The Classification of Lower Organisms
Ernst Hkinrich Haickei, in 1874
From Rolschc (1906).
By permission of Macrae Smith Company.
C f 3
The Classification
of
LOWER ORGANISMS
By
HERBERT FAULKNER COPELAND
\
PACIFIC ^.,^,kfi^..^ BOOKS
PALO ALTO, CALIFORNIA
Copyright 1956 by Herbert F. Copeland
Library of Congress Catalog Card Number 56-7944
Published by
PACIFIC BOOKS
Palo Alto, California
Printed and bound in the United States of America
CONTENTS
Chapter Page
I. Introduction 1
II. An Essay on Nomenclature 6
III. Kingdom Mychota 12
Phylum Archezoa 17
Class 1. Schizophyta 18
Order 1. Schizosporea 18
Order 2. Actinomycetalea 24
Order 3. Caulobacterialea 25
Class 2. Myxoschizomycetes 27
Order 1. Myxobactralea 27
Order 2. Spirochaetalea 28
Class 3. Archiplastidea 29
Order 1. Rhodobacteria 31
Order 2. Sphaerotilalea 33
Order 3. Coccogonea 33
Order 4. Gloiophycea 33
IV. Kingdom Protoctista 37
V. Phylum Rhodophyta 40
Class 1. Bangialea 41
Order Bangiacea 41
Class 2. Heterocarpea 44
Order 1. Cryptospermea 47
Order 2. Sphaerococcoidea 47
Order 3. Gelidialea 49
Order 4. Furccllariea 50
Order 5. Coeloblastea 51
Order 6. Floridea 51
VI. Phylum Phaeophyta 53
Class 1. Heterokonta 55
Order 1. Ochromonadalea 57
Order 2. Silicoflagellata 61
Order 3. Vaucheriacea 63
Order 4. Choanoflagellata 67
Order 5. Hyphochytrialea 69
Class 2. Bacillariacea 69
Order 1. Disciformia 73
Order 2. Diatomea 74
Class 3. Oomycetes 76
Order 1. Saprolegnina 77
Order 2. Peronosporina 80
Order 3. Lagenidialea 81
Class 4. Melanophycea 82
Order 1 . Phaeozoosporea 86
Order 2. Sphacelarialea 86
Order 3. Dictyotea 86
Order 4. Sporochnoidea 87
V ly
Chapter Page
Orders. Cutlerialea 88
Order 6. Laminariea 89
Order 7. Fucoidea 91
VII. Phylum Pyrrhophyta 94
Class Mastigophora 95
Order 1. Cryptomonadalea 96
Order 2. Adiniferidea 98
Order 3. Cystoflagellata 99
Order 4. Cilioflagellata 102
Order 5. Astoma 105
VIII. Phylum Opisthokonta 110
Class Archimycetes Ill
Order 1. Monoblepharidalea Ill
Order 2. Chytridinea 113
IX. Phylum Inophyta 119
Class 1. Zygomycetes 121
Order 1. Mucorina 121
Order 2. Entomophthorinea 124
Class 2. Ascomycetes 125
Order 1. Endomycetalea 129
Order 2. Mucedines 130
Order 3. Perisporiacea 131
Order 4. Phacidialea 133
Order 5. Cupulata 134
Order 6. Exoascalea 137
Order 7. Sclerocarpa 137
Order 8. Laboulbenialea 140
Class 3. Hyphomycetes 140
Order 1. Phomatalea .... 141
Order 2. Melanconialea 141
Order 3. Nematothecia 141
Class 4. Basidiomycetes 142
Order 1. Protobasidiomycetes 146
Order 2. Hypodermia 147
Order 3. Ustilaginea 149
Order 4. Tremcllina 149
Order 5. Dacryomycetalea 150
Order 6. Fungi 150
Order 7. Dermatocarpa 152
X. Phylum Protoplasta 157
Class 1. Zoomastigoda 157
Order 1. Rhizoflagellata 158
Order 2. Polymastigida 163
Order 3. Trichomonadina 166
Order 4. Hypcrmastiglna 168
Class 2. Mycetozoa 171
Order 1. Enteridiea 171
Order 2. Exosporea 177
vi
Chapter Page
Order 3. Phytomyxida 177
Class 3. Rhizopoda 179
Order 1. Monosomatia 183
Order 2. Miliolidea 185
Order 3. Foraminifera . . . 185
Order 4. Globigerinidea 187
Order 5. Nummulidnidea 188
Class 4. Heliozoa 189
Order 1. Radioflagellata 190
Order 2. Radiolaria 194
Order 3. Acantharia 195
Order 4. Monopylaria 198
Orders. Phaeosphaeria 198
Class 5. Sarkodina 200
Order 1. Nuda 201
Order 2. Lampramoebae 205
XI. Phylum Fungilli 206
Class 1. Sporozoa 207
Order 1. Oligosporea 209
Order 2. Polysporea 211
Order 3. Gymnosporidiida 211
Order 4. Dolichocystida 214
Orders. Schizogregarinida 215
Order 6. Monocystidea 215
Order 7. Polycystidea 216
Order 8. Haplosporidiidea 218
Class 2. Neosporidia 219
Order 1. Phaenocystes 219
Order 2. Actinomyxida 221
Order 3. Cryptocystes 222
XII. Phylum Ciliophora 223
Class 1. Infusoria 228
Order 1. Opalinalea 228
Order 2. Holotricha 229
Order 3. Heterotricha 230
Order 4. Hypotricha 233
Order 5. Stomatoda 233
Class 2. Tentaculifera 235
Order Suctoria 235
List of Nomenclatural Novelties 237
Bibliography 238
Index 271
VII
LIST OF ILLUSTRATIONS
Portrait of Ernst Heinrich Haeckel Frontispiece
Figure Page
1. Structure of cells of blue-green algae 13
2. Photographs of Escherichia coli . . . 15
3. Caulobacterialea; Myxobactralea; Cristispira Veneris 26
4. Coccogonea; Gloiophycea 32
5. Bangialea 42
6. Nuclear phenomena in Polysiphonia violacea 45
7. Heterocarpea 48
8. Ochromonadalea 54
9. Ochromonadalea; Silicoflagellata 56
10. Vaucheriacea 64
11. Choanoflagellata 68
12. Hyphochytrialea 70
13. Bacillariacea 72
14. Oomycetes 78
15. Stages of nuclear division in Stypocaulon 84
16. Familiar kelps of Pacific North America 90
17. Microscopic reproductive structures of Laminaria yezoensis ... 92
18. Cryptomonadalea 97
19. Cystoflagellata; Cilioflagellata 104
20. Astoma 106
21. Astoma 108
22. Monoblepharidalea 114
23. Chytridinea 116
24. Zygomycetes 122
25. Ascomycetes 132
26. Ascomycetes 136
27. Mycosphaerella personata 138
28. Basidiomycetes 144
29. Fruits of Agaricacea 153
30. Rhizoflagellata 160
31. Polymastigida; Trichomonadina 164
32. Hypermastigina 170
33. Mycetozoa 176
34. Ceratiomyxafruticulosa 178
35. Life cycle of "Tretomphalus" i. e., Discorbis or Cymbalo por a . . . 180
36. Shells of Rhizopoda 184
37. Radioflagellata 192
38. Radiolaria; Acantharia; Monopylaria; Phaeosphaeria 196
39. Chaos Protheus 200
40. Sarkodina 204
41. Life cycle of Goussia Schuhergi 208
42. LUe cycle of Plasmodium; Babesia bigemina 212
43. Life cycle of Myxoceros Blennius 220
44. Infusoria, order Hypotricha 232
45. Tokophrya Lemnarum 234
ix
Chapter I
INTRODUCTION
The purpose of this work is to persuade the community of biologists that the ac-
cepted primary classification of living things as two kingdoms, plants and animals,
should be abandoned; that the kingdoms of plants and animals are to be given definite
limits, and that the organisms excluded from them are to be organized as two other
kingdoms. The names of the additional kingdoms, as fixed by generally accepted
principles of nomenclature, appear to be respectively Mychota and Protoctista.
These ideas originated, so far as I am concerned, in the instruction of Edwin
Bingham Copeland, my father, who, when I was scarcely of high school age, admitted
me to his college course in elementary botany. He thought it right to teach freshmen
the fundamental principles of classification. These include the following:
The kinds of organisms constitute a system of groups; the groups and the system
exist in nature, and are to be discovered by man, not devised or constructed. The
system is of a definite and peculiar pattern. By every feature of this pattern, we are
inductively convinced that the kinds of organisms, the groups, and the system are
products of evolution. It is this system that is properly designated the natural system
or the natural classification of organisms. It is only by metaphor or ellipsis that these
terms can be applied to systems formulated by men and published in books.
Men have developed a classification of organisms which may be called the taxo-
nomic system. Its function — the purpose for which men have constructed it — is to
serve as an index to all that is known about organisms. This system is subject to cer-
tain conventions which experience has shown to be expedient. Among natural groups,
there are intergradations; taxonomic groups are conceived as sharply limited. Natural
groups are not of definite grades; taxonomic groups are assigned to grades. When we
say that Pisces and Filicineae are classes, we are expressing a fact of human conven-
ience, not a fact of nature. The names assigned to groups are obviously conventional.
Since the taxonomic system represents knowledge, and since knowledge is ad-
vancing, this system is inherently subject to change. It is the right and duty of every
person who thinks that the taxonomic system can be improved to propose to change
it. A salutary convention requires that proposals in taxonomy be unequivocal: one
proposes a change by publishing it as in effect; it comes actually into effect in the
degree that the generality of students of classification accept it. The changes which
are accepted are those which appear to make the taxonomic system, within its conven-
tions, a better representation of the natural system. Different presentations of the
taxonomic system are related to the natural system as pictures of a tree, by artists of
different degrees of skill or of different schools, are related to the actual tree; the
taxonomic system is a conventionalized representation of the natural system so far as
the natural system is known.
These statements are intended to make several points. First, as a personal matter,
advancement of knowledge of natural classification, and corresponding improvement
of the taxonomic system, have been my purpose during the greater part of a normal
lifetime. Secondly, I have pursued this purpose, and continue to pursue it, under the
guidance of principles which all students of classification will accept (perhaps with
variations in the words in which they are stated). In the third place, I have tried to
answer the question which scientists other than students of classification, and likewise
the laity, are always asking us: why can one not leave accepted classification undis-
2 ] The Classification of Lower Organisms
turbed? One proposes changes in order to express what one supposes to be improved
knowledge of the kinds of organisms which belong together as facts of nature. If here
I place bacteria in a different kingdom from plants, and Infusoria in a different king-
dom from animals, it is because I believe that everyone will have a better understand-
ing of each of these four groups if he does not think of any two of them as belonging
to the same kingdom.
The course of evolution believed to have produced those features of the natural
system to which the present work gives taxonomic expression is next to be described.
Life originated on this earth, by natural processes, under conditions other than
those of the present, once only. These are the opinions of Oparin ( 1938) 1, and appear
sound, although some of the details which he suggested may not be. When the crust
of the earth first became cool, it was covered by an atmosphere of ammonia, water
vapor, and methane, and by an ocean containing the gases in the atmosphere above
it and minerals dissolved from the crust. This is to state the hypotheses that organic
matter in the form of methane is older than life; and that whereas conditions on the
face of the earth tend now to cause oxidation, they tended originally to cause reduc-
tion. In a medium of the nature of the supposed primitive ocean, spontanous chemical
changes will occur and produce organic compounds of considerable complexity: this
has repeatedly been demonstrated by experiment. To convert a solution of ammonia,
methane, and minerals into protoplasm, Oparin postulates a very long series of
changes, producing successively more complicated compounds and mixtures, and re-
quiring perhaps hundreds of millions of years. The changes are conceived as acci-
dents; they are supposed to have been probable accidents, like throwing a seven at dice,
not events which could only very rarely occur by accident, like throwing twenty sevens
in succession. By supposing that some of these processes used up the m.aterials neces-
sary for them, Oparin provides an explanation of the single origin of life: we are
confident that all life is of one origin, because all protoplasm is of the same general
nature, and all life consists of essentially the same processes. The course of events
described would have yielded, as the original form of life, anaerobic saprophytes; this
is in harmony with the fact that anaerobic energesis is in a sense the basic metabolic
process. The original organisms would scarcely have possessed nuclei: Oparin's
theories indicate, as the most primitive form of life which has been able to survive,
the anaerobic bacteria. The anaerobic bacteria are indeed very far removed from any
lifeless things; their protoplasm and their metabolism are fundamentally the same
as ours.
Life requires energy. Under anaerobic conditions, an organism can obtain energy
by converting sugars to alcohol, but it can not use alcohol as a source of energy. This
example means that anaerobic energesis yields energy in strictly limited quantity and
produces incompletely oxidized compounds. So long as all life was anaerobic, it was
engaged in converting the organic matter upon which it depended into forms which
it could not use; life under these conditions, at least if they persisted for any great
period of time, was surely very sluggish. A further scries of changes in the metabolic
system, occurring accidentally in certain organisms and preserved by natural selec-
tion, brought photosynthesis into existence. The purple bacteria are believed to rep-
resent stages in the evolution of photosynthesis, which exists in its fully developed
form, involving the release of elemental oxygen, in the blue-green algae. Once photo-
^ Dates in parentheses are references to works which have been consulted and listed in
the bibliography.
Introduction [ 3
synthesis was established in certain organisms, aerobic energesis became possible both
to these and to others. This made possible a manner of life more vigorously active
than before. The inconsiderable groups of autotrophic bacteria — the organisms which
live by oxidizing inorganic matter — appear to be secondary developments dependent
upon the existence of photosynthesis.
The organisms whose origin has been suggested thus far — the ordinary bacteria,
anaerobic and aerobic, the autotrophic bacteria, the purple bacteria, and the blue-
green algae — are relatively simple in structure and function; all consist of minute
physiologically independent cells. The first step in the evolution of more complex
organisms was the evolution of the nucleus.
Morphologically, the nucleus is a part of a protoplast which is set apart by a mem-
brane and which originates ordinarily by division of a pre-existent nucleus in the
manner called mitosis. In this process, a definite number of definite chromosomes
appear and undergo equal division. The nucleus exercises control over the protoplast
in which it lies. Its controlling action depends upon the chromosomes which go into
it, and mitosis has the effect that all nuclei which are derived from one original nu-
cleus strictly by normal processes of mitosis are identical in the controlling effects
which they exert. Thus the nucleus serves for the precise transmission of a compli-
cated heredity. Beside mitosis, there are two other processes — two only — meiosis and
karyogamy, by which nuclei may produce other normal and enduringly viable nuclei.
In a sequence of generations of individuals sexually produced, these processes occur
alternately, each one at one point in each cycle of sexual i-eproductlon. Mendelian
heredity is produced by changes, in the sets of chromosomes (or parts of chromo-
somes) in individual nuclei, which occur during meiosis and karyogamy. The role of
the nucleus in sexual reproduction is one of its essential characters: the nucleus is re-
lated to sexual reproduction, including Mendelian heredity, as structure to function.
The existence of organisms without nuclei shows that the nucleus evolved after life
did: it did not evolve at the same time as protoplasm. The essential uniformity of
the nucleus and of its association with sexual reproduction shows that these things
evolved only once, and together. There are a very few organisms, as Porphyridium
and Prasiola, in which the presence or absence of nuclei is not certain; there is ac-
cordingly scant evidence for speculation as to the manner of this evolution. As to the
tim.e, we know only that microfossils representing nucleate organisms occur in the
uppermost strata of the Proterozoic era.
By making possible the precise transmission of a complicated heredity, the nucleus
has made possible the development of complexities of structure and function exceed-
ing by far anything occurring in non-nucleate organisms. It appears that as soon as
the nucleus was in existence, organisms provided with it entered upon evolution in
many characters and gave rise to many distinguishable groups. Among these groups,
those which consist respectively of the typical plants and the typical animals are the
greatest. There is, however, neither any a priori reason, nor any evidence from nature,
for a belief that all groups of nucleate organisms must naturally belong to one or the
other of these two. Several other groups, in general much less considerable than these,
are thoroughly distinct and appear equally ancient.
E. B. Copeland understood the history of life very much as it has just been pre-
sented. In his teaching, he treated the bacteria and blue-green algae as standing en-
tirely apart both from plants and from animals, and pointed out several other groups
which are not as a matter of nature either plants or animals. It was his opinion that
these groups should be treated as a series of minor kingdoms; he excused himself
4 ] The Classification of Lower Organisms
from the attempt to formulate a definite and comprehensive system. This teaching
was the original stimulus which has led to the present work. I bear witness that E. B.
Copeland taught these things in 1914; he did not publish them until he had ceased
to teach (1927).
In the year 1926, when the teaching of elementary botany was first fully my own
responsibility, I came to the conclusion that the establishment of several kingdoms
of nucleate organisms in addition to plants and animals is not feasible; that all of
these organisms are to be treated as one kingdom. This is one of the few points of
originality which I claim for my work. It is true that the kingdom thus described is
not very different from the third kingdom which various early authors proposed and
which Haeckel (1866) named Protista. Haeckel, however, in his varied presentations
of the kingdom Protista, included always the bacteria. By setting apart the bacteria
and blue-green algae as yet another kingdom, one meets, at least in part, the objection
to the "third kingdom" that it is heterogeneous beyond what can be tolerated.
