1856-1895) the descriptions of which are
so meager that some of the forms must remain as species inquirendae.

MacCallum (1902) found and described Heronimus chelydrae from the
lungs of the snapping turtle. Barker and Parsons fifteen years later re-
described this form without reference to the work of MacCallum under the
name Aorchis extensus. Barker and Laughlin (1911) described Notocotyle
quinqueserialis from the intestine of Fiber zibethicus and later Barker (1916)
described from the same host the new genus and species Nudocotyle novicia.
These with the report by Cole of Monostoma faba (later named Collyricum
colei by Ward) are the major items from North America. More recently
Stunkard (1919) has shown that Heronimus chelydrae of MacCallum and
Aorchis extensus of Barker and Parsons are identical, and Tyzzer (1918)
from the study of Collyriclum colei Ward and comparison of it with the
description of Collyriclum faba Bremser by Kossack, provisionally regards
them as identical. Tyzzer's paper affords a detailed account of the
anatomy of this species and a study of the maturation, fertilization and
development of the miracidia within the eggs. His views regarding the
identity of the two forms are discussed in detail later (p. 62-65).

SPECIES PREVIOUSLY DESCRIBED FROM NORTH AMERICA

Most descriptions of North American species of this group are inade-
quate and as a result of this it is impossible to determine with any degree
of accuracy their rightful systematic position without re-examination of
the material. This has been lost in a number of cases and in other instances
all effort to locate certain specimens has proved futile. The following list
is an attempt to bring together all references to date on the species pre-
viously described from North American fresh water hosts. Names in-
dented represent later descriptions of the same form arranged in chrono-
logical order.

Monostomum ornatum Leidy 1856:43

Monostomum ornaHim Brandes 1892:504-510
Haematoloechus? Stafford 1902:724

Monostomum incommodum Leidy 1856:43
Distoma oricola Leidy 1884:47
Distoma oricola Leidy 1891:414

Monostomum affine Leidy 1858:43

Notocotyle? affine (Leidy 1858) Barker 1916:183

12 ILLINOIS BIOLOGICAL MONOGRAPHS [230

Monostomum spatulatum Leidy 1858:111

Monostomum spathulatum Leidy of Diesing 1859:426

Monostomum asperum Vaillant (nee. Nitzsch) 1863:6-7; 1863:347-348
Monostomulum asperum Vaill. of Brandes 1892:504-511
Monostomum aspersum Vaill. of Pratt 1902:966
Monostomum aspersum Vaill. of Pratt, Ward 1918:382

Monostomum mutabile Zeder of Leidy 1885:9
Cycloccelum leidyi nov. spec, of this paper

Monostomum obscurum Leidy 1887:24

Cyclocoelum obscurum (Leidy) of this paper

Monostomum amiuri Stafford 1900:402

Cyclocoelum vicarium (Arnsdorff 1908) Kossack 1911:518
Monostomum vicarium Arnsdorff 1908:362

Notocotyle quinqueserialis Barker and Laughlin 1911:261-274

Catatropis filamentis Barker 1915: PI. I, fig. 6
Catatropis fimbriata Barker 1915:190

Nudocotyle novicia Barker 1916:175-184

Eeronimus chelydrae MacCallum 1902:632-636

Aorchis extensus Barker and Parsons 1914:193-194

Collyriclum colei Ward 1917:2-3

Monostoma faba Cole 1911:42-48
Collyriclum f aba Tyzzer 1918:267-292

NEW SPECIES DESCRIBED IN THIS PAPER

CyclocoeHdae

Cyclocoelinae

Cyclocoelum leidyi nov. spec.
Cyclocoelum pseudomicrostomum nov. spec,
Cyclocoelum macrorchis nov. spec.
Cyclocoelum wilsoni nov. spec.
Cyclocoelum halli nov. spec.
Cyclocoelum triangularutn nov. spec.
Cyclocoelum cuneatum nov. spec.
Notocotylidae

Notocotylinae

N otocotylus urbanensis (Cort)

(Cercaria only known previously)
Paramonostomum echinum nov. spec.

231] NORTH AMERICAN MONOSTOMES 13

FAMILIES OF MONOSTOMIDAE

Altho in another section of this paper the monostomes are shown to
be more closely related to other groups than to each other it is thought
best to preserve the group classification until further evidence is secured
which definitely proves the suggested relationship. For this reason the
key to the monostome families is included and it is expected to serve only
for the rapid determination of specimens rather than to furnish full diag-
nostic characters.

KEY TO FAMILIES

1(2) Intestinal crura anastomosing in the posterior end

Family Cyclocoelidae

2(1) Intestinal crura ending blindly near the posterior end 3

3(4) Testes lateral to crura Family Notocotylidae

4(3) Testes within crura 5

5(6) Excretory pore posterior and terminal .... Family CoUyriclidae
6(5) Excretory pore dorsal; in anterior body half . Family Heronimidae

14 ILLINOIS BIOLOGICAL MONOGRAPHS [232

DESCRIPTION OF FAMILIES

Validity of the older name. — The question raised by Liihe (1901)
regarding the use by Looss (1899) of the family name Monostomidae
Cobbold 1866 [erroneously credited to MonticelH by Llihe (1901) and by
Stossich (1902)] without a type genus Monostomum and the extended
discussion of Looss (1902) is a needless one if the rules of the International
Code be applied. Article V of this code deals specifically with the family
name which by the acceptance of the genus name Cyclocoelum becomes
Cyclocoehdae.

CYCLOCOELIDAE Kossack 1911

Diagnosis. — Endoparasitic trematodes of large to middle sized muscu-
lar bodies. Mouth opening terminal or subterminal surrounded by
muscular sucker usually much reduced. Ventral acetabulum sometimes
present, pharynx large, muscular; esophagus long. Intestinal crura
simple or possessing internal ceca, anastomosing in the posterior end of
the body. Excretory bladder between posterior intestinal arch and end
of body with median dorso-terminal pore. Genital pore median, usually
ventral to pharynx. Copulation organs present, well developed; seminal
vesicle in cirrus pouch. Vitellaria in general lying between body wall and
intestinal crura, sometimes surrounding the latter. Genital glands between
intestinal crura, simple or lobed, forming the points of a triangle. Laurer's
canal wanting; receptaculum seminis present. Uterus strongly developed
lying in more or less regular folds between intestinal crura over which they
sometimes extend, usually filling entire space between crura. Eggs numer-
ous, without polar filament, usually containing well developed miracidia
with characteristic double eye spots.

Parasitic in body cavity, lungs and nasal cavities of water birds.

Type and only American genus Cyclocoelum. Other genera Haemato-
trephus, Hyptiasmus, Typhlocoelum, Tracheophilus, Ophthalmophagus,
Spaniometra and Bothriogaster.

The foregoing diagnosis differs from that of Kossack in noting that an
oral sucker and a receptaculum seminis have been found in two genera of
this family, namely Cyclocoelum and Haematotrephus. In the genus
Cyclocoelum these organs have been found in the species described in this
paper as well as in the following European species: Cyclocoelum prohlema-
ticum Stoss., Cyclocoelum ovopunctatum Stoss., Cyclocoelum brazilianum
Stoss., and in Cyclocoelum tringae (Brandes) as well as in Haematotrephus
similis Stoss.

233j NORTH AMERICAN MONOSTOMES 15

During the study of the Cyclocoelidae and the attempt to prepare a
key the writer was impressed with the segregation of the genera into dis-
tinct groups. These groups are here designated as sub-families. The group
composed of Cyclocoelum, Haematotrephus and Hyptiasmus are according
to this grouping left in the sub-family Cyclocoelinae created by Stossich
(1902) for the entire group. A second group is that formed by Typhlocoe-
lum Stossich and Tracheophilus Skrjabin (1913) which on the basis of the
strongly developed vitellaria and diverticula of the intestine deserve rank
as a distinct sub-family to which I desire to assign the name Typhlocoeli-
nae. As stated in a later section of this paper Cohn (1904) would ally this
group with the Fasciolidae on the basis of a rudimentary sucker found in
Typhlocoelum flavum (Mehlis). A third group is that formed by Ophthal-
mophagus and Spaniometra and is based on the position of the ovary in
the intestinal arch with the testes anterior thereto, and on the position
of the vitellaria which in these genera are ventral to the intestine. Bothrio-
gaster is also placed in this group although it presents some striking
differences, especially with respect to the vitellaria which in this genus are
markedly like those of Cyclocoelum. However with respect to the repro-
ductive glands it conforms more nearly to Spaniometra. Other peculiar-
ities of this genus will be discussed elsewhere in this paper.

KEY TO SUB-FAMILIES AND GENERA

1(11) Ovary between testes or on a level with posterior testis 2

2(8) Intestinal crura without diverticula

Sub-family Cyclocoelinae Stossich 1902 .... 3

3(6) Right and left sides of vitellaria separated at posterior end. . . .4
4(5) Uterus loosely folded; long loops surrounding genital glands. . . .

Haematotrephus Stossich 1902

5(4) Uterus compact; loops short not surrounding genital glands. . . .

Cyclocoelum Stossich 1902

6(3) Right and left sides of vitellaria continuing into one another at

posterior end 7

7 Uterine loops extending laterally over intestinal crura and

vitellaria to body wall Hyptiasmus Kossack 1911

8(2) Intestinal crura with distinct diverticula

Sub-family Typhlocoeline .... 9

9(10) Testes strongly lobed Typhlocoelum Stossich 1902

10(9) Testes round Tracheophilus Skrjabin 1913

11(1) Ovary posterior to testes, situated in intestinal arch

Sub-family Ophthalmophaginae .... 12

12(15) Vitellaria ventral to crura 13

13(14) Testes in posterior body half .Ophthalm.ophagus Stossich 1902
14(13) Testes in anterior body half Spaniometra Kossack 1911

16 ILLINOIS BIOLOGICAL MONOGRAPHS [234

15(12) Vitellaria lateral to crura

Ventral acetabulum conspicuously developed

Bothriogaster Fuhrman 1904

Up to the present the only genus in this family represented in North
America is Cyclocoelum and for this reason the other genera are not dealt
with in the following pages.

CYCLOCOELUM

Historical. — The question of the type for this genus is indeed a com-
plicated one and as Stiles (1908) has said "well represents a ship without
a rudder." Liihe accepts the designation of Hoyle (1888) and takes issue
with Looss who by elimination accepts Monostomum prismaticum as type.
Stiles (1908) presents a third view in which he says that the name Mono-
stomum is a synonym to Festucaria by priority rule, since Zeder deliber-
ately renamed the genus Festucaria Schrank (1788).

A critical study of the literature involved brings to light certain facts
on this point which seem to the writer to be worthy of space here. The
name Festucaria was proposed by Schrank (1788) for two species Festu-
caria anatis and Festucaria strigis. These species were described by Goeze
(1782) in Klasse II of his family (genus) Planaria as "rundlichte oder
walzenformige Plattwurm" of which he says there are two genera. Only
one of them is of concern here and is described by Goeze (1782:173-174) as
having a single mouth, ("mit einfacher Mlindung").

From the context of the description one is led to believe that these
worms were collected by Goeze since he says, "Ich, * * * habe sie nur
in zwo Arten von Vogeln gefunden; in einer zahmen Ente, und in einer
grauen Weideneule." He follows this with a description of the worms in
situ as well as when he placed the intestine of the host in water where the
worms "loosed themselves."

The type of the genus Festucaria becomes by page precedence Festu-
caria anatis Schrank (1788). This purports to be the same worm described
by Goeze (1782:174) as follows: — "In den Dedarmen einer Ente sassen
hin und wieder kleine gelbliche Knotgen. Da ich einige abnehmen wollte,
merkte ich, dass sie vest anklebten. Als ich sie ins Wasser brachte, gaben
sie sich los, und ich erkannte sie fiir rundlichte Plattwiirmer mit einer
Miindung." The description of his figure 9 plate 13 adds "(a) die einfache
Miindung mit 2 Punkten; (b) eine Art von Maul dariiber." From the
description and the figures this appears to be the crown of spines which
Goeze mistook for a mouth.

The description of Schrank (1788), based on Goeze (1782), runs as
follows : —

Splitterwurm — Festucaria

Ein langlichter, einformiger Wiirm vorne mit einer einzigen Sauge-
miindung.

235] NORTH AMERICAN MONOSTOMES 17

"Enten Sp. 54, Cylindri-formed, columnar, walzenformig." "Zween
erhabene Punkte in der Mundoffnung, Festucaria anaiis." "Goeze Eingew.
174 tab. 13, figs. 8-11; in Enten."

Gmelin (1790) classifies the same worm as Fasciola anatis. Zeder (1800)
renames the genus Festucaria, Monostoma, and includes under the name
the following species;

Monostoma elaphi

Monostoma prismaticum

Monostoma ocreatum

Monostoma miitabile

Monostoma verrucosum

Monostoma bombynae
In the same work the species of Schrank {Festucaria anatis) (Fasciola of
Gmelin) is described as an Echinostome (Distoma anatis). Rudolphi (1801)
recognized the same genus Festucaria Schrank with Monostoma Zeder as
a synonym and later in the same paper (p. 62) stated that he permits the
name Festucaria to stand even though Zeder had proposed a new one.
This name he retained until 1809 when he rejects it for the denomination
of Zeder which he says "omnino praeferenda sit." In his later paper
(1819) Rudolphi still adhered to the genus name of Zeder. Subsequent to
this time helminthologists have accepted without comment the genus
Monostoma of Zeder. Consequently Hoyle (1888) de ignated Monostoma
mutabile as type of this genus. Four years later Brandes (1892) in his
Revision der Monostomiden grouped together what he beheved to be
members of this genus, "als gute Arten," Monostoma mutabile Zeder, M.
flavum Mehhs, M. arcuatum Brandes, M. tringae Brandes and M. ellipti-
cum Rudolphi and suggested their separation from the remaining species
under the name Cyclocoelum with the following diagnosis: "Diese fiinf
Spezies scheinen ihrer Organisation und Lebensweise nach zusammenzu-
gehoren, jedenfalls die ersten 4, die sammtlich in der Leibes- oder Infra-
orbital-hohle von Wasservogeln schmarotzen, wahrend M. ellipticum in
der Lunge von Rana maculata gefunden wurde, * * * . Die Enden der
Darmschenkel mit einander verschmelzen, sodass der Darmtractus einem
Ring darstellt." Looss (1899) accepted the revision of Brandes and
named M. mutabile as type of the genus of Brandes. Later in the same
notable work the author used the term Monostomidae to characterize a
certain group of worms. But his contemporary Liihe (1900) in a re-
view of the work of Looss objected on the ground of the nomenclature law
to the use of the family name Monostomidae without a type genus Mono-
stoma and called attention to the priority of the "vollig verschollenen"
genus Festucaria Schrank. One year later Looss (1901) in explanation
and confirmation of his earlier work showed by elimination that Mono-
stomum prismaticum is type of the genus Monostoma Zeder. In regard to

18 ILLINOIS BIOLOGICAL MONOGRAPHS [236

the right of priority of Festucaria Schrank he stated that he found no
reason to call back from oblivion that absolutely meaningless name.
"Ich habe mich bisher nicht veranlasst gef uhlt, diesen auch mir bekannten,
aber in der That ganzlich der Vergessenheit anheimgefallenen und prak-
tisch absolut bedeutungslosen Namen wieder ins Leben zuriickzurufen;
ich empfinde dafur auch heute noch keine Neigung und iiberlasse deshalb
die Entscheidung der Frage mit Vergnugen denjenigen, die sich mehr
dafur interessieren," Liihe continued the discussion on the basis of the
priority of Festucaria Schrank and the deUberate renaming of Zeder. He
stated, however, that he believes Festucaria anatis Schrank (based on Goeze
1782, Taf. 13, Figs. 8-11) to be Echinostomum echinatum. However Liihe
continued to use the genus name Monostoma Zeder.

Stossich (1902) accepted Monostoma mutahile Zeder (1800) as the
earhest record of the group and approved its removal by Brandes (1892)
to the genus Cyclocoelum. He does not consider the priority of Festucaria
or the renaming by Zeder (1800) but Zeder says distinctly that Schrank
(1788) placed those worms with one sucker in a genus under the name
Festucaria and that he (Zeder) in conformance to the German system of
nomenclature wished to introduce the Greek term Monostoma which he
considered a more appropriate and characteristic name.

Kossack (1911) cited the positions of Hoyle, Luhe, Looss and Stiles
previously mentioned; he then avoids the real situation by accepting only
the more generally recognized works and thus believes himself dealing
with a necessity, and also to be in full accord with the law of priority.
On this basis he omits from the discussion the genus name Festucaria and
substitutes for the old name Monostoma the more significant name
Cyclocoelum.

It seems to the writer that the case at hand is clearly covered by
Article 32 of the International Rules of Zoological Nomenclature; and
after a careful study of the case the writer is led to accept Stiles' view and
to abide by the ruling of the International Commission according to which
the genus Monostoma Zeder (1800) becomes a synonym of Festucaria
Schrank (1788). The fact that Zeder did not include the original species,
Festucaria anatis, in his genus does not affect the case since the genus
name would remain with the type species; hence the redescribing of this
form by Zeder under the name Distotna anatis does not change the case
since in that event Festucaria falls with Monostoma into synonomy. Then
Festucaria strigis having been removed to the genus Strigea, whether
Festucaria anatis, is a distome (Zeder 1800), or an echinostome (Zeder
1803, Rudolphi 1809, and Liihe 1901), the case remains the same and can
be analyzed only as a direct renaming of the original form. Accordingly
the species of Monostoma Zeder fall in the genus Festucaria Schrank 1788.
The type is clearly Festucaria anatis.

237] NORTH AMERICAN MONOSTOMES 19

The opinion expressed by Liihe (1901) that he believes Festucaria ana-
tis to be an echinostome, is indeed not a new view for in fact it is suggested
in the description of Goeze when he describes the orifice as having above it
a sort of a mouth. As was mentioned earlier in this work Goeze probably
erroneously described the crown or ring of spines for the mouth. Also
his description of the worms in situ in the intestine of a domestic duck
suggests the probability of an echinostome. Zeder (1800) evidently
recognized the distome nature of these worms for he described in his
group Echinis under the name Distoma anatis worms which he terms
identical with Cucullanus conoideus Bloch (1782), Planaria enten sp.
Goeze (1782), Festucaria anatis Schrank (1788) and Fasciola anatis Gme-
lin (1790). Characteristic points of his description are the large aceta-
bulum and the ring-like swelling around the head armed with spines.
In 1803 this worm was described by the same author as Distoma echinatum.
Rudolphi (1809) described Cucullanus conoideus Bloch, Planaria (teres
poro simplici) Goeze, Fasciola anatis Gmelin, Festucaria anatis Schrank,
Distoma anatis Zeder (1800) and Distoma echinatum Zeder (1803) as identi-
cal. Likewise Diesing (1850) and Baird (1853) interpret Festucaria anatis
as identical with Distoma echinatum. This with the opinion of Liihe
already given appears to be sufficient evidence to determine the probably
Echinostome nature of Festucaria anatis Schrank. In any event the species
described by Zeder (1800) as Monostoma mutabile which we now know as
Cyclocoelum mutabile is quite strikingly different anatomically and hence
clearly not closely related to Festucaria anatis based on the opinions of Zeder
(1800, 1803), Rudolphi (1809, 1819), and Liihe (1901). From a study of
the literature the writer is in agreement with the opinion of Liihe, Rudolphi
and Zeder that Festucaria anatis is probably Echinostomum echinatum
Rud. (1809). Whether these authors studied the same material can be
only a matter of conjecture.

An additional fact, however, is furnished by our present knowledge of
the normal habitat of these worms which with few exceptions are confined
to partially closed cavities of the body, the infraorbital sinus, and areal
sacs for Cyclocoelum and Hyptiasmus, and the trachea for Typhlocoelum.
As has been previously noted the habitat of Festucaria anatis is the intes-
tine of a domestic duck.

The genus Cyclocoelum Brandes (1892) was formulated to include the
following species Monostoma mutabile, M. flavum, M. arcuatum, M.
tringae, and M. ellipticum with the following description: first four species
collected from the body cavity and infraorbital sinus of water birds the
fifth Monostoma ellipticum, from the lungs of Rana maculaia, with intes-
tinal crura anastomosing in the posterior end.

In the acceptance of the genus name Cyclocoelum Brandes (1892) the
writer realizes that he is subject to criticism on account of the genus

20 ILLINOIS BIOLOGICAL MONOGRAPHS [238

Cyclocoela Dujardin (1845) and in view of this fact calls attention to the
recommendation of the International Zoological Commission in the Inter-
national Code which reads: — "It is well to avoid the introduction of new
generic names which differ from generic names already in use only in ter-
mination or in a slight variation in spelling which might lead to confusion.
But once introduced, such names are not to be rejected on this account."

Structure of the Genus

The description of Stossich (1902) states that these worms are covered
with spines which that author interprets of sufficient importance to be a
specific characteristic. Although Zeder (1803) observed these granula-
tions he was not able to determine their real nature and suggests that they
are either cellular structures or glands underlying the thin cuticula. It
was not until the work of Fuhrmann (1904) was published that the true
nature of these, which are really pits became known. This author de-
scribed and figured these pits as found on the ventral surface of Bothrio-
gaster variolaris and stated that he has observed similar depressions on
both the dorsum and venter of Cyclocoelutn mutabile. However, he raises
the question whether these are not a product of preservation. Kossack
(1911) verified the work of Fuhrmann by similar observations on several
species belonging to at least three distinct genera of this family. He sup-
ported the view of Fuhrmann as to their origin and added as evidence of
their nonexistence in living material that the authors who had opportunity
to study living material — notably von Siebold and Van Beneden — did not
mention their presence.

There can be no doubt that Zeder (1800, probably also 1803) made
observations on living material. That the particular observation men-
tioned above was made from living material of course is only a matter of
conjecture. It is evident, however, that at least one author who studied
living material did observe these pits. Hence the lack of mention by von
Siebold and Van Beneden is not sufficient evidence to prove their non-
existence in living material.

That the state of contraction at the time of fixation is responsible for
the degree of depth and apparent frequency of these pits remains unques-
tioned. But unless they occur in the living specimen it would be impos-
sible to find them so regularly in preserved material. In addition if they
are artifacts of preservation one would expect to find them in other trema-
tides of similar size and structure.

While the writer has not had opportunity to study living Cyclocoelidae
he has found the above mentioned "Grubchen" of Kossack, the "ovale
depressionen" of Fuhrmann, constantly in more than one hundred and
fifty specimens belonging to at least nine species of the genus Cyclocoelum
and in Eaematotrephus similis. While in the study of more than a hundred

239] NORTH AMERICAN MONOSTOMES 21

Specimens of Heronimtis chelydrae MacCallum (1902) both living and pre-
served not a single instance has been found. For these reasons the writer
feels safe in saying that these pits are characteristic of the living animal
and are only emphasized by the state of contraction at the time of preser-
vation. The various worms being fixed in different states of contraction
would consequently show these pits more conspicuously in the more con-
tracted specimens.

The body wall is composed of at least five layers. From outside inward
they are as follows, cuticula, basement membrane, circular muscle, longi-
tudinal muscle, and epithelium (Fig. 25). These compose what is com-
monly known as the dermomuscular sac. The disposition of the parts of
the dermomuscular sac of this group agrees in most respects with the inter-
pretation of Monticelli (1888) and Blochmann (1896). It differs however
from the observation of Fuhrmann (1904) in which he says that the body
musculature is differentiated into outer longitudinal, inner circular muscle
layers and inside of this the layer of bands of diagonal muscles, in that
the outer muscle layer is formed by circular muscles (Fig. 25). The single
statement of Fuhrmann mentioned above is so strikingly different from
all comprehensive works on this subject that the writer is lead to believe
that it is a lapsus calami and that in reality the muscle layers of Bothrio-
gaster variolaris are identical with those of other trematodes.

Zeder (1803) states that these worms have a single sucker on the for-
ward end. His description of this organ is scanty and lacks the points
which distinguish the sucker from the pharynx so that one is lead to believe
in the light of present knowledge that he interpreted the pharynx in the
Cyclocoelidae to be the same as the sucker in the Notocotylidae. Von
Siebold (1835), the first to give a clear account of the anatomy of the
Monostomidae in his description of Cyclocoelum {Monostomum) mutabile
(Zeder), speaks of the mouth as a transverse oral opening leading to a
funnel shaped canal which narrows gradually posteriorly and terminates
in the so-called pharynx. No trace of a sucking organ was observed by
this author.

Following this Van Beneden (1858) referred to the above work fre-
quently but stated that the monostomes have only a mouth sucker situated
in the anterior region. In another paragraph of the same work he speaks
of the digestive system of Trematodes as showing generally an anterior
sucker in the bottom of which is situated the mouth. This he says opens
into a second enlargement similar to the preceding sucker, the pharyngeal
bulb. In his figures of Cyclocoelum {Monostomum) mutabile (Zeder) the
structure termed pharyngeal bulb above is indexed as buccal bulb. These
show the pharyngeal bulb with no anterior sucking musculature surround-
ing the mouth opening. In a later paper (1861) the same author describing
again this same species spoke of the bulb and the region preceding it which

22 ILLINOIS BIOLOGICAL MONOGRAPHS [240

he said is seen with difficulty. A little later in this work in the description
of Notocotyle {Monostomum) verrucosunt he employed the term "la ven-
tuose anterieure ou plutot le buccal," applying it to the spherical muscular
bulb at the extreme anterior of this worm a Notocotylid, evidently mistak-
ing this structure for the same structure termed the pharyngeal bulb in
the earlier work. His descriptions show clearly that the same organ which
in the distomes is termed pharynx is here termed anterior sucker or buccal
bulb.

Some years later Monticelli (1892) described the mouth as small in
Monostomum mutabile and Monostomum expansum; of greater or less size
in Ogmogaster plicatum and Monostomum galeatum; circular in Monostomum
hippocrepis and Monostomum trigonocephalum ; ellipsoidal in Monostomum
cymhium and Monostomum ornatum. It is usually ventral and generally
situated in the extreme anterior end. When present a prepharynx of
variable length is situated directly in front of the pharyngeal sucker, the
"anterior sucker or buccal bulb" of Van Beneden. Monticelli thus dis-
tinguishes between the funnel shaped tapering canal of von Siebold (1835)
and the adjoining posterior structure; and designates it as a prepharynx.
He says that in Notocotyle and some other genera of this family the
prepharynx is wanting and that then the pharynx is anterior and plays the
role of an anterior sucker. For this reason he designates this structure in
these genera as a sucker pharynx.

Braun one year later, refers to the description of Monticelli and sug-
gests that a sucking organ has been developed out of the pharynx. In
1901 the same author refers to the "bulbus buccalis" of Van Beneden, or
pharynx of Monticelli, as a Mundsaugnapf which he says is followed by
the esophagus. In another paragraph of the same work when describing
Monostomum trigonocephalum Rud. (since removed to the genus Prono-
cephalus by Looss) collected from the intestine of the sea turtle he says
that the sucker is 0.12 mm long and 0.09 mm broad and again states
that it is followed by a straight esophagus 0.3 mm long, without a phar-
ynx. Thus Braun has construed the muscular structure at the extreme
anterior in the Notocotylidae, Pronocephalidae and other families of this
group to be a development of a structure similar to that termed pharynx
by Monticelli in the Cyclocoelidae.

Barker and Laughlin (1911) accept this view without comment and
describe the worms Notocotyle quinqueserialis as clinging to the intestine
of the muskrat tenaciously with the well developed oral sucker. They
found no evidence of a pharynx.

Taschenberg (1879) describes the mouth in the genus Didymozoon
as an opening followed by a funnel shaped duct leading to the pharynx.
This he states to be generally characteristic of the entire group. Lonnberg
(1891) found in Didymozoon lampridis the well developed sucker (pharynx of

241] NORTH AMERICAN MONOSTOMES 23

Taschenberg) and just posterior to it a very small muscular bulb the phar-
ynx. Odhner (1907) finds in Didyniozoon scombri Tschbg. a similar pharyn-
geal bulb which he figures and proves beyond doubt that the pharynx of
Taschenberg is a very strongly developed sucker followed by an extremely
small pharynx. In this same notable work he says that in Cydocoelum
(Monostomum) mutabile (Zeder) and other parasites where only a pharynx
is present that there is always a region anterior to it which he terms the
"Mundrohr" or "Mundhohle" (prepharynx of Monticelli), a structure
which by the contraction of the inner walls changes the pharynx into a
sucker. This same region is shown in the figures of Odhner for Didynio-
zoon scombri Tschbg.

Looss (1899) speaks of the mouth sucker but gives no equivalent for
the pharynx of Monticelli. Later, however, he interprets the swelling at
the beginning of the esophagus in Microscapha reticularis as a pharynx.
Cohn (1904) calls this to account when he states that this swelling is
nothing more than the esophageal sphincter which is present in many
species. The same author interprets the oral sucker of Looss, or buccal
bulb (anterior sucker) of Van Beneden, as a pharynx and adds that it is
his opinion that soon monostomes will be found with a well formed sucker
adjoining a typical pharynx. The evidence given in support of this con-
sists of the statement that Haplorchis cahirinus Looss has a strongly devel-
oped pharynx preceded by a rudimentary sucker and that he has observed
in Cydocoelum (Monostomum) mutabile (Zeder) and in one other species
(to be published later) a rudimentary sucker. On the contrary Looss
(1899) figures Haplorchis cahirinus with a well developed but small oral
sucker followed by a somewhat smaller but perfectly developed pharynx
and in his description of this species states specifically that the oral sucker
and the pharynx are well developed structures. He regards the rudimentary
structure occurring on the ventral side as an acetabulum. He adds also
that the nerve commissure which according to Braun lies in all the Digenea
more or less bent around the dorsal side of the oral acetabulum and the
pharynx, is in front of the muscular sucking organ in the Monostomidae
and therefore that organ is a true pharynx, tho this organ serves both as a
sucker and a pharynx.

Stossich (1902) calls the pharynx of Monticelli an "inner sucker" which
he says serves the same function as the mouth sucker of other trematodes.
Odhner (1907) states that those that acquire holdfast organs in the
least degree are the parasites of the respiratory organs; the group contain-
ing Cydocoelum mutabile being entirely suckerless. Kossack following the
decision of Monticelli says that the question can be determined only by
a study of the position and distribution of the nerve ganglia. Conse-
quently he termed the anterior muscular structure a true pharynx. Ward
(1918) calls the same structure the oral sucker and says that no pharynx is

24 ILLINOIS BIOLOGICAL MONOGRAPHS [242

present. The interpretation of Braun, Looss, Ward and others is indeed
well exemplified in many cases where the anterior region is telescoped
posteriad over the opened anterior portion of the pharynx (Fig. 46).

Records of an oral sucker in this group are few, aside from that
found in Cyclocoelum mutabile by Cohn (1902). Wedl (1857) gives an ac-
count of the oral sucker in Monostomum lanceolatum as follows: — "Der
kleine Mundnapf liegt an der Bauchseite des zugeschmalerten Vorder-
theiles des Thieres (Fig. 15a) und ist nach riickwarts von einem dickfleisch-
igen Bulhus oesophagus (b) begrenzt, * * * >' These with the account
of the writer (Harrah 1921) in which the oral sucker was described
in two species of this genus constitute the evidence produced to demon-
strate the presence of the oral sucker in this group.

In the light of the foregoing the question brought out by a long con-
tinued controversy remains unsettled. Is the structure termed the phar-
ynx by Monticelli (1892) phylogenetically a pharynx or an oral sucker?
While Monticelli attempted to prove by the distribution of the anterior
nerves that the muscular bulb, or pharynx as he termed it, was a true
pharynx, this has not been generally accepted and hence remains a matter
of much controversy. In the opinion of the writer the brain commissure
which lies distinctly anterior to the pharynx (Fig. 8) can be used as a land
mark only and in a different state of contraction might have its relative
position changed. The innervation is no doubt distributed to the other
anterior structures as well. Although the nerve commissure has the
same relative position in the distomes this alone does not prove the phy-
logenetic origin of the pharynx, and when a muscular sucking apparatus
is found and proof established of such an organ anterior to and adjoining
the pharynx, as predicted by Cohn (1904), then and only then can these
organs be safely designated as oral sucker and pharynx.

In this study the writer has examined more than one hundred speci-
mens of the genus Cyclocoelum Brandes comprising at least fifteen different
species. In this material different conditions are found. In Cyclocoelum
obliquuni Harrah 1921, Cyclocoelum halli nov. spec. Cyclocoelum obscurum
(Leidy), Cyclocoelum triangiilarum nov. spec, a very weak and scarcely
distinct oral sucker is present. The concentration of tissue is scarcely
discernible except under the best optical conditions and even in sections
there appears only a concentration of tissue at this point (Figs. 36-42).
The outer circular band or sheath is found to be very light and not a con-
tinuous band as in Cyclocoelum elongatum. The above condition has
been found to obtain in Cyclocoelum problematicum Stoss. and Cyclocoelum
tringae (Brandes). In these species the sucker musculature is easily over-
looked and when not taken into account the mouth opening agrees well
with the structure so clearly described by von Siebold (1835) and van Bene-
den (1858) and that named prepharynx by Monticelli (1892), On the

243] NORTH AMERICAN MONOSTOMES 25

Other hand in Cyclocoelum pseudomicrostomum an intermediate condition
is found. On first observation the mouth opening appears like the one
just described; however, on more careful scrutiny a light but well formed
sucker is discernible. In this case, however, the longitudinal and oblique
muscles are actually increased immediately surrounding the buccal duct
which leads inward toward the very muscular pharynx (Fig. 43). From
this more or less muscular wall, radial muscles extend outward having their
origin in a weak band of circular muscles. This outside covering of the
sucker is held in place by the same sort of transverse muscles as in the
distomes. Were this the only case found one could perhaps accept Cohn
(1902:715) who has observed what he terms a rudimentary mouth sucker
in Cyclocoelum mutabile (Zeder).

The maximum condition observed by the writer was that found in
Cyclocoelum elongatum, in which posterior to the opening of the mouth,
which is downward as before, is seen a large weak oral sucker scarcely vis-
ible in toto preparations. It is from one-third to one-half larger than the
pharynx posterior to it and extends from the extreme anterior of the
animal to well over the anterior portion of the pharynx. It measures
314/i in length by 463/z in width. The musculature is much less strongly
developed than that of the pharynx and consists of an outer circular layer
connected by radial, longitudinal, and oblique muscles to a much heavier
inner circular band which forms the muscular walls of the mouth (Figs.
44, 45, 47, 48). This muscular body is suspended by much lighter strands
of transverse muscle having their origin in the musculature of the body
wall and their insertion at times in the outer circular band of muscles
covering the sucking musculature and again in the radial muscles of the
sucker itself. In general the position of this sucker is such that it opens
downward but suspended as it is a slight contraction of the dorsal sus-
pensory muscles and at the same time a relaxation of the ventral ones
could easily give to the sucker a different position so that its aspect would
be changed from that of its true antero-ventral one (Fig. 29).

In all specimens studied the writer has found evidence of the oral
sucking mechanism and believes it to be a universal character in this
family. In a study of Eaematotrephus similis Stossich a sucker almost
as heavy as that of Cyclocoelum elongatum was found (Fig. 28), and in the
former species the sucker is considerably more prominent than in Cyclo-
coelum pseudomicrostomum and Cyclocoelum mutabile. Cyclocoelum micro-
stomum could not be obtained for study of this feature.

Following the oral sucker is a thin walled, lightly muscular tube extend-
ing posteriad and ventrad to the pharynx and opening into it on the
ventral side. The writer believes this to be a condition due to the state of
contraction at the time of preservation and that in a fully extended speci-
men the oral opening would enter the pharynx from the anterior face,

26 ILLINOIS BIOLOGICAL MONOGRAPHS [244

thus leaving some 150 to 200 ju between the oral sucker and the pharynx.
This portion is termed the prepharynx (Fig. 29).

As will be seen by the description which is to follow later the structure
of the pharynx is practically identical in many of the species of this and
other families and therefore must necessarily have developed from a
similar tissue in a similar manner. This being the case there remains the
functional differentiation which may come about in the absence of a well
developed oral sucking structure. No doubt the pharynx aids the weak
sucker in drawing in food and it is highly probable that the suction pro-
duced by these heavy muscular walls is indeed very great. As was stated
earlier in this discussion the position of the oral sucking apparatus relative
to the pharynx in all those species which do not possess a strong oral
sucking apparatus strongly indicates this view.

As was stated before the muscular bulb, or pharynx of Monticelli, is
identical in structure and has no doubt arisen in the same manner in all the
species of this family. Even though it may function as a sucking organ in
some instances this structure has not been modified and does not appear
to be typical sucker tissue as it has been described in other trematodes.
If, however, this muscular organ is present in one or more species of this
genus at the same time with a distinct anterior musculature which is
typical of that tissue as found in other groups of trematodes and is con-
stant in these species, then one is justified in designating the anterior
musculature a sucking organ or oral sucker and the musculature posterior
to it a pharynx as it was termed by Monticelli (1892).

Immediately posterior and dorsal to the prepharynx is a muscular
structure developed around the wall of the posterior prepharynx or an-
terior esophagus. On first observation the pharynx appears as two bean-
shaped halves lying one on either side of the anterior portion of the
esophagus. On more careful study, however, it is found to consist of heavy
muscular halves bound together on the edges by smaller bands of fibers
so that in the true cross section it appears as a cylinder with an elongated
dorso-ventral slit passing through it.

The pharynx is variable in size and form within the species as well as in
different species, and may be in some species distinctly elongated while
in others it is noticeably expanded laterally and still in others it is spherical.
It measures in Cyclocoelum pseudomicrostomum 77SiJ. and is slightly longer
than broad. In Cyclocoelum ohscurum it is in general spherical and meas-
ures 215 to 298 )U. In Cyclocoelum macr orchis it is distinctly longer than
broad and measures 264 to 314;u in length by 198 to 248/x in width. In
Cyclocoelum elongatum the pharynx is oblong measuring 264 to 331^1 in
length by 215 to 281 /x in width. While the range of measurements gives
a general idea of the size and shape of the pharynx this study has shown

245] NORTH AMERICAN MONOSTOMES 27

that th,e individual which has the longest pharynx is not always the one
which has the narrowest one and vice versa.

As stated above the size and form of the pharynx is not constant as
might be indicated by an average or range of measurements. Cydocoelum
ohscurum shows eight instances in which the pharynx is round, two with a
greater width than length, one with a greater length than width; this gives
an average for eleven specimens of 230/i in length and 231 )u in width,
agreeing closely with the record that eight of the eleven cases cited show
the pharynx to be spherical. From the range one would infer from the
maximums that the pharynx is spherical but from the minimums of less
transverse diameter than longitudinal. In Cydocoelum macrorchis and
Cydocoelum elongatum every specimen measured shows for the pharynx
a greater length than width while in Cydocoelum pseudomicrostomum and
Cydocoelum ohscurum some were spherical while the majority have a
greater length than width. From this comparison one can see readily that
the pharynx possesses such variability in size and form in this genus that
it can be termed at best only spheroidal.

The musculature of the pharynx is, as stated previously, similar in
all the species of this genus and consists of numerous fibers bound into bun-
dles that are so interlaced as to make a very powerful organ and in the
absence of the strong oral sucker it is quite probable that it serves as a
sucking organ as was stated earlier in this work.

The musculature is quite characteristic of this organ as it has been
described in other trematodes and consists of circular, radial and longi-
tudinal fibers. The circular muscles are most numerous and constitute
approximately 75 percent of the entire structure. They are pierced by
bundles of radial muscles which have their origin in the outer layer of
circular muscles and their insertion in the inner layer of the same muscles
whose fibers intertwine all the muscles of the circular and radial type, par-
ticularly at their origin and insertion.

In most cases this muscular bulb stands with the anterior end open
thus forming a continuation of the funnel shaped mouth opening to the
posterior end of the pharynx which in all instances observed by the writer
is closed just anterior to the esophagus that leads caudad from this organ.

The esophagus is a thin walled tube of varied length. In Cydocoelum
elongatum it measures 347 to 463/x; in Cydocoelum ohscurum 331 to 662)u;
in Cydocoelum halli 483ju on the average. The wide range of variation in
length is due to the fact that the esophagus takes an S-shape (Fig. 29)
which is no doubt due to a state of partial contraction taken at fixation.
This condition makes it impossible to secure the exact length of this organ.
Because of the different states of contraction the esophagus is more sinuous
in some specimens than in others and consequently shows a much greater
variation in length. In view of this fact the writer can place very little

28 ILLINOIS BIOLOGICAL MONOGRAPHS [246

weight on the exact length of the esophagus as a specific characteristic.
In fact in some cases where the toto specimen showed the esophagus to be
very short, sections of the same specimen gave evidence of a relatively
long but much folded tube due no doubt to contraction.

In the case of Cyclocoelum obscurum stated above the length varies
from 298 to 662/i making an average for the eleven specimens of 419m.
Under the conditions mentioned the average does not represent the actual
length of the esophagus. It was noted from a study of this collection that
the esophagus of one individual was more than twice as long as that of
another. However, the maximum length of the esophagus is not repre-
sented by the maximum measurements secured since in no case has the
writer observed the esophagus when it could be said to form a straight
line from the oral sucker to the pharynx. The normal condition is repre-
sented in figure 29 where the esophagus arises at the dorsal portion of the
pharynx and from there takes a winding course both laterally and dorso-
ventrally to the intestinal bifurcation into which it empties directly from
the ventral side of the latter organ. The esophagus bifurcates at the
posterior portion of the first body sixth to form the intestinal crura. One
branch passing to the right, the other to the left side of the body, thus
forming an arch which lies in the anterior one-sixth of the body. The
crura extend from this parallel to the body wall, being separated from it
only by the vitellaria, to the extreme posterior end where they anastomose
forming a posterior arch. This is separated from the posterior body wall
by the excretory bladder. The crura are usually large with a Ifirge lumen.
However, in some cases they are extremely irregular and show in a few
instances more or less distinct pouches which in some individuals appear
as distinct diverticula and in fact are as strongly exemplified as some of
those shown by Stossich (1902) and Kossack (1911) for the genus Typhlo-
coelum. This feature, however, is not constant for any species of the genus
Cyclocoelum observed by the writer and is probably due to the pressure
produced by expanded uterine loops which fill out the space between the
crura and in many instances reach over the crura. Anterior to the intes-
tinal bifurcation in Cyclocoelum macrorchis and Cyclocoelum obscurum
there is an evagination which makes a pronounced undivided neck to the
crura into which the esophagus opens. This appears in every specimen of
the fifty-six in the two collections. This condition is in decided contrast to
the other species studied. In these specimens, however, the intestinal
crura are comparatively empty and show a generally relaxed condition
whereas in most of the other specimens the crura are well filled and appar-
ently well extended. It is probable in view of the relaxed and empty
condition of the crura of the former species that the neck of the bifurcation
is the result of the condition rather than a constant character for the
species.

247] NORTH AMERICAN MONOSTOMES 29

The excretory system of these worms cannot be fully made out in
preserved material and at most one can obtain accurate knowledge of only
a small part of this system without having had opportunity to study
developmental forms while living. In general the excretory bladder can
be made out in all species as a simple flattened sac, except in Cyclocoelum
elongatum, lying between the posterior arch of the intestine and the body
wall, usually closely approached on either side by the posterior extremities
of the vitellaria. It opens to the exterior on the median dorsal surface by
a small duct surrounded at its exterior opening by a strong sphincter.
Two main branches of the system open into the excretory bladder one on
either side. These branches follow in general the course of the intestinal
crura and are joined by many secondary branches which anastomose
freely forming a vast net-work ramifying the entire body. The ducts of
the system are composed of relatively heavy muscular walls which aid
in the movement of the excretory products toward the bladder.

The nervous system so far as has been made out from the material at
hand does not diiifer materially from the description of Lang (1880). The
cephalic ganglia are situated one on either side of the pharynx and are
connected by a dorsal commissure which spans the anterior alimentary
organs between the pharynx and the sucker. In the miracidia are seen the
beehive-shaped eye-spots first observed by Van Beneden (1861) and later
described by Faust (1918).

The vitelline glands in this family are very much alike in all genera
being composed of two main canals which lie parallel to and usually outside
of the intestinal crura. From these main stems side branches go out both
dorsally and ventrally. The glands are made up of small follicular bodies
arranged around the ducts so as to form grape-like clusters. The develop-
ment of these glands is constant within a species and may be built up on
the main stem with relatively few branches. The strongest development
of the secondary branches is found in Cyclocoelum niicrostomum and Cyclo-
coelum pseudomicrostomiim and in the genus Hyptiasmus where the
branches of the vitelline glands form a net-work closely applied to the
intestinal crura. The vitelline substance is conveyed to the ootype by a
duct from each side in the region of the ovarian complex; these pass to a
point posterior to the shell gland where they unite. At the point of union
is usually found an enlargement, the vitelline reservoir. The duct then
passes to the dorsal side of the shell gland where it becomes embedded in
the latter organ; however, it emerges from this to re-enter it on the anterior
dorsal surface and just after its entry joins the oviduct at the point of its
enlargement to form the ootype.

The cirrus pouch as stated by Kossack (1911) shows little variation.
It is a clubshaped muscular pouch containing a relatively large vesicula
seminalis to which unites a short weakly developed pars prostatica. This

30 ILLINOIS BIOLOGICAL MONOGRAPHS [248

gives rise to a relatively large cylindrical unarmed ductus ejaculatorius.
In most individuals the ductus ejaculatorius and pars prostatica are much
coiled inside the cirrus sac and can be made out only with considerable
diflSculty. The writer has had the good fortune to have for study two
specimens of Cyclocoelum brazilianum Stoss. one of which has the ductus
ejaculatorius protruded from the body. The other one with the ductus
ejaculatorius extended into the uterus in a case of evident self-copulation.
(Figs. 31, 35). In these specimens the ductus is in excellent position for
study and appears as described above. Through the union of the end
portion of the cirrus pouch and the uterus a relatively large genital atrium
is formed.

The form of the genital glands throughout the family is in general very
similar. They are usually round or elliptical, sometimes flattened from
pressure of the surrounding parts, with the exception of Cyclocoelum vicar-
ium (Arned.) and in the genus Typhlocoelum in which species the testes
are lobed. The genital glands of the genus Cyclocoelum, the only Amer-
ican genus thus far known belonging to this family, are spherical in form.
The testes are located so the posterior is in or near the posterior intes-
tinal arch and the anterior, a greater or less distance from this, separated
often by uterus loops. Exceptions to this are found, however, in Cyclo-
coelum oculobium (Cohn) and Bothriogaster variolaris Fuhrmann in which
the ovary occupies the posterior intestinal arch while the testes are situ-
ated in the middle region of the body. The vasa efferentia given off from
the testes unite a short distance anterior and median to the anterior
testis in Cyclocoelum obscurum, to form the vas deferens which makes its
way between the uterine loops to the cirrus pouch previously described.
The ovary is spherical and communicates by a short duct to the compact
closely lying shell gland.

The presence of the receptaculum seminis and Laurer's canal have been
held in question since the earliest accurate work on the anatomy of these
worms, that of von Siebold (1835) who described in Monostomum mutabile
as organs contributing to the formation of the egg four distinct glands, the
vitellaria, which he interpreted as the ovary as follows: — "Die Ovarien
bilden kurze blinde Schlauche, die unter einander anastomosiren und den
Darmkanal, nachdem er vom Oesophagus aus die Seitenrander des Leibes
erreicht hat, in seinem ganzen weiteren Verlaufe wie ein Netz umgeben.
Es ist dies eine eigenthiimliche Anordnung, die ich bis jetzt noch bei keinem
anderen, zu den Trematoden gehorigen Wurme angetroffen habe." The
three other parts having to do with the formation of the shell are de-
scribed by the fo]lowing characteristic statements: — "An der zweiten
Abtheilung der weiblichen Geschlechtstheile, die zur Bildung der Eier-
haute bestimmt zu sein scheint, lassen sich deutlich drei eigenthiimliche
Organe erkennen. a) Erstens fallt hier ein runder, weissgelber Korper

249] NORTH AMERICAN MONOSTOMES 31

in's Auge, der zur rechten Seite dicht neben und vor dem hinteren Hoden
liegt, und an Umfang etwas kleiner als dieser ist. b) Diesem runden
Korper hangt zweitens nach innen ein ovaler, noch kleinerer und ebenfalls
weissgelb gefarbter Korper an, der mit ersterem durch einen kurzen, an-
fangs weiteren, nachher engeren Kanal in Verbindung steht. c) Endlich
liegt drittens, theils unter diesen Organen, theils zwischen dem runden
Korper und dem hinteren Hoden eine durchsichtige, fast farbelose und
unregelmassig umgranzte, feinzellige Masse, in die der gemeinschaftliche
kurze Ovariengang einmiindet, und aus der der eierfiihrende Uterus her-
vortritt. Es ist mir bis jetzt nicht gelungen, eine Verbindung dieses un-
regelmassigen Organes mit dem grosseren runden Korper bestimmt nach-
zuweisen; doch bin ich iiberzeugt, dass eine solche wirklich existirt."
Van Beneden (1861) in his study of Monostomum mutahile after character-
izing the vitellaria (vitellogene Van Beneden, Ovarien von Siebold) and
the ovary (germigene Van Beneden, one of the shell forming organs of von
Siebold) says (p. 74), that besides the testes and the ovary only one other
organ has been observed by him in the posterior region i.e., the vitelline
duct which dilates to form the vitelline reservoir. In regard to the other
organs described by von Siebold (1835) he says "Sont-ce la les deux autres
organes que M. von Siebold signale et qui contribuent a la formation des
oeufs? Cela est probable!"

More recently Braun (1892) stated that he is able to find the recep-
taculum seminis only in Aploblema, Cephalogonimus and in the dis-
tomes, and that on the basis of his own research Laurer's canal is wanting
in Monostomum mutabile. One year later he states that in the monostomes
a Laurer's canal appears to be wanting. Cohn (1902) reported the absence
of both these organs in Monostomum oculobium. On the contrary Stossich
in the same year after a thorough study of the group confirmed the pre-
sence of these organs in several species belonging to at least three genera
of this family. Arnsdorff (1908) describes for Monostomum vicarium a
small receptaculum seminis. Kossack (1911) after a study of a large
number of specimens belonging to different genera of this family, viz:
Cyclocoelum, Typhlocoelum, Haematotrephus, Hyptiasmus, states con-
trary to the finding of Stossich that both receptaculum seminis and Laurer's
canal are wanting in this family. S. J. Johnston (1916) makes no mention
of either the receptaculum seminis or Laurer's canal in any one of the
three genera studied, Cyclocoelum, Haematotrephus and Hyptiasmus.

In a study of a considerable number of specimens belonging to several
species of the genus Cyclocoelum the writer has observed the presence of
the receptaculum seminis. In addition to the observations made on the
American material the writer has been given the opportunity through the
efforts of Professor Henry B. Ward and the courtesies extended him by the
Curators of the museums of Berlin, Gottingen, and Vienna, to study Cyclo-

32 ILLINOIS BIOLOGICAL MONOGRAPHS [250

caelum mutabile, Cyclocoelum problematicum, Cyclocoelum ovopunctatum,
Cyclocoelum brazilianum, Cyclocoelum tringae, and Haematotrephus similis.
In all of these species a small spherical receptaculum seminis has been
found, in position median and dorsal. to the ovary (Figs. 20-24, 26 and 27).

On the same dorsal level with the receptaculum semnis and posterior
to the latter and the ovary one finds the compact shell gland, in size ap-
proximately equal to the ovary. It is composed of unicellular glands closely
packed together, each of which empties its secretion into the ootype by
means of a short straight canal; these ducts form the inner portion of the
gland. The ootype in the central portion of this gland gives rise to the
uterus which in turn expands, immediately upon emerging, into a large
receptaculum semnalis uterinum. During the sexual activity of the worm
this pouch is filled with spermatozoa. In the genus Cyclocoelum, the uterus
forms relatively short closely packed loops, in general going out from the
middle line of the body. Relatively few stretches go directly across the
body. The uterus fills out the entire space between the crura, then by a
relatively straight stretch it spans the distance to the genital atrium in the
region of the pharynx. In lateral extent the uterus reaches in general to
the middle or outer wall of the intestine or rarely even out to the body wall,
as in Cyclocoelum halli. This species differs in this respect from other
species of the genus and according to the generic limits of Kossack does not
belong here if this condition is a diagnostic factor. Cyclocoelum halli,
however, conforms so closely to the genus in other respects and does not
conform to the genus Haematotrephus in that the uterine loops do not
bend around the genital organs so that the writer feels justified in placing
it in the genus Cyclocoelum. Evidently there is little justification for the
creation of a new genus based largely on the extent of the uterus and hence
the limits of the genus Cyclocoelum have been extended to include this
species.

The condition known as a situs inversus is a common feature of this
family. Looss (1899) asserted that in all forms in which the genital pore
is median there is a possibility that sexual amphitypy will occur and
suggested that in Monostomes where this is the case situs inversus will
probably be found. Cohn (1902) was the first to record this condition
in the group when he found in Spaniometra oculobia (Cohn) a situs inversus
of the genital glands in a ratio of 9:5. Kossack (1911) added to this by his
observations on Cyclocoelum problematicum. Stoss., Cyclocoelum ovopunc-
tatum Stoss., and Cyclocoelum vicarium (Arnsd.) in which he says that on
the average the right and left positions are equally frequently present.

The writer has found a similar condition to exist in all species of
Cyclocoelum represented in North America as well as being able to verify
the observations of Kossack on Cyclocoelum. problematicum and Cyclocoelum
ovopunctatum. Reference to the following table shows that the right and

251]

NORTH AMERICAN MONOSTOMES

3Z

left positions are on the whole equal in number and where significant
differences occur they are perhaps due to the small number of individuals
studied. In Cyclocoelum ohscuriim with sixty specimens in the lot the
difference is relatively small. The total of ninety-four specimens in all
show an approximately equal count for each arrangement.

Ante-
rior
testis

Cirrus
pouch

Number of uterine loops
between testis

Total uterine

loops on side of

anterior testis

Rt.

Lt.

Rt.

Lt.

0

1

2

3

4

5

6

7

8

9

25-
30

30-

35

35-
40

40-
45

45-
50

C. obscurum . . .

34

26

28

32

1

1

5

4

6

3

1

9

23

22

5

C. elongatum . .

4

4

.1

4

1

5

1

1

2

3

7

C. pseudomicro-
stomum. . .

3

1

2

2

1

1

1

1

1

2

1

C. macrorchis . .

8

14

7

15

1

6

3

6

1

3

2

1

4

12

5

Total numbers .

49

45

40

54

III ■ 1

The variation in the position of the testes from one another is equally
of little importance from the standpoint of specific diagnosis. However,
since Stossich used this as the means of separation of species in Cyclocoe-
lum and Haematotrephus the writer feels it worth while to give here the
result of observations on American material. Kossack raised objections
to the importance assigned this point by Stossich and showed that the
variation within a species was even greater than that between the genera
before mentioned.

The study of the American material has served to substantiate the
view of Kossack (1911) reached by study of the European material.
Reference to table shows the number of uterine loops to vary from none
where the testes lie contiguous to one another to nine while the entire
number of uterus loops, counted always on the side of the anterior testis,
is likewise variable and appears to bear no relation to the total number of
uterine loops between the testes of the individual since the specimen
which showed nine loops between the testes has in all only 38 loops while
in another individual of the same lot of Cyclocoelum obscurum, which had a
total of 40 loops only 3 could be found between the testes. Other examples
are 8 and 43; 3 and 44; 7 and 39 as compared to 1 and 39. These are a few
examples taken from Cyclocoelum obscurum. Other species show the same
to be true except in Cyclocoelum lialli and Cyclocoelum triangularum where
the relative positions of the genital glands appear to be constant. How-

34 ILLINOIS BIOLOGICAL MONOGRAPHS [252

ever, since the number of specimens here is small the writer is inclined
to attach relatively little importance to this feature.

The life history of this group is relatively unknown, although the ear-
liest record of a monostome larva is said to date from 1817. An early
account is found in Filippi 1859. This author obtained from Bythinia
tentaculata (L) [= Paludina impiira of Filippi] a larva he named Cerceria
lophocerca which he described (1859:5) as follows: "EUe est caracterisee
par un bulbe pharyngien assex fort, par la presence de deux yeux ou taches
pigmentaires semilunaires avec une petite lentille dans la concavite, et
par une queue munie d'une crete membraneuse longitudinale. Dans I'in-
terieur du corps on voit des rudiments d'organes sexuels sous la forme de
trois masses vesiculaires."

According to Luhe (1909) the determination of Filippi is doubtful
since the description and figures show a close resemblance to Cercaria
fuhopunctata Ercol. which is an undoubted distome larva. Cort (1915)
on the other hand recognized distinct features in this larva and stated
that it is "entirely different from all other monostomes known." As such
it stands alone and unidentified.

While the description of Filippi is meager and bereft of many diagnos-
tic characters a few outstanding features point to its alliance to this group.
This relationship is shown by the absence or at most only poorly devel-
oped oral sucker (Filippi, pi. I, fig. 3), by the presence of a strongly devel-
oped pharynx, and by the position of the three "rudimentary" sex organs.

Key to species of Cyclocoelum

1(6) Uterus restricted to the intercecal zone 2

2 (5) Testes unequal in size 3

3(4) Pharynx larger than oral sucker; ratio of posterior testis to

ovary 2:1 Cyclocoelum mutabile (Zed.)

4(3) Oral sucker and pharynx small, approximately equal in size;

ratio of anterior testis to ovary 4:3

Cyclocoelum cuneatum nov. spec.

5(2) Testes equal in size. Oral sucker larger than pharynx; ratio

of testes too vary 3:1 Cyclocoelum leidyi nov. spec.

6(1) Uterus not restricted to intercecal zone 8

7(26) Uterus folding around the crura both dorsally and ventrally . . . 9

8(21) Cental glands separated by uterine loops 10

9(16) Testes equal in size 11

10(13) Pharynx larger than sucker; vitellaria extending laterally

beyond medial wall of crura 12

11(12) Ratio of testes to ovary 5:2

Cyclocoelum pseudomicrostomum nov. spec.

12(11) Ratio of testes to ovary 4:3. . .Cyclocoelum microstomum (Crepl.)

253] NORTH AMERICAN MONOSTOMES 35

13(10) Sucker larger than the pharynx 15

14(15) Vitellaria strongly developed, extending to inner wall of

intestine; ratio of testes to ovary 2:1

Cyclocoelum macrorchis nov. spec.

15(14) Vitellaria weakly developed, rarely reaching middle of

crura; ratio of testes to ovary 2:1

Cyclocoelum vicarium (Arnsd.)

16(9) Testes unequal in size 18

17(20) Oral sucker twice as large as pharynx 19

18(19) Ratio of posterior testis to ovary 2:1

Cyclocoelum ohscurum (Leidy)

19(18) Ratio of posterior testis to overy 3:1

Cyclocoelum ovopunctatum Stoss.

20(17) Oral sucker not twice the size of pharynx. Ratio of poste-
rior testis to ovary 3:1 Cyclocoelum problematicum Stoss.

21(8) Genital glands contiguous, not separated by uterine loops. . .24
22(25) Testes unequal in size; oral sucker twice as large as pharynx 25

Cyclocoelum wilsoni nov. spec.

24(23) Ratio of posterior testis to ovary 2:1

Cyclocoelum tringae (Brandes)

25(22) Testes equal in size; oral sucker and pharynx approximately

equal. Ratio of testes to ovary 10:7

Cyclocoelum triangularum nov. spec.

26(7) Uterus passing dorsally over intestinal crura and vitellaria
to body wall; vitellaria moderately developed, rarely ex-
tending beyond middle of crura 30

27(28) Sucker \]/2 times pharynx; testes unequal; ratio posterior

testis to ovary 3:2 Cyclocoelum brazilianum Stoss.

28(29) Sucker and pharynx equal in size; testes equal; ratio of

testes to ovary 2:1 Cyclocoelum halli nov. spec.

Description of species

CYCLOCOELUM LEIDYI nov. spec.

[Figures 1, 2, ZZ]

Syn: Monostomum mutabile Leidy 1885, nee Zeder 1800

This collection contains five specimens described by Leidy (1885) as
follows: "From the thoracic cavity of a Gray Snipe, Gallinago wilsoni,
Dr. Warren (of Westchester) obtained five Flukes, 18mm long, by 4mm
broad. These appear to be Monostomum mutabile."

These specimens are readily recognized as belonging to the genus
Cyclocoelum Brandes (1892) but are distinctly different from Cyclocoelum
mutabile and represent a new species. The following description shows the
characteristic differences between the two species.

36 ILLINOIS BIOLOGICAL MONOGRAPHS [254

These flukes are 16 to 18 mm long by 4 to 4 . 5 mm broad. The margins
of the body are practically parallel in the posterior two-thirds of the body.
The anterior one-third tapers to a weakly rounded point. The subterminal
mouth is surrounded by a weakly developed sucking musculature which
is approximately three times the size of the small pharynx. The small
ovoid pharynx measures 231 /i broad by 281 ^ long. The slender esophagut
which is curved in the form of an S measures 331// in length. It opens inso
the middle of the dorsal side of the intestinal bifurcation. The voluminous
intestinal crura run parallel to the margins of the body throughout their
entire course. The vitellaria are moderately developed and occupy the
region lateral to the crura. They extend laterally to the inner wall of the
crura and fold both dorsally and ventrally around them. They extend an-
teriorly not quite to the most anterior portion of the intestinal bifurcation
and are separated at the posterior end only by the excretory bladder. The
uterus fills the entire space between the intestinal crura. It lies in the
mid dorso-ventral region, is profusely coiled and does not extend out
beyond the inner wall of the intestine. Genital glands are confined to
the posterior intestinal arch. The posterior testis is situated in the middle
line of the body and lies directly in the arch formed by the intestine.
The anterior testis is a little removed and is contiguous to the crura. They
are equal in size and measure 877 to 910^t in diameter. The much smaller
ovary lies on the side of the body opposite to the anterior testis and in a
transverse plane between the two testes. It is spherical and measures from
380 to 390/z in diameter. Dorsally and on the inner posterior side of the
ovary is a small spherical receptaculum seminis 82 to 99/i in diameter,
which joins by a short duct the oviduct shortly after its emergence from
the ovary. Laurer's canal is not present. As the oviduct passes poste-
riorly into the adjacent shell gland aggregate it is joined by the vitelline
duct. At this point the oviduct enlarges to form the ootype. Just after
the uterus emerges from the compact shell gland a second enlargement is
seen, the receptaculum seminalis uterinum. It extends posteriorly to the
crura where it doubles on itself and pursues its coiled winding course to the
genital pore which is situated ventral to the middle region of the pharynx.
The cirrus pouch is 182)u broad by 331^ long and reaches to the middle
of the intestinal crura at their bifurcation. The genital pores open sep-
arately into a small genital atrium. Eggs thick shelled ovals, 66 by \\1 n
when fully mature. They contain in the anterior portion of the uterus well
developed miracidia as evidenced by the dark eye spots.

Habitat: Thoracic cavity Host: Gallinago wilsoni

Locality: Westchester, Pa. Date: 1885

Collector: Dr. H. W. Warren No. 106 Leidy Collection.

A comparison of this material with the data given by Stossich (1902:
13) and by Kossack (1911:510) for Cyclocoelum mutabile (Zeder) as well as

255] NORTH AMERICAN MONOSTOMES 37

comparison with specimens of Cyclocoelum mutabile obtained from the
Gottingen museum demonstrates clearly that this is a distinct species.
It is similar to Cyclocoelum mutabile in the size and form of the body, in
the lateral extent of the uterine loops, the extent and development of the
vitellaria, the size of the ovary, and the size and shape of the eggs.

It differs from this species in having a much smaller pharynx, a much
larger sucker, a longer esophagus, larger testes, and a relatively heavier
and more irregularly folded uterus. This species is similar to Cyclocoelum
prohlematicum Stossich in the size of the testes and the extent of the
vitellaria but differs from that species in having a smaller pharynx and a
broader, thinner and less muscular body in proportion to its length. For
comparison with this species a figure (Fig. 3) of Cyclocoelum mutabile
(Zed.) is placed beside that of Cyclocoelum leidyi.

CYCLOCOELUM PSEUDOMICROSTOMUM nov. spec.
[Figures 4, 27, 30, 43]

Large monostomes 13 to 14.5 mm in length by 4 to 4.5 mm in greatest
width which is found at the beginning of the posterior body third. From
this point forward the side walls taper gradually to the end of the anterior
body third, at which point they bend inwardly more sharply to form a small
obtusely rounded end. The posterior end is bluntly rounded. The mouth
is subterminal, surrounded by an external banding musculature which
measures 662 to 745 ju in diameter. This is followed by a large heavy
slightly elongate pharynx 778 to 910ju in length by 745 to 844/i in width.
The genital pore lies median and ventral to the forward end of the pharynx.
From this point the cirrus pouch stretches posteriad almost to the posterior
wall of the intestinal bifurcation. The vitellaria extend from the posterior
end of the cirrus pouch to the excretory bladder in the posterior end of the
body. It is even more strongly developed than that of Cyclocoelum
microstomum and in its lateral extent passes the inner wall of the crura and
over the lateral folds of the uterine loops which in this species rarely pass
over the inner wall of the crura. The testes as in other species of this genus
lie in the posterior region of the body and within the intestinal crura. The
posterior testis does not fill the entire intestinal arch, is antero-posteriorly
flattened and measures 827 to 910m in width by 993 to 1192)u in length,
while the anterior testis which is separated from it by uterine loops meas-
ures 745 to 993 ;u in width by 1076 to 1324/x in length. The ovary lies on
a level with the anterior margin of the posterior testis and adjacent to the
cecum opposite to the anterior testis, is much smaller and spherical,
measuring 413 to 496;u in diameter. Dorsal to the ovary is the spherical
receptaculum seminis 148 to 165 /x in diameter. The shell gland is similar
in size to the ovary and occupies a position dorsal and posterior to that
structure. As was stated above the uterus does not usually pass over the

38 ILLINOIS BIOLOGICAL MONOGRAPHS [256

inner wall of the intestine and fills out entirely the space between the
crura. The eggs are thick shelled ovals, 51 to 66/x in width by 102 /x in
length.

Habitat: Lung Host: "Wild duck"

Locality: Omaha, Nebr. Date: 1903

Collector: C. E. Stringer No. 1041 Ward Collection

This species is also found in the Leidy Collection vial no. 186 which has
been dried out and is in a poor state of preservation. John C. Johnson
collected this species from Fulica americana taken at Golden Gate Park,
San Francisco, Cal. in January 1919.

Cydocoelum pscudomicrostomiim finds its nearest relative in Cyclocoe-
lum microstomum (Crepl.). The two species are of nearly equal size.
The pharynx of the former is distinctly larger, the testes are not of equal
size as in C. microstomum and are slightly broader than long probably due
to pressure from the closely packed uterus. The ovary of Cydocoelum
pseudomicrostomum is noticeably smaller than that of Cydocoelum micro-
stomum, while the lateral extent of the uterus in the former species is more
restricted. With respect to the development of the vitelline glands
Cydocoelum pseudomicrostomum presents the heaviest development found
in any known species of this genus.

CYCLOCOELUM HALLI nov. spec.
[Figures 5, 11, 20, 36-42]

Large monostomes varying in length from 11 to 14 mm in width by 3
to 4 mm in greatest width which is found slightly posterior to the middle
of the body. From this point the body tapers anteriorly to almost a
point and posteriorly only a little, forming an obtusely rounded end.
The body is muscular, dorsally convex, and ventrally flat or slightly con-
cave. The body wall is entirely covered with numerous small pits ob-
served by Zeder (1803) in Monostoma mutahile. The subterminal mouth
leads by a funnel-shaped tube to the pharynx. This tube or mouth
proper is surrounded by a concentration of musculature which on the
outer margin is formed into circular bands, the outer covering of the sucker
(Figs. 36 to 42). The sucker is spherical in form and measures 387/1 in
diameter. It is separated from the smaller (263 /x) but more heavily muscu-
lar, spherical pharynx, by the nerve commissure. The esophagus is 483 /x
in length and extends from the posterior portion of the pharynx to the
dorsal side of the intestinal bifurcation. The latter a simple tubular struc-
ture lies along the margins of the body for its entire length and anasto-
moses at the posterior end. As previously described the excretory system
is composed of a system of tubules ramifying the entire body in this
species, as in Cydocoelum elongatum. These tubules anastomose and
empty into the excretory bladder. In this species a single thin walled sac

257] NORTH AMERICAN MONOSTOMES 39

which opens to the exterior by a small dorsal pore. The genital glands are
situated in the posterior fifth of the body where they are closely packed in
the posterior arch of the intestine, the posterior testis almost entirely
filling this space. It is slightly flattened antero-posteriorly and measures
in its greatest dimension 1052 /z and in an axis at right angles to this, 894^.
The anterior testis situated a short distance cephalad to the posterior,
is spherical in shape and a little smaller, having a diameter of 894//. The
two testes are separated by a particularly long loop of the uterus which
extends to the intestinal arch, and in some instances even beyond, and
usually folds back part way forming a double loop. The vasa efferentia
given off from the anterior margins of the two testes unite cephalad and
mesad to the anterior testis to form the vas deferens which takes a fairly
straight course to the cirrus pouch with which it unites.

The cirrus pouch is a rather large oblong sac extending from the
pharynx to a short distance beyond the anterior wall of the intestinal
bifurcation. It opens into a small genital atrium which in turn opens to
the exterior, ventral to the pharynx.

The ovarian complex is situated opposite to and on a level with the
anterior testis. The ovary is very much smaller than the testes, measuring
434/1 in diameter. Dorsal to this is the receptaculum seminis, the duct
from which joins the oviduct before it enters the shell gland. The shell
gland, a compact spherical organ, is situated dorsal and posterior to the
ovary. It has a diameter of 388 /i. The vitellaria lie between the lateral
body wall and the external wall of the digestive crura over which they
seldom pass. They extend from the posterior pharyngeal region to the
extreme posterior end where they are separated by the excretory bladder.
The vitelline glands are made up of small follicular grape-like clusters
arranged along a main stem, which in the region of the anterior testis
gives rise to the vitelline ducts. These pass mesad and unite a short
distance posterior to the shell gland to form the common duct which
passes straight to the shell gland. It enters this at the posterior side
and passes through the outer portion of this organ to its union with the
oviduct just after the entrance of the latter into the shell gland. The
oviduct then enlarges to form the ootype. On emergence from the shell
gland the uterus enlarges to form a large pouch, the receptaculum
seminalis uterinum. From this the uterus makes a few short loops and then
the long loop, previously mentioned, which separates the testes. From
this point forward it lies in more or less regular transverse folds which
extend out to the vitellaria, tho in some instances, particularly in the
posterior three-fifths of the body, these loops extend to the body wall.
In the anterior fifth of the body the uterine loops are not so long and
here fill out entirely the space between the intestinal crura. From the

40 ILLINOIS BIOLOGICAL MONOGRAPHS [258

bifurcation of the intestine the uterus reaches in a relatively straight
stretch to the genital atrium.

The eggs are large, thick shelled ovals, measuring 161 /z in length by 99 /u
in breadth. The double, dark eyespots give evidence of the developing
miracidium within the eggs before they have passed in their course anterior
to the middle region of the body.

Habitat: Abdominal air-sacs. Locality: host taken in Raleigh

Host: Totanus melanoleucus (?) N. C.

Date: April 7, 1894 Collector: W. C. Hall

No. 21.90 Ward collection

Habitat: Liver? or Lung? Locality: Creston, Iowa

Host: Totanus soUtarius Collector: W. C. Hall

Date: Aug. 30, 1895 No. 21 . 763 Ward collection

Filed among the records of Hall is a statement that he collected 5
specimens from the air sacs of Totalnus flavipes Sept. 4, 1895, which he
believed to be this species. The material from this host has not been found.
This species is most closely related to Cyclocoelum brazilianum Stossich
and resembles that form in the lateral extent of the uterus, the relative
position of the genital glands, and in the size of the ovary. It differs,
however, in the size of the testes, the more weakly developed vitellaria,
and relative size of the oral sucker and pharynx, these being of equal size
in C. halli while in C. brazilianum the sucker is distinctly larger than the
pharynx.

CYCLOCOELUM WILSONI nov. spec.
[Figure 6]

Medium sized monostomes 12 mm long by 3 mm wide in maximum
which is found at the beginning of the posterior one-fifth of the body.
Posterior and bluntly rounded. Anterior to the point of greatest width
the body tapers gradually to a blunt but relatively small point. The
mouth sucker measures 374^i in diameter and is one-third larger than
the oval pharynx which measures 298/x in length by 269/1 in width. The
esophagus is relatively long and gives rise by bifurcation to the simple
intestinal crura, which as in other species run parallel to the body wall
and anastomose in the posterior end of the body. The genital aperture
is ventral to the middle of the pharynx. The cirrus pouch extends from
this point to the middle of the intestinal bifurcation. The follicular yolk
glands extend from the anterior wall of the intestinal bifurcation almost
to the excretory bladder at the posterior end. In lateral expanse they pass
over the external wall of the intestinal crura to the middle of that organ
where they meet the furthest expanse of the uterus. The uterus in this
species lies entirely anterior to the anterior testis and fills out the space
between the crura, folding both dorsally and ventrally over the walls of

259] NORTH AMERICAN MONOSTOMES 41

these to the middle region of the same. The genital glands lie contiguous
to one another in the posterior arch of the intestine. The posterior testis
lies a little posterior to and a little more nearly in the middle of the arch
than the anterior one which is smaller and contiguous to the former,
filling out the opposite portion of the intestinal arch. The testes are not
separated by uterine loops as is generally true in this genus. The anterior
testis is spherical, 910/x in diameter while the posterior testis is slightly
elongated and measures 992) jj. in length by 910^ in width. The ovarian
complex lies anterior to and adjacent to the posterior testis. The ovary and
shell gland are spherical, equal in size, and measure 413 to 447 ^ in diame-
ter. The small spherical receptaculum seminis lies partially embedded in
the shell gland and measures 150/i in diameter. The eggs are thick shelled
ovals and measure 150;u in length by 76/x in width.

Habitat: Intestine Host: Gallinago wilsoni

Locality: Creston, Iowa Date: August 4, 1894

Collector: W. C. Hall No. 21.89 Ward Collection

The direct relationship of this species is not so readily apparent. As
to position of the genital glands it holds a place close to Cyclocoelum
tringae (Brandes) and Cyclocoelum triangularum nov. spec, and tho it is
much larger presents in general the same characteristic features namely:
genital glands contiguous, uterine loops directed backward, vitellaria
moderately deveoped, oral sucker larger than pharynx. However, in the
size of the body and the relative size of the pharynx and sucker as well as
the relative size of the genital glands Cyclocoelum wilsoni is clearly distinct
and must be recognized as a proper species.

CYCLOCOELUM CUNEATUM nov. spec.

[Figures 7, 24]

Medium sized worms 10.5 to 12 mm in length by 2.5 to 3.5 mm in
greatest width which is found at the beginning of the posterior body
fourth. From the point of maximum width the margins of the body run
approximately parallel to the level of the anterior testis at which point
they narrow abruptly to form the obtusely rounded posterior end. From
the point of greatest width the margins of the body converge cephalad in
almost straight lines to a very narrow and pointed anterior end, which
gives the impression of a well formed wedge. At the pointed anterior end
is found a very small weakly developed sucker, 198 to 215/i in diameter.
This is followed by an oblong pharynx of approximately the same width
as the sucker. It measures 150 to 198^t in width by 215 to 231 /x in length.
The esophagus is three and one-half to four times the length of the pharynx.
The intestinal crura are simple. The vitelline glands extend from the
middle region of the bifurcation of the intestine to the excretory bladder

42 ILLINOIS BIOLOGICAL MONOGRAPHS [260

in the posterior end. In lateral extent they rarely traverse the outer
wall of the crura. The genital opening is located anterior to the pharynx.
The cirrus pouch extends to the anterior wall of the intestinal bifurcation.
The uterus fills out the entire space between the crura and its loops occa-
sionally span the outer wall of the same organ. The genital glands are
relatively small and occupy the positions so common to the species of this
genus. The posterior testis, not filling out the intestinal arch, is antero-
posteriorly flattened and measures 413ju in width by 496 to 612)U in length.
The anterior testis is separated from the posterior by several uterine
loops and is smaller and spherical, measuring 331 to 413^1 in diameter.
The spherical ovary is situated in a transverse plane anterior to and about
equally distant from the two testes. It is one-fourth smaller than the
testes measuring 215 to 331/x in diameter. The shell gland lies median
and posterior to the ovary, is spherical or only slightly ovoid and equal
in size to the latter organ. The receptaculum seminis although indis-
tinguishable in toto mounts is clearly seen in sections. It is anterior and
dorsal to the ovary. Wax reconstructions show it to be spherical and
approximately one-half the size of the ovary, measuring ISO^t in diameter.
The ovarian complex is separated from the testes by uterine loops. The
eggs are thick shelled ovals, 66 ^ wide by 115;u to 122 ju long.

Habitat: Abdominal cavity Host: Gallinago delicata (Oid.)

Locality: ? Date: ?

Collector: ? No. 08. 172 Ward collection.

The relationships of this species is not so evident as it partakes of the
characteristics of a number of species. With respect to the lateral extent
of the uterine loops it is more nearly like Cydocoelum mutahile and Cyclo-
coelum leidyi while in development of the vitellaria it simulates Cydocoelum
halli. The pharynx and oral sucker are noticeably smaller than in any
known species of this genus and this is a feature of this species as is also
the small size of the genital glands and the proportionate size of the testes
to the ovary, a ratio of 4:3.

CYCLOCOELUM OBSCURUM (Leidy)

[Figures 8, 21]
Syn: Monostomum ohscurum Leidy 1887
This species was described by Leidy as follows: "Elongated, elliptical,
flattened, obtusely angular in front, obtusely rounded behind, oral and
genital and other aperatures scarcely distinguishable. Length 4 to 8 Hues;
width 1 line."

"Numerous specimens in the stomach of a Jew-fish, Megalops thris-
soides."

The host name Megalops thrissoides used for the Jew-fish by Leidy
1887 is evidently a lapsus calami. However this is corrected by Stiles and
Hassall (1894) to Stereolepis sp?

261] NORTH AMERICAN MONOSTOMES 43

Brandes (1892) enumerates this among other species which he has
not had opportunity to study and he justly says that it is inadequately
described. Monticelli (1892) and Braun (1893) in spite of the meager
description retain it as a valid species.

The original material of this species is found in the Army Medical
Museum under Number 1035 Comparative Anatomy Series with the
description "Flukes Monostotnum obscurum from the stomach of a Jew-
fish (Stereolepis)." From this I have written the following description.

Monostomes of medium size measuring from 6 to 13 mm in length and
from 1.5 to 3 mm in maximum width which is found a little posterior to
the middle of the body. The margins of the body are almost parallel for
the greater part of their length, tapering gently to the more pointed
anterior end and abruptly to form the obtusely rounded posterior end. As
in most species of this genus the body is convex dorsally and flat or slightly
concave ventrally. The subterminal mouth is surrounded by a weakly
developed sucking musculature which measures 115ju in diameter. This
leads to the spherical or slightly elongated pharynx, measuring 115 to
264 /x in width by 115 to 298 ^t in length. Following this the slender esopha-
gus, 500 to 750/i in length, leads to the intestinal bifurcation. The crura
are quite variable in size as well as in the character of the median wall.
In some cases they show a tendency to the formation of internal ceca; these
appear to be due to the pressure from the closely packed uterus which fills
the space between the crura. The excretory system in this species has not
been made out except for the single termnal excretory vesicle situated as
in the other species of this genus between the posterior arch of the intes-
tinal crura and the posterior body wall. It opens to the exterior by a single
dorsal pore. The genital organs lie within the intestinal crura in the
posterior end of the body. The posterior testis, filling the posterior arch
of the intestine, is flattened anteriorly by the closely packed uterine
loops and is slightly larger than the anterior being 300 to 877/1 in width
by 480 to lOOOju in length. The anterior testis is usually more nearly
spherical and measures 380 to 82 7 /i in width by 462 to 827ju in length.
It lies obliquely anterior to the posterior and adjacent to the crura. The
vasa efferentia are short and unite a short distance anterior to the anterior
testis to form the vas deferens which for the most part passes dorsal to the
uterus to the cirrus pouch. This organ is of medium size 248 to 579 fx in
length by 115 to 199 /z in width. In general its posterior limit lies on a
level with the middle of the intestimal bifurcation. The ovarian complex
lies between the testes and on the side opposite to the anterior testis.
It is composed of a spherical ovary, 275 to 463;u in diameter, a spherical
receptaculum seminis, 132 to 148/x in diameter, and a shell gland in size
and form similar to the ovary. The position of these is clearly shown in
figure 21. Beginning in the shell gland the uterus immediately upon

44 ILLINOIS BIOLOGICAL MONOGRAPHS [262

emergence enlarges to form the large receptaculum seminalis uterinum.
It fills the intercecal space with more or less regular loops which in general
go out from the middle of the body. These loops fold around the inner
surface of the crura and usually do not pass beyond the outer wall of the
latter organ. The vitellaria for the most lie outside the crura and extend
from the pharynx to the excretory bladder. Laterally they reach to the
middle of the crura and in exceptional instances to the inner wall of that
organ.