It has been necessary to meet also the objection that the "third kingdom" substi-
tutes, for an acknowledgedly vague boundary between plants and animals, two vague
boundaries: it has been necessary to recognize characters by which sharp definition
can be given to plants and animals. It is my contention that these characters have
long been known. The kingdom of plants, as the taxonomic representation of a
natural group, is to be defined by the system of chloroplast pigments described by
Willstatter and Stoll (1913), and also by the production of certain carbohydrates
which occur only sporadically elsewhere. The kingdom of animals is defined by em-
bryonic development through the stages called blastula and gastrula, as pointed out
by Haeckel (1872). It is believed that no organisms exhibit both of these sets of
characters; the "third kingdom" includes the nucleate organisms which exhibit
neither. The kingdoms of plants and animals as here defined are essentially those
which are traditionally and popularly accepted. They include all the creatures which
Linnaeus listed as plants and animals, with the exceptions of forms of which he knew
little, and which he listed superficially at the ends of his treatments of the respective
kingdoms.
Of course, the definitions are not warranted to describe the kingdoms without ex-
ception. For one thing, each is supposed to have come into existence by evolution
through a line of organisms which exhibited its characters imperfectly. For another,
evolution can erase what it has created; it is proper to include in a group organisms
which have by degeneration lost its formal characters. These things are true of all
taxonomic groups.
In due form, then, the system of kingdoms here maintained is as follows:
Kingdom I. Mychota. Organisms without nuclei; the bacteria and blue-green
algae.
Kingdom II. Protoctista. Nucleate organisms not of the characters of plants and
animals; the protozoa, the red and brown algae, and the fungi.
Kingdom III. Plantae. Organisms in whose cells occur chloroplasts, being plastids
of a bright green color, containing the pigments chlorophyll a, chlorophyll h, carotin,
and xanthophyll, and no others; and which produce sucrose, true starch, and true
cellulose.
Kingdom IV. Animalia. Multicellular organisms which pass during development
through the stages called blastula and gastrula; typically predatory, and accordingly
consisting of unwalled cells and attaining high complexity of structure and function.
This system has twice been given brief publication (1938, 1947). I am glad to say
Introduction [ 5
that Barkley (1939, 1949) and Rothmaler (1948) maintain a system of kingdoms
which differs from this in a single significant detail.
Assuming that this system is tenable as a matter of reason, it will nevertheless not
be accepted among taxonomists unless they have some knowledge of what it means
in detail. No person is called upon to recognize the kingdoms Mychota and Protoc-
tista until systems of their subordinate groups are available. The bulk of the present
work consists of such systems. Complete systems of divisions or phyla, classes, and
orders are presented. Groups of lower rank are presented in part, as examples. As a
matter of facility, the groups of lower rank are presented more fully in the smaller or
better known groups than in the larger or more obscure.
The preparation of this work has taken more than ten years. In the course of it I
have received much help. Among those who have answered queries, or who have in
various drafts scrutinized the whole work or parts of it for faults of every degree of
significance, are Dr. G. M. Smith of Stanford University; Dr. A. S. Campbell of St.
Mary's College; Dr. Herbert Graham, formerly of Mills College; Dr. Lee Bonar, Dr.
G. L. Papenfuss, and Dr. H. L. Mason of the University of California at Berkeley;
Dr. E. R. Noble of the University of California at Santa Barbara; and Dr. H. C. Day
of Sacramento Junior College. The counsel of E. B. Copeland has not been withheld.
It is a matter of grief that two distinguished zoologists of the University of California,
Dr. S. F. Light and Dr. Harold Kirby, have passed away during the long course of
this work; as have two colleagues who were my closest friends, Dr. H. J. Child and
Dr. C. C. Wright.
The portrait of Haeckel which is my frontispiece is used by permission of Macrae
Smith Company, Philadelphia. Two figures of Chrysocapsa are used by permission
of the Cambridge University Press. Numerous figures have been taken from the
Archiv filr Protistenkunde with the gracious permission of Prof. Dr. Max Hartmann.
We do well to realize our indebtedness to libraries and librarians. To a great extent,
this work has been made possible by the unstinted hospitality of the Biology Library
of the University of California at Berkeley.
Two statements appear regularly in prefaces; they are of truths which are strongly
impressed upon authors. In the first place, those who have given help have made the
work better; the author alone is responsible for deficiencies. The foregoing list of
good friends and good scholars does not claim them as proponents of the thesis of
this work.
In the second place, the work is not offered as perfect or nearly so. The scholar in
a strictly limited field may become master of the available knowledge. One who at-
tempts studies in a broad field realizes that he is dealing with many subjects of which
others know far more than he; that he has not wrung dry the existing literature; that
some of the problems which puzzle him will be solved if he will wait a little longer.
His colleagues have a right to raise these matters as criticisms. But surely, it is not
desired that studies in broad fields be never attempted or indefinitely delayed.
A matter which is particularly likely to arouse criticism is that of the names which
are here applied to the groups. The principles according to which this has been done
are set forth in the following chapter. I beg my colleagues, in dealing with this chapter
and with the names subsequently applied, not to imagine that I have acted without
grave thought. I have decided, that as in classification, so also in nomenclature, I
should set before the community of biologists an experiment in the application of
principles; among which principles there are surely some whose strict application
will be to the good of our science.
Chapter II
AN ESSAY ON NOMENCLATURE
Whoever sets forth a system of groups finds himself under the necessity of making
responsible decisions as to names. The kingdoms have received more names than one
(Table 1 ), and so have nearly all of the major groups within them: it has here been
necessary to decide as to the validity and application of the names Flagellata and
Mastigophora, Rhodophyceae and Florideae, Rhizopoda and Sarcodina, and many
others.
TABLE 1. Names Applied by Various Authors to the Kingdoms
OF Systems of Four Kingdoms
Authors
Kingdoms
Copeland,
1938, and
Rothmaler,
Copeland,
Haeckel, 1894
Barkley, 1939
1948 1947 and here
I Protophyta
Monera
Anucleobionta
Mychota
II Protozoa
Protista
Protobionta
Protoctista
III Metaphyta
Plantae
Cormobionta
Plantae
IV Metazoa
Animalia
Gastrobionta
Animalia
In dealing with plants, with animals, or with bacteria, it is necessary to observe
the codes of nomenclature enacted by international congresses for the respective
groups: the botanical code (Fournier, 1867; Lanjouw, 1952), with amendments
enacted in 1954; the zoological code of 1889 as amended in 1948 and 1953 (issue of
an edition incorporating the amendments is expected; Hemming, 1954); and the
bacteriological code (Buchanan et al., 1948). Breach of the appropriate code renders
an author liable to the penalty of having his work treated as nullity.
The existence of three sets of rules for one thing, and the continual amendment of
the older codes, are evidence of imperfection. It will not be purely destructive to
point out certain anomalies in the codes as they stand.
The zoological code pretends to overrule the principles of grammar in treating
specific epithets as names. It is true that some of these words are names: the Catus in
Felis Catus is a name of the cat, and the Mays in Zea Mays is a name of maize. But
the great majority are adjectives; the sapiens in Homo sapiens is not by itself a des-
ignation of man, and the vulgarc in Hordeum vulgarc is not a name of barley. It is a
further offense against grammar that the code prescribes, as the names of all families
of animals, adjectives in the feminine. Applied originally to families of birds, Aves,
these names were unobjectionable; but the names of the kingdom and of the over-
whelming majority of its subordinate groups are neuter.
The botanical code as published with its appendages makes a book of more than
two hundred pages. A statement of principles, in which the last clause provides for
exceptions, occupies two pages. The definite rules and recommendations occupy
about thirty-five pages; one who studies them critically will find that they prescribe
more than one procedure not warranted by principle. A list of names maintained or
rejected irrespective of principle occupies about seventy pages. These things mean
that current botanical nomenclature is only within limits a matter of rule; it is to a
considerable extent governed by enactments of the nature of ex post facto laws and
bills of attainder.
An Essay on Nomenclature [ 7
The bacteriological code is for the most part a condensation of an earlier edition
of the botanical code. It includes the odd feature that the name of a genus of bacteria
is to be changed if it had previously been used either among plants or among Protozoa.
Since there is an earlier Phytomonas among flagellates, bacteriologists have given a
new name to the bacterium Phytomonas. The avoidance of homonyms which they
desire will not, however, be attained: no zoologist will allow a new name for the
flagellate Klebsiella on account of an earlier Klebsiella among bacteria.
The grounds upon which these things are treated as wrong are provided by a
passage in the botanical laws of 1867 which is believed to define the legitimate
authority of congresses and codes:
"Les regies de la nomenclature ne pouvent etre ni arbitraires ni imposees. Elles
doivent etre bassees sur des motifs assez clairs et assez forts pour que chacun les
comprenne et soit dispose a les accepter."
It is implied by this statement that principles, appealing to the reason and found
sound by the trial of experience, were in existence when it was written; and this is
the truth. By this statement, the legitimate powers of congresses are those of courts
of common law, which avoid the explicit making of law, but discover the law, inter-
pret it, and apply it. Congresses and codes may legitimately (a) state explicitly
corollaries of the principles when they are not obvious; and (b) determine arbitrarily
matters which are necessarily determined arbitrarily, not being within the range of
principle. One would not in theory deny a power (c) to validate breaches of principle
when these are of an expedience verging on necessity; but its use by botanical con-
gresses to produce a roll of exceptions of twice the bulk of the text of the code leads
one to doubt the expedience of this admission. It has been through failure to recog-
nize the legitimate limits of their powers — through a conception that their powers
are sovereign or plenary — that international congresses have come to enact codes
conflicting with each other and giving incomplete satisfaction in themselves.
Under these circumstances, a nomenclature of superior legitimacy can be applied
in groups treated as removed from the jurisdiction of the codes. Not without diffi-
dence, this assumption is extended to the bacteria; it will be agreed that the nomen-
clatural practice applied to the bacteria must be the same as that which is applied
to the blue-green algae.
Here one attempts a brief formulation of those principles, appealing to reason
and proven sound in practice, to which all nomenclature must conform.
1. Scientific names are words of the Latin language. They are not "of Latin form"
or "construed as Latin"; they are Latin. This is to treat Latin as a living language and
scientific names as subject to the rules of its grammar. They are not code-designa-
tions, nor words of any language or none, as chemical names are.
2. The name of a group of the kind called a genus is a proper noun in the singular.
Linnaeus replaced all generic names which were adjectives; all of us his successors
should do likewise.
3. The names of groups of genera are proper nouns, or adjectives used as proper
nouns, in the plural.
The foregoing principles are of pre-Linnaean origin; beginning with his first sig-
nificant work (1735), Linnaeus took them for granted. For the principle next to be
stated, authority is the practice of Linnaeus in later works (1753 and subsequently) :
4. The name of a species consists of the name of the genus to which it belongs fol-
lowed by one epithet, ordinarily an adjective, occasionally a noun in apposition or
in the genitive.
8 ] The Classification of Lower Organisms
A fifth principle represents Linnaean practices as subsequently modified:
5. Named taxonomic groups are necessarily of certain fixed ranks called categories,
i.e., lists. There are seven principal categories, specified as follows. Every individual
organism belongs to a group conceived as the single kind and called a species. Every
species belongs to a genus; every genus to a family; every family to an order; every
order to a class; every class to a division or phylum; ever)' division or phylum to a
kingdom. These conventions have the effect that the groups of each principal category
embrace the entire range of the kinds of organisms.
The categories of genera and species come down from classic antiquity. Linnaeus
originated orders; he originated classes in the sense of named definite groups; and it
appears that he is responsible for kingdoms: the writer knows of no earlier authority
for the traditional three kingdoms of nature. The category next below that of king-
doms has been variously called; originally it was emhranchements (Cuvier, 1812).
The history of the category of families is somewhat involved. It originated in the
work of Adanson (1763); in the following year, Linnaeus (1764) treated the groups
which Adanson had called families as natural orders. Botanists for a long time held
that families and orders are the same thing. Zoological practice gradually made fam-
ilies a separate category. Authority for the list of seven principal categories as given
is Agassiz (1857).
Nothing prevents the assignment of groups to categories other than these, to sub-
classes, tribes, and the like. These may be called subordinate categories. The groups
of any subordinate category embrace only fragments of the range of kinds of
organisms.
The work of Linnaeus was largely innovation, and he did not have the face to de-
clare binding the generally accepted rule of priority. Definite authority for the rule
is de Candolle (1813). As currently applied, it may be stated as follows:
6. The valid name of a group is its oldest published name, conforming to the rules,
and not previously applied in the same kingdom.
As corollaries of the rule of priority, when groups are combined, the oldest name
of any of them must be applied to the whole, and when a group is divided, its name
must be retained for one of the parts. The part to which the original name is to be
applied is determined by the method of types, formulated by Strickland and his as-
sociates (1843) :
7. When a group is divided, its name must be applied to the portion which includes
whatever part of it the original author would have regarded as typical. The part thus
specified is the nomcnclatural type of the group.
In the application of these principles to the naming of the groups of Mychota and
Protoctista, the following practices appear expedient.
A name is applied by publication in such fashion that the community of biologists
may reasonably be held responsible for knowing of its existence and recognizing the
entity to which it is to be applied. This means that it is to be printed in a technical
book or journal and defined in a language for which the generality of biologists will
not require an interpreter, namely Latin, English, French, or German. Any regulation
more detailed than this is an excuse for breaches of priority. Definition is not neces-
sarily by description: nearly all of the Linnaean genera of plants were established
by the listing of species in the Species Plantarum.
When two or more groups published in the same work at the same time are to be
combined, their names are of equal priority. The choice of one of their names by the
first author who combines them is binding.
An Essay on Nomenclature [ 9
A type as specified in the original publication of a group, or as implied by the in-
clusion of a single subordinate group, is unchangeable. Linnaeus and his immediate
successors had no conception of the device of types, and it is practically impossible
to be certain of the elements which they would have regarded as typical in some of
their groups. It remains necessary that the type system be applied to these groups. In
some of them, it may be expedient that international authority, proceeding with due
caution, declare types arbitrarily. An individual scholar will do better to call what he
supposes to be the type of a group by a difTerent term, namely standard (Sprague,
1926) : the standard of a group is a supposed type which remains open to debate. The
framers of codes have undertaken to make binding the choice of a type by the first
author who divides a group. On various occasions, however, this action has been
demonstrably mistaken.
Certain venerable names, as Vermes and Algae as used by Linnaeus, were applied
to altogether miscellaneous collections of organisms among which the selection of a
standard would be purely arbitrary. Such names are called nomina confusa, and are
to be abandoned.
It follows from the principle of the binomial nomenclature of species that no genus
is named until one or more of its species are designated by binomial names. It fol-
lows also that in works in which the nomenclature of species is not definitely binomial
no names are of any standing. Hence, the point of time from which priority is effective
is that of the introduction of binomial nomenclature, namely 1753. The enactment of
other starting points for the nomenclature of particular groups is pretended law
which is not law, like the pretended laws of American states which attempt to regu-
late interstate commerce under the appearance of doing something else.
The original spelling of names, so far as it is tolerable Latin, is not to be changed.
Errors of gender or number, obvious mistakes of spelling, and misprints, are to be
corrected. Good Latin is written without diacritical marks: a German Umlaut in a
name as published is corrected by inserting an e; accents, cedilles, and other barbar-
isms are dropped. The codes err in prescribing changes in spelling beyond those
which are here admitted. If they should establish uniformity in the future, it would
be at the expense of divergence from the most respected works of the past.
Specific epithets are capitalized if they are ( 1 ) names in the nominative, in ap-
position with the generic names; (2) names of persons, places, or organisms in the
genitive; (3) adjectives derived from names of persons.
Transfer of groups from one kingdom to another does not warrant any meddling
with names. When a group is transferred from one kingdom to another, its valid name
in the former — its oldest name not previously used in the kingdom in which it was
originally published — has priority from the date of its original publication.
Names of groups higher than genera are in the plural. Some are proper nouns; the
remainder are adjectives used as proper nouns, agreeing in gender with the names of
the kingdoms in which they are included; either expressing characters of the groups
which they designate, or consisting of generic names modified by terminations signi-
fying "resembling" or "of the group of." Plurals of generic names are not tenable
(de Candolle, 1813) : Ericae means the species of the genus Erica; it does not mean,
and can not be used to designate, the genus together with its allies. Names consisting
of words other than generic names modified by terminations signifying "resembling"
or "of the group of" are not tenable, because they are nonsense: the name Conifer-
inae, applied by Engler to a class, is an adjective with an additional adjectival termi-
nation superimposed.
10 ] The Classification of Lower Organisms
A name once applied in any principal category may not be transferred to another,
unless it be of a form barred in the former and prescribed in the latter. The main
clause of this statement is a consequence of the rule of priority. The exception is a
concession to the practice of using names with uniform endings in certain categories.
Names of groups not of principal categories do not have priority as against names
applied in principal categories. This practice, which denies to names in subordinate
categories the full sanction of priority, is justified by the fact that groups in these cate-
gories are of concern only to specialists in the groups in which they occur; one is not
in reason responsible for being aware of their names in groups outside of ones own
specialty.
Almost all families of plants have had names with the uniform ending -aceae from
the point of time at which the category of families was distinguished from that of
orders. Such names were applied to algae, liverworts, and mosses by Rabenhorst
(1863) and to higher plants by Braun (in Ascherson, 1864). They are adjectives in
the feminine, agreeing with the name of the kingdom Plantae. It is altogether expe-
dient that names of this form be held obligatory throughout the kingdom of plants.
A uniform termination for names of families of animals has been in use for many
years, but these names are not equally positively sound both grammatically and by
priority. There has been a strong tendency to apply uniform terminations to the names
of groups of other categories. So far as concerns groups of subordinate categories —
suborders, subfamilies, and so forth — this practice appears expedient; these groups
being of concern only to experts in the groups in which they occur, it is as well that
their designations be of the nature of code designations rather than names. In at-
tempting to put this practice into effect, some zoologists have made the mistake of
applying the same adjective in different genders to different groups; they have not
realized that Amoebida is the same word as Amoebidae. Meanwhile, uniform termi-
nations for names of phyla, classes, and orders, beside involving wholesale violation
of priority, is something of an insult to the intelligence.
The terminations of ordinal names in -ales and of family names in -aceae, currently
in use among the Mychota, are here changed to -alea and -acea to agree with the
neuter name of the kingdom. A change of the gender of an adjective does not create
a new word, and the original authorities for the names will stand. Accordingly:
The name of an order of Mychota, if based on that of a genus, must bear the termi-
nation -alea. Names of this form are valid in no other category of this kingdom, and
may be reapplied to orders. They have priority and authority by publication explicitly
as orders. Such names do not supersede older ordinal names not based on names of
genera.
The name of a family of Mychota is formed of the stem of a generic name (not
necessarily a valid name, but never a later homonym) by adding the termination
-acea. Names of this form are not valid in any other categor)', and may be reapplied
to families. They have priority and authority by publication explicitly as families.
The names of families of Protoctista, unlike those of Mychota, of plants, and of
animals, do not have by priority prevalently a uniform termination. Many of the
oldest were first named in -ina. Those of flagellates and myxomycetes have double
sets of names, respectively in -aceae and -idae, in current use. It is not expedient to
impose uniform terminations on the names of these groups, at least not in the present
work. Accordingly:
Each group of Protoctista is called by its oldest name of tenable form in the cor-
rect category, barring any previously used in other principal categories, irrespective
An Essay on Nomenclature [ 1 1
of termination. All names which are adjectives are used in the neuter, but ascribed to
the original authors.
The practices described have resulted in the use of many names which will seem
strange, producing lists which are undeniably heterogeneous. A friendly critic notes
as an example of these things the Hst of classes, Heterokonta, Bacillariacea, Oomy-
cetes, and Melanophycea, on page 55. It will be realized that the three among these
names which are adjectives must be in the feminine if the groups are construed as
Plantae, neuter if Protoctista. Taking this fact into account, these are actually the
first names, not previously used in other principal categories, applied to these groups
as classes. What other names could one use? Everyone will know what groups are
intended. Would any person understand them better if new names had been created
by applying a uniform termination to the old roots?
Enough about nomenclature. We should begin to deal with organisms.
Chapter III
KINGDOM MYCHOTA
Kingdom I. MYCHOTA Enderlein
Stamm Moneres Haeckel Gen. Morph. 2: xxii ( 1866), in part.
ScHizoPHYTAE Cohn in Beitr. Biol. Pfl. 1, Heft 3: 201 (1875).
Class ScHizoPHYTA or Protophyta McNab in Jour, of Bot. 15 : 340 ( 1877 ) ; not sec-
tion Protophyta nor cohors Protophyta Endlicher (1836).
Kingdoms Protophyta and Protozoa Haeckel Syst. Phylog. 1: 90 (1894), in part;
not Protophyta Endlicher nor class Protozoa Goldfuss (1818).
Subdivision Schizophyta Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: iii (1900).
Division Schizophyta Wettstein Handb. Syst. Bot. 1 : 56 ( 1901 ).
Phylum Protophyta Schaffner in Ohio Naturalist 9: 446 (1909), in part.
Kingdom Mychota Enderlein Bakt.-Cyclog. 236 (1925).
Kingdom Monera Copeland f. in Quart. Rev. Biol. 13: 385 (1938).
Kingdom Anucleobionta Rothmaler in Biol. Zentralbl. 67: 248 (1948).
Organisms without nuclei.
The common name of Mychota in general is bacteria, but those which contain
chlorophyll together with other pigments which make the green color impure are
called blue-green algae.
The cells of Mychota are always separate or physiologically independent: multi-
cellular bodies with distinct tissues do not occur. The cells are of various shapes; most
often they are cylindrical, being of diameters from a fraction of one micron to a few
microns, rarely more. Except in the groups of myxobacteria and spirochaets, they
are walled; the thickness of the walls is of the order of 0.02^ (Knasyi, 1944). The
walls may contain cellulose, but consist chiefly of pectates, compounds of slightly
oxidized polysaccharides with sulfate, calcium, and magnesium (Kylin, 1943). These
compounds are readily rendered gelatinous by hydration or hydrolysis, and the cells
are often imbedded in gelatinous layers called sheaths or capsules.
In describing the Mychota as lacking nuclei, one commits himself to one side of a
controversy of many years duration. Because of the greater size of the cells of the
blue-green algae, the facts are more easily ascertained in this group than in the proper
bacteria.
The cells of blue-green algae (Gardner, 1906; Swellengrebel, 1910; Haupt, 1923)
are divided into outer and inner parts which are not sharply distinct. Pigments occur
in a dissolved or colloidal condition in the outer part, which contains also granules
of stored food. The granules are not carbohydrate, although a form of glycogen dis-
tinct from that of higher organisms has been extracted (Gardner; Kylin, 1943). The
inner part contains rods and granules, some of which stain like chromatin, while
others ("red granules of Biitschli") are stained red by methylene blue. Cell division
is by constriction. Olive (1904) interpreted the inner part of the cell as a nucleus
continually in process of mitosis, and accordingly without a membrane. It is true that
in series of disk-shaped cells one may recognize series of corresponding granules.
Where the cells are more elongate, the rods and granules of the interior are divided
at random. Haupt expressed the impropriety of calling any part of these cells a
nucleus.
Kingdom Mychota
[13
Recent studies of typical bacteria by conventional microtechnical methods (Rob-
inow, 1942, 1949; Tulasne and Vendrely, 1947) and by the electron microscope (Hil-
lier, Mudd, and Smith, 1949) have made it possible to recognize the essential identity
of the structure of their cells with those of the blue-green algae. The protoplast con-
sists of outer and inner parts. The outer part, considered as a substance, may be
called ectoplasm (Knasyi, 1930), and the inner, considered as a body, may be called
the central body (Biitschli, 1890). The ectoplasm is very thin, occupying usually less
than one fifth of the radius of the cell. The spiral bands which have often been seen
%
Bi^i
Ss;.-
Z#
)ii-i'-
"ji;-^
Irx
•■■•;i©\ ^->
'■l-i"
V ^ .?.. ■fw .^,.'^» ■;
^vWfe
■4 m
Fig. 1.- — Structure of cells of blue-green algae, a, Symploca Muscorum after
Gardner (1906). b, Oscillatoria Princeps after Olive (1904). C, Lyngbya sp. from
a slide prepared by Dr. P. Maheshwari, x 1,000. d, Anabacna circinnalis after
Haupt (1923) x 2,000.
in cells of bacteria, and which Swellengrebel ( 1906) mistook for a nucleus, are thick-
enings of the ectoplasm. Specific stains for nucleoprotein (chromatin), as Feulgen
or Giemsa, usually color uniformly the entire central body. If the cells are exposed to
hydrochloric acid, a part of the nucleoprotein, containing ribonucleic acid, dissolves.
The remainder, containing desoxyribonucleic acid, persists in the form, basically, of
a single fairly large granule in each cell. In rod-shaped bacteria, this granule appears
usually to divide by constriction before the cell begins to divide, and may redivide,
so that the cell may contain two dumb-bell shaped bodies. De Lamater and Hunter
(1951) succeeded in a partial de-staining of the dumb-bell shaped bodies and inter-
preted them as dividing nuclei containing centrosomes and definite numbers of
chromosomes; typical chromosomes, however, are never as small as the bodies they
describe, and are not imbedded in bodies of nucleoprotein from which they can be
distinguished only by the most refined technique. Enderlein (1916) observed in rod-
shaped bacteria series of granules of which some at least are identical with the dumb-
14 ] The Classification of Lower Organisms
bell shaped bodies. He named these granules mychits. It might be held that the
mychit is a chromosome, and the central body of bacteria a nucleus of a single
chromosome, if it were not true that the blue-green algae contain comparable bodies
of variable form and indefinite number.
Many bacteria swim by means of flagella. The diameter of the flagella, as revealed
by the electron microscope, is of the order of 0.02 [J.. Their positions and lengths were
made known, before the invention of the electron microscope, by the technique of
Loeffler (1889), which consists essentially of depositing upon them a heavy layer
of tannic acid. By the absence or presence and arrangement of flagella, bacteria are
classified as of four types: atrichous, without flagella; monotrichous, with one flagel-
lum at one end; lophotrichous, with a tuft of flagella at one end; peritrichous, with
flagella on the sides.
Myxobacteria, spirochaets, and such blue-green algae as are sheathless filaments,
are capable of bending movements (some spirochaets, observed with the electron
microscope, are found also to have flagella at the ends of the cells). Spirochaets swim
vigorously; in myxobacteria and blue-green algae, the bending movements are a mat-
ter of slow writhing. Filaments and cells of blue-green algae are capable also of a
moderately rapid gliding movement. The mechanism of this movement has been
the subject of much speculation, reviewed by Burkholder ( 1934), but remains uncer-
tain. The appearance of the movement is as though it were caused by local secretion
of substances affecting surface tension.
The normal reproduction of Mychota is by constriction of the cells, each into two
equal daughter cells; whence the various names in schizo- (Greek axi^co, to split).
Henrici (1928) studied the changes undergone by bacteria during multiplication. As
the cells become numerous, decreasing the food supply and producing substances
harmful to themselves, they begin to attain greater length before dividing. Subse-
quently there is a gradual transition to enlarged and distorted forms called involution
forms, which divide irregularly, cutting off minute fragments. These observations
suggest the idea that the involution forms are the true normal forms of bacteria, the
so-called normal forms being a temporary stage adapted to rapid multiplication
under favorable conditions.
In many rod-shaped bacteria, when conditions cease to be ideal, the protoplasts
produce within themselves walled bodies of dehydrated protoplasm called spores
(endospores). In general, each cell produces only one spore. No experiment has
definitely shown how long these spores can remain alive; it is surely a matter of cen-
turies, doubtfully of millenia.
Lohnis and Smith (1916, 1923) observed of Azotobactcr that numbers of proto-
plasts might escape from their walls and unite in a common mass, which they named
the symplasm. The existence of this stage has never been confirmed by other authori-
ties. If the symplasm exists, it is a device for achieving the effect which nucleate or-
ganisms attain by sexual reproduction, that is, combination of the heredity of differ-
ent lines of ancestry.
Tliat Mychota can actually combine characters from different linos of ancestry
was first demonstrated beyond question by Tatum and Ledcrberg (1947). They
mixed cultures of pairs of varieties of Escherichia coli, differing in two or more
physiological characters, and isolated from the mixtures races having characters de-
rived from both components. Further Mork, reviewed by Ledcrberg and Tatum
(1953), has abundantly demonstrated phenomena analogous to typical sexual
reproduction.
Kingdom Myrhola
[15
'9^
^ V*
••% ^
^ ^^ m^ W 9m w%
#^., li.^
Fig. 2. — Photographs of Escherichia coli by Dr. C. F. Robinow, reproduced by
Hillier, Mudd, and Smith (1949); left, stained to show the ectoplasm, in which
there are thickenings which tend to be spiral; right, stained to show the large re-
peatedly dividing granule in the central body. About x 2,000. By courtesy of Dr.
Robinow and of the Society of .\merican Bacteriologists.
Kingdom Mychota [17
The metabolic systems of the Mychota are remarkably diverse. The most super-
ficial list of physiological types would include the following: (a) anaerobic parasites
and saprophytes; (b) facultatively aerobic parasites and saprophytes; (c) the vinegar
bacteria, being apparently the only known organisms which, while requiring organic
matter, are incapable of anaerobic energesis; (d) the autotrophic bacteria, the only
organisms which maintain life by oxidation of inorganic matter; (e) organisms living
by incomplete photosynthesis; and (f) organisms capable of typical photosynthesis.
Geologically, the Mychota are ancient. Iron deposits and certain other formations
believed to have been produced by them occur in Archeozoic rocks estimated as more
than a billion years old.
More than five thousand names have been applied to species of bacteria, but in
the attempt to distinguish them, only about fifteen hundred are enumerated (Ber-
gey's Manual, 6th ed., 1948). The species of blue-green algae are probably fewer
than one thousand.
The classification of this group is inescapably highly tentative. The morphology
is simple and not highly varied; the physiological characters likewise appear simple,
but are highly varied, including many which are not known in other groups. The
antiquity of the Mychota makes it probable that many groups which appear to be-
long together consist actually of parallel developments. The undoubted antiquity of
the apparent main groups would lead one to place them in the category of divisions
or phyla; but it is not expedient to make many divisions of a group of 2500 species:
this would produce too many divisions of a single class or classes of a single order.
The kingdom is accordingly treated as a single phylum, and its main divisions as
classes.
Phylum ARCHEZOA Haeckel
yhylB. Archephyta and Archezoa Haeckel Syst. Phylog. 1:90 (1894); not Phylum
Archephyta Haeckel (1866).
Phylum Myxophyceae Bessey in Univ. Nebraska Studies 7: 279 (1907).
Phyla Dimychota and Monomychota Enderlein Bakt.-Cyclog. 236 (1925).
Bacteriophyta and Cyanophyta Steinecke (1931).
Stamme Cyanophyta and Schizomycophyta Pascher in Beih. bot. Centralbl. 48,
Abt. 2: 330 (1931).
Divisions Cyanophyta and Schizomycetae Stanier and van Niel in Jour. Bact. 42:
464 (1941).
Characters of the kingdom.
Archezoa is Haeckel's name, at the point cited, for the bacteria. The name had
been applied othervv^ise by Perty (1852), but not in a principal category. It will not
be considered inappropriate, if it be remembered that the meaning of zoe is as much
life as animal.
The conventional division of the group into two classes, bacteria and blue-green
algae, is not perfectly natural. All of the recognized blue-green algae belong together;
but the recognized bacteria are a wide miscellany, some of them belonging with the
blue-green algae. Here three classes are recognized.
1. Cells without internal pigment, heterotrophic
or living by chemosynthesis; not usually pro-
ducing filaments with prominent sheaths.
18 ] The Classification of Lower Organisms
2. Cells with firm walls, non-motile or
motile by means of flagella Class 1. Schizophyta.
2. Cells with thin walls or none, motile by
means of changes of shape, also some-
times by flagella Class 2. Myxoschizomycetes.
1. Cells mostly with internal pigment, living by
photosynthesis or chemosynthesis, exception-
ally heterotrophic; often producing filaments
with prominent sheaths Class 3. Archiplastidea.
Class 1. SCHIZOPHYTA (Cohn) McNab
Schizomycetes Nageli ex Caspary in Bot. Zeit. 15: 760 (1857).
Class Schizophyta or Protophyta McNab in Jour, of Bot. 15: 340 (1877).
Class Schizomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt, 1:
33 (1879).
Class Schizomycetae SchafTncr in Ohio Naturalist 9: 447 (1909).
Classes Holocyclomor pha and Hemicyclomorpha Enderlein Bakt.-Cyclog. 236
(1925).
Dependent or chemosynthetic Mychota, with walled cells, without photosynthetic
pigments and not producing sheathed filaments.
This class includes as orders the typical bacteria and two minor groups.
1. Cells solitary or loosely gathered into clusters
or filaments, spherical, rod-shaped, or spiral,
not differentiated along the axis Order 1. Schizosporea.
1. Consisting of branched filaments not divided
into cells Order 2. Actinomycetalea.
1. Cells attached by stalks, the attached and
free ends differentiated Order 3. Caulobacterialea.
Order 1. Schizosporea [Schizosporeae] Cohn in Hedwigia 11: 17 (1872).
Order Schizomycetes (Nageli) McNab in Jour, of Bot. 15: 340 (1877).
Order Eubacteria Schroter 1886.
Order Haplobacteriacei Fischer in Jahrb. wiss. Bot. 27: 139 (1895).
Orders Cephalotrichinae and Peritrichinae Orla-Jensen in Centralbl. Bkt. Abt.