Habitat: Stomach Host: Stereolepis sp?

Locality: ? Date: ?

Collector: ? Cat. No. 1035 Comparative

Anatomy series.

Habitat: ? Host: Symphaemia semipalmata

Locality: Lincoln, Nebraska Date: ?

Collector: ? Cat. No. 08. 179 Ward collection

Habitat: ? Host: Unknown

Locality: Spokane, Wash. Date: ?

Collector: W. E. Allen Cat. No. 08 . 183 Ward collection

Cyclocoelum obscurum is most closely related to Cyclocoelum ovopuncta-
tum Stossich and differs from that species in the more slender form, the
more heavily developed vitellaria and the relative size of the testes to the
ovary which in Cyclocoelum obscurum are twice as large as the ovary while
in Cyclocoelum ovopunctatum they are three times as large.

CYCLOCOELUM MACRORCHIS nov. spec.
[Figure 9]

This species varies in length from 7 to 15 mm and in maximum width,
which is found just posterior to the middle of the body, from 2 to 4 mm.
From this point the body tapers towards both ends, the posterior being
obtusely rounded while the anterior is considerably more attenuated.
It forms a moderately rounded point. The margins of the body lie nearly
parallel in the middle region of the body. The subterminal mouth is sur-
rounded by a weakly developed musculature, the oral sucker, which is
only a little larger than the well developed pharynx just posterior to it,
and measures 255 ju in diameter. The pharynx is oval in shape being about
one-fifth longer than wide and measures on the average 271 m long and
238/x wide. The esophagus in this species is on the whole well extended
and ranges from 331 /i in the state of least extension to 662 /z in that of
greatest extension exhibited in preserved material. At its posterior end
the esophagus turns ventrad and bifurcates forming the voluminous
crura present in this species (Fig. 9). These as in other species of this
genus lie parallel to the margins of the body and anastomose at the poster-

263] NORTH AMERICAN MONOSTOMES 45

ior end. The excretory system as far as can be made out in preserved
material conforms in this species to the description given previously in
having a single thin walled vesicle in the extreme posterior end of the body
into which the anastomosing tubules empty. It opens to the exterior
slightly dorsal to the posterior end. The genital glands in general occupy
the intercecal zone and fill entirely that space. The posterior testis occupies
the intestinal arch, is usually spherical in form and measures 783 to 984^t
in diameter. The anterior testis is usually removed from the posterior by
several uterus loops. It is spherical and approximately the same size as
the posterior measuring 730 to 860;u in diameter. As in other species of
this genus the vasa efferentia unite cephalad and mesad to the anterior
testis. From this point the vas deferens takes its course among the uterine
folds to the posterior end of the cirrus pouch which is situated at the
middle of the intestinal bifurcation. The cirrus pouch extends from the
genital atrium caudad to the middle of the intestinal bifurcation. From
this point the club-shaped cirrus pouch extends cephalad to the genital
atrium and lies ventral to the anterior end of the pharynx. The ovarian
complex is situated between the testes and adjacent to the crura opposite
the anterior testis. The ovary is spherical and measures 413 to 463 ju in
diameter. Dorsal and posterior to the ovary is the spherical receptaculum
seminis, 165 ju in diameter. The duct of this unites with the oviduct
before it enters the shell gland. The shell gland is approximately the same
size as the ovary and is situated posterior and dorsal to that organ (Fig.
9). The well developed follicular yolk glands occupy the region of the
body lateral to the intestinal crura and extend from the anterior-most
part of the intestinal bifurcation to the posterior end where they are
separated only by the small excretory bladder. The follicles are arranged
in clusters on secondary branches from the main stem and in this manner
extend laterad around the crura both dorsally and ventrally, in many
cases reaching out as far as the inner wall of the crura. The vitelline ducts
are given off in the region of the shell gland and pass mesad to a point
just dorsal to the shell gland where they unite to form the vitelline reser-
voir. From this the common vitelline duct passes dorsal to the shell gland
and joins the oviduct just before its entrance into that organ. Immediately
upon entering the shell gland the oviduct enlarges to form the ootype.
Upon emergence from this the uterus enlarges to form the receptaculum
seminalis uterinum. From this point the much folded uterus fills out the
entire space between the crura and passes over the bifurcation in a relative-
ly straight stretch to the genital atrium. The numerous eggs which fill the
uterus are thick shelled ovals, measuring from 122 to 153/* in length by 56
to 66 n in width. In general the eggs are smaller in the beginning of the
uterus than they are near the genital orifice. The eggs in the anterior
region show well developed miracidia with double dark eye spots.

46 ILLINOIS BIOLOGICAL MONOGRAPHS [264

Habitat: In lung and along the Host: Straight-billed Curlew

back in the abdomen.
Collector: W. E. Allen No. 08.180 Ward collection

Cyclocoelum macrorchis differs from Cyclocoelum mutahile in that the
uterus in the former is not restricted to the intercecal space and that the
oral sucker is larger than the pharynx, where as in Cyclocoelum mutabile
the vitellaria are more heavily developed and the genital glands are smaller.
Cyclocoelum macrorchis, however, has a much more muscular and much
thicker and heavier body.

CYCLOCOELUM TRIANGULARUM nov. spec.

[Figure 10]

Medium sized worms 8 mm long by 2 . 5 mm wide in maximum. Body
lanceolate in form. Oral sucker weak, 260ju in diameter, only a little
larger than the pharynx which is longer than broad, measuring 248 ^u in
length by 215 ;u in breadth. The length of the esophagus is approximately
one-tenth of the entire body length. Intestinal crura simple. Genital
pore ventral to the posterior end of the pharynx. Cirrus pouch extending
to the middle of the intestinal bifurcation. The vitellaria extends from the
posterior end of the cirrus pouch almost to the excretory bladder at the
posterior end of the worm. In lateral extent they pass over the wall
of the crura to the middle of that organ. The genital glands lie in the
posterior arch of the intestine and are not separated by uterine loops.
The two testes lie on the same level, one in either side of the arch, are
spherical and equal in size. They measure 41 3 ^t in diameter. The ovary
lies anterior to the testes and in the middle line of the body. It is a little
more than one-half the size of the testes, is spherical in form and measures
248 M in diameter. The shell gland lies between the ovary and testes also
in the middle line of the body and measures 264 ju in width by 314 jj. in
length. The receptaculum seminis is situated anterior to the shell gland
and dorsal to the ovary. Like the ovary the receptaculum seminis is
spherical in form and measures 132^t in diameter. The eggs are thick
shelled ovals 132 ^Li long by 75 /z wide.

Habitat: Abdominal air sacs Host: Tringa maculata

Locality: Creston, Iowa Date: September 4, 1895

Collector: W. C. Hall No. 21.88 Ward collection

As was stated in another section of this paper Cyclocoelum triangularum
shows a striking similarity to Cyclocoelum wilsoni and is distinguished from
that species by the generally smaller body, the more nearly equal sucker
and pharynx, the testes of equal size and the relative size of the testes and
ovary which in Cyclocoelum triangularum have a ratio of 10:7. It is more
nearly equal in size with Cyclocoelum tringae but is distinguished from

[265 NORTH AMERICAN MONOSTOMES 47

this species by the larger pharynx, smaller sucker, the testes being equal
in size and also smaller in proportion to the size of the ovary.

CYCLOCOELUM VICARIUM (Arnsdorff 1908) Kossack 1911
Syn: Monostomwn vicarium Arnsdorff

The writer has not had opportunity to study specimens of this species
and hence must rely upon the descriptions of Arnsdorff (1908) and Kossack
(1911). Since the description of Kossack made after having studied the
original material is at variance with the original description only in minor
detail the writer has based the following description on the work of Arns-
dorflf.

Monostomes varying between 10.5 and 14.4 mm in length and 3 to
3.1 mm in maximum breadth. The body is opaque, flattened; ventral
surface flat, dorsal slightly swollen. The sidelines of the body diverge
from the small pointed anterior end to the height of the testes. From here
they form the bluntly rounded posterior end. The dorsal surface of the
body is quite wrinkled. These folds appear in optical section to form pap-
pillae. The mouth opening is terminal. The strongly muscular pharynx is
an elongate oval with a long diameter of 460/i and a breadth of 270/t.
The thickness of its wall is 130^1. The esophagus, 640 ^i long, leads to the
intestinal crura which run parallel to the side walls of the body and anas-
tomose in the posterior end. The genital organs lie in the broad hinder
end. The posterior testis is flattened antero-posteriorly and measures
1190 /i in length by 732 /z in breadth. It lies in the middle line of the body
with its forward margin reaching to the vitelline duct. The anterior testis
is spherical in form with a diameter of 835 to 878 /z. It is removed anteriorly
from the posterior end and lies adjacent to the intestinal crura. The ovary
lies adjacent to the crural wall, opposite to the anterior testis and between
the positions of the testes. It is spherical and has a diameter of 402 /z.
Between it and the posterior testis is found the relatively small recep-
taculum seminis.

The vitellaria are composed of numerous follicles which lie parallel to
the body wall and between that and the intestinal crura; they extend from
the region of the intestinal bifurcation to the posterior end where they are
separated by a short interval. The numerous transverse uterus loops
fill out the space between the crura and in the posterior half of the body
overlap them. The genital pore is situated just posterior to the pharynx.
The club-shaped cirrus pouch is small and does not reach the intestinal
bifurcation. The eggs are numerous, elliptical in form and measure 102 ju in
length and 68 /i in breadth. The ripe eggs hold a well developed embryo
as can be recognized by the black eye spots in the miracidium.

48 ILLINOIS BIOLOGICAL MONOGRAPHS [266

The comparison of Arnsdorff with the figures and description of Stos-
sich for closely related species show a striking similarity to Cyclocoelum
problematicum. Cyclocoelum vicarium differs from this species, however,
in the relative size of the worms, the extent of the cirrus pouch and the
size of the eggs.

Habitat: Intestine Host: Arquatella maritima mari-

tinia
Locality: North East Labrador Date: September 14, 1906

Collector: Hantzsch Konigsberg Museum

NOTOCOTYLIDAE Luhe 1909

Of the monostome families which have up to this time been widely
studied no other family is of greater interest than the Notocotylidae. It
is in this family that the earliest records of the monostomes are found in
Catatropis verrucosa (Frolich 1789) collected from the rectum of A7ias
domestica. These worms were classed by Frolich and Gmelin as Fasciola.
Ten years later Zeder (1800) removed them to the genus Monostoma.
They were later separated from the remaining monostomes by Diesing
(1839) and placed in a new genus Notocotylus which genus remains as the
type of the family. Although Diesing included this earliest known form in
his genus Notocotylus, it has been found in more recent time by Odhner
(1905) to be distinct from the Diesing type species, Notocotylus triserialis,
and was removed to the new genus Catatropis which place it holds at pre-
sent as type of that genus. A further study of Notocotylus triserialis Diesing
by Kossack (1911) revealed its identity with Monoslomuni attenuatum
Rud. 1809. Thus the species name becomes a synonym to Notocotylus
attenuatus.

In regard to the early records of this family in America there still
remains a question. Some authors notably Barker and Laughlin would
place Monostomum affi-ne Leidy 1858 as the earliest American record while
others doubt this determination and still others reserve opinion on the
matter. Barker (1916) questions the determination of Leidy and expresses
the opinion that Monostomum affine Leidy belongs to the genus Notocoty-
lus. While the description of Leidy (1858:110-112) is insufficient for an
accurate determination of the systematic position of this species certain
facts given in his description are distinct and seem sufficient to show that
this worm is not Notocotylid in character. The length of Monostomum
affine as given by Leidy is 6}/^ lines or 13.5 mm which is two and one-half
to three times longer than any known species of this family. Likewise on
the same basis the Leidy species is at least three times wider than the
largest known Notocotylid. However, more important, intrinsic charac-
ters are the presence of a pharynx, an echinate penis and eggs prolonged at
one pole only. In addition these worms were taken from the gall-bladder

267J NORTH AMERICAN MONOSTOMES 49

and bile ducts of the muskrat while the Notocotylidae are normally-
inhabitants of the intestine and rectum. It is not impossible that a Noto-
cotylid species may have ascended the gall-duct of this host and become
modified and adjusted to the new conditions. Yet on the basis of the facts
noted above the writer cannot agree with the opinion of Barker regarding
the Notocotylid character of these worms.

More recently Hassall (Stiles & Hassall 1894) collected from Arvicola
riparius and Fiber zibethiciis taken in Maryland in 1892 specimens deter-
mined as Monostomum sp. and one year later from Aix sponsa and Dafila
acuta worms also determined as Monostomum sp. On observation and
study by the writer the specimens taken from Aix sponsa, Dafila acuta
and Fiber zibethicus have been found to be distinctly different from forms
previously described and on the basis given in a later section must be
recognized as a new species. On account of the similarity to the immature
stages of Cercaria urbanensis Cort it is believed to be the adult form of this
species. The material taken from Arvicola riparius has been found to
agree with Notocotylus quinqueserialis (Barker and Laughlin 1911). Other
American records are those of Barker (1915, 1916) in which he records
Catatropis filamentis from Fiber zibethicus taken in Nebraska and Nudoco-
tyle novicia from the same host taken in the same region.

Additional records presented in this paper are Notocotylus urbanensis
(Cort 1914) from the black and domestic swan taken at Golden Gate
Park, San Francisco, Cahfornia by John C. Johnson in February 1919 and
Paramonostomum echinum nov. spec, from the intestine of Fiber zibethicus
taken at Wray, Colorado by C. H. Gable, October 1916.

DIAGNOSIS OF FAMILY

Small monostomes tapering at both ends, posterior end broadly
rounded, anterior slightly more attenuated. Generally with rows of papil-
lae formed of unicellular dermal glands. Esophagus short, without
pharynx; intestinal ceca with short diverticula, extending entire length of
body. Genital pore median, except in Nudocotyle where it is distinctly
lateral, usually near oral sucker. Cirrus sac elongate. Testes symmetrical,
extracecal, near posterior end. Ovary between testes. Vitellaria lateral,
anterior to testes. Uterine coils between cirrus sac and genital glands,
transverse, regular, usually not extending outside intestinal crura. Eggs
small with long p olar filament on each end.

KEY TO SUBFAMILIES AND GENERA

1(5) Genital pore anterior median. . .Sub-family Notocotylinae . . 2

2(5) With ventral glands 3

3(4) Ventral glands protrusible Notocotylus Diesing 1839

4(3) Ventral glands not protrusible Catatropis Odhner 1905

50 ILLINOIS BIOLOGICAL MONOGRAPHS [268

5(2) Ventral glands wanting Paramonostomum Liihe 1909

6(1) Genital pore marginal posterior without ventral glands ....

Sub-family Nudocotylinae . . 7

Uterus in anterior body half Nudocotyle Barker 1916

The Notocotylidae are up to the present represented in North America
by two species of Notocotylus and one species of each of the other genera in
this family.

The Notocotylidae were subdivided by Kossack (1911) into two sub-
families; Notocotylinae including Notocotylus Diesing (1839) Catatropis
Odhner (1905) and Paramonostomum Liihe (1909), and Ogmogasterinae
represented by a single species Ogmogaster pUcatus (Creplin 1829) Jager-
skiold 1891. A third subfamily Nudocotylinae was created by Barker
(1916) to hold Nudocotyle novicia. In this Barker would include Barisomum
erubescens Linton (1910).

NOTOCOTYLINAE Kossack 1911

Small to medium sized Notocotylidae with thin cup-shaped body; two
to five rows of prominent papillae on ventral surface. Genital pore median,
near intestinal bifurcation. Cirrus pouch enclosing only a small part of
seminal vesicle. Vitellaria well developed occupying a region posterior
to middle portion of body anterior to testes and lateral to intestinal crura.
Ovary and testes symmetrical, in extreme posterior part of body. Ovary
between testes and separated from them by intestinal crura. Uterus regu-
larly coiled, between intestinal crura.

Type genus Notocotylus. Other American genera Catatropis and
Paramonostomum.

NOTOCOTYLUS Diesing 1839

Syn: Notocotyle Diesing 1850

The genus Notocotylus was formed by Diesing in 1839 to include
Fasciola verrucosa Frolich, Fasciola anseris Gmelin, Festucaria pedata
Schrank and Monostoma verrucosum Zeder. It was characterized by the
author as follows: "Corpore oblonga-ovato, depressiuscula, antice parum
attenuate, postice rotundato, ore terminali orbiculari; acetabulis suctoriis
dorsalibus numerosis, serie triplici longitudinali; cirro longo spirali ven-
trali." In 1850 the author changed the name to Notocotyle with only a
slightly modified diagnosis as follows: "Corpus oblongum depressum.
Caput corpore continuum. Os subterminale anticum. Acetabula numerosa
(24-50) juxta totam dorsi convexiusculi longitudinem treseriata sissilia,
orbicularia, limbo callosa. Penis ventralis superus longi spiralis. Porus
excretorius .... In avium intestinis crassis et coecis endoparasita!"
Under this caption Diesing included his former genus Notocotylus. Altho
Monticelli, Barker and others adhere to the more recent form of the name

269] NORTH AMERICAN MONOSTOMES 51

the writer feels justified under Article 32 of the International Rules of
Zoological Nomenclature in accepting with Kossack, Ward and others the
older name Notocotylus.

This genus is up to the present represented in America by a single spe-
cies Notocotylus quinqueserialis (Barker and Laughlin). Altho Barker
(1916) would place Monostomum affine Leidy in this group, his determina-
tion seems to be unwarranted on the basis of the description of Leidy which
shows distinct anatomical differences namely a small pharynx, echinate
penis, a well marked excretory canal traceable to the beginning of the
oviduct, and sub-pyriform eggs prolonged at one pole only. In addition to
the anatomical differences Monostomum affine was found parasitic in the
gall bladder and gall ducts of Fiber zibethicus whereas Notocotylus has been
taken only from the intestine and ceca of the muskrat and water birds.

NOTOCOTYLUS URBANENSIS (Cort 1914)
[Figures 12, 14, 17, 18, 19]

Syn: Monostoma sp. Stiles and Hassall 1894

Medium sized worms 2.5 to 3 . 5 mm long by 0 . 5 to 1 mm wide, having
three rows of ventral glands each row containing 13 to 14 glands. Oral
sucker strongly muscular 112 to 153ju followed by a short esophagus with-
out pharynx; intestinal crura provided with numerous short diverticula
both externally and internally. Genital pore just posterior to intestinal
bifurcation. From this point the cirrus pouch extends caudad to the end
of the first body third or a little beyond this level. Vagina one-half the
length of the cirrus. Usually about ten uterine loops anterior to the most
anterior part of the vitellaria which as in other species of the genus lie
lateral to the ceca and extend from the middle of the body to the lobed
testes in the posterior end. The irregularly lobed ovary is situated between
the testes and is separated from them by the crura. Eggs numerous pos-
sessing two long polar filaments. Eggs without filaments measure 20^1 in
length, and are approximately one-half that in width.

Habitat: Intestine Host: Dafila acuta

Locality: Maryland Collector: A. Hassall

Date: January, 1893 No. 5772 U.S.N.M.

Habitat: Cecum Host: Fiber zibethicus

Locality: Maryland Collector: A. Hassall

Date: June 23, 1892 No. 5769 & 5770 U.S.N.M.

Habitat: Intestine Host: Aix sponsa

Locality: Maryland Collector: A. Hassall

Date: August, 1893 No. 5771 U.S.N.M.

Notocotylus urbanensis agrees with Notocotylus attenuatus in size and
form, and in the relative length of the cirrus pouch and vagina. With
respect to the number of papillae in each row it conforms more closely

52 ILLINOIS BIOLOGICAL MONOGRAPHS [270

to Notocotylus aegyptiacus; it diflfers, however, from this species in the
relative length of the cirrus pouch and vagina. In the position of the
genital pore it agrees with Notocotylus seineti Fuhr. and in this respect it
dififers from other known species of this genus.

Stages in the Life History

Three collections No. 5769, 5770, and 5771 of the United States
National Museum contain immature forms of this species. Collection No.
5771 contains both immature and sexually mature worms. The very young
stages found in the two collections from the muskrat agree so well with
the immature forms from Aix sponsa that it is impossible to differentiate
the two forms and consequently they are taken to be identical.

The most immature specimens have apparently just burst out of the
cysts since the pigmentation can be seen quite as perfectly as in most
Notocotylid oercariae. The pigmentation in this species agrees generally
with that described by Cort (1914) for Cercaria urbanensis. The pig-
mentation remaining is arranged around the lateral eye spots (Fig. 19) and
the lateral pigmented lines extending from the eye spots to near the
posterior end. Aside from this there is a very diffuse pigmentation through-
out the entire body. From the time of encystment of the cercaria to the
youngest stages at hand considerable change has taken place. The
anterior eye spot has been lost and the general pigmentation as described
above is generally much reduced as compared with the heavily pigmented
Cercaria urbanensis. The locomotor pockets have been resorbed so that
no trace of them exists in the youngest stages at hand. Development of
the ventral glands is the most conspicuous change which has taken place.
Figure 14 shows diagrammatically the youngest stage studied in which
three ridges or keels are thrown out on the ventral side. The median one
being about twice as high as the lateral ones which are ventral in position
to the intestinal crura (Fig. 12). Along the median ridge the papillae
altho only partially differentiated are clearly seen. The lateral ridges
show indistinct irregularities which in section are clearly the beginnings
of the papillae. No trace of the outer rows of papillae which occur in
Notocotylus quinqueserialis have been observed. In more mature stages
the ventral papillae are distinctly seen (Figs. 17, 19).

Faust (1918) stated that in the Monostomata the paired ceca are
filled with a jell and are nonfunctional in the cercaria stage. In contrast
to this the ceca in the youngest stages studied, which are of course well
past the cercaria stages Faust studied, show that changes have taken
place in this feature. The ceca in these stages show the intestine as a tube
(Fig. 12) whose walls are surrounded by large nucleated cells. Totos in
this stage of development also show distinct but small internal and external
diverticula (Fig. 19).

271] NORTH AMERICAN MONOSTOMES 53

The genital glands show little development over that of the cercariae.
The ovary and testes are made out readily in the toto mount as well as the
cords of cells which are to dififerentiate into uterus, vagina, vas deferens
and cirrus. In more mature stages differentiation of the cirrus and vagina
is well started so that the relative length of the two organs can be deter-
mined.

In the more mature specimens represented in figure 17 the genital
glands have made a tremendous growth and appear very much as in the
sexually active worm. The uterus, however, is less distinct and probably
still non-functional. It is in this stage that the vitelline glands make their
first appearance and here appear as single celled isolated follicles. The
ducts of these follicles cannot be traced so that it is impossible to determine
if the relation found by Faust (1918), namely that the vitellaria of Notoco-
tylids are composed of five inner and three outer portions, obtains in this
species.

Of the seven larval Monostomata described from North America, viz.
Cercaria hyalocauda Haldemann 1842 Cercaria konadensis Faust 1918
Glenocercaria lucania Leidy 1877 Cercaria aurita Faust 1918

Cercaria urbanensis Cort 1914 Cercaria rohusta Faust 1918

Cercaria pellucida Faust 1918
the immature stages described above resemble more closely Cercaria ur-
banensis Cort (1914) than any other known monostome cercaria. Based on
the similarity of the excretory system, of the genital organs, on the pig-
mentation and on the late differentiation of the vitellaria, no trace of which
has yet been found in Cercaria urbanensis, it seems highly probable that
these forms can be actually connected. Hence while demonstration of the
life history by experimental methods has not at this time been given, it
seems justifiable to accept Cercaria urbanensis as the larval form of this
Notocotylid.

NOTOCOTYLUS QUINQUESERIALIS (Barker and Laughlin)

Syn: Monostoma sp. Stiles and Hassall 1894

Notocotyle quinqueseriale Barker and Laughlin 1911

Medium sized to large Notocotylids with wedge shaped body 2 . 5 to
4 mm long by 0 . 66 to 1.33 mm in maximum width which is found at the
level of the ovary. Anterior end more or less pointed, posterior end
rounded. Dorsal surface smooth, convex, ventral concave, unarmed but
possessing five longitudinal rows of adhesive glands or papillae. Each
row containing from 16 to 18 distinct wart-like projections. Mouth sub-
terminal, spherical 200 to 450^t in diameter; esophagus short without phar-
ynx. Intestinal crura irregular in shape and size with short internal and
external diverticula. Genital pore between mouth and intestinal bifurca-

54 ILLINOIS BIOLOGICAL MONOGRAPHS [272

tion. Cirrus pouch extending from this point into the beginning of the
second body third, being approximately one-third the length of the entire
body. Vagina two-thirds the length of the cirrus pouch. The much lobed
testes are situated in the posterior end, external to the intestinal crura.
Ovary irregularly lobed, on a level with and between the testes, intracecal
in position. Vitellaria extracecal, anterior to the testes, extending to the
middle of the body.

Habitat: Intestine Host: Arvicola riparius

Locality: Maryland Collector: A. Hassall

Date: 1892 No. 5773 U.S.N.M.

Habitat: Intestine Host: Fiber zihethicus

Locality: Baker Lake, Washington Collector: H. E. Metcalf
Date: August 13, 1915 No. 15.120 Ward collection

The description of Barker and Laughlin while adequate states that the
intestinal ceca are simple. A careful examination of the material at hand
shows small, short diverticula both externally and internally throughout
the length of the ceca. The writer has not had opportunity to study
the material described by Barker and Laughlin.

CATATROPIS Odhner 1905

Body elongate, anterior and posterior ends usually equally rounded.
Anterior half of ventral surface covered with three rows of non-protrusible
papillae; median row set on a ridge or keel; lateral rows each containing 8
to 12 glands. Vagina strongly developed, usually as long as the cirrus
pouch.

This genus was created by Odhner to hold Catatropis verrucosa (Fro-
lich) which Odhner found to differ from Notocotylus in the character of
the ventral glands, those of Notocotylus being protrusible while those found
in Catatropis verrucosa were very much reduced, being in the form of a
median ridge or keel in the median row and small embedded papillae or
glands in the outer rows. It is similar to Notocotylus in its inner organiza-
tion and differs from that genus in the fact that the ventral glands are not
protrusible.

CATATROPIS FILAMENTIS Barker 1915

Syn: Catatropis fifubriata Barker 1915

Thin flat worms, gradually tapering anteriorly, 2.2 to 3.3 mm long
by 0 . 56 to 0 . 7 mm wide at the level of the testes. Anterior half of the body
covered with needle like spines arranged in oblique rows. Three rows of
flattened papillae on the ventral surface, 12 to 13 in each row. Oral sucker
spherical, 66 to 99 /j. in diameter. Esophagus 105 to 132 ;u in length.
Pharynx wanting. Intestinal crura undulating. Testes two to four lobed,
external to the intestinal crura. Ovary globular or oval 132ju long by 105

273] NORTH AMERICAN MONOSTOMES 55

to 112/i wide, margin irregular. Shell gland ovoid, anterior to and a little
larger than the ovary. Cirrus pouch tubular, elongate, extending to the
beginning of the second body third. Prostate gland and cirrus covered
with papillae. Vagina straight, muscular, as long as the cirrus pouch.
Vitellaria external to the ceca, extending from the middle of the body
caudad to the testes. Excretory bladder forked, opening to the exterior
just dorsal to the posterior end. Eggs thick shelled, 20 to 22/i long by ll/>i
wide, having two long polar filaments, one at each end.

Habitat: Duodenum Host: Fiber zihethicus

Locality: Nebraska Collector: ?

PARAMONOSTOMUM Liihe 1910

This genus, created by Liihe to hold Monostomum alveatum (Mehlis)
Creplin, is characterized by Liihe as follows: Body compressed, egg shaped,
greatest breadth a little caudad from middle of body, posterior end broadly
anterior tapering and pointed; anterior half of ventral surface thick set
with short heavy spines. Ventral glands absent. Cirrus pouch weakly
muscular. Vagina usually one-half length of cirrus pouch.

Type species: Paramonostomum alveatum (Mehlis) Crepl.

American representative : Paramonostomum echinum nov. spec.

Barker (1916) criticises the erection of a new genus on the basis of the
absence of the ventral glands on the ground that the number of rows vary
from two in N otocotylus diserialis Ssinitzin to five in N otocotylus quinqueser-
ialis Barker and Laughlin. Yet the same author accepts the genus Catatro-
pis of Odhner founded on the non-protrusible character of these same
glands. There is apparently as much reason to accept the genus of Liihe
based on their absence as that of Odhner founded on their non-protrusi-
bility.

PARAMONOSTOMUM ECHINUM nov. spec.

Thin cup-shaped worms, 2 to 2.5 mm in length by 0.6 to 0.7 mm in
maximum width which is found at the beginning of the posterior third
of the body length. No ventral papillae have been found on these worms,
the anterior half of the ventral surface being covered with heavy spines
5ju in length. These curve caudad and are thick set according to the
definite pattern shown in figures 13 and 16. Mouth terminal, spherical,
102 to 125/i in diameter, followed by a short esophagus which bifurcates
to form the intestinal crura; these follow an undulating course to the
posterior end of the body where they end blindly. Crura provided with
short but definite internal and external diverticula. Genital pore situated
just posterior to the intestinal bifucation. Cirrus pouch extends from this
point into the beginning of the second third of the body. Vagina one-half
as long as the cirrus pouch. Prostate gland and cirrus without papillae.

56 ILLINOIS BIOLOGICAL MONOGRAPHS [274

Testes four lobed, extracecal, lying at the same level in the posterior end of
the body. Ovary between the testes and separated from them by the
intestinal crura. The three to four lobed ovary is usually elongated antero-
posteriorly. The uterus as in other members of this genus is coiled trans-
versely between the crura and extends from the level of the ovarian com-
plex to the posterior end of the cirrus pouch at the beginning of the second
body third. The vitelline glands occupy an extracecal position and extend
from the testes to the middle region of the uterine coils which is found in
the caudal portion of the second body third. Eggs numerous, medium
thick shelled, 20 /i in length and approximately twice as long as wide.
They possess a long polar filament at each end.

Habitat: Intestine Host: Fiber zibethicus

Locality: Wray, Colorado Collector: C. H. Gable

Date: October 30, 1916 No. 21.91 Ward collection

NUDOCOTYLINAE Barker 1919

Small cup shaped Notocotylidae with thick bodies, without ventral
glands. Genital pores separate, ventral, lateral, in posterior half of body.
Cirrus pouch pear-shaped, enclosing small portion of seminal vesicle.
Vitelline glands strongly developed compact masses, lateral to ceca and
anterior to testes. Uterus in transverse folds, in anterior half of body,
extending laterally over intestinal crura.

Type genus: Nudocotyle Barker 1916

Barker would include Barisomum Linton 1910 in this sub-family.
This is a doubtful decision since in the genus Barisomum the genital pore
is in the anterior body third and the shell gland lies posterior to the ovary.
It possesses in fact certain Notocotylid characters but conforms more close-
ly to the Pronocephalidae than to the Notocotylidae in the position of the
genital pore, and the character and position of the genital glands.

NUDOCOTYLE NOVICIA Barker 1916

Small thick oval worms, 709 to 899 )u in length; 500 to 657)u in breadth.
Anterior end tapering gradually, posterior markedly truncate. Dorsal
surface strongly convex, ventral concave. Body smooth, devoid of ventral
papillae or spines. Oral sucker subterminal, spherical 50 to 65 yu in diame-
ter; pharynx wanting; intestinal ceca undulating but without diverticula.
Male and female genital pores separate, ventral, lateral, in beginning of
posterior body third. Cirrus pouch large, club-shaped, about one-third of
body width in length. It lies transversely and median in anterior portion
of posterior body half. Cirrus without spines. Testes extracecal in posi-
tion, lying in same level in posterior fifth of body, frequently 2 to 5 lobed.
Ovary elongated, convoluted or lobed, in extreme posterior end of body
between testes and separated from them by intestinal crura. Shell gland

275] NORTH AMERICAN MONOSTOMES 57

compact, anterior to ovary. Laurer's canal and receptaculum seminis
were not observed by Barker. Eggs oval, twice as long as wide, 20 to 24 )u
long 10 to 13^t wide. With long polar filament on each end about five
times as long as egg itself.

HERONIMIDAE Ward 1917

This family was created by Ward (1917) to hold the two aberrant genera
Heronimus MacCallum (1902) and Aorchis Barker and Parsons (1914).
Ward called attention to the close resemblance of the two forms and
suggested that they might prove to be identical. He characterized the
family as follows: ''Moderate sized monostomes with thick, elongate,
soft body, slightly flattened, tapering toward both ends. Oral sucker weak;
pharynx large; esophagus short or absent; ceca simple, narrow, extending
to posterior tip but not united. Vitellaria compact, tubular. Uterus with
four longitudinal regions; genital pore ventral to oral sucker, near anterior
tip. Testis tubular, small; copulatory apparatus poorly developed. In
lungs of turtles, northern North America."

One year later (1918) the same author restated the family diagnosis
with the following addition: "Vitellaria compact tubular, shaped like an
inverted V. Testes tubular, lobed or with short branches, united into a
V-shaped organ with the apex anteriad," and again stated that the two
forms probably belonged to the same genus. About a year later Stunkard
(1919) presented a paper in which he showed that the apparent difference
in the two forms was due largely to the partially diagrammatic figure of
MacCallum (1902) and to the discrepancies in the description of Barker
and Parsons (1914) and that the two forms are identical, thus not only
belonging to the same genus as suggested by Ward but representing a
single species, Heronimus chelydrae MacCallum.

HERONIMUS CHELYDRAE W. G. MacCallum 1902

Syn: Aorchis extensus Barker and Parsons 1914
Aorchis extensus Ward 1917
Monostoma sp. Stiles and Hassall 1894

The genus was created by MacCallum (1902) to include worms collected
from the lungs and bronchi of Chelydra serpentina. The genus stands
according to MacCallum "in many respects far apart from the other
genera," especially in the position and nature of the genital opening, in the
complicated structure and course of the uterine tract, in the unusual
formation of the yolk glands, in the presence of but one testicle and in the
position of the excretory pore. The genus may be recognized by the
following diagnosis: Medium to large monostomes with semicyclidrical
body tapering slightly towards both ends; strongly muscular. Mouth

58 ILLINOIS BIOLOGICAL MONOGRAPHS [276

opening terminal, oral sucker small, pharynx weak but distinct. Esophagus
very short; intestinal ceca simple, ending blindly in the extreme posterior
of the body. Genital pore inconspicuous, median, ventral to the pharynx.
Ovary situated in the anterior one-fourth of the body, lateral, usually
intracecal; shell gland smaller than the ovary and posterior to that organ.
Receptaculum seminis present, usually about two-thirds as large as the
ovary. Laurer's canal absent. Uterus, except for the four longitudinal
loops, coiled around the intestinal crura from the level of the ovary to the
posterior end of the animal. Vitelline gland a coarse, compact U-shaped,
closed tubular structure, dorsal to the intestine. Testis U-shaped, closed
portion cephalad, about one-fourth the body length from the anterior
end. The tubular, irregularly lobed testicular mass extends caudad to a
level about one-eight the body length from the posterior end. Protrusible,
non-muscular cirrus present. Excretory pore median, anterior, dorsal to
the pharynx. Eggs large, ovoid, thin shelled, containing fully developed
miracidia in the metraterm.

The anatomy of this form is well described by MacCallum (1902) and
Barker and Parsons (1914, 1917), and with the additions and corrections
of Stunkard (1919) calls for no further anatomical discussion here.

The writer has been given an opportunity to examine the type specimen
of this species deposited in the United States National Museum and can
verify the statements of Stunkard on the specific identity of the two species.
MacCallum reported the original material from Chelydra serpentina
taken in the Grand river at Dunville, Ontario, Canada. Barker and
Parsons (1914) report from Chrysemys marginata taken in Lake Emily,
Minnesota, and the Mississippi river and later (1917) in the same host
taken in the Mississippi river near Fairport, Iowa. At this time they call
attention to the distribution in Illinois.

Ward (1917) reports this species from "various turtles" taken in Michi-
gan, Indiana, Illinois and Nebraska. Stunkard (1919) collected this species
from Chelydra serpentina taken in Illinois, Ohio, North Carolina and Texas;
in Chrysemys marginata taken in Iowa, Illinois, Missouri and Kentucky;
in Pseiidemys elegants and Malacoclemmys geographicus in Illinois; Aro-
mochelys odoratus and Kinosternum pennsylvanicum in North Carolina.
The specimen listed by Stiles and Hassall (1894:253) as "Monostoma sp. —
Chelonia gen. sp. (bronchi) — Illinois-Forbes-Leidy" belongs here.

The writer has found this species in Chelydra serpentina and Chrysemys
marginata taken in the drainage ditch at Urbana, Illinois; in Chrysemys
marginata taken in the Mississippi river near Fairport, Iowa; in Graptemys
geographicus taken near Chicago, Illinois; in Chelydra serpentina and
Chrysemys marginata taken in Minnesota and in Kinosternum pennsylvani-
cum, Kinosternon odoratus, and Chrysemys picta, a new host, taken in North
Carolina.

277] NORTH AMERICAN MONOSTOMES 59

While this species was reported by MacCallum as "not by any means a
constant parasite," he having found it in only one host infected of a number
examined, more recent data show this worm to be rather constantly present.
In seven specimens of Chrysemys marginata collected in the summer of 1911
from Lake Emily, Minn., Barker and Parsons found five infected, one of
them yielding thirteen worms from both lungs. The same authors (1917)
found that female turtles were more than three times as heavily infected
as males, Stunkard (1919) from the examination of "about three hundred
turtles" reports the heaviest infection in one host as six. On an examina-
tion of "more than fifty turtles" he found no difference in the relative in-
fections of males and females. The writer has examined one hundred and
two hosts of six different species and of these forty-four showed infection
with these worms. The highest percentage of infection for any species was
found in Cinosternum pennsylvanicum from North Carolina in which 34
out of 45 specimens or approximately 75 percent were infected. The highest
number of individuals from a single host of this species was eleven while a
single individual of Graptemys geographicus, and the only one of eighteen
which showed infection, carried twelve worms in both lungs.

During the past four years the writer has had opportunity to examine
more than one-hundred turtles as stated above. It was noted early in the
investigation that the collections made in different seasons showed no
striking difference in percentage of infection. It was then undertaken to
determine if possible the length of life of this parasite in the definite host.
It is generally understood that most intestinal paraties have an annual
cycle and depend on this for continuity of the species. However, data on
this point seems lacking in this group. The work on Heronimus chelydrae
consisted in the examination of a number of turtles collected in the same
region, Raleigh, North Carolina, at various seasons of the year.

Some of these dissected on arrival showed relatively heavy infections,
others which were kept in the laboratory aquaria for periods of six, twelve
and eighteen months still carried infection, and a single specimen of Chely-
dra serpentina which had been kept in an aquarium for more than three
years yielded two specimens of Heronimus chelydrae and a single nematode,
probably Camallanus americanns. As was stated above hosts examined
when taken, usually carry intestinal forms in addition to the lung fluke
already mentioned, while those which have been kept in aquaria for a
period of six months or more show a marked reduction in the number of
the intestinal forms. There is no apparent change in the number of the lung
flukes present.

Little is known regarding the condition of parasites during hibernation
of the host. Blanchard (1903) records that hibernating marmots do not
contain any intestinal parasites. Ward (1909) reports observations carried
out on the frog, Rana virescens. In this he says that parasitic infection
increases steadily up to hibernation, and does not decrease during the latter

60 ILLINOIS BIOLOGICAL MONOGRAPHS [278

period; that the parasites reach the climax of sexual maturity soon after
the host emerges from the winter quarters. They discharge their eggs
and pass out from the body of the host soon after the time of spawning,
and at the close of this period the hosts are relatively free from infection.

Observations of the writer verify the sexual inactivity of the parasite
Heronimus chelydrae during the early part of the winter and the copious
discharge of mature miracidia in early spring. On the other hand these
turtles did not become free from infection at any time during the period
of the experiment. It must be kept in mind however that the turtles used
were not subject to natural conditions, i.e., no opportunity was afforded
for hibernation and no eggs were deposited during this time. That trema-
todes adapted to partially closed cavities can live longer than for a single
reproductive phase is evident, since in the instance reported above in which
the host has been kept for a period of more than three years, the two para-
sites found were sexually mature, and were producing large quantities of
ripe eggs when the host was examined; this is true also in one other case
in which the host was kept for more than eighteen months.

COLLYRICLIDAE Ward 1917

This family was created to hold the genus Collyriclum of Kossack and
is circumscribed by Ward as follows:

"Small to moderate sized monostomes with discoidal compressed, not
muscular body, broader than long. Oral sucker weak; pharynx present;
ceca simple, long, capacious, not united. Genital pore ventral near center
of body. Vitellaria follicular, scanty, antero-lateral; ovary much lobed,
symmetrical. Uterus posterior, in irregular coils which show an antero-
posterior tendency, terminal region enlarged. Testes oval, symmetrical,
behind ovary. Eggs very small. Adults parasitic in dermal cysts on abdo-
minal surface of the skin of birds."

In the light of our present knowledge of these forms the family diagnosis
must be modified with respect to the condition of the testes. Tyzzer (1918)
has shown the testes of the American species to be irregularly lobed and
not oval as described for the European species, Collyriclum faba by both
Kossack (1911) and Jegen (1917). The writer has examined a number of
specimens of Collyriclum colei Ward and has found the observation of
Tyzzer stated above to be correct.

Type and only genus: Collyriclum Kossack 1911.

American representative: Collyriclum colei Ward 1917

COLLYRICLUM COLEI Ward 1917
Syn: Monostoma faba Cole 1911

Collyriclum faba Tyzzer 1918
Diagnosis: Small hemispherical worms, 4 to 5 mm in length and breadth
by 3 mm thick. Cuticula covered with spines 35 fx in length, arranged in
groups which form rather regular rows around the worm. Mouth terminal

279] NORTH AMERICAN MONOSTOMES 61

or slightly dorsally placed, surrounded by a muscular sucker 220/i long by
375 )u wide. Pharynx smaller 140^1 long by 125 m wide, adjacent to the
sucker, followed by short esophagus which bifurcates to form the large
voluminous simple intestinal crura; these end blindly at the end of the
middle body third. Genital orifice ventral, near center of body. Testes
near the ends of the crura, lobed, the main portion pear-shaped. Ovary
in intestinal bifurcation, three branched, each division containing from 5
to 10 lobes. Vitellaria well developed, imperfectly symmetrical with 5 to
7 groups on the left and 7 to 9 on the right. Uterus much coiled, generally
in the posterior half showing a tendency to antero-posterior coiling. Eggs
small, 19 to 22ju long by 10 to 12^t wide, containing in the end portion of
the uterus a fully formed miracidium.

The anatomy of these worms has been so thoroughly discussed by Tyz-
zer (1918) for the American species, and by Kossack (1911) and Jegen
(1917) for the European species that it does not seem necessary to enter
into a detailed discussion here. I desire instead to give a comparison of the
two forms since Tyzzer found reason based largely on the inconsistency
of Kossack's description and figures to declare the American material
identical with that found in Europe.

Ward (1917) after examination of the material reported by Cole (1911)
as Monostoma faba pointed out distinct differences between this and the
European form described by Kossack (1911). The following is his state-
ment, "As a cause of an epidemic among sparrows at Madison Wisconsin,
Cole (1911) reported under the name of Monostoma f aha a trematode that
in reality differs distinctly from the European species. The form of the
ovary, the extent of the vitellaria, the dermal spines, and other details of
structure disagree with the recent description of Kossack who, moreover,
assigned Rudolphi's species." [erroneously attributed to Rudolphi, really
Bremser (1831) ] "to his new genus Collyriclum. The American form
constitutes a new species in this genus and to it the name Collyriclum colei
may be given."

One year later (1918) the same author restated the differences in the
two species as follows: "These specimens differ clearly from the European
form in numerous minor details, such as ovary, yolk glands, dermal spines,
etc., and demand recognition as a distinct species under the name given
here."

Tyzzer (1918) made a comparison of the two forms based on Kossack's
description of Collyriclum faba and concluded that the two species were
identical despite certain distinct differences which he explained away on the
basis of the discrepancies between Kossack's description and figures. The
work of Jegen (1917) which reached America after Tyzzer's had appeared
verifies in a large measure the description of Kossack and leaves little
doubt that the American material is distinct from Collyriclum faba. The
following is the diagnosis of this species as given by Jegen: "In Cysten

62

ILLINOIS BIOLOGICAL MONOGRAPHS

[280

Table I. Comparison of Collyriclum faha with C. colei
After Tyzzer, 1918, with additions from Jegen, 1917

European Material

Collyriclum faba

American Material
Collyriclum colei

Shape

Somewhat hemispherical.

The same. Minor additional
points noted.

Size

4.2-4.8X4.5-5.46 mm
Jegen from P. domesticus
4.4-5.2X6-5.4.

4.1X4.8X2.9 mm.

Mouth

Ventrally placed.

Dorsal to margin of body. In
flattened specimens appears
ventral from over-riding of
larger dorsal surface.

Uterus

Similarly arranged in both.

Intestine

Identical in form.

and extent.

Vitellaria

Symmetrical, each with
seven rarely six or eight,
follicle groups.

Not perfectly symmetrical, five
to seven on left and seven to
nine follicle groups on right
side.

Ovary

T-shaped, each of three divisions
with four or five branches.
Jegen reports 5-7 lobes.

Of similar form, each division
from 5-10 lobes.

Testes

Oblong or saber-shaped.

Showing three or more large
processes and other minor
irregularities, and curved over
the blind ends of the intestine.
In gross specimens seen only
in part, appearing oblong or
saber shaped. Measurement
not feasible.

Genital orifices

General agreement in both.

Oral sucker

0.204,5 to .441, 2 mm.
Jegen, 0.3 to 0.45 mm.

Average, 0.219,8X0.375,3 mm.

Pharynx

0.129,1 to 0.193,7 mm.
Jegen, 0.113 to .145 mm.

Average 0. 140X . 124,5 mm.

Spines

"Arranged in lines, with the in-
dividual spines apparently
widely separated from one an-
other." Up to 35m in length.

Jegen: in groups of 4 to 8,
28 to 35m.

Set in annular rugae; maximum
distance separating latter, .45
to 53m. Up to 35,5m in
length. Average dorsal, 27,9m
long.

Ova

19,8X9,7m.

19 to 21,4X10,6-11,6m.
Average 20,5X1 1,3m.

281] NORTH AMERICAN MONOSTOMES 63

zu zweien vorkommende Trematoden. Korperform annahernd rund.
Dorsale Flache stark gewolbt, ventrale weniger gewolbt bis flach. Haut
mit Stacheln besetzt, die in Gruppen von vier bis acht Einzelstacheln
stehen. Mundsaugnapf endstandig. Darmschenkel einfach und zwei
Drittel der Breite des Korpers einnehmend. Bauchsaugnapf fehlt. Ter-
minal am Hinterende eine muskelreiche Partie, die bei der Fortbewegung
als Saugorgan Wirkt. Genital pori auf einer papillenartigen Erhohung,
median etwas vor der Korpermitte gelegen. Excretionsblase birnformig
und bedeutend uber die Mitte hinausreichend. Dotterstocke aus zwei
seitlich gelegenen Follikelgruppen (7) bestehend. Hoden dorsal, den
Darmschenkelspitzen genahert. Keimstock vor den Hoden, im ersten
Korperdrittel, aus drei lobosen Gruppen bestehend. Schalendriise unmit-
telbar neben und unter dem Keimstock. Laurersche Kanal vorhanden.
Receptaculum seminis fehlt. Uterusschlingen hauptsachlich im hinteren
Korperteil. Eier ohne Filamente, mit scharf abgesetztem Deckel und
einer kleinen, seitlichen Spitze am entgegengesetzten Pol, sehr zahlreich."

Because of the distinct differences shown by Tyzzer his table is incor-
porated with Jegen's corrections to Kossack's description of Collyridum
faba. It is printed on preceding page of this paper.

A study of the table shows clearly that the species in question differ
with respect to the asymmetry and extent of the vitellaria and the form
of the testes. In regard to the agreement of the American material with
Kossack's description Tyzzer says: "that the American material agrees
very closely in most respects with Kossack's description, there being
similarity of size and shape, in the appearance of the alimentary canal and
uterus, and in the position of the genital orifices. The measurements of the
oral sucker, pharynx, esophagus, spines, and ova correspond rather closely,
and such differences as occur appear to be within the limits of species
variation."

Regarding the grouping of the spines Kossack (1911:574) "Die Haut
ist mit Stacheln bedeckt, die in regelmassigen Reihen angeordnet sind.
Die einzelnen Stacheln sind ziemlich weit voneinander entfernt und
durchschnittlich 0.035 mm lang." Jegen describes the spines as being
arranged in groups in rows. Four to eight spines in a group, and says,
"Kossack erwahnt (S. 574) dass die Stacheln in regelmassigen Reihen ange-
ordnet seien. Ich glaube nun nicht, wie Odhner dies ausspricht, dass er
die Stachelgruppen iibersehen hatte, wenn sie iiberhaupt in seinem Mate-
rial vorhanden waren. Vielmehr liegt die Moglichkeit vor, dass er ein
Entwicklungsstadium vor sich hatte, bei dem die Gruppen noch nicht
vollstandig gebildet waren."

Tyzzer calls attention to the ovary which he says "presents more
lobules than was noted by Kossack." Jegen states that the ovary is very
strongly lobed and found: "An jedem der drei Aste sitzen fiinf bis sieben

64 ILLINOIS BIOLOGICAL MONOGRAPHS [282

solcher einzelner KnoUen." Tyzzer states further that "the basis for differ-
entiation of an American species at the present time appears therefore to
be rather inadequate." That he is dealing with the same species as that
reported by Cole (1911) and later named Collyriclum colei by Ward (1917)
cannot be doubted and according to his statement his material "is un-
doubtedly of the same species,"

Although Tyzzer failed to find any difference which would warrant a
second species of Collyriclum the present investigation has shown that
except in minor detail the work of Jegen agrees with that of Kossack, and
on the basis of the descriptions of these investigators the American mate-
rial is distinct from Collyriclum faba in that the testes are lobed and in the
asymmetry and extent of development of the vitellaria. It is impossible
to explain away these differences either as "artefacts in preservation" or
as "individual variation," nor is it probable that both Kossack and Jegen
have overlooked these features. On the basis of these difference the Amer-
ican material must be recognized as a distinct species under the name Colly-
riclum colei Ward.

A preliminary study of specimens of Collyriclum faba secured by Pro-
fessor Ward bears out the description of Kossack and Jegen as well as the
conclusions of the writer drawn from the study of their papers and compari-
son with American specimens of this genus. A more thorough study of the
European material is not feasible at this time but is anticipated at an
early opportunity.

REMARKS ON THE LIFE HISTORY

The life history of Collyriclum faba is doubtfully known. Tyzzer gives
a careful description of the development of the egg of Collyriclum colei
from the time of fertilization to maturity in the end portion of the uterus.
Regarding the mature eggs he says "The eggs stored in the terminal por-
tion of the uterus evidently contain miracidia, the morphological features
of which are not clear in fixed material owing to imperfect preservation
and shrinkage." He continues with a description of the "hair-like"
structures which have been distinguished. This adds evidence that a
miracidium is present.

The work of Jegen (1917) which is an attempt at the life history of
Collyriclum faba differs radically from the statements of Tyzzer in that he
finds the eggs contain two embryos which are not miracidia but young
trematodes; these need only to be incubated in the intestine of the host
that they may break out of the egg shell and freed with the excrement,
may wander into the feather follicle. He found also cysts (Dauercyste)
which after a longer period of incubation break open and the worm enters
the follicle of a feather. Jegen summarizes his work as follows: "Die
Eier von Collyriclum faba werden durch den Wirt mit dem Parasiten aufge-

283] NORTH AMERICAN MONOSTOMES 65

pickt und gelangen in den Vogeldarm, wo die Embryonem ausschliipfen.
Mit den Excrementen werden letztere ins Freie befordert, wo sie, sofern
die Moglichkeit zur Infektion vorhanden ist, direkt in die Federfollikel
der jungen Vogel einwandern. Im andern Fall bilden sich Dauercysten,
die nach langerer Entwicklungsruhe such auflosen und den eingeschlos-
senen Organismus frei lassen, so dass er ebenfalls in die Federfollikel ein-
wandern kann." This summary of Jegen is supported by experimental
evidence gained by incubation of the eggs in a portion of the intestine of
an infected bird as well as by numerous attempts to incubate the eggs
which had not passed through the intestine of the host altho he says these
gave negative results. The "Dauercysten" he found in excrement of infec-
ted birds, which was dried by exposure to air, and at other times in the
nest and on the feathers of young sparrows.

The outline of Jegen gives essentially a direct development which
omits the parthenogenetic stages observed in all cases where the life
history of digenetic trematodes is known. Observations of the writer
support the view of Tyzzer that a miracidium is present in the mature egg
in the uterine egg sac. The fully developed miracidium shows well devel-
oped germ balls. This is in direct contrast to the findings of Jegen who
says that the embryo contains two well developed germ balls with numer-
ous others which disappear later in course of development. Jegen found
that the eggs would develop only in the intestine of the host and when fed
to uninfected birds empty egg shells and embryos were found on the second
and third day. He neither states nor demonstrates that these experimental
birds became infected with the adult parasites.

The work of Jegen dealing with the life history of Collyriclum is full
of gaps. The life history as given is bridged over by supposition. The
infection of the sparrows by feeding of eggs is not demonstrated, only
the presence of embryos which may well be miracidia. Jegen does not
demonstrate beyond doubt that these cause the formation of the cysts in
the sparrow. The "Dauercysten" which h'C says infect directly the host
based on his experiment of the dried excrement certainly serve to protect
the parasite from dessication until it can reach the intermediate host;
nothing indicates whether this be a miracidium or another infective stage;
and the periodicity of occurrence of the adult parasite in correlation with
a rainy season tends to show that the former is the correct interpretation.

The work of Jegen is apparently a misinterpretation of the life history
and leaves much to be done in order to demonstrate the facts. The only
clear contribution of this author to the life history of this form lies in the
discovery of the "Dauercysten" which he apparently misinterpreted.
Many more extensive and careful experiments must be carried out in order
to demonstrate conclusively the complete life history of this form.

66 ILLINOIS BIOLOGICAL MONOGRAPHS [284

Species Inquirendae
CLINOSTOMUM (?) INCOMMODUM (Leidy)

Syn: Alonostomum incommodum Leidy 1856
Distoma oricola Leidy 1884
Distomum incommodum Leidy 1890
Monostomum incommodum Leidy 1904

Monostomum incommodum is described by Leidy as follows: "Body
compressed, above convex, below concave, sides parallel anteriorly convex,
posteriorly angularly convex. Head continuous with the body, obliquely
truncated. Mouth round, surrounded with a wide circular lip which is
emarginate below. Male generative aperature? communicating with a
hemispherical cavity (acetabulum?) one-fourth the length of the body from
the head. Length 9 lines, breadth 1^2 lines.

Habitat: Fauces Host: Alligator mississippiensis

Locality: Florida Collector: J. W. Bailey

Date: Previous to 1856
Leidy (1890) places this species as Distomum incommodum and as a
synonym of this species his Distoma oricola from the mouth of Alligator
mississippiensis. On the basis of the later determination of Leidy (1890)
his description of Distoma oricola is here included for the purpose of com-
parison and as evidence for the disposition of Monostomum incommodum.

Distoma oricola Leidy 1884

"Body elongated elliptical, moderately wider and thicker posteriorly and
ending in a blunt, angular extremity, convex dorsally and flat ventrally,
unarmed, smooth or minutely wrinkled transversely. Mouth subterminal,
and enclosed with a reniform lip succeeded by a linear annulus. Acetabu-
lum large, globular, included at the anterior fourth of the body, and open-
ing ventrally by a conspicuous central aperature. Generative orifice
ventral, at the posterior fourth of the body. Length, 15 to 20 mm; breadth,
3 mm. Eight specimens obtained from the mouth of the alligator, A.
mississippiensis, in Florida, by Mr. Stuart Wood."

Pratt (1902) surmises that this species is allied to the genus Clinosto-
mum. This view is supported by Ward (1918). Then if, according to this
view the determination of Leidy, that Distoma oricola is a synonym of
Monostomum incommodiim which he determined later (1890) as a Distome,
Distomum incommodum, be accepted, Monostomum incommodum is like-
wise allied to the genus Clinostomum.

285] NORTH AMERICAN MONOSTOMES 67

MONOSTOMUM ORNATUM Leidy 1856

Syn. — Monostomum ornatum Brandes 1892
Monostomum ornatum Braun 1893
Monostomum ornatum Diesing 1858
Monostomum ornatum Monticelli 1892
Monostomum ornatum Stafford 1902

This species was described by Leidy as follows:

"Body slightly compressed ovoidal, anteriorly broad; yellow variegated
with brownish red. Mouth infero-terminal, acetabuliform, transversely
oval. Penis conical, protruding a short distance below the mouth. Female
aperture a short distance below the penis. Length 1 to 1}^ lines, breadth
3^^ to ^ line, thickness }/ito}/2 Hne."

Habitat: Body cavity Host: Rana pipiens

Locality: Philadelphia Collector: H, W. Warren

Stafford (1902) questions the determination of these worms and con-
jectures that they belong either to the genus Haematoloechus sp. which
is commonly found in the lungs of frogs or to Dist. quietum or Cephalogoni-
mue amer. and have been liberated in the first case from the lung and in the
other cases from the small intestine of the host. He inclines to the former
view "on account of the ease with which the small ventral sucker may be
overlooked and the readiness with which worms may be freed from the lungs
without observation." He adds that, "it is unlikely to be Distomum quietum
from the position of the genital openings" and that "it could scarcely have
been Dist. retusum {Ceph. amer.?) since he reports it also in the same paper
although he does not describe it there but in an earlier number."

In regard to the habitat of Monostomum ornatum Stafford says that "of
the hundreds of frogs I have examined I have never yet found a Trematode
free in the body cavity and I doubt if anybody else has ever obtained one
that did not first get there by the accidental cutting or tearing of some
other organ."

The above assumption of Stafford that trematodes do not occur free in
the body cavity of frogs appears to be doubtfully correct. Osborn (1922)
and Cort (1913) report encysted Clinostomum in the body cavity. The
writer in examination of numbers of frogs has been able to observe these
and has several times found some of these worms out of their cysts and free
in the body cavity. These frogs were opened most carefully and worms
that might have been freed by the cutting of the body wall could not
have found their way to remote parts of the cavity in the time consumed
by the operation and examination. On the other hand it is hardly prob-
able that the dozen specimens recorded should escape from the lungs
unnoticed as suspected by Stafford and none be left to show the normal
habitat. However, as Stafford has noted and as was stated above, the worm
could scarcely have been Distomum retusum {Cephalogonimus americanus?)

68 ILLINOIS BIOLOGICAL MONOGRAPHS [286

since it is recorded in the same paper, Stafford, however, has failed to note
that Leidy recorded also in this same paper Distomum variegatum from the
lungs of Rana pipiens which he also describes in the same "earlier number"
in which he describes Distomum retusum. Hence Haematoloechus is ruled
out as well. From the habitat of these worms the writer suspects that it
may possibly be a larval stage of a species of Clinostomum. The descrip-
tion is so meager that it is impossible to place it in any definite manner.

MONOSTOMUM SPATULATUM Leidy 1858

Syn: Monostomum spathulatum Diesing 1859

The following description given by Leidy includes the only available
data. "Body flat, oblong ovate, narrowing anteriorly, obtuse posteriorly,
color white, with brown tortuous lines indicating the course of the oviduct.
Mouth acetabuliform, circular. Testes three, alternating on each side
posteriorly with the oviduct. Ovaries on each side finely lobulated.
Generative aperture small a short distance behind the mouth. Penis
undistinguishable. Length 3-4 lines: breadth }/2 line.

Habitat: Gall-bladder. Host: Fish species unknown

Locality: Eastern U. S. Collector: Jeffries Wyman

MONOSTOMUM AFFINE Leidy 1858
Syn: Notocotyle(?)affi,ne Barker 1916

Leidy described this species as follows: "Body spatulate narrowest
anteriorly, flat; posterior end obtuse, with an excretory orifice communi-
cating with a well marked canal traceable as far forwaid as the commence-
ment of the oviduct. Mouth round, oral acetabulum small, followed by
a smaller pharyngeal bulb. Intestine simple, traceable on each side to
the posterior end of the body. Testes four, posterior to the position of the
distended oviducts. Ovaries finely lobulated, situated on each side exter-
nal to the position of the intestine; oviduct transversely tortuous and dis-
tended with brown ova. Penis ensheathed, long, tortuous, echinate.
Generative aperture small, acetabuliform. Ova oval and prolonged at one
pole, or sub-pyriform. Length of body 6}/^ lines; breadth 1 line."

"Four specimens were obtained by Dr. J. N. Corse from the bile-ducts
and gall-bladder of the muskrat (Fiber zibethicus). Closely allied to M.
hippocrepis Diesing, but has no trace of the horse-shoe like collar to the
head."

As is stated previously in this work the suggestion of Barker appears
to be a misfit on the basis of the pharynx, the echinate penis and the
undivided excretory canal which Leidy says is traceable as far forward as

287] NORTH AMERICAN MONOSTOMES 69

the beginning of the oviduct. These characters are fundamental dififerences
such as are used to separate families and groups and on the basis of these
differences the supposition of Barker (1916) seems untenable.

MONOSTOMUM ASPERUM Vaillant 1863

Syn: Monostomulum aspersum Brandes 1892
Monostomum aspersum Pratt 1902

Vaillant described this species as follows: 1.4 to 1.9 mm long by 0.94
mm wide, elongated, both ends rounded; dorsal surface convex, ventral
flat; cuticula in the anterior % densely covered with small spines regularly
spaced and arranged in alternating rows. Mouth opening round. The
digestive system composed of muscular esophageal bulb, followed by a long
esophagus which extends to the middle of the body where it bifurcates to
form the two ceca which extend to the level of the excretory bladder. The
excretory bladder forms a semi-oval sac, which occupies the posterior end of
the animal. The genital organs are between the excietory bladder and the
ends of the intestinal ceca. The male organs are composed of a curved
spinous penis, with a testicle, a vesicula seminalis and a deferent canal.
The female organs are wanting or in some individuals incompletely devel-
oped and consist only of a granular cell mass.

Found in transparent sub-epidermal cysts of Siren lacertina.

In a later paper the same author (Vaillant, 1863a :347) expresses doubt
in regard to his interpretation of the sex organs which he says appear to
be only poorly developed.

On the basis of the descriptions of Vaillant, Brandes considered this
as a larval form but attempted no further indication for the disposition
of the species. Monticelli (1892) enumerates it without any attempt to
determine its rightful place. Pratt assigns to it the same place as did
Brandes.

Both accounts of this worm are such that the inference may be readily
drawn regarding its larval nature. Neither account gives evidence of any
individual in a sexually mature state. The sex organs are poorly developed
and no mention is made of a uterus or of eggs which would indicate sexual
maturity. The exceptional habitat assigned to this worm by Vaillant which
he says is comparable to that of Monostomum faba Bremser is not so rare
in the light of the present knowledge of the life histories of distomes and
is quite in contrast to what is now known of the m^onostome life history.
Many distomes are known to pass a portion of their life cycle encysted in
the skin and superficial layers of the body, while no such stage has yet been
found in the cycle of a monostome. The description of Vaillant as well as
his figures are so inadequate that it w^ould be hazardous to speculate on the
systematic position of this form more than to say on the basis of the digestive
system the cuticular spines and what has been given regarding the genital
glands that it is probably the immature stage of a distome.

70 ILLINOIS BIOLOGICAL MONOGRAPHS [288

MONOSTOMUM AMIURI Stafford 1900

This trematode taken from the swim bladder of Amiurus nebulosus by
Stafford, is described by him as follows: "About 5 mm long and 2.25
mm broad much flattened and is broadest in its posterior two-thirds, the
anterior third narrowing towards the mouth-sucker. The living animals
are very soft bodied, inactive creatures."

"The integument bears a cuticle and is apparently very thin the sub-
cuticular and glandular parts seem to have a similar structure to the same
parts of the Distomes with which I am most acquainted. The intestinal
system begins with the mouth whose thick muscular walls form the oral
sucker. Following this is a muscular pharynx and a narrow esophagus
which gives rise to two lateral intestinal ceca, extending as broad tubes to
near the posterior end of the body where they end blindly."

"The posterior excretory bladder unpaired."

"The reproductive system is of the usual complex type. Each individ-
ual dioecious. The testes are situated most posterior in the body, between
the median expulsion tube of the excretory system and the ends of the
intestinal ceca, the vasa efferentia rise out of their anterior ends and
proceed, by a direct course, to near the middle of the animal where they
meet in the vesicula seminalis. This runs forward and opens by a muscular
penis on the ventral surface of the worm, about one-third from its anterior
end. The ovary is located a little behind the middle of the animal. The
uterus filled with eggs occupies most of the posterior two-thirds of the body
and opens by a bulbous vagina immediately behind and to the right of the
penis. The two lobular yolk glands lie outside of the forked intestine and
extend from the level of the genital openings to the hind end of the animal.
A longitudinal yolk duct receives the yolk cells from the numerous follicles
on each side and conducts them, by a transverse tube in the region of the
ovary, to a yolk reservoir that communicates with the oviduct close by
the shell gland. The egg is 45 by 24ju in size and its blunt broader end is
provided with a short hooked filament."

The description and figure of Stafford shows a remarkable similarity
to the Heterophyidae with which the worm agrees in having simple sac-
like intestinal ceca extending to the posterior end, genital pore in immediate
neighborhood of the ventral sucker if that organ is what has been inter-
preted as the female genital opening by Stafford. Testes oval, symmetri-
cally arranged near the posterior end, seminal vesicle S-shaped, ovary oval,
median, anterior to the testes. Vitellaria lateral to the intestinal ceca;
extending from the level of the testes. The eggs also fall within the size
found in this family but differ in having a short hooked filament at the
blunt end. It differs also in that scales have not been recorded for this
species by Stafford.

289] NORTH AMERICAN MONOSTOMES 71

In the above comparison the bulbous vagina of Stafford has been con-
strued to be the ventral sucker and considering this as a sucker this species
agrees in every important anatomical feature with the Heterophyidae. The
absence of scales here if they have not actually been overlooked could be
considered only of specific import and according to the findings of Ward
and Hirsh (1915:148) "spines and scales are caducous in life and are easily
lost also if the specimens lie in preserving fluid for some time.
^ The fact that this species was found in the swim bladder of Amiurus
nebulosus is not sufficient reason for its separation from the Heterophyidae
since three genera of that family are known to have larval stages in fish,
namely, Metagonimus, Cryptocotyle, and Paracoenogonimus. Ransom
(1920:530) surmises that "Heterophyes," the genus to -which. Mono stomu7n
amiuri seems to be most closely related, "occur in their immature stages in
fish," the adults thus far having been found in fish eating birds and mam-
mals.

Habitat: Swim bladder Host: Amiurus nebulosus

Locality: probably near Toronto, Collector: Stafford
Canada

■ i&[q biiii (i.O<^I,i A■Jl^cOi'6

irij no aaioaqa airlT .8n'->''+'^^'
muW-

72 ILLINOIS BIOLOGICAL MONOGRAPHS [290

THE POLYPHYLETIC ORIGIN OF THE MONOSTOMES

From the earliest records of the Monostomata up to the present time
this group of parasites has served for a dumping ground for inaccurately
studied species in which the acetabulum has been wrongly interpreted or
overlooked entirely. Many species have since been studied more carefully
and consequently have been transferred to other genera. Out of this has
arisen the problem of the origin of the Monostomata. Accumulative
evidence has lead to the belief that these forms are directly related to var-
ious other groups. This evidence is presented below.

Certain investigators of recent time have come to consider the trema-
todes of polyphyletic origin. According to Faust (1918) these conclusions
are the result of "lack of study and consequent inability to recognize the
fundamental resemblance of the genital, excretory and nervous systems."

The first to suggest relationship between the Monostomata and the
Distomata was Monticelli (1893:149-150) when he called attention to the
similarity of Kollikeria and Didymozoon. More recently Ariola (1906)
reinforced this opinion by grouping Monostonia fillicolle Rud. and Distoma
okeni Kolliker together on the basis of their anatomical similarity even
though Monostonia fillicolle does not possess an acetabulum. MacCallum
and MacCallum (1916) on the basis of anatomical similarity grouped
together the two genera Kollikeria and Nematobothrium altho Kollikeria
shows in many cases well developed acetabula while Nematobothrium is
in that respect typically monostomatous.

Cohn (1904) in his study of Monostomum flavum Mehlis, worked over
by Stossich (1902) and placed in the new genus Typhlocoelum, found a
well developed but small ventral acetabulum which he figures in sagittal
sections. This species on the one hand is apparently very closely related
to the genus Cyclocoelum and was placed by Stossich in the same sub-
family, Cyclocoelinae. On the other hand Cohn would transfer this to
the Fasciolidae because of the presence of the ventral acetabulum which
he says is diminished and in other instances often lost because of the shut-in
habitat under which these worms live.

He adds as was stated previously the observation of a rudimentary
mouth sucker in Cyclocoelum mutabile and in one other species of this
group. Here he states that Cyclocoelum mutabile does not lack a primary
sucker in many cases and like the Cestodarian Amphilina has lost hold-fast
organs because of the lack of need for such organs in the cavities of the
body of the host in which habitat these worms are wont to live. According

291] NORTH AMERICAN MONOSTOMES 73

to this author Cyclocoelum mutahile and Typhlocoelum flavum are very
closely i ".ted and because of different stimuli in their respective habitats,
viz. : ?.\ veolar spaces, abdominal cavity and liver, occasionally the intestine
for Cyclocoelum, and trachea and bronchi for Typhlocoelum, the acetabu-
lum of Cyclocoelum has been lost while the oral sucker of Typhlocoelum
has atrophied.

In his earlier work (1902) Cohn described Monostomum oculobiiim
collected from Vanellus melanogastrus as having neither oral nor ventral
suckers and relates it to Cyclocoelum mutabile. Fuhrmann (1904) describes
a species, Bothriogaster variolaris collected from the intestine of Rostrhamus
sociabilis, a South American Falconid, which from his figures and descrip-
tion appears to be very similar to Monostomum oculohium of Cohn. He
states that in regard to the intestinal crura, absence of the oral sucker
and the presence of only a pharynx it is like Cyclocoelum mutabile. The
position of the genital glands is not the same but on the other hand is like
those of Monostomum oculobium of Cohn. But it differs from Monostomum
oculobium in that a ventral sucker is present which Fuhrmann believes
Cohn had overlooked in his species. Fuhrmann would place Bothriogaster
variolaris in the sub-family Syncoelinae of the Fasciolidae.

Odhner (1907) supports the view of Cohn and cites a number of in-
stances to demonstrate it. Chief among these are, first, his genus Apora-
cotyle which is a blood parasite. He says that this suckerless form has its
nearest relative in the distome Hapalotrema constrictum (Leared), a blood
parasite of the sea turtle. Second, those inhabiting the airsacs have
developed the hold-fast organs to the least degree. This includes the
Cyclocoelidae in which Odhner says all species are without a ventral
sucker except for the recent discovery by Cohn of an entirely rudimentary
acetabulum in Typhlocoelum flavum. He adds that in the Holostomes
and Hemistomes ventral suckers have been greatly reduced and in some
instances have disappeared entirely. In further support of this view he
cites the reduction of the sucker in the male Bilharzia with the complete
loss of this organ in Bilharzia kovalewski, and notes also the Echinostome
like genus Pegosomum which inhabits the gall duct and has lost entirely
the oral sucker. Odhner believes that the Monostomata will be finally
split up and appended to other trematode groups, i.e., to the Distomes,
Amphistomes, Holostomes and perhaps others.

In the light of the foregoing one may well ask, what is a Monostome?
The question has been aptly raised as to what characters or combinations
of characters afford a reliable and accurate basis for the natural classifica-
tion of the trematodes. On the basis of the examples given above the
presence or absence of an acetabulum can be considered of relatively
little importance and if such be the case other factors must be looked for
in an attempt to establish a natural s^'stem of classification.

74 ILLINOIS BIOLOGICAL MONOGRAPHS [292

In view of the facts stated above the writer believes that an accurate
knowledge of the more fundamental systems of organs will reveal group
relationships which have heretofore passed unnoticed. In the present
systematic groups homology of organs is a factor which has been generally
passed over. This can best be demonstrated by a careful study of the
developing organs in the early stages of the life history of the individual.
Up to this time little has been done on the life histories of the Monosto-
mata. Von Siebold (1835) and Van Beneden (1861) studied the early
stages of the parthenogenetic phase. La Valette St. George (1855) de-
scribed under the name of Monostomum flavum a cercaria which he believed
to belong to this species. From his descriptions and figures one can see
diagnostic points which tend to show that this cercaria belongs to the
Notocotylidae on the basis of the well developed oral sucker, the posses-
sion of the locomoter pockets described by Looss, Cort, and Faust for
Notocotylid species. The fact that La Valette St. George showed the
intestine anastomosed posteriorly does not furnish evidence to the contrary
since the crura of the Notocotylid approach each other in the posterior
end of the worm and this may easily be mistaken for anastomosis or on the
other hand the author may have misinterpreted the excretory ducts for
the crura of the intestine. The characteristic features are the absence of
the pharynx and the presence of the three eyespots which appear to be
characteristic of the Notocotylidae. In addition to these Haldemann,
Leidy, Cort and Faust have described some six or seven monostome
cercaria from American hosts all of which have been referred to the Noto-
cotylidae. While the writer has had opportunity to study immiature
stages of N otocotylus urbanensis, the preserved material has yielded only
few facts that can be interpreted as of phylogenetic importance. These
will be discussed in a subsequent section of this paper. As yet no Mono-
stome life history has been demonstrated experimentally and the develop-
ment of the important systems of organs has not been followed in the life
history of even a single species. This seems to the writer to be a necessary
step to be followed out in the major families in order to demonstrate the
phylogenetic relationships of the large groups.

INTERRELATIONSHIP OF THE MONOSTOME FAMILIES

Before entering into the discussion of the probable origin of the Mono-
stomes it seems fitting to discuss the interrelationship of the families as a
unit or natural group of Trematodes. A comparison of the family diagno-
ses, given earlier in this paper, shows a striking contrast in each of the
families discussed and of the families not included the same striking con-
trast may be drawn. No system of organs is the same in all of the families
save perhaps the nervous system, so far as it has been made out, which is
essentially the same in all the trematodes. The excretory system differs

293] NORTH AMERICAN MONOSTOMES 75

in these families; in the Cyclocoelidae it is composed primarily of a large
half-moon shaped bladder and ramifying network of anastomosed tubules.
In the Notocotylidae a single club-shaped bladder extends almost to the
ovary and sometimes a little posterior to the excretory pore {Notocotylus
quinqueserialis) ; at the anterior end of this bladder two branches are given
off which pass lateral to the ovary and from these numerous side branches
are produced. In the Heronimidae the bladder consists of a large median
sac with the pore in the anterior dorsal region, while in the Collyrichdae
the bladder is single, club-shaped, and branches near the center of the
body. In regard to the digestive system equally striking differences appear.
The Notocotylidae show no trace of a pharynx while the Cyclocoelidae,
Heronimidae and the Collyrichdae possess both sucker and pharynx. The
crura of the Cyclocoelidae anastomose in the posterior end of the body
while in the other families under consideration the crura end blindly.
Another characteristic difference is the presence of dermal glands in some
of the Notocotylidae, while such organs are unknown in the other families.
In the genital system are to be found equally great characteristic differ-
ences in the position of the glands with respect to the intestinal crura as
well as in the form of the glands themselves. These and other important
differences seem to indicate that these families have arisen from different
lines.

THE APPARENT RELATION OF THESE FAMILIES TO OTHER GROUPS

Since the great diversity of structure in this group seems to indicate
that the monostomes have arisen from different sources, there remains to be
considered in conjunction with this fact the close affinity of certain of these
famihes to widely separated groups.

The finding of a rudimentary acetabulum in Monostomum flaviim
Mehlis by Cohn and of the well developed acetabulum in Bothriogaster
variolaris by Fuhrmann in the same year together with the similarity in
structure seems to indicate a relationship to the Fasciolidae through the
Syncoelinae according to Fuhrmann (1904) to which Bothriogaster is
most closely related.

After a similar manner the Notocotylidae find their closest parallels
among marine forms where the genus Notocotylus has anatomically a very
close relative in Adenogaster serialis while those Notocotylids without the
ventral glands appear more like the genus Glyphicephalus.

The Heronimidae stand alone in their organization and do not show
close relationship to any known trematodes.

Attention was first called to the distome character of Collyriclum faba
by Braun (1892) and again by Kossack (1911:577) who points out the
relation of this species to Distomum gastrophilum but hesitates to decide
whether it is a natural one, thus: "Indessen wage ich es vorlaufig noch

76 . ILLINOIS BIOLOGICAL MONOGRAPHS [294

nicht zu entscheiden, ob hier natiirliche Verwandtschaft oder nur eine
Konvergenzerscheinung vorliegt."

More recently Odhner (1914) placed this genus in his new family Trog-
lotremidae which he characterizes as follows: Mehr oder weniger abge-
plattete "Distomen" oder "Monostomen" von gedrungener Korperform
und 2-13 mm Lange. Das ausserste Hinterende als einkleiner "Schwanz-
anhang" vorstreckbar. Bauchflache flach oder etwas ausgehohlt, Riicken-
flache gewolbt. Haut iiber und iiber mit spitzen Stacheln bewaffnet.
Muskulatur bei den Cystenbewohnern schwach entwickelt, auch in den
Saugnapfen. Darmapparat mit Pharynx, nicht allzu langem Oesophagus
and Darmschenkeln, die ein mehr oder weniger kurzes Stuck vor dem
Hinterende endigen. Exkretionsblase Y-formig oder einfach schlauchfor-
mig. Genitalporus dicht am Vorder- oder Hinterrand des ev. Bauch-
saugnapfes, median oder leicht lenkseitig. Cirrusbeutel meist fehlend,
[except Troglotrema] Pars prostatica und Samenblase immer unter-
scheidbar. Hoden symmetrisch, in oder hinter der Korpermitte, langs
gestellt. Ovar unmittelbar vor den Hoden, rechtseitig, [Except Paragoni-
mus westermanni in which amphytypy occurs] meist stark gelappt [except
Troglotrema]. Receptaculum seminis und Laurerscher Kanal vorhanden.
Dotterstocke meistens sehr stark entwickelt [except Collyriclum] und
dabei ausschliesslich oder hauptsachlich unter der Riickenflachs ausge-
breitet, nur einen medianen Streifen frei lassend. Uterus bald sehr lang
und stark hin und her gewunden, bald relativ kiirzer und mehr aufge-
knauelt. Eier im ersteren Falle klein, 0.017-0.025 mm lang, im letzteren
bedeutend grosser, von 0.063-0.085, nach einigen Angaben sogar bis 0. 12
(?) mm Lange. — Parasiten von carnivoren Saugetieren oder von Vogeln,
meistens paarweise in cystenahnlichen Hohlungen."