2,22: 334,344 (1909).
Order Eubacteriales Buchanan in Jour. Bact. 2: 162 (1917).
Mychota whose cells in the typical condition are without internal pigment, walled,
of the form of rods, spheres, or spirals, not differentiated along the axis. As this is a
numerous group, likely with advancing knowledge to require division, it will be well
to provide it with a nomenclatural standard, and to suggest as such Cohn's principal
discovery among bacteria, namely Bacillus sublilis.
These are the typical bacteria. As originally described by Leeuwcnhoeck (1677),
they were taken to be a few kinds of "animacules" distinguished only by extremely
small size. Only after many years were they shown to be numerous and varied, and
highly important as causes of diseases and of other natural phenomena.
The natural classification of the typical bacteria has been hard to discern. The
characters by which groups can be distinguished include forms of cells and of clusters
of cells; absence or presence and arrangement of flagella; non-formation or formation
Kingdom My c hot a [ 19
of endospores; metabolic products; and the peculiar character called Gram reaction.
The method of staining invented by Gram, 1884, consists of staining successively
with gentian violet and iodine. It gives an intense blue-black color. From some bac-
teria, this color is washed out by alcohol; others retain it; the former are said to be
Gram negative, the latter Gram positive. In practice one applies successively gentian
violet, iodine, alcohol, and safranine, the last being a red dye whose function is to
make the Gram negative bacteria visible. The substance stained by gentian violet
plus iodine is believed to be lipoid, such as occurs in all cells. The Gram positive
quality is believed to consist in a relatively low isoelectric point, a capacity, that is,
to combine with anions in a relatively acid medium. This quality lies in the ectoplasm
of the cells and disappears in aging cultures.
The classification given in Bergey's Manual (1923, 1925, 1930, 1934, 1939, 1948)
is accepted (at least among Americans) as standard. The following system of thirteen
families is a moderate rearrangement of the Bergeyan system, with certain ideas or
names from Enderlein (1917, 1925), Buchanan ( 1925), Pribram (1929) and Stanier
andvanNiel (1941).
1. Gram positive, with exceptions many of
which are intracellular parasites; atrichous or
peritrichous.
2. Spheres dividing in more planes than
one.
3. Gram positive Family 1. Micrococcacea.
3. Gram negative; intracellular patho-
gens in animals Family 2. Neisseriacea.
2. Rods, or spheres dividing in one plane.
3. Not producing endospores.
4. Atrichous.
5. Not intracellular parasites. . Family 3. Corynebacteriacea.
5. Intracellular parasites Family 4. Rickettsiacea.
4. Peritrichous Family 5. Kurthiacea .
3. Producing endospores Family 6. Bacillacea.
1. Gram negative.
2. Atrichous or peritrichous, requiring com-
paratively complicated organic food.
3. Not plant pathogens.
4. Not fixing nitrogen.
5. Capable of growth on or-
dinary media Family 7. Achromobacteriacea.
5. Requiring special media;
minute atrichous pathogens. Family 8. Pasteurellacea.
4. Fixing nitrogen Family 10. Azotobacteriacea.
3. Plant pathogens Family 9. Rhizobiacea.
2. Atrichous, monotrichous, or lophotrich-
ous; the atrichous representatives, and
many others, can survive with organic
foods simpler than carbohydrates, or
with none.
3. Mostly requiring at least carbo-
hydrates Family 11. Spirillacea.
20 ] The Classification of Lower Organisms
3. Not requiring carbohydrates.
4. Oxidizing alcohol to acetic
acid, and acetic acid to CO2
and H2O Family 12. Acetobacteriacea.
4. Not as above; many examples
strictly autotrophic Family 13. Nitrobagteriacea.
Family 1. Micrococcacea [Micrococcaceae] Pribram in Jour. Bact. 18: 370, 385
(1929). Family Coccaceae Zopf 1884; but the genus Coccus is a scale insect. Gram
positive spheres producing packets or irregular masses. Micrococcus, saprophytic or
parasitic, producing irregular masses of cells; the pathogenic species have been treated
as a separate genus Staphylococcus. Sarcina, saprophytic or commensal spheres pro-
ducing packets.
Family 2. Neisseriacea [Neisseriaceae] Prevot ex Bergey et al. Manual ed. 5 : 278
(1938). Family Neisseriacees Prevot in Ann. Sci. Nat. Bot. ser. 10, 15: 119 (1933).
Obligate parasites, the Gram negative spherical cells occurring chiefly in pairs within
leucocytes in the lesions of disease. Neisseria gonorrhoeae, the gonococcus; A^. ]Veich-
selbaumii Trevisan {N. intracellularis, N. meningitidis, Auctt.), the meningococcus.
Family 3. Corynebacteriacea [Corynebacteriaceae] Lehmann and Neumann 1907.
Family Corynebacteriidae Enderlein in Sitzber. Gess. naturf. Freunde Berlin (1917) :
314. Family Lactobacillaceae Winslow et al. in Jour. Bact. 2: 561 (1917). Family
Lactobacteriaceae Orla-Jensen 1921. Family Leptotrichaceae Pribram in Jour. Bact.
18: 372 (1929), not family Leptotrichacei Schroter 1886. Gram positive rods, or
spheres dividing in one plane and producing chains, non-motile.
Streptococcus, spheres in chains; saprophytes in milk, involved in the making of
butter and cheese; and commensals and serious pathogens causing, for example,
abscesses, septicemia, erysipelas, and pneumonia.
Diplococcus, spheres usually in pairs, encapsulated. D. pneumoniae occurs in many
immunologically distinct races which are the usual causes of pneumonia.
Lactobacillus, rods, microaerophilic, producing lactic acid. In milk, involved in
the making of butter and cheese; in the oral cavity, being the usual agent of dental
caries (Rosebury, Linton, and Buchbinder, 1929); common in sewage.
Leptotrichia, rods which become exceptionally long before dividing. Oral cavity
of man and beasts.
Corynebacterium, rods, becoming club-shaped, staining in a banded pattern. The
type species is the agent of diphtheria, C. diphthcriae; the genus includes also many
harmless commensals important only as making diagnosis difficult. The cells divide
in an exceptional fashion, by breaking violently from one side to the other near one
end; the cut-off end swings around beside the main body and proceeds to grow.
Repeated division in this manner produces clusters of parallel cells (Park, \V'iliiams,
and Krumweide, 1924).
Family 4. Rickettsiacea [Rickettsiaceae] Pinkerton 1936. Families Bartonellaceae
Gieszszykiewicz 1939 and Chlamydozoaceae Moshkovsky 1945. Minute obligate intra-
cellular parasites of varied form, commonly Gram negative but with Gram positive
granules.
There have been many observations of bodies of the characters stated, but a satis-
factory classification of them is not yet possible. Howard Taylor Ricketts showed
that Rocky Mountain spotted fever is transmitted by the tick Dcrmocentor, and
observed, in the cells of diseased tissues, minute irregularly staining bodies; in 1910,
Kingdo7n Mychota [21
in the course of further studies of the disease, he contracted it and died. Stanislas
Prowazek, called into the Austrian military medical service in 1914, began to study
typhus, which is transmitted by lice; observed similar intracellular bodies; contracted
typhus, and died in February, 1915 (Hartmann, 1915). The cause of Rocky Mountain
spotted fever is Rickettsia Rickettsii, and that of typhus. is R. Prowazekii. Several
other species are known. By serological methods, Anigstein (1927) showed that
R. Melophagi is closely related to Corynebacterium.
In cases of the disease of the west slope of the Andes called verruga peruana,
Oroya Fever, or Carrion's disease, there occur intracellular bodies named Bartonella
bacillijormis. Noguchi and others (192H) completed the demonstration that the
disease is transmitted by biting flies of the genus Phlebotoyniis. Good authority has
construed Bartonella as a sporozoan.
Students of flagellates, Sarkodina, and Infusoria have occasionally observed in
the cytoplasm or nuclei of these organisms minute bodies multiplying to form consid-
erable masses. These parasites have generally been construed as chytrids, but have
little in common with proper chytrids. The genus Caryococcus Dangeard includes at
least a part of them.
Family 5. Kurthiacea, fam. nov. Gram positive peritrichous rods, not producing
endospores. Kurthia, harmless; Listeria Pirie ex Murray in Bergey's Manual 6th ed.
408 (1948), pathogenic in sheep and man.
Family 6. BaciUacea [Bacillacei] Fischer in Jahrb. wiss. Bot. 27: 139 (1895).
The spore-forming rods, always Gram positive, mostly peritrichous, very numerous in
species, common, and important.
Bacillus Cohn 1872, is one of the oldest generic names of rod-shaped bacteria
which can be definitely applied: it can be definitely applied because the type species
B. subtilis was so described as to be recognizable. The genus has been used to include
rods in general or at random. Defined as aerobic spore-formers, as proposed by
Buchanan, 1917, it is a thoroughly natural group. As treated in the fifth edition of
Bergey's Manual, it included nearly 150 duly distinguished species; in the sixth
edition, this number is cut to thirty-three. The great majority are saprophytic. Ex-
ceptions, important pathogens, are B. anthracis; and B. alvei and other species causing
foulbrood of bees.
The anaerobic spore-formers constitute the genus Clostridium. The type species
wa? discovered and named three times in different connections. As an anaerobe
involved in the fermentations which give butter its flavor, it is C. butyricum Prazmow-
ski. As an organisms whose cells contain granules staining like starch, it is Bacillus
Amylobacter van Tieghem. It has the property of fixing nitrogen; discovered in this
capacity by Winogradsky (1902) it was named C. Pastorianum. The species of
Clostridium, as of Bacillus, are numerous. They are primarily saprophytic, but many
species produce powerful toxins and are serious pathogens. Examples are C. tetani;
C. botulinum; and C. septicum and a whole roll of other species, causing various
forms of gangrene, occasion for the study and distinction of which was found during
World War I.
Family 7. Achromobacteriacea [Achromobacteriaceae] Breed 1945. Family Bac-
teriaceae McNab in Jour, of Bot. 15: 340 (1877), based on a generic name which
must be abandoned as a nomen conjusum. Family Enterobacteriaceae Rahn 1937, not
based on a generic name. Gram negative rods which lack the dictinctive characters
of the families subsequently to be treated.
22 ] The Classification of Lower Organisms
The nine genera listed first occur normally in animals, mostly in the gut and
mostly as commensals; exceptions are important pathogens. Most of them produce
acid, and many of them produce gas, from sugar. These genera are the traditional
colon-typhoid-dysentery group.
Escherichia coli, the colon bacillus, and Aerohacter aerogenes, the gas bacillus, are
common commensals which produce acid and gas from dextrose and lactose. The
standard method of testing waters for contamination is essentially a test for the
presence of these organisms.
Klebsiella also produces acid and gas from sugars. It inhabits the respiratory
tract. The cells are heavily capsulated and non-motile. The type species K. pneumo-
niae is an important pathogen, the pneumobacillus of Friedlander.
Proteus vulgaris (this is at least the third genus to bear the name Proteus, but the
first in this kingdom) produces acid and gas from dextrose but not lactose, and
liquefies gelatine. It is usually isolated from spoiled meat.
Salmonella is distinguished from Proteus by non-liquefaction of gelatine. Many
of its species are harmless commensals; others cause paratyphoid fevers. Immunologi-
cal study of cultures of Salmonella from cases of disease and from waters have re-
sulted in the distinction of fully 150 races, mostly unnamed and identifiable only by
immunological reactions. Eberthella includes motile rods producing acid but not
gas from sugars, and belonging to the same immunological system as the various
races of Salmonella. Eberthella typhi causes typhoid fever.
Shigella is distinguished from Eberthella by non-motility. The Shiga bacillus,
S. dystenteriae, is the cause of dystentery.
Bacteroides is a numerous group of acid-producing gut bacteria, motile or non-
motile, generally harmless.^ distinguished from the foregoing as strictly anaerobic.
Alcaligenes fecalis, an apparently harmless organism isolated from intestinal con-
tents, does not produce acid from sugars; grown in milk, it produces an alkaline
reaction.
Numerous races of bacteria which have been isolated from soil and are capable
of attacking cellulose are assigned to the genus Cellulomonas. Bacteria which produce
an extracellular red pigment are Serratia (one of the oldest generic names for bac-
teria); those which produce yellow pigment are Flavobacterium; those which produce
blue, black, or violet growths are Chromobacterium. Cultures which lack the distinc-
tive characters of all of the above named genera (most such cultures have been
isolated from water) are called Achromohacter.
Family 8. Pasteurellacea nom. nov. Family Parvobacteriaceae Rahn; there is no
corresponding generic name. Minute non-motile Gram negative rods, pathogenic,
requiring special media for cultivation. Pasteurclla avicida is the cause of chicken
cholera, upon which Pasteur made important studies. Of greater direct importance
to man is Pasteurella pestis, the cause of plague. Hemophilus includes the agents
of whooping cough, soft chancre, and conjunctivitis. Brucella includes the organisms
which cause Malta fever, undulant fever. Bang's disease, contagious abortion. Pfeif-
ferella mallei is the cause of glanders.
Family 9. Rhizobiacea [Rhizobiaceae] Conn in Jour. Bact. 36: 321 (1938). Gram
negative rods, atrichous or peritrichous, parasites on plants. Cultured in the presence
of sugars, these organisms produce acid; they are evident allies of the colon group.
Erwinia commemorates Erwin F. Smith, the discoverer of many bacteria pathogenic
to plants. Typical species cause blights, wilts, or dry necroses. The discovery by
Burrill, 1882, of Erwinia amylovora, the cause of the fire blight of pears, should
Kingdom Mychota [ 23
have prevented the formulation of a theory, once entertained, that all bacteria
require neutral media, and are accordingly incapable of causing diseases of plants.
The species of Pectobacterium, as P. carotovorum, cause rots. Those of Agro-
bacterium cause galls; A. tumefaciens causes crown gall of many plants.
Rhizobium includes the species which produce little galls ("nodules") on the
roots of plants and which benefit their hosts by fixing nitrogen. The best known
hosts of Rhizobium are plants of the family Leguminosae; the relationship between
Leguminosae and Rhizobium is a classic example of symbiosis. There are several or
many species of Rhizobium, scarcely distinguishable morphologically, but living on
different groups of legumes. The race which was first recognized and isolated, R.
Leguminosarum Frank 1890 [Schinzia Leguminosarum Frank 1879; Bacillus Radicic-
ola Beijerinck 1888) is that which attacks plants of the pea tribe. Bewley and Hutch-
inson (1920) accounted for the variety of forms which Rhizobium can assume. In
the roots of plants it occurs as involution forms. In culture, it is a peritrichous rod,
but the flagella are often reduced to one, and it has been confused with the mono-
trichous bacteria (Conn and Wolfe, 1938).
Family 10. Azotobacteriacea [Azotobacteriaceae] Bergey, Breed, and Murray in
Bergey's Manual 5th ed., preprint, v and 71 (1938). These are the organisms which
were originally isolated by Beijerinck (1901) by inoculating with garden soil shallow
layers of a nitrogen-free nutrient solution containing mannite. The commonest species,
Azotobacter Chroococcum, is usually seen as ellipsoid cells, as much as \\x thick and
7[J. long, solitary, with peritrichous flagella, or forming non-motile clusters imbedded
in a heavy capsule. Beijerinck observed the occurrence of globular involution forms
as much as 15^ in diameter. Lohnis and Smith (1916) made a thorough study of
variations in form, and reported a remarkable variety of other stages, including the
symplasm.
The Pasteurellacea and Rhizobiacea are apparently reasonably close allies of
the Achromobacteriacea. The Azotobacteriacea stand somewhat apart. The remain-
ing families of the present order are more definitely distinct, being marked by mono-
trichous or lophotrichous flagella.
Family 11. Spirillacea [Spirillaceae] Migula 1894. Family Pseudomonadaceae
Winslow et al. in Jour. Bact. 2: 555 (1917). Rods and spirals, Gram negative, mono-
trichous or lophotrichous; not producing much acetic acid, and mostly heterotrophic.
Pseudomonas is a numerous genus of rods which may or may not produce a fluores-
cent pigment soluble in water; they do not produce a yellow pigment which is in-
soluble in water. The original species, P. aeruginosa, was isolated from pus, in which
it produces a blue-green discoloration; it is by itself weakly if at all pathogenic.
Other species have been isolated from fresh and salt waters and brines; the bacteria
which produce phosphorescence on salt fish are of this genus. Many further species
arc: pathogenic to plants, producing chiefly leaf spots.
Phytomonas Bergey et al. 1923 {Xanthomonas Dowson 1948) includes numerous
plant pathogens which in culture produce an insoluble yellow pigment; among them
are the causes of cabbage rot, walnut blight, and leaf spots on many plants.
Pacinia Trevisan 1885 includes monotrichous curved rods. The type species P.
cholerae-asiaticae is the cause of Asiatic cholera. Among numerous other species
the majority are harmless saprophytes in waters. Recent authorities have treated the
cholera organism as the type of the genus Vibrio Miiller (1773); their action is an in-
tolerable falsification of the usage of a full century preceding the discovery of the
cholera organism.