This family appears to be an unnatural grouping. Of the four genera
included three of them, Troglotrema, Paragonimus and Collyriclum,
show radical divergence from the family diagnosis as evidenced by the
fact that exceptions must be made in order to include them. In an appen-
dix the author adds to this new family Renicola pinguis which he says is
the closest relative of Collyriclum fab a.

Jegen (1917) cites the close relationship of Collyriclum to Brandesia
turgida. This author emends the family diagnosis of Odhner for the new
family Troglotremidae as follows in order to include these two genera,
Renicola and Brandesia. "Die Haut entweder mit spitzen Stacheln oder
mit Schuppen durchsetzt. Bauchsaugnapf entweder unmittelbar vor oder
hinter der Korpermitte."

The present investigations do not show the same close relationship
for Collyriclum as found by Odhner and Jegen in the family Troglotremi-
dae while it shows a distinct relationship to Brandesia. This genus is
strikingly different from other genera in Odhner's family Troglotremidae.

295]

NORTH AMERICAN MONOSTOMES

77

Table II. Probable Relationships of Coll^Ticlidae

Brachycoelinae

Collyrididae

Pleura genetinae

Shape

Egg-shaped or spheri-
cal

Hemispherical

Small to medium
slightly elongated

Integument

Naked rarely spinous

Spinous-in rows

Spiny or scaled

Oral sucker

Well developed

Terminal small

Weakly developed

Acetabulum

Well developed

Wanting

Weakly developed

Pharynx

Small

Smaller than sucker

Poorly developed

Crura

Thin, small

Thin, voluminous

Never reaching pos-
terior end

Excretory

V-shaped

Large Y-shaped

V or Y-shaped

Genital pore

Ventral between suck-
ers

Center of ventral sur-
face

Between suckers, lat-
eral

Copulatory organs

Present or absent

Present, moderately
developed

Well developed

Testes

Symmetrical oval

Symmetrical, irregu-
larly lobed

Symmetrical oval

Ovary

Lateral, near acetabu-
lum

Lateral, anterior, deep-
ly lobed

Lateral, near or an-
terior to acetabulum

Receptaculum
seminis

Present

Wanting (?)

Present

Laurer's canal

Present

Present

Present

Vitellaria

Single arborescent
group

Several arborescent
groups on each side

Simple arborescent
group anterior
to crura

Uterus

Usually posterior to
testes

In general in posterior
body half

Usually in posterior
body half

Eggs

Numerous about 22ju
long

Numerous 19 to 21)u
long

29 to 34m long
13 tol6M broad

Host

Mammals, birds and
reptiles

Birds, parasitic in pairs
in cysts

Amphibia and rep-
tiles. Renicola and
Loxogenes in pairs

78 ILLINOIS BIOLOGICAL MONOGRAPHS [296

The position of the genital pore as well as the organology depicts it as
distinct from Collyriclum. This study confirms the systematic position of
Brandesia given by Looss in that it is more closely related to the Pleu-
rogenetinae. Collyriclum is in some respects closely related to the Pleuro-
genetinae and in other respects to the Brachycoelinae. The diagnostic
characters of Collyriclum are so evenly distributed between these two sub-
families that it holds an intermediate place as shown in the table opposite.

Whether the interpretation given above is the correct one remains
indeed a matter of conjecture and certainly lacks much of confirmation.
It is difficult to ascertain from an anatomical study of adult forms as
to whether acetabula are vestigial. The most immature Cyclocoelidae
studied by the writer show no trace of such organs. These forms belong
to Cyclocoeluni ohscurum. In more advanced stages of Cyclocoelum halli
the sucker is found practically as well developed as in the adult. Since
the material of the immature stage of Cyclocoelum ohscurum was not well
preserved conclusions cannot be drawn from it. In the immature stages
of N otocotylus urhanensis studied by the writer the ventral glands were
found to develop after encystment of the cercariae and to show no rela-
tion in their development to an acetabulum. Much evidence on this
point can yet be obtained by the elucidation of this and other life histories.

It is worthy of note, however, that in the Notocotylidae the oral
sucker is well developed in the cercariae as well as in the adult, but instead
of a single well developed acetabulum the condition is somewhat varied.
In N otocotylus quinqueser talis five rows of small sucking discs are pro-
vided. In Notocotylus attenuatum three rows of similar organs are present,
while in Nudocotyle and Paramonostomum no such sucking organs are
present. These species Hve in a similar habitat (intestine of the muskrat)
and under this condition have developed in the first instance different
numbers of these organs while in the latter case no such structures have
been observed. Nudocotyle novicia, however, presents other striking differ-
ences which need not be considered here. In the cercariae of these forms
described notably by Cort (1914) and Faust (1918) no such organs are
found nor is there any indication of their early development, while in the
immature forms of N otocotylus urhanensis studied by the writer these glands
are found well along in development soon after being freed from the cyst.
The other organs of these cercariae correspond so well to the adult struc-
ture that there is little doubt as to the identity of the form. In this case
then the sucking discs are developed after the organism enters the defini-
tive host. A final decision of this question, however, must await further
evidence and experimental demonstration of the life history.

The cases of reduction of sucking musculature cited by Cohn (1904)
and Odhner (1907, 1911) lead again to the question raised previously and
in the light of the theory of Cohn and Odhner it is difficult to determine

297] NORTH AMERICAN MONOSTOMES 79

if this reduction is of phylogenetic import or subject to rapid change under
environmental stimuli. A comparison of similar worms under different
conditions leads to the belief that the sucking musculature is not subject
to such rapid and radical change as indicated by Odhner (1907). As shown
in an earlier part of this paper, Cyclocoelum wilsoni and Cyclocoelum
vicarium from the intestine of their host have poorly developed oral suckers
and no trace of an acetabulum while Cohn has found a rudimentary aceta-
bulum in Monostomum flavum, a species which inhabits the trachea.

In the Heronimidae which inhabit the lungs and especially the larger
bronchi of turtles, the oral acetabulum is well developed while any evi-
dence of the presence of a ventral sucker has not been obtained. This
stands in rather striking contrast to the statement of Odhner (1907)
referred to above. Fuhrmann (1904) cites an interesting case in his Bothrio-
ga.ster variolaris which was collected from the intestine of Rostrhamus
sociahilis. This species according to the author has a ventral acetabulum
but no oral sucker. A pharynx, however, is present. As stated pre-
viously his species is strikingly similar to Monostomum oculobium Cohn
taken from Vancllus mclanogastrus which Cohn (1902) says is devoid of
sucking apparatus. Why has the oral sucker of Bothriogaster variolaris
atrophied if reduction is due to habitat as stated by Cohn (1902) and
Odhner (1907)?

Another interesting correlation is found among the Notocotylidae and
the PronocephaUdae both of which inhabit the intestine. Among the
Notocotylidae are forms which late in life develop the ventral holdfast or
adhesive glands and those without such glands, both types of which have
been found in the alimentary tract of the muskrat. Linton (1910) reports
a species Barisomum eruhescens from the intestine of three tropical fish
which show no sign of any sucking organ save the oral. Of the several
genera of the Pronocephalidae taken from the intestine of Chelone midas
only Adenogaster shows any trace of accessory adhesive apparatus. On
the other hand Odhner (1911) found reason to ally the Angiodictyidae a
closely related family to the distomes on account of a weakly developed
acetabulum found in Haplorchis carhinus Looss.

As was stated previously Odhner (1907) proved beyond doubt the
presence of both pharynx and oral sucker in Didymozoon scombri Tschbg.
but found there no ventral sucker. Four years later Ariola (1906) asserts
the Didymozoon nature of Kollikeria (Distoma) okejii (Kolliker) which
he states is synonymous with Monostomum fillicolle Rud. In the Parona
material reported as Kollikeria in which there were many distomes accord-
ing to this author, he finds a single form without the acetabulum.

On the other hand Collyriclum faha (Bremser) and Collyriclum colei
Ward, other cyst living trematodes, have no acetabula but present, however,
a well developed oral sucker. It is difficult to interpret how worms which

80 ILLINOIS BIOLOGICAL MONOGRAPHS [298

live under such similar conditions as the Didymozoonidae and the Colly-
riclidae could by the effect of habitat alone become so totally different.

That the habitat in which the worm lives is a factor in the modification
cannot be disputed but is certainly only one factor which contributes to
the modification. What makes up the aggregate of factors in a given case
could be only a matter of conjecture. Recent experimental investigation,
however, has shown that factors influencing modifications of an individual
or race may be both external and internal and that the change produced
may be gradual or occur as a mutation in which case little or no trace may
be left to depict the ancestral route. While the writer is inclined to look
upon the loss of sucking musculature as a gradual change brought about
by a group of factors among which the habitat seems to have played an
important part, there is every reason to believe that sudden radical re-
arrangements have taken place and these are no doubt responsible for many
of the variant forms. While there is good reason to believe that the Mono-
stomata have arisen from highly divergent groups the evidence seems as
yet about equally divided and the final decision must necessarily rest on
further studies of the anatomy and life history of members of this and
other groups.

The systematic arrangement used in the earlier part of this paper is
essentially that employed by Liihe (1909), Kossack (1911) and Ward
(1918), which for the above stated reason it seemed best to preserve at
least for the present. It is hoped that interest in this aberrant group will
increase and that careful studies will result either in the preservation of
these forms as a natural taxonomic group or in their separation and subse-
quent distribution among other well organized units to which they are truly
related.

299] NORTH AMERICAN MONOSTOMES 81

SUMMARY

I. Monostomes from North American land and fresh water hosts have

been studied.
II. Nine new species have been described in detail,

III. The knowledge of their anatomy and life history is greatly increased.

1. Both oral sucker and pharynx are present in the Cyclocoelidae.

2. In the genera of Cyclocoelidae studied the receptaculum seminis

is a constant feature while Laurer's canal is not present.

3. The origin of the ventral glands in Notocotylus urbanensis is

clearly shown in early stages of development.

IV. The discovery of well developed miracidia in Collyridum colei renders

improbable Jegen's account of the life history of Collyridum fab a.
V. A careful study of the anatomy of the Cyclocoelidae, the Notocotyli-
dae, the Collyriclidae, and the Heronimidae demonstrates clearly
that they are not immediately related.

1. The Cyclocoelidae are probably allied to the Fasciolidae.

2. The Notocotylidae are closely related to the Pronocephalidae

not merely in general organology but especially in the presence
of peculiar ventral glands.

3. A study of the organology of the Collyriclidae demonstrates

clearly their direct relationship to the Pleurogenetinae.

4. The Heronimidae do not show immediate relationship to any

known trematodes.
VI. The monostomes are a heterogeneous group and must ultimately be
distributed among other groups in accordance with their funda-
mental relationships.

82 ILLINOIS BIOLOGICAL MONOGRAPHS [300

BIBLIOGRAPHY

Ariola, V.

1906. Monostoma filicolle Rud. e Distoma okeni Kolliker. Zool. Anz., 30:185-186.
Arnsdorff, a.

1908. Monostomum vicarium. Centralbl. Bakt. Par., Abt. I, Orig., 47:362-366; 2 fig.
Baird, W.

1853. Catalogue of the species of entozoa or intestinal worms contained in the collec-
tion of the British Museum. London; 2 pi.
Baeker, F. D.

1915. Parasites of the American Muskrat (Fiber zibethicus). Jour. Par., 1:184-197;
2 pi., 4 figs.

1916. A new Monostome Trematode Parasitic in the Muskrat with a Key to the
Parasites of the American Muskrat. Trans. Amer. Mic. Soc, 35 :175-184; 1 pi.

Barker, F. D. and Laughlin, J. W.

1911. A New Species of Trematode from the Muskrat, Fiber zibethicus. Trans. Amer.
Mic. Soc, 30:261-274; 1 pi.
Barker, F. D. and Parsons, Susanne.

1914. A new species of Monostome from the Painted Terrapin, Chrysemys marginata.
Zool. Anz., 45:193-194.

1917. A Monostome Lung Fluke from the Painted Terrapin, Chrysemys marginata Agas-
siz. Trans. Amer. Mic. Soc, 36:55-66; 2 pi.

Van Beneden, P. J.

1861. M6moire sur les vers intestinaux. Suppl. C. r. Acad. Sci., 2:1-376.
Blanchard, R.

1903. Experiences et observations sur la marmotte en hibernation. VI. Observations
sur les parasites en general. C. r. Soc. biol., 55:1124-26.
Bloch, M. E.

1782. Abhandlung von der Erzeugung der Eingeweidewiirmer. Berlin. 54 pp.; 10 pi.
Blochmann, F.

1896. Die Epithelialfrage bei Cestoden und Trematoden. Hamburg. 12 pp.; 2 pL,
Ifig.
Brandes, G.

1891. Zum feineren Bau der Trematoden. Zeit. wiss. Zool., 53:558-577; 1 pi.

1892. Revision der Monostomiden. Centrlbl. Bakt. Par., 12:504-511.
Braun, M.

1889-1893. Trematodes, in Bronn's Klass. Ordn. d. Thier-Reichs. 4, Abt. la:306-925,

1893. II. Bericht uber thierische Parasiten. Centrlbl. Bakt. Par., 13:59-68.

1893a. Die Wohnsitze der endoparasitischen Trematoden. Centrlbl. Bakt. Par., 13:

465-468.
1901. Trematoden der Chelonier. Mitt. zool. Mus. Berlin, 2, 58 pp.; 2 pi., 2 fig.
1901a. Zur Kenntniss der Trematoden der Saugethiere. Zool. Jahrb., Syst., 14:311-

348; 2 pi.
Bremser, M.

1831. Schmalz, XIX Tabulae Anatomian entozoorum illustrantes. Dresdae et P

Lipsiae; 11-16. (Not available.)
COBBOLD, T. S.

1860. Synopsis of the Distomidae. Jour. Proc. Linn. Soc. Lond. Zool., 5:1-56.

301] NORTH AMERICAN MONOSTOMES 83

1864. Entozoa; An introduction to the study of Helminthology. London. 480 pp.;
21 pi., 82 text fig.
COHN, L.

1902. Mittheilungen uber Trematoden. Zool. Anz., 25:712-718; 9 fig.
1904. Helminthologische Mitteilungen II, Arch. Naturg., 1:229-251.
Cole, L. J.

1911. A Trematode Parasite of the English Sparrow in the U. S. Bull. Wise. Nat. Hist.
Soc, 9:42-48; 2 pi.
CORT, W. W.

1913. Notes on the Trematode Genus Clinostomum. Trans. Amer. Mic. Soc, 32:
169-182; 1 pi.

1914. Larval Trematodes from North American Fresh- Water Snails. Jour. Par.,
1:65-84; 15 figs.

1915. Some North American Larval Trematodes. 111. Biol. Monogr., 1 :446-532; 8 pi.
Creplin, F. C. H,

1829. Filariae et Monostomi speciem novam in Balaena rosirata repertam. Nova
acta Phys.-med. Acad. Nat. curios., Bonnae, 14:871-882; 1 pi.
DiESING, K. M.

1839. Neue Gattimgen von Bmnenwiirmem. Ann. Wien. Mus. Naturg., 2:219-242;

7 pi.
1850. Systema Helminthum. Vienna; 1, 679 pp.

1858. Revision der Myzhelminthen: Trematoden. Sitz. Akad. Wiss. Wien, math.-
naturw. CI, 32:207-390; 2 pi..

1859. Nachtrage u. Verbesserungen zur Revision der Myzhelminthen. Sitz. Akad.
Wiss. Wien, math-naturw. Cl., 35:421^51.

DUJARDIN, F.

1845. Histoire natureUe des helminthes ou vers intestinaux. Paris. 654 pp.
Faust, E. C.

1918. Life History Studies on Montana Trematodes. HI. Biol. Monogr., 4:1-120; 9 pi.

FiLIPPI, F. DE

1856. Quelques nouvelles observations sur les larves des trematodes. Ann. sci. nat.,

Zool., (4) 6:83-86. [Translation, Ann. Mag. Nat. Hist., (2) 20:129-132.]
1859. Troisi6me memoire pour servir k I'histoire gen6tique des trematodes. Mem.
Accad. sci. Torino, (2) 18:201-232; 3 pl.
FrQuch, J. A. VON.

1789. Beschreibung einiger neuen Eingeweidewiirmer. Naturforscher., Halle, 24:101-
162; 4 pl.

FUHRMANN, O.

1904. Neue Trematoden. Centrlbl. Bakt. Par., Abt. I, Orig., 37:58-64; 4 fig.
Gmelin, J. F.

1790. Caroli a Linn6 Systema Naturae, Vermes. 13 ed., 1 :3021-3909.
Goeze, J. a. E.

1782. Versuch einer Naturgeschichte der Eingeweidewurmer thierischer Korper.
Blankenburg. 47 1 pp. ; 44 pl.
Harrah, E. C.

1921. Two new Monostomes from Asia. Jour. Par., 7:162-165; 2 fig.
HOYXE, W. E.

1888. Trematoda. Encycl. Brit., N. Y. 9 Ed., 23:535-540.
Jagerskiold, L. a.

1891. tlber den Bau des Ogmogasler plicatus (Creplin) {Monoslomum plicatum Creplin)
K. Svensk. Vetensk.-Acad. Handl., Stockhohn, 24:1-32; 2 pl.

84 ILLINOIS BIOLOGICAL MONOGRAPHS [302

Jegen, G.

1917. Collyriclum faba (Bremser) Kossack. Ein Parasit der Singvogel, sein Bau und
seine Lebensgeschichte. Zeit. wiss. Zool., 117:460-553; 2 pi.

Johnston, S. J.

1916. On the Trematodes of Australian Birds. Jour. Proc. Roy. Soc. N. S. W., 50:
187-261; 11 pi.

Kossack, W.

1910. Neue Distomen. Centrlbl. Bakt. Par., Abt. I, Orig., 56:114-120; 4 fig.

1911. tJber Monostomiden, Zool. Jahrb., Syst., 31 :493-590; 3 pi.

Lahille, F.

1918. Nota sobre Monostoma mutabile y la clasificacion general de los Trematodes.
Buenos Aires. Physis: Rev. Soc. Arg. Ci. Nat., 4:328-331.

Lang, A.

1880. Untersuchungen zur vergleichenden Anatomic und Histologic des Nervensystems
der Platyhelminthen. II. tJber das Ncrvcnsystem der Trematodcn. Mitt.
Zool. Stat. Ncapel, 2:28-52; 3 pi.

La Valette St. George, A. J. H. von

1855. Symbolae ad trcmatodum evolutionis historiam. Berolini, 39 pp.; 2 pi.

Leidy, Jos.

1856. A S3Tiopsis of Entozoa ai^d some of their Ectocongeners observed by the author.
Proc. Acad. Nat. Sci. Phila., 8:42-58.

1858. Contributions to Helmintholog>'. Proc. Acad. Nat. Sci. Phila., 10:110-112.

1877. On Flukes infesting Molluscs. Proc. Acad. Nat. Sci. Phila., 29:200-202.

1884. Distoma and Filariae. Proc. Acad. Nat. Sci. Phila., 36:47^8.

1885. On some parasitic worms of birds. Proc. Acad. Nat. Sci. Phila., 37:9-11.
1887. Notice of some parasitic worms. Proc. Acad. Nat. Sci. PhQa., 39:20-24.

1890. Notices of Entozoa. Proc. Acad. Nat. Sci. Phila., 42:410-418.

1904. Researches in helminthology and parasitology. Smithson. Misc. Coll., 46:1-281.

[Leidy, 1904, is an exact reprint of all his original papers.]
Linton, Edward.

1910. Helminth Fauna of the Dry Tortugas. II. Trematodes. Carnegie Inst. Pub.,

133:11-98; 28 pi.

LONNBERG, E.

1891. Mitteilungen iiber einige Helminthen aus dem zool. Museum der Universitat
zu Kristiania. Verb. biol. Ver. Stockholm, 3:64-78; 1 pi.

Looss, A.

1899. Weitere Beitrage zur Kenntniss der Trematoden-Fauna Aegyptens, zugleich
Versuch einer natiirlichen Gliedcrung des Genus Distomum Retzius. Zool.
Jahrb., Syst., 12:521-784; 9 pi.

1901. Natura doceri, cine Erklarung und Begriindung einiger Grundsatze, wclchc mich
bei meinem "Versuch einer natiirlichen Gleiderung des Genus Distomum Retzius"
geleitet. Centrlbl. Bakt. Par., Abt. I, 29:191-210.

1902. tJber neue und bekanntc Trematoden aus Seescluldkroten. Zool. Jahrb., Syst.,
16:411-894; 12 pi.

LtJHE, M.

1900. [Review of] Looss, A., Weitere Beitrage etc. Centrlbl. Bakt. Par., Abt. 1,
28:458-466.

1901. iihei Monoslomum orbiculare. Centrlbl. Bakt. Par., Abt. 1, 29:49-60; 5 fig.
1901a. Zwei neue Distomen aus indischcn Anuren. Centrlbl. Bakt. Par., Abt. 1,

30:166-177; 5 fig.

303] NORTH AMERICAN MONOSTOMES 85

1909. Parasitische Plattwiirmer. Brauer, Die Susswasserfauna Deutschlands, 17,
217 pp.; 188 fig.
MacCallum G. a. and MacCallum, W. G.

1916. The family Koellikeriadae (Didymozoidae) Mont. Zool. Jahrb., Syst., 39:141-
168; 3 pi.
MacCallum, W. G.

1902. Heronimus chelydrae, nov. gen., nov. sp. A new Monostome parasite of the
American snapping turtle. Centrlbl. Bakt. Par., Abt. I, Orig., 32:632-636; 2 fig.
MONTICELLI, F. S.

1888. Saggio di una morfologia dei trematodi. Tesi per ottenere la privata docenza
in zoologia nella R. Universita di Napoli. 130 pp.

1892. Studii sui trematodi endoparassiti. Dei Monostomum del Box salpa. Atti.
Accad. sci. Torino, 27:514-534; 1 pi.

1892a. Studii sui trematodi endoparassiti. Monoslomum cymbium Diesing. Mem.

Accad. sci. Torino, 42:683-727; 1 pi.
1892b. Studii sui trematodi endoparassiti; sui genere Notocotyle Diesing. Boll. Soc. nat.

Napoli, 6:28-46.

1893. Studii sui Trematodi endoparassiti. Primo contribute di osservazioni sui Dis-
tomidi. Zool. Jahrb., Suppl. 3; 229 pp.; 8 pi., 3 fig.

Odhner, T.

1900. Aporocotyle simplex n.g.n. sp., ein neuer T3T)us von ektoparasitischen Trematoden.

Centrlbl. Bakt. Par., Abt. 1, 27:62-66; 1 fig.
1905. Die Trematoden des arktischen Gebietes. Fauna Arctica, 4:291-372; 3 pi., 4 fig.
1907. Zur Anatomie der Didymozoen; ein getrenntgeschlechtlicher Trematode mit

rudimentarem Hermaphroditismus. Zool. Stud. TuUberg, p. 309-342; 1 pi., 6 fig.
1911. Zum naturlichen S3'stem der digenen Trematoden. I. Zool. Anz., 37:181-191.
1914. Die Verwandtschaftsbeziehungen der Trematodengattung Paragonimus Bm.

Zool. Beitr. aus Uppsala, 3:231-245; 5 fig.
OSBORN, H. L.

1911. On the Distribution and Mode of Occurrence in the United States and Canada of
Clinostomum marginatum, a Trematode Parasitic in Fish, Frogs, and Birds.
Biol. BuU., 20:350-366; 1 pi.
Pratt, H. S.

1902. Synopses of North American invertebrates. 12. The trematodes. Am. Nat.,
36:887-910, 953-979.^
Ranson, B. H.

1920. Synopsis of the Trematode family Heteroph\idae with descriptions of a new
genus and five new species. Proc. U. S. Nat. Mus., 57:527-573; 33 fig.
RtTDOLPm, C. A.

1801. Beobachtungen iiber die Eingeweidewiirmer. Arch. Zool. Zoot., 2:1-65.
1809. Entozoorum sive vermium intestinalium historia naturalis. 2, 457 pp.
1819. Entozoorum synopsis cui accedunt mantissa duplex et indices locupletissimi.
811 pp.

SCHSANZ, F. VON P.

1788. Verzeichniss der bisher hinlanglich bekannten Eingeweidewiirmer, nebst einer
Abhandlung iiber ihre Anverwandtschaften. Miinchen, 166 pp.
Seebold, C. T. E. von

1835. Helminthologische Beitrage. Archiv Naturg., 1 :4S-84; 1 pi.
Skrjabin, K. I.

1913. Vogeltrematoden aus Russich Turkestan. Zool. Jahrb., Syst., 35:351-388; 2 pi.

1913a. Tracheophilus sisowi n.g. n. sp. Centrlbl. Bakt. Par., Abt. I, Orig., 69:90-94; 1 pi.

86 ILLINOIS BIOLOGICAL MONOGRAPHS (304

Stafford, J.

1900. Some undescribed Trematodes. Zool. Jahrb., Syst., 13:399-414; 1 pi.
1902. Cephalogonimtis americanus (new species). Centrlbl. Bakt. Par., Abt. I, Orig.,
32:719-725; 1 pi.
Stiles, C. W. and Hassall, A.

1894. A preliminary catalogue of the parasites contained in the collections of the
United States Bureau of Animal Industry, United States Army Medical Museum,
Biological Department of the University of Pennsylvania (Coll. Leidy) and in
Coll. Stiles and Coll. Hassall. Vet. Mag., 1:245-253, 331-354.

1908. Index-Catalogue of Medical and Veterinary Zoology; Subjects Trematoda and
Trematode diseases. Hygienic Lab., Bull. 37 :401.

Stossich, M.

1902. H Monostomum mutabile Zeder e le sue forme affine. Boll. Soc. adriat. sci. nat.
Trieste, 21:1-40; 9 pi.
Stxjnkard, H. W,

1917. Studies on North American Polystomidae, Aspidogastridae, and Paramphistomi-
dae. 111. Biol. Monogr., 3-281-394; 11 pi.

1919. On the specific Identity of Heronimus chelydrae MacCallum and Aorchis exknsus
Barker and Parsons. Jour. Par., 6:11-18; 2 pi.
Taschenberg, O.

1879. Didymozoon, eine neue Gattung in Cysten lebender Trematoden. Ztschr. ges.
Naturw., 5:605-617.
Tyzzer, E. E.

1918. A monostome of the genus Collyriclum occuring in the European sparrow, with
observations on the development of the ovum. Jour. Med. Res., 38:267-292;
4 pi.

Vaillant, L.

1863. Note sur deux helminthes trematodes observ6s chez la sirfene lacertine. C. r. Soc.

biol., 4:6-7.
1863a. Note sur quelques helminthes de la sirSne lacertine. Ann. sci. nat., Zool., 19:
347-350, 1 pi.
Ward, H. B.

1909. The Influence of Hibernation and Migration on Animal Parasites. Proc. 7.
Internat. Zool. Cong., 12 pp.

1917. On the Structure and Classification of North American parasitic Worms. Jour.
Par., 4:1-12; 1 pi.

1918. Parasitic Flatworms. In Ward and Whipple, Fresh Water Biology. Pp. 365-
453; 114 fig.

1921. A New Blood Fluke from Turtles. Jour. Par., 7:114-129; 1 pi.
Ward, H. B. and Hirsh, E. F.

1915. The species of Paragonimus and their diflferentiation. Ann. Trop. Med. Par.,
9:109-162; 5 pi.
Wedl, C.

1857. Anatomische Beobachtungen Uber Trematoden. Sitz. Akad. wiss. Wien.,
math.-naturw. CI., 26:241-278; 4 pi.

WlLLEMOES-SUHM, R. VON

1873. Helminthologische Notizen III. (II Uber den Bau und den Embryo des Mono-
stomum faba Brs.) Zeit. wiss. Zool., 23:332-335; 1 fig.
Zeder, J.'G. H.

1800. Erster Nachtrag zur Naturgeschichte der Eingeweidewurmer, mit Zufassen und
Anmerkungen herausgegeben. Leipzig. 320 pp.

1803. Anleitung zur Naturgeschichte der Eingeweidewurmer. Bamberg. 432 pp.

88

ILLINOIS BIOLOGICAL MONOGRAPHS

[306

DESCRIPTION OF PLATES

LIST OF ABBREVIATIONS

c Cirrus

cm circular muscle

cp cirrus pouch

g ventral glands

ga genital atrium

i intestinal crura

Im longitudinal muscle

m mouth

0 ootype

e esophagus

om oblique muscle

OS oral sucker

ov ovary

p pharynx

pp prepharynx

rs receptaculum seminalis

ru receptaculum seminalis

uterinum
sg shell gland
u uterus
V vas deferens
vd vitelline duct
vr vitelline reservoir
w wall of excretory tubule

All drawings were made by the aid of either a camera lucida
or an Edinger drawing apparatus.

307] NORTH AMERICAN MONOSTOMES 89

PLATE I

90 ILLINOIS BIOLOGICAL MONOGRAPHS [308

PLATE I

Fig. 1. CyclococlumleidyiyAoTS^ivitvf. Xl2.
Fig. 2. Portion of the egg filled uterus of Cydocoebim leidyi. X45.

Fig. 3. Cydocoeliim mutahile (Zeder). Specimen No. 284 from Mehlis Collection, dorsal
view. Xl2.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE I

309] NORTH AMERICAN MONOSTOMES 91

PLATE II

92 ILLINOIS BIOLOGICAL MONOGRAPHS [310

PLATE II

Fig. 4. Cydocoelum pseudomicroslomu)n,wentT3\view. X12.
Fig. 5. Cydocoelum halli, ventral view. X 12.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE II

311] NORTH AMERICAN MONOSTOMES 93

PLATE III

94 ILLINOIS BIOLOGICAL MONOGRAPHS [312

PLATE III

Fig. 6. CydocoelumwUsonifVexiX.rsS.vi&vr. X12.
Fig. 7. Cyclocoelum cuneatum, dorsal view. X 12.
Fig. 8. Cyclocoelum obscurum, ventral view. Xl7,

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE III

^^^' NORTH AMERICAN MONOSTOMES

95

PLATE IV

96 ILLINOIS BIOLOGICAL MONOGRAPHS [314

PLATE IV

Fig. 9. Cyclocoelum macrorchis, ventral view. X 15.

Fig. 10. Cyclocoelum iriangularum, ventral view. Xl2.

Fig. 11. Cyclocoelum halli, young specimen, ventral view. X23.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

&

v^!':l

HARRAH NORTH AMERICAN MONOSTOMES PLATE IV

315] NORTH AMERICAN MONOSTOMES 97

PLATE V

98 ILLINOIS BIOLOGICAL MONOGRAPHS [316

PLATE V !

Fig. 12. Cross section, immature specimen of Notocotylus urbanensis. X 190. !

Fig. 13. Spines of Paramonoslomum echinum, front view. X 700.

Fig. 14. Notocotylus urbanensis, young specimen, partial lateral view. X95. <

Fig. 15. Paramonostomum echinum, ventral view. X 87.

Fig. 16. Spines oi Paramonoslonnmi echinum, lateral view. X700.

Fig. 17. Notocotylus urbanensis, immature stage, ventral view. X150.

Fig, 18. Notocotylus urbanensis, ventral view. X44. i

Fig. 19. Notocotylus urbanensis, young specimen, dorsal view. XI 20. \

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

6

i

13 ^^

ijii^

Di

J7

HARRAH NORTH AMERICAN MONOSTOMES PLATE V

317J NORTH AMERICAN MONOSTOMES 99

PLATE VI

100 ILLINOIS BIOLOGICAL MONOGRAPHS 1318

PLATE VI

All figures magnified 92 times

Fig. 20. Ovarian complex, Cydocoelum halli, drawing from wax reconstruction.

Fig. 21. Ovarian complex, Cydocoelum obscurum, drawing from toto, same specimen as in

figure 8.
Fig. 22. Ovarian complex Cydocoelum ohliquum, drawing from wax reconstruction.
Fig. 23. Ovarian complex, Cydocoelum problemalicum, drawing from toto mount of No.

2449 Berlin Museum.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE VI

319] NORTH AMERICAN MONOSTOMES 101

PLATE VII

102 ILLINOIS BIOLOGICAL MONOGRAPHS [320

PLATE VII

Fig. 24. Ovarian complex, Cydocoelum cunealum, anterior view. X92.

Fig. 25. Frontal section, showing structure of body wall. X 435.

Fig. 26. Ovarian complex, Haematolrephus similis, partial anterior view from above. X92.

Fig. 27. Ovarian complex, Cydocoelum pseudomicrostomum, dorsal view. X92.

Fig. 28. Frontal section, anterior end of Haematolrephus similis. X145.

Fig. 29. Drawing of wax reconstruction of anterior end of Cydocoelum elongatum. X 108.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

cp 29

HARRAH NORTH AMERICAN MONOSTOMES PLATE VII

32 1 1 NORTH A M ERIC A N MONOSTOMES 103

PLATE VIII

104 ILLINOIS BIOLOGICAL MONOGRAPHS [322

PLATE VIII

All figures magnified 44 times

Fig. 30. Ventral view of anterior end of Cyclocoelum pseudomicroslomiim.

Fig. 31. Ventral view of anterior end of Cyclocoelum braziliamim.

Fig. 32. Dorsal view of anterior end of Cyclocoelum mutabile.

Fig. 33. Ventral view of anterior end of Cyclocoelum leidyi.

Fig. 34. Ventral view of anterior end of Cyclocoelum elongatum.

Fig. 35. Ventral view of anterior end, of Cyclocoelum brazilianum, cirrus extruded.

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE VIII

323] NORTH AMERICAN MONOSTOMES 105

PLATE IX

106 ILLINOIS BIOLOGICAL MONOGRAPHS [324

PLATE IX j

\

Fig.36-42. Inclusive. Consecutive 20 micra sections of oral sucker, Cyc/ocoetow AaWi. X71. ]

Fig. 43, Cross section of oral sucker, Cydocoelum pseudomicrostomum. X90. '

Fig. 44. Cross section through anterior portion of oral sucker, Cydocoelum elongalum. X90. i

Fig. 45. Cross section through genital atrium, Cyc/ocoe/wm e/oMgo^MW. Xl2. ;

Fig. 46. VrontdX section, Cydocoelum elongatum. X105. i

Figs. 47 & 48. Cross sections of oral sucker of Cydocoelum elongalum, posterior to that in i

figure 44. X90. j

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

HARRAH NORTH AMERICAN MONOSTOMES PLATE IX

COPYKIGHT, 1923 BY THE UNIVERSITY OP ILLINOIS
DiSTKlEUTED FEBRUARY 9, 1923

A CLASSIFICATION OF THE LARVAE
OF THE TENTHREDINOIDEA

WITH FOURTEEN PLATES

BY

HACHIRO YUASA

CoDtributioDS from the

Entomological Laboratories of the University of Illinois

No. 69

THESIS

SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE
OF DOCTOR OF PHILOSOFHY IN ENTOMOLOGY IN THE GRADUATE
SCHOOL OF THE UNIVERSITY OF ILLINOIS

1920

TABLE OF CONTENTS

I. Introduction 7

II. Morphology 14

in. Taxonomy 35

Superfamily Tenthredinoidea 37

Families of Tenthredinoidea 38

Family Xyelidae 39

Family Blasticotomidae 42

Family Tenthredinidae 42

Subfamilies of Tenthredinidae 4^

Subfamily Diprioninae 45

Neodiprion Rohwer 46

Species of Neodiprion 47

Monoctenus Dahlbom 48

Diprion Schrank 49

Subfamily Emphytinae 49

Genera of Emphytinae 50

Subfamily Selandriinae 51

Genera of Selandriinae 52

Thrinax Konow 52

Species of Thrinax 53

Strongylogaster Dahlbom 53

Species of Strongylogaster 53

Selandria Leach 54

Subfamily Dolerinae 54

Dolerus Jurine 55

Species of Dolerus 56

Subfamily Phyllotominae 57

Tribe Phyllotomini 57

Genera of Phyllotomini 58

Endelomyia Ashmead 58

Caliroa Costa 58

Species of Caliroa 59

Tribe Phlebatrophini 59

Phlebatrophia MacGillivray 60

Subfamily Tenthredininae 60

Genera of Tenthredininae '. . . 61

Macrophya Dahlbom 62

Species of Macrophya 63

Subfamily Cimbicinae 64

Genera of Cimbicinae 65

Cimbex Olivier 65

Trichiosoma Leach 65

Abia Leach 65

Species of Abia 66

Subfamily Hoplocampinae 66

Genera of Hoplocampinae 67

Hemichroa Stephens 67

Marlattia Ashmead 68

Caulocampus Rohwer 68

Subfamily Dineurinae 69

Subfamily Cladiinae 70

Genera of Cladiinae 71

Trichiocampus Hartig 71

Sf>ecies of Trichiocampus 72

Priophorus Dahlbom 73

Species of Priophorus 73

Cladius Rossi 74

Subfamily Nematinae 74

Genera of Nematinae 75

Diphadnus Hartig 76

Pristiphora Latreille 77

Species of Pristiphora 77

Micronematus Konow 79

Lygaeonematus Konow 81

Pachynematus Konow 81

Species of Pachjoiematus 82

Nematus Panzer 82

Sj)ecies of Nematus 83

Croesus Leach 83

Amauronematus Konow 84

Species of Amauronematus 84

Pteronidea Rohwer 85

Species of Pteronidea 85

Pontania Costa 88

Species of Pontania 88

Subfamily Blennocampinae 91

Genera of Blennocampinae 92

Tomostethus Konow 92

Blennocampa Hartig 93

Erythraspides Ashmead 94

Monophadnus Hartig 94

Hypergyricus MacGillivray 94

Species of H>'pergyricus 94

Monophadnoides Ashmead 95

Isodyctium Ashmead 95

Subfaroily Fenusinae 96

Genera of Fenusinae 96

Kaliofenusa MacGillivray 97

Fenusa Leach 97

Subfamily Scolioneurinae 98

Metallus Forbes 98

Species of Metallus 99

Subfamily Hylotominae 99

Hylotoma Latreille 100

Species of Hylotoma 100

Subfamily Schizocerinae 101

Schizocerus Lepeletier 102

Subfamily Acordulecerinae 103

Acordulecera Say 103

Species of Acordulecera 103

Family Pamphiliidae 104

Species of Pamphiliidae 105

Family Cephidae 108

Genera of Cephidae 109

Janus Stephens 110

Species of Janus 110

Adirus Konow 1 10

Trachelus Jurine Ill

Cephus Latreille Ill

Species of Cephus Ill

Hartigia Schiodte 112

Family Xiphydriidae 112

Xiphydria Fallen 113

Family Siricidae 114

Tremex Jurine 115

Family Megalodontidae 116

Family Oryssidae 117

Oryssus Latreille 118

IV. Phylogeny 120

V, Summary 133

VI. BibUography 135

Vn. Explanation of Plates 141

VIII. Index 169

325] LARVAE OF THE TENTHREDINOIDEA—YUASA

I. INTRODUCTION

That the cardinal principle of modern taxonomy is based on the funda-
mental facts of evolution and that the essential problem of classification
is the phylogenetic relationship of organisms need no argument. In order
to ascertain genetic afl&nities, it is not sufficient to investigate the morpho-
logical characters alone, but all other attributes, physiological and biolog-
ical, must be considered. It is also evident that the immature stages of
organisms should receive as thoro consideration as the adult if taxonomy
of insects is to attain that degree of comparative perfection obtained in the
classification of other organisms.

Systematic entomologists, dealing as they do with animals of such
diversity and complexity morphologically and biologically, have from
early times recognized, at least to some extent, the taxonomic significance
and value of the developmental stages of insects, but the practical difficul-
ties in obtaining necessary materials, accurately determined and adequate
in quantity and range, have made progress in this phase of insect taxonomy
very tardy. A good start, however, has been made by recent workers as
was pointed out by Brues (1919), and their results vindicate both the
possibilitity and practicability of such investigations. There is, moreover,
an urgent demand for such studies from economic entomologists, who are
constantly confronted by the problem of identifying the immature stages
of economic species.

The present study is an attempt to deal with the larvae of the Ten-
thredinoidea from the standpoint of synoptic and, to some extent, genetic
classification. The systematic significance of the morphological charac-
ters will be discussed in part two; the taxonomic treatment of the families,
subfamilies, genera, and species will constitute part three; and, as full a
discussion of the phylogenetic relationship of the families as is possible
with the data at hand, will form part four. No one appreciates the inade-
quacy of this study, both in thoroughness and comprehensiveness, more
than the author, but it is hoped that he has opened a way for those who
will advance our knowledge of this highly interesting group of insects to
a more satisfactory condition in the future.

The taxonomic literature dealing with the adults of the Tenthredinoidea
is extensive. The historical development of the subject is interesting to
students of this group of insects but a detailed account is out of place here.
However, a brief statement of the history of the group is desirable.

8 ILLINOIS BIOLOGICAL MONOGRAPHS {326

Linnaeus in the fourth edition of the Systema Naturae (1744) estab-
lished the order Hymenoptera under the name of Gymnoptera and applied
to the order its present designation in the first edition of the Fauna Suecica.
The name Piezata was proposed by Fabricius (1775) for the order, but
this name never came into general use. Latreille (1796), following Lin-
naeus, divided the order into two sections, Terebrantia and Aculeata.
The first section included two groups, Phytophaga, which comprises the
Tenthredinoidea, and the Entomophaga or parasitic Hymenoptera. The
Ditrocha and Monotrocha of Hartig (1837) correspond approximately
with the two sections of Latreille. Gerstaecker clearly recognized the
Tenthredinoidea as a unique compact group and proposed in 1867 to
divide the order Hymenoptera into two suborders. He used the name
Symphyta for the Tenthredinoidea and Apocrita for the remainder of the
order. The term Symphyta thus antedates Konow's (1890) subordinal
name Chalastogastra. Various terms have been proposed for this group
of Hymenoptera and the following are coextensive with the superfamily
name Tenthredinoidea as used in the present paper: Phytophaga, Ses-
siliventres, Securifera, Serrifera, Symphyta, and Chalastograstra. Rohwer
and Cushman (1917) proposed a third suborder of Hymenoptera, Idiogas-
tra, for the family Oryssidae and placed it between the Chalastogastra and
Clistogastra of Konow.

Early students of the Tenthredinoidea divided the superfamily into
two groups, Phyllophaga for the Tenthredinidae or "Tenthredo" of
Linnaeus and Xyllophaga for the Siricidae or "Urocerus" of Geoffroy.
With the exception of Stephens (1835) and Andr6 (1879), who recognized
the additional families Xiphydriidae and Cephidae, respectively, besides
the two families mentioned above, the old system was followed for many
years. With the progress in studies of the world fauna of this group
of insects, modern writers l^iave proposed many elaborate schemes of classi-
fication. Konow in 1890 suggested one family and three subfamilies and
Dalla Torre (1894) catalogued one family divided into eighteen subfamilies,
while Ashmead (1898) proposed fifteen families and twenty-seven sub-
families. Enslin (1911) criticized Konow's three divisions as unnatural
and proposed four families, Oryssidae, Siricidae, Cephidae, and Tenth-
redinidae, thus reverting to a considerable extent to the scheme of the old
school as represented by Cameron (1882) and others. The recent and
more important systems are those proposed by Konow (1905), MacGilliv-
ray (1906), and Rohwer (1911). These systems, when compared, show a
great discrepancy in the number and rank of the groups which formerly
constituted the family Tenthredinidae, as is indicated graphically in
Plate XIV. MacGillivray, whose classification is based on a thoro-going
phylogenetic study of the wings, is of the opinion that the large complex
of genera obtained in this family are readily separable into a number of

327] LARVAE OF THE TENTHREDINOIDEA—YUASA 9

definite groups on structural differences, and that they are best dealt
with by considering them simply as subfamilies. In general the systems
of Konow and Rohwer are, with the exceptions noted below, more in
accordance with each other in their essential features than either one of
them is with that of MacGillivray. A comparison of these three schemes
brings out various points of interest. As far as the major groups are con-
cerned, (1) all are in agreement in associating Xiphydriidae with Siricidae;
(2) Konow and Rohwer agree in placing Siricidae and Siricoidea closer to
Lydidae and Megalodontoidea respectively than does MacGillivray, as
also in associating Megalodontidae with Pamphiliidae and Xyelidae and
Pamphiliidae with Cephidae; (3) Konow and MacGillivray agree in the
relation of the Blasticotomidae to Xyelidae and Pamphiliidae; (4) Rohwer
differs radically from them in his arrangement of the Blasticotomidea,
which he places between the Argidae and Tenthredinidae in his super-
family Tenthredinoidea; and, finally (5), Rohwer (1917) is unique in
creating a third suborder of Hymenoptera, Idiogastra, for the Oryssidae,
In respect to the arrangement of the subfamilies and tribes of the Tenth-
redinidae, as restricted by MacGillivray, the striking dissimilarity of these
authorities in assigning different rank to more or less related groups is
well illustrated in their treatment of the Emphytinae, Selandriinae, and
Lycaotinae. According to MacGillivray Konow's tribe Selandriades of his
family Tenthredinini corresponds to the three subfamilies just mentioned,
while according to Rohwer it embraces not only these subfamilies but also
another — Allantinae; that is, Rohwer considers the Emphytinae related
to the Tenthredmidae thru the tribe Allantini, which, together with the
tribes Taxonini and Eriocampini, constitute his subfamily Allantinae.
The Emphytinae and Lycaotinae are also related, according to Rohwer,
to the Blennocampinae thru his subfamily Empriinae, which contains,
besides Empriini and Lycaotini, the tribe Blennocampini. MacGillivray
and Rohwer agree in regard to the Cimbicinae to the extent that they
both consider it a compact group. Konow differs radically from these
writers by associating his tribe Syzygoniides with the Cimbicides and
Abiides. In regard to Konow's Nematides and Lobocerotides, the three
systems agree fairly well. The affinities of the Blennocampinae, Fenusinae,
and Scolioneurinae are recognized by MacGillivray and Konow. Rohwer
and Konow agree, as do all other systematists except MacGillivray, in
treating the Diprioninae and Monocteninae as allied groups inseparable
into subfamilies. Konow's subfamily Lophyrini, however, is a hetero-
geneous group and includes such widely separated groups as Acordule-
cerinae and Diprioninae.

The classification proposed by MacGillivray is based upon a critical
investigation of an essential structure, the wing, and is a logical conclusion
of the application of the taxonomic principles promulgated by Comstock

10 ILLINOIS BIOLOGICAL MONOGRAPHS [328

(1893). This system of classification is adopted in principle in the present
paper, for the writer believes that by judicious discrimination between the
palingenetic and the cenogenetic peculiarities, the characters manifested
by immature insects may be interpreted as indicating the genetic affinities
of the insects under consideration, and that in this sense the immature
stages of insects are of systematic importance. A classification based upon
such larval characters should agree in essential points with that based
upon the adult characters chosen for their phylogenetic value, and there-
fore it is interesting to see whether MacGillivray's alary system is justified
by a study of the larvae. It must be stated here that Dr. MacGillivray
does not accept all my conclusions as to the relation of the families based
upon a study of the larvae.

The immature stages of the Tenthredinoidea of more common occur-
rence seem to have attracted the attention of naturalists from an early
time. In Moufet's Theatorum Insectorum (1634) the adult saw-flies
are referred to the group "Vespa" and what seems to be a Tremex to
"Odonata," but no mention of the larvae is made. Goedart (1682) was
the first naturalist to make observations on the larvae of a saw-fly. His
records, rendered in interesting archaic terms, clearly indicate that he had
under observation the larvae of Cimbex or Trichiosoma on Salix, and Arge
on Rosa. Madame Merian (1730) pictured saw-fly larvae together with
those of the Lepidoptera, as may be seen on her plates 22, 25, and H.
Swammerdamm's (1737) figure 1, table XLIV, refers to galls on the
leaves and stems of Salix apparently made by Pontania or Euura. He
also figured a larva having fifteen segments with larvapods on abdominal
segments 3-8 and 11. Frisch (1766) figured about seven species and
recorded observations on their life-history. De Geer, as cited by Bergmann,
on the authority of Le Peletier (1823), had seen a larva with twenty-
two feet exclusive of the anal larvapods. According to the same authority,
Reaumur observed certain larvae with twenty-four feet, indicating a
xyelidan condition. Linnaeus (1758) recorded the food-plants of twenty-
two species of saw-flies out of the forty species of "Tenthredo" enumer-
ated, and spoke of the larvae in general of "Tenthredonis larvae pleraeque
folia plantarum exedunt, polypodae, seu pedibus plus quam XVI com-
muniter instructae," Following Linnaeus many students of the Tenthre-
dinoidea contributed much to a knowledge of the immature stages, altho
quite disproportionately to their larger contributions to that of the adults.
Among those who have done much towards the progress of our knowledge
of the larvae, either as original investigators or as compilers or as both,
the following are the more important: Andre, Brischke, Cameron, Costa,
Dahlbom, Dalla Torre, Dufour, Dyar, Fallen, Hartig, Kaltenbach, Kirby,
Klug, Konow, Latreille, Leach, Middleton, Newman, Oliver, Panzer,
Le Peletier, Snellen von Vollenhoven, Spinola, Westwood, and Zaddach.

3291 LARVAE OF THE TENTHREDINOIDEA—YUASA 11

Still, our knowledge of the immature stages of the Tenthredinoidea is
very meager in comparison to that of the adult. The larvae of only
418 species out of the total of 2701 species listed by Konow in his mono-
graph (1905) were dealt with in his artificial table for the larvae. This was
less than sixteen per cent of the described species of the world up to his
time, and many larvae included in this sixteen per cent were known to
this authority only thru literature. According to Dyar (1895) less than
twenty-five per cent of the North American species have been recognized
in the larval state. It is no exaggeration to say that the larvae of more
than eighty per cent of the Nearctic species are yet to be described.

Our knowledge of the physiology and morphology of the immature
stages of the Tenthredinoidea is exceedingly meager. The following list
includes most of the important literature: Graber (1890) on the embry-
ology of Arge berherides; Doncaster (1907) on the gametogenesis and
fertilization in Nematus ribesi; Buchner (1918) on the accessory chromo-
somes in Tenthredo, Allantus, Arge, etc.; Frenzel (1885) on the epithelial
regeneration of the alimentary canal of the larva of Cimbex; Holtz (1909)
on the histology and physiology of the digestive cells in the larva of
Nematus; Poletajew (1885) on the silk-glands of the larvae of Cimbex
and Tenthredo; Cholodkovsky (1897) on the blood and reflex bleeding of
the larva of Cimbex; and MacGillivray (1913) on the general external
anatomy of the larvae.

The biology of the Tenthredinoidea abounds in phenomena of great
interest to experimental evolutionists and presents many problems of eco-
logical importance. The list of papers dealing with the life-history and
habits, especially of economic species, is fairly extensive. Cameron
(1882) has published an excellent summary of general biological and
ecological observations. The biology of the Nearctic Tenthredinoidea
has been discussed in detail by MacGillivray (1913). Since the biological
and ecological studies of the Tenthredinoidea are beyond the scope of the
present paper, readers are referred to the last-mentioned publication.

Materials. — Four collections, designated for convenience as the Cor-
nell, Maine, MacGillivray, and Yuasa collections, contain most of the
materials used in this study. The writer has examined more than 2500
alcoholic specimens of larvae representing at least 400 species during
the course of this study. He has also, during the last few years, at Ithaca,
N. Y., and at Urbana, 111., made observations on the life-history and
habits of numerous species in his attempts to breed more than two hundred
and fifty species.

The Cornell collection consists of about forty species and belongs to
the Cornell University. Most of the specimens were collected by Dr.
MacGillivray and Mr. Chester Young in the vicinity of Ithaca, N. Y.
They were in rather a poor state of preservation and proved useful only

12 ILLINOIS BIOLOGICAL MONOGRAPHS |330

in checking up materials in other collections and in comparing the de-
scriptions of Young with his own material. This collection is designated
by the letter C.

The Maine collection consists of about 200 species, many of which were
bred and identified by Dr. MacGillivray. The collection belongs to the
Maine Agricultural Experiment Station and all the specimens were in
excellent condition. They were collected by Dr. MacGillivray with the
assistance of Mr. Earl Shaw during the summer of 1913 in the vicinity of
Orono, Maine. This collection together with the collecting and breeding
records were placed in the writer's hands, and they proved to be indis-
pensable to the present study. The collection is designated by the letter
M.

The MacGillivray collection consists of about thirty-five species
collected and identified by Dr. MacGillivray, together with larvae of
some unidentified species. The specimens came from Ithaca, N. Y.,
Orono, Me., Onekama, Mich., Urbana, 111., and a few other localities.
This collection is designated by the letter G.

The Yuasa collection consists of about two hundred and thirty species
including 98 bred species. A majority of the specimens were collected by
the writer at Ithaca, N. Y., during the summers of 1917 and 1918. Some
species were collected at Urbana, 111., and others came from different
parts of the United States and Canada thru the generosity of various
entomologists. This collection contains also the cocoons of practically all
the bred cocoon-making species and eggs and pupae of a limited number of
species. This collection is designated by the letter Y.

Besides the four collections just mentioned, a number of rare speci-
mens were generously loaned to me by several people, as subsequently
acknowledged, and were of great value in the preparation of this paper.

Identification. — All bred species in the Maine, MacGillivray and
Yuasa collections were identified by Dr. MacGillivray. Some of the
specimens in the Cornell collection bore labels, and when the larvae agreed
satisfactorily with the published descriptions the identifications were
accepted. In only a few cases has identification depended solely on
published descriptions of larvae.

Terminology and Nomenclature. — For the description of the external
anatomy of the head and mouth-parts of the larvae the terms used in my
paper (1920) dealing with the generalized insects have been used. Other
terms, some new, are used in part II. Taxonomic names have been
adopted from Rohwer (1911) and MacGillivray (1906).

Bibliography. — Works on taxonomic units are omitted altogether. A
complete bibliography of the Nearctic Tenthredinoidea was not under-
taken owing to space limitations, but the most important literature on
the subject is listed, as also that cited in the text.

3311 LARVAE OF THE TENTHREDINOIDEA—YUASA t3

Acknowledgments. — I am deeply indebted to various scientists who
have assisted me by donation of specimens, by encouragement, and by
criticism and suggestions. I am obliged to the former Director of the
Maine Agricultural Experiment Station, C. D. Wood, and to Dr. Edith
Patch of the same Station; to Dr. J. Chester Bradley, of the Cornell
University, and to Professor S. A. Forbes, of the University of Illinois, for
the loan of specimens and for the privilege of examining collections under
their charge. I wish to thank Dr. J. G. Needham for the privilege of
working in the entomological laboratories of Cornell University during the
summers spent at Ithaca, and Mr. S. A. Rohwer, of the U. S. Bureau
of Entomology, Washington, D. C, for his generous criticism and for the
donation of valuable reprints. I am also indebted for separates to Dr. W.
E. Britton, and to Messrs. H. E. Burke and William Middleton; for the
donation of specimens, to Messrs. H. G. Crawford, Wm. Baerg, C. C.
Hamilton, R. W. Harned, J. W. McCoUoch, J. R. Malloch, H. S. Smith,
H. B. Weiss, and to Drs. W. E. Britton, C. H. Kennedy, Edna Mosher, and
Alvah Peterson; and for the loan of rare specimens of Cephidae and Xiphy-
dria, and of Monoctenus and Trachelus to Dr. E. P. Felt and Mr. S. A.
Rohwer, respectively. I take pleasure in acknowledging my deep in-
debtedness to Dr. Albert William Bellamy, of the University of Chicago,
for constant encouragement. I am greatly obliged to Professor William
Trelease, of the University of Illinois, and to Professor K. M. Wiegand
and Mr. A. R. Bechtel, of the Cornell University, for the identification
of the host-plants. It gives me great pleasure to acknowledge my lasting
obligation to Professor Alexander Dyar MacGillivray, under whose
supervision this work was undertaken, for his constant interest, encour-
agement, and helpful suggestions thruout the entire course of this study
and for identifying all of the bred specimens in my collection, as well as
for the privilege of using his unpublished morphological terminology.

14 ILLINOIS BIOLOGICAL MONOGRAPHS ^332

II. MORPHOLOGY

The external anatomy of the larvae of the Tenthredinoidea has received
but little attention from entomologists. The only important contribution
on the subject is MacGillivray's study (1913), which deals with the anat-
omy and coloration as well as with the biology of the larvae. The writer
has made a comparative study of the external anatomy of the larvae of
representative Nearctic Tenthredinoidea in order to find characters upon
which both analytic and synoptic classifications of the larvae might be
based. Since MacGillivray has treated the general aspect of the anatomy,
only the more important structures and features will be discussed in the
following pages.

The larvae of the Tenthredinoidea (Figs. 1-25) are typically subcylin-
drical, cruciform, caterpillar-like, slightly flattened on the ventral aspect,
and usually taper slightly caudad. In the leaf-miners the body is de-
pressed. The body in its metamerism is well differentiated into a head
and a series of thirteen somites which are more or less similar in structure.
The segmentation is distinct. The first three segments compose the
thorax and are distinguishable on account of their position, form, and,
in podous larvae, more readily by the presence of three pairs of thoracic
legs. The abdomen consists of the ten remaining visible segments and in
polypodous larvae the presence of larvapods gives a characteristic appear-
ance to the uromeres. They are usually subdivided by transverse depres-
sions into annulets.

Head. — The head (Figs. 26-38) is typically subglobose, more or less
circular in frontal contour, strongly chitinized, and usually setiferous.
The mouth is directed ventrad or slightly ventro-caudad. In the leaf-min-
ers it is directed cephalo-ventrad as in the Fenusinae (Fig. 34) or cephalad
as in Phlebatrophia (Fig. 37). The surface of the head may be polished
and shiny as in Neodiprion (Fig. 28), or roughened, verrucose, or granu-
late, and divided into minute irregular areas as in Pteronidea (Fig. 30)
and the Cimbicinae (Fig. 29), or, in life, thinly coated with a waxy secretion,
as in some Emphytinae. It may be glabrous as in Metallus (Fig. 35) and
Phlebatrophia (Fig. 37), or it may be variously setiferous as follows:
microscopically and sparsely setiferous as in Tremex; with a few scattering
setae as in the Phyllotominae; with minute stiff peg-like setae as in Neodi-
prion; with numerous promiscuously distributed short setae as in Dolerus

3i3] TARVAE OF THE TENTHREDINOIDEA—YUASA 15

and the Tenthredininae; with long setae as in Pteronidea; or with abund-
ant conspicuous setae as in Monophadnoides and the Cladiinae. The
setae tend to be more numerous and longer on the ventral portion of the
head. The number and location of the setae on the head vary with the
individuals, excepting those on the clypeus and labrum, but their general
characteristics, such as relative abundance, manner of distribution,
and the kind of setae, are constant within genera and subfamilies. The
head may be pale, creamy white, or light brown, but oiften in life appears
as green or greenish white on account of the greenish blood showing thru
the cuticle, or it may be blackish or brownish with or without distinct color-
markings. The darker color is due to a deposition of colored pigments in
the cuticle and is generally permanent in alcoholic specimens. The color
and the coloration of the head are generally constant specific characters.
There are, however, ontogenetic changes in these respects. The very young
stages may be lighter in color and later stages darker, or vice versa as in
Cimbex, or all stages except the last instar may be darker and the ultimate
stage greenish as in Pteronidea ribesii. The color markings may be diffuse
in the young and become localized and definite in older stages, as in some
species of Dolerus, or they may vary from faint spots to general contiguous
markings as in certain species of Strongylogaster. The more common mark-
ings are brownish spots on the dorsum of the vertex, on the front, and often
caudad of each ocellera. There may be a stripe along the epicranial stem
and vertical furrows or dorsad of the ocellarae. When the head is darkly
colored the clypeus is usually lighter in color than the other parts of the
head.

The head is usually exposed, but there is a tendency in the leaf-miners,,
wood-borers, and a few others to have the cephalic end of the prothorax
produced into a broad fold on the dorsal and lateral aspects. This fold
covers the caudal portion of the head as in Tremex, Metallus (Fig. 35),
and Caliroa (Fig. 69).

The structures of the head will be discussed under two sections, one
dealing with the fixed parts, that is all the immovable parts of the head-
capsule, and the other dealing with the movable parts — the antennae and
mouth-parts. The fixed parts include the vertex, front, clypeus, labrum,
occiput, and postgenae, together with their bounding sutures, ocellarae,
and tentoria.

Epicranial Suture. — The inverted Y-shaped median suture of the head
is the epicranial suture. The stem (es) of the Y originates at the occipital
foramen, extends cephalad, dividing the vertex into halves, and bifurcates
on the cephalic aspect of the head. Each arm of the bifurcation (ea)
extends obliquely laterad for a short distance and then bends ventrad
to the ventral margin of the head, terminating near a precoila. The
epicranial suture is present in all larvae except those of the Xiphydriidae

16 ILLINOIS BIOLOGICAL MONOGRAPHS [334

(Fig. 45), where it is in part indistinct, and in the Siricidae and Oryssidae,
where it is obsolete. It is interesting to note that the most highly special-
ized genus of the Tenthredinidae, Phlebatrophia (Fig. 37), possesses this
suture. The relative length of the stem and arms varies in different
families and subfamilies. There is a depression near the bend of the arm,
as in Lygaeonematus (Fig. 31), which indicates the point of attachment of
muscles {ma), and should not be confused with a true tentorina (/>«).
Near the ventral end of each epicranial arm there is a thickening of the
surface and a pit. This pit is a pretenorina {pn) and the thickened piece
corresponds to the clypealia {d) of the larva of Corydalis. The preten-
torina and clypealia are constant in position and universal in occurrence in
the Tenthredinoidea. In the Siricidae the pretentorinae are distinct and
sometimes mistaken for ocellarae. In ecdysis the head is split along the
epicranial suture nearly to the ventral ends of the arms.

Vertex. — The large area on each side of the epicranial stem is the vertex
(»). It extends from the dorso-meson of the epicranium to the ventral
margin of the head, laterad of the epicranial arm, and cephalad of the
occipital suture when this is present, and bears an ocellara {o) and anten-
naria (ar). There is on the dorsal part of the vertex on each side a distinct
furrow which originates at the occipital foramen and extends cephalad
for some distance onto the lateral aspect of the head. This is the vertical
furrow {vf) and is characteristic of the larvae of the Tenthredinoidea.
It is wanting only in the leaf-miners and wood-borers. The nature of this
furrows is not known. There is a corresponding carina on the ental sur-
face of the head, and the major muscles of the retractor of the mandible
are attached to the ental surface of the vertex dorsad and ventrad of the
vertical furrow. The furrows usually converge at the cephalic end, but
sometimes are subparallel to each other.

Genae. — The portion of the vertex ventrad of an imaginary line drawn
ventrad of each ocellara parallel to the ventral margin of the head is a
gena (g). The extent of the genae varies, therefore, according to the
location of the ocularia. The setae on the genae are sometimes longer
than those elsewhere.

Ocularia. — The larvae of the Pamphiliidae, Xyelidae, Tenthredinidae,
and Cephidae possess a pair of ocellarae (e), one on each side of the head.
These organs of sight are remarkably uniform and constant in structure
and location. With the exception of Phlebatrophia and the Cephidae, the
ocellarae are usually clear, semiglobose or at least distinctly convex,
and are located on or near the center of the ocularia {ou), which are usually
circular and distinctly blackish. The ocularia are located on the vertex
dorsad of the antennariae in the Tenthredinidae and caudad of them
in the Pamphiliidae and Cephidae. In the Cephidae and Phlebatrophia
the ocularia are obsolete and the ocellarae are indicated by pigmented

335) LARVAE OF THE TENTHREDINOIDEA—YUASA 17

granules showing thru the cuticle. The ocellarae are obsolete in the
highly specialized families, Xiphydriidae and its allies. The life-habits
are correlated with the presence and the degree of development of the
organs of sight.

Front. — The area bordered by the epicranial arms on the cephalic aspect
of the head is the front (if). The ventral boundary is indicated by a
transverse depression connecting the ventral ends of the epicranial arms.
This depression is the fronto-clypeal suture (fcs). The depression is
usually concave dorsad and often obsolete at each lateral end. The
front is usually flattened or only slightly convex and bears scattered setae
which vary in number and arrangement in different genera and in arrange-
ment in different individuals. The extent of the front is determined by
the length of the epicranial arms. The front is usually subquadrate,
often wider than 1 ong, but in some cases, as in Caliroa (Fig. 53), it is
much longer than wide. In the absence of the epicranial stem the lateral
boundaries are indicated by the pretentorinae as in Tremex (Fig. 46).
Phlebatrophia (Fig. 37) is unique in possessing a distinct median longi-
tudinal furrow on the dorsal half of the front.

Clypeus. — The area ventrad of the front is the clypeus (c). Its ventral
boundary is the clypeo-labral suture (els), and the lateral margins are free,
oblique, and converge ventrad. The clypeus is usually much wider than
long and is divided usually into postclypeus (po) and preclypeus (pe) by a
difference in color and by a transverse row of setae. Sometimes the
clypeal suture (cs) is distinct, as in some Nematinae (Fig. 31). The clypeal
setae vary in number from two to eight or ten but are constant within a
genus and often within a subfamily. Four is the most common number.

Lahrum. — The small lobe attached to the ventral margin of the clypeus
is the labrum (/). It is usually transverse and has a median emargination
on the ventral margin. This emargination is usually shallow and broad
but occasionally very deep, as in Eriocampa and a few other Emphytinae.
Dolerus (Fig. 42) is characterized by the distinct asymmetrical median
emargination which makes the sinistral half of the labrum much smaller
than the dextral. The cephalic margin is nearly smooth and slightly
oblique in Tremex (Fig. 46). The labrum is very small in the Xiphydriidae
(Fig. 45). From two to several labral setae {Is) are borne on each side
of the meson, those near the meson being usually smaller than the lateral
ones. The number of labral setae are as a rule constant within a genus.
A row of setae which may be seen projecting from the ventral surface of the
labrum belongs to the epipharynx. The labrum is often divided into
halves by a distinct median longitudinal depression, as in Caliroa (Fig. 53),
Endelomyia (Fig. 48), and some Tenthredininae. This character is generic
in some subfamilies and only specific in others. The labrum in the Cim-

18 ILLINOIS BIOLOGICAL MONOGRAPHS [336

bicinae (Fig. 29) is unique in having a pair of longitudinal depressions on
each half which converge ventrad and bound a small median piece.

Postgenae. — The area mesad of the lateral boundary of the vertex on
the caudal aspect of the head is the postgena {pa). The dorsal boundary
is the vertical furrow and the mesal the occipital foramen. The ventral
margin is concave and is connected with the labicoria {Ic). It is usually
more or less flat and glabrous. The occipital suture {os) is sometimes
distinct, as in Pteronidea.

Tormae. — At each end of the clypeo-labral suture there is a chitinized
rod which extends onto the ventral surface as far as the epigusta. This is
the torma, present in all Tenthredinidae.

Occiput. — The narrow area on the dorsal third of the occipital foramen
between the vertical furrows is the occiput. The dorsal boundary is in-
distinct since the occiput merges with the vertex without any indication
of a suture.

Maxillariae. — The very narrow chitinized sclerites which form a sub-
circular collar around the dorsal and lateral margins of the occipital
foramen have been identified as the maxillariae {my). They are usually
only slightly developed and are continuous with the cervacoria. The
identity of these sclerites with the maxillariae of generalized adult insects
is uncertain, but they occupy the same position as the maxillariae and
consequently are considered as homologous with them. The dorsal third of
the maxillariae in the leaf-miners Metallus (Fig. 35), Phlebatrophia (Fig.
37) and Fenusinae (Fig. 34), are strongly chitinized, distinctly concave,
trough-like, and produced entad. Along this ental margin a part of the
muscles which control the movement of the head are attached.

Occipital Foramen. — The large opening in the caudal aspect of the
head thru which the internal organs of the head are connected with those
of the body is the occipital foramen {of). The ventral margin of the
occipital foramen is membranous and connected directly with the laba-
coria and cervacoria {cc).

Precoila. — The strongly chitinized acetabulum located near the
ventro-mesal angle of the vertex or the dorso-lateral angle of the clypeus
is the precoila {pr). The preartis of the mandible {py) articulates at this
point. The precoila is distinct in all Tenthredinoidea.

Mandibularia. — The small transverse whitish or light-colored area
ventrad of the ventral margin of the head on the cephalo-lateral aspect is
the mandibularia {mb). It is usually only slightly chitinized and merges
with the mandacoria without any indication of a suture. The extensacuta
{ec) of the extensatendon of the mandible is usually distinct. The mandi-
bulariae are sometimes very large, as in the Xyelidae (Fig. 27).

Postcoila. — The cup-shaped acetabulum on the latero-ventral angle
of the postgena and caudal angle of the mandibularia is the postcoila

337J I^iRVAE OF THE TENTHREDINOIDEA—YUASA 19

{ptt), where the postartis of the mandible articulates. The occipital suture
{os) when present originates in or near the postcoila. The postcoila is
always present.

Paracoila. — There is a slight projection at the mesal end of the caudo-
ventral margin of the head where the cardo of the maxilla articulates. This
is the paracoila {pi). It is not well developed but is present and discernible
in nearly all tenthredinid larvae.

Odontoidea. — The lateral cervical sclerite is articulated with the head
capsule on the mesal margin of the postgena some distance ventrad of the
origin of the vertical furrow. This point of articulation is an odontoidea
{od) and is rather indistinct in the larvae of this group of insects.

Tentorium. — The tentorium is very simple in tenthredinid larvae.
It consists of the metatentoria (w/), corpotentorium {ct), and pretentoria.
The supratentoria are apparently obsolete. The metatentorium is the
strongly chitinized conspicuous ental bar extending into the head capsule
from the ventro-mesal margin of each postgena. The two metatentoria
fuse on the meson and form the bridge, the corpotentorium, which gives
support to the caudo-ventral portion of the head. The position of each
metatentorina is indicated by a pit- or slit-like depression {mn). The loca-
tion of the pretentorinae {pn) has already been indicated. A strong
ental arm, much smaller than a metatentorium, extends ventro-mesad
from each pretentoria into the head capsule and fuses with the corpoten-
torium near the middle of the latter. This bar is a pretentorium. In
ecdysis the tentorium breaks in the middle of the corpotentorium, freeing
the mesal ends of the pretentoria and metatentoria. The tentorium in
the Xyelidae and Pamphiliidae is similar to that of the Tenthredinidae in
structure and location.

The movable parts of the head include all the appendages, that is, the
antennae, mandibles, maxillae, and labium.

Antennae. — The antennae are present in the larvae of all Tenthredin-
oidea, but their structure, size, position, and number of segments vary in
the different families and subfamilies. Each is borne by a distinct anten-
naria {ar) which is located in the ventro-lateral portion of the vertex; in the
generalized families they are located cephalad of the ocularia; in the
specialized, ventrad of them. The antennariae are usually subcircular or
subquadrate. The antacoria {an) is usually extensive, distinctly convex,
and whitish in color (Figs. 143-153). It is only occasionally narrow and
confined to the periphery of the antennaria, as in certain Nematinae
(Fig. 154). The antennae of the Pamphiliidae (Fig. 26) are setiform,
one-half as long as the head is wide, with seven cylindrical segments.
There is a circular sensorium on the ventral aspect of the distal portion of
the second, third, and fifth segments (Fig. 39). In the Xyelidae (Fig. 27)
the segments are shorter but thicker, and vary in number from six to seven

20 ILLINOIS BIOLOGICAL MONOGRAPHS (338

according to the genus. The antennae of the Tenthredinidae (Figs-
145, 147, 149, 150, 153) apparently represent a specialization from those of
the Xyelidae. They are much shorter and vary in number of segments
from one to five. The antennae vary in shape and are conical in the
Emphytinae and other generalized subfamilies, with five ring-like or
limpet-shaped segments, with four more or less irregular, incomplete, often
partly fused segments in the Nematinae (Fig. 145), or flattened and fused
into a single segment in the Schizocerinae or button-like and one-segmented
in the Cimbicinae, Fenusinae, and Metallus (Fig. 147), or subcorneal or
irregular as in some Nematinae and Phlebatrophia (Fig. 150). When
the antennae consist of five segments, they are usually cylindro-conical
and remarkably uniform in shape. The antennal segments are usually
strongly chitinized, more or less ring-like, successively smaller in diameter,
and the distal segment is conical or occasionally erect and peg-like as in the
Diprioninae. The segments do not always form a complete ring; one
side may be reduced to a mere line, or be entirely wanting as in some
Nematinae, in which cases the segment is said to be incomplete. Some-
times fusion of all or some of the segments may take place. Certain
segments are sometimes setiferous and also bear some sensoria. The
number of segments is constant for a subfamily. The relative length and
shape of the segments vary, but are constant in species in some cases, and
in others constant in genera. The antennae of the Cephidae are small,
with four or five segments, while in the Xiphydriidae they are three-
segmented and in the Siricidae and Oryssidae single-segmented. It is
possible, therefore, to arrange the families of Tenthredinoidea in an as-
cending series according to the number and size of the segments of the
antennae. The tenacity of the antennae is well Illustrated in the Oryssidae,
which in spite of the extreme modification of other structures still retains
one-segmented antennae.

Mouth-parts. — The larvae of the Tenthredinoidea possess well-devel-
oped mandibulate mouth-parts. They include the mandibles, maxillae,
and labium, and are remarkably uniform and constant in structure in the
different families. The modifications take place in the relative size of
parts and in the number of segments of the articulated parts.

Mandibles. — The mandibles (rnd) are always present, and are typically
thick, strongly chitinized, and sharply dentate, the dextral dissimilar to
the sinistral in the number and shape of the dentes, and in having one or
two and occasionally more mandibular setae on the lateral aspect. The
number and arrangement of the dentes, the number of mandibular setae,
and the relative size and shape of the mandibles are constant for certain
genera. The mandibles of the Schizocerinae are rather thin and flattened,
and in Phlebatrophia very thin and elongated with one triangular blade-

339] LARVAE OF THE TENTHREDINOIDEA—YUASA 21

like dentis. In this character the larvae of this genus are more specialized
than all other larvae, even including Oryssus.

Maxillae. — The maxillae (mx) are always present and typically consist
of cardo, stipes, subgalea, palp if er, palpus (mp), galea (gl), and lacinia
(la). The cardo is usually more or less chitinized and is divided into a small
subcardo and a larger triangular alacardo. The subcardo articulates with
the head in the paracoila. On the lateral margin of the alacardo the large
stipes is attached. The stipes is usually less chitinized than the cardo,
submenbranous, convex on the lateral aspect, and attached to the lateral
margin of the alacardo. The cephalic aspect is membranous and is con-
tinuous with the maxacoria. The caudal aspect along the mesal margin
is strongly chitinized and continuous with the elongate triangular subgalea.
The line of fusion is indicated by a distinct oblique chitinized ridge which
extends from near the proximal end of the subgalea to the lateral angle of
the lacinia. The latero-ventral angle of the stipes is often produced as a
small triangular lobe, the stipal angle of Crampton (1921) as in the
Tenthredininae. The palpifer is a more or less membranous, mound-like
lobe attached to the distal end of the cephalic margin of the stipes and
often bears one or more setae. The palpus is borne by the palpifer and
typically consists of four more or less conical segments. The relative
size and shape of the segments vary and afford good characters for the
separation of genera and species. The galea is typically strongly chitinized,
digit-like, conical or slightly curved mesad, bluntly pointed, unsegmented,
and usually smaller than the palpus. The lacinia is located mesad of the
galea and cephalo-ventrad of the subgalea. It is usually subtriangular,
slightly flattened, lobe-like, and bears a row of setae on its oblique mesal
margin. It is sometimes distinctly flattened, strongly chitinized, with a
stiff row of setae, as in the Emphytinae, or rounded and with minute spi-
nous setae as in Diprion, or with a sharp triangular compressed seta in
addition to an ordinary row of setae as in the Xyelidae. The galea and
lacinia are always present except in Oryssus but are reduced in size in
Tremex. It is interesting to note that in the leaf-miners the palpi are
reduced but the galea are usually normal in size and larger than the palpi.
In Oryssus the maxillae are fleshy lobes with all the component parts
obsolete and with a brownish area in which a few sensory papillae are lo-
cated (Rohwer and Cushman, 1917). The palpi are apparently two-
segmented in the Xiphydriidae and Siricidae.

Labium. — The labium (li) consists typically of submentum, mentum,
stipulae, palpiger, palpi (//>), and togaglossa — representing the fused
glossae and paraglossae. The submentum (sm) and mentum (w) is typ-
ically membranous, convex, with two or more setae, and very broad in
the larvae of most species. In some cases, as in the Diprioninae, part
of the mentum is chitinized. In the leaf-miners, such as Fenusa, Metallus.

22 ILLINOIS BIOLOGICAL MONOGRAPHS [340

and Phlebatrophia, the mentum is strongly chitinized and flattened.
There is a distinct median longitudinal depression in Metallus. The
palpiger is practically wanting. The palpus is typically three-segmented
(Fig, 157). The relative size and structure of the segments vary, but are
usually constant for a subfamily or a genus. In Tremex and Phlebatrophia
the palpi are apparently two-segmented and very minute, while in Oryssus
they are obsolete. The stipulae are typically membranous, broad, some-
times bearing two setae, and fused with the mentum without any indication
of a suture. The stipulae are flattened and chitinized in Metallus and the
Fenusinae. Totaglossa is typically membranous, subglobose or bluntly
pointed, fused with the stipulae without any indication of a suture. It is
readily identified on account of its median position and characteristic
shape and structure. There is a slit-like opening for the duct of the silk-
gland, the sericos {crv) on the meson near the caudo-ventral aspect of the
totaglossa. The shape, size, and location of the sericos vary but it is
always present and chitinized. The cephalic or dorsal aspect of the
totaglossa is strongly convex, membranous, and sometimes bears a few
minute setae and sensoria.

Prepharynx. — The prepharynx, the so-called "hypopharynx" of the
larvae of the Tenthredinoidea, is very simple in structure and the bound-
aries of the parts that can be identified in generalized insects (Yuasa,
1920) are obsolete. The propharynx consists of the epipharynx and epi-
gusta. The epipharynx is membranous, is of the same size and shape as the
labrum, and bears an oblique row of a few setae on each side slightly
dorsad of the ventral margin. The epigusta is membranous and is sup-
ported on each lateral portion by a torma. The ambipharynx is restricted
and membranous. The parapharynx consists of the basipharynx and
hypopharynx Qix). The basipharynx is subglobose or convex, often
slightly chitinized on the sides, sometimes having a few minute setae,
and usually converges ventrad. The portion ventrad of the constriction is
considered as belonging to the hypopharynx, but is usually membranous
and continuous with the cephalic surface of the totaglossa without any
indication of difi^erentiation. The laciniae fit against the sides of the
constricted part of the parapharynx. No striking modification in form of
the prepharynx appears in the different families.

Trunk. — The portion of the body caudad of the head is the trunk.
It consists of thirteen segments which connect with the head by means of
the cervacoria. The first three segments compose the thorax and the
remainder the abdomen.

Cervacoria. — The membrane {cc) which connects the thorax with the
head is rather broad and usually folded under the protruding cephalic end
of the prothorax. There is a chitinized sclerite on each side, the cephalic
end of which articulates with the head against the odontoidea and the

341] LARVAE OF THE TENTHREDINOIDEA—YUASA 23

caudal end with the epiraeron and coxa. This is the lateral cervical sclerite
{Ics), which is always present and usually distinctly colored — brownish or
blackish. The cervacoria is continuous with the submentura on the
ventral aspect, and is often produced on the meson as a small mound-like
setiferous protuberance, as in Strongylogaster.

Thorax. — The prothorax is usually constricted on the cephalic portion,
the dorsal aspect is declivous, and the lateral aspect is produced. The
dorsum is typically divided into a narrow cephalic portion and a wide
caudal one, which in turn may be subdivided into two or more annulets
(Fig. 65). The first division is usually setiferous on the lateral aspect,
while the second division is setiferous on the dorsal aspect. On the middle
of the lateral aspect there is a large spiracle. This is the mesospiracle
(fftsp), which has migrated onto the prothorax. There is another setiferous
area cephalo-ventrad of the spiracle. The prothoracic leg is attached to
the latero-ventral margin of the segment, ventrad of the setiferous sub-
spiracular area, which is usually produced as a lobe and which Crampton
(1918) has designated as the surcoxal plate. There is a small, usually
strongly chitinized sclerite cephalad of the leg. This is the episternum-
epimeron, or the eupleuron of Crampton. The dorsal aspect of the pro-
thorax and often the lateral one may be chitinized and colored, forming
shield-like areas as in the Xyelidae and certain leaf-miners. The proster-
num is usually membranous, subdivided into two or more annulets, but
sometimes it is flattened and strongly chitinized as in Metallus. There is
usually a small pit or chitinized rod near the caudal part of the segment on
the ventral aspect. This is the profurcellina {pfn) and marks the caudal
limit of the prosternum. The mesothorax and metathorax are more
or less similar in structure, frequently the largest segments in the body,
more or less ring-like and often distinctly annulate. The metaspiracles
(tsp) are located in the metacoria and are usually minute and f unctionless.
The mesothorax and metathorax resemble the prothorax in other details.
In the Fenusinae (Fig. 21) and Metallus (Fig. 22) the dorsum of the
mesothorax and metathorax is provided with an ovoid, fleshy, sucker-like,
low protuberance (scp) on each side of the meson. Its function is not
known. Dimorphopteryx is characterized by the presence of a pair of
prominent dorsal protuberances on the prothorax and a median protu-
berance on the mesothorax.

Thoracic Legs. — The larvae of the Tenthredinoidea, with the exception
of the Oryssidae, possess three pairs of thoracic legs. They are the most
persistent of all the thoracic and abdominal appendages and as a rule are
very similar in structure, both facts indicating a common origin. A typical
leg consists of five more or less well-chitinized segments: coxa, trochanter,
femur, tibia, and a distal segment representing the fused tarsus and tarsal
claw.

24 ILLINOIS BIOLOGICAL MONOGRAPHS (342

The coxa {ex) is usually the largest of all the segments, subcorneal, and
articulated to the ventro-lateral margin of the segment. The cephalo-
dorsal angle is strongly chitinized and articulates against the chitinized
end of the episternum-epimeron. There is a distinct oblique depression
extending from this angle to the middle of the dorso-distal margin of the
coxa. The distal ends of the coxae are usually chitinized and form ring-
like thickenings. The ventral half of the coxa is more or less mem-
branous. The trochanter {tr) is usually small, longer on the ventral
than on the dorsal aspect. The femur (Jm) is usually cylindrical and is
often dilated at the distal end. Its ventro-distal portion is usually mem-
branous and is sometimes produced, forming a pointed projection, the
femoral process (Jp), as in Dolerinae (Fig. 135) and in related subfamilies.
The tibia {t) is subcylindrical, narrower in diameter at the distal than at the
proximal end, and either longer or shorter, than the femur or subequal to it.
The distal segment is typically very short and claw-like. The apparent
claw {cw) represents a fusion of the tarsus and tarsal claw, and is usually
sharp and distinctly curved. The segments are usually setiferous and
more or less membranous on the ventral aspect and at the joints.

The general plan of structure of the legs is the same in a majority
of the larvae, but there are variations in the shape, size, arrangement of
setae, and in the number of apparent segments within the families and
subfamilies. The variations usually consist in the suppression of the
trochanter as in Phlebatrophia (Fig. 136) and the Fenusinae (Fig. 140),
or in the modification of the distal segment as in the Pamphiliidae (Fig. 130)
and Hylotominae, or in the reduction of the entire structure to a fleshy,
subconical, indistinctly segmented clawless protuberance as in Phleba-
trophia and certain highly specialized families. The absolute homology of
the segments in a modified leg can not be established, but when the number
of the segments is less than five it is probable that the trochanter is the
first one to disappear.

The Pamphiliidae are distinct from all other tenthredinoid larvae in
having setaceous legs with all segments cylindrical except the distal ones.
The distal segment is very slender, non-setiferous, straight, and sharply
pointed without indication of a claw. The Xyelidae (Fig. 131) possess
legs which are small but typical in structure and number of segments.
It is quite possible to derive the normal tenthredinid legs from those of the
Xyelidae. In the subfamilies of the Tenthredinidae, a series of modifica-
tions of the legs is found, altho the majority of the subfamilies and genera
are provided with typical five-segmented, well-developed claw-bearing
legs. The Phyllotominae (Figs. 141, 142) are characterized by very
short, stubby, chitinized legs which consist of four segments, including the
large strongly curved claw. Phlebatrophia (Fig. 136) is unique among the
Tenthredinidae in having fleshy, rudimentary clawless legs. The Fenusi-

343] LARVAE OF THE TENTHREDINOIDEA—YUASA 25

nae (Fig. 140) also possess small four-segmented legs but, unlike the
Phyllotominae, have simple and ring-like segments and claws normal in
form. The Hylotominae differ from the other subfamilies in possessing
apparently six-segmented legs. Their distal portion consists of a distinctly
separated tarsus and claw and bears an empodium-like fleshy lobe on the
caudal portion of the claw. The minimum number of segments is found in
the legs of the Schizocerinae (Fig. 139), where the mesothoracic and
metathoracic legs consist of only three simple cylindrical segments, while
the prothoracic legs are composed of four. There is a well-developed
fleshy subglobose lobe caudad of the claw. It is interesting to note that
the gall-makers, for example, Pontania have essentially the same type of
legs as the leaf -feeding Nematinae (Fig. 133), while the leaf-mining larvae
of the Fenusinae, Schizocerinae, and others, have modified legs. The
highly specialized families, Cephidae, Xiphydriidae, and Siricidae, possess
fleshy, indistinctly segmented rudimentary legs which are never provided
with claws. The most specialized family, Oryssidae, is entirely apodous.
The legs present, therefore, very good characters for differentiating the
families and the subfamilies of the Tenthredinoidea. The phylogenetic
significance of the thoracic legs is quite evident.

Abdomen. — The segments composing the abdomen, with the exception
of the two caudal segments, are more or less similar in structure, ring-like,
and usually subdivided into four to seven annulets. Segments 2-7 or 2-8
and 10 usually bear a pair of larvapods, the so-called "prolegs," on the
ventral aspect. The first and ninth abdominal segments never possess
larvapods except in the Xyelidae. The third abdominal segment is more
typical than the other segments and least modified, and for this reason
has been used as a type for description. In a typical larva this segment is
subdivided into a number of annulets on the dorsum and latus dorsad of
the spiracular line. The latus ventrad of the spiracular line is typically
lobe-like, setiferous, and distinguishable as two areas, the subspiracular
area {ssl) or the surpedal area {sdl) according to the location. When
these areas are distinctly lobe-like they are designated as subspiracular
lobe and surpedal lobe. The latter corresponds to the thoracic surcoxal
plate of Crampton. The subspiracular and surpedol lobes sometimes fuse
and extend the full length of the segment as an oblique fold, as in wood-
boring larvae and the Hylotominae. In the latter this lobe is conspicuously
produced laterad, making the segment distinctly flattened. The sub-
spiracular and surpedal lobes, when fused, are known as the sublaterai
lobe {sll). The larvapods are located on the ventral aspect some distance
from the meson. The sternum is divided into two or more annulets, the
annulation being usually distinct but its exact limits diflScult to determine
on account of the presence of the larvapods. The intersegmental coria
{cor) is distinctly indicated on the venter (Fig. 81). The segmentation

26 ILLINOIS BIOLOGICAL MONOGRAPHS [344

is distinct but the limits of the somites are not so readily determined. The
cephalic limit of a segment is usually indicated by the distinct depression
on the dorsal and lateral aspects and by the short ventro-lateral depressions
which terminates at the cephalic end of the subspiracular lobe. Thus
the cephalic limit of a segment is not a straight line, but curves caudo-
ventrad and then slightly cephalo-ventrad of the subspiracular lobe.
The typical annulation and arrangement of setae, tubercles, and glandubae
on the typical abdominal segment are indicated elsewhere (Figs. 73-79).
The ninth abdominal segment is readily distinguishable because of its
location and shape, and by the absence of spiracles and larvapods. It is
typically smaller than the preceding segments, tapers more or less caudad,
and usually has one less annulet on the dorsum than have the preceding
segments. Its caudal limit is usually distinctly indicated by a deep
depression. The tenth, or the apparent ultimate, segment is modified and
differs from the other abdominal segments because of the presence of the
anus, anal larvapods, and other structures peculiar to this segment (Figs.
89-103).

Tenth Urotergum. — The tergum of the tenth abdominal segment is
usually convex and often setiferous. It sometimes bears numerous spinous
processes, as in the Blennocampinae and Dimorphopteryx, or paired suranal
protuberances as in certain genera of Nematinae, or a median suranal
process {srp), as in the Cephidae and its allies. In these highly specialized
families the tenth abdominal segment is produced cephalad and fits into
the deep semicircular emargination of the ninth segment. The tergum
possesses a distinct, deep, median longitudinal depression extending
from the cephalic end of the tergum to the proximal end of the median
suranal process. In the Xyelidae the tergum is produced distinctly hunch-
like on the meson of the cephalic third caudad of the deep, broad transverse
depression. The tergum is produced caudad in certain species of Pachy-
nematus and forms a distinct caudal projection. In this genus the gland-
ubae are very conspicuous. The size, convexity, number, and arrangement
of the spinous protuberances, caudal processes, and setae are useful
characters in recognizing different subfamilies and genera.

Suranal Lobe. — The membranous lobe (srl) of the Tenthredinidae,
which forms the dorsal wall of the anal slit may represent the rudiment of
the dorsal half of the ultimate segment, the so-called "telson." It bears
numerous setae of varying size and number and is usually fused with the
tergum of the tenth segment. In the larvae of the Xiphydriidae and its
allies, which possess a median suranal process, the suranal lobe is distinct,
more or less chitinized in part, usually separated from the tenth tergum by
a ridge or by an oblique suture which extends from the chitinized depres-
sion dorsad of the suranal process to the lateral end of the anal slit (au).
The area cephalad of this oblique suture is the pleuron of the tenth seg-
ment.

345] LARVAE OF THE TENTHREDINOIDEA—YUASA 27

Subanal Lobe. — The membranous lobe {shl) which forms the ventral
wall of the anal slit may represent the rudiment of the ventral half of the
ultimate segment or "telson." It is never distinctly chitinized, always
indistinguishably fused with the tenth abdominal sternum, usually setifer-
ous, and is rather restricted in extent. In the Tenthredinidae the subanal
lobe is strongly convex and extends to the anal larvapods, the postpedes of
Crampton. If the subanal lobe represents a part of the telson, and if the
subanal appendages {sba) of the Pamphiliidae and Cephidae are the
appendages of the ultimate segment, then the subanal lobe possesses a
pair of genuine appendages. In the larvae of the Xyelidae there occur
sometimes distinct subglobose setiferous swellings dorso-caudad of the anal
larvapods. Their homology and function are unknown.

Tenth Urosternum. — The sternum of the tenth abdominal segment is
restricted in extent, more or less convex, and often glabrous. In the
Xyelidae and Tenthredinidae the anal larvapods occupy the greater part
of the caudal portion, which is thereby produced subconically ventrad.
This sternum is usually glabrous and more or less flattened in apodous
larvae. There are no annulations observable on this sternum.

Suranal Process. — In boring larvae, the ultimate segment is provided
with a strongly chitinized mesal suranal process {srp) oil the suranal lobe.
This process, which has been variously designated by different writers, is
characteristic of the families Cephidae (Figs. 108, 109, 112, 114, 115)
Xiphydriidae (Figs. 107, 110), and Siricidae (Figs. 113, 120, 122). The
size, shape, number, and arrangement of the dentiform tubercles and
setae vary in different families but they are constant within species and
often also within genera. It is undoubtedly an adaptive structure devel-
oped in connection with the boring habit of the larvae. It is interesting to
note that in some of the gall-making larvae of Pontania and leaf-stem
boring larvae of Caulocampus, the tergum of the ultimate segment is
produced on the caudo-meson and forms a distinct protuberance provided
with chitinized points on its caudal end. The larvae of the Pamphiliidae
also possess a minute hook-like process on the caudo-meson of the ultimate
tergum. The genetic connection of this hook-like tubercle and the distinct
suranal process of highly specialized families is doubtful, but it is not difficult
to surmise a common origin for the suranal process of the Cephidae,
Xiphydriidae, and Siricidae. The suranal process corresponds to the
postcornu of Crampton.

Caudal Protuberances. — In certain genera of the Nematinae and in
a few other genera of the Tenthredinidae, the tergum of the ultimate
segment is provided with two or more protuberances which vary in size,
shape, number, and position in different genera. A typical condition in
nematid larvae is found in Pteronidea (Figs. 126, 127), which possess a
pair of conical, pointed, well-chintinized processes {srp), one on each side.

28 ILLINOIS BIOLOGICAL MONOGRAPHS (346

on the caudal margin of the tergum dorsad of the membranous suranal
lobe. These processes are always two in number and more or less constant
in position in nematid larvae, but vary in size and shape altho constant
within species. They are conical, subconical, sharply or bluntly pointed,
truncate, or distinctly swollen at the distal end as in Pteronidea trilineata.
The tenth abdominal tergum of the spinous larvae of the Blennocampinae
is provided with several symmetrically arranged conspicuous spinous
processes on the caudal portion and in part along the caudal margin. The
larva of Dimorphopteryx is unique among the Emphytinae in the possession
of four very distinct, sharply pointed spinous protuberances along the
caudal margin of the ultimate segment dorsad of the suranal lobe. In
certain of the gall-making species of Pontania the caudal end of the tenth
abdominal tergum is produced caudad and forms a median prominence
which usually has two minute strongly chitinized points close together on
the meson. A similar protuberance is found in the larvae of Caulocampus
acericaulis. In the Siricidae a pair of minute sharply pointed solid chitin-
ized spines occurs on the tergum of the ultimate segment, one on each
side of the median longitudinal depression (Fig. 113).

These protuberances have been variously designated. Crampton
(1919) considers the paired protuberances and spines to be homologous
with the cerci of Orthoptera and Ephemeridae. If they represent rudi-
mentary cerci they must belong to the eleventh abdominal segment,
the telson of embryologists, since the true appendages of the tenth segment
are transformed into the anal larvapods. But this homology is open to
question because these protuberances are mere projections of the surface
and not at all appendages in a morphological sense, and, furthermore,
because in other larvae the number and position of the protuberances vary
considerably. No one would suggest that the caudal tubercles and spinous
processes of Dimorphopteryx, Blennocampa, Hypergyricus, Caulocampus,
and others are homologous with the cerci of generalized insects; yet, there
is no reason to assume that the caudal tubercles of these larvae are differ-
ent in origin, structure, and function — whatever that may be — from the
suranal paired processes of the nematid larvae. These protuberances
may or may not be at all related genetically to the suranal median process
of the Cephidae and its allies. At any rate our present knowledge does
not permit any definite conclusion regarding the true nature or homology
of these structures. The interpretation advanced by Middleton (1921)
seems more reasonable. He named these protuberances pseudocerci.

Subanal Appendages. — The larvae of the Pamphiliidae (Figs. 91, 95)
and Cephidae (Figs. 108, 109, 111, 116, 117, 118, 119) possess a pair of
subanal appendages on the ultimate segment, one on each side ventrad
of the lateral ends of the anal slit. These appendages are long, setaceous,
and three-segmented in the Pamphiliidae, but are rudimentary, papilliform,

347] LARVAE OF THE TENTHREDINOIDEA—YUASA 29

and only indistinctly segmented in the Cephidae. When the appendages
are long and setiform, the relative length and color of the segments differ
in different species. In the Cephidae the appendages may or may not be
provided with accompanying setae near the proximal end, and these
setae may or may not form a continuous group with the setae on the
sternum.

That these structures are true appendages of the segment is indicated
by the segmentation and by the fact that they are invariably articulated
at the proximal end against the surface, not being mere protuberances
like the caudal processes of the tergum. Some embryologists consider these
appendages to be homologous with the cerci of generalized insects and the
anal "prolegs" of lepidopterous larvae, and, therefore believe them to be
true appendages of the eleventh abdominal segment. It is obvious that
they can not be homologous with the anal larvapods of other tenthredinoid
larvae. Crampton (1919) designated them as arthrostyli on the ground
that they are apparently homologous with the styli of the Ephemeridae
and other insects. The opinions of entomologists differ, for example
Middleton (1921) homologizes the subanal appendages with the post-
pedes of Crampton. The homology and function of these appendages
need further investigation. However, it is significant that long distinctly
segmented appendages should occur in the Pamphiliidae and rudimentary
ones in the Cephidae.

Larvapods. — The embryological data seem on the whole to support
the view expressed by Korschelt and Heider (1899), who say that "the
abdominal appendages of the caterpillars of the Lepidoptera and Hymen-
optera are to be regarded as true limbs," and that "limb-rudiments first
form on all or most of the abdominal segments, but they very soon dis-
appear on those segments which in the larvae have no limbs, while on the
other segments they are transformed into the functional prolegs." Graber
(1890) has shown that the so-called anal prolegs of Hylotoma are the
appendages belonging to the tenth or true penultimate abdominal segment.
They are, therefore, not homologous with the anal "prolegs" of lepidop-
terous larvae, which are, according to Graber (1890), the appendages of
the ultimate segment.

The maximum number of larvapods {pig) occurs in the Xyelidae, where
each of the abdominal segments is provided with a pair. The first and
ninth pairs may be smaller than the others, as in Odontophyes, but they
are always discernible. The number of larvapods present in the Tenthredin-
idae varies from six to eight pairs. They are usually present on abdominal
segments 2-7 and 10 or 2-8 and 10, rarely on 2-6 and 10. In the Fenusinae
(Fig. 105) and Caulocampus the tenth pair is obsolete, and in Metallus
(Fig. 103) they are fused together forming a median protuberance.
Larvapods are entirely wanting in the other families of the Tenthredinoidea.

30 ILLINOIS BIOLOGICAL MONOGRAPHS [348

f A typical larvapod is a fleshy subconical protuberance narrowed toward
the distal end, and is usually subdivided into a larger but shorter proximal
portion and a smaller but longer distal portion. Sometimes the distal
end is dilated and turned mesad, as in Neodiprion (Fig. 82), or the cephalo-
ventral angle is pointed, as in Tenthredo. The larvapods are well devel-
oped in most of the free-living larvae but in the leaf-miners and
fruit-borers they are reduced and smaller. They are very small in the Hylo-
tominae and rudimentary in the Fenusinae (Fig. 86) and Schizocerinae,
while they are obsolete in Phlebatrophia. The degree of development of
the larvapods is closely correlated with the habits of the larva. The
larvapods are usually located on the middle of each lateral half of the
sternum but occasionally are very close together near the meson, as in
Neodiprion. The number of pairs of larvapods present is a convenient
character for dififerentiating the subfamilies of the Tenthredinidae. The
larvapods often bear a few setae on the cephalic and lateral aspects.
The setae, when present, are confined to the mesal aspect. Lack of setae
on the larvapods is often a generic character and the number and arrange-
ment of the setae is typical of the species.

Crampton (1919), with good reasons, proposed to substitute the term
uropods for the long-used but misleading term prolegs. The term uropoda
has been employed by students of the Crustacea in designating the abdom-
inal appendages, especially one of the posterior pairs of pleopods, and
according to Smith's glossary of Entomology the term refers to "any of
the abdominal feet of Arthropoda." These facts indicate the necessity of a
distinctive term, and the new term larvapods following the suggestion of
Dr. MacGillivray, is used until a happier term is created for these true
abdominal appendages of insect larvae.

Metamerism. — Graber (1890) has shown that the number of somites
which compose the body of the larvae of Hylotoma is fourteen exclusive
of the head. The first three somites belong to the thorax and the remaining
eleven to the abdomen. The ultimate segment, or the telson of the embry-
ologists, is difiScult to discern in larvae. It is probably represented by the
suranal and subanal lobes of the larvae, but the boundary between this
segment and the tenth somite is so obliterated, and the ultimate segment,
which is originally much smaller than the preceding somites, is in the
larval stage so much more reduced, that it is permissible and also con-
venient to speak of the abdomen as being composed of ten segments.
For this reason the tenth abdominal somite, which bears the so-called
anal prolegs, is designated as the ultimate segment in this paper. It is to be
noted that Nelson (1915:111) considers that there are eleven segments
and a telson in the abdomen of the embryos of Hymenoptera. In all the
larvae of the Tenthredinoidea examined, it is always possible to count
ten abdominal segments. The body is usually distinctly segmented.

349] LARVAE OF THE TENTHREDINOIDEA—YUASA 31

The exact limit of the somite in a larva is not easy to determine.
Entomologists seem to have paid little or no attention to this point.
Castle (1900) has made an excellent study of the metamerism of the
Hirudinea, but his conclusions are not directly applicable to the case of in-
sect larvae tho they are pregnant with important suggestions. He found
that the natural and true limits of a somite coincide with the limits of the
neuromere, and that both reduction and increase in the number of rings,
which correspond to the annulets in the saw-fly larvae, take place at the
ends of the segment. The classical work of Lyonet (1762) and the recent
study of Forbes (1914) on the musculature of lepidopterous larvae, as also
a study of Boving (1914) indicate that the musculature affords a reliable
criterion for determining the limits of a somite. From a careful examina-
tion of the musculature of various types of tenthredinoid larvae (Fig. 129),
the author has come to the conclusion that in these larvae the natural
and accurate determination of the extent of the somites composing the
body is best based upon the musculature. A detailed discussion of this
subject is out of place here. It is sufficient to say that a majority of the
longitudinal muscles, including the dorsal, lateral, and ventral retractor
muscles, originate on the cuticular fold, or coria, which, on the exterior, is
usually indicated by a deep depression (is). Only a few muscles of impor-
tance cross this fold, nearly all the muscles being attached either to the
cephalic or caudal part of the coria. This cuticular fold, therefore, is con-
sidered as the cephalic limit of the somite, and the annulets into which
the somite is subdivided are numbered consecutively, commencing at its
cephalic end. To assume, a priori, the spiracular annulet, or the annulet
which bears the spiracle, to be the first annulet is arbitrary and inaccurate
inasmuch as this annulet, according to the criterion of musculature, cor-
responds to any one of the first three annulets of the somite. The position
of the spiracular annulet is constant and definite within a genus or sub-
family as long as the number of annulets of the somite is constant. Some
of the annulets are usually setiferous and often bear in addition transverse
rows of glandubae. The position of such setiferous annulets is constant within
the genus or subfamily when the annulation is constant. The number of
annulets on the ninth abdominal segment is always smaller than that on
the preceding segments. Since the setiferous annulets have a definite
order, it is possible to determine which annulets are obsolete on the penul-
timate segment. The first annulet to disappear is a caudal one, and
ordinarily it is only the caudal annulet that is missing. The number of
annulets of the sternum is less than that of the tergum. The length of
the annulets varies but their relative size is constant within a species.
The primitive number of annulets of the abdominal segments is unknown.
If the annulation of the generalized Tenthredinoidea is assumed to be
representative of the primitive condition, then four is the primitive num-

32 ILLINOIS BIOLOGICAL MONOGRAPHS [350

ber of annulets. In the specialized Tenthredinoidea the number varies
from one to seven, but five to seven is of the most common occurrence.
The number becomes smaller in the highly specialized families, being
reduced to a single annulet in the Siricidae and Orysidae.

Spiracles. — The spiracles {spi) are present on the prothorax and the
first eight abdominal segments in all Tenthredinoidea. The pair on the
prothorax are always the largest. The abdominal spiracles are usually
uniform in size and shape except the last pair, which are often larger than
the others. The spiracles are definite in location in regard to the annulets
and are always situated on some one of the first three annulets of the
segments. The spiracular line is usually located slightly ventrad of the
middle of the lateral aspect of the body, but sometimes it migrates ven-
trad to the latero-ventral line as in Caliroa. The spiracles (Fig. 155) are
usually very simple in structure, vertical in position, never circular in
outline but narrowly ovate, rounded or pointed at both ends. The peri-
treme is narrow but strongly chitinized and brownish or blackish. The
labiae are narrow and the spiracular opening is usually closed and appears
like a dark line. The peritreme is sometimes distinctly thickened as in the
Hylotominae. There is often a semicircular or irregular chitinized colored
area on each side of the spiracles, as in certain nematid genera. These
areas vary in size and shape but are constant within species, and their
presence is usually constant within genera and often within a subfamily.
When these areas are present, the spiracles are said to be winged.

The true prothoracic spiracles are considered as wanting in adult and
larval insects. The spiracles found on the prothorax of Tenthredinoid
larvae are the mesothoracic spiracles {msp) which have migrated from
the mesocoria onto the prothorax. The metathoracic pair {tsp) is usually
functionless, very small, and located in the metacoria, or obsolete. In the
Cephidae and Siricidae, however, the metaspiracles are distinct, functional,
and as large as the abdominal spiracles. It is difiicult to explain the
rudimentary condition of this pair in the Xiphydriidae, since other charac-
ters indicate that they have a common origin with the Cephidae and
Siricidae. It is possible, however, in the course of evolution, to have one
structure of the body modified faster than another structure.

Setae. — The surface of the body is usually provided with some setae,
particularly on the head, thoracic legs, and the ultimate segment of the
abdomen on the suranal and subanal lobes. The number, size, arrange-
ment, and structure of the setae vary in different taxonomic units, accord-
ing to their location; and to some extent according to the stage of larval
growth. There is a tendency toward the loss of setae in the ultimate
stage or the last instar, as is Pteronidea, or to have fewer and smaller setae
in the leaf-mining and wood-boring larvae, as in the Fenusinae, Cephidae,
and others. There seem to be no definite setal patters, as in lepidopterous

351] LARVAE OF THE TENTHREDINOIDEA—YUASA 33

larvae, but when certain annulets of the segments possess setae their pres-
ence on these annulets in successive segments is constant if not in precisely
the same number and order. There is a tendency to have more and longer
setae on the lower half of the head than on the upper half. The number
and arrangement of the setae are variable on the vertex and front but are
fairly constant on the clypeus, labrum, and mandibles.

A spinal formula is an abbreviated expression of the arrangement of
the tubercles or spines on the various parts of the body. The figures of
the formula indicate the number of branches of a spine and are arranged in
order, beginning with the mesal spine in the case of those on an annulet
and with the cephalic spine in the case of the subspiracular areas. The
spinal formula of the prothoracic segment represents the arrangement of
the spines on the large or second annulet, on the first or smaller lateral
annulet, on the subspiracular area, and on the postsubspiracular area
respectively. The spinal formula of the third abdominal segment indicates
the arrangement of the spines on the first tubercle-bearing (usually 2d)
annulet, on the next small annulet if this is present, on the third tubercle-
bearing (usually 4th) annulet, on the subspiracular area, and on the post-
subspiracular or surpedal area, respectively.

The number of branches of the spines sometimes varies and the arrange-
ment of the spines also may show minor variations. The spinal formulae
represent the most typical arrangement.

Glands and Glandubae. — There are many types of glands opening to the
exterior found on the various parts of the body of the larvae of the Tenth-
redinoidea. The larvae of the Nematinae and Cladiinae are provided with
a series of ventral glands on the ventro-meson of abdominal segments
1-7, Sometimes a pair of eversible glands is found in the cervical region,
as in Megaxyela major. The larvae of the Cimbicinae possess a spiracular
gland located dorsad of each spiracle of abdominal segments 2-8, It is from
these glands that the yellowish fluid of these larvae is poured out when
disturbed. A peculiar sucker-like protuberance with a depressed center
occurs in the larvae of the Acordulecerinae on the sublateral area of
abdominal segments 2-4 or 5 and 8. The function of this structure is not
known but it is not improbable that it is secretory in nature. The wax
glands of the wax-secreting larvae such as certain Tenthredininae, Emphy-
tinae, Selandriinae, etc., are minute and located on various parts of the
body of these larvae, but their detailed structure has not been studied.
The most common type of these glands is found in the larvae of the
Diprioninae, Emphytinae, Selandriinae, Tenthredininae, some Nematinae,
and others. The cutaneous glands of these larvae are provided with chi-
tinized rings about their external openings. These chitinized openings are
known as glandubae. They may be located at the end of tubular protu-
berances, and in such cases they are spoken of as being stalked, or they

34 ILLINOIS BIOLOGICAL MONOGRAPHS [352

may be found flush with the body surface, when they are spoken of as being
sessile. The glandubae are especially conspicuous in the larvae of the
Diprioninae and Pachynematus. The slime-glands of Caliroa have semi-
sessile glandubae which are few in number. The glandubae are constant
in their type, presence, general arrangement, and location, within genera
and subfamilies.

Formulae of Segmented Appendages. — For convenience in designating
the size and relationship of various segmented appendages, the resort
has been made to various so-called formulae. The segments of an appen-
dage are numbered, beginning with the proximal segment. In a formula
numbers of the segments are arranged in the descending order of magni-
tude, and those of equal dimensions are placed in a parenthesis. For
example, the expression "antennal formula: (2,5), 3, 4, 1" shows that the
antenna is composed of five segments and that segments 2 and 5 are equal
in length but longer than the others, and that segment 4 is shorter than
segment 3 but longer than the first or proximal segment.

353] LARVAE OF THE TENTHREDINOIDEA—YUASA 35

III. TAXONOMY

Strictly speaking no classification of the Tenthredinoidea based upon
larval characters has hitherto been proposed. Attempts have been
restricted to the characterization of the different subdivisions included in
the superfamily. Among the earlier writers Le Peletier's (1823) work may
be mentioned. After a brief general account of the larvae, he gave a list
of eighteen divisions in which he grouped the species of the Tenthredinoidea
and stated whether the larvae of each division were known or unknown
and, if known, the number of the thoracic and abdominal legs present.
It is interesting to note that he mentioned a group of larvae the body of
which he characterized as "donkey-form" (aselliform) which he was
unable to place in any of his divisions. Dahlbom (1835) published careful
descriptions and a synopsis of larvae of sixty-three species. A synoptic
table for the larvae was compiled by Westwood (1840) from this work
and was published with additions. The characters used are the number of
abdominal legs and the feeding habits of the larvae. Norton (1867)
republished Westwood's table without additions. In the table given by
Cameron (1882) the larvae of more than ninety-five species are included.
The major groups are separated on the number of thoracic and abdominal
legs present. These subdivisions are segregated on biological characters
such as reflex bleeding, types of cocoons, and, finally, genera and species,
when known, are separated on the coloration, setae, food-plants, and feed-
ing habits. In 1895 Dyar, the most prolific and the only important Ameri-
can writer on the larvae of the Tenthredinoidea, published "A recognition
table for the known sawfly larvae of the North Atlantic States." The
larvae of one hundred and twenty-six species including forty-one not
specifically identified were considered in this synopsis. The characters
used are the number and location of abdominal legs, types of cocoon,
feeding habits, food-plants, and coloration. This last character was
employed extensively in separating different species. The next attempt
along this line was undertaken by Chester Young (1898), who was the
first to take into consideration the structural characters of the appendages
of the head. Unfortunately this work remains unpublished, but it is on
file as a baccalaureate thesis in the library of Cornell University. Konow
(1901) summarized the taxonomic information concerning the known
larvae of European and American species, four hundred and eighteen in
all, in the form of an analytical table. The presence or absence of abdom-
inal legs, number of antennal segments, and modifications and appendages

36 ILLINOIS BIOLOGICAL MONOGRAPHS [354

of the ultimate body-segment were used in separating the families and
subfamilies. The number and location of abdominal legs, the food-plants,
types of cocoon, and coloration furnished the basis for the separation of
tribes, genera, and species. Middleton (1915, 1917) has characterized the
larvae of the genus Dimorphopteryx and of the family Cephidae. It
must be noted that these writers were concerned only in the preparation
of recognition tables for the separation of the particular species they had
in hand, and, with the exception of Middleton, no one has attempted
to construct a synopsis of families, genera, and species as such.

In the following pages, the author has attempted to define and describe,
as far as possible with the materials at hand, families, subfamilies, genera,
and species by the use of larval characters. With a few exceptions, no
attempt has been made to incorporate data from previous writers for the
reason that the characters recorded by them were found in most cases of
little or no value for the present purpose — not because they were inaccu-
rate, altho that was true in many cases, but chiefly because they were not
of specific significance. For example, Dyar's descriptions of species are
usually very accurate and dependable but most of the characters noted
excepting coloration often proved to be only of family or subfamily sig-
nificance. The definitions given here are correct for the materials actually
studied, but it is not surprising if they do not hold good in many cases
when more materials become available for examination. It is obviously
impossible to attain perfection in the face of so many missing links in the
series of genera and species. These missing links will be filled in as rapidly
as accurately identified materials become available, but it must be remem-
bered that absolutely correct identification is only possible, in the majority
of cases, after carrying individual larvae of the species thru to the adult
stage, exuviae being saved for each instar.

In this study the classification of the Tenthredinoidea proposed by
MacGillivray in 1906, with later additions, has been adopted in the main
in arranging and restricting the families, subfamilies, genera, and species.
For generic synonymy, Rohwer's "Genotype of the Sawflies and Wood-
wasps" (1911) has been followed. In this section all references to the
bibliography of the different divisions and subdivisions have been omitted.

355] [ARVAE OF THE TENTHREDINOIDEA—YUASA 37

SUPERFAMILY TENTHREDINOIDEA

Larvae with exposed, well diflferentiated head, trunk consisting of three
thoracic and ten visible abdominal segments; spiracles always present on
prothorax and first eight abdominal segments; antennae and chitinized
dentate mandibles always present; ocellarae, when present, always one on
each side of the head; thoracic legs, when present, always three pairs,
a pair to each segment; larvapods, when present, always six pairs or
more, and except the Xyelidae, are never on the first and ninth abdominal
segments, but always on the second abdominal segment, and never with
crochets; mouth-parts, when normal, with mandibles strongly chitinized
with distinct dentes, dextral dentes differing in number, shape, and
arrangement from sinistral; maxillae with cardo, stipes, palpifer, palpus,
galea, and lacinia present, palpus typically with four segments, galea
conical, digit-like, and lacinia usually flattened, its cephalic margin with a
fringe of setae; labium with submentum, mentum, palpi, stipulae, and
totaglossa, palpi typically with three segments, totaglossa membranous,
bulbose, with a sericos on its meso-distal portion; general appearance of
body caterpillar-like or grub-like; free leaf -feeders, leaf-miners, web-
spinners, leaf-rollers, wood- and stem-borers, and parasitic larvae.

Free Leaf-feeders. — Body caterpillar-like; thorax with well-developed,
distinctly segmented legs, typically with five segments, coxa, trochanter,
femur, tibia, and tarsus and tarsal claw; abdomen typically with a pair of
larvapods on segments 2-7 or 2-8 and 10; ultimate segment sometimes
with caudal protuberances but never with a distinct suranal process or with
subanal appendages; head typically semiglobose; antennae typically mul-
tisegmented, segments one to five in number; ocellarae always present,
usually located dorsad of antennariae; mouth-parts well developed and
typical in structure; abdominal segments usually with five to seven annu-
lets, some of which bear transverse rows of setae and often some glandubae;
head, thoracic legs, and anal area usually setiferous; majority of Tenthre-
dinidae and Xyelidae.

Leaf-miners. — Body somewhat depressed, head sometimes distinctly
depressed and mouth-parts directed cephalo-ventrad; thoracic legs small,
modified, number of segments reduced to four or to one, legs sometimes
entirely fleshy, conical, or mamma-like, with or without tarsal claws;
abdomen with very small larvapods or larvapods nearly obsolete; mouth-
parts sometimes modified, labial and maxillary palpi with reduced number

38 ILLINOIS BIOLOGICAL MONOGRAPHS (356

of segments; annulation sometimes obsolete. A few subfamilies of
Tenthredinidae.

Nest-builders. — Thorax with seta-like segmented legs; abdomen without
larvapods; antennae long, setaceous, seven-segmented; mouth-parts nor-
mal; ultimate segment with distinct subanal appendages and a minute
hook-like caudal process on caudo-meson of tergum; ocellarae present;
web-spining leaf-rollers, Pamphiliidae.

Borers. — Thorax with rudimentary legs, tarsal claws never present;
abdomen without larvapods, ultimate segment with a distinct suranal
process or with a pair of subanal appendages; mouth-parts somewhat
modified, maxillary and labial palpi reduced in number of segments;
ocellarae wanting or with vestigial eye-spots; metaspiracles sometimes
functional, as large as abdominal ones; wood-borers and stem-borers,
Siricidae, Xiphydriidae, and Cephidae.

Parasites. — Body grub-like, thoracic and abdominal legs wanting;
mouth-parts modified, maxillary and labial palpi obsolete; ocellarae
wanting; antennae one-segmented; parasitic larvae, Oryssidae.

The larvae of the typical Tenthredinoidea are readily differentiated
from the larvae of other Entometabola by the presence of a single ocellara
on each side of the head and, usually, six or more pairs of larvapods, none
of which are provided with crotchets, and never occur on first and ninth
abdominal segments. The characteristic mouth-parts include four-
segmented maxillary palpi and three-segmented labial palpi. The character
of the antennae, the number of ocellarae and larvapods, the charac-
ter of the mouth-parts, especially maxillary and labial palpi, and the
presence of thoracic legs distinguish the leaf-miners and wood-borers of
the Tenthredinoidea from other leaf-miners and wood-borers, such as
certain Lepidoptera, Coleoptera, and Diptera. The larvae of Hymenop-
tera other than Tenthredinoidea are distinguishable from those of the
latter as follows: They are apodous, thoracic and abdominal legs being
always wanting; the mouth-parts are vestigial, maxillary and labial
palpi, if present, papilliform, never distinctly segmented; ocellarae are
never present; and suranal process and subanal appendages are always
wanting. The larvae of the Oryssidae are separable from other hymen-
opterous larvae on the basis of the characters used in the definition of that
family elsewhere.

FAMILIES OF TENTHREDINOIDEA

1(6) Thoracic legs present, either normal in form, distinctly segmented, or modified, if
modified, fleshy or conical, if conical, head and body distinctly depressed; larvapods
either present or wanting 2.

2(5) Thoracic legs normal in form, not seta-like, rarely mamma-like; larvapods usually
present; subanal appendages wanting; antennae usually with less than seven seg-
ments 3.

357] LARVAE OF THE TENTHREDINOIDEA—YUASA 39

3(4) Larvapods present on all abdominal segments; antennae with six or seven segments.

XYELIDAE.
4(3) Larvapods not present on all abdominal segments; antennae never with more than five

segments TENTHREDINIDAE.

5(2) Thoracic legs seta-like; larvapods wanting; subanal appendages present, setaceous;

antennae very long, with seven segments PAMPHILIIDAE.

6(1) Thoracic legs vestigial, not distinctly segmented, mamma-like or wanting, if mamma-
like, head and body never distinctly depressed; larvapods wanting 7.

7(12) Thoracic legs present; ultimate segment with suranal process 8.

8(9) Subanal appendages present, vestigial, papilliform; oceilarae present; antennae

with four or five segments CEPHIDAE.

9(8) Subanal appendages wanting; oceilarae wanting 10.

10(11) Antennae with three segments; metaspiracles functionless, very much smaller than

abdominal spiracles 11.

XIPHYDRIIDAE.
11(10) Antennae with one segment; metaspiracles functional, as large as abdominal spir-
acles SIRICIDAE.

12(7) Thoracic legs wanting; ultimate segment without suranal process and subanal
appendages ORYSSIDAE.

MacGillivray (1906) divided the superfamily Tenthredinoidea into
nine families. They are the Xyelidae, Pamphiliidae, Blasticotomidae,
Tenthredinidae, Xiphydriidae, Siricidae, Megalodontidae, Cephidae, and
Oryssidae. The first two families constitute his Generalized Tenthredinoi-
dea and the last six his Specialized Tenthredinoidea. Since the Blastico-
tomidae and Megalodontidae belong to the Palearctic fauna and are not
represented in North America, they are omitted from the foregoing table.

Family Xyelidae

Larvae (Fig. 6) of medium size, length 13-18 mm.; body caterpillar
like, subcylindrical, flattened on the ventral aspect, uniform in diameter'
except last two segments which are suddenly constricted, stout; segmenta-
tion and usually annulation distinct; cuticle smooth, tuberculate and
setiferous, but never slimy; color greenish, yellowish, whitish, or brownish;
tubercles, when present, brownish or blackish and setiferous; prothorax
sometimes with a pair of lateral eversible cervical glands in the cervacoria;
head circular in frontal contour, moderately large, width usually more
than one-half the diameter of the thorax; mouth directed ventrad; head
slightly overlapped by the prothorax, if any setae, sparsely and inconspi-
cuously setiferous; antennae long, conspicuous, with six, sometimes
seven, segments; ocularia about one-fifth the diameter of the antennaria
and located caudo-dorsad of it, elevated, oceilarae very small; epicranial
suture and vertical furrows present; mouth-parts normal in form; pro-
thorax with a large, often colored, shield-like area on the dorsum and
lateral aspects; legs in comparison with the size of the body very small,
normal in form, all three pairs subequal in size; larvapods present on all

40 ILLINOIS BIOLOGICAL MONOGRAPHS [358

abdominal segments including the first and ninth, where they are some-
times reduced in size; typical segments with four annulets; spiracles on the
second annulet; sublateral lobe produced ventrad, modified into a triangu-
lar lobe laterad of larvapod which it resembles in form; ninth abdominal
tergum with three annulets; tenth abdominal tergum constricted distinctly
and transversely on its cephalic fourth and with a distinct hump-like protu-
berance on the meson caudad of the cephalic constriction, concolorous with
the head and setiferous tuberlces; anal larvapods and ventral or subanal
lobes distinctly large, contiguous, forming a trilobate prominence on the
meson of the tenth sternum; subanal lobe with a pair of setiferous protu-
berances dorsad of larvapods; insects single-brooded, solitary, chiefly
exposed-feeders; pupate in earthen cells in the ground.

The Xyelidae is a small family consisting of seven genera and of
a limited number of species, most of which belong to the North American
fauna. The adults are readily distinguished from all other Hymenop-
tera by the presence of the free part of the vein R2 in the wings. On
venational characters, MacGillivray (1906) considers the members of this
family to be the most generalized Hymenoptera known, having, "departed
from the type of the wing assumed for the original progenitor of the
Hymenoptera only in the loss of the free part of vein Cu2." The genera,
at the same time, possess many features of prominent progressive speciali-
zations which have proceeded in each case in a different sequence so that
a linear arrangement of the genera does not express their true affinities.

Over twenty-five species have been reported from boreal America.
Of this number, four species belonging to as many genera have been recog-
nized in the larval state. Another unidentified species, feeding on pecan,
is added in this paper. Dyar (1898) described the larvae of Megaxyela
major and Xyela minor and gave a definition of the family based on charac-
ters found in these species. He pointed out that they are most nearly
related to the Pamphiliidae. The larvae of Odontophyes aviingrata were
described by the same author (1899). Konow (1901) overlooked Dyar's
1898 paper and gave in his analytical table for the larvae Odontophyes
aviingrata as the sole representative of the subfamily Xyelini. That
Konow was unfamiliar with any xyelid larvae may be reasonably assumed
from the fact that he classified them with those larvae which had no
larvapods and that he placed a question mark before the analytical item
"ohne Afterborsten." The larvae of Pleuroneura, Paraxyela and Proto-
xyela are unknown. The described larvae feed on the foliage of hickory,
butternut, pecan, elm, and the staminate flowers of pine.

The most important materials that I had in the study of this family
were received from Professor R. W. Harned, of the Mississippi Agricultural
College, but, altogether, the material at hand is so limited that it does

359] LARVAE OF THE TENTHREDINOIDEA—YUASA 41

not permit a characterization of the genera. The following key will serve to
separate the species:

1(8) Larvapods present on all abdominal segments, those on the first and ninth segment
sometimes rudimentary; thoracic legs normal in form; body tuberculate, tubercles
setiferous, concolorous with the head; head creamy or brownish or blackish; abdom-
inal segments typically with four annulets, first annulet smooth, non-tuberculate,
crescentic, and confined to the dorsal aspect, three following annulets convex, with
transverse row of setiferous tubercles; not on staminate flowers of conifers 2.

2(7) Head daik-colored, brownish or blackish; tenth abdominal tergum always and
prothoracic and ninth abdominal terga usually with dark-colored patches; tenth
tergum brownish or blackish; subanal lobe with a pair of wart-like brown tubercles
which bear blackish brown setae on the dorsal and lateral aspects, less densely on
the ventral aspect; dorsal tubercles arranged typically as follows: the second and
third annulets with four to five, the fourth annulet with two or three; the subspiracu-
lar and surpedal lobes, each with a tubercle; the dorsal tubercles of the second annu-
let with one to three setae 3.

3(6) Tubercks of typical abdominal segment arranged as follows: second and third
annulets with four tubercles above the spiracular line, each tubercle bearing one
seta, the fourth annulet with two tubercles, one on the spiracular line with two setae,
the other, dorsad of the line, with one seta; on hickory and pecan 4.

4(5) Prothoracic tergum with a large dark brown, median patch whose lateral margins
converge toward the cephalic margin, which is one-half as long as the caudal margin;
the ninth abdominal tergum with a large dark brownish patch on each side of the
meson, the patches converging toward the caudal margin so that the caudal halves
are nearly confluent on the meson with a pair of brownish tubercles btween the
cephalic halves; tenth abdominal tergum almost completely dark browTi in color;
the first annulet of typical segment v/ith the dorsal pair of tubercles quadrate, one-
half as long as the annulet; abdomen with a brownish, cloudy, longitudinal dorso-
lateral line involving the dorsal pair of tubercles and extending usually from the

second abdominal segment to the seventh; on hickorj^ and pecan; Y-226, G

Megaxayela major Cresson.

5(4) Prothoracic tergum without a large dark brownish median patch, but with a pair of
small blackish patches distinctly separated on the meson of the first and second
annulets, the cephalic pair larger, triangular, and their apices directed laterad, the
caudal pair subquadrate, further apart than the cephalic pair; the ninth abdominal
tergum without a pair of large dark brownish patches but with two pairs of small
blackish patches, distinctly separated on the meson, the cephalic pair smaller and
slightly further apart than the caudal pair; tenth tergum blackish, with its cephalic
constricted portion pale or creamy; first annulet of typical abdominal segment with
the dorsal pair of tubercles subcircular, one-third as long as the annulet; abdomen

without a brownish, cloudy longitudinal dorso-mesal lines; on pecan; Y-227

Megaxyela sp. 1.

6(3) Tubercles of a ty'pical abdominal segment arranged as follows: second and third
annulets with four to five tubercles above the spiracular line, each tubercle bearing
two setae; the fourth annulet with three tubercles, one on the spiracular line and the
other two dorsad of it, each tubercle with two setae; on hickory and butternut;
Y-228 Odontophyes aviingrala D\-ar.

7(2) Head light in color, creamy white or pale brown; the prothoracic and the ninth
and tenth abdominal terga without dark-colored patches; tenth tergum light brown;
subanal lobe with a pair of wart-like creamy tubercles which bear long light brown

42 ILLINOIS BIOLOGICAL MONOGRAPHS |360

setae, uniformly distributed on all aspects; tubercles arranged as follows: the second
and third annulets with four tubercles dorsad of the spiracular line, the second
tubercle dorsad of the ventral one sometimes rudimentary and often represented by
a single tiny seta, the fourth annulet with two tubercles, one on the spiracular line
and the other dorsad of it, the sublateral area divided into three lobes and each
with one tubercle; the dorsal pair of tubercles of the first annulet with four to five

setae; on Ulmus; G Macroxyela ferruginea Say.

8(1) Larvapods very small; thoracic legs rudimentary; body not tuberculate; head
creamy whi te ; abdominal segments typically with three annulets ; on staminate flowers
of pine (Dyar in 1898 described no larvapods) Xyela minor Dyar.

Family Blasticotomidae

The Blasticotomidae contains a single genus and species, Blasticotoma
filiceti Klug, which is confined to central and eastern Europe. It is an
archaic type. The systematic position of this unique species has been
considered differently by practically every writer who has studied it.
MacGillivray (1906) has shown, however, that it is in certain of its charac-
ters closely allied to the Xyelidae and Pamphiliidae, while in others it
approximates the Tenthredinidae, and that, it is intermediate in position
between these two groups.

Because of its taxonomic position, it is highly desirable to know the
characters of the larvae of this species, but unfortunately the literature is
void of information in regard to the immature stages, and this interesting
quest must await future discoveries.

Family Tenthredinidae

Larvae (Figs. 7-25) very small to very large, length 10-40 mm.; cater-
pillar-like, leaf -feeders, leaf-miners, or fruit-borers; body cylindrical,
thorax usually largest in diameter, body tapering caudad, sometimes
flattened on the ventral aspect, leaf-miners depressed; greenish or variously
colored with or without distinct markings; smooth, glabrous, setiferous,
tuberculate, or spinous; segmentation usually and annulation sometimes
distinct; third abdominal segment with 6, 7, 5, 4, 3, or 2 annulets, men-
tioned in the order of frequency; some of annulets usually setiferous and
often with glandubae; thoracic legs always present, usually well developed,
typically with five segments, sometimes with three, four, or six segments,
but always with distinct tarsal claws; legs rarely rudimentary, fleshy,
indistinctly segmented, and without tarsal claws; larvapods present
usually on abdominal segments 2-7 and 10 or 2-8 and 10, occasionally the
seventh and tenth pairs wanting, rarely with all larvapods obsolete; head
typically semiglobose, setiferous, with or without distinct markings or uni-
formly brownish, blackish, or greenish; antennae always present, never with
m,ore than five segments; ocellarae always present, one on each side;

361] LARVAE OF THE TENTHREDINOIDEA—YUASA 43

maxillary and labial palpi typically with four and three segments respec-
tively, never obsolete, number of segments rarely reduced; clypeus usually
with two or three setae on each side; mandibles usually with one to four
setae; tenth abdominal tergum without suranal process and sometimes
with caudal protuberances; subanal appendages never present; epicranial
suture and vertical furrows usually present; metaspiracles functionless,
obsolete, or very much smaller than abdominal spiracles; various glands
sometimes present.

The family Tenthredinidae according to MacGillivray contains twenty-
four subfamilies of which five are not represented in the Nearctic
fauna. The subfamilies found in the United States and Canada are as
follows: Diprioninae, Emphytinae, Selandriinae, Dolerinae, Phyllotominae,
Lycaotinae, Tenthredininae, Cimbicinae, Hoplocampinae, Dineurinae,
Monocteninae, Cladiinae, Nematinae, Blennocampinae, Fenusinae, Sco-
lioneurinae, Hylotominae, Schizocerinae, and Acordulecerinae. Of these,
Lycaotinae and Dineurinae have not been available for study.

SUBFAMILIES OF TENTHREDINIDAE

1(42) Thoracic legs normal in form, five-segmented; if modified, tarsal claws always

present; larvapods usually well developed 2.

2(23) Larvapods present on abdominal segments 2-8 and 10; antennae elongate, conical,

usually with five segments 3.

3(20) Thoracic legs with five segments, normal in form 4.

4(11) Third abdominal segment with six annulets on dorsum 5.

5(10) Antennae conical, with five segments 6.

6(9) Labrum bilaterally symmetrical; legs with tibia shorter than femur, tarsal claws

short, strongly curved 7.

7(8) Body rather slender, tapering caudad, without small distinct tubercles; tenth
abdominal tergum without small tubercles; head never shiny, jet-black, body never

yellowish white EMPHYTINAE (in part).

8(7) Body rather robust, uniform in diameter thruout, with small distinct tubercles; tenth
abdominal tergum with several small protuberances, if without, head shiny, jet-
black, body yellowish white BLENNOCAMPINAE (in part).

9(6) Labrum not bilaterally sjonmetrical, but distinctly asymmetrical; legs with tibia

longer than femur, tarsal claws slender, only slightly curved DOLERINAE.

10(5) Antennae not conical, with three segments, the third segment erect and peg-like.

DIPRIONINAE.

11(4) Third abdominal segment with more or less than six annulets on dorsum 12.

12(19) Third abdominal segment with seven annulets on dorsum; body without conspicuous

branched spines or tubercles 13.

13(18) Antennae conical, with five segments; labrum without secondary longitudinal

sutures 14.

14(15) Larvapods setiferous; clypeus with three setae on each side; mandible with two
setae; labrum without a median longitudinal depression. .SELANDRIINAE (in part).

EMPHYTINAE (in part).

15(14) Larvapods glabrous; clypeus with two setae on each side; mandible with 1-4 setae;

labrum with or without a median longitudinal depression 16.

44 ILLINOIS BIOLOGICAL MONOGRAPHS [362

16(17) Legs with tibia minute, distinctly shorter and smaller than femur; maxillae with

stipes without cephalo-ventral triangular projection; mandibles with two setae;

labrura without a median longitudinal depression 17.

SELANDRIINAE (in part).
17(16) Legs with tibia large, usuall}^ subequal to or longer than femur; maxillae with stipes

with cephalo-ventral triangular projection; mandibles with 1, 2, or 3-4 setae; labrum

with or without a median longitudinal depression TENTHREDININAE.

18(13) Antennae not conical, with one segment; labrum with secondary longitudinal

sutures; small but distinct crescentic glandubae dorsad of spiracles. CIMBICINAE.
19(12) Third abdominal segment with five, rarely three or four, annulets on dorsum;

body with conspicuous branched spines or tubercles. BLENNOCAMPINAE (in part).

20(3) Thoracic legs with four segments, modified .21.

21(22) Tenth urotergum and prothoracic and mesothoracic tergites with conspicuous fleshy

pointed protuberances; body not tadpole-like EMPHYTINAE (in part).

22(21) Tenth urotergum and prothoracic and mesothoracic tergites without conspicuous

fleshy pointed protuberances; body often distinctly tadpole-like.

PHYLLOTOMINAE (in part).
23(2) Larvapods on abdominal segments 2-7 and 10, rarely on segments 2-7 or 2-6 and

10 24.

24(39) Thoracic legs with five segments, normal in form; larvapods on segments 2-7 and

either with or without anal larvapods 25.

25(36) Larvapods present on the ultimate segment, either normal and separated or fused

on the meson, forming a single prominence 26.

26(35) Anal larvapods normal and separated 27.

27(28) Antennae with five segments; third abdominal segment with six annulets; tenth

abdominal tergum with several caudal protuberances. HOPLOCAAIPINAE (in part).

28(27) Antennae with four, rarely three, segments; third abdominal segment usually
with less than six annulets; tenth abdominal tergum with or without caudal protu-
berances 29.

29(32) Abdominal segments 1-7 on ventro-meson with an eversible gland; body often with
numerous conspicuous setae, setae arising from distinct tubercles; antennae with four
segments 30.

30(31) Body with numerous conspicuous multisetiferous tubercles, each tubercle bearing
several long setae, some of which are distinctly longer than others; third abdominal
segment with four annulets, annulet 1 with a transverse row of setae, annulets 2 and
3 with a transverse row of setiferous tubercles; tenth abdominal tergum never with
caudal protuberances altho with numerous long setae; setae barbed .CLADIINAE.

31(30) Body without numerous conspicuous multisetiferous tubercles, if tubercles present,
they do not bear several long setae some of which are distinctly longer than others;
third abdominal segment with varying number of annulets; tenth abdominal tergum
sometimes with caudal protuberances; setae not barbed NEMATINAE.

32(29) Abdominal segments 1-7 on ventro-meson without an eversible gland; body never
conspicuoush^ setiferous; antennae with three or four segments; third abdominal
segment with three or five annulets 33.

33(34) Antennae with four segments; third abdominal segment with five annulets; abdom-
inal segments 2-4 and 8 or 2-5 and 8 without a postsubspiracular sucker-like pro-
tuberance HOPLOCAMPINAE (in part).

34(33) Antennae with one segment; third abdominal segment with three annulets; abdominal
segments 2-4 and 8 or 2-5 and 8 each with a postsubspiracular sucker-like protuber-
ance ACORDULECERINAE.

363] LARVAE OF THE TENTHREDINOIDEA—YUASA 45

35(26) Anal larvapods united on the meson forming a single protuberance; antennae with
one segment; third abdominal segment with two annulets; prothorax often with
dorsal and ventral shields; vertical furrows wanting; head and body depressed,
glabrous SCOLIONEURINAE.

36(25) Larvapods wanting on ultimate segment; vertical furrows wanting 37.

37(38) Antennae with three segments; third abdominal segment with four annulets, annulets
2 and 3 setiferous; tenth abdominal tergum with a caudo-mesal protuberance; body-
not depressed HOPLOCAMPINAE (in part).

38(37) Antennae with 1-2 segments; third abdominal segment with two annulets, annulets
glabrous; tenth abdominal tergum without a caudo-mesal protuberance; body
depressed FENUSINAE.

39(24) Thoracic legs with 3-4 or 6 segments; larvapods on abdominal segments 2-7 and 10
or 2-6 and 10, very small 40.

40(41) Mesothoracic and metathoracic legs with six segments; prothoracic legs with six
segments; larvapods on abdominal segments 2-7 and 10 and occasionally with a
rudimentary eighth pair, or 2-6 and 10; body dilated laterad, sublateral lobe pro-
duced and conspicuous, often with numerous setiferous tubercles. HYLOTOMINAE.

41(40) Mesothoracic and metathoracic legs with three segments; prothoracic legs with
four segments; larvapods on abdominal segments 2-7 and 10 with an occasional
rudimentar)' eighth pair; body not dilated laterad, sublateral lobe not produced
and conspicuous; body never with numerous setiferous tubercles but with minute
protuberances SCHIZOCERINAE.

42(1) Thoracic legs not normal in form, but fleshy, indistinctly four-segmented, tarsal
claws wanting; larvapods vestigial on abdominal segments 2-8 and 10, ultimate

pair united on the meson, forming a single protuberance

PHYLLOTOMINAE (in part).

Subfamily Diprioninae

Larvae (Fig. 7) moderately large, length 18-25 mm.; body cylindrical,
somewhat robust, tapering gradually caudad; segmentation and annula-
tion distinct; third abdominal segment with six annulets, annulets 1, 2,
and 4 or 2 and 4 with setae and glandubae; larvapods on abdominal
segments 2-8 and 10, close together on the meson; thoracic legs normal,
well developed, with five segments; prothoracic legs distinctly smaller than
other legs; color of body usually yellowish or greenish, with grayish, or
brownish stripes or rows of black spots; antennae with three segments, seg-
ments 1 and 2 minute, flat, irregular, incomplete, segment 3 erect, peg-like,
strongly chitinized; head and legs usually with spinous, stiff setae; glandu-
bae prominent and numerous; ventral glands wanting; spiracles not winged;
cuticle microscopically spinulate; larvae feed on conifers.

The subfamily Diprioninae is represented in North America by three
genera, Diprion, Neodiprion, and Monoctenus. The Neartic species
formerly placed in the genus Diprion (Lophyrus) are placed by Rohwer
(1918a) in Neodiprion. Diprion simile Hartig of Europe has recently
become estabhshed in the United States. With the exception of Mac-
Gillivray, systematists agree in associating the genus Monoctenus with
Diprion and its allies.

46 ILLINOIS BIOLOGICAL MONOGRAPHS (364

The three genera studied can be separated as follows:

1(2) Body large, stout, longer than 24 mm.; markings mottled with dark brown and
yellowish irregular spots, without brownish stripes or black spots; setae on head
small, slender, those on genae similar to other setae, not spinous; head blackish.

Diprion Schrank.

2(1) Body smaller, slender, usually shorter than 24 nam. ; markings not mottled, but with
brownish longitudinal stripes or rows of black spots 3.

3(4) Third abdominal segment with annulets 1, 2 and 4 with setae and glandubae; setae

on head stiff, long, spinous, those on genae often very large and spinous

Neodiprion Rohwer.

4(3) Third abdominal segment with annulets 2 and 4 with setae and glandubae; setae on
head microscopic, and very few in number Monocknus Dahlbom.

Neodiprion Rohwer

Larvae rather large; length about 19-24 mm.; body slender, with longi-
tudinal brownish stripes or rows of black spots along subdorsal, supra-
spiracular, and sometimes subspiracular lines; head round in contour,
cephalic and caudal margins parallel in profile; front flattened, subpenta-
gonal, as high as wide; ocularia large; antennaria subequal in diameter to
ocularia, their own diameter apart; labrum semicircular, with small
crescentic median emargination; mandibles sharply dentate, the dextral
with four dentes and the sinistral with five; maxillary palpi large, four-
segmented, segments 1-3 ring-like, successively diminishing in diameter,
segment 4 suddenly and distinctly smaller than the preceding segment,
conical, bluntly pointed; galea chitinized, digit-like, smaller than palpi;
lacinia thick, lobate, bearing a minute triangular blade-like seta on ventro-
mesal angle, a stiff seta on dorso-mesal angle, and a row of three to five
minute setae on the oblique cephalo-mesal margin; labial palpi normal,
segment 2 usually longest, segment 3 conical, suddenly and distinctly
smaller than preceding segment; totaglossa with dorso-cephalic depression
on the meson and with several minute sensory pits; parapharynx distinct,
linguiform, constricted dorsad of the middle by a pair of chitinized pieces;
thoracic legs well developed, normal in form, usually blackish, segments
strongly chitinized, coxae largest, trochanter ring-like, chitinized on the
caudal two-thirds, with a whorl of setae, distad of setae membranous,
femur smaller than trochanter in diameter, entirely chitinized, wider than
long on dorsal aspect, increasing in diameter distad, tibia subequal in length
to femur but smaller in diameter, tarsal claws small, basal portion of claw
undeveloped, all segments of leg membranous on the ventral aspect,
prothoracic legs one-half the size of metalegs, mesolegs slightly smaller
than the latter; third abdominal segment with annulation formula, 2, 1,
(3, 5, 6), 5; spiracle on annulet 2, annulets 1, 2 and 4 with a transverse
row of slender, cylindro-conical glandubae, annulet 2 with a few micro-
scopic setae; substigmatal and surpedal lobes large, with several glandubae

365] LARVAE OF THE TENTHREDINOIDEA—YUASA 4.1

and a few setae; larvapods well developed, distal surface incurved mesad,
rather dilated, not pointed, distance between the pair at the base less
than the length of larvapod, venter with four annulets; tenth abdominal
tergum gradually convex, with glandubae; anal setae numerous, small;
spiracles elongate and oblong.

SPECIES OF NEODIPRION

1(10) Head black, light grayish or brownish, never uniformly reddish brown or orange;
body longitudinally striped or spotted 2.

2(3) Body with four longitudinal rows of black spots along subdorsal and supraspiracular
lines; tenth abdominal tergum with cephalic two-thirds entirely black; spots on each
side of dorso-meson elongate, wider at cephalic end; spots on supraspiracular line
large, subquadrate; spots on prothorax obsolete; body yellowish white ; length, 22 mm. ;
on Pinus; Y-221, G-573, C-1, 1-4101 abhotii Leach.

3(2) Body without four longitudinal rows of black spots along the subdorsal and supra-
spiracular lines, but with longitudinal colored bands along each side of the dorso-
meson; latus with longitudinal row of independent segmentally arranged blackish
or brownish spots or with continuous brownish bands 4.

4(5) Body with a row of brownish spots along supraspiracular lines, spots segmentally
arranged, one on each segment, often those on middle segments obsolete, sometimes
all spots obsolete; tenth abdominal tergum with a pair of large blackish or brownish
spots which are all sometimes contiguous on meson; subdorsal bands narrower than
the distance between them; larvapods marked faintly along pedal line when supraspir-
acular spots are distinct; head black; length, 23 mm.; G-1686-2, -5. Neodiprion sp, 1.

5(4) Body with broad longitudinal bands along supraspiracular lines, instead of rows of
segmentarly arranged spots 6.

6(7) Subspiracular lines with broad brownish bands; head black; pedal lines also marked
brown; tenth abdominal tergum faintly marked; subdorsal bands much wider than
the distance between them; these bands much lighter in color than those on latus;
body dull greenish; length, 21 mm.; on spruce; G-791 abielis Harris.

7(6) Subspiracular lines without brownish bands; head light brown, brown, or pale
creamy yellow; subdorsal bands very narrow or very wide; pedal lines with or
without bands 8.

8(9) Pedal lines with distinct brownish bands; tenth abdominal tergum unmarked except
along caudal margin; subdorsal bands very much wider than the distance between
them, lighter in color; head blackish or brownish with brown marks on vertex and
front or pale brown; length, 19 mm.; G-156 Neodiprion, sp. 2.

9(8) Pedal lines without distinct brownish bands; tenth abdominal tergum marked,
entirely concolorous with head ; subdorsal bands very narrow, usually narrower than
the distance between them; head light brown or pale creamy white with vertex

shaded grayish; length, 19 mm.; G- 1686-3, -6, -7 Neodiprion sp. 3.

10(1) Head reddish brown or orange; body spotted; subdorsal and supraspiracular lines

with black spots 11.

11(12) Subspiracular and pedal lines with black spots; spots on subdorsal lines tapering
caudad, partly broken between annulets; spots along supraspiracular lines large,
subquadrate; spots on subspiracular lines smaller, sometimes very small but never
obsolete; spots on pedal lines very small, sometimes obsolete; tenth abdominal
tergum with a pair of large black spots which sometimes fuse on the meson; length,
25-28 mm.; G-1554, -1686-8, -593, C-cue-315, C-Young-37 lecontei Fitch.

4S ILLINOIS BIOLOGICAL MONOGRAPHS [366

12(11) Subspiracular and pedal lines without black spots; spots on subdorsal lines small,
often not distinctly tapering caudad; tenth abdominal tergum with a pair of large
black spots; length, 20 mm.; G-133 Neodiprion, sp. 4.

MONOCTENUS DaHLBOM

Larvae rather small; length about 15 mm.; body slender, dorsum
with difiFuse brownish shade or with longitudinal stripes; third abdominal
segment with annulets 2 and 4 with setae and glandubae; head as in Neo-
diprion except that setae are minute and sparse; clypeus and labrum with
two setae on each side; labrum with small crescentic median emargination;
maxillary palpi large, rather slender, segments 1-3 ring-like, subequal in
length but successively smaller in diameter; galeae and laciniae as in Neo-
diprion; labial palpi rather slender, its segments subequal in length; man-
dible with one mandibular seta; antennae with segment 1 complete, very
narrow, oval, segment 2 flat, incomplete, irregular, segment 3 peg-like,
erect; glandubae conical, distinct; setae microscopic; sublateral lobes not
well developed; annulation typically (1, 2), (3, 4, 5, 6), anal setae numer-
ous, short, and minute; telson with glandubae obsolete.

MacGillivray established in 1906 the subfamily Monocteninae for
the genus Monoctenus associating it with the Cladiinae and Nematinae,
thus deviating from the universal practice of regarding the genus Monoc-
tenus as a member of the subfamily Diprioninae or its equivalent. On
the basis of the venation, MacGillivray's contention is quite justifiable,
and it is most interesting to know what larval characters would indicate
in regard to the relationship between Monoctenus and Diprion and its
allies. Marlatt (1887) published notes on the immature stages of Monoc-
tenus unicolor but his descriptions do not touch the detailed anatomy
necessary for the definition of the genus. Recently, however, I was
fortunate enough, thru the courtesy of Mr. Rohwer, to examine a speci-
men belonging to the United States National Museum which Rohwer
considered to belong to a new species of Monoctenus. A careful study
of this larva convinced me that [so far as this species is concerned] there
are no essential differences between the larvae of Monoctenus and those
of the typical Diprioninae to justify the creation of a new subfamily.
For this reason I have followed the universal practice and decided to
treat Monoctenus as a member of the Diprioninae.

Monoctenus n. sp. Rohwer. — Length, 14 mm., head- width 1.5 mm.;
head brownish; body on dorsum dorsad of spiracular lines, segments of legs,
episternum and epimeron, deep brown; glandubae elongate, conical,
minute, brownish at tip; on cedar.

A specimen bearing the label "5419 Sawfly on cedar, Cadek, Mo.,
June 10, 1892."

367] LARVAE OF THE TENTHREDINOIDEA—YUASA 49

DiPRION SCHRANK

As far as known, only one species is represented in North American
fauna. An examination of D. simile shows that this genus is not very
different from the other genera of Diprioninae but may be separated from
them by the characters of the setae and coloration of the head, and certain
minor points. It is not possible to characterize the genus with the material
at hand.

Diprion simile Hartig. — Body robust, length, 25 mm.; latus with a series
of yellowish or whitish spots on a uniformly grayish brown background,
dorso-meson with a narrow yellowish stripe bordered on each side by an
equally narrow grayish brown band; dorso-lateral lines broadly yellowish,
interrupted at each annulet by fine transverse lines; supraspiracular
lines with three yellow spots, size of spots increasing caudad; dorsad of these,
three smaller spots with the middle one largest and subequal to cephalic
spot of supraspiracular lines; three spots on subspiracular lines, the middle
one being the largest; pedal lines with a large spot on each segment;
larvapod with a brownish spot; tenth abdominal tergum and sternum
marked with grayish; the tergum with a deep constriction dorsad of suranal
lobe; head setae small, slender, hair-like, never spinous or stiff; setae on
genae similar to those on front, never stiff and spinous; legs with femur
sometimes longer than wide on the dorsal aspect; head black; body yellow-
ish gray, mottled; G.

Subfamily Emphytinae
Larvae (Fig. 8) small to moderately large, usually greenish, sometimes
striped; body cylindrical, slender, tapering caudad; segmentation distinct,
annulation fine, indistinct; third abdominal segment usually with six, rarely
seven, annulets, annulets 2 and 4 or 1, 3, 5 and rarely 1, 3, and 6 setiferous;
head greenish or brownish; sometimes with spots on vertex and front;
labrum with or without a mesal longitudinal depression, with 4-5 labral
setae on each side of the meson; clypeus with 2 or 3 setae on each side;
mandibles with one seta rarely with two; larvapods on abdominal segments
2-8 and 10, well developed, usually glabrous, rarely setiferous; ventral
glands wanting; glandubae small, conical, on annulets 1 and 3 or rarely
on 2 and 4; tenth abdominal tergum usually setiferous but without paired
caudal protuberances, rarely with conspicuous spines on the caudal margin,
if spines present, then prothorax and mesothorax on dorsum with two and
one protuberances respectively; antennae elongate-conical, with five seg-
ments, segments ring-like; thoracic legs usually normal in structure, with
femur subequal in length to or sHghtly longer than tibia, tibia well devel-
oped and normal, femur with its disto- ventral angle produced; legs, when
modified, short, stout, and trochanter obsolete; mouth-parts normal in form,
spiracles not winged; larvae leaf -feeders.

50 ILLINOIS BIOLOGICAL MONOGRAPHS [368

The Emphytinae is a large subfamily embracing a number of genera
and numerous species. MacGillivray considers this the second subfamily
of his generalized Tenthredinidae and places it between the Diprioninae
and Selandriinae. The larvae of this subfamily are found readily and in
general appearance and habitus resemble very closely the larvae of the
Selandriinae and Tenthredininae, but they can be separated by the number
of annulets, which in this subfamily, with the exception of Hemitaxonus
and Epitaxonus, is six, while in the other two subfamilies it is seven. The
two genera mentioned are characterized by the presence of seven annulets
on the typical abdominal segment and also by the setiferous larvapods,
thus resembling in these two particulars the larvae of the Selandriinae.
It is of interest to note that Rohwer would associate Hemitaxonus with
:;uch genera as Selandria, Eriocampoides, etc., in the tribe Selandriini of
his subfamily Selandriinae. Middleton (1915) has published a definition
of the genus Dimorphopteryx together with a key for the separation of
three species.

The writer has collected a large number of larvae belonging to this
subfamily, but on account of the difiSculty of breeding adults many
species remain unidentified. In the preparation of the synoptic key to the
genera and in discussions following, only bred or otherwise identified
species have been considered, the consequence being that future study may
require much modification in our conception of the various genera dealt
with.

GENERA OF EMPHYTINAE

1(4) Third abdominal segment with 7 annulets; larvapods setiferous 2.

2(3) Larvapods with 5-3-1 setae on cephalic, lateral, and caudal aspects respectively;
thoracic legs with femur longer than or subequal to tibia; labial palpi with segment

2 longer than segment 1 ; maxUlar>' palpi with segments subequal in length

Hemitaxonus Ashmead.

3(2) Larvapods with 8-5-1 setae on cephalic, lateral, and caudal aspects respectively;
thoracic legs with femur shorter than tibia; labial palpi with segments subequal to

each other in length; maxillary palpi with segment 2 longer than segment 1

Epitaxonus MacGillivray.

4(1) Third abdominal segment with 6 annulets; larvapods glabrous 5.

5(26) Tenth abdominal tergum and prothoracic and mesothoracic tergites without con-
spicuous fleshy pointed protuberances; thoracic legs normal in form, with trochanters
distinct 6.

6(9) Annulets 1, 2, and 4 setiferous 7.

7(8) Antennae with segment 5 longest; legs with femur longer than tibia; head usuall}'
without markings; labial palpi, if segments not subequal, segment 2 longer than
segment 1 Empria Lepeletier.

8(7) Antennae with segment 1 longest; legs with femur subequal to tibia; head usuallj'
with markings; labial palpi, if segments not subequal, segment 2 shorter than seg-
ment 1 Parataxonus MacGUlivray

9(6) Annulets 2 and 4 setiferous 10.

3691 LARVAE OF THE TENTHREDINOIDEA—VUASA 51

10(11) Clypeus with three setae on each side of meson; mandibles with two setae; maxillae
with palpifer produced dorsad as a triangular lobe; labrum with a deep median

emargination with a row of secondary setae caudad of the emargination

Erlocampa Hartig.

11(10) CI>-peus with two setae on each side of meson; mandibles with one seta; maxillae
with palpifer not produced dorsad as a triangular lobe; labrum without a deep
median emargination 12.

12(13) Thoracic legs with trochanter longer than tibia. . . .Strongylogaslroidea Ashmead.

13(12) Thoracic legs with trochanter distinctly shorter than tibia 14.

14(15) Body spotted or transversely striped; head dorsad of ocellarae entirely blackish or
brownish; very large and robust larvae Macremphytus ISIacGillivray.

15(14) Body never spotted or transversely striped; head dorsad of ocellarae not entirely
blackish or brownish; smaller larvae 16.

16(17) Labial palpi with segment 1 longer than segment 2; legs with femur always longer
than tibia; tibia usually twice as long as trochanter; body rather robust; head not

marked distinctly with brown; annulet 4 longest on third abdominal segment

Monoslegia Costa.

17(16) Labial palpi with segment 1 shorter than segment 2; legs with femur not alwaj's
longer than tibia; tibia always more than twice as long as trochanter; body rather
slender; annulet 4 not longest on third abdominal segment 18.

18(19) Head entirely pale, no markings; legs with femur longer than tibia; body on dorsum

not shaded darker than the venter; tenth abdominal tergum not marked

Monosoma MacGillivray.

19(18) Head not entirely pale, usually with brown spots or markings; legs with femur
not always longer than tibia, often subequal; body on dorsum sometimes shaded
darker than the venter; tenth abdominal tergum often marked 20.

20(21) Legs with femur always longer than tibia; tenth abdominal tergum usually marked ;
body on dorsum usually shaded darker than the venter Emphytus Klug.

21(20) Legs with femur subequal to or shorter than tibia; tenth abdominal tergum usually
unmarked; body on dorsum not usually shaded darker than the venter 22.

22(23) Tenth abdominal tergum marked with a spot; body on dorsum shaded darker than

the venter; labial palpi with segment 2 longer than segment 1

Unitaxomts MacGillivray.

23(22) Tenth abdominal tergum unmarked; body on dorsum not shaded darker than the
venter; labial palpi with segment 2 subequal to or shorter than segment 1 24.

24(25) Labrum with a distinct median depression Taxonus Hartig.

25(24) Labrum without a distinct median depression Phrontosoma MacGillivray.

26(5) Tenth abdominal tergum and prothoracic and mesothoracic tergites with con-
spicuous fleshy pointed protuberances; thoracic legs not normal in form, with
trochanters obsolete Dimorphopteryx Ashmead.

Subfamily Selandriinae

Larvae (Fig. 9) small to fairly large, length 18-26 mm.; body cylindrical
and gradually tapering caudad; segmentation and annulation distinct and
fine; larvapods on abdominal segments 2-8 and 10; third abdominal
segment with seven annulets, annulets 1, 3, and 5 setiferous, annulets 3
and 5 with glandubae; thoracic legs normal in form except tibia sometimes
very minute; body uniformly greenish, without colored markings; head
with or without brownish spots; antennae with five segments, long, conical;

52 ILLINOIS BIOLOGICAL MONOGRAPHS [370

mouth-parts normal in form, well developed, palpi large; mandibles with
two setae; clypeus with three, sometimes four, setae on each side, rarely
with two; stipes of maxillae with triangular cephalo-ventral projection;
glandubae minute, stalked; spiracles not distinctly winged; larvapods
setiferous, with about ten setae, rarely glabrous.

The Selandriinae includes a limited number of genera. On the basis of
the venation this subfamily is placed next to the Emphytinae where
practically all systematists have placed it. Rohwer's conception of this
subfamily is somewhat different from that of MacGillivray and therefore
he differs from the latter in the disposition of some of the genera. For
example, Hemitaxonus is assigned to the tribe Selandriini while Mac-
Gillivray placed it, together with Epitaxonus, in the Emphytinae. It may
be said that the larvae of these two genera are very closely related to the
Selandriinae and differ from all other Emphytinae in the typical number

of annulets.

GENERA OF SELANDRIINAE

1(4) Thoracic legs with tibia normal in form, never greatly reduced, usually subequal
in length to femur; larvapods setiferous; clypeus with three or four setae on each

side 2.

2(3) Tenth abdominal tergum with brown spots; larvapods usually with ten or more
setae; clypeus usually with three and often four setae on each side; labium deep
brown, with four to six setae on each side; glandubae slightly longer than one-half the
length of adjacent setae; spiracles not winged; larger larvae; length more than
24 mm Thrinax Konow.

3(2) Tenth abdominal tergum not marked with brown spots; larvapods usually with less
than ten setae; clypeus with three, rarely with four, setae on each side; labium pale
brown or whitish with three to six setae; glandubae usually' subequal in length to
adjacent setae; spiracles faintly winged; smaller lai-vae; length less than 24 mm.

Slrongylogaster Dahlbom.

4(1) Thoracic legs with tibia reduced, very much smaller than femur; larvapods glabrous;
clypeus with two setae on each side Sdandria Leach.

Thrinax Konow

Larvae comparatively large, long, length more than 24 mm.; body
slender, finely annulate, uniformly greenish; head and tenth abdominal
tergum with brown markings; thoracic legs normal, tibia and femur
cylindrical, tapering caudad, subequal in length; larvapods with about
ten setae, distributed as follows: 4-5 on cephalic and lateral aspects and
one on caudal; clypeus usually with three and often with four setae on each
side; labrum with four to six setae on each side, deep brown, without
median longitudinal depression; glandubae small, very short, slightly longer
than one-half the length of adjacent setae; spiracles not winged; maxillary
palpi with segments 2 and 4 subequal in length; labial palpi with segment 4
longer than segment 3 ; ninth abdominal tergum with six annulets, annulets
1, 3, and 5 setiferous, annulet 6 a little shorter than the second.

371] LARVAE OF THE TENTHREDINOIDEA—YUASA 53

SPECIES OF THRINAX

Head pale brown with vertex marked with brown on dorsum and caudad of ocellarae,
front with a round brown spot contiguous to ventral apex of the dorsal marking on
the vertex, frontal spot not reaching the fronto-clypeal suture; antennae, mouth-
parts, femur, tibia, and tarsal claw deep browm; tenth abdominal tergum with a
pair of small brown spots; annulation formula, 1, (5, 4, 3), 2, (6, 7); antennae, 5, 4,
3, 2, 1 ; labial palpi mth distal two segments subequal; legs with trochanter, femur, and
tibia with lengths to each other as 12, 15, and 16 respectively; tarsal claws with the
proximal portion shorter than the distal narrow curved portion; length 26 mm.;
wadth of head 2 mm.; on fern; M-18 impressatus Provancher.