24 ] The Classification of Lower Organisms
Spirillum includes the typical spirals, lophotrichous, a small number of species of
harmless saprophytes in foul waters.
Thiospira includes large lophotrichous spirals, colorless, containing granules of
sulfur. They are believed to live by chemosynthesis.
Family 12. Acetobacteriacea [Acetobacteriaceae] Bergey, Breed, and Murray
1938. As gross objects, growths of Acetobacter aceti Beijerinck have been known since
prehistoric times. With included yeasts they constitute mother of vinegar (the old
names Mycoderma mesentericum Persoon, Ulvina aceti Kiitzing, and Umbina aceti
Nageli designated the combination of bacteria and yeasts, and it seems proper to
reject them). Free-swimming cells with polar flagella have been observed; ordinarily
the cells appear as rods in chains, heavily encapsulated, or as involution forms.
The organic food required by Acetobacter is alternatively alcohol, which is oxidized
to acetic acid, or acetic acid, which is oxidized to carbon dioxide and water. These
processes are strictly aerobic: to make vinegar, one exposes wine to air; to preserve
it, one seals the vessels.
Family 13. Nitrobacteriacea [Nitrobacteriaceae] Buchanan in Jour. Bact. 2: 349
(1917). Organisms oxidizing the simplest organic compounds; or facultatively capa-
ble of chemosynthesis; or living strictly by chemosynthesis and strictly aerobic: mostly
Gram negative monotrichous or atrichous rods.
Methanomonas is capable of oxidizing methane; Carboxidomonas of oxidizing
carbon monoxide; Hydrogenomonas, of oxidizing elemental hydrogen. Thiobacillus
includes organisms which oxidize hydrogen sulfide or elemental sulfur.
Winogradsky had discovered chemosynthesis in the course of studies of Beggiatoa
and other sulfur-oxidizing organisms before he undertook to isolate bacteria which
cause nitrification, that is, the natural production of nitrates in soil and waters.
He achieved success (1890) by inoculating, with soil or sewage, media which con-
tained salts of ammonia but no food; he saw the nitrifying organisms first as minute
motile rods which he named Nitromonas. Further study and the use of solid media
showed that nitrification takes place in two stages and is the work of several kinds of
organisms. Winogradsky distinguished Nitrosomonas europaea and N. javaneyisis,
monotrichous rods from different regions as indicated, oxidizing ammonia to nitrites;
Nitrosococcus, non-motile spheres from South Amerca, effecting the same oxidation
as Nitrosomonas; and Nitrobacter, non-motile rods oxidizing nitrites to nitrates.
Subsequent authors have validated Winogradsky's names by creating the combina-
tions Nitrosococcus nitrosus and Nitrobacter VVinogradskyi. Subsequently, Winograd-
sky discovered yet other bacteria capable of the same oxidations.
The presence of nitrifying bacteria is necessary for the normal growth of most
crops. So active are the nitrifying bacteria that no more than traces of ammonia and
nitrites are found in normal soils, and so avidly do plants absorb nitrates that these
accumulate only in fallow fields.
Order 2. Actinomycetalea [Actinomycetales] Buchanan in Jour. Bact. 2: 162
(1917).
Organisms which consist typically of slender filaments not divided into cells,
but which are capable of producing conidia, that is, minute spherical or elongate
bodies cut off by constriction from the ends of the filaments, or of breaking up into
cells of the form of regular or irregular rods. Non-motile; Gram positive or Gram
negative; often of the staining character called acid fast.
The order may be treated as a single family.
Kingdom Mychota [ 25
Family Mycobacteriacea [Mycobacteriaceae] Chester 1907. Family Actinomyce-
taceae Buchanan in Jour. Bact. 3: 403 (1918). Family Streptomycetaceae Waksman
and Henrici 1943. Characters of the order. Three genera require discussion.
Streptomyces Waksman and Henrici 1943. The original name of this genus is
Streptothrix Cohn (1875); there is an older genus Streptothrix among plants, and
the numerous species of the present genus have generally been included in Actino-
myces. Cultures are readily isolated from air or soil. They appear as slowly growing
colonies which may at first be of various colors and have shiny surfaces. Their texture
is tough; a blunt needle will more often tear a colony from the medium than pene-
trate it. As the colonies grow, they become truncate; the exposed surfaces become
white and powdery; pigments, black, brown, red, or yellow, in various races, are
produced, and discolor the medium. The toughness of the colonies is a consequence
of their structure, of myriad crooked branching filaments about 1|J. in diameter,
without joints; the white and powdery surface is produced by myriad conidia released
in basipetal succession. The cultures are of an odor which may be described as that
of earth under the first rain after drouth: undoubtedly, this familiar odor is that of
Streptomyces in the soil. Drechsler (1919), from careful study of several species of
Streptomyces, concluded that they are fungi; their filaments are, however, much
finer than those of fungi, and no definite nuclei have been seen.
Certain species of Streptomyces cause a scabbiness of potatoes. Except for this, the
genus was for a long time regarded as quite unimportant. When the capacity of the
fungus Penicillium notatum to inhibit the growth of bacteria had been observed,
and had led to the discovery of the drug penicillin, Waksman, the leading authority
on the classification of Actinomycetalea, sought comparable drugs produced by
Streptomyces, and had the great success of discovering streptomycin.
Actinomyces Bovis Harz 1877 is one of several species of the same general nature
as Streptothrix which are pathogenic to animals. It causes lumpy jaw of cattle.
Mycobacterium Lehmann and Neumann 1896 is typified by M. tuberculosis, the
agent of one of the most important diseases of man, supposed originally to have
attacked cattle, and to have spread around the world with European cattle. It is a
chronic disease, destroying the tissues slowly and producing a nugatory sort of im-
munity which makes it possible to test for the disease, but does not check it. The
cells are recognized in sputum and in diseased tissues by the acid fast reaction: the
dye carbol fuchsin must be applied hot in order to color them; once it has done so,
it does not wash out in acid alcohol. It is cultivated with difficulty. The growth is
dry, powdery, wrinkled, with an odor described as sickening-sweet. It consists of
branching filaments which break up readily into rod-shaped or irregular fragments.
Lesions of leprosy contain acid fast organisms named Mycobacterium leprae. Gay
(1935) has discussed the results of attempts to cultivate this species. They have
yielded either "diphtheroid" cells or a "streptothrix." He concludes that most of
the reports are of the same organism reacting variously to various conditions.
Order 3. Caulobacterialea [Caulobacteriales] Henrici and Johnson in Jour. Bact.
29: 4 (1935).
Aquatic bacteria, the cells of most examples secreting gelatinous matter in such a
manner as to produce stalks. Henrici and Johnson provided a system of four families,
five genera, and nine species. Stanier and van Niel (1941) rejected the group as
artificial, placing some of the genera among Eubacteria and leaving others unplaced.
The order may be maintained for the accommodation of the latter and divided into
two families.
26]
The Classification of Lower Organisms
m
Fig. 3 — a-e, Caulobacterialea after Henrici and Johnson (1935) x 2,000: a,
Nevskia sp.; b, Caulobacter vibrioides; c, Caulobacter sp.; d, Pasteuria sp.; e, Blasto-
caulis sp. f. Various stages of Cytophaga Hutchinsonii [Spirochaeta cytophaga) after
Hutchinson and Clayton ( 1919). g-k, Myxobactralea after Thaxtcr (1892), the cells
X 1,000, in the fruits x 200. g, h, Cells and fruit of Chondromyccs crocatus; i, fruit
of C. aurantiacus; j,k, vegetative cells and spores, and fruit, of Myxococcus coralloi
des. I, m, Dividing cells of Cristispira Veneris after Dobell (1911) x 2,000.
Kingdom Mychota [ 27
Family 1. Leptotrichacea [Leptotrichacei] Schroter 1886. The cells not elongated
in the direction of the axis of the stalk.
Didymohelix ferruginea (Ehrenberg) Griffith (first named, and usually listed,
under Gallionella, which is a misspelling of the name of a genus of diatoms) occurs
in waters containing iron. Older authors described it as consisting of paired filaments,
less than 1^ in diameter, colored bright yellow with imbedded iron oxide, and coiled
about each other. In fact, the supposed paired filaments are the margins of a single
twisted band, which is not itself an organism but the stalk secreted by a terminal
cell. Spirophyllum Ellis is either the same species or a closely related larger one.
Leptothrix Kiitzing Phyc. Gen. 198 ( 1843) was inadequately described; the species
which was first named, and which is accepted as the type, was L. ochracea. It is be-
lieved that this name properly designates the masses of ochraceous matter seen in
iron springs. Under the microscope, this matter is seen to consist of fine yellow
filaments, straight and unbranched. Ellis (1916) described them as consisting of a
cylinder of protoplasm, not divided into cells, enclosed in a sheath. Almost surely,
these structures, generally recognized as of the same nature as Didymohelix, are like-
wise stalks secreted by minute terminal cells.
Siderocapsa Molisch and Sideromonas Cholodny, described as minute spheres or
rods imbedded in capsules colored by ferric oxide and attached to plants in waters
containing iron, are perhaps to be interpreted as stalkless members of the present
group.
Nevskia Famintzin, forming minute gelatinous colonies floating on water, does not
accumulate iron.
Family 2. Caulobacteriacea [Caulobacteriaceae] Henrici and Johnson 1. c. (1935).
The cells elongated in the direction of the long axes of the stalks. Caulobacter, Pas-
teuria, and Blastocaulis, colorless saprophytes in waters or parasites in aquatic
animacules.
Class 2. MYXOSCHIZOMYCETES Schaffner
Class Myxoschizomycetae Schaffner in Ohio Naturalist 9: 447 (1909).
Class Polyyangidae Jahn Beitr. bot. Protistol. 1: 65 (1924).
Class Spirochaetae Stanier and van Niel in Jour. Bact. 42 : 459 ( 1941 ) .
Parasitic or saprophytic Mychota, the elongate cells with thin walls or none,
capable of bending movements and sluggishly or actively motile. In many examples
there is a resting stage: the cell contracts generally, so as to diminish the surface,
and deposits a definite wall. The structure so produced is a spore of the type called
an arthrospore or chlamydospore.
The two orders Myxobactralea and Spirochaetalea have not previously been
combined to form a separate class. A certain species which Hutchinson and Clayton
(1919) described as a spirochaet, Spirochaeta cytophaga, has subsequently been
found to be a myxobacterium. The hint of relationship thus conveyed is confirmed
by the whole character of both groups, as may be seen from the discussions of them
by Stanier and van Niel (1941) and Knasyi (1944).
Order 1. Myxobactralea [Myxobactrales] Clements Gen. Fung. 8 (1909).
Order Myxobacteriaceae Thaxter in Bot. Gaz. 17: 389 (1892).
Order Myxobacteriales Buchanan in Jour. Bact. 2: 163 (1917).
The cells not definitely of spiral form, sluggishly motile. In typical examples, the
28 ] The Classification of Lower Organisms
cells occur in swarms imbedded in slime; the entire mass moves concertedly, and is
eventually converted into macroscopically visible fruiting bodies.
The group was first recognized by Thaxter. He took note that the fruiting bodies
of Chondromyces had already been described by Berkeley and Curtis as those of a
gasteromycete, and learned subsequently that Polyangium Link, also described as
of the puffball group, is an older name for his Myxobacter. The swarms of cells live
in air on damp substrata (commonly the feces of various kinds of animals), moving
across them and digesting and absorbing food as they proceed. Labratory culture is
fairly easy. As a reaction, apparently, to exhaustion of the available food, the cells
change into chlamydospores; the masses of spores held together by dried slime are
called cysts. These may be borne on simple or branched stalks built up from the
slime as a preliminary to the formation of the cysts and spores. The group is of
essentially no economic importance.
The accepted classification is that of Jahn (1924); to the four families which he
recognized, one more has been prefixed for the accommodation of the genus
Cytophaga.
family 1. Cytophagacea [Cytophagacae] Stanier 1940. The chlamydospores
formed sporadically by individual cells, not in cysts. Cytophaga Hutchinsonii Wino-
gradsky [Spirochaeta cytophaga Hutchinson and Clayton) is one of several species
discovered as active fermenters of cellulose. The slenderly spindle-shaped cells are
sluggishly motile, and produce ellipsoid chlamydospores resembling yeasts.
Family 2. Archangiacea [Archangiacae] Jahn op. cit. 66. Spores elongate in irregu-
larly extensive masses, not in cysts. Archangium, Stelangium.
Family 3. Sorangiacea [Sorangiaceae] Jahn op. cit. 73. Spores elongate, the
cysts angular, in masses, not stalked. Sorangium.
Family 4. Myxobacteriacea [Myxobacteriaceae] (Thaxter) E. F. Smith 1905.
Family Polyangiaceac Jahn op. cit. 75. Spores elongate, in distinct rounded cysts,
clustered or solitary, sessile or borne on simple or branched stalks. Polyangium
Link 1795 [Myxobacter Thaxter 1892), Stelangium, Melitangium, Podangium,
Chondromyces.
Family 5. Myxococcacea [Myxococcaceae] Jahn op. cit. 83. Spores spherical;
cysts indefinite or definite. Myxococcus, Chondrococcus, Angiococcus.
Order 2. Spirochaetalea [Spirochaetales] Buchanan in Jour. Bact. 2: 163 (1917).
Cells solitary, spiral in shape, actively motile.
The first known species of this group was Spirochaeta plicatilis, observed in foul
waters by Ehrenberg (1838). The next was the species now known as Borrelia recur-
rentis (Lebert) Bergey et al., observed in the blood of relapsing fever patients by
Obermeier, 1873.
During the last years of the nineteenth century, many attempts to identify the
agent of syphilis by standard bacteriological methods were unsuccessful. The German
government directed Schaudinn and Hoff'mann to continue this work. Fritz Schau-
dinn, 1871-1906 (Stokes, 1931), had attained distinction as a student of pathogenic
protozoa. Within a few weeks, by the microscopic examination of lesions, he attained
success where the bacteriologists had failed, and discovered Treponema pallidum
(Schaudinn and Hoffmann, 1905).
Spirochaets were first cultivated by Noguchi; few others have been successful in
this difficult practice. It requires a medium of aseptic, not sterilized, animal ma-
terial, under more or less anaerobic conditions. Each species requires its peculiar
variant of the conditions, to which it is quite sensitive.
Kingdom Mychota [ 29
Spirochaeta plicatilis and other saprophytic species, together with certain species
parasitic in mollusks, are fairly large. The species which are parasitic or commensal
in other animals may be extremely small. It is chiefly by study of the larger species
that the structure is known. The internal structure is septate. Dobell (1911) found
in Cristispira, at the margin of each septum, a whorl of granules staining like chroma-
tin, and interpreted these granules collectively as a nucleus. Noguchi (in Jordan and
Falk, 1928) saw in the interior of the smaller species no chambered structure, but a
lengthwise rod. This has been interpreted as a nucleus, as a locomotor or skeletal
structure, or as an artifact. The electron microscope has shown actual flagella at the
ends of cells of Treponema pallidum. Reproduction is normally by transverse divi-
sion into two. During division, the daughter cells may coil about one another, giving
a false appearance of lengthwise division. Gross (1913) observed that Cristispira is
capable of breaking up into cylindrical Stdhchen corresponding to the chambers.
The discovery of Treponema by an eminent protozoologist; the character of
spirochaetal diseases, several of which are spread by biting insects, and produce only
that nugatory immunity which makes diagnosis possible but does not check the
disease; and the supposed lengthwise division of the cells; led to the hypothesis that
the spirochaets are protozoa. Dobell was surely correct in dismissing this hypothesis,
insisting that the spirochaets are neither protozoa nor typical bacteria, but a group
sui generis.
The larger and smaller spirochaets are reasonably treated as separate families.
Family 1. Spirochaetacea [Spirochaetaceae] Swellengrebel 1907. The cells com-
paratively large, 80-500(1 long. Spirochaeta, Saprospira, Cristispira.
Family 2. Treponematacea [Treponemataceae] Robinson in Bergey Man. 6th ed.
(1948). Family Treponemidae Schaudinn 1905. The cells 4-15^ long.
Treponema Schaudinn. The cells comparatively loosely coiled. T. pallidum, the
agent of syphillis. T. pertenue, the agent of yaws. T. macrodentium and T. micro-
dentium, harmless commensals in the mouth.
Borrelia Swellengrebel is doubtfully distinct from the foregoing; Noguchi reduced
it. B. recurrentis and other species cause relapsing fevers. B. Vincenti causes Vincent's
angina (trench mouth). The fusiform cells always found associated with it and
supposed to be ordinary bacteria of a genus Fusiformis or Fusobacterium may be its
chlamydospores.
Leptospira Noguchi. The cells tightly coiled. L. icterohaemorrhagiae is the agent
of infectious jaundice. L. icteroides, isolated by Noguchi in South America, sup-
posedly from cases of yellow fever, is perhaps the same thing: it is now known that
yellow fever is caused by a virus. It was in pursuing in Africa his study of yellow
fever that Noguchi lost his life by this disease (Flexner, 1929; Eckstein, 1931).