Head pale brown with vertex marked with brown on dorsum and caudad of ocellarae,
front with a subquadrate spot contiguous to the ventral apex of dorsal marking of the
vertex, frontal spot reaching the fronto-cl3Tjeal suture; antennae, labrum, mouth-
parts, femur, tibia, and tarsal claw deep brown; tenth abdominal tergum with a
pair of large brown spots; anmJation, 1(5, 4, 3, 1), (6, 7); antennae, 5, 4, 1, 3, 2;
labial palpi with distal segment longer than the preceding; legs with trochanter,
femur, and tibia with lengths to each other as 12, 15 and 16 respectively; tarsal claws
with the proximal portion as long as the distal curved portion; length 25 mm.;
width of head, 2 mm.; on fern; Y-20-3-1, -20-1 pulatus MacGillivray.

Strongylogaster Dahlbom

Larvae small comparatively speaking, length less than 24 mm.; body

slender, finely annulate, uniformly green; head pale or light brown or

sometimes with a few spots; tenth abdominal tergum never distinctly

marked; larvapods usually with less than ten setae; thoracic legs normal in

form, femur and tibia subequal to each other or one longer than the other;

labrum pale brown or whitish, with three to six setae on each side; glandu-

bae usually subequal in length to adjacent setae; spiracles often faintly

winged.

SPECIES OF STRONGYLOGASTER

1(2) Head with blackish brown markings, vertex with a pair of diverging spots over the
vertical furrows directed toward ocellarae and a spot caudad of each ocellara;
vertical markings sometimes faint, sometimes very distinct and large, merging into a
continuous vertical marking; front never with spot; antennae, labrum, and mouth-
parts light brown; tenth abdominal tergum usually without markings, rarely with a
pair of faint spots; annulation, 1, 4, (2,3, 5), (6, 7,) ; antennae, 1, 5, (2, 3, 4); maxillary
palpi, 2, (1, 4), 3; labial palpi with two distal segments equal in length to each other,
shorter than distal segment of maxillary palpi; labrum with three or four setae on
each side; mandible with two, rarely three, setae; legs with trochanter, femur, and
tibia with lengths to each other as 10, 10, and 12 respectively; uropods with about
eight or nine setae, five on cephalic, 3-4 on lateral, and one on caudal aspect; glandu-
bae long, slender, subequal in length to adjacent setae; spiracles not winged; length
18 mm.; width of head 1.8 mm.; on Pteris aquilina; Y-21 annulostis Norton.

2(1) Head without blackish brown markings, uniformly pale brown 3.

3(4) Trochanter distinctly shorter than femur; labial palpi with distal segment as long
as the preceding segment; head uniformly pale; body and legs uniformly green
without markings; annulation, 1, (3, 4, 5), (6, 7, 2); antennae slender, 5, 1, (4, 3, 2);
maxillary palpi, (2, 1), (4, 3), distal segment longer than that of labial palpi; labrum

54 ILLINOIS BIOLOGICAL MONOGRAPHS (372

and clypeus with four and three setae respectively on each side; larvapods with about
6-8 setae, 4-5 on cephalic, 2-3 on lateral aspect; glandubae subequal in length to
adjacent setae; spiracles with faint brown wings; thoracic legs with trochanter, femur,
and tibia with lengths to each other as 8, 13, and 15 respectively; length, 19 mm.;

width of head, 1.6 mm.; on Pteris aquilina; Y-168-4 tacilus Say.

4(3) Trochanter not distinctly shorter than femur; labial palpi with distal segment shorter
than the preceding segment; head uniformly pale, rarely with a pair of faint spots on
dorsal part of vertex; body uniformly green; legs distad of trochanter brownish;
annulation, 1, (5, 4, 3), (2, 6, 7); antennae, slender, conical, (5, 1), 4, (2, 3); maxillary
palpi, 2, (4, 1), 3, distal segment subequal to that of labial palpi; labrum and clypeus
with four and three setae respectively on each side; thoracic legs with trochanter little
shorter than femur, tibia usually almost as long as femur; length, 21-23 nun.; width
of head, 1.8 mm.; on Pteris aquilina ;Y-l8-l, M-32 (in part), M-86. politm Provancher.

Selandria Leach

Larvae comparatively small, length less than 24 mm., usually about
15 mm.; body slender, finely annulate; head pale; body green; tenth
abdominal tergum unmarked; larvapods glabrous; legs very short, with
tibia conspicuously reduced in size, femur distinctly dilated at distal
end, bearing rudimentary tibia on dorsal margin; clypeus with two setae on
each side; mandible with two setae; labrum with three setae on each side;
glandubae very small; spiracles not winged.

Selandria flavipes Norton. — Legs with trochanter ring-like, femur dilated
at distal end, with ventro-mesal projection only slightly narrower in
diameter than trochanter; tibia very small, much smaller in diameter than
femur, appearing as if surrounded by fleshy part of the latter, deep brown
in color; trochanter, femur, and tibia with lengths to each other as 7, 11,
and 6, respectively; annulation, 1, 4, 2, (3, 5, 6, 7); antennae slender, coni-
cal, 5, (1, 2, 4, 3); maxillary palpi, (4, 1, 2), 3; labial palpi with distal
segment subequal in length to segment 1, longer than distal segment of
maxillary palpi; length, 15 mm.; width of head 1.3 mm.; on Pteris aqui-
lina: Y-168, M-70, C-S.f.

Subfamily Dolerinae

Larvae (Fig. 10) moderately large, length, 15-25 mm.; body slender,
cylindrical, tapering uniformly and gradually caudad, either uniformly
greenish or brownish, or dorsum colored darker than venter, never with
bright and distinct patterns; segmentation and annulation distinct; third
abdominal segment with six annulets, annulets 2 and 4 on dorsum seti-
ferous and with glandubae; larvapods on abdominal segments 2-8 and 10;
thoracic legs well developed; head large, as wide as thorax or nearly so;
vertical furrows distinct; antennae five-segmented, conical; labrum
distinctly asymmetrical, dextral part larger than sinistral; head, legs,
larvapods, tenth abdominal tergum, and sternum moderately setiferous;
glandubae present; spiracles not winged; cuticle distinctly, uniformly, micro-

373] LARVAE OF THE TENTHREDINOIDEA—YUASA 55

scopically verrucose on the dorsum and latus between spiracular lines; on
monocotyledonous plants.

The Dolerinae is a well-defined subfamily with a distinct habitus
and is closely allied to the Emphytinae and Selandriinae. The most
important adult character for differentiating the group from other sub-
families of the generalized Tenthredinidae is the coalescence of the cells
Ri and R5 due to the atrophy of the free part of the vein Ro. The sub-
family contains two genera, the old genus Dolerus and the recently de-
scribed genus Loderus. Leach separated, under the name of Dosytheus,
all those species having certain antennal peculiarities and, according
to Stephens, also having bright colors on the abdomen. This differen-
tiation was considered invalid by Hartig and his view was endorsed by
Norton and Cameron. Norton described a species under the name of
Dorytheus apricus var. alhijrons which is now placed in the genus Loderus.
The monobasic genus Pelmatopus of Hartig, based on P. minutus, is now
considered as congeneric with Dolerus. Since the larvae of Loderus are
unknown, the genus Dolerus alone is considered here.

Dolerus Jurine

Head viewed from cephalic aspect circular in contour in mature
specimens, epicranium semiglobose, front distinctly flattened; mouth-
parts directed caudo-ventrad; antennaria never circular, with obtuse
corners at the angles of their dorsal side; antennae with formula, 5, (3, 4,
2), 1, distal segment conical, apex less chitinized and obtusely rounded,
never sharply pointed, segments 2-5 well chitinized, segment 1 narrow but
distinctly larger in diameter than distal segments; front distinctly wider
than high; labrum asymmetrical, dextral part always larger than sinistral
or with pointed ventro-mesal angle; mandible very thick, large, dextral
with four distadentes and one curved sharp proxadentis, sinistral with
four distadentes and mesal surface deeply emarginate; parapharynx with
apex dilated and chitinized; maxillary palpi, galea, lacinia, and labial
palpi normal in structure and well chitinized; thoracic legs with femur
often produced papilla-like on its disto-ventral angle, tibia long, cylindrical,
tapering uniformly distad, distinctly longer than femur, tarsal claw
rather slender and straight; abdominal segments with six annulets, typical
formula, (1, 2), 3, 4, (5, 6), 1 =5-|-6; spiracles on annulet 2; annulets 2 and
4 with conical glandubae and tiny cylindrical truncate setae with large
calices; tenth abdominal tergum semiglobose, anal setae numerous; ventral
glands never present.

The genus Dolerus is represented in North America by more than
thirty species but none of them had been identified in the immature stages
until the writer reared the adult of D. similis Nort. at Ithaca, N. Y. The

56 ILLINOIS BIOLOGICAL MONOGRAPHS (374

larvae of this genus are easily obtained and readily identified because of
the marked asymmetry of the labrum.

The following key will serve to separate the species studied:

SPECIES OF DOLERUS

1(6) Head uniformly pale, creamy, or pale brown; body uniformly whitish or greenish,
without distinct dorsal band 2.

2(3) Body with small black spots on each segment along supraspiracular and pedal lines;
head pale brownish yellow; spots on ninth abdominal segment much smaller than
preceding ones; length, 25 mm.; on wheat and grasses; Y-11 7-1-1, G-d-1. Dolerus sp. 1.

3(2) Body without small black spots on each segment along supraspiracular and pedal
lines; head creamy; length less than 25 mm 4.

4(5) Distance between antennaria and mandibularia subequal to distance between anten-
naria and ocularium; length, 20 mm.; on Carex Irichocarpa; Y-24-5-2, -145-1 (?),-147.

Dolerus sp. 2.

5(4) Distance between antennaria and mandibularia twice or more than twice the distance
between antennaria and ocularium; length of prothoracic spiracles in relation to
vertical diameter of antennaria variable; femur with or without disto-ventral pro-
jection; front with or without pale brown spot; length 18-20 mm.; on grasses, sedge,
timothy; Y-29-11,-32-1, M-7, H,-41-l,-63,-225 Dolerus sp. 3"

6(1) Head spotted, banded, or distinctly brown, black, or purple; body uniformly longi-
tudinally banded or striped on dorsum, especially along dorso-lateral lines, rarely
uniformly whitish or greenish 7.

7(8) Head with a distinct blackish semicircular band extending from gena to gena dorsad
of front and involving ocellarae; body with a very fine brownish line along latero-
dorsal lines, more distinct on caudal segments; thoracic legs uniformly pale; length,
15 mm.; on grasses; Y-41-l-l,-41-2,-41-3,-8.47(?) M-41,-235 Dolerus sp. 4.

8(7) Head without distinct blackish semicircular band, but with spots or dark-colored
areas; thoracic legs with femur, tibia, and claws brown, not concolorous with coxa. .9.

9(12) Head pale brown, vertex with brown spots; body uniformly whitish or with light

dorsal band 10.

10(11) Vertex with two small spots, one dorso-mesad of each vertical furrow, variable in
size but never linear along the furrow; body uniformly whitish or creamy; length.

21 mm.; on Carpinus and Pkris aquilina (both doubtful) ; Y-74-1-1, M-82

Dolerus sp. 5.

11(10) Vertex with one minute spot at the origin of epicranial stem; light brown spot along

epicranial suture to ocellarae; body with dorsal band lighter on dorso-meson and

darker on supraspiracular lines, more distinct on caudal segments; length, 15 mm.;

on Equisetum arvense; Y-145-2 Dolerus sp. 6.

12(9) Head brownish with purplish or brownish markings on vertex, only rarely light
brown or yellowish, then vertical furrows with brownish streaks; body with distinct
dorsal band 13.

13(14) Head deep purphsh black with following parts whitish: proximal half of epicranial
stem, vertical furrows, vertex caudad of ocellarae to the middle of epicranial stem
very narrowly, epicranial arms, clj^ieus, and labrum; dorsal band lighter on dorso-
meson; pedal lines with a row of grayish patches; legs with femora without disto-
ventral projection; length, 19 mm.; on Equisetum arvense; Y- 146-1- 2. .similis Norton.

14(13) Head usually brownish or yellowish, vertex deeply brown, at least along vertical
furrows; tj^jically pale on vertex ventrad of vertical furrows and caudad of ocellarae;
front with or without brownish spot; tenth abdominal tergum on both sides usually

375] LARVAE OF THE TENTHREDINOIDEA—YUASA 57

more brownish than on meson; legs with femora without dis to- ventral projection;
length, 20-23 mm.; on sedges; Y-28-l,-27-l-l,-30,-210-l-8.32(?)-l-l, M-7,-8,-9,-35,
-64,-193,G-d-3 Dolerus sp. 7.

Subfamily Phyllotominae

Larvae (Fig. 11-12) small, length usually less than 15 mm.; body sub-
cylindrical or depressed, without colored patterns; larvapods on abdominal
segment with two or six annulets; antennae with 3-4 or 5 segments;
thoracic legs with four segments, short, stubby, with or without tarsal
claws.

The Phyllotominae can be divided into two distinct tribes on the basis
of the larval characters. The tribes can be separated as follows:

Thoracic legs with tarsal claws; head normal in form, not depressed; third abdominal

segment with six annulets; external feeders Phyllotomini.

Thoracic legs without tarsal claws; head depressed; third abdominal segment with
two annulets; leaf-miners Phlebatrophini.

The Phyllotominae is a distinct group and includes four genera, Phyl-
lotoma, Caliroa, Endelomyia, and Phlebatrophia. In the Nearctic region,
the last three genera are represented by a limited number of species.
MacGillivray considered this family as one of the five generalized sub-
families of the Tenthredinidae, quite apart from the Fenusinae and
Scolioneurinae, but Rohwer would associate them in his subfamily Mes-
sinae while Konow would include Hoplocampinae and Phyllotominae
in his tribe Hoplocampides. The subfamily is divisible into two distinct
groups according to the characters of the larvae. The remarkable speciali-
zation of structures due to the leaf-mining habit of the larva in one genus
where specialization has proceeded much further than in any of the other
leaf-miners, makes the division of the subfamily into two tribes desirable.

Tribe Phyllotomini

Body practically subcylindrical, thorax distinctly swollen, some-
times distinctly tadpole-like, tapering caudad; segmentation and annu-
lation indistinct, fine, subequal in length; third abdominal segment
with six annulets, annulets 2 and 4 microscopically and sparsely setiferous
or minutely tuberculate; tenth abdominal tergum with or without tubercles;
thoracic legs as long as head is wide, subequal in size, short, modified, with
four segments, stubby, with distinct tarsal claws, coxa conical, femur
cylindrical, as long as wide, tibia convex, wider than long, distal segment
very minute, with sharp incurved claw; larvapods on abdominal segment
2-8 normal in form, glabrous, distal lobe with a minute point on its cephalo-
ventral angle; ultimate segment with a pair of normal larvapods or without
any; head small, normal, not depressed, sparsely setiferous, longer than

58 ILLINOIS BIOLOGICAL MONOGRAPHS {376

wide, slightly pointed at dorsal apex; mouth directed slightly ventro-caudad;
antennae with four or five segments, slender, elongate, conical, or sub-
conical; mouth-parts normal in form; spiracles with or without wings;
ventral glands wanting; prothoracic glands sometimes present; glandubae
present, conical, tuberculate or sessile; cuticle usually microscopically
verrucose; larvae in life sometimes distinctly slimy; subgregarious; leaf-
skeletonizers.

GENERA OF PHYLLOTOMINI

Body without minute tubercles, tadpole-like, slimy; glandubae sessile Caliroa Costa.

Body with minute tubercles, not tadpole-like, not slimy; glandubae conical, tuberculate.

Endelomyia Ashmead,

Endelomyia Ashmead

Larvae small, length less than 15 mm., greenish yellow; body sub-
cylindrical, apparently almost glabrous, not tadpole-like, thorax
thickened, tapering caudad; third abdominal segment with six annulets,
annulets 2 and 4 tuberculate; tenth abdominal tergum with eight to ten
conical tubercles arranged approximately in three transverse rows; suranal
and subanal lobes with several rather long stiff setae; thoracic legs with
distinct tarsal claws; larvapods on ultimate segment normal in form,
separated; antennae with five segments, slender, elongate-conical; mandi-
bles with dentes; spiracles not winged; spiracles on sublateral lines; pro-
thoracic glands wanting; glandubae conical, tuberculate; body not slimy.

Endelomyia aethiops Fabricius. — Length, 13 mm, ; width of head, 1.2 mm. ;
head light brown; mouth-part's, labrum, and tarsal claws deep brown; body
greenish yellow to yellowish white; tubercles concolorous with body;
typical tubercular formula on prothorax: 3-6 on first annulet, 2-3 on pro-
subspiracular lobe, 2-5 on annulet 2; third abdominal segment with 2 and
3 tubercles on annulets 2 and 4 respectively, 1 on annulet 3 near the spiracle,
1 each on subspiracular and surpedal lobe; annulation, 1, 2, 3, 4, (5, 6);
antennae, 1, (2, 3, 4), 5; maxillary palpi, (1, 2, 3), 4; labial palpi, 1, 2, or
(1,2); subgregarious; on Rosa; Y-2, M-127.

Caliroa Costa

Larvae small, length 6-12 mm., whitish; body distinctly tadpole-like;
thorax conspicuously swollen, rounded on dorsum and flattened on venter;
tapering distinctly caudad; third abdominal segment with six indistinct
annulets, annulets 2 and 4 with a few glandubae; tenth abdominal tergum
without tubercles; suranal and subanal lobes often with a number of stiff
rather long setae; thoracic legs with distinct tarsal claws; larvapods on
ultimate segment obsolete, their position indicated by a small median
swelling; antennae with four segments, rather thick, elongate, distal
segment microscopic; mandibles with dentes; spiracles usually winged;

377] LARVAE OF THE TENTHREDINOIDEA—YUASA 59

spiracular line abnormally low in position, coinciding with latero-ventral
line; prothoracic glands present, large, triangular, fleshy, attached cephalo-
mesad of prothoracic legs; glandubae sessile;

SPECIES OF CALIROA

1(2) Anal setae not all of same type and length, longer ones arising from minute but
distinct tubercles, brown, apparently barbed, curved at tips, as long as labrum,
ten to twelve in number, arranged in a transverse row on suranal and subanal lobe;
spiracles distinctly winged, brown; clypeus light brown; head deep brown; thoracic
legs and antennae deep brown; smaller setae on anal area normal in form, much
shorter than the long barbed ones, scattered be3'ond the transverse rows; lengths of
front, cl>T)eus, labrum, and width of labrum to each other as 23, 10, 8 and 12 respec-
tively; length of body, 11 mm.; width of head, 1 mm.; on cherry, plum, Crataegus;
Y-209, M-260,-249,-115, C-551,-552 cerasi Linnaeus.

2(1) Anal setae all of same type and length, none arising from distinct tubercles and
barbed, all of normal tj-pe, much shorter than labrum; spiracles usually not winged .3.

3(8) Head blackish, deep brown, or brownish; legs brownish in part, not concolorous with
body 4,

4(7) Head blackish or dark brownish 5.

5(6) Head black or dark brownish black; anal setae scattered, pale, subequal in length,
about three-fifths as long as labrum; prothoracic legs distinctly lighter in color than
other legs, which are brownish; spiracles of cephalic segments faintly winged; clypeus
whitish; lengths of front, clypeus, labrum, and width of labrum to each other as
22, 9, 8, and 11, respectively; length of body, 10.5 mm.; width of head, 1 mm.; on
oak; M-157,-200,-245, G-553c, Y-mck Caliroa sp. 1.

6(5) Head deep brown; anal setae scattered, subequal in length; prothoracic legs higher in
color than other legs; spiracles never winged; anal setae about one-third as long as
labrum; clypeus pale brown; lengths of front, clypeus, labrum, and width of labrum
to each other as 18, 10, 8, and 11 respectively; length of body 6 mm.; width of head,
.9 mm.; on wild cherry; C obsokta Norton.

7(4) Head brov.nish or light brown; all legs pale brownish; anal setae scattered, subequal
in length, about one-third as long as labrum; clypeus pale brown; lengths of front,
cljijeus, labrum, and width of labrum to each other as 10, 9 ,7, and 11, respectively;
length of body, 8 mm.; width of head 1 mm.; on white oak and Crataegus; C-7,
M-230 quercus-alba Norton.

8{3) Head pale brown or whitish; legs whitish, concolorous with body; spiracles not
winged; anal setae scattered, subequal in length, about three-fifths as long as labrum;
lengths of front, clypeus, labrum, and width of labrum to each other as 21, 9, 7, and
10, respectively; length of body, 10.5 mm.; width of head, 1 mm.; Y-121, G-553c,
M-143,-201,-231,-236-242 quercus-coccinea Dyar.

Tribe Phlebatrophini

Body viewed from the side distinctly depressed, venter flattened,
thorax thickened, broadest on mesothorax, prothorax declivous cephalad,
caudal segments distinctly tapering; segmentation and annulation distinct;
third abdominal segment with two annulets, annulet 2 sparsely and incon-
spicuously setiferous; tenth abdominal tergum without tubercles; tenth
sternum small; thoracic legs modified, fleshy, indistinctly four-segmented.

60 ILLINOIS BIOLOGICAL MONOGRAPHS [378

tapering to distal end, distal segment microscopic, mamma-like, without
tarsal claw; larvapods vestigial, located on abdominal segments 2-8 and 10,
anal larvapods united on the meson, forming a single sucker-like pro-
tuberance; head distinctly depressed, pointed, subtriangular in contour,
partly overlapped by protruding prothorax; mouth directed cephalad;
vertical furrows obsolete; antennae with 3-4 segments, segments 1 and 2
large and conical, segments 3 and 4 subcylindrical and much smaller and
less in diameter than proximal segments; mouth-parts modified, mandibles
slender, sharply pointed, without dentes, blade-like; labium flattened and
large; spiracles not winged; ventral and prothoracic glands wanting;
glandubae obsolete; body not slimy; leaf-miners.

Phlebatrophia MacGillivray

Larvae very small, whitish; length less than 10 mm.; body distinctly
depressed, tapering much caudad, broadest on mesothorax, prothorax
declivous cephalad; lateral lobes somewhat prominent; tenth abdom-
inal tergum convex, almost glabrous, about half as wide as mesothorax;
suranal and subanal lobes semiglabrous; third abdominal segment with
two annulets, caudal annulet about four times as long as the cephalic,
microscopically and sparsely setiferous; thoracic legs fleshy, tarsal claws
wanting; larvapods rudimentary; ocularia protruding, located laterad
of antennariae; epicranial suture in part obsolete; spiracles not winged;
spiracular line normal in position; maxillary palpi with four segments,
labial palpi with two segments; totaglossa roundly protruding; stipes
elongate, subgalea long, slender, with distinct chitinized carinae; labrum
flattened; mandibles slender, sharply pointed, without dentes, blade-like;
leaf-miners.

Phlebatrophia mathesoni MacGillivray. — Length, 7 mm.; width of head,
1 mm.; mesothorax, 2.4 mm. wide; head light brown, mandibles and
carinae of subgalae deep brown; maxillary palpi typically 2, 1, 3, 4, distal
segment very minute; antennae with proximal segment or segments
larger in diameter and fleshy, conical, two distal segments together smaller
and shorter than the other two segments, but longer than labial palpi;
distal segment of labial palpus very minute; leaf-miners of birch; C and G.

Subfamily Tenthredininae

Larvae (Fig. 13) of medium to rather large size; body cylindrical,
slender, tapering uniformly and gradually caudad; segmentation and
annulation distinct, fine; third abdominal segment with seven annulets,
annulets 1, 3, and 5 setiferous and 3 and 5 with transverse row of glandu-
bae; abdominal segments 2-8 and 10 with larvapods; antennae with five
segments, slender, cylindro-conical; body uniformly greenish, with dark

379\ LARVAE OF THE TENTHREDINOIDEA—YUASA 61

dorsal band or with complicated color-patterns; tenth abdominal tergum
convex, suranal or caudal protuberances wanting; ninth abdominal tergum
with six annulets, annulet 6 as long as annulet 2; thoracic legs normal,
well developed, femur with ventro-distal, conical membranous projection;
clypeus with two setae on each side; labrum with 3-5 setae on each side,
with or without a median longitudinal depression; maxillary palpi slender,
normal; galea digit-like, large; lacinia flattened, with a row of 10-15 setae
on the oblique, truncate cephalic margin; stipes with a sharp triangular
cephalo- ventral projection; labial palpi long, slender, with segment 2
longer than segment 1; mandible with 1, 2, or 3-4 setae; head variously
marked, distinctly and densely setiferous; spiracles on third annulet, not
winged; larvapods glabrous; glandubae distinct, slender, elongate cylindro-
conical, sometimes longer than adjacent setae; cuticle microscopically and
densely spinulate; ventral glands wanting; larvae free leaf -feeders.

The Tenthredininae constitutes, according to MacGillivray, the second
subfamily of the series of specialized Tenthredinidae. Rohwer (1911)
would divide the subfamily into two tribes, Perineurini and Tenthredinini,
using the position of the propodeal spiracles and shape of the cephalic
margin of the scutellum as characters for differentiating them. In many
cases the larvae resemble those of the Emphytinae.

GENERA OF TENTHREDININAE

1(4) Mandibles with more than one seta; labrum with median longitudinal depression;

legs with dorsal aspect of femur usually less than twice as long as trochanter, but

often subequal to it 2.

2(3) Mandibles with two setae Macrophya Dahlbom.

3(2) Mandibles mth four, occasionally three, setae Tentliredo Linnaeus.

4(1) Mandibles with a single seta; labrum without median longitudinal depression;

legs with dorsal aspect of femur usually twice as long as trochanter 5.

5(6) Body with complexly patterned markings on the dorsum; distal segment of maxillary

palpi usually longer than that of labial palpi; head nearly black . . Tenthredopsis Costa,
6(5) Body without complexly patterned-markings on the dorsum; distal segment of

maxillary palpi usually not longer than that of labial palpi; head pale or light brown

Neopiis MacGLUivray

Tenthredopsis semilutea Norton. — Body on dorsum with complexly
patterned purplish-black markings extending to supraspiracular lines;
two lighter colored spots on dorso-meson, their apices directed cephalad,
cephalic one much larger than caudal; large spot with caudal emargination
on latus and contiguous to mesal triangles; subspiracular lobe with faint,
minute spots; otherwise ventral half of body including legs and larvapods
whitish; head purplish black excepting the following parts, which are
white: genae including antennae and antennariae, lower fourth of front,
vertex narrowly, laterad of vertical portion of epicranial arms, clypeus,
labrum, and mouth-parts except tips of mandibles, which are black;

62 ILLINOIS BIOLOGICAL MONOGRAPHS [380

fronto-clypeal suture sometimes black; in young specimens, head grayish
and body entirely whitish-green; in life head and body coated with a waxy
bloom; annulation, (3, 5, 1), (2, 4), 6, 7; antennae, 5, 2, (1, 3, 4); maxillary
palpi, (4, 2), 3, 1; labial palpi with distal segment twice as long as the pre-
ceding segment, but subequal in length to or shorter than distal segment
of maxillary palpi; labrum usually with three setae on each side and
without median longitudinal depression; mandible with one seta; legs
with trochanter one-half as long as tibia, femur slightly longer than
tibia; glandubae half as long as adjacent setae; length, 18 mm.; width of
head, 1.8 mm.; on Thalictrum polygamum; Y-8,-92-1-1.

Neopus 14-punctatus Norton. — Head pale creamy-white with brown
spots on dorso-meson of vertex, caudad of ocellarae, and on front, frontal
spots much darker; body whitish green, dorsum with a grayish shade, bor-
dered along supraspiracular lines with narrow grayish bands, dorso-meson,
especially on thorax, with fine double bands; venter including legs and
larvapods whitish; annulation, (1, 5, 3), (2, 4, 6, 7); antennae, (5, 1), 2,
(3, 1); maxillary palpi, 4, 2, (3, 1); labial palpi with distal segment one-
fourth longer than preceding segment, but subequal to distal segment of
maxillary palpi; labrum with three setae on each side, median longitudinal
depression wanting; mandible with single seta; legs with trochanter nearly
one-half as long as femur, tibia equal in length to femur; glandubae more
than half the length of adjacent setae; length of body, 18 mm.; width of
head, 1.8 mm.; on Podophyllum peltatum: Y-205-1-1.

Tenthredo bilineata MacGillivray. — Head whitish green with vertex
brown dorsad of genae except narrow line along epicranial stem, vertical
furrows, and caudad and dorsad of ocularia; body on dorsum with series of
triangular brownish markings; triangle with apex directed cephalad and
with minute deep brown spot at each basal angle, the triangle divided on
meson by a faint light line; supraspiracular line with light brownish in-
definite band; venter including legs and larvapods whitish; annulation,
(3, 1, 5), (7, 6, 2, 4); antennae, 5, (1, 2, 3, 4); maxillary palpi, (4, 2), 1, 3;
labial palpi with distal segment nearly twice as long as the preceding seg-
ment and shorter than distal segment of maxillary palpi; labrum with five
setae on each side and with median longitudinal depression; mandibles with
four setae; legs with trochanter nearly as long as femur, tibia longer than
femur; glandubae conical, large, subequal in length to adjacent setae;
length of body, 21 mm.; width of head, 2 mm.; on Geranium maculatum:
Y-175-2.

Macrophya Dahlbom

Body usually whitish green, on dorsum with or without grayish band,
latus sometimes with small black spots; head usually marked on vertex;
antennae with segment 1 or 2 longest or all segments subequal in length;

381] LARVAE OF THE TENTHREDINOIDEA—YUASA 63

maxillary palpi usually with segments 2 and 4 subequal in length; mandible
with two setae; labrum with three setae on each side, with median longitu-
dinal depression; trochanter distinctly shorter than femur; head and
body in life usually coated with a thin whitish waxy bloom; dorsal vessel
usually showing thru cuticle as a dark fine line; setae microscopic; glandu-
bae shorter or longer than adjacent setae; length, 16-21 mm.

SPECIES OF MACROPHYA

1(4) Head pale brown, with or without a minute brown spot at the caudal end of epi-
cranial stem; body on dorsum with light or pale grayish band, darker along supra-
spiracular lines, band sometimes obsolete; venter whitish, including legs and
larvapods; tenth abdominal segments unmarked; labial palpi with distal segment
twice as long as the preceding segment; glandubae very small, shorter than adjacent
setae; setae microscopic; head and body coated with thin waxy bloom; on Prunus
serotina; subgregarious 2.

2(3) Larger species, length, 20 mm.; width of head, 1.8 mm. ; annulation, (1, 5), 3, (4, 6, 7),

2; antennae, (2, 1), (3, 4, 5) ; maxillary palpi, 2, (1, 3, 4) ; Y-126,-126-3-C-l

flicta MacGillivray.

3(2) Smaller species, length, 16 mm.; width of head, 1.8 mm.; annulation, (5, 1, 3), 4, 6,

2, 7; antennae, 2, (5, 1, 4), 3; maxillary palpi, (2, 4), (3, 1); Y-59-3- 1,-59-4-1

fistula MacGillivray.

4(1) Head with a large blackish spot on dorsal part of vertex, often with black spots
caudad of ocellarae; body with or without distinct grayish dorsal band, on latus with
rows of black or yellowish spots; venter usually whitish; pedal line sometimes with
fine gray markings 5.

5(8) Body entirely whitish; head on vertex usually with a dorsal spot not expanding
distad ; without spots caudad of ocellarae 6.

6(7) Body with a row of small black spots along supraspiracular lines, two spots to each
segment, cephalic spot larger than caudal; tenth abdominal tergum unmarked;
annulation, (1, 3, 5) (4, 7), (6, 2); antennae, (4, 1), 2, 3; legs with femur more than
twice as long as trochanter, tibia shorter than femur, coxa with grayish marking;
glandubae longer than adjacent setae; length of body, 22 mm.; width of head, 2.3
mm.; on Sambucus; Y-8. 11,-11 tibiator Norton.

7(6) Body without a row of small black spots along supraspiracular lines, but with
yellowish spots on latus instead, which are obsolete in alcoholic specimens; annula-
tion, (3, 5, 1), (2, 7, 4, 6); antennaCj 1, (2, 3, 4, 5); maxillary palpi, (4, 2), (1, 3);
labial palpi with distal segment only one-fourth longer than the preceding segments;
legs with trochanter slightly shorter than tibia, femur sUghtly longer than tibia;
length of body, 22 m.; width of head, 2.2 mm.; on Sambucus racemosa; Y-8. 11
-2(?)-3 epinota Say.

8(5) Body not entirely whitish, dorsum with dark dorsal band; latus with one or more
rows of distinct black spots; head on vertex with a dorsal spot expanding distad,
broadly T-shaped, caudad of ocellarae with spots; tenth abdominal segment with a
minute black spot on caudo-meson; venter lighter in color 9.

9(10) Body on latus with grayish band darker in color than dorsal band and with only
one row of distinct black spots; supraspiracular lines with a row of distinct spots;
dorso-lateral lines with a row of smaller, inconspicuous spots; subspiracular line
with very faint grayish spots, nearly obsolete ; pedal lines with grayish linear markings ;
annulation, 1, (3, 5), (7, 2, 4), 6; antennae, (1, 2, 3, 4, 5); maxillary palpi, (2. 4),
1, 3; labial palpi with distal segment not quite twice as long as the preceding segment;

64 ILLINOIS BIOLOGICAL MONOGRAPHS [382

legs with femur twice as long as trochanter and slightly longer than tibia; length of

body, 19 mm.; width of head, 2 mm.; on Aster prenanthoides ; Y-8, 81(?)

lineata Norton.
10(9) Body on latus with grayish band lighter in color than grajdsh-purple dorsal band and
with three rows of distinct black spots; dorso-lateral and supraspiracular lines with
rows of small spots, two spots to each segment, with caudal spot much smaller and
sometimes nearly obsolete; spots on supraspiracular line largest; pedal line with a
row of spots, two to each segment, with cephalic spots smaller than caudal; head
with black spots on vertex large, sometimes coalesced, covering entire vertex except
genae and vertical furrows; front with faint gray spot; annulation, (3,5, 1), (7,2,4,6);
antennae, 2, (1, 5), (3, 4); maxillary palpi, (4, 2), (1, 3); labial palpi with distal
segment not quite twice as long as the preceding segment; legs with trochanter more
than one-half as long as tibia, femur equal in length to tibia; length of body, 21.5
mm.; width of head, 2.1 mm.; on Solidago juncea and Rudbeckia laciniata; Y-160-2,
-160-1 pulchdla Klug.

Subfamily Cimbicinae

Body cylindrical (Fig. 14), tapering uniformly caudad, apparently
glabrous, pro thorax narrowed; segmentation indistinct; annulation fine, 7,
(2, 3, 4), (1, 5, 6), annulets 2, 4, and 7 microscopically setiferous; thoracic
legs normal in form, with five segments, femur slightly longer than tibia;
larvapods on abdominal segments 2-8 and 10, divided into two unequal
lobes on the distal surface, few setae on the dorso-caudal aspect, and none
on the cephalic aspect as viewed from side; tenth abdominal tergum with-
out suranal protuberances; suranal and subanal lobes with several short
setae; head large, rather thickly setiferous, normal in form; labrum sub-
divided by diverging depression into median lobe and two lateral lobes,
sometimes asymmetrical; antennae with a single segment, button-like but
chitinized; ventral glands wanting; glandubae microscopic or minute and
stalked, sometimes distinct conical tubercles; spiracles distinctly winged;
conspicuous spiracular glands located dorsad of each spiracle of abdominal
segments 2-8; cuticle microscopically and densely spinulate or verrucose;
mouth-parts normal in form, maxillary and labial palpi rather slender,
galea very thick, curved mesad at distal end, sericos large, distinctly
chitinized, pear-shaped, U-shaped, or V-shaped; stipes with a distinct
triangular projection on the dorsal or cephalic margin; body covered
with waxy bloom in life; larvae ejecting yellowish fluid from spiracular
glands when disturbed; free leaf-feeders, sometimes semigregarious.

The Cimbicinae is a small compact subfamily consisting of few genera
and a limited number of species in the Nearctic region. Systematists are
in accord in the general conception of the group, but Konow includes the
Perginae in his subfamily Cimbicini. Cimbex americana, with its several
varieties, is a well-known representative of this subfamily.

383] LARVAE OF THE TENTH REDINOIDEA—YU ASA 65

GENERA OF CIMBICINAE

1(4) Body in general whitish, without minute colored spots 2.

2(3) Dorsum of body with a black median stripe Cimbex Olivier.

3(2) Dorsum of body without a black median stripe Trichiosoma Leach.

4(1) Body in general not whitish, but with minute colored spots Abia Leach.

Cimbex Olivier

Larvae very large, distinctly robust, largest of all saw-fly larvae, 40-50
mm. in length; body whitish with a distinct dorso-mesal black line; labrum
distinctly asymmetrical, dextral portion larger than sinistral, median
cephalic or ventral emargination distinct and deep; body with distinct
minute warts or conical glandubae, cuticle microscopically verrucose;
spiracular glands semicircular; sericos of labium pear-shaped with its nar-
row neck directed dorsad or cephalad; lateral lobes small but prominent with
several conical tubercles; maxillary palpi, 2, 4, 3, 1; labial palpi, 1, 2;
antennae mound-like, wider than high

Cimbex amerkana Leach. — Length, 50 mm.; head, 5 mm. wide; head
white, creamy, microscopically brownish verrucose; black median stripe on
dorsum extending from prothorax to the middle of eighth abdominal
segment; conical tubercles or glandubae on sublateral lobes larger than
elsewhere and 4-7 in number; larvae solitary; on willow, elm, poplar,
maple, alder, linden, etc.; Y-68, -8, -182, M-99.

Trichiosoma Leach

Larvae very large, somewhat slender, entirely white; labrum slightly
asymmetrical, right side larger than sinistral, median ventral emargina-
tion deep, but not reaching the median lobe; body not warty; glandu-
bae microscopic, stalked, arising from very low swellings; cuticle micro-
scopically verrucose; spiracular glands semicircular; sericos of labium
U-shaped, narrower on dorsal or cephalic end; lateral lobes not prominent,
without conical tubercles; maxillary palpi 2, 4, 3, 1; labial palpi, 1, 2;
antennae conical, longer than wide.

Trichiosoma sp. — Length, 38 mm. ; width of head, 4 mm. ; head creamy
white, body whitish; solitary; on willow, poplar, alder, wild cherry;
M-78, -44.

Abia Leach

Larvae large, rather plump, greenish or grayish green with minute
yellowish or blackish spots; labrum symmetrical, median ventral emargi-
nation only slightly indicated, broad; body without warts or tubercles;
cuticle microscopically and densely spinulate; glandubae stalked and
minute, not arising from swellings; spiracular glands semicircular; sericos
of labium V-shaped, widest at dorsal or cephalic end; sublateral lobes

66 ILLINOIS BIOLOGICAL MONOGRAPHS [384

inconspicuous, with several glandubae; maxillary palpi, (4, 2) 3, 1; labial
palpi with segments subequal; antennae conical, usually as wide as long.

SPECIES OF ABIA

1(2) Body with a broad dorso-mesal yellowish stripe between subdorsal lines; dorsum
above supraspiracular lines shaded brownish gray; head brownish gray except
front, genae, clypeus, and labrum which are lighter; ocellarae white; body dorsad of
supraspiracular lines shaded grayish or brownish, venter pale whitish-green; dorsum
between subdorsal lines yellowish white; body with four rows of minute black
spots; middorsal row with a large spot on annulet 7 and a smaller spot on annulets 2
and 4; subdorsal row with a large spot on annulet 7, with a bright yellow spot between
two black spots mentioned above; supraspiracular row with a spot on annulets 2
and 4 and a very small spot on annulet 7; subspiracular row with a spot at the
ventral ends of annulets 3 and 4; a yellow spot caudad of each spiracle; length,

26 nam.; width of head, 2.7 mm.; on honeysuckle; semigregarious; Y-8.13

americana Cresson.

2(1) Body with a broad dorso-mesal yellowish stripe between subdorsal lines; dorsum
above supraspiracular lines not shaded brownish-gray, but body generally greenish . . 3.

3(4) Pedal line without distinct more or less continuous smoky brownish gray band;
black spots directly ventrad of spiracles never present; second annulet without minute
but distinct spots dorsad of lateral large spots; head grayish brown except front,
genae, and clypeus, labrum lighter; ocellarae white; body entirely grayish green,
dorsum between subdorsal lines concolorous with other parts, the dorso-meson
with a narrow yellomsh line; five rows of minute black spots: meso-dorsal row with
a larger spot on annulet 7 and a smaller spot on annulets 2 and 4; subdorsal row with a
larger spot on annulet 7 und very minute and often obscure spot on annulet 2, and
larger, distinct, often subdivided, spot on annulet 4, the latter two nearer to meson
than the regular subdorsal spots; lateral row with a large spot between annulets
2 and 3; supraspiracular row with a smaller spot on annulets 4 and 7; subspiracular
with a larger spot at the ventral end of annulets 3 and 4; a bright yellow spot between
mesal and subdorsal black spots on annulet 7; length, 28 mm.; width of head, 2.8
mm.; on Triosleum aurantiac-nm; Y-8.13-2, M-196 inflata Norton.

4(3) Pedal line with distinct, more or less continuous smoky brownish gray band; black
spots directly ventrad of spiracles always present, at least on majority of segments;
one or two minute black spots on second annulet dorsad of large lateral spots always
present; length, 30 mm.; width of head, 2.9 mm.; on honeysuckle; Y-104, G-583,
both from Urbana, 111. ; larvae resemble A . inflata in markings and general appearance
but differ as above Abia sp. 1.

Subfamily Hoplocampinae

Larvae (Fig. 15) small; body cylindrical, slender; segmentation and
annulation sometimes obsolete; third abdominal segment with four or five
annulets, annulets 2 and 3 or 2 and 4 setiferous or all annulets glabrous;
larvapods present on abdominal segments 2-7 and 10, glabrous or setiferous,
sometimes very rudimentary; ventral glands usually present on the meson
of abdominal segments 1-7; body greenish or yellowish, striped or spotted
or without any markings; tenth abdominal tergum with or without suranal
protuberances, if present, often more than two in number; antennae

385J LARVAE OF THE TENTHREDINOIDEA—YUASA 67

conical, with five segments, or flattened, with four, sometimes apparently
with three; larvae free leaf -feeders and borers in fruits or petioles of leaves.

The Hoplocampinae as defined by MacGillivray contains at present
five genera, Marlattia, Hoplocampa, MacGillivrayella, Hemichroa, and
Craterocercus, and represents, together with Dineurinae, a series in which
the anal veins have been modified before the loss of the radial cross-vein.
Formerly Rohwer (1910, 1911a, 1913b) considered Hoplocampa and
MacGillivrayella as constituting a subfamily Hoplocampinae but later
(1918c) he abandoned this idea and united these genera with six other
genera and subgenera to form the tribe Hemichroini of his subfamily
Nematinae, as others have done. With the exception of two genera,
Platycampus and Anoplonyx, Rohwer's tribe Hemichroini becomes
coextensive with our Hoplocampinae. Cameron (1883) considered this
subfamily as forming "a connecting link between the Selandrides and
Nematides." There are reasons for indicating a close relation between
this subfamily and Nematinae. A study of the larvae confirms the con-
tention of Rohwer (1918c) that the grouping of Caliroa and Phyllotoma with
the Hoplocampinae, as was done by Konow and Enslin, is untenable.

It must be stated here that since only three genera, each represented
by a single species, were available for this study, the preceding definition
of the subfamily is necessarily incomplete.

GENERA OF HOPLOCAMPINAE

1(2) Tenth abdominal tergum without caudal protuberances Marlattia Ashmead,

2(1) Tenth abdominal tergum with caudal protuberances 3.

3(4) Caudal protuberances more than two in number; larvapods well developed; third

abdominal segment with five annulets; free leaf-feeders Hemichroa Stephens,

4(3) Caudal protuberances two in number on caudal projection; larvapods rudimentary;

third abdominal segment with four aimulets; leaf-petiole borer. Caulocampiis Rohwer.

Hemichroa Stephens

Larvae small, greenish; length less than 18 mm.; body slender, tapering
uniformly caudad; third abdominal segment with five annulets 2 and 4
setiferous; tenth abdominal tergum with several conical caudal protuber-
ances on its caudal margin; antennae distinctly conical, with five segments,
as long as the longest diameter of antennaria; antennal segment 1 cres-
centic, dorsal in position, extending nearly the entire length of antennaria,
segment 2 complete or incomplete, reduced to mere line on cephalic
aspect, segments 3 and 4 ring-like tho reduced in length on cephalic portion,
segment 5 conical or peg-like, bluntly pointed at apex; thoracic legs
with tibia subequal in length to femur; larvapods glabrous; spiracles faintly
winged; glandubae distinct and large; larvae free leaf -feeders.

Hemichroa dyari Rohwer. — Larvae yellowish green; length, 16 mm.;
head blackish; body with blackish dorso-lateral lines and interrupted

68 ILLINOIS BIOLOGICAL MONOGRAPHS [386

blackish lines on subspiracular and pedal lines; tenth abdominal tergum
with six to seven conical protuberances, suffused with brown; surpedal and
subspiracular lobes with two setae and two glandubae; glandubae with
diameter twice that of anal setae; annulation, (1, 2, 4) 3, 5; on alder; Y-39-
-1-1,-8.73 (?) -l,-8.73(?)-2, C-8.

Marlattia Ashmead

Larvae comparatively very small, greenish, with or without stripes
or spots; body cylindrical, tapering caudad; third abdominal segment
with five annulets, annulets apparently glabrous; tenth abdominal seg-
ment without caudal protuberances; antennae with four segments, flat-
tened; segment 1 small, incomplete; segments 2 and 3 complete but
reduced to narrow line on cephalic aspect; segment 4 minute, mamma-
like; head sparsely setiferous, setae increasing in length on the lower
portion; thoracic legs normal in form, tibia subequal in length to femur; lar-
vapods with a few setae; spiracles minute, unwinged; glandubae micro-
scopic.

Marlattia laricis Marlatt. — Head pale yellowish, body greenish with
faint subdorsal lines; third abdominal segment with five annulets, (1, 2)
4, 3, 5; larvapods with three setae; suranal lobe with a few setae on caudo-
ventral aspect; small larvae, length, 10 mm.; M-57.

Caulocampus Rohwer

Larvae very small; length less than 10 mm.; body subcylindrical, taper-
ing at prothorax, constricted suddenly on the ultimate segment, not
spotted or striped, but whitish, sparsely and microscopically setiferous;
third abdominal segment with four annulets, annulets 2 and 3 with very
minute setae; thoracic legs minute, with five segments, tibia longer than
femur; abdominal segments 2-7 with rudimentary larvapods represented by
spinulate swellings; tenth abdominal tergum much smaller in diameter
than preceding segments, strongly converging caudad, caudal margin
produced and chitinized, a pair of minute brownish caudal protuberances
on the caudal margin of the projection; head small, sparsely and minutely
setiferous; ocellarae represented by pigmented spots, distinct, ring-like,
ocularia wanting; antennae apparently with three segments, conical, seg-
ments 1 and 2 incomplete, segment 3 slender, peg-like; mouth-parts
normal and conspicuous tho small; spiracles not winged; glandubae
obsolete; in young specimens dorsum of abdominal segments 1-7, with a
pair of protuberances on annulet 2; larvae borers in leaf -petiole.

This genus is monotypic and unique in the reduction of larvapods
and ocellarae and in the possession of modified caudal projection on the
ultimate segment with rudimentary caudal protuberances. In this last

387] LARVAE OF THE TENTHREDINOIDEA—YUASA 69

character the larvae of this genus resemble those of certain species of
Pontania. The modifications of the body are undoubtedly correlated with
the boring habit of the larvae. MacGillivray still considers this genus
as without question belonging to the Cladiinae, but Rohwer regards it as
belonging to his tribe Hemichroini. It is dealt with here under the Hop-
locampinae because the larvae more closely resemble larvae of this sub-
family than they do those of the Cladiinae.

Caulocampiis acericauUs MacGillivray. — Length, 8 mm.; width of head,
.8 mm.; head light brown, body straw-yellow; resembles larvae of weevils
in general appearance; mouth-parts normal in form; in young specimens
head yellowish and body whitish; annulation, 2, 1, 3, 4; maxillary palpi,
3, 2, 1, 4, segments brown, slender; galea digit-like, very small; thoracic
legs with very small tibiae, tibia subequal in length to maxillary palpus;
tenth abdominal tergum with many minute setae evenly and promiscu-
ously scattered, not concentrated on subanal lobe; larvae bore into the
petioles of maple-leaves; Y (generosity of Dr. W. E. Britton, New Haven,
Conn.).

Subfamily Dineurinae

Larvae small; body subcylindrical, flattened on venter or cylindrical,
usually tapering toward the caudal end, greenish or yellowish, often with
dorsum darker, never with bright-colored markings; glabrous or setiferous;
head small, light greenish or yellowish, never with distinct markings;
ocellarae blackish; mouth-parts usually brownish; thorax wider than the
remainder of body, thoracic legs well-developed, caudal pairs larger than
the cephalic, directed laterad; segmentation distinct; annulation indistinct;
larvapods on abdominal segments 2-7 and 10, sometimes rudimentary;
intersegmental coria often distinct and whitish; larvae feed on under side
or upper side of leaves, eating the parenchymatous layers only or feeding
on edges of leaves or mining in the leaves; ultimate stage glabrous and
yellowish; pupation in single-layered parchment-like cocoons in the
ground; some species with nauseating odor.

The Dineurinae as limited by MacGillivray contains three genera,
Dineura, Mesoneura, and Pseudodineura, and includes not over twenty-
five species, which are mostly distributed in Europe and North America.
This subfamily resembles in wing-type the Hoplocampinae. Systematists
do not agree in the exact position of the small European genus Pseudo-
dineura. Konow would place Dineura in his tribe Nematides but both
Mesoneura and Pseudodineura in his tribe Blennocampides. Rohwer,
on the other hand, would associate Dineura and Mesoneura in his tribe
Nematini, and is not quite certain whether Pseudodineura also belongs
to this tribe or not. Cameron, who described the larva of Pseudodineura
parvulus under the name of Dineura despeda, altho aware of the differences

70 ILLINOIS BIOLOGICAL MONOGRAPHS [388

between this species and its allies and other species of Dineura, hesitated
to agree with Thompson in associating it with Blennocampa. The vena-
tion certainly indicates close relationship between Dineura and Pseudo-
dineura. The latter is characterized in its larval stages by the leaf-mining
habit and structural modifications due to this mode of life, altho appar-
ently these do not constitute very striking distinctions if one may judge from
published records. Since previous authors failed to study the structures
of the head more carefully, and since these are of great taxonomic im-
portance, and also on account of the discrepancy between the larval
habits of the three genera in question, it is impossible to pass judgment on
their affinity until more is known with regard to their larval structures
and habits, particularly in the case of Pseudondineura.

The life history of the subfamily has been recorded by Girard, Cameron,
and Brischke and Zaddach. The larvae of the American species are un-
known, and as none of the European species have been available for study
the definition of the Dineurinae here given is tentative, and is based on
descriptions and figures published by Cameron (1882) and by Brischke
and Zaddach (1883).

Subfamily Cladiinae

Larvae (Fig. 16) of small to medium size; body rather flattened, wider
than high, slightly tapering caudad, conspicuously hairy, greenish or with
segmen tally arranged spots on dorsum darkly shaded; segmentation and
annulation usually distinct; third abdominal segment with four annulets,
annulets 1, 2, and 3 setiferous, setae, especially on annulets 2 and 3,
arising from wart-like tubercles, long, often curved, always microscopically
barbed, never branched, some of the setae distinctly longer than others;
annulet 4 narrow and glabrous; larvapods present on abdominal segments
2-7 and 10 well developed, long, distal portion often dilated, appearing as if
subdivided, often curved mesad, always with few setae; ventral glands small
but always present on abdominal segments 1-7; tenth abdominal tergum
without caudal protuberances but with many setae of varying length;
thoracic legs spreading flat laterad; femur with a ventro-distal projection,
subequal in length to tibia; antennae with four segments, subconical, large;
segment 1 complete or incomplete, segment 2 complete, thicker, dorsal or
caudo-dorsal portion with clear spaces, segment 3 smaller and narrower
than segment 2, segment 4 minute, conical; spiracles never winged; glandu-
bae small or obsolete; sericos usually very wide, occupying nearly four-
fifths of the width of the totaglossa; larvae external leaf -feeders.

The Cladiinae is a small subfamily and according to MacGillivray
consists oi six genera; Anoplonyx, Platycampus, Priophorus, Cladius, and
Trichiocampus. The first three genera are placed in the tribe Hemichroini
of his subfamily Nematinae by Rohwer (1911, 1918), who states that

389] LARVAE OF THE TENTHREDINOIDEA—YUASA 71

"the characters of both the adult and the larva pomt out subfamily
difference between Caulocampus and Priophorus" and that the former
superficially resembles Hoplocampa in all stages but is really related to
Craterocerus. The diflference of opinion is due to the different value
placed by these two writers on the presence or absence of the radial cross-
vein in differentiating the subfamilies. The genus Caulocampus has been
discussed in connection with the Hoplocampinae because the larvae of this
genus are very different from those of the Cladiinae both morphologically
and biologically and because they are more naturally associated with the
larvae of the Hoplocampinae. Anoplonyx is represented in the Neartic
region by a single species. Platycampus includes four American species,
two of which have been recognized in the immature stages. P. americana
feeds on Populus and P. juniperi on juniper. None of these larvae have
been examined.

GENERA OF CLADIINAE

1(2) Body spotted, with a row of blackish or brownish spots on subdorsal, supraspiracular
or subspiracular lines; setae usually recurved exceedingly long, longest ones longer
than one-half the height of the head; annulet 1 usually with one seta on each side of
meson Trichiocampus Hartig.

2(1) Body never spotted; setae usually straight, long, but the longest ones never distinctly
longer than half the height of the head; annulet 1 always with more than one seta on
each side of meson, usually with four to six setae 3.

3(4) Head with spots, usually with blackish patches on dorso-meson of vertex and caudad
of each ocellara; body dorsad of spiracular lines usually shaded darker than the
venter; body sometimes pinkish; postsupraspiracular tubercles usually with three
setae, never with more than four Priophorus Dahlbom.

4(3) Head never with spots, usually uniformly greenish; body dorsad of spiracular lines
never shaded darker but concolorous with venter; body never pinkish but greenish
yellow or whitish; postsupraspiracular tubercles usually with six setae, never with
less than four Cladius Rossi.

Trichiocampus Hartig

Larvae small to moderately large, length from 10 to 25 mm., distinctly
hairy, with segmentally arranged spots; body with a longitudinal row of
blackish or brownish spots along subdorsal, supraspiracular, or subspiracu-
lar lines; annulet 4 shortest, annulet 1 usually with one and sometimes two
setae on each half of body, annulet 2 with tubercles bearing 2-5 setae,
annulet 3 with three warts, two dorsal ones bearing 4-5 setae and ventral
one with 6-9 setae, postspiracular tubercle usually with two setae, sub-
spiracular lobe v/ith 8-9 setae, surpedal lobe with 6-10 setae; setae usually
recurved, variable in length, longest setae nearly subequal in length to
the height of head; warts or tubercles with setae of varying length, those
on annulet 1 among the shortest.

72 ILLINOIS BIOLOGICAL MONOGRAPHS [390

SPECIES OF TRICHIOCAMPUS

1(2) Body with three pairs of longitudinal rows of blackish or brownish segmentally
arranged spots along subdorsal, supraspiracular, and subspiracular lines; tenth
abdominal tergum not entirely black but white except a pair of minute spots; head
light brown with brownish spots, with following parts dark brown : dorso-meson of
vertex, dorsal two-thirds of front, and vertex dorso-caudad of each ocellara including
gena; preclypeus whitish, other parts pale, including occiput between vertical
furrows; a row of brownish spots from mesothoracic to ultimate segment along each
side of dorso-meson; a row of larger spots along supraspiracular line from prothoracic
to penultimate segments; another row of much smaller spots along subspiraciJar
lines from mesothoracic to eighth abdominal segment; mesothoracic and metathoracic
subspiracular spots more than twice as large as supraspiracular spots of same seg-
ments; the former with circular white areas around the proximal end of setae;
prothoracic supraspiracular spots small and indistinct; third abdominal segment
with following setal map: 1, 3, 5, 1, 2, 5, 4, 8, 9-10, 8-9; tenth abdominal tergum
white, except a pair of minute spots; subdorsal spots not involving tubercles 2 and 3;
supraspiracular spot involving tubercles 4 and 5; subspiracular spot on caudal half of
subspiracular tubercle; maxillary palpi (2, 3), 1, 4; head in younger specimens black-
ish except near the mouth; body without spots; in older specimens, supraspiracular
spots appear first, then subspiracular, beginning with caudal segments, setae on tuber-
cles sometimes one or two less than in mature specimens; on Populus; length of
body, 12 mm.; width of head, 1.5-1.6 mm.; G-Onekama and Orono on oak, length of
body, 13 mm.; width of head, 1.7 mm., M-207. (The latter resembles the former
so closely and indistinguishably, altho its setae may be slightly fewer in number,
that they are considered as identical.) />aeiw/i« MacGillivray.

2(1) Body with two pairs of longitudinal rows of blackish or brownish segmentally
arranged spots along supraspiracular and subspiracular lines; tenth abdominal
tergum entirely black; head blackish, with paler areas 3.

3(4) Mesothoracic supraspiracular spots subequal in size to subspiracular spots; supra-
spiracular spots of abdominal segments not involving postsupraspiracular tubercles;
prothoracic supraspiracular spots minute and indistinct; preclypeus, labrum, and
genae blackish; metathoracic subspiracular spots with minute but distinct circular
whitish areas around proximal end of setae; head blackish, paler along vertical
furrows, epicranial arms, and postclypeus; body with a row of blackish spots from
mesothoracic to penultimate segment along supraspiracular and subspiracular lines;
tenth abdominal tergum black; third abdominal segment with the following setal
map: 1, 3, 3, 0, 2, 4, 4, 5-6, 8-9, 6-8; subspiracular spot involving few setae directly
ventrad of spiracles; head in younger specimens entirely blackish and spots on
subspiracular lines wanting; maxillary palpi, (2, 3), 1, 4; length of body, 18 mm.;
width of head, 1.3 mm.; on SalLx; Y-151-1-1,-151-1-3; M-100,-261. (The specimens
in the Maine collection are practically identical with my specimens except in the
number of setae on subspiracular and surpedal lobes, which may exceed the number
given byoneor two) /lo/cAice MacGillivray.

4(3) Mesothoracic supraspiracular spots not subequal to but distinctly larger, twice or
more, than subspiracular spots; supraspiracular spots of abdominal segments in-
volving postsupraspiracular tubercles; prothoracic supraspiracular spots large and
distinct; preclypeus, labrum, and genae not blackish but pale brown; metathoracic
subspiracular spots without minute but distinct whitish circular areas around the
proximal end of setae; head black, paler along vertical furrows, epicranial arms,
clypeus, labrum, and genae; body with a row of blackish or brownish spots from
prothorax to penultimate segment along supraspiracular and subspiracular lines:

391 LARVAE OF THE TENTHREDINOIDEA — YUASA 73

tenth abdominal tergum black; third abdominal segment with the following seta I
map: 2, 2, 3, 1, 2, 4, 4-5, 7-8, 8-9, 9-10; mesothoracicsupraspiracular spot more than
twice as large as subspiracular spot; prothoracic supraspiracular spots moderately
large and distinct; subspiracular spot involving few setae in caudal portion of sub-
spiracular tubercle; mesothoracic and metathoracic subspiracular spots without circu-
lar whitish areas around the proximal end of setae; maxillary palpi usually 3, 2, 1, 4;
head in young specimens pale brownish, with a row of very smaU supraspiracular
spots; head in older larvae blackish, body spotted like mature specimens but spots
smaller; on Populus; length of body, 21-23 mm.; width of head, 2-2.1 mm.; Y-172-1-1.

Trichiocampus sp. 1.

Priophorus Dahlbom

Larvae small, hairy; length less than 17 mm.; body with dorsal half,
at least in part, usually with grayish or olivaceous shade; never with
spots; annulet 1 with a transverse row of several setae; annulet 2 with
two warts, each bearing 4-5 setae; annulet 3 with three warts, dorsal two
bearing 5-6 setae each, ventral with 8-10 setae; postsupraspiracular
wart usually with three setae; subspiracular lobe with 12-15 setae and sur-
pedal lobe with 6-9; setae usually more or less straight, usually of two
dififerent lengths, longer ones usually less than one-half the height of the
head; warts bearing setae of two varying lengths, those on annulet 4
being among the shortest setae.

SPECIES OF PRIOPHORUS

1(2) Front with a distinct blackish or fuscous spot; vertex with a dorso-mesal fuscous spot
occupying nearly the entire space between vertical furrows; body never pinkish but
whitish; head caudad of ocellarae fuscous; body dorsad of spiracular lines from meso-
thorax to penultimate segment olivaceous or grayish, color becoming dilute on caudal
segments; third abdominal segment with following setal map: 4-5, 4, 5, 1, 3, 6, 8, 12,
13, 8-9; maxillary palpi 2, (3, 4), 1; in younger specimens dorsal grayish shade con-
fined to cephalic segments; on hazel; length, 15 mm.; width of head, 1.5 mm.; M-109.

modestius MacGiUivray.

2(1) Front without a distinct blackish or fuscous spot, but with a light or pale brown spot;
vertex with a dorso-mesal fuscous or blackish spot, not nearly occupying the entire
space between vertical furrows; body sometimes pinkish; not on hazel 3.

3(4) Head brown ; vertex with blackish spot occupjing about one-half the distance between
vertical furrows; body pinkish; head with blackish spot caudad of ocellarae; dorsum of
mesothorax to first abdominal segment shaded gray, prothorax whitish; tliird
abdominal segment with the following setal map: 5, 4, 4, 1, 3, 5, 5, 10, 10-12, 6-2;
maxillary palpi, 2, (3, 4), 1; in younger specimens body whitish; length, 14 mm.;
width of head, 1.2 mm.; on SalLx; Y-154-1-2 palliolatus MacGillivray.*

4(3) Head pale brown; vertex on dorso-meson with blackish spot occupying two-thirds
the distance between vertical furrows; spot also caudad of ocellarae; body not
pinkish but whitish, dorsad of spiracular lines from mesothorax to penultimate

*This species was described as Trichiocampus palliolatus n. sp. by Dr.MacGillivray in the
Entomological News, vol. XXXII, 1921, page 49, but the characters of the larva place it in the
genus Priophorus. Upon reexamination of the adult specimens. Dr. MacGillivray agrees
with me in the change I here make.

74 ILLINOIS BIOLOGICAL MONOGRAPHS (392

segment distinctly and uniformly olivaceous-gray; third abdominal segment with
following setal map: 5-6, 5, 5, 1-2, 3, 6, 6, 10, 15, 8-9; maxillary palpi, 2, 3, (4, 1);
in younger specimens black spots on head larger and olivaceous shade on dorsum of
body restricted to cephalic segments; length, 16 mm.; width of head, 1.6 mm.;
on Prunus virginicus; Y-138-3. (This species resembles P. solitaris according to Dyar's
description but the latter feeds on Alnus, Prlophorus sp. 1.

Cladius Rossi

Larvae rather small; length less than 15 mm.; body slightly flattened,
greenish or yellowish green; never with spots or shaded on dorsal half; annu-
let 1 with a transverse row of several setae, annulet 2 with two warts bearing
5-6 setae, annulet 3 with three warts, dorsal two bearing 6-7 setae, ventral
with 10 setae; postsupraspiracular tubercles with 5-6 setae; subspiracular
lobe with 14-17 setae, surpedal lobe with 10 setae; setae usually straight,
usually of two lengths, longer ones less than half the height of head; setae
on annulet 1 among the shortest.

Cladius pectinicornis Fourcroy. — Length, 12-14 mm.; head pale brown-
ish or yellowish, microscopically verrucose, with brownish spots; front
touched with light brown; body hairy, uniformly greenish or greenish
yellow; third abdominal segment with the following setal map: 5-6, 5, 6,
1, 5-6, 6, 7, 10, 14-17, 10; maxillary palpi, (2, 3), 1, 4; on Rosa; Y-3, M-244.

Subfamily Nematinae

Larvae (Fig. 18) small to moderately large; body cylindrical, slender, or
abdomen increasing in diameter; segmentation and annulation usually
distinct; third abdominal segment with 4, 5 or 6 annulets, annulets 1, 2, 3,
or 1, 2, 4, or more usually 2 and 4, setiferous; larvapods present on abdom-
inal segments 2-7 and 10, setiferous or sometimes glabrous; ventral glands
present on the meson of abdominal segments 1-7; thoracic legs normal in
form; body uniformly greenish or darker colored, striped or spotted,
tuberculate, setiferous or smooth; antennae with four segments, conical,
subconical, limpet-shaped, or flattened; antennal segments sometimes
incomplete or in part fused together; tenth abdominal tergum with or
without a pair of caudal protuberances; glandubae sometimes distinct,
conspicuous, and stalked; spiracles winged or not; larvae free leaf -feeders,
gall-makers, and leaf-rollers.

The Nematinae is a large subfamily of several genera and numerous
species and is characterized by the coalescence of the cells 2d A and 3d A
due to the atrophy of the free part of the 3d anal vein. The absence of the
radial cross-vein and the cell 1st 2d A distinguishes the adults of this
subfamily from those of the Hoplocampinae and Cladiinae respectively.
Rohwer (1912), who would unite the Nematinae, Hoplocampinae, and
three genera of the Cladiinae in one subfamily, Nematinae, states that the

393] LARVAE OF THE TENTHREDINOIDEA—YUASA 75

subfamily contains two types of larvae and that most of the aberrant larvae
belong to his tribe Hemichroini. It may be pointed out that the Nema-
tinae as defined by MacGillivray also contains two types of larvae, which
are separable on the presence or absence of the caudal protuberances on
the ultimate segment. There are, however, other morphological and
biological characters of the larvae which suggest that this subfamily con-
tains a number of genera of wide diversity, and that such genera as Ptero-
nidea and Pontania might profitably be subdivided into more genera.

GENERA OF NEMATINAE

1(18) Tenth abdominal tergum without caudal protuberances 2.

2(11) Antennae conical or subcorneal, antenna! segments 2 and 3 always complete, segment
4 peg-like or conical, at least as long as wide at proximal end; third abdominal
segment always with six annulets 3.

3(4) Annulets 2 and 4 glabrous; larvapods glabrous; glandubae obsolete; thoracic legs
with coxae in part always colored brownish Nematus Panzer.

4(3) Annulets 2 and 4 not glabrous; larvapods usually not glabrous; glandubae usually
not obsolete; thoracic legs with coxae usually in part not colored 5.

5(6) Antennae with segment 3 ring-like, its cephalic portion subequal in length to caudal
portion, segment 2 complete, its cephalic portion not reduced to a mere line; body
increasing in diameter to abdominal segments 5-6; spiracles usually winged; larva-
pods glabrous or with 2-4 or more setae as viewed from lateral aspect

PrisHphora Latreille.

6<5) Antennae with segment 3 not ring-like, its cephalic portion not subequal in length
to caudal portion; segment 2 usually complete but its cephalic portion reduced to a
mere line 7.

7(8) Body increasing in diameter to abdominal segments 5-6, not uniformly cylindrical;
spiracles usually winged; larvapods with 4-6 setae as viewed from lateral aspect.

Diphadnus Hartig.

8(7) Body not increasing in diameter to abdominal segments 5-6 but uniformly cylindrical;
spiracles never winged; larvapods with 1-2 setae as viewed from lateral aspect 9.

9(10) Thorax distinctly swollen; head pale brownish green; maxillary palpi with segment 2
as long on its lateral aspect as on its mesal aspect; legs with femur and tibia con-

colorous with body, whitish; body with dorsum not shaded bluish green

Pkronidea Rohwer (in part).

10(9) Thorax never distinctly swollen; head not pale brownish green but blackish; maxillary
palpi with segment 2 three times as long on its lateral aspect as on its mesal aspect;
legs with femur and tibia not concolorous with body but blackish; body with dorsum
shaded bluish green Lygaeonematus Konow.

11(2) Antennae not conical or subcorneal, but flattened; antennal segments 2 and 3 not
always complete, segment 4 never peg-like or conical, never as long as wide at
proximal end; third abdominal segment not always with six annulets 12.

12(13) Segments with four annulets, annulets 1, 2, and 3 setiferous; gall-makers

Pontania Costa (in part).

13(12) Segments with more than four annulets, annulet 1 not setiferous; not gall-makers . . 14.

14(15) Segments with five annulets, annulets 2 and 3 setiferous; antennae with all segments
fused together; larvapods with two setae as viewed from lateral aspect; body-setae
very long Micronenmlns Konow.

76 ILLINOIS BIOLOGICAL MONOGRAPHS [394

15(14) Segments with five or six annulets, annulets 2 and 4 setiferous; antennae usually not
with all segments fused together; larvapods usually with many more than two
setae as viewed from lateral aspect; setae on body not very long; glandubae usually
conspicuous and stalked 16.

16(17) Segments with six annulets; larvapods with 7-10 setae as viewed from lateral aspect;

surpedal areas at least in part always marked with gray; latus of abdomen never

with numerous brownish spots; tenth abdominal tergum sometimes distinctly

pointed and produced caudad and with many conspicuous glandubae near the caudal

margin; larvae usually feeding on monocotyledonous plants. . .Pachynematus Konow.

17(16) Segments with five annulets; larvapods with 3-5 setae as viewed from lateral aspect;
surpedal areas not marked with gray; latus of abdomen sometimes with numerous
brownish spots; tenth abdominal tergum never distinctly pointed and produced
caudad; larvae usually feeding upon willow ^«/aMroMewo/«^ Konow (in part).

18(1) Tenth abdominal tergum with a pair of caudal protuberances 19.

19(20) Gall-makers and leaf-rollers; segments with four annulets; annulets 1, 2, and 3
setiferous; antenae flattened; body setae more than twice as long as spiracles;
caudal protuberances normal in form and position or loidimentary or borne on a
small caudo-mesal projection, if normal in form and position, the tergum with
paired colored markings Pontania Costa (in part) .

20(19) Free leaf-feeders; segments usually with five or sLx annulets, if four, annulets 2 and 4
usually setiferous; antennae conical or flattened 21.

21(22) Antennae conical, segment 2 complete, segment 3 ring-like, its cephalic portion
subequal in length to caudal portion; body on latus with eleven conspicuous blackish-
brown spots, surpedal and subspiracular areas of abdominal segments 1-9 and venter
between larvapods of abdominal segments 2-8 similarly marked; spiracles not winged;
caudal protuberances of ultimate segment small, blunt, not longer than wide at
proximal end; segments with five or four annulets, with annulets 2 and 3 or 2 and 4
setiferous Croesus Leach.

22 (2 1 ) Antennae conical or flattened ; if conical, segment 3 not ring-like but reduced in length
on cephalic aspect; body not marked as in Croesus; spiracles winged or not winged;
caudal protuberances of ultimate segments not blunt, minute, but usually much
longer than wide at proximal end 23.

23(24) Segments apparently with five annulets; annulets 2 and 3 setiferous; antennae
flattened, with segment 3 complete; body-setae subequal in length to spiracles; smaller
larvae, length, 13-15 mm Amauroncmatus Konow (in part).

24(23) Segments with four or six annulets; annulets 2 and 4, rarely 1, 2, and 3, setiferous;
antennae conical or flattened, segment 3 complete or incomplete; body-setae often
longer than the length of abdominal spiracles; small to moderately large larvae,
length, 15-23 mm Pteronidea Rohwer (in part).

DiPHADNUS HARTIG

Larvae small, greenish; length less than 14 mm.; body cylindrical, in-
creasing in size to abdominal segments 5-6, tapering at each end; thorax not
swollen; tenth abdominal tergum without a pair of caudal protuberances;
third abdominal segment with six annulets, annulets 2 and 4 setiferous;
spiracles distinctly and usually equally winged; head marked with black or
brown streaks along epicranial stem and dorsad of ocellarae; somewhat
compressed cephalo-caudad; labrum with median emargination broad and
deep and with median longitudinal depression; antennae distinctly conical,

395) LARVAE OF THE TENTHREDINOWEA—YUASA 77

with four segments, segment 1 incomplete, segment 2 usually complete
tho reduced to a mere line on cephalic part, segment 3 narrowed on cepha-
lic part, segment 4 short, conical, or peg-like; larvapods setiferous, setae
4-6 in number as viewed from lateral aspect; anal larvapods rather large;
glandubae sessile; cuticle microscopically spinulate; suranal and subanal
lobes with numerous setae; larvae free leaf -feeders.

Diphadnus append! culatus Hartig. — Length, 13 mm.; width of head,
1.35 mm.; body green, venter glossy white; tenth abdominal tergum not
marked; head greenish white with blackish brown streak along epicranial
stem, except near the occiput, and continuing to the dorsal two-thirds of
front, vertex dorsad of each ocellara not quite reaching the epicranial
suture; surface of head with minute brownish spots; following parts
brown: mandibles at apices, antennae, cervical sclerites, coxae at proximal
third, tarsal claws, and spiracles; setae minute, with conspicuous calices,
blackish; glandubae smaller at distal end than the calyx of seta; larvapods
on cephalic and lateral aspects with 4-6 setae and with a single ventral
glanduba; annulation, 1, (6, 2), (3, 5), 4; subspiracular lobe with 5-6
setae and without glandubae; surpedal lobe with about 4 setae and one
subsessile glanduba; on gooseberry; Y-158, -159, M-128.

Pristiphora Latreille

Larvae small, greenish; length less than 15 mm.; body cylindrical
usually slightly enlarged at abdominal segments 5-6, tapering at each
end; thorax not swollen; tenth abdominal tergum without the paired
caudal protuberances; third abdominal tergum with six annulets, annulets
2 and 4 microscopically setiferous, setae sometimes obsolete; annulet 1
always longest and annulet 3 and 4 always shortest; spiracles usually
winged, caudal wing usually much smaller than cephalic; head marked
usually with a blackish or brownish streak along epicranial stem, surface
with minute brown spots; labrum with distinct mesal emargination and
longitudinal depression; antennae conical or limpet-shaped, usually with
four distinct segments, segment 1 always minute, incomplete on caudal
side, segment 2 complete, but usually narrower on cephaUc side, segment 3
uniform in length, segment 4 short, conical; larvapods usually glabrous
and with a single ventral glanduba, if setiferous, setae microscopic, 2-4
in number as viewed from side; anal larvapods rather conspicuous, glandu-
bae sessile or stalked; cuticle microscopically spinulate, suranal and
subanal lobes with numerous setae; tenth abdominal tergum as seen
from side notched dorsad of suranal lobe; free leaf-feeders.

SPECIES OF PRISTIPHORA
1(8) Larvapods setiferous; antennae always conical; head always with a brownish streak
dorsad of each ocellara; spiracles always with cephalic wing distinctly larger than

78 ILLINOIS BIOLOGICAL MONOGRAPHS |396

caudal wing; setae with calices surrounded by minute brownish areas; glandubae
always sessile, sometimes microscopic, diameter never more than half the diameter
of calices of setae 2.

2(3) Glandubae minute but distinct, about one-third as large as calices of setae or micros-
copic in smaller specimens; annulation, 1, (5, 6), 2, 3, 4; subspiracular area with 3-4
setae; surpedal area with 3-4 setae; larvapods with 2-4 setae as viewed from side;
head light brownish-green, marked with fuscous streaks on both sides of entire
length of epicranial stem, front with dorsal two-thirds light brown; following parts
light brown: labrum, antennae, mandibles, cervical sclerites, tarsal claws, and spir-
cles; body greenish, caudal segments pinkish or bluish, dorsal vessel dark green with
distinct fine white line on eash side; Length, 14 mm.; on Salix; Y- 143-2-2,-155, M-96.

murtfddtiae Marlatt.

3(2) Glandubae microscopic, difficult to detect 4.

4(5) Second annulet of abdominal segments always longer than either fifth or sixth
annulet; larvapods with about two setae as viewed from side; subspiracular areas
with 4-5 setae, surpedal areas with 4 setae; annulation, 1, (5, 6), 2, 4, 3; on birch;
M-24 Prisliphora sp, 1.

5(4) Second annulet of abdominal segments always shorter than either annulet 5 or 6;
larvapods with 1-2 or 2-4 setae as viewed from side; subspiracular areas with four
setae; on alder or willow 6.

6(7) Larvapods with 1-2 setae as viewed from side; annulation 1, 2, (4, 5, 6), 3; surpedal
areas with 3-5 setae; body green, cylindrical; head pale brownish-green, marked with
brown streak along entire length of epicranial stem and dorsal two-thirds of front
and vertex, narrowly dorsad of each ocellara; following parts brown: antennae,
labrum, mandibles, maxillary palpi, cervical sclerites, tarsal claws, and spiracles;
antennae conical, segment 4 short, conical; spiracle with cephalic wing distinctly
longer than caudal; setae without brown areas surrounding calices; glandubae
minute, sessile, microscopic; M-74 Pristiphora sp. 2.

7(6) Larvapods with 2-4 setae as viewed from side; surpedal areas with 3-5 setae; annula-
tion, 1, 2, 6, 5, 4, 3; on wolly willow; M-90 Pristiphora sp. 3.

8(1) Larvapods glabrous; antennae conical or limpet-shaped; head with or without a dark

I streak dorsad of each ocellara; spiracles winged or not winged, if winged, wings

subequal in size, or cephalic wings larger than caudal; setae usually with calices

surrounded by minute but distinct brownish areas; glandubae sessile or stalked,

sometimes microscopic 9.

9(10) Head distinctly and uniformly brownish; spiracles indistinctly winged; all setae
ventrad of subdorsal lines with calices surrounded by minute brownish areas; an-
tennae conical; glandubae subequal to calices in diameter; annulation, 1, 5, 6, 2, 3, 4;

on birch; M-132 Pristiphora sp. 4.

10(9) Head never distinctly and uniformly brownish 11.

11(12) Body on dorso-meson from prothorax to fifth abdominal segment with a distinct
blackish line; spiracles winged, cephalic wings usually larger than caudal; all setae
ventrad of subdorsal lines with calices surrounded by minute brownish areas; an-
tennae conical; glandubae minute, about half as large as calices in diameter; head
with distinct line along epicranial stem; vertex without a dark streak dorsad of each

ocellara; annulation, 1, 2, (5, 3, 4, 6) or 1, 2, (3, 4, 5, 6) ; on Prunus; M-128

Prisliphora sp. 5.

12(11) Body on dorso-meson from prothorax to fifth abdominal segment without a distinct
blackish line 13.

13(14) Head entirely pale or light brown; vertex without dark streak along epicranial stem,
antennae distinctly conical; spiracles winged, wings subequal in size; setae never with

397] LARVAE OF THE TENTHREDINOIDEA—YUASA 79

calices surrounded by distinct brownish areas; glandubae probably obsolete, micro-
scopic; annulation, 1, 5, 6, 2, 3, 4; on oak; M-20 Pristiphora sp. 6.

14(13) Head never entirely pale or light brown; vertex always with a dark streak along
epicranial stem, sometimes indistinct but never entirely wanting 15.

15(22) Spiracles winged ; setae ventrad of spiracular lines with calices surrounded by distinct
minute brownish areas; antennae always conical; vertex with or without a dark
streak dorsad of each ocellara 16.

16(19) Spiracles with cephalic wings always larger than caudal; glandubae always sessile;
glandos microscopic, difficult to detect; setae on latus dorsad of spiracular lines
with calices surrounded or not surrounded by distinct brownish areas 17.

17(18) All setae on latus dorsad of spiracular lines with calices surrounded by brownish
areas; vertex dorsad of each ocellara with a brownish streak; annulation, 1, (5, 6),
2, 4, 3; body green; head marked as in P. hivittata; length of body, 12-13 mm.; on
Salix; M-72 sychophanta Walsh.

18(17) All setae on latus dorsad of spiracular lines with calices not surrounded by brownish
areas; vertex dorsad of each ocellara without a brownish streak, uniformly pale;
annulation, 1, (2, 5, 6), 4, 3; body enlarged on abdominal segments 5-6, greenish,
with dorsal vessel dark green with white line on each side; head greenish with brown-
ish streaks along entire length of epicranial stem, expanding on the dorsal half of
front; following parts brown: labrum, mandibles, antennae, cervical sclerites, tarsal
claws, and spiracles; antennae conical, segment 4 elongate, conical; on Spiraea latifolia
and S. iomentosa; M-4 Pristiphora sp. 7.

19(16) Spiracles with wings subequal in size; glandubae stalked ; glandos subequal in diameter
to calices; setae on latus dorsad of spiracular lines never withcalices surrounded by
distinct brownish areas 20.

20(21) Vertex with a distinct dark streak dorsad of each ocellara ; annulation, 1, 2, (5, 6), 4, 3;
length, 16-17 mm.; body cylindrical, increasing in size to abdominal segments 5-6,
green, dorsal vessel dark green with fine white line on each side of it; head green,
slightly brownish, with fuscous streaks; following parts light brown : dorsal two- thirds
of front, labrum, maxillary palpi and galea, labial palpi, and cervical sclerites; follow-
ing parts brown: antennae, mandibles, spiracles, glandubae, and setae; antennae
conical, distinctly with four setae, segment 4 conical; spiracles with wings; ven-
tral glands subequal in size to larvapods; glandos in diameter subequal to or larger

than calices of setae; setae on dorsimi very minute; on Spirea; Y, M-14,-88

bivittata Norton.

21 (20) Vertex without a distinct dark streak dorsad of each ocellara; annulation, 1, 2, 6, 5, 3,
4; on Potentilla; M-10 Pristophora sp. 8.

22(15) Spiracles not winged; setae ventrad of spiracular lines never with calices surrounded
by distinct minute brownish areas; antennae limpet-shaped, segments indistinguish-
ably fused; vertex always with a brown streak dorsad of each ocellara; glandubae
sessile; glandos microscopic, difficult to detect; annulation, 1, (2, 5, 6), (3, 4); length
12-13 mm.; body green, with broad dark green dorsal vessel bordered on each side
by a fine white line; head greenish brown; setae stifiF, comparatively long, without
brown areas surrounding calices; on Geum canadensis; Y -2X2. .Pristiphora sp. 9.

MiCRONEMATUS KONOW

Larvae small; length less than 15 mm.; body subcylindrical, tapering
at both ends; mesothorax distinctly and metathorax slightly swollen;
lateral lobes prominent and swollen; segmentation distinct; annulation
indistinct; third abdominal segment with five annulets, annulets 2 and

80 ILLINOIS BIOLOGICAL MONOGRAPHS (398

3 setiferous; thoracic legs spreading laterad, normal in form; larvapods
setiferous, normal in form, except the anal pair, which is reduced in size,
only one-half as large as the other pairs; tenth abdominal tergum with-
out the paired caudal protuberances; head circular, smaller than thorax
in width and height, front flattened; antennae apparently with four seg-
ments, flattened, all segments fused together; ventral glands very large;
setae sparse, very long; spiracles not winged; suranal and subanal lobes
multisetiferous; glandubae subsessile; free leaf-feeders.

Konow in 1905 listed three species of Micronematus: abbreviatus,
calijornicus, and monogyniae. The second species properly belongs to
Diphadnus. Of the two remaining species, M. abbreviatus was recognized
in the larval stages a long time ago by Snellen von Vollenhoven (1868).
This genus is European and is represented in North America by a single
species, Micronematus gregarius Marlatt. That this species does not
belong to Pachynematus can be readily seen from the structure and
biological characters of the larvae as was suggested by Dyar (1897)
in his original description of the immature stages. Three facts distinguish
this species from all other known species of Pachynematus: (1) the larva
has five annulets instead of six; (2) the anal larvapods are very much reduced
in size; and (3) the larva feeds on willow instead of grasses. On the other
hand M. gregarius has certain characters in common with M. abbreviatus
as recorded by Vollenhoven. Since this species can not be referred to
Pachynematus or to any other known American genus, and since it has
characters which are peculiar to Micronematus, and since the adult
characters, according to Ashmead (1898), would place it in this genus,
this species is here considered as belonging to Micronematus. If future
study should prove this position untenable, a new genus should be erected,

Micronematus gregarious Marlatt. — Length, 12 mm.; body shiny,
yellowish white; alimentary canal showing thru as green tube; head pale
testaceous with a broad blackish band across the front between ocellarae
and a narrow band dorsad of each ocellara to and along vertical furrow;
mouth-parts, cervical sclerites, and legs except coriae brownish; abdominal
segments 1-8 with postspiracular and subspiracular areas swollen, mound-
like, and tinted fuscous; abdominal segments 2-7 with colored postspiracu-
lar areas larger than subspiracular areas; those on segments 1 and 8 much
smaller; thoracic segments with two colored patches on latus, one larger
and more ventral than the other; larvapods with about two setae on the
cephalo-lateral aspect; ventral glands nearly three times as large as larva-
pods; setae slender, at least twice as long as spiracles; spiracles not winged;
subspiracular area with two setae and single glanduba, surpedal area
with 3-4 setae and single glanduba; larvae gregarious; on Salix; Y.

399] LARVAE OF THE TENTHREDINOIDEA—YUASA 81

Lygaeonematus Konow

Larvae comparatively speaking moderately large, length 18-20 mm.;
body cylindrical, tapering uniformly and slightly caudad; thorax not
swollen; abdomen never swollen; dorsum grayish green and venter pale,
head shiny black, only slightly narrower than thorax; antennae conical,
with four distinct segments, segment 1 minute, incomplete, and with single
sensory pit, segments 2 and 3 narrower on cephalic part than on caudal
part, segment 4 conical; labrum with mesal emargination shallow and broad;
maxillae with galea larger than labial palpi; maxillary palpi with segment 2
three times as long on lateral margin as on mesal; third abdominal segment
with six annulets, annulet 1 largest, annulets 2 and 4 setiferous; larvapods
well developed, setiferous, with 1-2 setae and single ventral glanduba;
glandubae sessile; glandos smaller than calyx of a seta; ventral glands
subequal to or larger than larvapods in size; spiracles not winged; tenth
abdominal tergum as seen in profile not notched dorsad of suranal lobe and
without caudal paired protuberances; setae usually arising from minute
fleshy mound-like protuberances; cuticle microscopically spinulate; free
leaf-feeders.

Lygaeonematus erichsoni Hartig. — Length, 18-20 mm.; body somewhat
shiny, greenish gray dorsad of spiracular lines, ventrad of them opaque
bluish- white, head black; following parts fuscous to blackish: mouth-parts,
cervical sclerites, femur, tibia, and claws; following parts grayish: surpedal
areas, coxa, and trochanter in part, swellings between legs, and abdominal
surpedal areas faintly; annulation, 1, 2, 6, (3, 5, 4); glandos half as large
in diameter as caUces of adjacent setae; surpedal areas on abdomen with
8-10 setae and two glandubae; subspiracular area with 6-7 setae, usually
without glandubae; cuticle with distinct microscopic brownish spinulae;
on larch; Y-162, M-105, -165.

Pachynematus Konow

Larvae comparatively speaking moderately large; length, 15-23 mm.;
body cyhndrical, tapering uniformly and distinctly caudad; thorax rarely
and abdomen never swollen; third abdominal segment with six annulets,
annulets 2 and 4 setiferous, annulet 1 longest, annulet 4 usually shortest;
tenth abdominal tergum without a pair of caudal protuberance, and without
a notch dorsad of suranal lobe as seen in profile; the tergum sometimes
produced distinctly caudad and pointed; suranal and subanal lobes with
numerous setae; head brownish or greenish, sometimes with fuscous
streak along epicranial stem, semiglobose, subequal in width to thorax
or only slightly narrower; labrum with median longitudinal depression,
cephalic emargination shallow; maxillae with galea always larger than
labial palpi, sometimes more than twice as large; antennae with four seg-
ments, flattened, never distinctly conical, segment 1 usually minute, in-

82 ILUNOIS BIOLOGICAL MONOGRAPHS [400

complete, segments 2 and 3 with cephalic parts distincty reduced to mere
lines, sometimes all four segments fused; larvapods always setiferous, with
8-11 setae on cephalic and lateral aspects and a single ventral glanduba;
glandubae conspicuous, cylindro-conical, at least twice as long as wide
at proximal end; spiracles usually not winged, if winged, rather indistinctly;
setae stiff, brown, arising from fleshy mound-like minute tubercles; in
younger specimens third abdominal segment with five annulets, annulets
1, 2, and 3 with stiff brown setae; glandubae obsolete; cuticle microscopi-
cally spinulate; larvae usually feed upon grasses.

This generic description is based upon two identified and several un-
identified species. The known species can be separated as follows:

SPECIES OF PACHYNEMATUS

Head with narrow fuscous or blackish streak along epicranial stem and vertical furrows;
tenth abdominal tergum with a broad fuscous streak on the meson and the caudal margin
produced distinctly caudad and bluntly pointed, with many conspicuous glandubae; antennae
with four fused segments; annulation, (2, 1), 6, (3, 4, 5); larvapods with 7-9 setae on cephalic
and 1 seta on lateral aspect; subspiracular lobe with 5-6 setae and 1-2 glandubae; surpedal
lobe with 5-7 setae and 1-3 glandubae; body on dorso-lateral lines with narrow, interrupted
longitudinal fuscous, bands; head as wide as thorax; length, 20 mm. width of head, 2 mm.;

on Carex; Y-150 subalbalus Norton.

Head with narrow fuscous or blackish streak along epicranial stem and vertical furrows;
tenth abdominal tergum without a broad fuscous streak on the meson, the caudal margin
only slightly produced caudad, rounded, with many glandubae; antennae with four segments
not fused; annulation, (1, 2), 6, (3, 4, 5) ; larvapods with about ten setae on cephalic and one se-
ta on lateral aspect; subspiracular lobe with about six setae and a single glanduba; surpedal
lobe with six setae and three glandubae; head as wide as thorax; length, 16 mm.; width of
head, 1.6 mm.; on Carex; Y-177 repertus MacGillivray.

Nematus Panzer

Larvae comparatively small, length about 10 mm.; body cylindrical,
uniform in diameter thruout the entire length excepting the caudal end,
where it is tapering, dull green, apparently glabrous; annulation indis-
tinct; third abdominal segment with six annulets, annulets glabrous,
formula: 1, 5, 2, 6, 3, 4; thoracic legs normal in form; coxa usually fus-
cous on proximal half; larvapods small, diminishing in size gradually on
the caudal segments, not setiferous; ventral glands subequal in size to larva-
pods; tenth abdominal tergum without a pair of caudal protuberances,
with a notch dorsad of suranal lobe as seen in profile, caudal margin trun-
cate or slightly emarginate on the meson, not produced; suranal lobe with
several minute setae on ventral aspect; subanal lobe with moderately
numerous minute setae; abdomen with lateral lobes inconspicuously
swollen; head circular in outline, rounded in profile on dorsal half, front
flattened; ventral half of head with several longer and larger setae; dorsal
half nearly glabrous or with microscopic setae; vertex usually with a fuscous

401] LARVAE OF THE TENTHREDINOIDEA—YUASA 83

Streak caudad and dorsad of each ocellara; antennae four-segmented,
subconical or sublimpet-shaped, sometimes in part fused; segment 1
minute, segments 2 and 3 reduced to mere line on cephalic aspect, segment
4 minute, subconical; mouth-parts and cervical sclerites fuscous to black-
ish; maxillary palpi with two distal segments distinctly smaller than
segment 2, which is sometimes longer than all the other segments taken
together; galea larger than labial palpus; sericos distinct, circular; sub-
spiracular and surpedal lobes with a few microscopic setae; glandubae
probably microscopic, usually obsolete; spiracles usually winged; cuticle
microscopically spinulate; free leaf-feeders.

SPECIES OF NEMATUS

1(4) Vertex with a fuscous streak caudad and dorsad of each ocellara; maxillary palpi
with segment 2 shorter than all the other segments taken together 2.

2(3) Vertex with fuscous streaks extending dorsad of vertical furrows; antennae with all
segments fused and almost completely filling antafossae; spiracles distinctly winged;
tenth abdominal tergum with caudal margin shallowly emarginate on the meson;
head with dorsal half entirely glabrous ; length, 1 1 mm. ; on oak ; M-21 . . chloreus Norton.

3(2) Vertex with fuscous streaks not extending dorsad of vertical furrows; antennae with
all segments distinct, but not nearly completely filling antafossae; spiracles not
distinctly winged; tenth abdominal tergum with caudal margin not emarginate on the
meson; head with dorsal half sparsely setiferous; length, 9 mm. ; Y-133. Nemalus sp. 1.

4(1) Vertex without a fuscous streak caudad and dorsad of each ocellara; maxillary palpi
with segment 2 slightly longer than all the other segments taken together; antennae
with four distinct segments; spiracles winged; length, 10.5 mm.; on oak; M-2.

Nematus sp. 2

Croesus Leach

Larvae moderately large, length about 25 mm., body cylindrical tapering
caudad on abdominal segments 7-10 and also at cephalic end of prothorax;
segmentation distinct; annulation indistinct; third abdominal segment
with four or five annulets, annulets 2 and 3 setiferous; tenth abdominal
tergum with a pair of low, conical, bluntly rounded caudal protuberances,
usually as wide as or wider than high; head flattened on front, rounded,
shiny black; antennae distinctly conical with four segments: segment 1
incomplete, three times as long as wide, segments 2 and 3 usually complete
and not reduced to mere lines on the cephalic aspect, segment 4 peg-like,
bluntly pointed, longer than wide at proximal end; maxillary palpi with
segment 1, 3, and 4 subequal in length, segment 2 twice as long as segment
1 ; larvapods setiferous; spiracles not winged, indistinctly colored; glandubae
subsessile or with a short stalk, very wide in diameter more than three
times the diameter of shaft of setae; maxacoria developed into a distinct
triangular swelling dorso-caudad of cardo, and covered with dark spinulae.

Croesus latitarsus Norton. — Length, 25 mm.; annulation, 3, 2, 1, 4 or
2, 1, 5, 3, 4; larvapods with 4-5 setae and a single glanduba as viewed
from side, no markings; tenth abdominal tergum with dark-colored area

84 ILLINOIS BIOLOGICAL MONOGRAPHS [402

contiguous to concolorous suranal protuberances; suranal and subanal
lobes with numerous setae, which increase in length with the distance
from the anus; body yellowish green or brownish, with colored patches on
subspiracular and surpedal areas and on subdorsal line from mesothorax
to ninth abdominal segment; abdominal segments 2-8 on ventro-meson
each with a large colored patch; in younger stages body fuscous with
few distinctly delimited colored patches or none; M-1, -26,-50,-5 1,-48,

Amauronematus Konow

Larvae small to moderately large; length 15-20 mm.; body cylin-
drical, often tapering distinctly caudad; latus with subspiracular and
surpedal areas colored or with numerous, minute colored spots; head
blackish, brownish, or greenish; third abdominal segment with five annu-
lets, annulets 2 and 3 or 2 and 4 setiferous; tenth abdominal tergum with
or without caudal paired protuberances; body setae much longer or
shorter than the length of spiracles; larvapods with 2-3 or 3-5 setae as
viewed from side; spiracles not winged; larvae free leaf -feeders.

The larval stages of five species of Amauronematus have been described
by Dyar. Of these, larvae of luteotergum, dyari, oregonensis, and similis
have not been available for study, but according to Dyar's descriptions,
the last three apparently lack the caudal paired protuberances. I have a
large number of larvae collected on willow by Dr. MacGillivray which
also lack the paired caudal protuberances and probably belong to this
genus. The species may be separated as follows :

SPECIES OF AMAURONEMATUS

1(4) Tenth abdominal segment with paired caudal protuberances 2.

2(3) Abdomen on ventro-meson with a row of blackish spots; caudal paired protuberances
and head blackish; gregarious; on alder luteotergum Norton.

3(2) Abdomen on ventro-meson without a row of blackish spots; paired caudal protuber-
ances and head not blackish but pale; solitary; on Azalea azaleae Marlatt.

4(1) Tenth abdominal segment without paired caudal protuberances 5.

5(6) Mature and also younger larvae with antennae flattened and their segments fused
in part, with no discernible or with very small antacoria; younger larvae (14 mm. or
less) with annulets 1, 2 and 4 setiferous; annulets 2 and 4 with transverse row of
warty protuberances each bearing 2 or 3 stiiBf stout setae; older larvae (14 mm. or
more) without numerous brownish spots on dorsum and latus and without brownish
interrupted and diffuse dorso-mesal and dorso-lateral lines; on sweet fern; M-85.

Amauronefnatus sp. 1.

6(5) Mature larvae with antennae whose segments are all distinctly separated by distinct
antacoria ; young larvae with antennae like those of preceding species; younger larvae
(14 mm. or less) with annulets 1, 2, and 4 setiferous; annulets 2 and 4 without
transverse row of warty protuberances the setae being minute and arranged singly;
older larvae (14 mm. or more) mth numerous brownish spots on dorsum and latus
with brownish, interrupted, and diffuse dorso-mesal and dorso-lateral lines; on willow;
M-10 virendus MacGillivray.

403] LARVAE OF THE TENTHREDINOIDEA—YUASA 85

Another species, resembling the preceding very closely and, difficult to distinguish except
by the colored spots which are more or less darker and shghtly more numerous than in
virendtts: M-112 vescus MacGillivray.

Pteronidea Rohwer

Larvae small to moderately large; length 15-25 mm.; greenish, often
spotted or banded transversely or longitudinally; body cylindrical, slender,
uniformly tapering caudad, thorax rarely conspicuously swollen; head and
trunk setiferous, often tuberculate; head blackish, brownish, or greenish;
antennae with four segments, sometimes with segments in part fused,
conical, subconical, or flattened; third abdominal segment with 4-6
annulets, more commonly 5-6, annulets 2 and 4, rarely 1, 2, and 3 setiferous;
tenth abdominal tergum with or without a pair of small but distinct suranal
caudal protuberances, if without, body swollen on thorax, protuberances
pointed, bluntly rounded, truncate, or swollen at distal end; larvapods
setiferous, setae few in number; spiracles winged or unwinged; glandubae
subsessile or obsolete; leaf-feeders, sometimes gregarious.

The genus Pteronidea is rich in number of species. The author has
examined a large number of specimens representing at least thirty species
and including much bred material, and has prepared the following synoptic
key for differentiating species. It may be stated here that Pteronidea,
together with a few allied genera, is readily separated from all other Tenth-
redinidae by the presence of a pair of suranal caudal protuberances on the
lateral portion of the caudal margin of the ultimate tergum. Pteronidea
thoracica Harrington is unique in lacking the caudal paired protuberances,
but is easily distinguished by its characteristic, somewhat elongate, tad-
pole-like body, and also by its white head and body and its spreading legs.
The color and coloration and the presence of setiferous tubercles and their
arrangement are useful characters in separating species.

SPECIES OF PTERONIDEA
1(2) Tenth abdominal tergum without suranal processes; thorax conspicuously swollen;

thoracic legs spreading out flat laterad; body entirely greenish white; on Primus

virginiana; Y-141 thoracica Harrington.

2(1) Tenth abdominal tergum with suranal processes; thorax not conspicuously swollen. ?>.

3(50) Head black or brov/n; body usually with numerous colored patches 4.

4(5) Body entirely blackish with distinct yellowish spots on latus; on Salix, Popitlns

balsamifera, etc.; Y-8.45, M-104, M-182, Y-5-2 ventralis Say.

5(4) Body not entirely blackish, without distinct yellowish spots on latus 6.

6(7) Body entirely green; head light brown; suranal processes short, mere swellings; on

Ribes sp.; Y-1 ribcsi Scopoli (ultimate stage)

7(6) Body not entirely green; suranal processes usuall}' distinctly pointed, more than

mere swellings g.

8(9) Body yellowish with 11 transverse black markings extending between subdorsal

lines across the venter. Young collection 55 Pkronidia, sp. 1.

9(8) Body not yellowish, without 11 transverse blackish markings 10.

86 ILLINOIS BIOLOGICAL MONOGRAPHS [404

10(17) Body without numerous small tuberculate areas, usually without fine longitudinal
series of sub-adjacent patches; thorax, at least, always suffused on dorsum with
darker shade 11.

11(12) Body shiny metallic fuscous; dorsum of all segments grayish brown; tenth abdominal
tergum uniformly black; Y-132-2-1 (on Carpinus caroliniana), Y-132m-l-2 (on
Morus alba), Y-193 (on Alnus rugosa), erylhrogaska Norton.

12(11) Body not shiny metallic fuscous; dorsum of none of segments grayish-brown; tenth
abdominal tergum not uniformly black 13.

13(14) Abdomen on dorso-meson with a fine longitudinal line; tenth abdominal tergum
fuscous on dorso-meson; suranal processes distinctly fuscous; on Alnus; M-114.

Pteronidea sp. 2.

14(13) Abdomen on dorso-meson without a fine longitudinal line; tenth abdominal tergum
not fuscous on dorso-meson; suranal processes not distinctly fuscous IS.

15(16) Tenth abdominal tergum with a fuscous triangular mark on each side of meson; head
more or less uniformly brown; on Salix rostrata; Y-169, C-427-45. .Pteronidea sp. 3.

16(15) Tenth abdominal tergum without a fuscous triangular mark on each side of meson;
head not uniformly brown but epicranial stem, vertex dorsad of ocellarae, front in the
center, and labrum distinctly darker; on hazel (?); M-153 Pteronidea sp. 4.

17(10) Body with numerous small blackish tuberculate areas ; sometimes with fine interrupted
longitudinal lines composed of irregular subadjacent colored areas; thorax on dorsum
never suffused with darker shade 18.

18(27) Body on dorso-meson with a fine more or less continuous black line independent of
greenish or pale dorsal vessel, line occasionally faint but never entirely obsolete;
latus usually with fine more or less continuous longitudinal lines 19.

19(20) Abdomen on ventro-meson with fuscous spots; larvapods on cephalic surpedal
areas never with fuscous spots; on Alnus; M-232 Pteronidea sp. 5.

20(19) Abdomen on ventro-meson never with fuscous spots; larvapods on cephalic surpedal
areas often with fuscous spots 21.

21(24) Antennae with antacoriae distinct, not limited to periphery of antafossae; antennal
segment 1 very minute, never more than twice as long as wide; segment 2 usually
incomplete, if complete, ventral portion never more than a mere faint line; segment 3
usually complete, cephalic portion reduced to a line 22.

22(23) Larvapods on cephalic aspect with minute irregular blackish or brownish spots near
setae; on Salix; Y-8.48(?)-l Pteronidea sp. 6.

23(22) Larvapods on cephalic aspect without minute irregular black spots near setae; on
Salix spp.; Y-6-1, -6-6,-44-1-1, M-156 (in part), C-140, C-649 odoratus Dyar.

24(21) Antennae with antacoriae indistinct, limited to periphery of antafossae; antennal
segment 1 complete, or if incomplete, never less than twice as long as wide; segments
2 and 3 always complete, their cephalic and dorsal portions never reduced to mere
lines; segment 4 cylindro-conical; all four segments often fused together in part and
filling antafossae almost completely; larvapods on cephalic aspect with irregular
black spots near setae, spots sometimes very minute but never wanting from all
larvapods 25.

25(26) Antennae with all 4 segments fused together in part and filling antafossae almost
completely; on Salu spp.; Y-95-1-1, Y-8.45 (?)s-l-l cornelU Marlatt.

26(25) Antennae without all 4 segments fused together in part; segment 1 always dis-
tinctly separated from the other segments; antacorriae always distinct; on Populus;
M-156 (in part) Pteronidea sp. 7

27(18) Body on dorso-meson never with a fine more or less continuous black line independent
of greenish or pale dorsal vessel; latus usually without fine more or less continuous
longitudinal lines 28.

405J LARVAE OF THE TENTHREDINOIDEA—YUASA 87

28(29) Latus with distinct more or less continuous longitudinal lines; venter on meson
without black spots; C-713 Pkronidea sp. 8.

29(28) Latus without distinct more or less continuous longitudinal lines; venter on meson
usually with black spots 30.

30(31) Larvapods always with black spots on cephalic aspect; dorsum with setiferous
black tubercles more or less uniform in size; abdomen on ventro-meson without
black spots; on Ribes sp.; Y-1; M-135 ribesi Scopoli.

31(30) Larvapods never with black spots on cephalic aspect; dorsum with setiferous black
tubercles never uniform in size; abdomen usually with black spots on ventro-meson.32.

32(33) Larvapods on abdominal segments 4-7 usually with minute black spots on mesal
aspect, those on sixth abdominal segment never wanting; abdomen on ventro-
meson without black spots; on Salix; Y-95-1; M-155 (in part); M-140

Pkronidea sp. 9.

33(32) Larvapods on abdominal segments 4-7 without minute black spots on mesal aspect;
abdomen on ventro-meson with black spots 34.

34(47) Abdominal segments 7-9 on ventro-meson with black spots 35.

35(40) Ninth abdominal tergum with colored patches on first three annulets, 4, 6, and 4
patches respectively 36.

36(39) Dorsum not shaded grayish-browTi 37.

37(38) On Populus; G-14; M-158 effeia MacGillivray,

38(37) On hazel; M-110 effusa MacGillivray.

39(36) Dorsum shaded grajnsh-brown; on Salix; Y-8.45(?)-2-l Pteronidca sp. 10.

40(35) Ninth abdominal tergum with colored patches on first three annulets 41.

41(44) Colored patches on first three annulets 2, 6 and 4 in number respectively 42.

42(43) Body small, less than 13 mm. in length; Young-49 Pkronidea sp. 11.

43(42) Body large, more than 15 mm. in length; on Populus balsa in if era; M-182; G-pop.

Pkronidea sp. 12.

44(41) Colored patches on first three annulets 2, 4, and 4 in number respectively 45.

45(46) On birch; M-139 emcrita MacGillivray.

46(45) On Populus; Y-45 lomhardae Marlatt.

47(34) Abdominal segments 2-8 on ventro-meson with black spots 48.

48(49) Large larvae, 20-23 mm. in length; dorsum without dark shade; head much smaller
than thorax in width and height; on Salix; Y-8.45(?)s-5-2. .fulvicrus Provancher.

49(48) Moderately large larvae, less than 20 mm. in length; dorsum always with dark shade;
head comparatively large, only slightly smaller than thorax in width and height; on
Salix;M-119 CTaww/a MacGillivray.

50(3) Head not black or brown, usually greenish with few linear markings; body usually
without numerous colored patches 51 .

51(54) Body with a pair of distinct fine latero-dorsal lines; larvapods with 1-2 setae as
viewed from side 52.

52(53) Head with a black line extending the entire length of epicranial stem to the occiput;
on Salix; M-12 eriidita MacGillivray.

53(52) Head without black line extending the entire length of epicranial stem to the occiput;
on Salix; M-190 Pkronidea sp. 13.

54(51) Body without a pair of distinct fine dorso-lateral lines; larvapods usually with 3-5

setae as viewed from side 55.

55(58) Head entirely green 58.

56(57) Suranal process sharply pointed ; on Rhododendron canadense; M-46

Pkronidea sp. 14.
57(56) Suranal processes bluntly rounded; on Salix; M-133 Pt^onidea sp. 15.

88 ILLINOIS BIOLOGICAL MONOGRAPHS [406

58(55) Head not entirely green, usually with a blackish or brownish line along epicranial

stem and one dorsad of each ocellara 59.

59(64) Suranal process sharply pointed 60.

60(61) Body tapering caudad uniformly and slightly; uniformly green; on Salix cordata;

Y-153-1-1,-153-? mmdica Walsh.

61(60) Body not tapering caudad uniformly and slightly; abdomen swollen on segments

5-7; body not uniformly green 62.

62(63) Latus of each segment with a large fuscous patch; tenth abdominal tergum with

caudal margin between suranal processes straight; on Alnus; M-71

equina MacGillivray.
63(62) Latus of each segment without a large fuscous patch; tenth abdominal tergum with

caudal margin between suranal processes convex; on birch; M-61. Pteronidea sp. 16.

64(59) Suranal processes never sharply pointed but enlarged at distal ends 65.

65(66) Antennae with segment 3 complete, altho reduced to a mere line on cephalic aspect;

on birch; M-60 Pteronidea sp. 17.

66(65) Antennae with segment 3 incomplete; Y-120 (on Gledilsia triacantkos); Y-143-1

on Salix cordata trilineata Norton.

PoNTANiA Costa

Larvae comparatively small, whitish or greenish, usually 10-15 mm.
in length; gall-makers or leaf-edge-rollers; body cylindrical, thorax usually
not swollen; tenth abdominal tergum usually with a pair of suranal pro-
tuberances; when the paired caudal protuberances are normal in position,
i. e., near the lateral ends of caudal margin of the segment, the tergum
usually with paired blackish or brownish markings; caudal protuberances
sometimes very minute and borne on the caudal margin of the produced
median projection; spiracles winged or not winged; third abdominal
segment with four annulets, annulets 1-3 setiferous; head usually dark
brown or blackish in younger specimens and yellowish or pale brown in
older specimens; labrum with mesal emargination shallow or obsolete;
maxillary palpi with segment 2 longest, usually nearly equal in length to
segments 3 and 4 taken together; antennae with four segments, segments
fused or separate, segments 1 and 2 usuallly incomplete and separate,
segment 3 sometimes complete, often fused with segment 4; larvapods
setiferous, with four or more setae.

Most Nematinae with gall-making or leaf-folding larvae belong to this
genus. The different types of galls are supposed to be specific and are
considered of systematic value. Many species are indistinguishable in the
immature stages except by the morphology of the galls they produce.

SPECIES OF PONTANIA
1. Gall-Makers
1(18) Tenth abdominal tergum with or without the paired caudal protuberances, if present,
small, blunt, usually not longer than wide at pro.ximal end, sometimes borne on
caudal projection of suranal lobe; tergum without paired blackish or brownish
markings ; ninth abdominal tergum with distinct paired markings or transverse rows of
minute spots 2.

407] LARVAE OF THE TENTHREDINOIDEA—YUASA 89

2(3) Tenth abdominal tergum without paired caudal protuberances; tibia distinctly
longer than femur; head brownish or yellowish with space between ocellarae across

the front paler; Y-8-4- 1,-8-4-3,-8-6-1, C-y67, M-212; on Salix poynum Walsh.

3(2) Tenth abdominal tergum with paired caudal protuberances; tibia not distinctly

longer than femur 4.

4(5) Suranal lobe not produced caudad, as a small mesal projection, but with a pair of
caudal protuberances, small but normal in position, near the lateral ends of the
caudal margin; head blackish or blackish brown, space along epicranial suture paler;
spiracles usually not winged; dorsum of segments not transversely marked with

gray; on Salix; Y-7-1,-7-4-1,-8.8, C-cu 201, M-92 hyalina Norton.

5(4) Suranal lobe produced caudad, forming a small m.esal projection which bears rudi-
mentary paired protuberances; head light brown with front and vertex dorsad of
each ocellara darker; spiracles usually winged; dorsum of segments transversely
marked with gray; antennae with segments 3 and 4 fused; labrum with mesal emar-

gination obsolete 6.

6(7) Gall not transected by the leaf, but attached to one surface, point of attachment
showing as discolored scar; greater part of gall free from the leaf; two or more galls

sometimes adjacent; woolly small-leaved willows; M-148 Pontania sp. 1.

7(6) Gall transecting the leaf; usually only one gaU on a leaf 8.

8(9) Gall involving the midrib; surface of gall irregularly constricted; sometimes 2 or

more galls adjacent; M-226 devincta MacGillivray.

9(8) Gall not involving the midrib tho extending to it 10.

10(15) Long axis of gall parallel to midrib; leaf transecting the gall into two subequal

parts 11.

11(12) Gall kidney-shaped, strongly convex, about 14 mm. in length; M-213. Pontania sp. 2.

12(11) Gall not kidney-shaped 13.

13(14) Gall bean-shaped, slightly convex; 12-14 mm. in length; Y-191-1-1

demissa McGillivray.
14(13) Gall bean-shaped, strongly convex; 14-15 mm. in length; M-262. .Pontania sp. 3.

15(10) Long axis of gall transverse to midrib 16.

16(17) Gall transected by leaf into two subequal semiglobose parts, surface not constricted

by a furrow; M-211,-216 Pontania sp. 4.

17(16) Gall transected bj'^ leaf into two unequal parts; surface constricted by a furrow;
M-93 Pontania sp. 5.

2. Leaf-folders

18(1) Tenth abdominal tergum always with the paired caudal protuberances
which are sharply pointed, normal in position, and longer than wide at proximal
end; tergum usually with paired blackish or brownish markings; ninth abdominal
tergum with or without distinct paired markings or transverse rows of minute
spots 19.

19(20) Tenth abdominal tergum without distinct paired blackish or brownish markings;
head entirely yellowish; M-l 50 Pontania sp. 6.

20(19) Tenth abdominal tergum with distinct paired blackish or brownish markings 21.

21(26) Ninth abdominal tergum without any markings or spots; antennae with segment 3
sometimes incomplete 22.

22(25) Tenth abdominal tergum with markings extending entire length and dumbbell-
shaped; antennae with segments 2 and 3 usually complete 23.

23(24) Folded portion of leaf irregularly wrinkled; usually both edges folded; M-175.
1-1 Pontania sp. 7.

24(23) Folded portion of leaf not irregularly wrinkled; usually one edge folded; Y-31-1-1,
-8, 46(?)-2-2 Pontania sp. 8.

90 ILLINOIS BIOLOGICAL MONOGRAPHS (408

25(22

26(21

27(32
28(29;

29(28

30(31

31(30

32(27
33(34;

34(33

35(36;

36(35

37(42
38(39

39(38;
40(41
41(40
42(37

Tenth abdominal tergum with markings not extending the entire length and not
dumbbell-shaped; antennae with segments 2 and 3 incomplete; folded portion of leaf

irregularly wrinkled; often both edges folded; Y-139-1-1 Pontania sp. 9.

Ninth abdominal tergum with some markings or spots; antennae with segment 3

usually complete 27.

Ninth abdominal tergum with two transverse rows of minute colored spots 28.

Tibia longer than femur; front concolorous with vertex; setae on abdominal terga

more than three times as long as spiracles; Y-166-1-1 Pontania sp.lO.

Tibia subequal in length to femur; front not concolorous with vertex but darker; setae

on abdominal terga less than three times as long as spiracles 30.

Head blackish; mature larvae 9 mm. in length; on Salix; Y-8.46(?)-l-2

Pontania sp. 11.
Head brownish; mature larvae 11 mm. in length; on Populus; M-166

Pontania sp. 12.
Ninth abdominal tergum without two transverse rows of minute colored spots .... 33.
Ninth abdominal tergum with two pairs of transverse blackish or brownish markings,
interrupted on meson; head brownish with area along epicranial suture distinctly

clear and paler; Y-142-1 derosa MacGillivray.

Ninth abdominal tergum with one pair of transverse blackish or brownish markings,
interrupted on meson; head blackish or brownish in younger specimens, yellowish
or light brown in older specimens with area along epicranial suture not distinctly

paler 35.

Tenth abdominal tergum with markings not reaching the paired caudal protuberances
but broken on caudal half into minute spots; head yellowish with two minute brown
spots on front; leaf-edge folder, folded edges not irregularly wrinkled; M-146.

Pontania sp. 13.
Tenth abdominal tergum with markings reaching the paired caudal protuberances,
not broken on caudal half into minute spots; yellowish or brownish without two

minute brown spots on front 37.

Ninth abdominal tergum with paired markings on its cephalic half 38.

Ninth abdominal tergum with a transverse row of minute spots caudad of paired
markings; leaf-edge-folder; both edges often folded and folded portion irregularly

wrinkled; M-145 Pontania sp. 14.

Ninth abdominal tergum without a transverse row of minute spots caudad of paired

markings 40.

Antennae with segment 3 incomplete; only one edge of leaf-folded, folded portion not

irregularly wrinkled; M-89,-147 Pontania sp. 15.

Antennae with segment 3 complete; both edges of leaf folded, folded portion irregu-
larly wrinkled; M-116,-144 Pontania sp. 16.

Ninth abdominal tergum with paired markings on caudal half; antennae with
segment 3 complete; single edge-folder, folded portion not irregularly wrinkled;
Y-8.46(?)-2-2 (in part) Pontania sp. 17.

Nematid genus 1. — Larvae small, greenish; body cylindrical, tapering
uniformly toward caudal end; segmentation distinct; annulation indistinct;
third abdominal segment apparently with four annulets, annulets 1, 2, and
3 setiferous; thoracic legs conspicuously long, nearly as long as thorax,
slender, with trochanter longer than femur; larvapods well developed,
setiferous; tenth abdominal segment without the paired caudal protuber-
ances; head circular, front flattened, smaller than thorax in width and

409] LARVAE OF THE TENTHREDINOIDEA—YUASA 91

height; antennae with four segments, segments small, incomplete, with
large sensory (?) pits, segments sometimes fused in part; spiracles not
winged.

This genus is represented by a single unidentified species collected by
Chester Young on Salix at Ithaca, New York. The larva is unique in the
character of the legs in that the trochanter is longer than the femur.
That this species belongs to the Nematinae is unquestionable but it is not
closely related to any genus in particular except perhaps to Pontania.
It may represent an undescribed genus.

Species 1. — Length, 10.5 mm.; body greenish; head brownish with dorsal
half of front dark fuscous; labrum semicircular with slight mesal emar-
gination; maxillary palpi with segment 1 nearly as long as segment 2
which is cylindrical and as wide at distal end as at proximal, segment 3
much smaller, segment 4 minute, peg-like, two distal segments curved
mesad; galea conical, only slightly larger than labial palpi; thoracic legs
with coxae subequal in length to tibiae, with trochanter slightly shorter
than coxa, and as long on dorsal margin as on ventral, femur shorter
than trochanter, cylindrical, three-fourths as wide as long, tarsal claws
slightly curved; larvapods with 1-2 setae near cephalic aspect; setae
slender, not stiff, rather sparse; tenth abdominal tergum rounded on
caudal margin, with few setae; subanal lobe with several setae; abdominal
segments with subspiracular areas with two setae and surpedal areas with a
single setae; on Salix nigra; C-c.y.-77.

Subfamily Blennocampinae

Larvae (Figs. 19-20) moderately large; body subcyHndrical, sometimes
rather robust, tapering uniformly caudad, venter more or less flattened,
usually distinctly spinose; segmentation distinct; annulation indistinct;
third abdominal segment with five or six annulets, rarely apparently with
four; thorax sometimes thickened; thoracic legs well developed, normal,
tibia shorter than or subequal to femur; femur produced ventro-distad as
pointed membranous projection; larvapods on segments 2-8 and 10, normal
in form, glabrous, subsegmented, distal lobe truncate on distal margin
and often curved mesad; tenth abdominal segment usually with several
spines arranged in a transverse row along caudal margin; suranal and sub-
anal lobes with several setae; head small, sparsely setiferous, narrower than
thorax, front slightly convex; antennae with five segments, slender, elon-
gately conical; ventral glands wanting; glandubae sometimes present;
spiracles rarely winged; spines often very long, furcate, with two, three, or
five branches, barbed, or represented by conical tubercles or sometimes
reduced to short bifurcate tubercles; cuticle microscopically and densely
spinulate; ultimate stage occurs, in which all setae and spines are lost and

92 ILLINOIS BIOLOGICAL MONOGRAPHS (410

body becomes colorless and glabrous; free leaf-feeders; sometimes gregari-
ous.

The Blennocarapinae as restricted by MacGillivray is a large sub-
family rich in genera and species, and is related to Fenusinae and Scolion-
eurinae. This is in agreement with Konow's statement altho this author
makes his tribe Blennocampides the third in his subfamily Tenthredinini.
Rohwer would group the majority of the genera under consideration in
his subfamily Empriinae, but take out the genera Phymatocera and Tomos-
tethus from the subfamily and place them in a subfamily by themselves.
This arrangement has an advantage in classifying the larvae because of
the fact that the larvae of Tomostethus lack the characteristic spines which
readily distinguish the Blennocampinae from all other groups in the larval
stages. The following key will separate the genera examined, with two addi-
tional ones, Erythraspides and Periclista, whose diagnostic characters
are taken from Dyar's paper (1898b).

GENERA OF BLENNOCAMPINAE

1(2) Body without spines; with six annulets, annulets 2 and 4 each with a transverse
row of minute but stalked glandubae Tomostethus Konow.

2(1) Body with spines 3.

3(10) Third abdominal segment with six distinct annulets; spines usually unbranched but
conical, if branched, very short and minute, tenth abdominal tergum with small
conical unbranched spines 4.

4(7) Body spines conical and not bifurcate, blackish 5.

5(6) Spiracles with distinct black wings Monophadnus'H.a.xtig.

6(5) Spiracles without distinct black wings Ilypergyricus MacGillivray

7(4) Body-spines not conical but bifurcate, blackish or whitish 8.

8(9) Spines whitish; tenth abdominal tergum not marked Blennocampa Hartig.

9(8) Spines black; tenth abdominal tergum marked with black. .Erythraspides Ashmead.
10(3) Third abdominal segment with five annulets, rarely apparently with four; spines in

part usually bifurcate, long, never short and conical 11.

11(12) Tenth abdominal tergum with a mesal spine cephalad of caudal marginal row of
spines; subdorsal spines of prothorax with five branches; prothoracic spinal formula:
5-2-1:5:1-2; third abdominal segment:2-2-2:0:3-2-2:2-l:l-2; ultimate tergum 1-1-1:
2 :2 M onophadnoides Ashmead.

12(11) Tenth abdominal tergum without a mesal spine cephalad of caudal marg^al row of
spines; subdorsal spines of prothorax with three branches at most 13.

13(14) Second annulet of third abdominal segment with three spines dorsad of spiracular
line; host-plants not confined to Quercus species; prothoracic spinal formula: 2-2-2:
2-3:1-2; third abdominal segment 2-2-2:1:2-2-2:2-1:1:1 Isodictium Ashmead.

14(13) Second annulet of third abdominal segment with two spines dorsad of spiracular
line; host-plants confined to species of Quercus; otherwise resembling the preceding
genus Periclista Konow.

Tomostethus Konow
Larvae moderately large, length 17-21 mm., rather robust, yellow-
ish white; body subcylindrical, tapering little caudad, venter flattened,

411) LARVAE OF THE TENTHREDINOIDEA—YUASA 93

without spines, sparsely and microscopically setiferous; head black, shiny,
much smaller than thorax; third abdominal segment with six annulets,
annulets 2 and 4 each with a transverse row of few stalked glandubae;
tenth abdominal segment without spines, truncate on caudal margin;
spiracles not winged, but with a pair of faint ventral crescentic brown
marks; antennae, (5, 4, 3), 2, 1 in older larvae and 5, (4, 3), 2, 1 in younger
larvae; maxillary palpi, 4, 2, 1, 3, pointed; labial palpi rather slender, (1, 2);
ultimate stage entirely whitish.

Tomosthethiis bardus Say. — Length, 18 mm.; width of head, 1.9 mm.;
body whitish with yellowish tinge, in older specimens yellowish white;
head shiny black with clypeus alone lighter in color; legs blackish brown;
larvae gregarious; on ash; G-2, Y-8.14.

Tomostethus tnulticinctus Rohwer. — According to Sasscer's description
(1911) the larvae of this species are indistinguishable from the preceding
species but the bred adults have been assigned to this species by Rohwer.
Larvae have not been examined.

Blennocampa Hartig

Larvae rather small, length 15-20 mm.; greenish; body subcylin-
drical, slender, tapering uniformly caudad; spines small, bifurcate, tubercle-
like or conical; head very small, third abdominal segment with six an-
nulets, annulets 2 and 4 spinose; tenth abdominal segment spinose, spines
conical, unbranched, numerous, arranged in four rows, 1: 1: 1-1-1:
1-1-1-1-1; typical prothoracic spinal formula: 2-2-1-1-1: 1-1-1: 1: 1-1;
third abdominal segment, 2-2-1: 1:2-2-1: 1-1-1: 1-1-1; antennae, 5, (1, 2, 3,
4); maxillaray palpi, (2, 3, 4), 1, slender, pointed; labial palpi (1, 2),
nearly equal to two distal segments of maxillary palpi taken together;
legs with femur longer than tibia; larvapods normal in form, more or
less rounded at distal end.

Blennocampa spiraeae Dyar. — Length, 16.5 mm.; width of head, 1.2
mm.; body greenish; head pale brown; legs concolorous with body; ocel-
larae entirely black, tips of mandibles and tarsal claws brown; maxillary
palpi with segment 4 and labial palpi with segment 2 deep brown; pro-
thoracic formula of spines variable, 2-2-1-1-1 or 2-3-1-1, or 2-1-1-1; ab-
dominal segment on surpedal lobe with from 2-1 to 2-2 spines; tenth
abdominal tergum with small, short, conical spines arranged in four rows
as follows: (1) two pair of spines on each side of meson, (2) two spines on
each side of the meson, near the center of the tergum, (3) lateral pairs
sometimes with additional spines, and (4) the last and caudal row of
five spines on each side of meson along the caudal margin of tergum, the
three lateral spines closer together than the others; on Spiraea; not bred:
M-28.

94 ILUNOIS BIOLOGICAL MONOGRAPHS [412

Erythraspides Ashmead

Larvae comparatively speaking small, inconspicuously spinose,
greenish; third abdominal segment with five annulets, annulets 2 and 4
each with three minute bifurcate spines.

According to Dyar's Key (1898) the larvae of Erythraspides pygtnaeae
is distinguished from those of Blennocampa spiraeae by the black head
and spines of the former. Record is meager, and without specimens no
adequate diagnosis can be given.

MONOPHADNUS HaRTIG

Larvae rather small, length less than 15 mm., spotted; body rather
robust, only slightly and uniformly tapering caudad; tubercles conical,
small, blackish, not furcate; third abdominal segments with six annulets,
annulets 2 and 4 tuberculate; tenth abdominal tergum with two rows of
tubercles, some of which are bifurcate; prothoracic spinal formula:
1-1:1:1:1; third abdominal segment, 1-1-1:1:1-1-1:1:1; antennae 5, (4, 3,
2), 1; maxillary palpi (4, 2), 1, 3; labial palpi (1, 2); palpi rather thick and
conical; spiracles with distinct black wings.

Monophadnus nubilipennis Norton. — Length, 14 mm.; width of head,
1.3 mm.; head blackish brown, clypeus alone lighter; body dirty white
with yellowish tinge; legs grayish; on hellebore; Y-42,-8.42.

Hypergyricus MacGillivray
Larvae rather large and robust, spotted; length 16-20 mm., body
subcylindrical, tapering but slightly and uniformly caudad; tubercu-
late, tubercles conical, short, stout, usually not furcate; third abdominal
segment with six annulets, annulets 2 and 4 tuberculate; tenth abdominal
segment with two rows of few tubercles; prothoracic spinal formula
variable, but with a single tubercle on supraspiraculararea and two tubercles
ventrad of it; third abdominal segment also with variable number of
tubercles, only two on subspiracular area; tubercles not furcate; spiracles
with faint ventral crescentic brown marks but without definite wings;
antennae rather obtusely rounded, segments decreasing in diameter from
proximal to distal, but increasing in length; suranal and subanal lobes
strongly convex, with numerous short recumbent setae; legs with femur
distinctly longer than tibia; distal portion of femur dilated and produced
ventro-mesad.

SPECIES OF HYPERGYRICUS
Head bJack, genae, antennariae, ventral half of front and clj^^eus lighter in color; tubercles
blackish; body whitish, faint grayish shade and yellowish tinge in older stages;
legs grayish brown; antennae with segment 3 and 4 subequal in length; prothorax
with spines, 2-2:1:1:1; third abdominal segment, 1-1:0:1-1:1:1; latus of body along
supraspiracular lines with broken band of grayish shade, marks distinct and square

413] LARVAE OF THE TENTHREDINOIDEA—YUASA 95

dorsad of each spiracle; on leaves and fruits of Smilacina racemosa; length, 18 mm.;

width of head, 1.8 mm.; Y-73,-29-10,-8.73, M-181 fumtpennis Norton.

Head brownish yellow; length, 17 mm.; width of head, 1.9 mm.; spots on dorsum variable;
body whitish with faint yellowish tinge; legs concolorous with body; supraspiracular
lines without smoky-black band; prothoracic spines 1-1-1 or 1-1-1-1:1:1:1 or 3; third
abdominal segment with 1-1 ; 1 :1-1-1 :1 :1 ; any of the dorsal tubercles may be wanting;
otherwise similar to the preceding species; Y; 20 specimens collected by Mr. J. R.
Malloch on Smilacina in Illinois Hypergyricus sp. 1.

MONOPHADNOIDES ASHMEAD

Larvae small, distinctly spinose, greenish; length less than 17 mm.;
body subcylindrical, tapering caudad, rather slender; spines furcate,
with two, three, or more branches; third abdominal segment with five,
apparently four, annulets, annulets 2 and 3 spinose; prothorax with spines,
5-2-l:(5 or 3):1:2; third abdominal segment, 2-2-2:0:(3 or 2)-2-2:2-l:l-2;
tenth abdominal tergum with a mesal furcate spine cephalad of caudal
marginal row of spines; legs rather slender, femur slightly longer than
tibia, not dilated at distal end; spiracles unwinged; maxillary palpi,
(1, 4), (2, 3); labial palpi, (1, 2); maxillary and labial palpi slender, pointed;
antennae, 5, (1, 2, 3, 4), sharply pointed and slender; semigregarious.

Monophadnoides ruhi Harris. — Length, 16 mm.; width of head, 1.6 mm.;
head pale brownish green, distinctly setiferous, spines whitish, branches
sometimes light brown; length of longer branches subequal to the width
of head as seen in profile; tenth abdominal tergum with mesal spines with
2-3 branches, on the caudal third of the tergum cephalad of the marginal
row of spines, sometimes another smaller unbranched spine cephalad of
the furcate mesal spines; marginal row of spines, beginning with mesal
spine, consists of two simple, one bifurcate and lower, and lateral simple
spine on each side of meson; on Rubus and also on giant Ragweed; Y-8.17,
M-19,-183, G-562, 500-3.

ISODYCTIUM ASHMEAD

Larvae rather small, usually distinctly spinose; length less than 17 mm.;
body subcylindrical, tapering caudad, rather slender; third abdominal
segment with five annulets, annulets 2 and 4 spinose, sometimes appar-
ently four-annulate, with annulets 2 and 3 spinose; spines furcate, usually
with two to three branches, branches usually conspicuously long, some-
times small, short, but always sharply pointed at distal end; spiracles never
winged; thoracic legs normal, femur subequal in length to or longer than
tibia; head blackish or spotted on vertex and front or uniformly greenish;
spines on prothorax, 2-2-2-1:2-2:1:2; on third abdominal segment 2-2-2:1:
2-2-1:1:1; tenth abdominal segment with a marginal row of spines, usually
four on each side of meson.

96 ILLINOIS BIOLOGICAL MONOGRAPHS [414

Several species have been examined. None of the material has been
identified altho the following larva may belong to /. suhgregarium.

Isodyctium sp. 1. — Length, 15.5 mm.; head, 1.5 mm. wide; head marked
on vertex and front with large confluent brownish spots, those on front
sometimes separate; body uniformly greenish; legs concolorous with
body, femur subequal in length to tibia; spines blackish, very long and
furcate, with large proximal end, those ventrad of spiracular lines whitish;
those one-half as long as head are wide as seen in profile; prothorax with
spines, 2-2-2-1:2-2:1:2; third abdominal segment, 2-2-2:1:2-2-2:2-1:1-1;
tenth abdominal tergum, 2-1:2:2, the mesal pair sometimes with confluent
bases; maxillary palpi, (4, 2), 3, 1 ; labial palpi, 1,2; ultimate stage: entirely
whitish, vertex pale brown, third abdominal segment with five distinct
annulets, setiferous but not spinose; on oak; M-6.

Subfamily Fenusinae

Larvae (Fig. 21) very small; body semicylindrical, venter flattened,
depressed, tapering caudad, glabrous; segmentation distinct; annulation
indistinct; third abdominal segment with either one or apparently 2-4
annulets; thorax slightly swollen, prothorax sometimes with dorsal and
ventral shields; legs small, short, apparently with four segments, spreading
cephalo-laterad; larvapods on abdominal segments 2-8, vestigial, merely
mound-like; anal larvapods obsolete; tenth abdominal tergum glabrous
without suranal processes or caudal protuberances, sometimes with small
mesal projections; suranal and subanal lobes glabrous; liead sparsely
setiferous, depressed, subtriangular in outline, wedge-shaped in profile
narrower than thorax and overlapped on caudal third by prothorax; ver-
tical furrows wanting; antennae apparently with single segment; antacoria
large; ocellarae minute, located dorso-caudad of antennariae; ventral
glands wanting; glandubae wanting; spiracles indistinctly winged; cuticle
sometimes with microscopic but distinct chitinized dentiform spines;
larvae, leaf-miners.

The Fenusinae is a small subfamily represented by four genera in the
Nearctic region. Systematists have always considered this group as
closely related to the Scolioneurinae and Blennocampinae. MacGillivray
is the only one who would assign them subfamily rank. Konow listed three
European species, dohrni, ulmi, and pumila, under the old generic name
Kaliosysphinga. The first two are now considered as types of distinct
genera. They have been introduced into this country and are liable to do
considerable damage at times. The genera studied are separable as follows:

GENERA OF FENUSINAE

Caudal end of body rounded, without a mesal suranal protuberance; sternum of ninth abdom-
inal segment with a pair of swellings covered with distinct microscopic dentiform spines:

4151 LARVAE OF THE TENTHREDINOIDEA—YUASA 97

tenth sternum strongly convex, much shorter than ninth sternum

Kaliofenusa MacGillivray.

Caudal end of body truncate, with a mesal suranal protuberance; sternum of ninth abdominal

segment flattened, uniformly and microscopically spinulate; tenth sternum slightly

convex, nearly as long as the ninth sternum Fenusa Leach.

Kaliofenusa MacGillivray

Larvae very small, greenish white; length less than 12 mm.; thorax
broadest on mesothorax; declivous on dorsum from metathorax toward
the head; head only slightly depressed, front convex; mouth-parts normal
except labium, which is flattened; mandibles rather thick in profile;
dorsum and venter covered with brownish, irregular, microscopic dentiform
spines, larger on the center of segment and larvapods; caudal end of body
rounded; tenth abdominal tergum without mesal protuberance; sternum
of ninth segment with a pair of microscopically dentate swellings; tenth
sternum strongly convex, much shorter than ninth sternum.

Kaliofenusa ulmi Sundevall. — Length, 10 mm.; width of head, .9 mm.;
head light brown, legs brown, ocularia, mandibles, maxillary palpi, deep
brown; sternum of ultimate segment with transverse depression on the
caudal third distinct; leaf -miners, on Ulmus, feeding on all tissues except
upper and lower epidermis; Y-4-1, -4-2,-8.4,

Fenusa Leach

Larvae very small, length less than 12 mm., greenish white; body rather
uniform in width except at caudal end which tapers suddenly and dis-
tinctly; head strongly depressed; front flattened; mouth-parts protruding
beyond the distal end of mandibles; body uniformly and microscopically
spinulate; no distinct localized microscopic dentiform spines; caudal
end of body truncate; tenth abdominal tergum with a distinct mesal coni-
cal suranal protuberance; sternum of ninth segment rather flattened,
without microscopically dentate swellings; tenth sternum slightly convex,
nearly as long as ninth.

Fenusa dohrni Tischbein. — Length, 10-11 mm., width of head, 1.1 mm.;
head brown; prothorax with dorsal and ventral shields indicated; thoracic
legs light brown, trochanter wanting, femur subequal in length to tibia;
tibia with proximal end subequal in diameter to distal end of femur;
tenth abdominal sternum with transverse depression in caudal fourth
indistinct; body often greenish, green adipose tissues being visible thru
the cuticle; dorsal vessel showing thru as light-colored line; leaf-miners on
Alnus vulgaris, feeding habit similar to that of Kaliofenusa ulmi; Y-4. A,
-43-1-2.

98 ILLINOIS BIOLOGICAL MONOGRAPHS [416

Subfamily Scolioneurinae

Larvae (Fig. 22) very small; body semicylindrical, somewhat de-
pressed, flattened on venter, broader on thorax, tapering caudad, glabrous,
greenish, never with bright patterns; segmentation distinct; annulation
indistinct, third abdominal segment with two annulets; larvapods rudi-
mentary, mere swellings on abdominal segments 2-7, the anal pair adjacent
on meson, forming a single protuberance; thorax thickened, prothorax
often with distinct dorsal and ventral chitinized shields; thoracic legs
small, slender, distinctly five-segmented, directed laterad; head depressed,
sub triangular, wedge-shaped in profile, narrower than thorax; mouth-
parts flattened and protruding, labium large, with submentum and mentum
strongly chitinized; antenna one-segmented; vertical furrows usually
wanting; tenth abdominal tergum abbreviated, glabrous; spiracles usually
winged; glandubae obsolete; ventral glands wanting; cuticle often with
minute dentiform tubercles; leaf-miners.

The Scolioneurinae is a small subfamily containing six genera, four of
which are peculiar to the Nearctic region. All six genera are represented
in the North American fauna. Prior to the recognition of the subfamily
by MacGillivray, the species belonging to it were referred to the genera
Fenusa and Blennocampa. Konow first segregated a species of Entodecta
and later more of Scolioneura from Blennocampa, placing them, together
with Fenusa and its allies, in other genera of his tribe Blennocampides.
Rohwer would separate the genera of the Scolioneurinae from those of
the Blennocampinae but unite them with those of the Fenusinae in the
tribe Messini of his subfamily Messinae. The close affinity of the Fenu-
sinae and ScoHoneurinae is evident from the fact that all known larvae
of these subfamilies are leaf-miners and that they possess similar types of
structural modifications. The definitions here given are based on obser-
vations on two American species of Metallus supplemented by writings of
European students — Cameron, Brischke, and Zaddach.

Metallus Forbes

Larvae small, length 10-13 mm., whitish or pale brownish; body de-
pressed, rather stout, mesothorax broadest; pleuron of each segment
tuberculate; cuticle with microscopic irregular chitinized dentiform
tubercles, those on center of dorsum and venter largest; head directed
ventro-cephalad, much narrower than thorax, attached to the ventral
part of prothorax; vertical furrows wanting; front twice as long as wide,
labrum subtriangu'ar, small; antennae mamma-like; ocellarae incon-
spicuous, about one-fifth the diameter of the antennaria; mouth-parts
small but distinct, slightly modified; mandibular dentes sharp; maxillary
palpi three-segmented, stipes elongate, galea digit-like, slightly curved

417] LARVAE OF THE TENTUREDINOIDEA—YUASA 99

mesad and subequal in size to palpi, lacinia thin, small, plate-like; labium
comparatively large, flattened; labial palpi inconspicuous, apparently
2-segmented; ligula globose and proturding; thoracic legs with coxa large,
stump-like, following segments suddenly smaller and slender, trochanter
ring-like, femur as long as wide, tibia longer than femur, cylindrical,
chitinized, tapering gradually to distal end, tarsal claws broadly curved,
sharp; mesothorax and metathorax with a dorsal membranous swelling
on each side of the meson; spiracles distinctly and semicircularly winged;
tenth abdominal tergum small, convex; anal setae wanting; subanal lobe
small, prominently convex.

SPECIES OF METALLUS

Head browiiish, not concolorous with body, epicranial suture very distinct; dorsal and ventral
shields distinct and brown; dorsal shield transverse, covering dorsum of prothorax,
proventral shield very large, occupying entire venter between prothoracic legs caudad of
and continuous with brown cervacoria, mesoventral and metaventral shields small,
transverse, triangular between legs; labium with submentum strongly chitinized, brown
with dark carina on meson and along caudal margin, mentum bro^vn, longer than wide;
larvapods with crescentric brownish band on cephalic aspect; length, 11-12 mm.; on
Rubus; Y ruhi Forbes.

Head pale or whitish, concolorous with bod)', at least not distinctly colored; dorsal and
ventral shields obsolete, only rarely faintly indicated; labium with submentum broad
and without dark median longitudinal carina, depression on meson rarely present,
never dark and distinct; larvapods without crescentric brownish band on the cephalic
aspect; length, 10 mm.; on Rubus; Y bethunei MacGillivray.

The larvae of Metallus rubi were described by Forbes, but the original
specimens are apparently lost. The description given here is based on
specimens in the Cornell Collection. Forty-three larvae of M. bethunei
were examined thru the courtesy of Mr. H. G. Crawford of Guelph,
Ontario, Canada.

Subfamily Hylotominae
Larvae (Fig. 23) moderately large; body semicylindrical, venter
flattened, distinctly wider than high, widest on abdominal segments 1-3,
tapering caudad, caudal segments only one half the width of widest seg-
ments; yellowish green, spotted or not; segmentation distinct; third
abdominal segment with three annulets, all setiferous, often tuberculate;
thoracic legs large, spreading laterad, apparently six-segmented inclusive
of claws; claws sharply curved, large, distinctly separated from fifth
segment by suture and with a large pulvillus-like swelling; larvapods
setiferous, on abdominal segments 2-6 and with rudimentary 7th pair, or on
2-7 and 10 with rudimentary 8th pair; antennae one-segmented, either
conical or button-like, if conical, larvapods on abdominal segments 2-6
and 10; spiracles distinctly winged; glandubae obsolete; tenth abdominal
tergum without suranal processes.

100 ILLINOIS BIOLOGICAL MONOGRAPHS (418

The Hylotominae in the adult stage is closely allied to the Schizocerinae.
but in the larval stages their affinities, if extant, are not so manifest.
The laravae are peculiar in possessing six-segmented 'thoracic legs and
varying numbers of larvapods. The shape and general appearance of the
larvae are so characteristic that they alone are sufficiently reliable for
identification in the field. Two genera are represented in the Nearctic
region, the genus Atomacera, which includes only a few species, has not
been studied.

Hylotoma Latreille

Head, conspicuously shiny black, brown or yellowish, or with a brown
median streak; body with minute blackish tuberculate setiferous spots
forming interrupted longitudinal rows along some or all of the subdor-
sal, latero-dorsal, supraspiracular, subspiracular, and pedal lines; annu-
lation usually 2, 3, 1; annulet 1 with a few minute setae, annulets 2 and
3 with a transverse row of a few stiff brown setae together with very
minute setae scattered around the larger setae; subspiracular tubercles
obsolete; pedal area prominent, produced laterad and oblique, extending
entire length of segment; maxillary palpi normal, four-segmented, segment
1 flattened, with distinct mesal projection, segments 2 and 3 cylindrical,
segment 4 minute, peg-like; labial palpi normal, three-segmented; larva-
pods located close together near the meson, proximal portion chitinized, seti-
ferous, distal portion small, membranous, non-setiferous, bluntly rounded,
those on seventh or eighth abdominal segment lacking membranous
distal portion.

The genus is divisible into two sections by the structure of antennae and
number of larvapods. Owing to the incompleteness of published records
of larvae of this genus, involving in some cases confusion in specific identi-
fication, it is not possible to determine many of the specimens collected.
The following key will separate the species represented in the collections
studied.

SPECIES OF HYLOTOMA

1(10) Antennae distinctly conical or peg-like, twice as long as wide; larvapods on abdominal
segments 2-6 and 10, seventh segment with rudimentary pair; head always uniformly
blackish, brownish, or yellowish; body always with numerous minute blackish spots
arranged longitudinally along subdorsal, dorso-lateral, supraspiracular and pedal
lines 2.

2(5) Head always black, thoracic legs with all segments blackish; tenth abdominal tergum
blackish 3.

3(4) Tenth abdominal segment on ventral half blackish; area between larvapods with
minute colored spots; area ventrad of pedal folds with minute spots; latus of each
segment with a few very minute secondary setiferous spots besides regular tubercu-
late spots; on Crataegus Y-194-3 194-2; on Prunus? Y-194-5 Hylotoma sp. 1.

4(3) Tenth abdominal segment on ventral half unmarked , whitish ; area between larvapods
without minute colored spots; area ventrad of pedal folds without minute spots;

419] LARVAE OF THE TENTH REDINOIDEA—YU ASA 101

latus of each segment without minute setiferous spots; on Alnus: Y-194-4

Hylotonia sp. 2.

5(2) Head not always blackish but usually yellowish or brown; thoracic legs not with all
segments blackish, segments distad of coxa usually brownish or brownish yellow;
tenth abdominal tergum not always blackish 6.

6(7) Head blackish; tenth abdominal tergum yellowish; subanal lobe yellowish; spots
between larvapods numerous and brownish; latus of each segment with many very
minute secondary setiferous spots; tuberculate spots of body yellowish with brown
border; on oak; Y-214-1-2 Hylotoma sp. 3.

7(6) Head yellowish or reddish brown; tenth abdominal tergum blackish; subanal lobe
whitish 8.

8(9) Area ventrad of pedal folds with many minute setiferous spots; area between larva-
pods sometimes with spots; spots on body not uniformly blackish or brownish but
blackish on cephalic and also usually on caudal portion of the body those on middle
portion brownish or yellowish with brown border sometimes spots aU pale yellowish

brown; head reddish or yellowish brown; on Crataegus; Y-222

scapularis Klug.

9(8) Area ventrad of pedal fold without spots; area between larvapods never with spots;
spots on body uniformly blackish or brownish; head yellowish or yellowish brown;

on elm; C-C.U. 668 Y-29-24 Hylotoma sp. 4.

10(1) Antennae button-like, usually wider than long; larvapods on abdominal segments
2-7 and 10, eighth segment with rudimentary pair; head blackish brownish or with a
distinct median streak from occiput to front; body sometimes not spotted except

on each side of the meson on cephalic segments 11.

11(14) Head blackish or brownish; body distinctly and regularly spotted; tenth abdominal
tergum and subanal lobe usually blackish; thoracic legs usually with coxae blackish

or brownish and other segments brownish or grayish 12.

12(13) Tenth abdominal tergum and subanal lobe whitish; all colored areas or spots of body
brownish; area between larvapods with few setiferous spots; area ventrad of pedal
fold with many setiferous spots; on Salix discolor; C- Young 61 . . . .Hylotoma sp. 5.
13(12) Tenth abdominal tergum and subanal lobe blackish or brownish; all colored areas
or spots blackish or brownish; area between larvapods usually without setiferous
spots; area ventrad of pedal fold with varying number of spots; on Prunus; Y-1946-1

G-Onekama No. 23; Maine 1915; C-C.U. 656 C.U. sub. 64 Hylotoma sp. 6.

14(11) Head yellowish or light brownish with brown streak along epicranial stem; body not
regularly and distinctly spotted except along dorso-meson on cephalic segments;
tenth abdominal tergum never blackish or brownish; thoracic legs always con-
colorous with body; Y-185-1-2 (Azalea) Y-185 (Willow) M-113 (birch bred) M-108
(hazel) macleayi Leach.

Subfamily Schizocerinae

Larvae (Fig. 24) small; body subcylindrical, flattened on venter,
tapering caudad, mesothoracic and metathoracic segments somewhat
swollen, prothorax distinctly tapering cephalad; whitish or creamy white,
never spotted or striped; segmentation and annulation indistinct; third
abdominal segment with three annulets, all annulets provided with a
transverse row of tubercles; spiracles on annulet 1; tenth abdominal
tergum without suranal or caudal protuberances; larvapods on abdominal
segments 2-8 and 10, the venter of ninth segment with a pair of minute

102 ILLINOIS BIOLOGICAL MONOGRAPHS [420

protuberances; thoracic legs modified, prothoracic legs apparently four-
segmented, distal segment pad-like, tarsal claws wanting; mesothoracic
and metathoracic legs apparently three-segmented, distal segment pad-
like with a sharp claw on cephalic side; antennae apparently one-segmented,
segment large with several clear spots; antennaria ventrad or slightly
caudad of ocularium; spiracles winged; glandubae obsolete; in younger
specimens larvapods sometimes very indistinct; tubercles often indistinct;
leaf-miners.

The Schizocerinae is represented in the Nearctic region by a single
genus, Schizocerus, and includes a limited number of species. Modern
systematists have always associated this subfamily with the Hylotominae,
the two being separated by the presence or absence of the free part of
Sc2 in the front wing. The larvae of this subfamily are unique in having
thoracic legs and maxillary and labial palpi modified by reduction in the
number of segments.

ScmzocERUS Lepeletier

Larvae small, length less than 15 mm., creamy- whitish; head small,
pale brown, higher than wide, sparsely and microscopically setiferous;
annulation 1, 2, 3; annulets above spiracular lines with tubercles, 2 on
annulet 1, four on annulet 2, and three on annulet 3, annulet 1 with a row
of tubercles on venter; a row of tubercles between larvapods and surpedal
area; subspiracular area not tuberculate or warty; tenth abdominal
tergum small, only slightly convex, tubercules almost obsolete, with
several stiff short setae on caudal margin; venter of ultimate segment
with distinct anal larvapods, subanal lobe with a pair of long and short
tubercles on each latero-caudal margin; maxillary palpi apparently three-
segmented, distal segment very minute, formula, 2, 1, 3; labial palpi
apparently two-segmented; mandibles as viewed from side narrow and
slender; totaglossa of labium large; spiracles winged, wings small, oblong;
spiracular line dividing the latus into two subequal dorso- ventral parts.

Schizocerus zabriskiei Ashmead. — Head pale brownish-green, body
whitish; in younger specimens head brownish, body with minute tubercles
touched with brown; legs, and venter between legs brownish; tubercles
ventrad of surpedal area and dorsad of larvapods usually three in number
in mature larvae and two in younger larvae; mature specimen, length,
13 mm.; width of head, 1.3 mm.; on Portulaca; Y, G.

Schizocerus sp. 1. — Larvae indistinguishable from the preceding species.
This species was collected at Muncie, 111., by Dr. Edna Mosher, who
bred adults. It is considered by Dr. MacGiUivray to be a new species.

421] LARVAE OF THE TENTHREDINOIDEA—YUASA 103

Subfamily Acordulecerinae

Body (Fig. 25) subcylindrical, tapering caudad, venter flattened,
thorax distinctly swollen; segmentation distinct; annulation indistinct,
third abdominal segment with apparently three annulets, all annulets
tuberculate, setiferous; thoracic legs 5-segmented, spreading laterad, distal
segment consisting of a sharp recurved claw and caudal membranous
globose swelling; prothoracic legs one-half as large as metathoracic;
larvapods rudimentary on abdominal segments 2-7 and 10, increasing in
size from cephalic pair to sixth pair; tenth abdominal segment truncate,
small, without suranal processes; antennae flattened, apparently 1-seg-
mented; abdominal segments 2-4 or 2-5 and 8 with crescentic sucker-like
protuberances, one on each postsubspiracular protuberance; ventral
glands wanting; spiracles not winged; glandubae wanting.

The Acordulecerinae, according to MacGillivray (1906), is represented
on the North American continent by a single genus, Acordulecera. Rohwer
would divide the subfamily into two tribes, Acordulecerini and Conocoxini,
the former including besides Acordulecera, Pantherix and possibly
Thulea and the latter Conocoxa and Nithulea. A radically different
arrangement is that of Konow who regarded Acordulecera as one of
fourteen genera included in his tribe Lobocerotides which was one of four
tribes constituting his subfamily Lophyrini. Konow seems to have been
unfamiliar with the larvae of the Nearctic genus Acordulecera, for he
speaks of the larvae of Lophyrini as "mit 16 Abdominalbeinen; an Coni-
feren." — a characterization not at all applicable to the genus under
consideration. The host-plant of two species of Acordulecera have been
recorded, and one species recognized in the larval stage. I have examined
larvae of several unbred species.

Acordulecera Say
Larvae very small, length less than 12 mm., greenish, never spotted or
striped; head usually brownish or pale; annulation, 3, (2, 1); prothorax
constricted; annulets with transverse row of slight tubercles, each bearing
slender peg-like setae; lateral lobes distinct; tenth abdominal tergum with
several setae; suranal and subanal lobes with several setae; maxillary palpi
rather slender, uniformly tapering distad; mandible thick; labial palpi
small; ligula dilated, rounded; free leaf-feeders; gregarious.

SPECIES OF ACORDULECERA

1(4) Sucker-like protuberances on abdominal segments 2-5 and 8 2.

2(3) Head with vertex blackish brown, front distinctly lighter in color; legs whitish
concolorous with body; sucker-like protuberances with three marginal setae; length
9.5 mm.; width of head 1.2 mm.; on butternut; Y-8.93(?)-2-2. . .Acordulecera sp. 1.

3(2) Head with vertex light brown, epicranial stem deep brown, front sometimes deep
brown; in young specimens head entirely dark brown; legs brown; sucker-like

104 ILLINOIS BIOLOGICAL MONOGRAPHS [422

protuberances with three marginal setae; length 11 mm.; width of head 1.3 mm.;
on chestnut and butternut; Y-93 C-cu 667 Acordulecera sp. 2.

4(1) Sucker-like protuberances on abdominal segments 2-4 and 8 5.

5(8) Sucker-like protuberances with three marginal setae; on oak 6.

6(7) Head light brown with epicranial stem deep brown or brownish with vertical furrows
lighter in color; legs brownish; length 9 mm.; width of head 1.2 mm.; Y-131 -137,
C-cu 680, M-37 Acordulecera sp. 3.

7(6) Head yellowish or light brown, with epicranial stem concolorous with body; length,
7 mm.; width of head, 1.1 mm.; M-239, M-243, C-yll dorsalis Say.

8(5) Sucker-like protuberances with five marginal setae; head deep brown, vertical
furrows and epicranial arms lighter in color; legs whitish, concolorous with body;
length, 9 mm. ; width of head, 1.2 mm.; on hickory; Y- 144-5 w«5/a MacGillivray.

Family Pamphiliidae

Larvae (Fig. 1) of medium size; body subcylindrical, slightly flattened
on the ventral aspect; sublateral lobe on the ventro-lateral margin dis-
tinct, moderately large; body slender to robust; segmentation and annula-
tion distinct; cuticle microscopically setiferous, appearing smooth, often
delicate, transparent; color usually greenish or creamy white; head semi-
globose, prominent, as wide as thorax; color creamy or brownish or blackish;
mouth directed ventrad; head completely exposed, sparsely setiferous;
epicranial suture and vertical furrows present; antennae extremely long,
setaceous, conspicuous, seven-segmented; ocularia located ventro-laterad
of antennariae; mouth-parts normal, sericos produced and prominent;
prothorax with shield-like, usually brownish, broad patches on the dorsal
and lateral aspects; thoracic legs modified, setiform, sharply pointed,
segments cylindrical, distal segment very long, slender, straight; third
abdominal segment with four annulets on dorsal and ventral aspects;
spiracles on the second annulet; larvapods wanting; eighth abdominal
segment on the venter mesad of the lateral lobe with a fleshy protuber-
ance resembling a larvapod, ninth segment cylindrical, smaller than the
preceding; tenth segment depressed, rounded on the caudal margin,
usually setiferous, sometimes conspicuously so, often with colored patches,
always with a median hook-like suranal process near the caudal margin
of the tergum; subanal lobe with a pair of setiform, three-segmented
conspicuous subanal appendages; insects one- or two-brooded; spinning
silken web or rolling leaves for their nests; solitary or gregarious; pupate in
the ground.

The Pamphiliidae is an easily circumscribed family of eight or nine
genera and a large number of species which are peculiar to the Northern
hemisphere. The adults differ from all other Hymennoptera except the
Xyelidae in the preservation of the subcostal vein in the hind wings.
In this character it is more generalized than the Xyelidae altho it is
more specialized than this family in other features of venation. Brischke

423] LARVAE OF THE TENTHREDINOIDEA—YUASA 105

and Zaddach (1865) discussed the immature stages and biology of eleven
European species of the Pamphiliidae and pointed out that certain of
their habits may be of taxonomic significance. The larvae of these species
fall into one or the other of two groups according to the type of nests they
build. The first group contains those whose larvae build nests by tying the
leaves of their food-plants together with threads of silk and are either
sohtary, as Lyda, or gregarious, as Cephaleia and Neurotoma. The
second group consists of those whose larvae build nests by rolling the
edge of the leaves of their food-plants and live inside the tubes so formed,
as Pamphilius. Some of this latter group make portable nests out of
detached pieces of leaves, as Pamphilius inanitus on Rosa. The adults,
however, are so closely related to each other that Rohwer (1911) con-
sidered such a subdivision impractical. Konow (1901), in his analytical
table of the larvae, included sLxteen species, representing four genera, but
did not register any Nearctic species. According to this writer (1905) the
larvae of Lyda and Cephaleia feed on coniferous plants while those of
Neurotoma and Pamphilius attack deciduous plants. Nothing is known
concerning the biology of Anoplolyda. Dyar (1895) included in his table
ten Nearctic pamphilids but, excepting Pamphilius ocreatus, all were uni-
dentified and many were taken from descriptions given by Packard (1890).
Of about fifty-five Nearctic species representing seven genera, Acan-
tholyda, Itycorsia, Cephaleia, Caenolyda, Neurotoma, Pamphilius, and
Anoplolyda, only five species have been identified in the larval stage,
and the food-plants of about six species recorded.

Four identified and several unidentified species have been examined.
It is not possible to define the genera with this limited material; the
species studied can be separated as follows.

SPECIES OF PAMPHILIIDAE

1(14) Subanal appendages with the second segment longer than or subequal to the third
segment, never distinctly shorter; head usually dark-colored; tenth abdominal tergum
with colored patches 2.

2(3) Subanal appendages with the second segment subequal in length to the third, all
segments black; first segment longer than the two distal segments taken together;
head black; bodj' olive-green with yellowish lateral lobes; on spruce; Packard
fl890)-35 Pamphiliid sp. 1.

3(2) Subanal appendages with the second segment longer than third, the third usually
darker in color than the others; first segment longer than or nearly equal to the two
distal segments taken together; head dark brown to green 4.

4(5) Head green; tenth abdominal tergum without colored patches; body green; subanal
appendages with the second and third segments black; on Pinus strobus; Packard-83.

Pamphiliid sp. 2.

5(4) Head not green, usually brownish; tenth abdominal tergum with or without colored
patches; body not greenish, sometimes reddish or olivaceous; subanal appendages
usually with segments brownish; on Pinus and Abies 6.

106 ILLINOIS BIOLOGICAL MONOGRAPHS [424

6(13) Head brown to yellowish 7.

7(12) Head dark brown 8.

8(9) Head paler along epicranial arms; subanal appendages with first segment as long as
ail the other segments taken together; all segments pale to light brown; length,25mm. ;
width of head, 3 mm.; body robust, large; following parts dark brown: head except
along epicranial arms, prothoracic shields, thoracic surpedal lobe, coxae, sterna
between thoracic legs, cervical sclerites, markings on the tenth abdominal segment,
median suranal process, and antenae; ocellarae black; suranal process short and
erect; tenth abdominal tergum with a pair of lateral patches, not connected along
caudal margin; tenth abdominal sternum with a median brown patch, distinctly
rounded on cephalic margin; subanal appendages light brown, first segment equal
in length to other two taken together, segment 2 longer than 3; thoracic legs with
fifth segment as long as fourth, and third taken together; antennal formula: (2, 5),
3, 6, 1, (4, 7); cuticle distinctly reticulate, each area brownish; three specimens in
MacGillivray collection bearing label "Cephalaeia sp. No. 839? Maine." (The
identification is open to question, but, since these larvae are readily distinguished
from the larvae of Pamphilius and Neurotoma by the characters used in the table
here, they may be considered, tentatively at least, as representing the genus Cephal-

eia.) Cephaleia sp. 1 .