Class 3. ARCHSPLASTIDEA Bessey
Myxophykea Wallroth 1853.
Myxophyceae Stizenberger 1860.
Division (of Class Algen) Pkycochromaceae and order Gloiophyceae Rabenhorst
Krytog.-Fl. Sachsen 1: 56' (1863).
Cyanophyceae Sachs Lehrb. Bot. ed. 4: 248 (1874).
OrAtx Cyanophyceae or Pkycochromaceae yicNdLhm]o\iT. oi'Qot. 15: 340 (1877).
Schizophyceae Cohn 1879, not suborder Schizophyceae Rabenhorst Deutschland's
Kryptog.-Fl. 2, Abt. 2: 16 (1847).
30 ] The Classificatio7i of Lower Organisms
Order Schizophyceae Schenck in Strasburger et al. Lehrb. Bot. 1894.
Class Schizophyceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
la: iii (1900).
Class Archiplastideae Bessey in Univ. Nebraska Studies 7: 279 (1907).
Class Cyanophyceae Schaffner in Ohio Naturalist 9: 446 (1909).
Class Myxophyceae G. M. Smith (1918).
Subclass Myxophyceae Setchell and Gardner in Univ. California Publ. Bot. 8,
part 1: 3 (1919).
Cyanophyta Steinke ( 193 1 ) .
Stamm Cyanophyta Pascher in Beih. Bot. Centralbl. 48, Abt. 2: 330 (1931).
Mychota most of which live by phytosynthesis of primitive or typical character,
many of them, and most of the saprophytic and chemosynthetic organisms included
with them, being of the form of sheathed filaments.
This is primarily the group of the blue-green algae. Blue-green algae are familiar
things as forming dark scums in water and on wet surfaces. Rabenhorst (1863) ap-
pears first to have recognized them as a group definitely distinct from green algae;
he named most of the recognized families. Revisions by Thuret (1875), Bomet and
Flahault (1886-1888), and Gomont (1892) failed to provide a satisfactory system
of the group; Kirchner's revision (in Engler and Prantl, 1898) is the accepted system.
One of the important contributions of Cohn was his suggestion that the bacteria
and blue-green algae belong together. He emphasized this view by mingling the
genera of the two groups in two new groups, "tribes," named in effect slime-formers
and thread-formers (1875). In this he went too far; but some of the arrangements
which he suggested appear natural. Beggiatoa, the type of order Thiobacteria of
Migula, appears to be a variant of the common blue-green alga Oscillatoria, differing
from it in living by chemosynthesis. Most of the so-called iron bacteria, family
Chlamydobacteriaceae of Migula, fall readily into scattered places among the blue-
green algae. Only the genus Sphaerotilus remains at loose ends. It is credibly reported
to produce cells swimming by means of flagella; no proper blue-green algae do this.
A variety of purple bacteria — bacteria, that is, which contain a red pigment —
have been discovered from time to time. Engelmarm (1888) observed that they swim
toward the light, and convinced himself that they live by photosynthesis. Van Niel
confirmed this, and showed that photosynthesis is in this group of a peculiar character;
it requires the presence of reducing agents and does not release oxygen. This type of
photosynthesis appears, in fact, to represent a stage of the evolution of typical photo-
synthesis; the group in which it occurs appears to represent the ancestry of the
typical blue-green algae. The poorly known green bacteria appear to belong with
the purple bacteria.
Various members of this class have been proved capable of fixing nitrogen (Sisler
and ZoBell, 1951; Williams and Burris, 1952).
Four orders may be distinguished:
1. Possessing a red ("purple") intracellular
pigment, or a green pigment not masked by
others Order 1 . Rhodobacteria.
l.With green pigment masked by others, or
colorless.
2. Producing cells with flagella; non-pig-
mented sheathed filaments not accumu-
lating ferrugineous matter Order 2. Sphaerottlalea.
Kingdom Mychota [ 31
2. Never producing cells with flagella.
3. Cells dividing in more planes than
one, growing to full size before re-
dividing; unicellular or colonial, not
filamentous Order 3. Coccogonea.
3. Cells dividing in one plane, and
accordingly producing filaments;
exceptional examples reproducing
by budding (unequal division) or
by repeated division into minute
spores Order 4. Gloiophycea.
Order 1. Rhodobaeteria Molisch Purourbakterien 27 (1907).
Rods, spheres, and spirals, solitary or colonial, with red or green pigment, per-
forming in the presence of light and reducing substances a sort of photosynthesis in
which no oxygen is released.
These organisms have generally been included in Thiobacteria, but do not include
Beggiatoa, the type of that order. Molisch divided them into two families, Thiorho-
daceae, aerobic, accumulating granules of sulfur, and Athiorhodaceae, microaero-
philic or anaerobic, not accumulating granules of sulfur. The green bacteria are to be
placed as a third family. The names originally applied to the families are not tenable.
Family 1. Chromatiacea (Migula) nomen familiare novum. Subfamily Chro-
MATiACEAE Migula. Family Rhodobacteriaceae Migula; Family Thiorhodaceae
Molisch; the family does not include genera with corresponding names. Purple bac-
teria, aerobic, accumulating granules of sulfur. Chromatium Perty includes the or-
ganism of foul waters which was originally named Monas Okenii. It is a plump rod,
often bent, sometimes exceeding \0\Ji in length, monotrichous or lophotrichous. There
are a dozen other genera, rods, spheres, and spirals [Thio spirillum., which belongs
here, is to be distinguished alike from Spirillum, Thiospira, and Rho do spirillum),
solitary or colonial, motile or non-motile. Most of them were discovered by Wino-
gradsky.
Family 2. Rhodobacillacea nom. nov. Family Athiorhodaceae Molisch. Molisch
named in this family a genus Rhodobacterium, but the name Rhodobacteriaceae had
already been applied by Migula to the preceding family. Purple bacteria, anaerobic,
not accumulating granules of sulfur. Molisch discovered all known members of the
present family. The method of culture was to place a mass of organic matter, for
example an egg, in the bottom of a cylinder of water (the original account specified
water of the River Moldau), cover the surface with oil, place in a north window, and
wait several weeks. This method yielded organisms which were assigned to seven
genera. Those of spiral form are Rhodospirillum. All others are by van Niel treated
as a single genus, which may be called Rhodobacillus Molisch {Rhodopseudomonas
van Niel).
Family 3. Chlorobiacea nom. nov. Family Chlorobacteriaceae Geitler and Pascher
ex van Niel in Bergey's Manual ed. 6: 869 (1948). Geitler and Pascher (in Pascher
Siisswasserfl. Deutschland, 1925) did not place this group in a definite category
and name it unequivocally: they called it Cyanochloridinae or Chlorobacteriaceae.
Minute spherical or elongate cells with a green pigment different from typical
chlorophyll, anaerobic, non-motile, producing irregular or regular gelatinous colonies.
Chlorobium, Pelodictyon, Clathro Moris, with a half a dozen known species. Certain
32]
The Classification of Lower Organisms
Fig. 4. — Coccogonea: a, Chroococcus sp.; b, C, Achromatiuni oxalijerum. Gloio-
phycea: d, Oscillatoria splendida; e, Phormidium sp.; f, Beggiatoa sp.; g, Chamae-
siphon incrustans; h, Anabaena inacqualis; \, Cylidrospcrmum majus; j, Chlarnydo-
thrix ochracea; k, 1, m, Clonothrix fusca after Kolk (1938); n, Dermocarpa protea
after Setchell and Gardner ( 1919); o, Crenothrix polyspora after Kolk ( 1938). All
X 1,000.
Kingdom Mychota [33
organisms of this group, occurring in symbiotic combinations with larger bacteria or
with protozoa, have been named as additional genera; one of these is Chlorobacterium
Lauterborn, but the name is a later homonym.
Order 2. Sphaerotilalea nom. nov.
Order Desmobactcrialcs Pribram in Jour. Bact. 18: 376 (1929); there is no cor-
responding generic name.
Cells colorless, elongate, in sheathed filaments which branch freely in the manner
called "false": the cells divide strictly in one plane; those at a distance from the tip
may so multiply as to break the continuity of the series by pushing a growing point
laterally out of the sheath. The cells may escape from the filaments, become lophotri-
chous, and function as swarm spores. There is a single family:
Family Sphaerotilacea [Sphaerotilaceae] Pribram 1. c. There is probably only
one species, Sphaerotilus natans Kiitzing {Cladothrix dichotoma Cohn). It is found
as minute gelatinous colonies fioating on stagnant water; cells 2-4^ in diameter.
Order 3. Coccogonea [Coccogoneae] (Thuret) Campbell Univ. Textb. Bot. 84
(1902).
Tribe Chroococcaceac [Coccogoneae) Thuret in Ann. Sci. Nat. Bot. ser. 6,
1: 377 (1875).
Subclass Coccogoneae Engler in Engler and Prantl Nat. Pfianzenfam. I Teil,
Abt. la: iii (1900).
Order Coccogonales Atkinson 1903.
Orders Chroococcales and Entophysalidales Geitler in Pascher et al. Siisswasserfl.
Deutschland 12: 52, 120 (1925).
Cells solitary or colonial, not filamentous, never flagellate; mostly of blue-green
color and living by photosynthesis.
Kirchner (in Engler and Prantl, 1898) placed here two families, Chroococcaceac
and Chamaesiphonaceae, but the second belongs to the following order. A proper
second family includes the colorless organisms of genus Achromatium.
Family 1. Chroococcacea [Chroococcaceac] (Nageli) Rabenhorst Kryptog.-Fl.
Sachsen 1:69 (1863). Order Chroococcaceac Nageli Gatt. einzell. Alg. 44 (1849).
Unicellular or colonial blue-green algae. Chroococcus, Gloeocapsa, Merismopedia,
Coelosphaerium, Gomphosphaeria, etc., occur as plankton or as masses on damp
surfaces or the bottoms of bodies of water. Certain species occur as symbionts or
parasites within the cells of the green algae Glaucocystis and Gloeochaete. The re-
sulting bodies, having the color of blue-green algae with the structure of green
algae, resisted classification until Geitler (1923) explained their nature.
Family 2. Achromatiacea [Achromatiaceae] Buchanan. Cells solitary, large, ellip-
soidal, without flagella, non-pigmented; protoplasm alveolar, with or without
granules of sulfur in the alveoli. Half a dozen species have been described; Bersa
(1920) was probably correct in reducing all to the original one, Achromatium
oxaliferum Schewiakoff. It occurs on mud under still waters rich in organic matter.
Order 4. Gloiophycea [Gloiophyceae] Rabenhorst Kryptog.-Fl. Sachsen 1 : 56
(1863).
Tribe Nostochineae {Hormonogoneae) Thuret in Ann. Sci. Nat. Bot. ser. 6,
1: 377 (1875).
34 ] The Classification of Lower Organisms
Family Hormogoneae Bomet and Flahault in Ann. Sci. Nat. Bot. ser. 7, 3 : 337
(1886).
Subclass Hormogoneae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: iii(1900).
Order Hormogoneae Campbell Univ. Textb. Bot. 84 (1902).
Order Hormogonales Atkinson 1905.
Blue-green algae whose cells divide predominantly in one plane, so that filaments
are produced, together with related colorless organisms.
So far as cell division is strictly in one plane, any branching of the filaments is of
the type called "false": it occurs by breaks in the continuity of the series of cells,
followed by the outgrowth, beside the original series, of the newly formed tips. In
some members of the group, however, the cells are not strictly confined to division in
one plane, with the result that "true" branching is possible. There are a few appar-
ently derived examples in which cell division takes place freely in all planes.
Many of these algae produce spores of the type called arthrospores by the direct
conversion of normal cells into thick-walled resting spores. Many (almost but not
quite exactly the same ones which produce arthrospores) produce peculiarly differ-
entiated cells called heterocysts (the word means "different cells"). These are en-
larged thick-walled cells with colorless contents; their most obvious function is to
furnish breaking points for the filaments. They are believed to be variants of the
arthrospores; they have been seen to germinate and give rise to normal filaments.
Ten families may be distinguished as follows:
1. Cells dividing strictly in one plane; branch-
ing none or of the "false" type.
2. The filaments not branching nor taper-
ing nor producing spores or heterocysts.
3. Filaments elongate.
4. Pigmented, blue-green Family 1. Oscillatoriacea.
4. Colorless organisms accumu-
lating sulfur Family 2. Beggiatoacea.
3. Filaments reduced to single cells
which reproduce by budding Family 3. Chamaesiphonacea.
2. Filaments branching or tapering, or pro-
ducing spores or heterocysts, or showing
several of these characters.
3. Filaments not tapering.
4. Filaments not branching Family 4. Nostocacea.
4. Filaments branching.
5. Blue-green algae mostly
producing heterocysts Family 5. Scytonematacea.
5. Minute colorless filaments
without heterocysts Family 6. Chlamydotrichacea.
3. Filaments tapering Family 7. Rivulariacea.
1. Cells dividing in more planes than one, usual-
ly after a preliminary filamentous phase.
2. Pigmented, blue-green.
3. Producing extensive filaments with
heterocysts Family 8. Sirosiphonacea.
Kingdom Mychota [35
3. Filaments more or less reduced, re-
producing by minute spores (gon-
idia) formed by repeated division
in all planes Family 9. Pleurocapsacea.
2. Colorless; filamentous, reproducing by
gonidia Family 10. Crenotrichacea.
Family 1. Oscillatoriacea [Oscillatoriaceae] Harvey 1858. Blue-green algae con-
sisting of unbranched filaments, not tapering, without spores or heterocysts; mostly
actively motile by mechanisms as yet unknown. In the commonest genus, Oscillatoria,
the filaments are straight and lack sheaths. Lyngbya and Phormidium produce
sheathed filaments, in the latter genus very slender. Microculeus and Hydrocoleum
have more than one filament in each sheath. In Arthrospira and Spirulina the fila-
ments are coiled; those of Spirulina are not visibly septate, and are said to be uni-
cellular.
Family 2. Beggiatoacea [Beggiatoaceae] Migula 1895. Beggiatoa Trevisan includes
slender colorless filaments, actively writhing, containing granules of sulfur, found
in foul waters and sulfur springs. The species were originally included in Oscillatoria.
Winogradsky (1887) showed that they live by chemosyn thesis, and discovered the
related genera Thiothrix and Thioploca. From the time of these discoveries, these
organisms were construed as bacteria of an order Thiobacteria. Under the current
hypri thesis that chemosynthesis is a derived character, we are free to believe that the
position originally assigned to the species of Beggiatoa was the natural one.
Family 3. Chamaesiphonacea [Chamaesiphonaceae] Borzi 1882. Order Chamaesi-
pkonales Smith Freshw. Alg. 74 (1933). The only genus is Chamaesiphon, minute
organisms epiphytic on freshwater plants. The ellipsoid cells are attached at one
end and are enclosed in tenuous sheaths. They reproduce by transverse division, which
cuts loose small cells from the free ends. By the time two or three such cells are
produced, the sheath is ruptured at the free end, and the small cells drift away to
repi educe the organism elsewhere.
Family 4. Nostocacea [Nostocaceae] (Nageli) Rabenhorst Kryptog.-Fl. Sachsen
1: 95 (1863). Order Nostocaceae Nageli 1847. Of this family the most familiar genus
is Nostoc, seen as gelatinous bodies, usually globular, green, blue-green, yellow, or
brown, of sizes from barely visible to the naked eye up to 10 cm. or more in diameter,
in fresh water or on damp earth. Under the microscope, these bodies or colonies are
seen to consist of myriad crooked and tangled filaments of bead-like cells imbedded
in a gelatinous matrix. Heterocysts are always, and spores usually, present.
If in water one finds filaments of much the same structure as those of Nostoc,
but comparatively short, straight, and free or at least not in definite colonies, these
represent the genus Anabaena. Filaments floating on water, with cylindrical spores
not confined to the ends of the filaments, are Aphanizomenon. Filaments each with
one heterocyst and one spore at one end are Cylindrospermum.
Family 5. Scytonematacea [Scytonemataceae] Rabenhorst op. cit. 106. Members
of this family produce heavily sheathed filaments like those of Lyngbya, with the
difference that heterocysts are usually present. The multiplication of the cells of a
filament may produce the result that the cell next to a heterocyst is driven out of line
and forced obliquely through the sheath. With further growth, the file of cells ending
in one which was forced out of line may appear to be the main axis of a system of
branches, while the original summit of the filament appears to be a lateral branch.
The description of "false" branching thus given applies particularly to Tolypothrix.
36 ] The Classification of Lower Organisms
In Scytonema, the pressure of multiplying cells causes waves of the filament to break
laterally through the sheath and produce branches in pairs. Plectonema branches
like Tolypothrix but has no heterocysts.