9(8) Head not paler along epicranial arms, but uniformly brownish 10.

10(11) Head pitch-brown; body pinkish to reddish brown; on spruce; Packard-36

Pamphiliid sp. 3.

11(10) Head horny brown; body horny brown; subanal appendages with first segment
longer than other two taken together, third segment brown; on Pinus strobus;
Packard-84 Pamphiliid sp. 4.

12(7) Head yellowish to brownish yellow; body pale brick-red or yellowish, each segment
with a large reddish spot on the spiracular line; subanal appendages with first segment
nearly equal in length to other two taken together; all segments blackish ; on Pinus
strohus; Packard-85 Pamphiliid sp. 5.

13(6) Head pale red, with a black spot on the front; body reddish olive-green with purplish
meso-dorsal line; on Austrian pine; closely related to Lyda campestris; Packard-82.

Pamphiliid sp. 6.

14(1) Subanal appendages with the second segment distinctly shorter than the third; head
variously colored, blackish to pale; tenth abdominal tergum either with or without
colored patches 15.

15(18) Tenth abdominal tergum with a colored patch on each side, median triangular patch
between the cephalic ends of the lateral patches wanting; lateral patches connected
along the caudal margin of tergum; sternum with a large colord patch 16.

16(17) Head blackish or dark brownish, in younger specimens pale brown; subanal append-
ages with first segment longer than other two segments taken together, third longer
than second, all segments brown, the third darker; cervical sclerites pale brown,
dorsal and lateral sliields of prothorax pale to light brown, patches on the ultimate
segment brownish; body with a pink dorsal line on the meson; antennae in mature
specimens with formula, (2, 3, 5), 1, 7, (4, 6) ; ocularium about one-third the diameter
of antennaria; ocellarae small, eccentric, near the mesal margin of ocularium;
mouth-parts brownish; length, 19 mm.; width of head, 1.9 mm.; on wild cherry;
G Neurotoma fasciata Norton.

17(16) Head lighter in color, yellowish, sometimes brownish yellow; subanal appendages with
first segment nearly equal to other two taken together, all segments black; following
parts blackish: prothoracic shields, thoracic legs, markings on ultimate segment,
subanal appendages, ocularia, thoracic sterna; the black markings of tenth abdominal

425] LARVAE OF THE TENTHREDINOIDEA—YUASA 107

tergum connected along the caudal margin; black marking of tenth sternum large,
completely covering the sternum, and with its cephalic margin between the subanal
appendages straight; antennae with formula, (3, 5), 2, 1, 7, (6, 4); length, 18 mm.;
width of head, 1.7 mm.; on plum; G NeuroiomainconspicuaNoTton.

18(15) Tenth abdominal tergum either with a colored patch on each side and with a median
triangular patch between the cephalic ends of the lateral patches alwaj^s present, the
lateral patches sometimes not connected along the caudal margin of the tergum,
the sternum with a large colored patch; or the tergum without colored patches and
the sternum either with a pair of small round spots ot without markings 19.

19(22) Tenth abdominal tergum with two lateral triangular patches and one mesal patch
and sternum with a large colored patch 20.

20(2 1 ) Head blackish ; body greenish-white; subanal appendages with the first segment longer
than the other two taken together, all segments pale brown, the third sometimes
darker; following parts blackish: head uniformly, antennae, prothoracic shields,
thoracic lateral lobe, markings on ultimate segment, suranal hook, cervical sclerites,
and coxae; antennae with formula, (3, 2), 5, 1, 7, (4, 6); eighth uromere with a pair
of marginal ventral glands near caudal portion of the sublateral lobe, both pale
brown; in life, body glossy white with diffused fleshy-reddish tint; antennae and
subanal appendages whitish; length, 19 mm.; width of head, 1.9 mm.; on Comus;
nests made by rolling edges of leaves cut across from the margin to the midrib;
gregarious; Y-125 Pamphilius sp. 1.

21(20) Head light brown, with a black spot on the apex of front; subanal appendages with
the first segment nearly equal in length to the other two taken together, two proximal
segments creamy, third deep brown; body whitish green; ocularia brown, a black spot
at the origin of the epicranial arms; following parts brown: thoracic shields, cervical
sclerites, and markings on the ultimate segment; tenth abdominal tergum with a
median cephalic triangular patch between the lateral patches; subanal appendages
with second segment longer than half the length of the third; thoracic legs pale or
creamy; antennae, beyond the first segment, deep brown, vnth formula: (2, 3, 5),

1, 7, (4, 6); length, 19 mm.; width of head, 1.8 mm.; on blackberry; G

Pamphilius dentatus MacG.

22(19) Tenth abdominal tergum without colored patches 23.

23(26) Tenth abdominal sternum without a pair of small colored patches 24.

24(25) Head unifomJy pale brown; subanal appendages wdth the first segment as long as
other two taken together, the second shorter than the third, all segments pale brown;
ocularia large, about one-third the diameter of antenaria, black; following parts
blackish brown: narrow prothoracic shields, cervical sclerites, and antennae; suranal
hook brown without colored base; antennae with formula, (2, 3), 5, 1, 7, (4, 6);
length, 12 mm.; width of head, 1.5 mm.; host unknown; Y-163-3. (This species
closely resembles P. persicus MacG.) Pamphilius sp. 2.

25(24) Head not uniformly pale brown, vertex brown, front with a brown spot; subanal
appendages with the first segment as long as the other two taken together, distal
two segments brownish; following parts brown: prothoracic shields, cervical sclerites,
antennae, and suranal process; legs pale brown, darker in young specimens; length,
17 mm.; width of head, 1.8 mm.; on blackberry; M-69 Pamphilius sp. 3.

26(20) Tenth abdominal sternum with a pair of small colored patches; head pale to light
brown, front sometimes with a brown spot, usually without it; subanal appendages
with the first segment nearly equal in length to other two taken together, the second
segment shorter than the third, all segments pale to brownish; ocularia and mandible
at distal end blackish brown; suranal process brown with its base pale; antennae

108 ILLINOIS BIOLOGICAL MONOGRAPHS [426

brown; tenth abdominal sternum with small brown round spot on each side; length,
16 mm.; width of head, 1.6 mm.; larvae solitary, leaf-edge rollers on Comus; M.

Pamphilius ocreatus Say.

Family Cephidae

Body (Fig. 2) cylindrical, sometimes slightly depressed, enlarged at
thorax, slightly and uniformly tapering caudad, slender or moderately
stout; segmentation usually distinct; annulation sometimes indistinct;
cuticle smooth or verrucose, microscopically and very sparsely setiferous;
color generallly pale or creamy white, never with distinct bright marks;
head circular in contour, semiglobose, moderately large, narrower than
thorax, caudal portion concealed by prothorax, pale brown or concojorous
with body, sparsely setiferous; mouth-parts directed ventrad, normal
in form, brownish; antennae with four or five segments, conical; ocellarae
small, with ocularia less than one-fifth the diameter of antennaria and
located latero-caudad of it; epicranial suture and vertical furrows present;
mesothorax distinctly, and metathorax with dorsal and lateral aspects
somewhat swollen; thoracic legs vestigial, fleshy, mamma-like, tarsal
claws wanting; third abdominal segment with two or three annulets,
sometimes indistinct; venter with three annulets; larvapods wanting,
sometimes with slight swellings in normal position of larvapods; lateral
lobes prominent, extending the entire length of the segment; tergum of
ultimate segment with mesal longitudinal broad depression and distinct
suranal process; sternum of ultimate segment with a pair of inconspicuous
vestigial, papilliform subanal appendages ventrad of the cephalic end of
the anal slit; internal feeders, boring into the stem of monocotyledonous
and herbaceous plants and bushes; pupation in tunnels in the host-plants.

The Cephidae contains about fourteen genera and is moderately rich
in number of species, some of them of intercontinental distribution.
Nine genera are represented in North America. Practically all systematists
have considered this group as a distinct aggregate worthy of family rank.
Rohwer (1911), however, has expressed the opinion that future studies
may possibly make it advisable to unite this group with the Xyelidae
and to treat each of them as subfamilies. There have so far been no facts
or reasons brought to light which call for such a step. On the other hand,
MacGillivray's study has emphasized the fact that "so far as the wings
are concerned, they (Cephidae) are the most distinct of any group of the
Tenthredinoidea, and are only indirectly related to any of the other
families." They are generalized in the manner of the origin of media
but are specialized in other features of the wings. On the basis of larval
characters this family is related to the Pamphiliidae and is quite unrelated
to the Xyelidae.

The systematic position of the osculant genus Syntexis is unsettled.
In the original description Rohwer (1915) stated that this genus has

427) LARVAE OF THE TENTHREDINOIDEA—YUASA 109

affinities with both the Cephidae and Xiphydriidae and on the basis of
venation, it probably belongs to the latter as defined by MacGillivray.
Certain features other than venation, however, led Rohwer to place it in
the Cephidae. It is not safe to venture any opinion without careful
examination of the larvae, but their antennae and their lack of papilliform
subanal appendages suggest close affinity with the Xiphydriidae.

At least six out of sixteen American species have been recognized
in the larval stages and their host-plants recorded. Janus integer and
Cephus cinctus are economic pests. Konow's tribe Macrocephides includes
the species whose larvae bore into the pith of shrubs and woody twigs,
and his tribe Cephides embraces those whose larvae bore into the stalks
of Graminaceae.

Middleton (1917) published descriptions and keys for distinguishing the
larvae of five species representing four genera, Adirus, Janus, Cephus, and
Hartigia together with a definition of the groups based on "characters
common to all the genera studied and probably to the family." He
reserved for future discussion the question of the systematic position of
the Cephidae, but pointed out the obvious affinity of this group with the
Siricidae on one hand and the Pamphiliidae on the other. Gahan (1920)
described the larva of Tracheitis tabidus (Fab.) and added a key for sepa-
rating this larva from that of Cephus cinctus and C. pygmaeus.

M^

GENERA OF CEPHIDAE

1(10) Papilliform subanal appendages present on the ultimate abdominal segment 2.

2(3) Suranal process depressed on the distal portion, oval in cross-section, with strongly
chitinized dentiform tubercles on the proximal portion; antennae five-segmented.

Janus Stephens.

3(2) Suranal process not depressed on the distal portion, circular in cross-section, with or
without strongly chitinized dentiform tubercles on the proximal portion; antennae
four-segmented 4.

4(5) Suranal process proximad of distal cj^lindrical portion with strongly chitinized
dentiform tubercles Adirus Konow.

5(4) Suranal process proximad of distal cylindrical portion without strongly chitinized
dentiform tubercles 6.

6(7) Terga of eighth and ninth abdominal segments setiferous, each with a transverse row
of distinct setae; tenth abdominal tergum as viewed from side strongly convex and
truncate on the caudal aspect; suranal process with stiff setae not arising from distinct
chitinized bases Trachelus Jurine.

7(6) Terga of eighth and ninth abdominal segments glabrous, each without a transverse
row of distinct setae; tenth abdominal tergum as viewed from side not strongly
convex and not truncate on the caudal aspect but gradually declivous caudad;
suranal process with stiff setae arising from distinct chitinized bases 8.

8(9) Suranal process with distal chitinized portion very short and ring-like, or very
long and cj^inder-like, more than twice as long as wide; suranal process with a
single transverse row of setae proximad of the distal chitinized portion or with two or
more whorls of setae; the distal margin of the distal chitinized portion not distinctly
serrate Cephus Latreille.

no ILLINOIS BIOLOGICAL MONOGRAPHS [428

9(8) Suranal process with distal chitinized portion, as long as wide, cylinder-like; suranal
process proximad of the distal chitinized portion with two or more whorls of setae;

distal margin of the distal chitinized portion minutely but distinctly serrate

Hartigia Schiodte.
10(1) Papilliform subanal appendages on the ultimate abdominal segment wanting.

Syntexis Rohwer.

Should Cephus and Hartigia (Middleton, 1917) and Trachelus (Gahan,
1920) prove to possess five-segmented antennae, as reported, contrary to
the observations of the present writer, the genus Janus can be separated
from them by the chitinized dentiform tubercles and Trachelus by the
glabrous eighth and ninth abdominal terga.

Janus Stephens
Antennae distinctly with five segments; segments 1 and 2 short,
ring-like, segment 3 longer, but half as wide as segment 1, segment 4
cylindrical, longer than wide, segment 5 elongate-conical, twice as long as
wide, subequal in length to or shorter than segment 4; suranal process
short, strongly chitinized, nearly as long as wide at proximal end, denti-
form tubercles with circular rows of stiff setae on the proximal half, distal
half small, depressed, narrowly oblong in cross-section; subanal appendages
peg-like, with a minute seta at the distal end, apparently segmented,
without setae near the proximal end; lateral area of suranal lobe with
2-3 setae; mandible with four dentes.

SPECIES OF JANUS

Subanal appendages distinctly two-segmented, segment 1 ring-like, much larger in diameter
than segment 2; antennae with segment 5 about one-half as long as segment 4; bores in
Salix and Populus abbreviatus Say.

Subanal appendages indistinctly two-segmented, segments subequal in diameter; antennae
with segment 5 subequal in length to or only slightly shorter than segment 4; bores in
Ribes species (currant); C integer Norton.

Adirus Konow

Antennae distinctly with four segments; segment 4 long, peg-like,
bluntly pointed, nearly three times as long as wide, twice as long as
segment 3; suranal process large, distal fourth strongly chitinized, sud-
denly constricted, circular in cross-section, with chitinized dentiform
tubercles; subanal appendages peg-like, fleshy, apparently unsegmented,
with setae near the proximal end, these setae separated from the remainder
of the setae on the sternum; lateral area of suranal lobe with 20-28 setae.

Adirus trimaculatus Say. — Middleton has described the larvae in
detail. They bore in the stems of blackberry and rose. Thru the kindness
of Dr. E. P. Felt, one mature and two young damaged specimens from

429] LARVAE OF THE TENTHREDINOIDEA—YUASA 111

the New York State Museum, labeled "a 2766" and "a 2261" were exam-
ined by me.

Trachelus Jurine

Antennae apparently with four segments — according to Gahan five
segments present; body thickest dorso-ventrad in meso thorax, second
and third abdominal segments widest, the ninth and tenth tapering
suddenly; suranal process rather short, tapering caudad gradually, chit-
inized dentiform tubercles wanting, with or without stiff short setae,
if setae present they do not arise from chitinized bases; ultimate tergum
setiferous on both sides of the median depression, convex as viewed from
side, truncate on the caudal aspect, not declivous caudad; third abdominal
segment with three annulets, annulet 2 largest and with a transverse row
of setae; subanal appendages peg-like, unsegmented.

Trachelus tabidus Fabricius. — Suranal process without distinct con-
striction, with or without brownish setae, if present, setae arranged in a
irregular circle in the proximal half; subanal appendages brownish with 2
minute setae at the distal end, and with 1 or 2 setae which are separated
from the remainder of the setae on the sternum; lateral area of suranal
lobe with 5-7 setae; antennae with segment 4 bluntly rounded, segments
2 and 3 ring-like, segment 1 narrow but large in diameter with a seta on
the dorsal margin, segments all brown; bore in the stalks of wheat, rye,
and barley (Gahan 1920).

Cephus Latreille

Antennae apparently with four segments — according to Middle-
ton five segments present — segment 4 less than twice as long as wide;
abdominal segments 5-8 with ventral swellings corresponding in position
to larvapods; suranal process without chitinized dentiform tubercles, either
with a narrow ring-like distal chitinized portion and a semicircular row of
setae or with a long cylindrical distal portion and two or more irregular
rows of whorls of setae on the proximal portion, the distal margin of the
distal chitinous portion entire and smooth; subanal appendages papilla-
like, cylindrical, more than twice as long as wide, bluntly rounded at the
distal end, with one or more setae near the proximal end; lateral area of
suranal lobe with 7-15 setae; mandible with three dentes.

According to Rohwer (1917) only two species of this genus are known
to occur in North America. They can be separated as follows:

SPECIES OF CEPHUS

Suranal process with the distal chitinized portion cylindrical, twice as long as wide, setae on
the proximal portion arranged in two or three irregular whorls; subanal appendages with
several setae; lateral area of suranal lobe with about 15 setae; the only true Nearctic
species known; bores in stalks of Elymus, Agropyron, Phleum, and wheat, cinctus Norton.

112 ILLINOIS BIOLOGICAL MONOGRAPHS (430

Suranal process with distal chitinized portion ring-like, shorter than wide, setae on the
proximal portion arranged in a semicircular row on the dorsal aspect; subanal append-
ages with one or rarely two setae, lateral area of suranal lobe typically with seven setae;
introduced from Europe; bores in stalks of wheat pygmaeus Linnaeus.

Hartigia Schiodte

Antennae apparently with four segments — according to Middleton
(1917) with five segments — segment 4 longer than segment 3, elongate,
conical; suranal process twice as long as wide at proximal end, without
strongly chitinized dentiform tubercles, with several whorls of spinous
setae; subanal appendages two-segmented, sometimes segmentation
indistinct, with accompanying setae separated from the remainder of
setae of the sternum; lateral area of suranal lobe with 15-20 setae.

Hartigia creisoni Kirby. — The larvae of this species have been described
in detail by Middleton. They bore in the stems of Rubus in California.
The preceding generic definition was based upon specimens obtained thru
the courtesy of Mr. Harry S. Smith, of Sacramento, California, and does
not quite agree with that given by Middleton (1917).

Family Xiphydriidae

Larvae (Fig. 3) small; body subcylindrical, thorax and two caudal seg-
ments distinctly swollen; segmentation distinct; annulation obsolete;
creamy white, no markings; glabrous; thoracic legs rudimentary, fleshy,
mamma-like, without tarsal claws; larvapods wanting; ultimate segment
with distinct suranal process, without subanal appendages; ocellarae
wanting; mouth-parts modified; maxillary palpi apparently two-segmented;
antennae apparently with three segments; metaspiracles functionless, very
much smaller than abdominal spiracles; cuticle on dorsum smooth, on
venter microscopically, sharply, and densely spinulate; tenth abdominal
tergum with deep meso-dorsal depression; wood-borers.

The Xiphydriidae contains four genera, Derecyrta, Brachyxiphus,
Xiphydria, and Konowia, which may be divided into two groups on
the presence or absence of the radial cross-vein in the wings. Systematists
have generally considered this group as a subfamily of the Siricidae, but
MacGillivray (1906) has elevated it to its present standing on the vena-
tional characters, which he has proven to be the most generalized of the
specialized Tenthredinoidea. Rohwer (1911) would divide the family
into two subfamilies, Xiphydriinae and Derecyrtinae, the latter being
monobasic. Of the four genera, two are represented in the Nearctic fauna.

In a recent synopsis of the Nearctic wood-wasps, Rohwer (1918b)
tabulates eight species of Xiphydria. He considers that X. walshii West-
wood, which MacGillivray (1916) assigned to the genus Konowia belongs
to the original genus, although the species was unknown to him, and
states that "it is possible that it is provancheri Cresson." Rohwer suggests

431] LARVAE OF THE TENTHREDINOIDEA—YUASA 113

that the specimens which Patten (1878) reared from Betula nigra and
regarded as X. attenuata Norton do not belong to this species but are
similar to an undescribed female bearing a Bradley manuscript name.
Nothing is known concerning the immature stages of Konowia basalts
Say. The larvae of Xiphydria are wood-borers and confine their attacks
to dead and decaying wood of deciduous trees. European species infest
willows, poplar, elm, and birch, and American species, maple, hickory,
and birch. Konow (1901) listed four species, Xiphydria prolongata, X.
camelus, X. longicollis, and X. abdominalis in his key to the larvae of
Tenthredinoidea.

Xiphydria Fallen

Larvae comparatively small; thorax distinctly swollen, the meta-
thorax being the largest segment of the body; abdominal segments 1-2
cylindrical, subequal in diameter; third abdominal segment with a single
annulet; two caudal segments somewhat globose; sublateral lobe moder-
ately large, extending the entire length of the segment; suranal process
comparatively long, with dentiform tubercles near the base; tenth ab-
dominal sternum much smaller than the tergum; head and ultimate
segment with long setae; head semiglobose, vertical furrows present;
epicranial suture in part indistinct; antennariae distinct; antennae three-
segmented, segments 1 and 2 ring-like, segment 3 conical, longer but of
much smaller diameter than the preceding segments; labium very small,
without median emargination; mandibles with distinct dentes; maxillae
with small modified palpi, apparently two-segmented; galea conical,
smaller than palpi, lacinia fleshy, tubercle-like, with several setae; labium
with submentum and mentum large, convex, membranous, palpi appar-
ently with three segments, small, median lobe large, flattened on venter,
suboblong; spiracles large, oblique, not winged; glandubae wanting.

Xiphydria provancheri Cresson. — Length, 12 mm.; width of head,
1.6 mm.; body yellowish white; head creamy white; mouth-parts brownish;
suranal process arising from large brownish strongly chitinized suranal
lobe, deep brown, at proximal end, more than twice as long as wide,
distal two-thirds suddenly and distinctly smaller in diameter than proximal
third, with two circular rows of dentiform tubercles and setae on the
proximal third and two distinct ventral dentiform tubercles on the distal
third, one caudad of the other, these without setae; on birch.

This description is based upon a rare specimen collected at Saranac
Inn, New York State, Aug. 20, 1900, and generously loaned by Dr. E. P.
Felt. The larva bores into partly decayed heartwood of standing birch
and makes a gallery about 2.5 mm. in diameter. The burrows are invari-
ably filled with the borings, except a short curved portion thru which the
adult makes its way to the surface. A parasite, Pammegischia xiphydriae
Ashmead, was reared by Dr. Felt from larvae of this species.

114 ILLINOIS BIOLOGICAL MONOGRAPHS [432

Dr. Felt (1906) also found another larva making moderately large
cylindrical burrows in decaying birch and considered it as probably
belonging to X. attenuata. He refers to the rearing of this species by
Patton and suggests Rhyssa humida Say as its parasite. Since the identity
of Patton's specimen is questioned and since the adult was apparently
not reared, it is not possible to identify the specimen under consideration.
If it were really X. attenuata, then it should be known as X. abdominalis
as proposed by Konow (1905) and Rohwer (1918).

Family Siricidae

Body large, 30-40 mm., cylindrical, uniform in diameter thruout
(Fig. 4), fleshy, plump; integument smooth, transparent, non-setaceous,
light in color; head circular, half as high as thorax; mouth directed ventrad,
mostly exposed, but slightly overlapped by pro thorax; antennae incon-
spicuous, apparently one-segmented; ocellarae wanting; epicranial suture
wanting and vertical furrow indistinct; mouth-parts not normal in form,
light in color; prothorax large; mesothorax and metathorax short in
comparison with abdominal segments; legs rudimentary, mamma-like,
subequal in size, borne on fleshy conical lobes; larvapods wanting; typical
segment with two indistinct annulets, sublateral lobe distinct but not
prominent; tenth abdominal segment semiglobose in profile; tenth tergum
distinctly depressed by a median furrow; suranal lobe on the meson with
dark colored, chitinized, suranal process; subanal appendages wanting; in-
ternal-feeder, bores in the trunks of deciduous and evergreen trees.

The Siricidae contains five genera and about fifty species, most of
which are confined to the northern hemisphere. The recognition of the
fact that these insects constitute a well-circumscribed group dates back
to the time of Linnaeus (1758) who described five species of Siricidae among
those of his heterogeneous genus Ichneumon, three of the five having
become the types of three modern genera. Systematists have universally
agreed in considering this group worthy of family rank. The family
falls into two natural divisions, Siricinae, including three genera, Sirex,
Urocerous, and Xeris, and Tremecinae, embracing two genera, Tremex
and Teredon. The genus Xeris was associated with the genera composing
the Tremecinae both by Ashmead (1898) and Konow (1905), but Rohwer
(1911) proposed a more natural arrangement, placing this genus in the
Siricinae. Bradley (1913) definitely divided the two groups on the number
of segments of labial palpi and the retention of cerci in the adults.

According to Bradley (1913) there are twenty species reported for
North America, representing all the known genera. The specific charac-
ters of some common species, as Sirex nigricornis, -Urocerus cressoni, and
Tremex columba, are subject to a wide range of variation and several
varieties have been described. So far as known, the larvae of the Siricinae

433] LARVAE OF THE TENTHREDINOIDEA—YUASA 115

are wood-borers attacking conifers, those of the Tremecinae boring in
deciduous trees. Tremex columba, or three races of this species which
infest maple, elm, apple, pear, beech, oak, and sycamore, are the best-
known examples. The food-plants of only three American species are
known, these including Slrex cyaneus and Urocerus albicornis. Larvae of
the Siricinae have not been examined.

Tremex Jurine

Larvae conspicuously large; body cylindrical, slightly flattened on
the venter; large, robust, usually bare except on head and tenth abdominal
segment; whitish or creamy white; setae microscopic; head semiglobose,
slightly wider than high on cephalic aspect, semicircular in profile, pro-
duced to the ventral half of the front, then suddenly truncated, pale brown-
ish; mandibles and coilae deep brown; antennae apparently one-segmented,
conical; antacoria partly chitinized and bearing a few small setae; ocel-
larae wanting; depression mesad of antennaria which is sometimes called
the "eye" is a pretentorina; vertical furrows concealed by overhang-
ing prothorax; clypeus small, light in color; labrum transverse, con-
vex, thick, asymmetrical, without median emargination, but with a
notch on the right third of slightly oblique cephalic margin; mandibles
strong; mandacuta distinct, brown; mandibularia narrow, inconspicu-
ous, maxilla fleshy except subgalea, stipes large; palpi two-segmented,
small; galea conical, brown, small, arising from broad shoulder which
bears a few tiny setae on the lateral portion; lacinia round, lobe-like,
bearing three rows of brown setae, which decrease in length on cephalic
or dorsal side; labium compact, submentum narrow, transverse, mem-
branous, mentum convex, lobe-like, deeply emarginate on cephalic margin,
ligula round, fitting into the emargination of mentum, palpi small, two-
segmented, second segment much smaller than first, conical and brown,
sericos large, transverse, distinct, crescentic; prothorax large, produced
dorsad and cephalad, overlapping the caudal third of the head; mesothorac-
ic and metathoracic segments about one-half the length of abdominal seg-
ments except the first abdominal which is only little longer than the meta-
thorax; thoracic legs rudimentary, mamma-like, short, tipped with tiny
chitinized spot, borne on fleshy conical pedal lobe; cervical sclerites want-
ing; sternum with transverse subtriangular lobes which meet on the meson
in front of median lobe between and slightly cephalad of prothoracic legs;
metaspiracles as large as abdominal spiracles; abdomen slightly and uni-
formly tapering to the caudal end; annulation indistinct on dorsum,
apparently with but one annulet, the venter with two annulets, the
second annulet larger than the first; sublateral lobe prominent, extending
the entire length of segment as a single oblique elevation; spiracles large,
brown; ninth abdominal segment a little shorter than the eighth; tenth

116 ILLINOIS BIOLOGICAL MONOGRAPHS [434

tergum convex, lateral area of suranal lobe broad, suranal process promi-
nent, deep brownish, strongly chitinized, compressed, with two pairs of
small but distinct teeth.

The foregoing definition of the genus is based on one species, Tremex
columba.

Tremex columba Linnaeus. — Length, 40 mm.; width of head, 4 mm.;
ultimate segment with setae as follows: tergum near the caudal margin
on each side of the median furrow with a small, brown, sharp, hook-like
spine, with tiny setae which arise from large calices; ventral side of suranal
lobe with such setae; tenth sternum small; small brown spot at the lateral
end of anal slit; subanal lobe non-setiferous; subanal appendages wanting;
1-8396; G-.

The eggs of the Pigeon Tremex are oblong-oval, pointed at both ends
about 1.2 mm, in length, deposited singly, but in limited area, close to each
other; oviposition takes place in early summer, female sometimes fails to
withdraw ovipositor and dies in situ; larvae on hatching in the wood make
a gallery and feed for probably one season; transformation takes place in
the burrow; adults emerge thru circular hole, about 8 mm. in diameter.
The larvae are parasitizec by Thalessa lunator and also by Megarhyssa
atrata Fabricius, according to Champlain (1921),

Felt (1906) suggests as remedial measure against this insect, the cutting
down and burning of all trees badly infested. Keeping the trees in vigor-
ous health is supposed to be sufficient to prevent injury as the larvae work
only in weakened or partly decaying wood.

Family Megalodontidae

Antennae long, conspicuous, multisegmented, located above or near
the ocellarae; larvapods wanting; last abdominal segment rounded, with a
pair of bristle-like segmented subanal appendages; larvae feed on herba-
ceous plants.

This family contains four genera, Rhipidioceros, Megalodontes,
Melanopus, and Tristactus, and about thirty-five species, which are
distributed in Europe, Asia, and North Africa, Systematists have invari-
ably associated this family with the Pamphiliidae, but that this position is
unnatural has been conclusively shown by MacGillivray. He has pointed
out that it represents a line of specialization very similar to that found
in the Siricidae, and that while it is more closely related to this family
than to any other, an abundance of characters justify one in considering
it as a distinct group.

Only one species, Megalodontes spissicornis Klug has been recognized
in the larval stage. The larvae, according to Hiendlmayr (1878), are found
on Lasperpitium latifolium L. in central Europe from the end of July to

435] LARVAE OF THE TENTH REDINOIDEA — YU ASA 117

the beginning of August. In the younger stages, they are gregarious
and live in a common nest like many Pamphiliidae, but they spin an
individual web when half-grown. There is one generation a year.

This interesting species was not available for study, and the foregoing
definition is abstracted from Konow (1901) and may be found by later
students of Httle value in defining the family. This family, so far as the
recorded larval characters are concerned, seems to be closely associated
with the Pamphiliidae in possessing the bristle-like subanal appendages
and long conspicuous antennae. Future observations, however, may
possibly reveal more important characters, not given in the brief synopsis
of Konow.

Family Oryssidae

Body (Fig. 5) eruciform, grub-like, subcyhndrical, slightly depressed,
swollen in the middle of the abdomen, tapering at each end; segmentation
distinct; annulation obsolete; creamy white, without colored markings;
spiracles on prothorax and first eight abdominal segments; thorax increas-
ing in size caudad, thoracic legs obsolete; larvapods wanting; fourth
abdominal segment largest in diameter, size of segments decreasing rapidly
caudad, last segment smallest; suranal process and subanal appendages
wanting; head white, compressed cephalo-caudad, circular in frontal
contour, narrower than thorax; antennae with a single segment, papilla-
like; mandibles tridentate; maxillae and labium vestigial, fleshy, lobe-like,
without palpi; ocellarae wanting; larvae parasitic on wood-boring larvae of
Coleoptera; pupation in the pupal cells of the hosts.

The Oryssidae contain six genera and a limited number of species
distributed thruout the world. The genus Oryssus alone is represented
in the Nearctic region. In former years the family has been associated
with the Siricidae, but recently writers are in accord in regarding it as an
extremely specialized compact group. MacGillivray (1906) came to the
conclusion that "so far as their wings are concerned the presence of the
second anal cell in the front wings is the only structure that would place
the genus Oryssus in the superfamily Tenthredinoidea"; The group is
not only highly specialized in the adult characters but a recent discovery
of the parasitic habit of the larvae isolates these Hymenoptera from all
other Tenthredinoidea as a unique class. In fact Rohwer and Cushman
(1917) have gone so far as to propose a new suborder, Idiogastra, placing it
"intermediate between the suborder Chalastogastra — where adult would
place it — and the suborder Clistogastra — with which the larva would ally
it." Whether this arrangement is acceptable or not, the fact that this
group is remarkably well circumscribed and that it represents the summit
of an extremely isolated line of specialization in the Tenthredinoidea
can not be doubted.

118 ILLINOIS BIOLOGICAL MONOGRAPHS [436

Only one species, Oryssus occidentalis Cresson, has been recognized in
the immature stages. The definitions given here are based on the descrip-
tions and figures of this species published by Rohwer and Cushman (1917).
It is quite possible that future studies may prove them inadequate for
the identification of the larvae of other genera and species yet to be dis-
covered.

Oryssus Lateeille

Larvae small; epicranial suture faint, arms obsolete; clypeus cres-
centic, narrow; fronto-clypeal and clypeo-labral sutures distinct; labrum
more than twice as wide as long, with shallow mesal emargination; an-
tennaria distinctly elevated, antacoria extensive, mound-like; antennae
small, mamma-like; mandibles strongly chitinized, curved, narrow mesal
dentis larger than lateral dentes, these subequal in size, sharp; maxillae
fleshy, sub triangular, unsegmented lobes; annulation on dorsum indistinct,
with apparently two annulets, venter with one; sublateral lobes distinct,
extending the entire length of the segments; spiracles visible from dorsal
aspect; segments transversely raised and with a few minute tubercles.

Orysius occidentalis Cresson. — Color white with mandibles and chit-
inized ridges near the mouth brown; head one-third as wide as the widest
segment of the body — the fourth abdominal segment; maxillae with minute
brownish spots bearing about three sensory papillae; labium with about
four stout setae on each side of meson; prothorax declivous toward the
head, forming straight line with the latter in profile, on dorsum subequal
in length to mesothorax; metathorax half as long as mesothorax; lengths
of abdominal segments as follows: 8, (1, 9), (2, 7), (4, 6), 5, 10 on dorsum,
(6, 7, 8), 5, 1, 3, (2, 4), 9, 10 on venter, 8, 7, 4, (1, 2, 5, 6), 3, 9, 10 on latus;
tenth abdominal segment one-fourth as wide and one-third as high as the
fourth and sixth segments respectively; dorso-cephalic margin of mesothor-
acic and eighth abdominal segments distinctly, and of metathoracic and
abdominal segments 5, 6, 7, 9, 10 slightly convex, and of abdominal
segments 2 and 3 concave; ventro-cephalic margin of abdominal segments
1, 6, 7, 8 distinctly, and of 2 and 9 slightly, concave, of 3, 4, and 5 convex,
of 1 and 6 with a distinct convex emargination on each side of meson;
abdominal segments with the distance from spiracles to dorsal surface
uniform on lateral aspect and much shorter than the distance from spiracles
to the ventral surface, the latter variable and increasing caudad to the sixth
segment and diminishing thereafter; "each thoracic and abdominal segment
has dorsally at each side of the middle a low, transverse elevation sur-
mounted by a transverse row of four or five short, stout back-pointing
spines"; setiferous elevations on abdominal segments 1-7 and 9 and on
metathorax near the caudal margin of the segments, those of prothoracic

437j LARVAE OF THE TENTHREDINOIDEA—YUASA 119

and mesothoracic and eighth abdominal segments being in the middle;
tenth abdominal segment with small pointed protuberances directed cau-
dad; venter of segments with brownish spots in place of legs; larvae para-
sitic on the larvae of Buprestis confluens Say, B. laevenlris Le Conte, and
possibly other species of Buprestidae.

120 ILLINOIS BIOLOGICAL MONOGRAPHS (438

IV. PHYLOGENY

A classification based on phylogeny is one of the essential concerns of
philosophical taxonomy. In order to ascertain the genetic relationship
of organisms, synthetic as well as analytic, consideration of evidence
drawn from all the branches of biological science is imperative. The
indissoluble relation of morphology, embryology, and paleontology to tax-
onomy is so manifest and familiar that no comments are needed. The
time has come, however, when a critical examination of the phylogenetic
significance and the taxonomic value of the physiological and biological
attributes of animals must be made. Whatever evidence comparative
physiology, biochemistry, and genetics may offer should be incorporated
as far as possible with the data obtained in other more commonly ex-
ploited fields of research. Only in this way is it possible to arrive at a
comprehensive, systematic and complete summation of knowledge of
animals. This is the primary function of philosophical taxonom}?^, and in
this sense the saying of W. S. Jevons that "science can extend only so far
as the power of accurate classification extends," is true.

There are good reasons to believe, however, that even to-day morphol-
ogy, as of old, holds its supreme place in systematic investigations as it
offers fundamental assistance in determining the genetic affinities of
organisms. The success of a study of phylogeny based on morphological
evidences depends on the ability of the investigator to select the proper
structures, to determine the direction and nature of changes undergone by
these structures, and to draw legitimate conclusions by judicious interpre-
tation of the facts observed. Data obtained from studies of the external
anatomy of the larval stages of entometabolous insects are of necessity
incomplete of themselves for determining the phylogeny of the group;
yet, in the absence of other means of approach to the problem, they
constitute essential facts significant enough to merit careful consideration.

The opinions of scientists in regard to the systematic importance of
the characters based upon the immature stages of Entometabola have
been divided. There are some who ascribe no importance whatsoever to
them and entirely ignore this phase of taxonomy. There are others who
recognize the importance of the larvae from the viewpoint of synoptic
classification as they are primarily interested in the practical purpose of

439] LARVAE OF THE TENTHREDINOIDEA—YUASA 121

synoptic descriptions and keys. There are still others who believe in the
intrinsic importance of the immature stages in the study of phylogeny.
The reasonableness of the oft-repeated objection which was voiced by
Comstock (1918) that the larvae of insects exhibit a cenogenetic develop-
ment and, therefore their ontogeny bears little or no relation to the phylog-
eny of the race, must be admitted in regard to certain structures which
are entirely too adaptive and too much modified by environmental factors
in meeting the trophic requirements of particular species or genera. But
admission of this fact is not incompatible with a belief in palingenesis
of other structures. Besides, the warning that the cenogenetic peculiari-
ties, which may be of value as distinguishing characters, are of no phylo-
genetic significance and must, therefore, be judiciously and discriminately
distinguished from more important palingenetic characters, applies not
only to the classification of the larvae but to the taxonomy of the adults as
well. This objection alone does not invalidate a belief in the intrinsic
importance of the immature stages from the phylogenetic point of view.
While the writer does not minimize the danger of a too confident expecta-
tion of finding phylogenetic indices in the successive ontogenetic stages in
entometabolous insects, yet he is equally reluctant to abandon his hope in
regard to the taxonomic value of the characters of immature insects.
The present study is a partial justification of his contention.

Students of the Tenthredinoidea have recognized the practical im-
portance of the larvae in determining the systematic position of different
taxonomic units. Norton (1867) stated that "Mr. Walsh has shown
that in some species of Euura and Nematus bred by him, it was almost
impossible to detect any difference in the imago, while the larvae varied
greatly. Doubtless our opinion will be greatly modified by future dis-
coveries." Cameron (1882) was of the opinion that the larvae were of
great value in differentiating the tribes and subtribes altho they appeared
to be of little use in regard to the genera. MacGilHvray (1913) goes further
and states that "it was hoped from a study of the immature stages of the
Tenthredinoidea that some information might be obtained as to the
validity of the species based on obscure anatomical details." Rohwer
also often uses the characters of the larvae as collateral evidence in decid-
ing the systematic position of certain subfamilies and genera.

Nothing definite is known in regard to the ancestors of the Hymenop-
tera beyond the probability that they have somehow arisen from a primi-
tive type of some neuropteroid-like Palaeodictyoptera. The order is
considered to be one of the most, if not the most, highly specialized of all
insects. Systematists are unanimous in regarding the Tenthredinoidea
as the most generalized of the Hymenoptera. It is difficult, if not impossi-
ble, to conjecture the primitive larval type of the Tenthredinoidea.
Judging, however, from what are universally considered to be generalized

122 ILLINOIS BIOLOGICAL MONOGRAPHS (440

conditions in insects in general and in the Tenthredinoidea in particular,
the probable ancestral type of larva may be characterized as follows:
body cylindrical; segmentation distinct; annulation indistinct, annulets
few in number; head exposed, subglobose, distinct from the trunk; thorax
and abdomen more or less similar in structure excepting the three pairs of
thoracic legs, which are well developed and consist of five segments,
tarsal claws distinct; abdomen with twelve segments including the telson;
larvapods present on abdominal segments 1-10; antennae long, composed
of several segments; ocellarae present, one on each side of head; mouth-
parts well developed, maxillary and labial palpi segmented; tenth abdom-
inal tergum without caudal protuberances or suranal process; eleventh
abdominal sternum with a pair of segmented subanal appendages; ten
pairs of functional spiracles present, including metaspiracles; larvae free
leaf-feeders.

There are, as was pointed out by Comstock (1893), two kinds of char-
acters of phylogenetic importance. "First, characters indicating differ-
ence in kind of specialization; and second, characters indicating difference
in degree of specialization of the same kind. The former will indicate
dichotomous divisions of lines of descent; the latter merely indicate
degrees of divergence from a primitive type."

In determining the probable genetic affinities of the families of the
Tenthredinoidea, the following structures have been taken into considera-
tion: thoracic legs, larvapods, subanal appendages, ocellarae, antennae,
mouth-parts, suranal process, and metathoracic spiracles. The list does
not by any means exhaust the structures which might be employed for
this purpose, but it is believed that the structures listed offer the most
reliable and essential basis for the determination of a phylogeny based
upon larval characters. The significant changes in these structures are:
addition or reduction of parts; difference in degrees of development of
existing parts; and modifications in length, size, shape, and degree of
chitinization of the parts. These modifications have been interpreted
according to the Comstockian principles quoted above.

The thoracic legs are among the most persistent structures in the
adult and larval stages of insects in general, and their absence is unques-
tionably an indication of specialization by reduction. It is likewise
reasonable to assume that any modification of the typical, simple, cylin-
drical, five-segmented condition as regards the form or the number of
segments is a sign of specialization. The legs of the larvae of the Pam-
philiidae approximate most closely the primitive condition in the number,
shape, and structure of the segments. The tarsal claw is straight and
very slender. In the Xyelidae the legs assume a condition different from
that of the Pamphiliidae. The differentiation of segments in size and shape
has proceeded further and the tarsal claws have become distinctly claw-

441] LARVAE OF THE TENTHREDINOIDEA—YUASA 123 '

I

like. The legs are very small compared with the size of the body. The i

Tenthredinidae present a series of conditions which illustrate beautifully
cases of modification both by reduction and addition. The typical, well-
developed, five-segmented legs undoubtedly represent the normal sequence i
in development from the condition found in the Xyelidae. The apparently I
six-segmented condition of the Hylotominae, four-segmented legs of the !
Fenusinae, and three-segmented condition of the Schizocerinae, together I
with the development of distal fleshy lobes in the first- and last-named i
subfamilies are cases of specialization. The fact that the specialization
by reduction of segments has not proceeded at the same rate in the last ,
two subfamilies is indicated by the difference in the structure of the |
segments. It is interesting to note that the prothoracic legs of the Schizo- i
cerinae still retain four segments in spite of the fact that in the two caudal
pairs the number of segments has been reduced to three. The osculant
genus Phlebatrophia is unique among all other Tenthredinidae in having
the legs modified to such an extent as to lose all resemblance to normal j
segmented legs. They have become mere fleshy, indistinctly segmented, ;
clawless protuberances. In this character this genus resembles highly |
specialized families such as the Cephidae and its allies. The Cephidae, i
Xiphydriidae, and Siricidae represent a series of modifications in which the i
changes have resulted in fleshy, vestigial, entirely clawless legs with or with- I
out indication of segmentation. Judging from the size and degree of '
segmentation, the Xiphydriidae is more generalized than the Siricidae j
and more specialized than the Cephidae, The Oryssidae is entirely apo- ,
dous, and the fact that it is parasitic on buprestid larvae leaves no doubt
as to its extreme specialization.

The larvapods are considered as true appendages of the abdominal
segments. Their presence is highly significant from a phylogenetic point

of view. The larvae of the Tenthredinoidea are divisible into two types i

according to the presence or absence of the larvapods. The Xyelidae i

and Tenthredinidae represent the type with polypodous larvae and the '

other five families represent the type with apodous larvae. In the first j

group the Xyelidae possess the maximum number, or ten pairs, of larva- j

pods, while the Tenthredinidae are provided with six to eight excepting I

certain specialized genera which possess very vestigial or no larvapods, ]

It has not been possible to determine the reason for the invariable absence '

of larvapods upon the first and ninth uromeres in the Tenthredinidae. It i

may be that the same mechanical factors which have caused the fusion i

of anal larvapods in boring larvae like Caulocampus are also responsible :

for this condition. The size and, to some extent, the structure and position i

of the larvapods vary within the Tenthredinidae as in the Schizocerinae, !

Hylotominae, and Fenusinae. It is interesting to note that the gall-making ,

genus Pontania retains normal larvapods as well as thoracic legs, while the ;

124 ILLINOIS BIOLOGICAL MONOGRAPHS (442

leaf-miners have both thoracic and abdominal legs reduced in the number
and size of their segments. The small size and reduced number of seg-
ments are correlated with the well-developed thoracic legs of the Hylo-
tominae. In this subfamily the claws are very large, sharply curved, and
provided with empodia-like distal structures, indicating a great adapta-
tion for clinging to leaves. This fact is sufficient to account for the reduc-
tion of the larvapods. The Pamphiliidae and four specialized families
in the same line of development are entirely without larvapods. It is
highly desirable to determine whether this apodous condition signifies
a common origin of all five families. Upon this question hinges much of
the interpretation of the phylogeny of the Tenthredinoidea.

In characterizing the larvae of the hypothetical primodial Ten-
thredinoidea the abdomen was considered as provided with the maximum
number of appendages, including ten pairs of larvapods and a pair of
subanal appendages. This assumption is based upon the fundamental
fact that the progenitor of insects having evolved from a typical arthro-
podan organism possessed the typically arthropodan character, abdominal
appendages. This assumption is justifiable in view of the following facts:
(1) the possession of appendages on all of the abdominal segments is a
fundamental arthropodan characteristic; (2) the embryos of practically all
insects exhibit at some time during their development rudiments of
abdominal appendages; (3) appendages are present on all or some of the
abdominal segments in the postembryonic stages of the Apterygota;
(4) the gonapophyses of the Exometabola represent the true abdominal
appendages in this group of insects; (5) the larvapods and other appenda-
ges are present in the larvae of the Mecoptera, Lepidoptera, generalized
Hymenoptera, and, possibly, in some other orders, — all these facts indi-
cating the wide occurrence and fundamental continuity of abdominal
appendages in the Hexapoda. It is, therefore, not unreasonable to assume
that the progenitor of insects, at least in some stage of its development,
possessed appendages on all of the abdominal segments. The same
argument supports the contention that the ancestors of the Hymenoptera
undoubtedly closely resembled the remoter ancestors of the Insecta.
The larvae of the progenitor of the Hymenoptera for this reason have
been considered as provided with the maximum number, or ten pairs,
of larvapods, a pair on each of the first ten abdominal segments and a
pair of subanal appendages on the eleventh abdominal segment. If this
assumption is true, the larvae of the Xyelidae, which possess ten pairs of
larvapods, must be considered as representing the most primitive condition
found in the Hymenoptera. Graber (1890) has shown that in the larvae
of Hylotoma the larvapods arise from the embryonic limb-rudiments and
are directly evolved from them during the development and, therefore,
the larvapods are the true appendages of the abdomen, homodynamous

4431 LARVAE OF THE TENTHREDINOIDEA—YUASA 125

with the thoracic legs and homologous with the abdominal appendages of
generalized insects. There is no reason for considering the larvapods of
the Xyelidae as embryologically and morphologically different from those of
Hylotoma, consequently the larvapods of the Xyelidae must be the
true appendages of the abdomen; and since the larvae of this family
are provided with the maximum number of larvapods, they must be
considered as the most generalized of the Tenthredinoidea. The Ten-
thredinidae with six to eight pairs of larvapods and certain other mor-
phological and biological characters are unquestionably related to the
Xyelidae and probably represent a line of evolution from a xylelid-like
ancestral stock. Among the Tenthredinoidea with apodous larvae, the
Pamphiliidae, with a pair of segmented subanal appendages, is undoubtedly
the most generalized of all five families. The origin of the Pamphiliidae
is consequently an important question. For the reasons already stated
in connection with the larvapods, the progenitor of the Hymenoptera has
been considered as possessing a pair of subanal appendages on the caudal
segment of the body. In this character as well as in all others the Pamphilii-
dae approach most nearly the primitive condition and, except for the
absence of larvapods, unquestionably represents the most generalized
condition found in the Tenthredinoidea, ovitranking even the Xyelidae.
The loss of larvapods in this case is just as difficult to explain as the loss
of subanal appendages in the case of the Xyelidae. These structures, the
larvapods and subanal appendages, must have been lost during the course
of phylogeny since the progenitor undoubtedly possessed both of these
structures, and these two families, in spite of their generalized conditions,
must represent the end-products of evolution in their particular lines.
It is, then, natural and proper to assume that there have taken place two
distinct lines of development from the ancestral type of the Hymenoptera.
In the one, the specialization consisted in the suppression of the develop-
ment of larvapods, as in the Pamphiliidae, and in the other in the suppres-
sion of the development of subanal appendages, as in the Xyelidae. These
two families, then, represent two independent lines of evolution and are
the most generalized families not only of the Tenthredinoidea but of the
Hymenoptera. Whether the Xyelidae is more generalized than the
Pamphiliidae, or vice versa, must, from the very nture of the case, remain
a question till the advancement of our knowledge shall perhaps make the
answer possible. There are, however, a few things that should be pointed
out regarding this question. If the suppression of the development of
larvapods is considered of equal phylogenetic significance with the sup-
pression of the development of the subanal appendages, and if the head
and the appendages of these two families alone are compared, there is no
doubt that the Pamphiliidae are more generalized than the Xyelidae.
But since the subanal appendages are true abdominal appendages homo-

126 ILLINOIS BIOLOGICAL MONOGRAPHS [444

dynamous with the larvapods, and since it is natural to believe that the
process of reduction has taken place very slowly by gradual suppression
of the appendages, it is not unreasonable to assume that the apodous
condition found in the Pamphiliidae represents a much later stage of
specialization than the condition of the polypodous larvae of the Xyelidae.
The biology of the Pamphiliidae also indicates that this family is perhaps
more specialized than the Xyelidae. However, these considerations
counterbalance each other, and, when all is said, it is difficult to decide
between the two families as to their relative degrees of specialization.
This somewhat drawn-out discussion leads to the following conclusions:
(1) the progenitor of the Hymenoptera possessed a pair of larvapods on
each of the first ten abdominal segments and a pair of segmented subanal
appendages on the eleventh segment; (2) the progenitor gave rise to dis-
tinct stocks which resulted in the production of larvae with larvapods in
one case and with subanal appendages in the other; (3) the Xyelidae
represents the former line of evolution and the Pamphiliidae the latter;
and (4) the question as to whether the Pamphiliidae is more generalized
than the Xyelidae or vice versa is by its nature unanswerable. To the
above conclusions it may be added that it is only natural and reasonable to
consider the Tenthredinidae as representing the further evolution of the
primitive stock from which the Xyelidae had evolved, and the Cephidae,
Xiphidriidae, Siricidae, and Oryssidae, in turn, as evolving from the
original stock which gave rise to the Pamphiliidae.

The subanal appendages are present only in the Pamphiliidae and
Cephidae. In the former they are rather long, setiform, well developed,
distinctly three-segmented; in the latter they are minute, vestigial, often
fleshy, papilla-like, and indistinctly segmented. Since the embryonic
history of these appendages has not been studied, their true nature is not
known. There is little doubt but that they are true appendages. If they
represent the appendages of the ultimate segment, as has been suggested
by certain writers, and correspond to the so-called style of generalized
insects, then their presence is an indication of a primitive condition. There
is hardly any question as to the common origin of the subanal appendages
in the Pamphiliidae and Cephidae, and if these structures represent what
they are assumed to represent these two families must have a close affinity.

The ocellarae are present in the Pamphiliidae, Xyelidae, Tenthredin-
idae, and Cephidae. They are well developed, and are usually accompanied
by well-defined ocularia in the first three families. In the Cephidae the
ocellarae are vestigial and represented by localized pigmented granules,
and lack ocularia. It is significant that the ocellarae are unmodified in
the gall-makers and leaf-miners of the family Tenthredinidae, except in
Phlebatrophia, where they are reduced in size and the ocularia indistinct.
The atrophy of the ocellarae is undoubtedly correlated with the mining
habit of the larvae.

445] LARVAE OF THE TENTH REDINOIDEA—YU ASA 127

The antennae are present in all larvae of the Tenthredinoidea. Judg-
ing from the condition obtaining in generalized insects, it is reasonable to
consider the antennae of the Pamphiliidae as representing the primitive
ancestral type. They are long and setiform in this family and consist of
seven cyhndrical well-chitinized segments. The Xyelidae is closely
related to the preceding family in antennal characters altho a shortening
of the length has taken place. In the Tenthredinidae the antennae undergo
much modification both in the number and form of the segments. They
may be conical, limpet-shaped, or flattened, and the number of segments
varies from five to one. The antennae of the Cephidae resemble those of
the Xyelidae and some of the Tenthredinidae in shape and number of
segments. The antennae undergo steady reduction in size and number of
segments in the three remaining families, reaching the extreme of reduc-
tion in the Oryssidae, where each is represented by a button-like swelling.
The trend of specialization in the antennae is orthogenetic so far as the
families are concerned but quite diverse in the subfamilies of the Ten-
thredinidae.

The mouth-parts, which include the mandibles, maxillae and labium,
afford a fertile field for characters which are of interest from a systematic
point of view. The mandibles, like the antennae, are the most persistent
and ever-present structures in the head of all larvae of the Tenthredinoidea.
The maxillary and labial palpi are typically four-and three-segmented
respectively. The change is in the reduction in number and size of the
segments. The Cephidae is normal in this respect but gradual change
takes place in the Xiphydriidae and Siricidae, while in the Oryssidae the
change has proceeded so far as to completely obliterate the maxillary and
labial palpi. The palpi of Phlebatrophia resemble those of the specialized
famihes. The families represent different stages of specialization, and
their relative systematic position can be indicated by the degree of changes
in the mouth-parts.

The suranal process which is located on the meson of the suranal
lobe or the tenth urotergum is characteristic of the larvae of the Cephidae,
Xiphydriidae, and Siricidae. It should not be confused with the caudal
protuberances of certain Tenthredinidae, as these two structures are
of an entirely different nature. There is a minute hook-like process on the
caudo-meson of the tenth abdominal tergum of the Pamphiliidae. It
should be noted that in certain larvae of Pontania and Caulocampus the
caudal portion of the ultimate tergum is produced caudad as a blunt more
or less strongly chitinized protuberance which undoubtedly serves the
same function as the suranal process of the specialized families. These
two structural modifications of the caudal end of the body, however, are
not homologous with each other. The suranal process is undoubtedly

128 ILLINOIS BIOLOGICAL MONOGRAPHS [446

an adaptive structure which has arisen in response to the habit of the
larvae and does not represent the true appendages of the segment, to
which the suranal lobe belongs. For this reason the caudal process is of
less significance phylogenetically than the subanal appendages of the
Pamphiliidae and Cephidae.

The metathoracic spiracles of the larvae are either obsolete or vestigial
in the majority of the Tenthredinoidea. The larvae of the Cephidae and
Siricidae differ from all others in that the metaspiracles are functional
and as large as the abdominal spiracles. It is important to ascertain the
original condition of the metaspiracles in these families because upon the
interpretation of their primitive condition depends their phylogenetic
value and hence the relationship between these two families and also
between them and other families. It is considered reasonable to assume
that the progenitor of insects and hence the ancestor of the Hymenoptera
possessed functional spiracles on all the segments of the body including the
metathorax, and that their metaspiracles must have been as large as the
abdominal spiracles. The closed minute functionless metaspiracles found
in the Pamphiliidae, Xyelidae, and others, indicate a condition of atrophy
rather than a rudimentary condition, and so far as this character is con-
cerned the Cephidae and Siricidae represent the unmodified primitive
condition and some sort of relation between these two families must be
assumed. But on the basis of other characters it is not conceivable that
these two families evolved one from the other in a linear sequence, apart
from and independent of other families; they must have descended from
a common stock which also gave rise to other families which exhibit
vestigial metaspiracles. If this is true there must have taken place a
series of dichotomies starting with functional metaspiracles, one line of
development resulting in the loss of this primitive character and the other
line of evolution retaining the original condition. By assuming four such
successive dichotomies in the line of evolution, the origin and significance
of the metaspiracles of the Cephidae and Siricidae can be reasonably
explained. At each of the four successive dichotomous divisions which
produced respectively the pamphiliid-like progenitor and Xyelidae,
Pamphiliidae and the cephid-like progenitor, Cephidae and the xiphydriid-
like progenitor, and Xiphydriidae and Siricidae, one line of descent always
carried the original character and the other line lost it until this peculiarity
was generally sifted out, being retained unmodified only in the Cephidae
and Siricidae. In this way the metaspiracles are here considered to be
the direct descendant of the primitive structures which remained unmod-
ified thruout the course of evolution of these families. The two families
are generalized in this respect indicating a close genetic relation.

447]

LARVAE OF THE TENTHREDINOIDEA—YUASA

129

The morphological characters discussed are summarized in the following
table:

CoMPA«isoN OF Various Steuctujies in the Families of the Tenthsedinoidea

Structure

Pamphi-
liidae

Cephidae

Xiphy-
driidae

Siricidae

Oryssidae

Xyelidae

Tenth-
redinidae

Thoracic legs

S-seg-
mented

Fleshy

Fleshy

Fleshy

Wanting

S-seg-
mented

5-, 4-. 3-

«egmented

Larvapods

Wanting

Wanting

Wanting

Wanting

Wanting

10 pairs

6-8 pairs

Subanal

appendages

Distinct,
long

Vestigial

Wanting

Wanting

Wanting

Wanting

Wanting

Ocellarae

Distinct

Vestigial

Wanting

Wanting

Wanting

Distinct

Distinct

Antennae

7-seg-

5- and 4-

3- seg-

1-seg-

1-seg-

7- and 6-

5-, 4-. and 1-

mented

segmented

mented

mented

mented

segmented

segmented

Moulh-parts

Typical

Typical

Modified

Modified

Vestigial

Typical

Typical, rare-
ly modified

Suraual process

Wanting

Distinct

Distinct

Distinct

Wanting

Wanting

Wanting

Meta thoracic

spiracles

Vestigial

Functional

Vestigial

Functional

Vestigial

Vestigial

Vestigial

The Pamphiliidae, with its long seven-segmented antennae, setiform
three-segmented subanal appendages, setiform five-segmented thoracic
legs, well-developed typical mouth-parts, together with the absence of
larvapods, is unquestionably one of the most generalized families of the
Tenthredinoidea. This family differs from the hypothetical type only
in the absence of larvapods and reduced metaspiracles.

The Xyelidae, with its fairly long seven-and six-segmented antennae,
five-segmented thoracic legs, well-developed typical mouth-parts, together
with the presence of ten pairs of larvapods and the absence of subanal
appendages, is undoubtedly a very generaUzed family, quite different
from the preceding. The only striking difference from the hypothetical
type is the absence of the subanal appendages and functional metaspiracles.

The Tenthredinidae, with its one- to five-segmented antennae, well-
developed thoracic legs, and six to eight pairs of larvapods, together with
the absence of the subanal appendages, is unquestionably related to the
Xyelidae, and if it has not been evolved directly from the latter the two
families must have arisen from a common stock. The Tenthredinidae is a
phylogenetic complex in itself, and some of the more specialized genera are
further removed from the more generalized genera, biologically as well as
morphologically, than the latter are from the Xyelidae or their xyelid-
like ancestors.

The Cephidae, with its segmented antennae, vestigial subanal append-
ages, vestigial thoracic legs, normal mouth-parts, absence of larvapods,
presence of suranal process, vestigial ocellarae, and large functional
metaspiracles, is considered an offshoot of the ancestral stem from which

130 ILLINOIS BIOLOGICAL MONOGRAPHS [448

the Pamphiliidae had previously evolved. The specialization is indicated
by the vestigial condition of the ocellarae, subanal appendages, and
thoracic legs, on the one hand, and the development of suranal processes
on the other. The presence of the functional metaspiracles is of phy-
logenetic importance. So far as the head characters are concerned, this
family resembles the Tenthredinidae to a limited extent, and in some of
the generalized genera of the latter the thoracic legs and the caudal portion
of the tenth abdominal segment undergo some modifications which in a
remote sense simulate the condition in the Cephidae. But since this
family differs from the Tenthredinidae, and resembles the Pamphiliidae in
the absence of larvapods and the presence of subanal appendages, it is
considered more reasonable to ascribe to it a closer relationship to the
Pamphiliidae than to the Tenthredinidae.

The Xiphydriidae, with its somewhat modified mouth-parts, three-
segmented antennae, fleshly thoracic legs, suranal process, absence of
larvapods, the general shape of the body, and its biology, resembles the
Cephidae but differs from it in theabsence of subanal appendages, ocellarae,
and in the vestigial functionless metaspiracles. The absence of the subanal
appendages may point to one of the two possibilities in regard to the origin
of the Xiphydriidae. This family might have evolved from the cephid-
like ancester but have lost the subanal appendages by the completion of
the process of atrophy which had already reduced the original distinctly-
segmented appendages (similar to those of the Pamphiliidae) to the
vestigial papilliform appendages of the Cephidae. The two families
under consideration might, on the other hand, have had a common stem
which possessed subanal appendages, ocellarae, and vestigial metaspiracles.
In the absence of positive support for the first possibility, it is more
expedient to consider the second possibihty as nearer to the true rela-
tionship of the two families, Xiphydriidae and Cephidae.

The Siricidae, with its greatly reduced thoracic legs and mouth-parts
together with certain other characters, is considered more specialized
than the Xiphydriidae. The presence of the functional metaspiracles
and its genetic significance have already been discussed. For the same
reason which suggests a common origin for the Cephidae and Xiphy-
driidae, the Siricidae is considered to have arisen from a common stock
which gave rise also to the Xiphydriidae. In the degree of specialization
by reduction as well as by addition, this family outranks the Xiphydriidae.

The Oryssidae, with its vestigial mouth-parts, absence of ocellarae,
thoracic and abdominal legs, subanal appendages, suranal process, caudal
protuberances, and functional metaspiracles, together with its parasitic
habit, is unquestionably the most highly specialized family of the Ten-
thredinoidea. Its morphological and biological characters are so different
from other families that it is not easy to ascertain the systematic position

449] LARVAE OF THE TENTHREDINOIDEA—YUASA 131

of the family. There are, however, certain considerations which suggest
a possible relationship between this family and the Siricidae. Morpho-
logically the oryssid larvae are more closely related to the apodous boring
larvae of the Cephidae, Xiphydriidae, and Siricidae than to the polypodous
free-living larvae of the Xyelidae and Tenthredinidae. The siricid larvae
are more closely related to those of the Oryssidae than to those of the
Cephidae and Xiphydriidae. The parasitic habit also suggests a closer re-
lation to the wood-boring larvae since it is more plausible to imagine the
possibility of a wood-boring larvae becoming parasitic on other wood-
boring insect larvae under some unknown but not entirely inconceivable
circumstances than to imagine the development of a parasitic habit de novo
in free-living leaf-feeders. Since the oryssid larvae are parasitic on the
larvae of Buprestis inhabiting plants which are also infested by the larvae
of the Siricidae, if any transformation of habit of the larvae has taken
place, it is more natural to expect the larvae of the Siricidae or some
siricid-like insect to become parasitic than any other larvae. The recent
investigation by Baumberger (1919) on the role of microorganisms in
the physiology of insect nutrition offers a valuable suggestion in regard to
the possibility of radical changes in food habits. For these reasons it is
considered reasonable to ascribe a common progenitor to the Siricidae
and Oryssidae, at least for the time being. It may be added that it is
not entirely unreasonable to assume an independent line of evolution
for the Oryssidae apart from all other Tenthredinoidea and consider
this family as having no close relation to any of the modern families of the
Tenthredinoidea. In that case, the Oryssidae must have arisen from the
ancestral stock before the Pamphiliidae and Xyelidae had their origin.
There is, however, no clear evidence in support of such relation and
since the relation is reasonably explained by associating the Oryssidae
with the Siricidae, the former is considered the most highly specialized
family of the Tenthredinoidea with a common origin with the ancestor of
the Siricidae.

The conclusions on the systematic position and relationship of the
different families of the Tenthredinoidea based exclusively on larval
characters and derived entirely independent of the opinions of the spe-
cialists who have paid more attention to the adults are of necessity not
the final words on the subject. The true significance of such conclusions
lies in their complemental and collateral value. It is interesting on this
account to compare the writer's opinion with the conclusions of the
modern authorities on this group of the Hymenoptera.

The relationship suggested here supports in its essential points the
three more important systems of classification proposed by Konow (1905),
MacGillivray (1906), and Rohwer (1911). MacGillivray considered the
Xyelidae the most primitive because of the venational character but

132 ILUNOIS BIOLOGICAL MONOGRAPHS (450

recognized the Pamphiliidae as the most generalized from his study (1913)
of the immature stages. Had he placed the Pamphiliidae before the
Xyelidae and the Cephidae before the Xiphydriidae and Siricidae, his
system would coincide exactly with the system based exclusively on the
immature stages. Konow and Rohwer both associate the Cephidae
with the Pamphiliidae and Xyelidae. This arrangement is partly supported
if the afl&nity between the Pamphiliidae and Cephidae, as suggested in
this study, is upheld. The genetic continuity of the Xyelidae and Ten-
thredinidae is clearly recognized by MacGillivray (1913). The true
systematic position of the Tenthredinidae or its equivalent is difficult
to express in linear arrangement. The important point to be noted is the
fact that these authors and also Morice (1919) consider the Xyelidae and
Tenthredinidae as different and apart from the other families of the
Tenthredinoidea. The Oryssidae unquestionably merits at least separate
family rank. In the absence of requisite knowledge of the larval characters
of the Hymenoptera other than the Tenthredinoidea, it is not expedient
to venture any opinion on the suggestion made by Rohwer and Cushman
(1917) to establish a third suborder, Idiogastra, for the reception of the
Oryssidae. Enslin (1911) differs from the authors already mentioned
not only in his arrangement of the groups in a descending order but also in
treating the Xyelidae and Pamphiliidae as subfamilies of his Tenthredini-
dae, on a level with the Cimbicini, Lophrini, and others. This study does
not support his arrangement. Morice (1919) suggested that "the Lydini
(Pamphiliidae Ensl.) may represent a primitive group of Tenthredinidae
which had branched off from the main stock before it had develoi>ed
certain characters," such as abdominal legs. Handlirsch (1908) considered
the Siricidae as having evolved from the osculant Juracic group, Pseudo-
siricidae. The antiquity of the Siricidae is accepted by Morice who
expresses his idea of the relationship of the families of the Tenthredinoidea
as follows: "We may suppose that the Siricidae are the earlier group,
but whether the Tenthredinidae and Lydini had Siricid ancestors, or
whether the Siricidae +Cephini+ Oryssidae and Tenthredinidae-!- Lydini
are respectively earlier and later branches of a common stock are questions
which must be left unanswered."

4511 LARVAE OF THE TENTHREDINOIDEA—YUASA 133

V. SUMMARY

The larvae of the Tenthredinoidea have proved to be of great value in
aflFording important evidence in regard to the probable phylogenetic
relationship of the families included in this superfamily. The more
significant conclusions reached in this study are summarized in the form
of a synoptic key as follows:

FAMILIES OF TENTHREDINOIDEA

Larvapods present, thoracic legs present, well developed, distinctly segmented.

Larvapods present on all abdominal segments Xyelidae

Larvapods never present on 1st and 9th abdominal segments Tenthredinidae

Larvapods wanting, thoracic legs present or wanting.
Thoracic legs present.
Thoracic legs and subanal appendages well-developed and distinctly segmented.

Pamphiliidae.

Thoracic legs vestigial, indistinctly segmented.

Subanal appendages and ocellarae present Cephidae.

Subanal appendages and ocellarae wanting.
Metaspiracles vestigial, much smaller than abdominal spiracles . . . Xiphydriidae.

Metaspiracles functional, as large as abdominal spiracles Siricidae.

Thoracic legs wanting Oryssidae.

A synopsis such as the foregoing is necessarily inadequate and some-
what misleading in indicating the aJEnities of the families. A better idea
is gained by means of the customary phylogenetic tree, altho such a
scheme also has its limitations. In the following diagram the relation
between the families of the Tenthredinoidea is shown. Here the relative
vertical positions are intended to represent approximately the degree
of specialization; and the continuous lines, the affinities.

The larvae of the Tenthredinoidea are thus divisible into two distinct
groups. The first group includes the larvae characterized by the presence
of both the thoracic and abdominal legs, and by the absence of the subanal
appendages and suranal process, and is represented by the Xyelidae
and Tenthredinidae. The second group consists of the five families,
Pamphiliidae, Cephidae, Xiphydriidae, Siricidae, and Oryssidae, and is
divisible into two subgroups. The first subgroup contains the first four
families and is characterized by the absence of abdominal legs, by the

134

ILLINOIS BIOLOGICAL MONOGRAPHS

1452

presence of vestigial clawless thoracic legs in the last three families, and
by the presence of either subanal appendages or suranal process or both.
The second subgroup contains a single family, Oryssidae, which is charac-
terized by the absence of both thoracic and abdominal legs, suranal

Highly
Specialised

Oryssidae

.^ ^. ^. .J Siricidae /
Tenthredinxdae / /

(specialiaed)l Xiphydriidae / /

Specialized

Tenthredijiidae / //
(generalized) / // Oephidae

Generalized

'"■"xH i/""'""-

Primitive

1

Fro-Syaphyta

Pbylogenetic tree indicating the probable affinities of various families of the Tenthredinoidea

process, subanal appendages, and segmented maxillary and labial palpi.
The Xyelidae and Pamphiliidae are undoubtedly the most primitive of the
first and second groups respectively.

The Tenthredinoidea, therefore, is considered to have developed
from a common ancestral stock along two distinct lines of evolution.
The first line of development led to the evolution of the Xyelidae and
Tenthredinidae and the second line produced the Pamphiliidae, Cephidae,
Xiphydriidae, Siricidae, and Oryssidae.

453] LARVAE OF THE TENTHREDINOIDEA—YUASA 135

VI. BIBLIOGRAPHY

Andre, E.

1879. Species des hym^nopetres d'Europe et d'Alg^rie. Vol. 1. Beaune.
ASHMEAD, W. H.

1898. Classification of the horn tails and sawflies, or the suborder Phytophaga. Can.
Ent., 30:141-145, 177-183, 205-213, 225-232, 249-257, 281-287, 305-316.
Baumberger, J. p.

1919. A nutritional study of insects with special reference to microorganisms and their
substrata. Jour. Exp. Zool., 28:1-82.
BoviNG, A. G.

1914. On the abdominal structure of certain beetle larvae of the campodeifonn type.
A study of the relation between the structure of the integument and the muscles.
Proc. Ent. Soc. Wash., 16:51-61.

Bradley, J. C.

1913. Siricidae of North America. Jour. Ent. and Zool., 5:1-35.
Brischke, C. G. a., and Zaddach, G.

1883. Beobachtungen iiber die Arten der Blatt- und Holzwespen.
Britton, W. E.

1906. The maple leaf -stem borer or sawfly. Can. Ent., 17:313-321.

1915. A destructive pine sawfly introduced from Europe. Jour. Ec. Ent., 8:370-382.
Brues, C. T.

1919. The classification of insects on the characters of the larva and pupa. Biol. Bull.,
37:1-21.

BOCHNER, P.

1918. Vergleichende Eistudien. 1. Die akzessorischen Kerne des Hymenoptereneiens.
Archiv f. mikrosk. Anat., Bd. 91, 3-4 Heft. 1-202.
Burke, H. E.

1917. Oryssus is parasitic. Proc. Ent. Soc. Wash., 19:87-89.
Cameron, P.

1882-1883. A monograph of the British phytophagous Hymenoptera. The Ray Society,
London, 1882-83, Vols. 1-4.
Castle, W. E.

1900. Metamerism of the Hirudinae. Proc. Am. Acad. Arts and Sci., 35:285-303.
CHAMPLAINf, A. B.

1921. The curious mating habit of Megarhyssa atrata (Fab.) Ent. News, 32:241.
Cholodkovsky, N.

1897. Entom. Miscellen. V. Uber Spritzapparate der Cimbicidenlarven. Horae Soc
Ent. Rossicae, 30.

1897. Uber das Bluten der Cimbicidenlarven. Horae Soc. Ent. Rossicae, p. 2.
COMSTOCK, J. H.

1893. Evolution and taxonomy. Wilder Quarter-Century Book. pp. 37-113. Ithaca.

1918. Nymphs, naiads, and larvae. Ann. Ent. Soc. Am.. 11:222-224.

136 ILLINOIS BIOLOGICAL MONOGRAPHS t454

Ckampton, G. C.

1918. The thoracic sclerites of immature pterygotan insects, with notes on the relation-
ships indicated. Proc. Ent. Soc. Wash., 20:39-65.

1919. The genitalia and terminal abdominal structures of males, and the terminal
abdominal structures of the larvae of "chalastogastrous" Hymenoptera. Proc.
Ent. Soc. Wash., 21:129-255.

1921. The sclerites of the head, and mouth-parts of certain immature and adult insects.

Ann. Ent. Soc. Am., 14:65-110.
Cresson, E. T.

1887. Synopsis of the famihes and genera of the Hymenoptera of America, North of

Mexico, together with a catalogue of the described species, and bibliography.

Trans. Am. Ent. Soc, Suppl. Vol. 1887, pp. 1-351.
Dahlbom, G.

1835. Conspectus Tenthredinidum, Siricidum, Oryssinorum, Scandinaviae quas

HjTimopterorum familiae. Hafniae.
Dalla Torre, C. G.

1894. Catalogus Hymenopterorum hucusque descriptorum systematicus et sjoionymi-

cus. Vol. 1. Tenthredinidae incl. Uroceridae.
De Geer, C.

1752. M6moires pour servir a I'histoire des insectes. Stockholm.
Doncaster, L.

1907. Gametogenesis and fertilization in Nematus rihesii. Quar. Jour. Micr. Sci.,

51:101-113.
Dyar, H. G.

1893a. Notes on two species of Tenthredinidae from Yosemite, Calif. Can. Ent.,

25:195-196.
1893b. Descriptions of the larvae of certain Tenthredinidae. Can. Ent., 25:244-248
1894. Descriptions of the larvae of certain Tenthredinidae. Can. Ent., 26:42-45,

185-189.
1895a. Descriptions of the larvae of certain Tenthredinidae. Can. Ent.., 27:191-196,

208-212.
1895b. On the larvae of some Nematoid and other sawflies from the northern Atlantic

states. Trans. Am. Ent. Soc, 22:301-312.
1895c. The larvae of the North American sawflies. Can. Ent., 27:337-344.
1896. Notes on sawfly larvae. Can. Ent., 28:235-239.
1897a. On the larvae of certain sawflies (Tenthredinidae). Jour. New York Ent. Soc,

5:18-30.
1897b. New Sawflies (Tenthredinidae) with descriptions of larvae. Jour. N. Y. Ent.

Soc, 5:190-201.
1897c. Some structural points in sawfly larvae. Proc. Ent. Soc. Wash., 14:218-220.
1897d. A new sawfly. Proc. Ent. Soc. Wash., 4:262-263.
1898. Notes on some sawfly larvae, especially the Xyelidae. Can. Ent., 30:173-176.

1898. On the larvae of certain Nematinae and Blennocampinae, with descriptions of
new species. Jour. N. Y. Ent. Soc, 6:121-138.

1899. Larvae of Xyelidae. Can. Ent., 31:127.

1900. On the larvae of Atomacera and some other sawflies. Jour. N. Y. Ent. Soc, 8:
26-31.

Enslin, E.

1911. Zur systematik der Chalastogastra. Deutsch. Ent. Zeitschr., 1911, 434-439.

1912. Die Tenthredinoidea Mitteleuropas. 1. Deutsch. Ent. Zeitschr., 1912, Beiheft
EssiG, E. O.

1914. The cherry fruit sawfly. Month. Bull. Cal. State Comm. Hort., 1914, 31.

455] LARVAE OP THE TENTHREDINOIDEA—YUASA 137

Fabrictus, J. C.

1775. Systema Entomologiae sistens Insectorum classes, ordines, genera, species,
adjectis synonjTnis, locis, descriptionibus, observationibus. Lipsiae.
Felt, E. P.

1906. Insects afTecting park and woodland trees. Memoir N. Y. State Museum, No. 8.
Fernald, H. T.

1903. The plum webbing sawfly. Ent. News, 14:298-302.
Forbes, W. T. M.

1914. A structural study of the caterpillars. III. Somatic muscles. Ann. Ent. See.
Am., 7:109-124.
Foster, S. W.

1913. The cherry fruit sawffy. Bull. U. S. Bur. Ent., No. 116, Part III.
Frenzel, J.

1885. Einiges iiber den Mitteldarm der Insekten, sowie iiber Epithelregeneration.
Arch. mikr. Anat., 26:229-306.
Frisch, J. L.

1766. Beschreibung von allerley Insecten in Deutschland, etc. Vol. 1.
Gahan, a. B.

1920. Black grain-stem sawfly of Europe in the United States. Bull. U. S. Dept. Agr.,
No. 834.
Gerstaecker,

1867. GattungOxybelus. Arch. Naturg., 1867, 20.
Goedart, J.

1682. Johannes Goedartius on insects, etc. York.
Graber, V.

1890. Vergleichende Studien am Keimstreif der Insekten. Denkschr. d. Math. Natur-
wiss. Classe d. Akad. d. Wiss. Wien, 57:621-734.

Hall, W. B.

1917. Notes on the immature stages of Hemitaxonus mullicinctus Rohwer. Proc. Ent.
Soc. Wash., 19:28.
Handlirsch, a.

1908. Die Fossilen Insekten und die Phylogenie der rezenten Formen. Leipzig.
Hartig, T.

1837. Die Familien der Blattwespen und Holzwespen, nebst einer allgemeinen Ein-
leitung zur Naturgeschichte der Hymenopteran. Berlin.
Hewitt, C. G.

1912. The large larch sawfly, with an account of its parasites, other natural enemies
and means of control. Bull. Div. Ent. Dept. Agr. Dom. of Canada, No. 10.

HiENDLMAYR, A.

1878. Tarpa spissicornis KLlug. Mt. MOnch. Ver. II, 163. (Kriechbaumer, Ent. Nachr.
4:169-170.)

HOLTZ, H.

1909. Von der Secretion und Absorption der Darmzellen bei Nematus. Anat. Hefte.
Wiesbaden, Abt. 1, Bd. 39:683-696.

Houghton, C. O.

1910. The blackberrj' leaf-miner. Bull. Delaware Agr. Exp. Sta., 87.
Jack, J. G.

1891. Notes on three species of Hylotoma. Psyche, 6:10-11.

1893. Notes on Taxonus nigrisoma and T. dubiiatus. Can. Ent., 25:183-184.
Jack, R. W.

1917. The turnip &&w&y (A thaltaflacca). Rhodesia Agr. Jour., 14:206-212.

138 ILLINOIS BIOLOGICAL MONOGRAPHS [456

KoNOw, F. W.

1890. Catalogus Tenthredinidarum Europae. Deutsche ent. Zeitschr. 1890, 241-254.
1901. Systematische Zusammenstellung der bisher bekannt gewerdenen Chalastogastra

Mecklenburg.
1905a. Hymenoptera. Fam. Lydidae. Wytsman's Genera Insectorum, Fasc. 27.
1905b. Hyemnoptera. Fam. Siricidae. Ibid., Fasc. 28.
1905c. Hyemnoptera. Fam. Tenthredinidae. Ibid., Fasc. 29.

KORSCHELT, E., UND HeIDER, K.

1892. Lehrbuch der vergleichenden Etnwicklungsgeschichte der wirbellosen Thiere.
Heft 2. (English translation, 1899).
Latreille, a.

1796. Pr6cis des Caracteres gen6riques des insectes, disposes dans un ordre natural par
le Citoyen Latreille. Bourdeaux.
Lepeletier de Saint Fargeau, A.

1823. Monographia Tenthredinetarum, synonymia extricata. Paris.
1836. Histoire naturelle des Insectes. Hymenopteres. Vol. IV. Paris.
Linnaeus, C.

1744. Systema Naturae. 4th ed. Paris.
1746. Fauna Suecica sistens animalia Sueciae regni, etc.
1758. Systema Naturae. 10th ed. Homiae.
Lyonet, p.

1762. Traite anatomique de la Chenille, qui rouge le Bois de Saule, etc La Haye.
MacGillivray, a. D.

1906. A study of the wings of the Tenthredinoidea, a superfamily of Hymenoptera.

Proc. U. S. Nat. Museum, 29, 569-654.
1913. The immature stages of the Tenthredinoidea. Ann. Rept. Ent. Soc. Ontario,
44: 54-75.

1916. Tenthredinoidea. Conn. Geol. Nat. Hist. Survey. Bull. No. 22:25-175.
Marlatt, C. L.

1895. The hibernation of nematids, and its bearing on inquilinous species. Proc. Ent.

Soc. Wash., 3, 263-267.
1887. Notes on a red cedar sawfly. Trans. Kansas Acad. Sci., 1885-6, 10:80-83.
1890. The final molting of tenthredinid larvae. Proc. Ent. Soc. Wash., 2:115-117.
Merian, Maria S.

1730. Histoire des Insectes de I'Europe, dessines d'apres nature et expliqu6s par Marie

Sibille Merian, etc. La Haye.
MiDDLETON, W.

1915. Notes on some sawfly larvae belonging to the genus Dimorphopteryx. Proc.

U.S. Nat. Mus. 48:497-501.

1917. Notes on the larvae of some Cephidae. Proc. Ent. Soc. Wash., 19:174-179.
1921a. LeConte's sawfly, an enemy of young pines. Jour. Agr. Res., 20:741-760.
1921b. Some notes on the terminal abdominal structures of sawfly. Proc. Ent. Soc.

Wash., 23:139-144.

1921c. Some suggested homologies between larvae and adults in sawflies. Proc. Ent.
Soc. Wash., 23:173-192.

1922. Sawflies injurious to rose foliage. U. S. Dept. Agr. Farmers' Bull. 1252.
MORICE, F. D.

1919. Notes on Australian sawflies, especially the "authors' types" and other speci-
mens in the British Museum of Natural History and the Hope Collections of the
Oxford University Museum; with diagnostic synopsis of the genera and species,
and photographs illustrating their structural characters. Trans. Ent. Soc.
London, 1918:247-333.

457) LARVAE OF THE TENTHNEDJNOIDEA—YUASA 139

MOUTET, T.

1634. Insectorum sive minimorum animalium theatrum, etc.
Nelson, J. A.

1915. The embryology of the honey-bee. Princeton Univ. Press.
Norton, E.

1867-69. Catalogue of the described Tenthredinidae and Uroceridae of North America.
Trans. Am. Ent. Soc, 1867-69, 1: 31-84, 193-280; 2:211-242, 321-368.
Packakd, a. S.

1883. Third Report U. S. Ent. Comm., 307, 324, 435.

1890. Fifth Rept. U. S. Ent. Comm., being a revised and enlarged edition of Bull.
No. 7, on insects injurious to forest and shade trees.

Parrott, p. J., AND Fulton, B. B.

1915. Cherry and hawthorn sawfly leaf-miner. Jour. Agr. Res., 5:519-528.
Pearsall, R. F.

1902. Life history- of Lyda fasciata (Norton), Fam. Tenthredinidae. Can, Ent., 34:
214-216.
Poletajew, N.

1885. Ueber die Spinndriisen der Blattwespen. Zool. Anz., 8:22-23.
Riley, C. V., and Marlatt, C. L.

1891. Wheat and grass sawflies. Insect Life, 4:168.
ROHWER, S. A.

1910. Hoplocampa. Can. Ent., 42:242-244.

1911. A classification of the suborder Chalastogastra of the Hymenoptera. Proc.
Ent. Soc. Wash., 13:215-226.

1911a. Genotypes of the sawflies and woodwasps, or the superfamily Tenthredinoidea.

BuU. Tech. Series, U. S. Bur. Ent. No. 20, Part 2.
1911b. Additions and corrections to "the genotypes of the sawflies and woodwasps, or

the superfamily Tenthredinoidea." Ent. News, 22:218-219.

1912. Notes on sawflies, with descriptions of new species. Proc. U. S. Nat. Mus.,
43:205-251.

1915. A remarkable new genus of Cephidae. Proc. Ent. Soc. Wash., 17:114-117.
1917. The American species of the genus Cephus Latreille. Proc. Ent. Soc. Wash.,

19:139-141.
1918a. New Sawflies of the subfamily Diprioninae. (Hym.). Proc. Ent. Soc. Wash.,

20:79-90.
1918b. Notes on, and descriptions of the Nearctic woodwasps of the genus Xiphydria

Latreille (Hym.) Ent. News, 29:105-111.
1918c. Notes on, and descriptions of sawflies belonging to the tribe Hemichroini (Hym.).

Proc. Ent. Soc. Wash., 20:161-173.

ROHWER, S. A. AND CuSHMAN, R. A.

1917. Idiogastra, a new suborder of Hymenoptera with notes on the immature stages
of Oryssus. Proc. Ent. Soc. Wash., 19:89-98.
Sasscer, E. R.

1911. Notes on a sawfly injurious to ash. Proc. Ent. Soc. Wash., 13:107-110.
SCHWARZ, E. A.

1909. Illustrations of the life history of a sawfly (Hylotoma Pectoraiis Leach) injurious
to willow. Proc. Ent. Soc. Wash., 11:106-108.
Snellen von Vollenhoven.

1868. Die inlandsche Bladwespen. Tijdschr. voor Ent., 11:
Stephens, J. F.

1835. Illustrations of British entomology; or synopsis of indigenous insects, etc.
London.

140 ILLINOIS BIOLOGICAL MONOGRAPHS [458

SWAMMERDAM, J.

1737. Biblia naturae, etc. Leydae.
SWENK, M. R

1911. A new sawfly enemy of the bull pine in Nebraska. 24th Ann. Rept. Neb. Agr.
Exp. Sta., 1911.

TOWNSEND, H. T. '

1894. Notes on the Tenthredinid gall of Euura orbitalis on Salix and its occupants.
Jour. N. Y. Ent. Soc, 2:102.
Walden, B. H.

1912. A new sawfly pest of the blackberry, Pamphilius dentatus MacGillivray. Rept.
Conn. Agr. Exp. Sta., 1912, 236-240.

Walsh, B. D.

1866. Imported insects; the gooseberry sawfly. Pract. Ent., 1:117-125.
Webster, F. M.

1900. The purslane sawfly, Schizocerus zabriskei Ashm. Can. Ent., 32:51-54.
Westwood, J. O.

1840. An introduction to the modem classification of insects. London.
Young, Chester.

1898. The larvae of the Tenthredina. Baccalaureate thesis, Cornell University.

1899. Descriptions of sawfly larvae. Can. Ent., 31:41-43.
Yuasa, H.

1920. The anatomy of the head and mouth-parts of Orthoptera and Euplexoptera. Jour.
Morph., 33:251-307.

459] LARVAE OF THE TENTHREDINOIDEA—YUASA 141

PLATE 1

142 . ILLINOIS BIOLOGICAL MONOGRAPHS (460

EXPLANATION OF PLATE
LARVAE OF TENTH REDINOIDEA

Fig. 1 — Pamphiliidae. Pamphilius sp. Y-125.

Fig. 2 — Cephidae. Janus integer.

Fig. 3 — Xiphydriidae. Xiphydria sp.

Fig. 4 — Siricidae. Tremex columba.

Fig. 5— Oryssidae. Oryssus occidenialis

(After Rohwer and Cushman, 1917).
au anus

msp mesothoracic spiracle
sba subanal appendage
spi abdominal spiracle
srp suranal process
//g^ mesothoracic leg
tsp metathoracic spiracle

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

8 S 10 „

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE I

4611 LARVAE OF THE TENTHREDINOIDEA—YUASA 143

PLATE II

144

ILLINOIS BIOLOGICAL MONOGRAPHS

1462

EXPLANATION OF PLATE
LARVAE OF XYELIDAE AND TENTHREDINIDAE

Fig. 6 — Xyelidae.
Fig. 7 — Tenthredinidae.
Fig. 8 — Tenthredinidae.
Fig. 9 — Tenthredinidae.
Fig. 10 — Tenthredinidae.

Megaxyela major.

Diprioninae. Neodipriron lecontei.
Emphj^inae. Emphytus apertus.
Selandriinae. Strongylogaster anmilosits.
Dolerinae. Dolerus similis.

au
nisp

Pk

spi
tig'
tsp

anus

mesothoracic spiracle
larvapod

abdominal spiracle
mesothoracic leg
metathoracic spiracle

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE II

4dJl LARVAE OF THE TENTHREDINOIDEA—YUASA 145

PLATE III

146

ILLINOIS BIOLOGICAL MONOGRAPHS

[464

Fig. 11 — Phyllotominae.
Fig. 12 — Phyllotominae.
Fig. 13 — Tenthredininae.
Fig. 14 — Cimbicinae.
Fig. 15 — Hoplocampinae.

EXPLANATION OF PLATE
LARVAE OF TENTHREDINIDAE

Caliroa cerasi.
Phlebatrophia mathesoni.
Tenthredo sp.
Ahia inflata.
Hemichroa americana.

au
msp

pig
spi
Ug'
tsp

anus

mesothoracic spiracle
larvapod

abdominal spiracle
mesothoracic leg
metathoracic spiracle

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE III

465] LARVAE OF THE TENTHREDINOIDEA—YUASA 147

PLATE IV

148

ILUNOIS BIOLOGICAL MONOGRAPHS

[466

Fig. 16 — Cladiinae.
Fig. 17 — Nematinae.
Fig. 18 — Nematinae.
Fig. 19 — Blennocampinae.
Fig. 20 — Blennocampinae.

EXPLANATION OF PLATE

LARVAE OF TENTHREDINIDAE

Cladius peclinicornis.
Pkronidea ventralis.
Pteronidea ribesi.
Monophadnoides rubi.
Tomostethus bardits.

au anus

tnsp mesothoracic spiracle

pig larvapod

spi abdominal spiracle

srp suranal process

tig* mesothoracic leg

tsp metathoracic spiracle

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE IV

4671 LARVAE OF THE TENTHREDINOIDEA—YUASA 149

PLATE V

150

ILLINOIS BIOLOGICAL MONOGRAPHS

[4«8

EXPLANATION OF PLATE

LARVAE OF TENTHREDINIDAE

Fig. 21 — Fenusinae.

Kaliofenusa ulmi.

Fig. 22 — Scolioneurinae.

Metallus rubi.

Fig. 23 — Hylotominae.

Hylotoma sp.

Fig. 24 — Schizocerinae.

Schizocerus zabriskei.

Fig. 25 — Acordulecerinae.

Acordulecera sp.

a7i anus

msp mesothoracic spiracle

pig larvapod

'

scp sucker-like protuberance

spi abdominal spiracle

tlg^ mesothoracic leg

tsp metathoracic spiracle

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

7 8

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE V

469] LARVAE OF THE TENTHREDINOIDEA—YAUSA 151

PLATE VI

132

ILLINOIS BIOLOGICAL MONOGRAPHS

[470

EXPLANATION OF PLATE

CEPHALIC ASPECT OF THE HEAD

Fig. 26 — Pamphilius sp.

Fig. 27 — Megaxyela major.

Fig. 28 — Neodiprion leconki.

Fig. 29 — Abia americana.

Fig. 30 — Pteronidea ribesi.

Fig. 31 — Lygaeonematus erichsoni.

Fig. 32 — Endelomyia aethiops.

Fig. 33 — Macremphytus varianus.
Fig. 34 — Kaliofenusa ttlmi.
Fig. 35 — Metallus rubi.
Fig. 36 — Schizocerus zabriskei.
Fig. 37 — Phlcbatrophia mathesoni.
Fig. 38 — Dolerus similis.

a

antenna

an

antacoria

or

antennaria

c

clypeus

cl

clypealia

els

clypeo-labral suture

crv

sericos

cs

clypeal suture

ea

epicranial arm

es

epicranial stem

f

front

fcs

f ronto-clypeal suture

si

galea

hx

hyj)opharynx

inl

line of invagination

I

labrum

ip

labial palpus

Is

labral setae

ma

muscular attachment

mb

mandibularia

md

mandible

mds

mandibular setae

mp

maxillary palpus

mx

maxilla

0

ocellara

ou

ocularia

pe

preclypeus

pn

pretentorina

Po

postclypeus

Pr

precoUa

py

preartis

V

vertex

vf

vertical furrow

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE VI

471] LARVAE OF THE TENTHREDINOIDEA—YUASA 153

PLATE VII

154

ILLINOIS BIOLOGICAL MONOGRAPHS

(472

EXPLANATION OF PLATE

VENTRAL ASPECT OF THE HEAD AND PROTHORAX

Fig. 39 — Pamphilus sp.

Fig. 45 — Xiphydria sp.

Fig. 40 — Megaxyda major.

Fig. 46 — Tremex columba.

Fig. 41 — Kaliofemisa tdmi.

Fig. 47 — Lygeonematus erichsoni.

Fig. 42 — Dolerus similis.

Fig. 48 — Endelomyia aethiops.

Fig. 43 — Neodiprion lecontei.

Fig. 49 — Metallus rubi.

Fig. 44 — Macremphytm varianus.

a

antenna

I

labrum

an

antacoria

la

lacinia

c

clypeus

Ics

lateral cervical sclerite

cc

cervacoria

It

labium

cl

clypealia

ip

labial palpus

ds

cljTpeo-Iabral suture

mb

mandibularia

cm

sericos

md

mandible

cw

tarsal claw

mp

maxillary palpus

ex

coxa

mx

maxilla

ea

epicranial arm

0

ocellara

ec

extensacuta

Pfn

profurcellina

es

epicranial stem

pn

pretentorina

f

front

Pr

precoila

fcs

fronto-clypeal suture

py

preartis

fm

femur

t

tibia

S

gena

tr

trochanter

il

galea

Ug'

prothoracic leg

hx

hypopharynx

V

vertex

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE VII

473] LARVAE OF THE TENTHREDINOIDEA—YUASA 155

PLATE VIII

156

ILUNOIS BIOLOGICAL MONOGRAPHS

[474

EXPLANATION OF PLATE

VENTRAL AND DORSAL ASPECTS OF HEAD AND PROTHORAX

Fig. 50 — Janus integer.

Fig. 51 — Schizocerus zabriskei.

Fig. 52 — Phlebalrophia matJiesoni.

Fig. 53 — Caliroa cerasi.

Fig. 54 — Pamphilius sp.

Fig. 55 — Dolerus similis.
Fig. 56 — Megaxyela major.
Fig. 57 — Caliroa cerasi.
Fig. 58 — Neodiprion lecontei.
Fig. 59 — Endelomyia aethiops.

a

antenna

la

lacinia

c

dypeus

le

labicoria

cc

cervacoria

It

labium

eg

cervical gland

IP

labial palpus

els

cl3^eo-labral suture

m

mentum

crv

sericos

mb

mandibularia

cw

tarsal claw

md

mandible

ex

coxa

tnp

maxillary palpus

ea

epicranial arm

msp

mesothoracic spiracle

ee

extensacuta

mx

maxilla

es

epicranial stem

0

ocellara

f

front

pfn

profurcellina

fm

femur

pn

pretentorina

g

gena

t

tibia

gl

galea

Ug'

prothoracic leg

hx

hypopharynx

ts

tarsus

is

intersegmental line (limit of samite)

V

vertex

I

labnim

vf

vertical furrow

ILLINOIS BIOLOGICAL MOXOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE VIII

4751 LARVAE OF THE TENTHREDINOIDEA—YUASA 157

PLATE IX

15S

ILLINOIS BIOLOGICAL MONOGRAPHS

1476

EXPLANATION OF PLATE

DORSAL AND LATERAL ASPECTS OF HEAD AND PROTHORAX

Fig. 60 — Macremphytus varianus.
Fig. 61 — Monophadnoides nibi.
Fig. 62 — Meiallus ruhi.
Fig. 63 — Fampkilius sp.
Fig. 64 — Megaxyela major.
Fig. 65 — Neodiprlon leconlei.
Fig. 66 — Dolerus siniilis.

Fig. 67 — Macreviphytus varianus.
Fig. 68 — Endelomyia aethiops.
Fig. 69 — Caliroa cerasi.
Fig. 70 — Pteronidea ribesi.
Fig. 71 — Lygeonematus erichsoni.
Fig. 72 — Abia americana.

a

antenna

Ics

lateral cervical sclerite

c

clypeus

li

labium

cc

cervacoria

IP

labial palpus

<-g

cervical gland

mb

mandibularia

cw

tarsal claw

md

mandible

ex

coxa

mp

maxillary palpus

ea

epicranial arm

msp

mesothoracic spiracle

cc

extensacuta

mx

maxilla

es

epicranial stem

0

ocellara

f

front

Pr

precoila

fm

femur

py

preartis

S

gena

t

tibia

hx

hypopharynx

tr

trochanter

is

intersegmental line (limit of somite)

V

vertex

I

labrum

vf

vertical furrow

la lacinia

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE IX

4771 LARVAE OF THE TENTHREDINOIDEA—YUASA 159

PLATE X

160

ILLINOIS BIOLOGICAL MONOGRAPHS

[478

EXPLANATION OF PLATE
LATERAL AND VENTRAL ASPECTS OF TmRD ABDOMINAL SEGMENT

Fig. 73 — Pamphilius sp.
Fig. 74 — Endelomyia aethiops.
Fig. 75 — Dolerus similis.
Fig. 76 — Megaxyela major.
Fig. 77 — NeodipHon lecontei.
Fig. 78 — Macremphytus varianus.
Fig. 79 — Caliroa cerasi.
Fig. 80 — Pamphilius sp.

Fig. 81 — Dolerus similis.
Fig. 82 — Neodiprion leconki.
Fig. 83 — Endelomyia aethiops.
Fig. 84 — Megaxyela major.
Fig. 85 — Caliroa cerasi.
Fig. 86 — Kaliofenusa ulmi.
Fig. 87 — Metallus ruhi.
Fig. 88 — Macremphytus varianus.

al-a7 annulets 1, 2, 3, 4, 5, 6, 7.

ch chitinized area.

cor intersegmental coria

pig larvapod

sdl surpedal lobe or area

spi abdominal spiracle

ssl subspiracular lobe or area

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA

LARVAE OF THE TENTHREDINOIDEA PLATE X

479] LARVAE OF THE TENTH REDINOIDEA—YUASA 1«1

PLATE XI

162

ILLINOIS BIOLOGICAL MONOGRAPHS

[480

EXPLANATION OF PLATE
LATERAL AND VENTRAL ASPECTS OF CAUDAL ABDOMINAL SEGMENTS

Fig. 89 — Megaxyela major.

Fig. 90 — Megaxyela major.

Fig. 91 — Pamphilius sp.

Fig. 92 — Macremphylus varianus.

Fig. 93 — Macremphytus varianus.

Fig. 94 — Caliroa cerasi.

Fig. 95 — Pamphilius sp.

Fig. 96 — Dolcrus similis.

Fig. 97 — Caliroa cerasi.

Fig. 98 — Endelomyia aethiops.

Fig. 99 — MetaJlus rubi.

Fig. 100 — Neodiprion lecontei.

Fig. 101 — Neodiprion lecontei.

Fig. 102 — Dolerus similis.

Fig. 103 — Metallus rubi.

Fig. 104 — Phlebatrophia mathesoni.
Ventral aspect of third ab-
dominal segment.

au

anus

al

annulet 1

aZ

annulet 2

a6

annulet 6

ch

chitinized area

pig

larvapod

sba

subanal appendage

sbl

subanal lobe

sdl

surpedal lobe or area

spi

abdominal spiracle

srl

suranal lobe

ssl

subspiracular lobe or area

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE XI

481] LARVAE OF THE TENTHREDINOIDEA—YUASA 163

PLATE XII

164

ILLINOIS BIOWGICAL MONOGRAPHS

[482

EXPLANATION OF PLATE
CAUDAL ASPECT OF HEAD, ABDOMEN, ABDOMINAL APPENDAGES

Fig. 105 — Kaliojenusa ulmi. Ventral
aspect of caudal abdominal segment.

Fig. 106 — Phlebatrophia mathesoni.

Fig. 107 — Xiphydria sp.

Fig. 108 — Cephus pygmaeus.

Fig. 109 — Adirus trimactilakis.

Fig. 110 — Xiphydria sp.

Fig. 1 1 1 — Hartigia cressoni.

Fig. 112 — Janus integer.

Fig. 113 — Tremex columba.

Fig. 114 — Cephus pygmaeus. Caudal end
of the abdomen enlarged.

Fig. 115 — Adirus trimaculatus.

Fig. 116 — Adirus trimaculatus. Subanal ap-
pendage enlarged.

Fig. 117 — Hartigia cressoni. Subanal
appendage enlarged.

Fig. 118 — Janus sp. Subanal appendage
enlarged.

Fig. 119 — Cephus Pygmaeus. Subanal
appendage enlarged.

Fig. 120 — Tremex columba. Lateral aspect of

suranal process enlarged.
Fig. 121 — Janus integer. Lateral aspect of

suranal process enlarged.
Fig. 122 — Tremex columba. Dorsal aspect of

suranal process enlarged.
Fig. 123 — Janus integer. Dorsal aspect of

suranal process enlarged.
Fig. 124 — Pteronidea ribesi. Ventral aspect

of third abdominal segment.
Fig. 125 — Pteronidea ribesi Lateral aspect

of caudal end of abdomen.
Fig. 126 — Pteronidea ribesi. Lateral aspect

of caudal end of abdomen.
Fig. 127 — Pteronidea ribesi. Dorsal aspect of

caudal end of abdomen.
Fig. 128 — Abia americana. Caudal aspect of

the head.
Fig. 129 — Neodiprion lecontei. Musculature

of third abdominal segment, semidia-

grammatic. Annulets numbered.

au

anus

my

maxillaria

cc

cervacoria

od

odontoidea

ch

chitinized area

of

occipital foramen

cht

chitinized tubercle

OS

occipital suture

cor

coria

pa

postgena

ct

corpotentorium

Pgr

postgenal ridge

es

epicranial stem

Pl

paracoila

li

labium

pig

larvapod

IP

labial palpus

ptl

postcoila

Isr

lateral area of suranal lobe

pa

postgena

mb

mandicoria

sba

subanal appendage

mc

maxacoria

sbl

subanal lobe

mn

metatentorina

sll

sublateral lobe

mp

maxillary palpus

spi

abdominal spiracle

mt

metatentorium

srp

suranal process

mx

maxilla

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

129

127

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE XII

4831

LARVAE OF THE TENTHREDINOIDEA—YUASA 165

PLATE XIII

166 ILLINOIS BIOLOGICAL MONOGRAPHS [484

EXPLANATION OF PLATE

ANTENNAE, LEGS, SILK-GLANDS ,

Fig. 130 — Pamphilius sp. Metathoracic leg.

Fig. 131 — Megaxy da major. Metathoracic leg. \

Fig. 132 — Macremphytus varianus. Metathoracic leg.

Fig. 133 — Pteronidea ribesi. Mesothoracic leg.

Fig. 134 — Neodiprion lecontei. Metathoracic leg.

Fig. 135 — Dolerus similis. Metathoracic leg.

Fig. 136 — Phlebatrophia mathesoni. Metathoracic leg.

Fig. 137 — Metallus rubi. Metathoracic leg.

Fig. 138 — Pteronidea ribesi. Mesothoracic leg. Dorsal aspect.

Fig. 139 — Schizocerus zabriskei. Metathoracic leg.

Fig. 140 — Kaliofenusa ulmi. Metathoracic leg.

Fig. 141 — Caliroa cerasi. Metathoracic leg.

Fig. 142 — Endelomyia aeihiops. Metathoracic leg.

Fig. 143 — Megaxyela major. Antena.

Fig. 144 — Adirus trimaculattts. Antenna.

Fig. 145 — Schizocerus zabriskei. Antenna.

Fig. 146 — Cephus pygmaeus. Antenna. i

Fig. 147 — Metallus rubi. Antenna. i

Fig. 148 — Tremex columba. Antenna. j

Fig. 149 — Kaliofenusa ulmi. Antenna. i

Fig. 150 — Phlebatrophia tnathesoni. Antenna. |

Fig. 151 — Janus integer. Antenna. ]

Fig. 152 — Hartigia cressoni. Antenna. >

Fig. 153 — Thrinax impressatus. Antenna. I

Fig. 154 — Pteronidea ribesi. Antenna. j

Fig. 155 — Tremex columba. Mesothoracic spiracle. )

Fig. 156 — Abia americana. Silk-glands. '

Fig. 157 — Abia americana. Labium, hypopharynx, and portion of silk-glands, enlarged. j

Fig. 158 — Abia americana. Lateral aspect of cephalic portion of silk-glands.

Fig. 159 — Abia americana. Dorsal view of silk-press, hypopharynx removed. i

Fig. 160 — Abia americana. Cephalic view of labium and hypopharynx.

oi-od antennal segments 1, 2, 3, 4, 5, and 6

an antacoria li labium

crv cericos Ip labial palpus

cw tarsal claw M arrow pointing mesad

ex coxa sd duct of sUk-glands

D arrow pointing dorsad sgc cells of silk-gland

fm femur sgd small duct of silk-glands

fp femoral process slpr silk-press

hx hypopharynx

ILLINOIS BIOLOGICAL MOXOGRAPHS

VOLUME VII

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE XIII

485i LARVAE OF THE TENTHREDINOIDEA—YUASA 167

PLATE XIV

168 ILLINOIS BIOLOGICAL MONOGRAPHS I486

EXPLANATION OF PLATE

Chart representing graphically the relationship of various taxonomic units or groups
according to the more important modern systems of classification of the Tenthredinoidea as
proposed by Konow (1905), MacGillivray (1906), and Rohvrer (1911-1918).

ILLINOIS BIOLOGICAL MONOGRAPHS

VOLUME VII

"~S

MORF. IMPORTANT MODRRN 8YSTKM.S OF CLASSIWCAnnHOFTHRTRmHRRDinOinRi

KONOW
1903

MACGILLIVRAY

ROHWER^
19 11 ~l 8

CHALASTOGASTKA TENTHREDINOIDEA CHALASTOGASTRA

I LYDIDAB
Lydioi
Me^Alodootidfs

Lvdidcs
CepKioi
Xyelini
& last! cotomi III

ftSIRICID>iE
Xiphydrmi
Siricini

Tieinccldes
Ory^sini

nrTENTHREOmiDAt

Cimbicini
Cimbicides
Abi '\(\es
5yzygor\iid«s

Ahjioi

5chizocerid«s j
LophyHni
Loboccrotid<

Perrcyides
Lophyr-ides

TentKn?dinini

HcmatidC^

Hap)<7cevinpid«

5|ennocam[ride

SelandriadCi-

Dolcridcs

Teothrcdin^s

7

(I XYEUDAB
la PAMPHILIIDAE
/ID DLASTlCOTOrilDAE
\crTEHTHRED)HIDAE
/Di prion inae (fji^l

/SelaodriioaC

Dolerioac
^Pbyllotoininae

/ /Lyca«rfio&e
/i JTen1*>fK<li"'n»
/( ICi'mbicinae

Heplocampin&e

Dioenrioae

^lonoct^m^^ac^

Cladiir\ae
Hemaiirsike
^lenooCampinaf \
FeouSinaC
Scolioneurinae,
Hylotofnir«ve
^ ScKiioceri nac

Tei'»-«yiir\ac

Lpbocerin»e
Acordulcccrinac^

Pt « ry3 ophorinaer^

rV XIPHYDRIIDAE
VI.SIWCIDAE

Siricina^
TremecioA^

vnriEGALODOriTIDAE

vidcephidae

aORYSSIDAE

MEGALODONTOIDEA 1
riEGALOOOMTIDAE

Pa«n(>hiliir>ae

XYEUDAE
CEPHIDAE
ORmOIDEA-IDIO-TU

SIRECOIPEA ('^'^^ittl
XIPHYDRIIDAE.
XipWdriinac
Dcrecystinat?
SIRECIDAE
5ii-ccina^
Sirecioi
Kcri I oi
\rvmec\nB>e

TEHTHREDINOIDEA Hi
CIMBECIDAE

Cimbecini
Pbenaiopfi^irti
Z&t-arinae

Rwudoclayffllar;,-,,^

PERREYIIDAE

Perreyi iliac

Phi I o masti^ i n >v^

ARGIDAE.

Slericli phorinaff
\BLASTICOTOniOAE

\

\

NTHREDIrtlDAE
, 1 DipriooinAc )«»iai
\\ AlJantiDAc
Y Taxonini
EHocampirii
■ Allarrtini
Dol^rioac
Totbrcd inina^
P*riocurii>i
TrnHirrdinini

V P'TrMotomir*!
\ * Messioi
VAtbaliinat?
V Empriir***'
Empriini
Lycatrlioi
PJenimca mptni
Pl>ytn&tacerinat>
Sclaiwjfiioo^
Strtandriini

'Cladi inaf

Ncrr>a+i_r»af

Htina^iMi ... „,
Henicht-oirtt (iqia)

PTERY&OPHORIPAE
Pt^ r-y^opKo ri r» »*

Phyla<-topKa9ir\ae
Euriinae 121

PERGIDAE
^LOBOCERIDAE

YUASA LARVAE OF THE TENTHREDINOIDEA PLATE XIV

4871

LARVAE OF THE TENTHREDINOIDEA—YUASA

169

INDEX

Page

abbotti 47

abbreviaius 1 10

abdominalis 113

Abia 65

Abia, species of 66

abietis 47

abdomen 25

acericaulis 69

Acordulecera 103, 104

Acordulecera, species of 103

Acordulecerinae 44

Acordulecerinae, subfamily 103

Adirus 109, 110

aethiops 58

albifrons 55

americana 65, 66

Amauronemaius 76, 84

Amauronematus, species of 84

annulosus 53

antennae 19, 127

appendicuiatus 77

apriciis 55

arthrostyli 29

attenuata 114

aviingrata 41

azaleae 84

bardus 93

basalts 113

bethunei 99

bilineata 62

bivUtata 79

Blasiicotoma 42

Blasticotomidae, family 42

Blennocampa 92, 93

Blennocampinae 43, 44

Blennocampinae, genera of 92

Blennocampinae, subfamUy 91

borers 38

Caliroa 58

Caiiroa, species of 59

caudal protuberances 27

Caulocampus 67, 68, 69

Page

Cephaleia 106

Cephidae 129

Cephidae, family 108

Cephidae, genera of 109

Ceplius 109, 111

Cephus, species of Ill

cerasi 59

cerv^acoria 22

ckloreus 83

Cimbex 65

Cimbicinae 44

Cimbicinae, genera of 65

Cimbicinae, subfamily 64

cinctus 109, 111

Cladkis 71, 74

Cladiinae 44

Cladiinae, genera of 71

Cladiinae, subfamily 70

clypeus 17

columba 116

cornelli 86

corpotentorium 19

cressoni 112

Croesus 76, 83

demissa 89

deniatus 107

derosa 90

despecta 69

devincta 89

Dimorphopteryx 51

Dineura 69

Dineurinae, subfamily 69

Diphadnus 75, 76, 77

Diprion 46, 49

Diprioninae 43

Diprioninae, subfamily 45

dohrini 97

Dolerus 55

Dolertis, species of 56

Dolerinae 43

Dolerinae, subfamily 54

dorsalis 104

%..

170

ILUNOIS BIOLOGICAL MONOGRAPHS

[488

Page

Dorytheus 55

dyari 67

efeta 87

effliisa 87

emerita 87

Empkylus 51

Emphytinae 43, 44

Emphytinae, genera of 50

Emphytinae, subfamily 49

Empria 50

Endelomyia 58

epicranial suture 15

epinota 63

Epitaxonus 50

equina 88

erkhsoni 81

Eriocampa 51

erudita 87

Erythraspjdes 92, 94

erylhrogaslra 86

evanida 87

fasciaia 106

Fenusa 97

Fenusinae 45

Fenusinae, genera of 96

Fenusinae, subfamily 96

ferruginea 42

flida 63

filiceti 42

fistula 63

flavipes 54

formulae of segmented appendages 34

free leaf-feeders 37

front 17

fulmcrus 87

fumipennis 95

gall-makers 88

genae 16

glands 33

glandubae H

gregarious 80

Hartigia 110, 112

head 14

Hemichroa 67

Hemitaxonus 50

Hoplocampinae 44, 45

Hoplocampinae, genera of 67

Hoplocampinae, subfamily 66

hyalina 89

Hylotoma 100, 101

Page

Hylotoma, species of 100

Hylotominae 45

Hylotominae, subfamily 99

Hypergyricus 92, 94, 95

Hypergyricus, species of 94

impressatus 53

inanitus 105

inconspicua 107

inflata 66

integer 110

Isodictiutn 92, 95

Janus 109, 110

Janus, species of 110

Kaliofenusa 97

Konoma 113

labium 21

labrum 17

laricis 68

larvapods 29, 123

latitarsus 83

leaf-folder 89

leaf-miners 37

lecontei 47

lineata 64

lombardae 87

longicollis 1 13

luteotergmn 84

Lygaeonematus 75, 81

macleayi 101

Macremphylus 51

Macrophya 61 , 62

Macrophya, species of 63

M acgilliv-rayella 67

Macroxyela 42

major 41

mandibles 20

mandibularia 18

Marlattia 67, 68

mathesoni 60

maxillae 21

maxillariae 18

Megalodontes 116

Megalodonlidae, family 116

Megaxyela 41

mendica 88

Metallus 98

Meiallus, species of 99

metamerism 30

metatentoria 19

metathoracic spiracles 128

489]

LARVAE OF THE TENTHREDINOIDEA—VUASA

171

Page

Micronematus 75, 79, 100

minor 42

minutus 55

ftivdestius 73

Monoctenus 46, 48

Monophadnoides 92, 95

Monophadnus 92, 94

Monosoma 51

Monostegia 51

mouth-parts 20, 127

muUicindus 93

murtfeldtiae 78

musta 104

Nefnatus 75, 82

Nematus, species of 83

Nematinae 44

Nematinae, genera of 75

Nematinae, subfamily 74

Neodiprion 46, 47, 48

Neodiprion, species of 47

Neopus 61, 62

nest-builders 38

Neurotoma 106, 107

nubiUpennis 94

obsoleia 59

occidentalls 118

occipital foramen 18

occiput 18

ocellarae 126

ocreatus 105, 108

ocuiaria 16

Odontoidea 19

Odontophyes 41

odoratus 86

Oryssidae 130

Oryssus 118

Oryssidae, family 117

Pachynematus 76, 81

PachynematuSy species of 82

paetidus 72

palliolatus 73

pamphiliid 105

Pamphiliidae 129

Pamphiliidae, family 104

Pamphiliidae, species of 105

Pamphilius 105, 107, 108

parasites 38

paracoila 19

Parataxonus 50

parvuliis 69

Page

patchiae 72

pectinicornis 74

Peridista 92

Phlebatrophia 60

Phlebatrophini 57

Phlebatrophini, tribe 59

Phrontosoma 51

Phyllotominae 44, 45

Phyllotomini 57

Phyllotomini, genera of 58

Phyllotominae, subfamily 57

poliliis 54

pomwn 89

Pontania 75, 76, 88, 98

Ponlania, species of 88

postcoUa 18

precoila 18

postpedes 29

pretentoria 19

prepharynx 22

Priophorus 71, 73, 74

Priophorus, species of . 73

Pristiphora 75, 77, 78

Pristiphora, species of 77

prolegs 29, 30

Prolongaia 113

postgenae 18

provanckeri 113

Pseiidodineura 69

Pkronidea 75, 76, 85, 86

Pteronidea, species of 85

pulatus 53

ptdchella 64

pygmaeae 94, 109, 112

quercus-alba 59

qu^cus-coccinea 59

14-punctalns 62

reperlus 82

ribesi 85, 87

rubi 95,99

scapidaris 101

Schizocerinae 45

Schizocerinae, subfamily 101

Schizocerus 102

Scolioneurinae 45

Scolioneurinae, subfamily 98

Selandria 52, 54

Selandriinae 43, 44

Selandriinae, genera of 52

Selandriinae, subfamily 51

172

ILUNOIS BIOLOGICAL MONOGRAPHS

[490

Page

semilutea 61

setae 32

simile 49

similis 55, 56

Siricidae 130

Siricidae, family 114

spiracles 32

spiraeae 93, 94

spisskornis 116

Strongylogaster 52, 53

Strongylogaster, species of 53

Slrongylogastroidea 51

subalbaius •. . 82

subanal appendages 28, 126

suranal lobe 26, 27

suranal process 27, 127

sychophanta 79

Syntexis 1 10

tabidus Ill

tacitus 54

Taxontis 51

Tenthredinidae 129

Tenthredinidae, family 42

Tenthredinidae, subfamilies of 43

Tenthredininae 44

Tenthredininae, genera of 61

Tenthredininae, subfamily 60

Tenthredinoidea, families of 38, 133

Tenthredinoidea, superfamily 37

Tenthredo 61, 62

Tenthredopsis 61

tenth urostemum 27

Page

tenth urotergum 26

tentorium 19

thoracica 85

thoracic legs 23, 122

thorax 23

Thrinax 52

Thrinax, species of 53

tibialor 63

Tomostethus 92, 93

tormae 18

Trachelus 109, 111

Tremex 115, 116

Trichiocampus 71, 73

Trichiocampus, species of 72

Trichiosoma 65

trilineata 88

trimactdatus 110

tnmk 22

ulmi 97

unicolor 48

Unitaxonus 51

ventralis 85

vertex 16

vescus 85

virendus 84

Xiphydria 113

Xiphydriidae 130

Xiphydriidae, family 112

Xyela 42

XyeUdae 129

Xyelidae, family 39

zabriskiei 102

I

-1

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII January, 1922 No. i

STUDIES ON GREGARINES II

SYNOPSIS OF THE POLYCYSTID GREGARINES

OF THE WORLD, EXCLUDING THOSE FROM

THE MYRIAPODA, ORTHOPTERA,

AND COLEOPTERA

WITH FOUR PLATES ?

<

BY

MINNIE WATSON KAMM

Price $1.00

Published by the University op Illinois Press

UNDER THB AUSPICES OF THE GRADUATE SghOOL

Urbana, Illinois

UNIVERSITY OF ILLINOIS STUDIES PUBLISHED
ILLINOIS BIOLOGICAL MONOGRAPHS
The Illinois Biological Moijogsaphs is a serial published quarterly by. the Univer-
sity of Illinois. It is the purpose of the publication to present in monographic form research
contributions of especial merit and of somewhat extended character. Each volume will
include about 500 pages and 20 or more full page plates. The series may be obtained at
a subscription price of three dollars yearly. Individual numbers will be sold separately at
prices determined by the size and amount of illustration contained in the particular number.

, '.' ■ ■;/ ■■;.■',■ ■ 'Vol.1.,'; /,■:
Nos. 1 and 2. A Revision of the Cestode family Proteocephalidae. With 16 plates. Ey

G. R. LaRue. $2.00.
No. 3. Studies on the Cestode family Anoplocephalidae. 6 plates. By H. Douthitt. 80 cts.
No. 4. Some North American Larval Trematodes. With 8 plates. By W, W. Cort. $1.20.

Vol.11

No. 1. Classification of Lepidopterous larvae. With 10 plates. By S. B. Fracker. $1,50.

No. 2. On the Osteology of some of the Loricati. With5plates. ByJ. E.Gutberlet. 50 cts.

No. 3. Studies on Gregarines, including twentyrone new species and a synopsis of the
Eugregarine records from the MyriapOda, Coleoptera, and Orthoptera of the
world. With 15 plates. By Minnie E. Watson. s$2.C0.

No. 4. The Genus Meliola in Porto Rico. , With 5 plates. By Frank L. Stevens. 75 cts.

Vol. Ill
No. 1. Studies on the factors controlling regeneration. By Charles Zelen)'. $1.25.
No. 2. The head-capsule and mouth-parts of Diptera. 25 plates. By A. Peterson. $2.00.
No. 3. Studie« on North American Polystomidae, Aspidogastridae, and Pararrtphistomidae.

With 11 plates. By Horace W. Stunkard. $1.25.
No. 4. Color and color-pattern mechanism of Tiger Beetles. With 29 black and 3 colored

plates. By Victor E. Shelford. $2.00.

Vol. IV
No. 1. Life history studies on Montana Trematodes. By E. C. Faust. 9 plates. $2.00.
No. 2. The Goldfish as a test animal in the stydy of toxicity.. By E. B. Powers. $1.00.
No. 3. Morphology and biology of some TurbeUaria from the Mississippi basin. With 3

plates. By Ruth Higley. $1.25.
No. 4. North American pseudophyilidean Cestodes from fishes. With 13 plates. By

A. R. Cooper. $2.00.

Vol. V
No. 1. The Skull of Amiurus. With 8 plates. By J. E. Kindred. ^1.25.
No. 2. Contributions to the Life Histories of Cordhis robicslus I^idy and Pamgordius

varius (Leidy). By Henrj' Gustav May. With 21 plates. $1.50,
Nos. 3 and 4. Studies of M5rsosporidia. A Synopsis of Genera and Species of MyTcosporidia.

By Rokusaburo Kudo. With 25 plates and 2 textfigures. $3.00.

Vol. VI

No. 1.; The Nasal Organ in Amphibia. By G. M. Higgins. With 10 plates. $1,00.

Nos. 2 and 3. Revision of the North American and West Indian Species of Cuscuta. With

13 plates. By Truman George Yuncker. $2.00.
No. 4. The'Larvae of the Coccinellidae. With 6 plates. By J, Howard Gage. 75cenis.

Vol. VII
No. 1. Studies on Gregarines, II. Synopsis of the polycystid Gregarines. 4 plat.

Minnie Watson Kamm. $1.00.
No. 2. The mollusk fauna of the Big Vermilion River, Illinois.. By F. C. Baker, . (In press.)
No. 3. North American Monostomes. By E. C. Harrah. (In press.) '■ '

Entered as second-class matter July 27, 1915, at tlie post-office at Urbana, Illinois, under the Act

of August 24, 1912. Acceptance for mailing at the special rate of postage provided for

in section 1102 Act of October 3, 1917, authorized July 31, 1918.

UNIVERSITY OF ILLINOIS STUDIES IN LANGUAGE AND LITERATURE

The Studies in Language and Literature are designed to include monographs in gen-
eral linguistics and comparative literature; the classical languages and Sanskrit; the Romance
languages; and English, the Scandinavian, and other Germanic languages. The title of the
ssries will be so construed as to admit the publication of such researches in the history of
culture as may throw light on the processes of language and the interpretation of literar
ture. This series is published quarterly; the annual subscription price is three dollars.

' Vol. IV
No. 1. Madame de Stael's literary rfeputation in England. By Robert C. Whitford. 75 eta.
No, 2, 3, 4. Index verborum quae in Senecae fabulis necnon in Octavia Praetexta reperiuntur

a Guilielmo Abbott Oldfather, Arthuro Stanley Pease, Howardo Vernon Canter

confectus. s$i.50. Part IL $1.50. Part IIL $1.50.

VOL. V

No. 1, 2. The influence of Christianity on the Vocabulary of Old English poetry. Part I.

By Albert Kaiser. $1.50.
No. 3. Spenser's Defense of Lord Gray, By H. S. V. Jones. $1.00.
No. 4, Ysopet and Avionnct, Latin and French texts. Kenneth McKenzie and W. A.

Oldfather. $1.50,

Vol. VI
No. 1. La Colleci6n Cervantina de la Sociedad Hispdnica de America (The Hispanic Society

of America) : Ediciones de Don Quijote. By Horaero Seris. $1 . 50.
No. 2, 3. M. Tulli Ciceronis De Divinatione. Liber Primus. With commentary. By A. S.

Pease. $3.00.
No. 4. De Fragmenti suetoniani de grammaticis et rhetoribus; codicum fide et nexu. By

R. P. Robinson. (In press.)

Vol. VII
No. 1. Sir R. Howard's comedy "The Committee." With introduction and notes. By

C. N. Thuxber. $1.50.
No. 2. The Sepulchre of Christ in art and liturgy. By N. C. Brooks. With plates. $1.50
No. 3. The language of the Konungs Skuggsja. By G. T. Flom. (In press.)

UNIVERSITY OF ILLINOIS STUDIES IN THE SOaAL SCIENCES

The University of Illinois Studies in the Social Sciences are designed to afford a means
of publishing monographs prepared by graduate students or members of the faculty in th«
departments of history, economics, political science, and sociology. Each volume will con-
sist of about 600 printed pages annually. The subscription price is three dollars a year.

Vol. II
Ito, 1. Taxation of corporations in Illinois, other than railroads, since 1872. By J. R.

Moore. 55 cts.
Nos, 2ajid3, The West in the diplomatic negotiations of the American Revolution. By P.

C, PhilHps. $1.25.
No, 4. The development of banking in Illinois, 1817-1863, By G. W. Dowrie, 90 cts.

Vol. xll
Nos. 1, and 2, The history of the general property tax in Illinois. By R. M. Haig. $1.25 .

Vol. IV
No. I. The Illinois Whigs before 1846. By C. M. Thompson. 95 cts.
Nos. 3-4. The history of the Illinois Central railway to 1870. By H. G. Brownson. $1.25 ,

Vol. V

No.l. Enforcement of infernatioru. I L'.v by mMnicipiiJ )...-^. ,.., ..QAVright $1.25.
No.2. The life of Jesse W. Fell. By Frances ]vi Mor('.hut'5<?. 60 cts.
No.4, Mine taxation in; the United States. By 1,. '". Voan;^. $1.50.

UNIVERSITY OF ILLINOIS STUDIES IN THE SOCIAL SCIENCES.— Continued

Vol. VI
Nos. 1 and 2. The veto power of the Governor of Illinois. By N. H. Debel. $1 .00.
No. 3. Wage bargaining on the vessels of the Great Lakes. By H. E. Hoagland. $1 .50.
No. 4. The household of a Tudor nobleman. By P. V. B. Jones. $1 .50.

Vol. vn

No. 4. The Journeymen Tailors' Union of America. By C. J. Stowell. $1.00.

Vol. vm

No. 2. Cumtilative voting and minority representation in Illinois. By B. F. Moore. $0.75.
Nos. 3 and 4. Labor problems and administration in the United States diuing the World
War. ByG. Watldns. $2.00.

Vol. IX
No. 1, 2. War powers of the executive in the United States. ByC. A. Berdahl. $2.25.
No. 4. The economic policies of Richelieu. By F. C. Palm. (In press.)

Vol. X
No. 1. Monarchial tendencies in the United States. By Louise B. Dunbar. {In press.)

UNIVERSITY STUDIES
General Series, Vol. I, II, III, and IV
Partly out of print. A detailed list of numbers will be sent on request.

PUBLICATIONS OF THE UNIVERSITY OF ILLINOIS

Following is a partial list of the publications issued at the University:

1. The University of Illinois Stxtdies in Language and Literature. Published
quarterly. Three dollars a year.

2. The Illinois Biological Monographs. Quarterly. Three dollars a year.

3. The UNiVERSiry or Illinois Studies in the Social Sciences. Monographs in his-
tory, economics, political science, and sociology. Quarterly. Three dollars a year.

4. The- University Studies. A series of monographs on miscellaneous subjects.

5. The Journal of English and Germanic Philology. Quarterly. Three dollars
a year. *

(For any of the above, address 161 Administration Building, Urbana, Illinois.)

6. The Bulletin of the Engineering Experiment Station. Reports of the re-
search work in the Engmeering Experiment Station. Address Director of Engineering
Experiment Station, University of Illinois.

7. The Bulletin of the Agricultural Experiment Station. Address Director of
Agricultural Experiment Station, University of Illinois.

8. The Bulletin of the State Laboratory of Natural History. Address Direc-
tor of State Laboratory of Natural History, University of Illinois.

9. The Bulletin of the State Geological Survey. Address Director of State
Geological Survey, University of Illinois.

10. The Bulletin of the State Water Survey. Address Director of State Water
Survey, University of Illinois.

11. The Report of the State Entomologist. Address State Entomologist, Uni-
versity of Illinois.

12. The Bulletin of the Illinois Association of Teachers of English. AddrMs
301 University Hall, University of Illinois.

13. The Bulletin of the School of Education. Address 203 University Hall,
University of Illinois.

14. The Law Bulletin. Address 207 Law Building, University of Ulinoia.

15. The general series, containing the University catalog and circulars of special depart-
ments. Address The Re^strar, University of Illinois.

} ■ 7 ■ -. ' -^

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII April, 1922 No. 2

THE MOLLUSCAN FAUNA OF THE BIG 1

VERMILION RIVER, ILLINOIS \

With Special Reference to its Modification as the Result of
Pollution by Sewage and Manufacturing Wastes

WITH FIFTEEN PLATES

BY

FRANK COLLINS BAKER

Price $1.25

PuBLisHKD BY THE University OF Illinois Press \

under the auspices of the graduate school \

Urbana, Illinois |

UNIVERSITY OF ILLINOIS STUDIES PUBLISHED

ILLINOIS BIOLOGICAL MONOGRAPHS

The Illinois Biological Monoghaphs is a serial published quarterly by the Univer-
sity of Illinois. It is the purpose of the publication to present in monographic form research
contributions of especial merit and of somewhat extended character. Each volume will
include about 500 pages and 20 or more full page plates. The series may be obtained at
a subscription price of three dollars yearly. Individual numbers \vill be sold separately at
prices determined by the size and amount of illustration contained in the particular number.

Vol. I
Nos. 1 and 2. A Revision of the Cestode family Proteocephalidae. With 16 plates. By

G. R. LaRue. $2.00.
No. 3. Studies on the Cestode family Anoplocephalidae. 6 plates. By H. Douthitt. 80 cts.
No. 4. Some North American Larval Trematodes. With 8 plates. By W. W. Cort. $L20,

Vol. II

No. 1. Classification of Lepidopterous larvae. With 10 plates. By S. B. Fracker. $1.50.

No. 2. On the Osteology of some of the Ix)ricati. With 5 plates. By J. E. Gutberlet. 50 cts.

No. 3, Studies on Gregarines, including twenty-one new species and a s3Tiopsis of the
Eugregarine records from the Myriapoda, Coleoptera, and Orthoptera of the
world. With 15 plates. By Minnie E. Watson. $2.00.

No. 4. The 'Genus Meliola in Porto Rico. With 5 plates. By Frank L. Stevens. 75 cts.

Vol. Ill
No. 1. Studies on the factors controlling regeneration. By Charles Meny. $1.25.
No. 2. The head-capsule and mouth-parts of Diptera. 25 plates. By A. Peterson. $2.00.
No. 3. Studies on North American Polystomidae, Aspidogastridae, and Paramphistomidae.

With 11 plates. By Horace W. Stunkard. $1.25.
No. 4. Color and color-pattern mechanism of Tiger Beetles. With 29 black and 3 colored

plates. By Victor E. Shelford. $2.00,

Vol. rv

No. 1. Life history studies on Montana Trematodes. By E. C. Faust. 9 plates. $2.00.
No. 2. The Goldfish as a test animal in the study of toxicity. By E. B. Powers. $1.00.
No. 3. Morphology and biology of some TurbeUaria from the Mississippi basin. With 3

plates. By Ruth Higley. $1.25.
No. 4. North American pseudophyllidean Cestodes from fishes. With 13 plates. By

A. R. Cooper. $2.00.

Vol. V
No. 1. The Skull of Amiurus. With 8 plates. By J. E. Kindred. $1.25.
No. 2. Contributions to the Life Histories of Gordius rohustus Leidy and Paragordius

mriiw (Leidy). By Henry Gustav May. With 21 plates. $1.50.
Nos. 3 and 4. Studies of Myxosporidia. A Sjiiopsis of Genera and Species of Myxosporidia.

By Rokusaburo Kudo. With 25 plates and 2 textfigures. $3.00.

Vol. VI
No. 1. The Nasal Organ in Amphibia. By G. M. Higgins. With 10 plates. $1.00.
Nos. 2 and 3. Revision of the North American and West Indian Species of Cuscuta. With

13 plates. By Truman Georj'.e Yuncker. $2.00.
No. 4. The Larvae of the Coccinellidae. With 6 plates. By J. Howard Gage. 75 cents.

Vol. VII
No. 1. Studies on Gregarines, II. Synopsis of the polycystid Gregarines. 4 plates. By

Minnie Watson Kamm. $1.00.
No. 2. The moUuscan fauna of the Big Vermilion River, Illinois. By F. C. Baker, $1.25.
No. 3. North American Monostomes. By E. C. Harrah. {In press.)

Entered as second-class matter July 27, 1915, at the post-office at Urbana, Illinois, under the Act

of August 24, 1912. Acceptance for mailing at the special rate of postage provided for

in section 1102 Act of October 3, 1917, authorized July 31, 1918.

UNIVERSITY OF ILLINOIS STUDIES IN LANGUAGE AND LITERATURE

The Studies in Language and Literature are designed to include monographs in gen-
eral linguistics and comparative literature; the classical languages and Sanskrit; the Romance
languages; and English, the Scandinavian, and other Germanic languages. The title of the
series will be so construed as to admit the publication of such researches in the history of
culture as may throw light on the processes of language and the interpretation of litera-
ture. This series is published quarterly; the annual subscription price is three dollars.

Vol. IV
No. 1. Madame de Stael's literary reputation in England. By Robert C. Whitford. 75 cti.
No. 2, 3, 4. Index verborum quae in Senecae f abulis necnon in Octavia Praetexta reperiuntur

a Guilielmo Abbott Oldfather, Arthuro Stanley Pease, Howardo Vernon Canter

confectus. $1.50. Part II. $1.50. Part III. $1.50.

VOL. V
No. 1, 2. The influence of Christianity on the Vocabulary of Old English poetry. Part I.

By Albert Keiser. $1.50.
No. 3. Spenser's Defense of Lord Gray. By H. S. V. Jones. SI, 00.
No. 4. Ysopet and Avioimet. Latin and French texts. Kenneth McKenzie and W. A.
Oldfather. $1.50.

Vol. VI
No. 1. La Colleci6n Cervantina de la Sociedad Hispdnica de America (The Hispanic Society

of America) : Ediciones de Don Qui jote. By Homero Seris. $1 . 50.
No. 2, 3. M. Tulli Ciceronis De Divinatione. Liber Primus. With commentary. By A. S.

Pease. $3.00.
No. 4. De Fragmenti suetoniani de grammaticis et rhetoribus; codicum fide et nexu. By
R. P. Robinson. {In press.)

Vol. vn

No. i. Sir R. Howard's comedy "The Committee." With introduction and notes. By

C. N. Thurber. $L50.
No. 2. The Sepulchre of Christ in art and liturgy. By N. C. Brooks. With plates. $1.50.
No. 3. The language of the Konungs Skuggsjd. By G. T. Flom. $1.50.

UNIVERSITY OF ILLINOIS STUDIES IN THE SOCIAL SCIENCES

The University of Illinois Studies in the Social Sciences are designed to afford a means
of publishing monographs prepared by graduate students or members of the faculty In tht
departments of history, economics, political science, and sociology. Each volume will con-
sist of about 600 printed pages annually. The subscription price is three dollars a year.

Vol. II
No. 1. Taxation of corporations in Illinois, other than railroads, since 1872. By J. R.

Moore. 55 cts.
Nos. 2 and 3. The West in the diplomatic negotiations of the American Revolution. By P.

C. Phillips. $1.25.
No. 4. The development of banking in Illmois, 1817-1863. By G. W. Dowrie. 90 cts.

Vol. xll
Nos. 1. and 2. The history of the general property tax in Illinois. By R. M. Haig. $1.25.

Vol. IV
No. 1. The Illmois Whigs before 1846. By C. M. Thompson. 05 cts.
Nos. 3-4. The history of the Illinois Central railway to 1870. By H. G. Brownson. $1.25.

Vol. V
No. 1 . Enforcement of international law by municipal law. By P. Q Wright $1 .25.
No. 2. The life of Jesse W. Fell. By Frances M. Morehouse. 60 cts.
No. 4. Mine taxation in the United States. By L. E. Young. $1 .50.

UNIVERSITY OF ILLINOIS STUDIES IN THE SOCIAL SCIENCES.— Continued

Vol. VI
Nos. 1 and 2. The veto power of the Governor of Illinois. By N. H. Debel. SI. 00.
No. 3. Wage bargaining on the vessels of the Great Lakes. By H. E. Hoagland. $1.50.
No. 4. The household of a Tudor noblem^in. By P. V. B. Jones. $1 .50.

Vol. VII
No. 4. The Journeymen Tailors' Union of America. By C. J. Stowell. $1.00.

Vol. VIII
No. 2. Cumulative voting and minority representation in Illinois. By B. F. Moore. $0.75.
Nos. 3 and 4. Labor problems and administration in the United States during the World
War. By G. Watkins. $2.00.

Vol. IX
No. 1, 2. War powers of the executive in the United Sta:tes. By C. A. Berdahl. $2.25.
No. 4. The economic policies of Richelieu. By F. C. Palm. $1.50.

Vol. X
No, 1. Monarchial tendencies in the United States. By Louise B. Dunbar. (/» press.)
No. 2. Fixed Price Associations. By M. W. Nelson. (In press.)

UNIVERSITY STUDIES
General Series, Vol. I, II, III, and IV
Partly out of print. A detailed list of numbers will be sent on request.

PUBLICATIONS OF THE UNIVERSITY OF ILLINOIS

Followiiag is a partial list of the publications Issued at the University:

1. The Univeksity op Illinois Studies in Language and Literature. Published
quarterly. Three dollars a year.

2. The Illinois Biological Monographs. Quarterly. Three dollars a year.

3. The University of Illinois Studies in the Social Sciences. Monographs in his-
tory, economics, political science, and sociology. . Quarterly. Three dollars a year.

4. The University Studies. A series of monographs on miscellaneous subjects.

5. The Journal of English and Germanic Philology. Quarterly. Three dollars
a year.

(For any of the above, address 161 Administration BuUding, Urbtina, Illinois.)

6. The Bulletin of the Engineering Experiment Station. Reports of the re-
search work in the Engineering Experiment Station. Address Director of Engineering
Experiment Station, University of Illinois.

7. The Bulletin of the Agricultural Experiment Station. Address Director of
Agricultural Experiment Station, University of Illinois.

8. The Bulletin of the State Laboratory of Natural History. Address Direc-
tor' of State Laboratory of Natural Kistory, University of Illinois.

9. The Bulletin of the State Geological Survey. Address Director of State
Geological Survey, University of Illinois. "

10. The Bulletin of the State Water Survey. Address Director of State Water
Survey, University of Illinois.

11. The Report of the State Entomologist. Address State Entomologist, Uni-
versity of Illinois.

12. The Bulletin of the Illinois Association of Teachers of English. Address
301 University Hall, University of Illinois.

13. The Bulletin of the School of Education. Address 203 University Hall,
University of Illinois. ,

14. The Law Bulletin. Address 207 Law Building, University of Illinois.

15. The general series, containing the University catalog and circulars of special depart-
ments. Address The Registrar, University of lUinois.

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII July 1922 No. 3

NORTH AMERICAN MONOSTOMES

PRIMARILY FROM FRESH WATER HOSTS
WITH NINE PLATES

BY 1

1

E. C. HARRAH i

Price $1.25

Published by the UNivERSiTy or Illinois Press ;

under the auspices of the graduate school ^

Urbana, Illinois |

UNIVERSITY OF iLtllSfois STUDIES PUBLISHED

ILLINOIS BIOLOGICAL MONOGRAPHS
The iLtmbis Biological Monographs is a serial published quarteriy by the Univer-
Bity of Illinois. It is the purpose of the publication to present in monographic form research
contributions of especial merit and of somewhat extended character. Each volume wiU
include about 500 pages and 20 or more full page plates. The series may be obtained at
a subscription price of three doUars yearly. Individual numbers will be sold separately at
prices determined by the size and amount of illustration contained in the particular number.

Vol. I
Nos. 1 and 2. A Revision of the Cestode family Proteocephalidae. With 16 plates. By

G. R. La Rue. $2.00.
No. 3. Studies on the Cestode family Anoplocephalidae. 6 plates. By H. Douthitt. 80 cts.
No, 4. Some North American Larval Trematodes. With 8 plates. By W. W. Cort. $1.20.

Vol. II

No. 1. Classification of Lepidopterous larvae. With 10 plates. By S. B. Fracker. $1.50.

No. 2. On the Osteology of some of the Loricati. With 5 plates. By J. E. Gutberlet. 50 cts.

No. 3. Studies on Gregarines, including twenty-one new species and a synopsis of the
Eugregarine records from the Myriapoda, Coleoptera, and Orthoptera of the
world. With 15 plates. By Miimie E. Watson. $2.00.

No. 4. The Genus Meliola in Porto Rico. With 5 plates. By Frank L. Stevens. 75 cts.

Vol. Ill
No. 1. Studies on the factors controlling regeneration. By Charles ZelenV. $1.25.
No. 2. The head-capsule and mouth-parts of Diptera. 25 plates. By A. Peterson. $2.00.
No. 3. Studies on North American Polystomidae, Aspidogastridae, and Paramphistomidae,

With 11 plates. By Horace W. Stunkard. $1.25.
No. 4. Color and color-pattern mechanism of Tiger Beetles. With 29 black and 3 colored

plates. By Victor E. Shelford. $2.00.

Vol. IV
No. 1. Life history studies on Montana Trematodes. By E. C. Faust. 9 plates. $2.00.
No. 2. The Goldfish as a test animal in the study of toxicity. By E, B. Powers. $1.00.
No. 3. Morphology and biology of some Turbellaria from the Mississippi basin. With 3

plates. By Ruth Higley. $1.25.
No. 4. North American pseudoph3'llidean Cestodes from fishes. With 13 plates. By

A. R. Cooper. $2.00.

Vol. V
No. 1. The Skull of Amiurus. With 8 plates. By J. E. Kindred. $1.25.
No. 2. Contributions to the Life Histories of Carditis robuslus Leidy and Paragordius

varius (Leidy). By Henr>' Gustay May. With 21 plates. $1.50.
Nos. 3 aOd 4. Studies of Myxosporidia. A Synopsis of Genera and Species of Myxosporidia.

By Rokusaburo Kudo. With 25 plates and 2 textfigures. $3.00.

Vol. VI
No. 1. The Nasal Organ in Amphibia. By G. M. Higgins. With 10 plates. $1.00.
Nos. 2 and 3. Revision of the North American and West Indian Species of Cuscuta. With

13 plates. By Truman George Yuncker. $2.00.
No. 4. The Larvae of the Coccinellidae. With 6 plates. By J. Howard Gage. 75 cents.

Vol. vn

No. 1. Studies On Gregarines, JI. S3Tiopsis of the polycystid Gregarines. 4 plates. By

Minnie Watson Kamm. $1.00.
No. 2. The molluscan fauna of the Big Vermilion River, Illinois. By F. C. Baker. $1.25.
No. 3. North American Monostomes. By E. C. Harrah. $1,25
No. 4. A Classification of the Larvae of the Tenthredinoidea. ByHachiro Yuasa. (In Press.)

Entered as second-class matter July 27, 191S, at the post-office at Urbana, Illinois, under the Act

of August 24, 1912. Acceptance for mailing at the special rate of postage provided for

in section 1102 Act of October 3, 1917, authorized July 31, 1918.

ITNTVERSITY OF ILLINOIS STUDIES IN LANGUAGE AND LITERATURE

The Studies in Langu^e and Literature are designed to include monographs in gen-
eral linguistics and comparative literature; the classical languages and Sanskrit; the Romance
languages; and English, the Scandinavian, and other Germanic languages. The title of the
series will be so construed as to admit the pubUcation of such researches in the history of
culture aa may throw light on the processes of language and the interpretation of litera-
ture. This series is published quarterly; the annual subscription price is three dollars.

Voi.rv

No. 1. Madame de Stagl's literary reputation in England. By Robert C. Whitford. 75 cts.

No. 2, 3, 4. Index verborum quae in Senecae fabulis necnon in Octavia Praetexta reperiuntur
a Guilielmo Abbott Oldfather, Arthuro Stanley Pease, Howardo Vernon Canter
confectus. $1.50. Part n. $1.50. Part III. $1.50,

VOL. V
No. 1, 2. The influence of Christianity on the Vocabulary of Old English poetry. Part I.

By Albert Keiser. $1.50.
No. 3. Spenser's Defense of Lord Gray. By H. S. V. Jones. $1.00.
No. 4. Ysopet and Avionnet. Latin and French texts. Kenneth McKenzie and W. A.

Oldfather. $1.50.

Vol. VI
No. 1. La Colleci6n Cervantina de la Sociedad Hispinica de America (The Hispanic Society

of America) : Ediciones de Don Quijote. By Homero Seris, $1 . 50.
No. 2, 3. M. Tulli Ciceronis De Divinatione. Liber Primus. With commentary. By A. S.

Pease. $3.00.
No. 4. De Fragmenti suetoniani de grammaticis et rhetoribus; codicum fide et nexu. By

R. P. Robinson. {In press.)

Vol. VII
No. 1. Sir R. Howard's comedy "The Committee." With introduction and notes. By

C. N. Thurber. $1.50.
No. 2. The Sepulchre of Christ in art and liturgy. By N. C. Brooks. With plates. $1.50.
No. 3. The language of the Konungs Skuggsji. By G. T. Flom. $1.50.

UNIVERSITY OF ILLINOIS STUDIES IN THE SOCIAL SCIENCES

The University of Illinois Studies in the Social Sciences are designed to afford a meant
of publishing monographs prepared by graduate students or members of the faculty in tht
departments of history, economics, political science, and sociology. Each volume will con-
sist of about 600 printed pages annually. The subscription price is three dollars a year.

Vol.n
h9o. L. Taxation of corporations in Illinois, other than railroads, since 1872. By J. R>

Moore. 55 cts.
Nos. 2 and 3. The West in the diplomatic negotiations of the American Revolution. By P.

C. Phillips. $1.25.
No. 4. The development of banking in Illinois, 1817-1863. By G. W. Dowrie. 90 cts.

Vol. m
Nob. 1. and 2. The history of the general property tax in Illinois. By R. M. Haig. $1.25 .

Vol. IV
No. !. The. Illinois Whigs before 1846. By C. M. Thompson. 95 cts.
Nos. 3-4, The history of the Illinois Central railway to 1870. By H. G. Brownson. $1.25 .

Vol. V
No. 1 . Enforcement of international law by municipal law. By P. Q. Wright $1 . 25.
No. 2. The life of Jesse W. Fell. By Frances M. Morehouse. 60 cts.
No. 4. Mine taxation in the United States. By L. E. Young. $1 . 50.

UNivERsrry of Illinois studies in the social sciences.— continued

Vol. VI
No6. 1 and 2, The veto power of the Governor of Illinois. By N. H. Debel. $1 .00.
No. 3, Wage bargaining on the vessels of the Great Lakes, By H. E. Hoagland. $1 .50.
No. 4. The household of a Tudor nobleman. By P. V. B. Jones. $1 .50,

Vol. vn

No. 4. The Journeymen Tailors' Union of America. By C. J. Stowell. $1,00.

Vol. vm

No. 2. Cumulative voting and minority representation in Illinois. By B. F, Moore, $0.75.
No«. 3 and 4. Labor problems and administration in the United States during the World
War. ByG. Watkins. $2.00.

Vol. rx

No. 1, 2. War powers of the executive in the United States, By C. A. Berdahl. $2.25.
No. 4. The economic policies of Richelieu. By F. C. Palm. $1.50.

Vol. X
No. 1. Monarchial tendencies in the United States. By Louise B, Dunbar. (In press.)
No. 2. Fixed Price Associations. By M, W. Nelson. (In press.)

UNIVERSITY STUDIES
GisNERAL Series, Vol, I, n, III, and IV
Partly out of print. A detailed list of numbers will be sent on request.

PUBLICATIONS OF THE UNIVERSITY OF ILLINOIS

Following is a partial list of the publications issued at the University:

1. The University of Illinois Studies in Language and Literature, Published
quarterly. Three dollars a year.

2. The Illinois Biological Monographs. Quarterly, Three dollars a year.

3. The University of Illinois Studies in the Social Sciences, Monographs in his-
tory, economics, political science, and sociology. Quarterly, Three dollars a year,

4. The University Studies. A series of monographs on miscellaneous subjects,

5. The Journal of Engush and Germanic Philology. Quarterly. Three dollars
a year.

(For any of the above, address 161 Administration Building, Urbana, Illinois,)

6. The Bulletin of the Engineering Experiment Station. Reports of the re-
search work in the Engineering Experiment Station, Address Director of Engineering
Experiment Station, University of Illinois.

7. The Bulletin of the Agricultural Experiment Station. Address Director of
Agricultural Experiment Station, University of Illinois.

8. The Bulletin of the State Laboratory of Natural History. Address Direc-
tor of State Laboratory of Natural History, University of Illinois.

9. The Bulletin of the State Geological Survey. Address Director of State
Geological Survey, University of Illinois.

10. The Bxtlletin of the State Water Survey, Address Director of State Water
Survey, University of Illinois.

11. The Report of the State Entomologist, Address State Entomologist, Uni-
versity of Illinois.

12. The Bulletin or the Illinois AssoaATioN of Teachers of English, Address
301 University Hall, University of Illinois,

13. The Bulletin of the School of Education. Address 203 University Hall,
University of Illinois.

14. The Law Bulletin. Address 207 Law Building, University of Illinois.

15. The general series, containing the University catalog and circulars of special depart-
ments. Address The Registrar, University of Illinois.

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. VII October, 1922 No. 4

A CLASSIFICATION OF THE LARVAE
OF THE TENTHREDINOIDEA

WITH FOURTEEN PLATES

BY

HACHIRO YUASA

Price $2.00 1

Published by the University of Illinois Press

under the auspices of the graduate school

Urbana, Illinois

UNIVERSITY OF ILLINOIS STUDIES PUBLISHED

ILLINOIS BIOLOGICAL MONOGRAPHS

The IixiKOis Biological Monographs is a serial published quarterly by the Univer-
sity of Illinois. It is the purpose of the publication to present in monographic form research
contributions of especial merit and of somewhat extended character. Each volume will
include about 500 pages and 20 or more full page plates. The series may be obtained at
a subscription price of three dollars yearly. Individual numbers will be sold separately at
prices determined by the size and amount of illustration contained in the particular number.

Vol.1
Nos. 1 and 2. A Revision of the Cestode family Proteocephalidae. With 16 plates. By

G. R. La Rue. $2.00.
No. 3. Studies on the Cestode family Anoplocephalidae. 6 plates. By H. Douthitt. 80 cts.
No. 4. Some North American Larval Trematodes. With 8 plates. ByW.W.Cort. $1.20.

Vol. II

No. 1. Classification of Lepidopterous larvae. With 10 plates. By S. B. Fracker. $1.50.

No. 2. On the Osteology of some of the Loricati. With 5 plates. ByJ.E. Gutberlet. SOcts.

No. 3. Studies on Gregarines, including twenty-one new species and a sjuopsis of the
Eugregarine records from the Myriapoda, Coleoptera, and Orthoptera of the
world. With 15 plates. By Minnie E. Watson. $2.00.

No. 4. The Genus Meliola in Porto Rico. With 5 plates. By Frank L. Stevens. 75 cts.

Vol. Ill
No. 1. Studies on the factors controlling regeneration. By Charles Zeleny. $1.25.
No. 2. The head-capsule and mouth-parts of Diptera. 25 plates. By A. Peterson. $2.00.
No. 3. Studies on North American Polystomidae, Aspidogastridae, and Paramphistomidae.

With 11 plates. By Horace W. Stunkard. $1.25.
No. 4. Color and color-pattern mechanism of Tiger Beetles. With 29 black and 3 colored

plates. By Victor E. Shelford. $2.00.

Vol. IV
No. 1. Life history studies on Montana Trematodes. By E. C. Faust. 9 plates. $2.00.
No. 2. The Goldfish as a test animal in the study of toxicity. By E. B. Powers. $1.00.
No. 3. Morphology and biology of some Turbellaria from the Mississippi basin. With 3

plates. By Ruth Higley. $1.25.
No. 4. North American pseudophyllidean Cestodes from fishes. With 13 plates. By

A. R. Cooper. $2.00.

VolV
No. 1. The Skull of Amiurus. With 8 plates. By J. E. Kindred. $1.25.
No. 2. Contributions to the Life Histories of Cordius robtistus Leidy and Paragordius

varins (Leidy). By Henrj' Gustav May. With 21 plates. $1.50.
Nos. 3 and 4. Studies of Myxosporidia. A Synopsis of Genera and Species of Myxosporidia.

By Rokusaburo Kudo. With 25 plates and 2 textfigures. $3.00.

Vol. VI
No. 1. The Nasal Organ in Amphibia. By G. M. Higgins. With 10 plates. $1.00.
Nos. 2 and 3. Revision of the North American and West Indian Species of Cuscuta. With

13 plates. By Truman George Yuncker. $2.00.
No. 4. The Larvae of the Coccinellidae. With 6 plates. By J. Howard Gage. 75 cents.

Vol. vn

No. 1. Studies on Gregarines, II. Synopsis of the polycystid Gregarines. With 4 plates. By

Minnie Watson Kamm. $1.00.
No. 2. The molluscan fauna of the Big Vermilion River, Illinois. With 15 plates. By F. C.

Baker. $1.25.
No. 3. North American Monostomes. With 9 plates. By E. C. Harrah. $1.25.
No. 4. A Classification of the Larvae of the Tenthredinoidea. With 14 plates. By Hachiro

Yuasa. $2.00.

Entered ss second-class matter July 27, 1915, at the post-office at Urbana, Illinois, under the Act

of August 24, 1912. Acceptance for mailing at the special rate of postage provided for

in section 1102 Act of October 3. 1917, authorized July 31, 1918.

UNIVERSITY OF ILLINOIS STUDIES IN LANGUAGE AND LITERATURE

The Studies in Language and Literature are designed to include monographs in gen-
eral linguistics and comparative literature; the classical languages and Sanskrit; the Romance
languages; and English, the Scandinavian, and other Germanic languages. The title of the
series will be so construed as to admit the publication of such researches in the history of
culture as may throw light on the processes of language and the interpretation of litera-
ture. This series Is published qiiarterly; the aimual subscription price is three dollars.

Vol. IV
No. 1. Madame de Stall's literary reputation m England. By Robert C. Whitford. 75 cts.
No. 2, 3, 4. Index verborum quae in Senecae fabulis necnon in Octavia Praetexta reperiuntur
a Guilielmo Abbott Oldfather, Arthuro Stanley Pease, Howardo Vernon Canter
confectus. $1.50. Part U. $1.50. Part IIL $1.50.

Vol. V
No. 1, 2. The influence of Christianity on the Vocabulary of Old EngL'sh poetry. Part I.

By Albert Keiser. $1.50.
No. 3. Spenser's Defense of Lord Gray. By H. S. V. Jones. $1 .00.
No. 4. Ysopet and Avionnet. Latin and French texts. Kenneth McKenrie and W. A.

Oldfather. $1.50.

Vol. VI
No. 1. La Colleci6n Cervantina de la Sociedad Hispdnica de America (The Hispanic Society

of America) : Ediciones de Don Quijote. By Homero Sens. $1 . 50.
No. 2, 3. M. Tulli Ciceronis De Divinatione. Liber Primus. With commentary. By A, S.

Pease. $3.00.
No. 4. De Fragment! suetoniani de grammaticis et rhetoribus; codlcum fide et nexu. By

R. P. Robinson. $2.00.

Vol. vn

No. L Sir R. Howard's comedy "The Committee." With introduction and notes. By

C. N. Thurber. $1.50.
No. 2. The Sepulchre of Christ in art and liturgy. By N. C. Brooks. With plates. $1.50.
No. 3. The language of the Konungs SkuggsjA. By G. T. Flom. $1.50.
No. 4. The significant name in Terence. By J. C. Austin. $2.00.

UNIVERSITY OF ILLINOIS STUDIES IN THE SOCIAL SCIENCES

The University of Illinois Studies in the Social Sciences are designed to afford & means
of pnblishing monographs prepared by graduate students or members of the faculty In the
departments of history, economics, political science, and sociology. Each volume will con-
sist of about 600 printed pages annually. The subscription price is three dollars a year.

VoLH
No. 1. Taxation of corporations in Illinois, other than railroads, since 1872. By J. R.

Moore. 55 cts.
Koi. 2 and 3. The West in the diplomatic negotiations of the American Revolution. By P.

C. PhilUps. $1.25.
No. 4. The development of banking in Illinois, 1817-1863. By G. W. Dowrie. 90 cts.

Vol. m
Nos. 1. and 2. The history of the general property tax in Illinois. By R. M. Haig. $1.25.

Vol. IV
No. 1. The Illinois Whigs before 1846. By C. M. Thompson. 95 cts.
No*. 3-4. The history of the Illinois Central railway to 1870. By H. G. Brownson. $1.25.

Vol. V
No. 1. Enforcement of international law by municipal law. By P. Q. Wright. $1 .25.
No. 2. The life of Jesse W. Fell. By Frances M. Morehouse. 60 cts.
No. 4. Mine taxation in the United States. By L. E. Young. $1 .50.

UNIVERSriY OF IIXINOIS STUDIES IN THE SOCIAL SCIENCES.— Continued

Vol. VI
No«. 1 and 2. The veto power of the Governor of IllinoU. By N. H. Dcbel $1 .00.
No. 3. Wage bargaining on the vessels of the Great Lakes. By 11. E. Hoagland. $1 . 50.
No. 4. The household of a Tudor nobleman. By P. V. B. Jones. $1 .50.

Vol. VU
No. 4. The Journeymen Tailors' Union of America. By C. J. StowelL $1.00.

Vol. VIII
No. 2. Cumulatire voting and minority representation in Illinois. By B. F. Moore. $0.75.
Not. 3 and 4. Labor problems and administration in the United States during the World
War. ByG. WatWns. $2.00.

Vol. DC
Na 1, 2. War powers of the executive in the United States. By C. A. Berdahl. $2.25.
No. 4. The economic policies of Richelieu. By F. C. Palm. $1.50.

Vol. X
No. 1. Monarchial tendencies in the United States. By Louise B. Dunbar. (In press.)
No. 2. Fixed Price Associations. By M. W. Nelson. (/» press.)

UNIVERSITY STUDIES
General Series, Vol. I, II, III, and IV
Partly out of print. A detailed list of numbers will be sent on request.

PUBLICATIONS OF THE UNIVERSITY OF ILLINOIS

Following is a partial list of the publications issued at the University:

1. The University of Illinois Studies in Language and Literature. Published
quarterly. Three dollars a year.

2. The Illinois Biological Monographs. Quarterly. Three dollars a year.

3. The University of Illinois Studies in the Social Sciences. Monographs in hia-
tory, economics, political science, and sociology. Quarterly. Three dollars a year.

4. The University Studies. A series of monographs on miscellaneous subjects.

5. The Journal of English and Germanic Philology. Quarterly. Three dollars
a year.

(For any of the above, address 161 Administration Building, Urbana, Illinois.)

6. The Bulletin of the Engineering Experiment Station. Reports of the re-
search work in the Engineering Experiment Station. Address Director of Engineering
Experiment Station, University of Illinois.

7. The Bulletin of the Agricultural Experiment Station. Address Director of
Agricultural Experiment Station, University of Illinois.

8. The Bulletin of the State Laboratory of Natural History. Address Direc-
tor of State Laboratory of Natural History, University of Illinois.

9. The Bulletin of the State Geological Survey. Address Director of State
Geological Survey, University of Illinois.

10. The Bulletin of the! State Water Survey. Address Director of State Water
Survey, University of Illinois.

11. The Report of the State Entomologist. Address State Entomologist, Uni-
versity of Illinois.

12. The Bulletin of the Illinois Association of Teachers of English. Address
301 University Hall, University of Illinois.

13. The Bulletin of the School of Education. Address 203 University Hall,
University of Dlinois.

14. The Law Bulletin. Address 207 Law Building, University of Illinois.

15. The general scries, containing the University catalog and circulars of special depart-
ments. Address The Registrar, University of Illinois.

•'K

MBl, WHOI UBRARV

^JH lASB U

■; !■»;;,' \:i:\ v.: ■■;v.

'l.i