Family 6. Chlamydotrichacea [Chlamydotrichaceae] Pribram in Jour. Bact. 18:
377 (1929). Aquatic organisms consisting of colorless cylindrical cells in sheathed
filaments, without heterocysts but exhibiting false branching, the sheaths of young
filaments thin and colorless, those of older ones thick and yellow to brown. Chlamydo-
thrix ochracea Migula was intended as a new name for Leptothrix ochracea Kiitzing,
but the entity to which it is believed to apply is totally different from the one to which
the latter name was applied above. Chlamydothrix is a filament of definite cells
about 1 ^ in diameter. The only other definitely characterized species of this family is
Clonothrix fusca Roze, the cells about 2^ in diameter, those near the tips of the fila-
ments dividing repeatedly (always in one plane) to produce spherical non-motile
gonidia (Kolk, 1.938).
Family 7. Fdvulariacea [Rivulariaceae] Rabenhorst op. cit. 101. The filaments
include heterocysts and exhibit the false branching of Tolypothrix; the outgrowth
of the filament below each heterocyst gives to the original terminal part the appear-
ance of a branch of which the heterocyst is the basal cell. The ends of the filaments
become attenuate and colorless. In Calothrix the filaments are mostly solitary; in
other genera they remain together in gelatinous colonies. Rivularia is without spores;
in Glocotrichia there is a large cylindrical spore next to each heterocyst.
Family 8. Sirosiphonacea [Sirosiphonaceae] Rabenhorst op. cit. 114. Family
Stigonemataceae Kirchner 1898. This family takes its name from the ancient generic
name Sirosiphon Kiitzing 1843, which turned out to be identical with Stigonema
Agardh 1824. The cells divide at first in one plane and produce filaments. Presently
they exhibit a capacity to divide in other planes, and may produce true branches or
multiseriate filaments or both. Heterocysts and spc^es are generally produced.
Family 9. Pleurocapsacea [Pleurocapsaceae] Geitler in Pascher et al. Siisswasser-
Fl. Deutschland 12: 124 (1925). This group was formerly included in Chamaesi-
phonacea, but it appears probable that Chamae siphon is related to Oscillatoria, and
the present group to Stigonema. Most of the Pleurocapsacea are marine, epiphytic
on seaweeds. Their apparently typical behavior, as exemplified by Hyella and
Radaisia, consists of the production of branching filaments whose terminal eel's be-
come enlarged, after which their contents undergo division in many planes to produce
numerous minute spores called gonidia. In Pleurocapsa and Xenococcus there is no
filamentous phase; the gonidium gives rise to a cluster of cells all of which produce
gonidia. In Dermocarpa the gonidium gives rise to a single vegetative cell which
divides only to produce gonidia.
Family 10. Crenotrichacea [Crenotrichaceae] Hansgirg. This family includes the
single known species Crenothrix polyspora Cohn, one of the traditional iron bacteria.
There is every appearance that it is a colorless variant of the Pleurocapsacea. A germi-
nating gonidium gives rise to an unbranched filament of cells, about 2^ in diameter,
in a sheath which is at first thin and colorless, later becoming thicker and discolored
by ferric oxide. Some cells may burst from the free end of the sheath as macrogonidia.
Others may begin to divide lengthwise. These may at first grow before re-dividing,
and may swell the sheath to a fusiform or trumpet-like shape. By further division
they produce numerous microgonidia, which may sift out of the sheath or be re-
leased by its decay.
Such are the Mychota, the organisms which may properly be characterized as
lacking nuclei.
Chapter IV
KINGDOM PROTOCTISTA
Kingdom !l. PROTOCTISTA Hogg
Regne Psycho diaire, Psychodies, Bory de Saint Vincent Diet. Class Hist. Nat. 8:
246 (1825), 14: 329 (1828).
Kingdom Protozoa Owen Palaeontology 5 (1860), not class Protozoa Goldfuss
(1818).
Regnum Primigenium seu Protoctista Hogg in Edinburgh New Philos. Jour. n.s.
12: 223 (1860).
Kingdom Acrita or Protozoa Owen Palaeontology ed 2: 6 (1861).
Kingdom Primalia Wilson and Cassin in Proc. Acad. Nat. Sci. Philadelphia 1863:
117 (1864).
Kingdom Protista Haeckel Gen. Morph. 2: xix (1866).
Kingdom Protobionta Rothmaler in Biol. Centralbl. 67: 243 (1948).
Nucleate organisms other than Plantae and Animalia: the marine algae and the
fungi and protozoa. Amiba diffluens may be construed as the standard.
The name Protista, of Haeckel, is the most familiar among those which have been
applied to the kingdom here to be discussed, but it is not the earliest. Among fol-
lowers of Cuvier, the animal kingdom consisted necessarily of four branches. Presum-
ably, it was this tradition that induced Owen to refer the Infusoria and Amorphozoa
(sponges) to a separate kingdom, which he called Protozoa. A year later, Owen pub-
lished an alternative name for this kingdom; but Hogg had already published modi-
fications of two of Owen's names, Protoctista and Amorphoctista(KTi^co,to establish,
create), for the reason that names in -zoa appeared inappropiiate to groups excluded
from the animal kingdom.
The limits here given to the kingdom Protoctista were proposed by the present
author (1938, 1947). They have been accepted, with exception in a single significant
point, by Barkley (1939, 1949) and Rothmaler (1948).
It is assumed that the evolutionary origin of the Protoctista consisted of the evolu-
tionary origin of the nucleus, and that all nuclei are essentially the same thing. Kofoid
(1923) insisted that enduringly viable nuclei originate among protozoa, as among
plants and animals, regularly by mitosis, never by binary or multipe fragmentation,
nor by aggregation of stainable granules. He did not recognize the nucleus as essen-
tially a device for sexual reproduction. Several considerable groups of protozoa, how-
ever, which Kofoid listed as not known to reproduce sexually, have been found to
do so. Here, then, it is maintained that all nuclei, in this kingdom as among plants
and animals, are the same thing; and that the nucleus is essentially a device for sexual
reproduction, that is, for processes of reproduction which involve always one act of
meiosis and one of karyogamy, and which produce Mendelian heredity as an effect.
Photosynthesis is believed to have evolved only cnce. As it occurs both among non-
nucleate and nucleate organisms, the nucleus is believed to have evolved in organisms
living by this function. The closest approach between non-nucleate and nucleate or-
ganisms is believed to be between the blue-green algae and the primitive red algae
(Smith, 1933; Tilden, 1933). Thus it appears that the original nucleate organisms
were not capable of swimming by means of flagella. Flagella appear to have evolved
in unicellular nucleate photosynthetic organisms as a device for dissemination (Bes-
38 ] The Classification of Lower Organisms
sey, 1905). The flagella of nucleate organisms are not homologous with those of
bacteria; they are much larger and of much more complicated structure.
The origin of flagella was apparently associated with a simplification of the system
of photosynthetic pigments, by the loss of chromoproteins, leaving systems of chloro-
phylls and carotinoids. The association of these two courses of evolution may have
been merely coincidental; Tilden suggested the idea that the loss of chromoproteins
may have been occasioned by increasing illumination of the waters of the face of the
earth.
Organisms of the body type of solitary walled cells, having chlorophylls and caro-
tinoid pigments but not chromoproteins, and producing flagellate reproductive cells,
appear to have undergone radiating evolution, producing a wide variety of types of
organisms, distinguished by different specific chlorophylls and carotinoids, different
types of flagella, and different specific metabolic products. The types of flagella oc-
curring in nucleate organisms are here particularly to be noted.
Loeffler (1889), in the original publication of the standard method of staining the
flagella of bacteria, remarked that he had applied this method also to certain larger
organisms. He found that the flagellum of Manas bears numerous lateral appendages,
and that the cilia of a certain infusorian bear solitary terminal appendages. Loeffler's
method is difficult, and has not been much used. Fischer (1894) used it and coined
terms, Flimmergeisseln and Peitschengeisseln, designating structures of the respective
types seen by Loeffler. Petersen (1929), having applied Loeffler's method to a reason-
able variety of flagellates, introduced refinements of terminology. Flagella of the
type of the larger flagellum of Monas (the organism bears also a minute simple flagel-
lum) became allseitswendige Flimynergeisseln; those of Euglena, which bear a single
file of appengages, became einseitswendige Flimmergeisseln.
Deflandre ( 1934) devised a different method for seeing the appendages on flagella,
and substituted, for the Teutonisms just quoted, French terms based on Greek. These
may be Anglicised as follows. ( 1 ) The acroneme flagellum bears a single terminal
appendage. The flagellum without appendages is said to be simple; so far as it ap-
pears among nucleate organisms, it appears to be a variant of the acroneme type.
(2) The pantoneme flagellum bears appendages on all sides. (3) The pantacroneme
flagellum bears both terminal and lateral appendages. It is a rarity, known only in
the collared monads, and may be supposed to be a variant of the pantoneme type.
(4) The stichoneme flagellum bears a single file of appendages.
The point in which Barkley and Rothmaler take exception to the limits here given
to kingdom Protoctista is this, that they include in this kingdom the green algae. In
the present work, scant attention is given to organisms whose plastids are bright
green, containing chlorophylls a and b, carotin, and xanthophyll, and no other pig-
ments; whose motile stages have acroneme flagella, more than one (usually two),
and equal; and which produce essentially pure cellulose, true starch, and sucrose.
These organisms represent the undoubted evolutionary origin of the higher plants;
a classification which attempts to represent nature includes them necessarily in the
plant kingdom.
Rothmaler set up a system of only four phyla, being the red organisms, basically
without flagella; those which are typically yellow to brown, having pantoneme flagel-
la; those with acroneme flagella, including the green algae; and the euglcnid group,
which have stichoneme flagella. The non-pigmented Protoctista were distributed
among these groups. The system appears unsound by the fact that large blocks of
non-pigmented organisms are placed where only portions of them belong.
Kingdom Protoctista [ 39
In the present work, a less symmetrical system of phyla is offered. Its basis is an
ingenuous system of red algae, brown algae, fungi, and the four traditional groups of
protozoa; this has been radically modified in view of the great accumulation of
knowledge subsequent to the formulation of these groups. The phylum Pyrrhophyta
as here limited is tentative; the phylum Protoplasta, marked only by negative char-
acters, amounts to a dumping ground for groups whose relationships are altogether
obscure.
1. Living by photosynthesis, which takes place
in plastids containing red or blue chromo-
protein pigments; never producing flagellate
cells Phylum 1. Rhodophyta.
1. Without chromoprotein pigments.
2. Typically living by photosynthesis,
brown, yellow, or green in color.
3. Producing flagellate cells each with
one pantoneme or pantacroneme
flagellum, often with additional
acroneme flagella Phylum 2. Phaeophyta.
3. Producing flagellate cells whose fla-
gella are never pantoneme or pan-
tacroneme, often stichoneme Phylum 3. Pyrrhophyta.
2. Dependent; motile cells with acroneme
flagella or cilia, or amoeboid, or none.
3. Not producing cilia, i. e., structures
of the nature of acroneme flagella,
numerous and widely distributed
on the surfaces of the cells.
4. Cells walled in the vegetative
condition.
5. Producing motile cells
with single posterior fla-
gella; bodies mostly with
tapering rhizoids Phylum 4. Opisthokonta.
5. Producing no motile cells;
bodies filamentous Phylum 5. Inophyta.
4. Cells not walled in the vegeta-
tive condition.
5. Mostly predatory, flagel-
late or amoeboid or with
flagellate or amoeboid stages Phylum 6. Protoplasta.
5. Parasitic in animals, pro-
ducing flagellate cells only
as rare exceptions Phylum 7. Fungilli.
3. With cilia .Phylum 8. Ciuophora.
Chapter V
PHYLUM RHODOPHYTA
Phylum 1. RHODOPHYTA Wettstein
Order Floridees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 115 ( 1813) .
Florideae C. Agardh Synops. Alg. Scand. xiii (1817).
Order pLORroEAE C. Agardh Syst. Alg. xxxiii (1824).
Division (of order Algae) Rhodospermeae Harvey in Mackay Fl. Hibern. 160
(1836).
Class Heterocarpeae Kiitzing Phyc. Gen. 369 (1843).
Class Florideae J. Agardh Sp. Alg. 1: v (1848).
Rhodophyceae Ruprecht in Middendorff Sibirische Reise 1, Part 2: 200 (1851).
Stamm Florideae Haeckel Gen. Morph. 2: xxxiv (1866).
Phylum RHODOPHYTA Wettstein Handb. syst. Bot. 1: 182 (1901).
Division Rhodophyceae Engler Syllab. ed 3: 18 fl903).
Phylum Carpophyceae Bessey in Univ. Nebraska Studies 7: 291 (1907).
Phylum Rhodophycophyta Papenfuss in Bull. Torrey Bot. Club 73: 218 (1946).
Definitely nucleate organisms {Porphyridium and Prasiola doubtfully so); with
few exceptions living by photosynthetic processes involving red and blue pigments
(phycocyanin and phycoerythrin) as well as green and yellow (chlorophylls a and d
and carotinoids); not producing true starch, and producing cellulose only in small
quantity, the cells walled chiefly with modified carbohydrates which tend to become
gelatinous; never producing flagellum-bearing cells, but sometimes producing cells
which move in water without the use of definite organelles.
Tilden (1933) and Smith (1933) are authority for placing the red algae next to
the blue-green algae, thus suggesting the inference that they include the most primi-
tive of nucleate organisms. The resemblances between blue-green and red algae
are in the following points. Both groups possess, along with the chlorophylls and
carotinoids usual in photosynthetic organisms, other pigments, both blue and red. To
these pigments as found in both groups, the same names, phycocyanin and phycoery-
thrin, are applied; they are not, however, the same chemical species (Kylin, 1930).
Neither group produces true starch; carbohydrate is stored as substances of the
general nature of dextrin or glycogen (occuring in the red algae as solid granules
called floridcan starch). Both groups produce cellulose only in scant quantities
(Miwa, 1940; Kylin, 1943); the cell walls consist chiefly of materials, of the general
nature of carbohydrates, which tend to become gelatinous. They share the negative
character of never producing flagella, and the positive one of producing cells which
call move actively upon surfaces, without motor organelles, by a mechanism as yet
unknown (Roscnvinge, 1927).
The phylum is divisible into two classes:
1. Cells of most examples each with one central
plastid, without protoplasmic interconnec-
tions, in aggregates of indefinite extent or or-
ganized as filaments or thalli with intercalary
growth; zygotes producing spores directly by
division Class 1. Bangialea.
Phylum Rhodophyta [41
1. Cells with protoplasmic interconnections,
containing except in the lowest examples sev-
eral parietal plastids, organized as filaments
with apical growth, the filaments usually
massed as thalloid bodies; zygotes giving rise
to spores indirectly Class 2. Heterocarpea.
Class 1. BANGIALEA (Engler) Wettstein
Subclass Bangioideae de Toni Sylloge Algarum 4: 4 (1897).
Subclass Bangiales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2 :
ix (1897).
Class Bangiales Wettstein Handb. syst. Bot. 1: 187 (1901).
Class Bangioideae and orders Bangiales and Rhodochaetales Bessey in Univ. Ne-
braska Studies 7: 291 (1907).
Class Bangieae SchaflFner in Ohio Naturalist 9: 448 (1909).
Protoflorideae Rosenvinge in Mem. Acad. Roy. Sci. Lett. Danemark, ser. 7,
Sciences 7: 55 (1909).
Abtheilung (of Stamm Rhodophyta) Bangiineae Pascher in Beih. bot. Centralbl.
48, Abt. 2: 328 (1931).
Subclass Protoflorideae Smith Freshw. Algae 120 (1933).
Red algae (exceptionally green or of other colors), the cells with solitary central
plastids (exceptionally with multiple parietal plastids), lacking protoplasmic inter-
connections, in irregular colonial masses or forming filaments or thalli with intercalary
growth; the zygote produced in sexual reproduction dividing to produce spores
directly.
The group is of one order, five families, about fifteen genera; the number of
known species is about eighty.
Order Bangiacea [Bangiaceae] Nageli 1847.
Characters of the class.
1. Cells forming irregular aggregates Family 1. Porphyridiacea.
1. Cells forming filaments or thalli.
2. Vegetative cells becoming spores with-
out dividing Family 2. Rhodochaetacea.
2. Vegetative cells undergoing division to
produce spores.
3. Organisms red, purplish, etc Family 3. Porphyrea.
3. Organisms green Family 4. Schizogoniacea.
2. Spores formed solitary in special cells Family 5. Compsopogonacea.
Family 1. Porphyridiacea [Porphyridiaceae] Kylin in Kungl. Fysiog. Sallsk.
Forhandl. 7, no. 10: 4 (1937). Order Porphyridiales Kylin 1. c. The only well known
species is Porphyridium cruentum (C. Agardh) Nageli (1849). It is widely dis-
tributed in damp climates, forming extensive red patches like blood on damp earth or
stone. The spherical cells are reported as varying widely in diameter (5-24^), and
Geitler (1932) and Kylin (1937) have distinguished additional species.
Porphyridium has been classified among blue-green, red, and green algae. Lewis
and Zirkle (1920) found in each cell a central red plastid, occupying most of its
volume, and having rays extending to the cell membrane. Within the plastid there is
42]
The Classification of Lower Organisms
Fig. 5. — a, Porphyra laciniata, thallus x 1/2. b-g, Porphyra tenera after Ishikawa
(1921); b, cells; c, cell dividing to produce sperms; d, sperms; e, fertilization;
f, "carpospores," i.e., cells produced by division of the zygote; g, stages of nuclear
division x 2,000. h, i, Porphyra umbilicaris after Dangeard ( 1927); h, fertilization;
i, stages of nuclear division x 2,000. All figures x 1,000 except as noted.
Phylum Rhodophyta L 43
a moderately large stainable granule; outside the plastid, a single additional granule
can usually be found. When a cell is to divide, the granules break up into consid-
rable numljers of smaller ones, some of which become organized as a system of strands
forming an irregular network on the surface of the plastid. The protoplast, the
network, and the plastid undergo constriction; the processes by which the daughter
cells return to the original structure were not clearly seen. Interpretation of these
observations is difficult. It is possible that the granule outside of the plastid is a
nucleus of the type of those which have been observed in Bangia and Porphyra.
Family 2. Rhodochaetacea [Rhodochaetaceae] Schmitz in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 2: 317 (1896). Family Goniotrichaceae Smith Freshw.
Algae 121 (1933). Branching filaments, sometimes becoming multiseriate by length-
wise division, the vegetative cells capable of escaping and functioning as spores.
Sexual reproduction unknown. Asterocystis, uncommon, in fresh water; the remain-
ing genera marine, epiphytic on other algae. Goniotrichum. Rhodochaete and Gonio-
trichopsis, the cells with numerous plastids.
Family 3. Porphyrea [Porphyreae] Kutzing (1843). Family Porphyraceae Raben-
horst 1868. Family Bangiaceae (Nageli) Schmitz (in Engler and Prantl, 1896).
Filaments or thalli of a red or purple color; the cells, in producing spores, may re-
lease their protoplasts as wholes or may undergo division into many. Rosenvinge
(1927) observed the active motion of these spores.
The most important genus is Porphyra; the individuals are thalli up to several
centimeters in diameter, on rocks or other algae in ocean water along coasts. They
are called purple lavers, tsu'ai, amanori; they are used as food, for making soup or in
condiments, and are extensively cultivated in Japan (Tseng, 1944). Bangia is either
freshwater or marine; in structure it differs from Porphyra in having filementous
bodies, uniseriate or pluriseriate.
During nuclear division in Porphyra tenera as described by Ishikawa (1921), polar
appendages form at both ends of the nucleus, which becomes elongate and appears
to consist of three strands. The strands break transversely, and each set of three fuses
into a mass. Dangeard (1927), dealing with Porphyra umbilicaris and Bangia fusco-
purpurea, observed nuclei 5^ in diameter, each consisting of a karyosome, that is, a
mass of chromatin, lying in a clear space surrounded by a membrane. In mitosis, the
membrane and the unstained matter disappear. Polar appendages grow out from the
karyosome, and their tips become cut off as granules which may be regarded as cen-
trosomes. The remainder of the karysome becomes organized as two masses, evidently
chromosomes, connected to the centrosomes by fibers. Each chromosome divides into
two; the daughter chromosomes move to the centrosomes and fuse with them to form
karyosomes about which new membranes appear. This description represents a defi-
nite, if primitive, process of mitosis.
Sexual reproduction, here where we first encounter it, involves differentiated ga-
metes. Naked sperms, indistinguishable from spores, move to the surface of other
cells which function as eggs. A strand of protoplasm grows through the gelatinous
wall of the egg from the sperm to the egg protoplast, and the protoplast of the sperm
migrates through the passage thus formed. The zygote divides two or three times,
producing spores. During the first two divisions, the two masses of chromatin which
appear are somewhat different in appearance from the vegetative chromosomes
(Dangeard, op. cit.); it may be supposed that these masses are tetrads and diads, and
that the divisions are meiotic. Evidently, this is a life cycle of the primitive type,
in which all cells except the zygotes are haploid.
44 ] The Classification of Lower Organisms
Family 4. Schizogoniacea [Schizogoniaceae] Chodat. Family Prasiolaceae West.
Family Blastosporaceae Wille. Filamemous or thallose algae, freshwater or marine,
of the structure of Porphyrea, but of a green color; sexual reproduction unknown.
Kylin (1930) found the pigmentation to be that of green algae rather than of red.
Copeland (1955) was unable to discern nuclei. The sole genus Prasiola {Schizo-
gonium represents a stage of development) is of about fifteen species. Setchell and
Gardner (1920) and Ishikawa (1921) suggested the place in Bangiacea here given to
this group.
Family 5. Compsopogonacea [Compsopogonaceae] Schmitz in Engler and Prantl
Nat. Pflenzenfam. I Teil, Abt. 2: 318 (1896). Family Erythrotrichiaceae Smith
Freshw. Algae 122 (1933). Filaments, unbranched or branched, uniseriate or pluri-
seriate, or thalli. Spore-formation is accomplished by the division of a vegetative cell,
by an oblique wall, into two unequal cells; the protoplast of the smaller is released as
a spore. Rosenvinge observed the spores of Erythrotrichia cornea to move as far as
140[i per minute. Sexual reproduction is much as in Porphyrea. Erythrotrichia.
Erythrocladia. Compsopogon, in fresh water, the cells with numerous parietal plastids.
Class 2. HETEROCARPEA Kutzing
Class Heterocarpeae Kiitzing Phyc. Gen. 369 (1843).
Class Florideae (C. Agardh) J. Agardh Sp. Alg. 1 : v ( 1848).
Subclass Florideae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2:
ix (1897).
Subclass Euflorideae de Toni Sylloge Algarum 4: 4 (1897).
Abtheilung (of Stamm Rhodophyta) Floridineae Pascher in Beih. bot. Centralbl.
48, Abt." 2: 328 (1931).
As this is the type group of phylum Rhodophyta, most of the synonymy of that
name applies to this one also.
Red algae whose bodies consist essentially of filaments growing apically, the cells
with protoplasmic interconnections, the plastids (except in some of the lowest ex-
amples) of the form of multiple parietal disks; the filaments commonly compacted
into cylindrical or thallose bodies; zygotes not dividing to form spores directly, pro-
ducing spores by budding or indirectly by processes of growth of various degrees of
complexity.
In undertaking to describe the varied, and often highly complicated, reproductive
processes of the typical red algae, one notes that these organisms occur as haploid
individuals, and that the majority occur as distinct male and female haploid individ-
uals. Sperms (commonly called spermatia) are minute naked protoplasts released
from small cells commonly occurring in patches on the surfaces of thalli. The egg
is called a carpogonium (Schmitz, 1883). It is the terminal cell of a specialized fila-
ment, the carpogonial filament, and bears a filiform terminal extension, the tri-
chogyne (Bornet and Thuret, 1867), whose function is to receive the sperms. The
cell, often diflferentiatcd, from which the carpogonial filament grows, is the support-
ing cell [Trugzelle).
In the more primitive members of the class, the zygote gives rise by budding to a
mass of cells called the cystocarp. The cells of the cystocarp release their protoplasts
as spores called carpospores. These on germination produce haploid individuals like
the original ones. The zygote nucleus is the only diploid nucleus in the life cycle; its
first divisions arc meiotic.
Phylum Rhodophyta
[45
In more advanced examples, the first step of development after fertilization con-
sists of the establishment of protoplasmic contact between the zygote and other cells.
These may be adjacent cells, reached directly, or distant cells, reached by the out-
growth of connecting filaments from the zygotes. In the generality of the group, the
cells with which contact is made give rise to cystocarps producing carpospores; in
this situation, the cells in question are called auxiliary cells. In some examples, the
connecting filaments, after making contact with cells called nurse cells, themselves
give rise to the cystocarps. The carpospores, in all of these more advanced examples,
give rise to diploid individuals. The diploid individuals are of the same vegetative
'ymW''^- :^^>^^
^■^y
i^
•«!»
Fig. 6 — Nuclear phenomena in Polysiphonia violacea after Yamanouchi (1906).
a, b, c. Stages of mitosis, d, e. Stages of homeotypic division.
structure as the haploid individuals, but do not produce spermatia, carpogonia, or
cystocarps. Certain cells, commonly scattered and imbedded in the body, produce
sets of four spores which are accordingly called tetraspores; these give rise to haploid
individuals.
This account means that these algae occur typically in somata of four types: male
and female haploid individuals; cystocarps, being a preliminary, parasitic, multipli-
cative phase of the diploid stage (Janet named this stage the carposporophyte; Drew,
1954); and free-living diploid individuals, reproducing by tetraspores. The produc-
tion of carpospores and tetraspores by different individuals of identical vegetative
structure explains the oldest name applied to this class, namely Heterocarpea.
Understanding of the life cycle of typical Heterocarpea has been reached only by
much labor and after a certain amount of confusion. The first significant observations
were by Bornet and Thuret (1867). Schmitz (1883) showed that the zygote makes
protoplasmic contact with other cells. He supposed that the contact of the zygote
with an auxiliary cell is a second sexual fusion {Copulation) following upon proper
46 ] The Classification of Lower Organisms
fertilization. Oltmanns (1898) disproved this: he showed that the nuclei of auxiliary
cells are inert, and that the nuclei of carpospores are derived entirely from zygote
nuclei. Yamanouchi (1906) showed that the chromosome number of carposporic
individuals of Polysiphonia violacea is 10, and that that of tetrasporic individuals is
20; and reported much more of the cytology. Centrosomes appear de novo during the
earlier stages of mitosis, and fade out and disappear during the later stages. The
mitotic spindle is formed, and the chromosomes take their place upon it, within an
intact nuclear membrane, which fades out in later stages. In meiosis, which produces
the nuclei of tetraspores, the tetrads and diads divide within the original nuclear
membrane, which becomes tetrahedrally lobcd, and then disappears except where
the haploid groups of chromosomes lie against it, with the result that the membranes
of the tetraspore nuclei are partly old and partly new.
There are some 2500 species of Heterocarpea, including comparatively few in
fresh water, but the majority of the marine algae. Many of them are beautiful; their
variety and beauty contribute to the pleasure which people find on coasts. Exper-
ienced naturalists can identify many genera by gross structure, but the systems of
orders and families based on gross structure, such as those of Kiitzing (1843) and J.
Agardh (1851-1863), have been found artificial and abandoned. A proper respect
for the principles of nomenclature makes it necessary, however, to apply many of the
names used in these systems. Schmitz applied his morphological studies to a classifica-
tion of the typical red algae as four groups ( 1889) ; Engler ( 1897) made these groups
definitely orders. Subsequent scholars have found this system sound in principle, but
have found it necessary, on the basis of studies of additional examples (for example,
by Kylin, 1923, 1924, 1925, 1928, 1930, 1932; Papenfuss, 1944; Sjostedt, 1926;
Svedelius, 1942) radically to rearrange the families and genera. At least four orders
in addition to those of Engler have been proposed but reductions have decreased
the number currently recognized to six.
The following key to the orders is a rather considerable modification of those pub-
lished by Kylin (1932) and Smith (1944).
l.All free-living individuals haploid; tetra-
spores not produced, or produced as carpospores. . Order 1. Cryptospermea.
1. Free-living individuals of two types, the one
producing gametes (the zygotes giving rise
to carpospores), the other producing tetraspores.
2. Without specialized auxiliary cells or
nurse cells, the lower cells of the carpo-
gonial filaments, or normal vegetative
cells, serving as auxiliary cells Order 2. Sphaerococcoidea.
2. With specialized nurse cells, the carpo-
spores produced from filaments which
have made contact with these Order 3. Gelidialea.
2. With specialized auxiliary cells from
which the carpogonia develop.
3. The auxiliary cells being intercalary
cells in specialized filaments homol-
ogous with the carpogonial filaments. . . . Order 4. Furcellariea.
3. The auxiliary cells terminal in fila-
ments which grow from the support-
ing cells of the carpogonial fila-
ments before fertilization Order 5. Coeloblastea.
Phylum Rhodophyta [ 47
3. The auxiliary cells originating after
fertilization as branches of the sup-
porting cells of the carpogonial
filaments Order 6. FLORroEA.
Order 1. Cryptospermea [Cryptospermeae] Kiitzing Phyc. Gen. 321 (1843).
Order Periblasteae Kutzing op. cit. 387, in part.
Orders H elmint hoc lade ae J. Agardh Sp. Alg. 2: 410 (1851), Chaetangieae op.
cit. 456 (1851), and Wrangelieae op. cit. 701 (1863).
Order Batrachospermaceae'R.ahtnhovstKxy^X.og.-Yl.^dichstn 1: 278 (1863).
Nemalioninae Schmitz in Flora 72: 438 (1889).
Order Nemalionales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
2: ix (1897).
Heterocarpea normally without diploid bodies, the carpogonium arising from the
zygote or from an adjacent cell serving as an auxiliary cell, the carpospores produc-
ing haploid bodies like the original ones. Certain genera which are exceptional to
these characters are noted below. Batrachospermum may be regarded as the standard
genus.
In all recent literature, this order is called Nemalionales. Eight families are rec-
ognized. The forms consisting of mere filaments, Acrochaetium, Rhodochorton, and
others, are family Acrochaetiacea [Acrochaetiaceae] Fritsch (Family Chantransi-
aceae Auctt., but Chantransia DC. as originally published included no members of
this family; Papenfuss, 1945). In the remainder of the order, the filaments are
differentiated, or, with or without differentiation, organized as bodies of definite
form, simply cylindrical, branched, or flattened. Fresh-water examples (the only
fresh-water Heterocarpea) include Batrachospermum, Lemanea, and Thorea. These
organisms are not red, but bluish, green, or brown. Marine examples include Nemalion
and Cumagloia.
In Liagora tetrasporifera and certain other species tetraspores are produced in the
place of carpospores. Within this genus, then, there has been a change in the time of
meiosis (which could be established, presumably, by a single mutation) from im-
mediately after fertilization to the end of the cystocarp stage.
Galaxaura is a genus of tropical marine algae which are calcified, which is to say
that they deposit much calcium carbonate in the tissues; they were originally classi-
fied as corals. They have distinct sexual and tetrasporic stages. Svedelius (1942) as-
certained their life cycle. Carpospore-bearing filaments arise both from the zygote
and from other cells, previously undifferentiated, which serve as auxiliary cells. The
genus has the structure of the present order, and is to be placed here, in spite of ex-
hibiting in unspecialized form the life cycle of the following orders.
Order 2. Sphaerococcoidea [Sphaeroccoideae] J. Agardh Sp. Alg. 2: 577 (1852).
Family Gigartineae Kiitzing (1843).
Orders Gigartineae and Chaetangieae J. Agardh op. cit. 229, 456 (1851).
Gigartininae Schmitz in Flora 72: 440 (1889).
Order Gigartinales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
2: X (1897).
Order Nemastomatales Kylin in Kgl. Fysiog. Sallsk. Handl. n. f. 36, no. 9 : 39
(1925).
Order Sphaerococcales Sjostedt in Kgl. Fysiog. Sallsk. Handl. n. f. 37, no. 4:
75 (1926).
48]
The Classification of Lower Organisms
(Legend on bottom of page 49)
Phylum Rhodophyta [ 49
This order, in all recent literature called Gigartinales, is a numerous and varied
one. The bodies are generally erect; they may be cylindrical or flattened, unbranched
or branched. In some examples, Haliarachnion, Rhodophyllis, Sebdenia, the zygote
sends out extensive filaments, which make contact with unspecialized cells scattered
in the body. In other examples, the zygote makes contact with a lower cell of the
carpogonial filament. In either case, the cells with which contact is made are auxiliary
cells and give rise to cystocarps; these produce carpospores, and the carpospores pro-
duce tetrasporic individuals. Certain species of Phyllophora, Gymnogongrus, and
Ahnfeldtia are exceptional in producing tetraspores in the place of carpospores; these
species have no free-Uving tetrasporic generation. In these organisms, as contrasted
with Liagora tetrasporifera, it is believed that this type of Ufe cycle has been estab-
lished by reduction of a longer one.
Kylin (1932) assigned twenty families to this order. Gracilaria is a minor source
of agar agar. Gigartina mammilosa and Chondrus crispus (Irish moss or carageen)
are well known as yielding a jelly, carageenin, resembling but distinct from agar agar
(Tseng, 1945).
Various abnormal growths on red algae have been found to be parasitic red algae,
almost always on hosts closely related to themselves (Setchell, 1914). To the present
order belong Gardneriella and its host Agardhiella; Plocamiocolax and its host Plo-
camium; Gracilariophila and its host Gracilaria (Wilson, 1910).
Order 3. Gelidialea [Gelidiales] Kylin in Kgl. Svensk. Vetensk.-Akad. Handl.
63, no. 11: 132 (1923).
Family Gelidieae Kiitzing (1843).
Order Gelidieae J. Agardh Sp. Alg. 2: 464 (1851).
Heterocarpea in which the zygote sends out a single elongate filament which makes
contact successively with several chains of nurse cells and gives rise to carpospores;
bodies consisting of branched filaments, the ultimate tips of the lateral branches
compacted into a firm layer covering a branching body, cylindrical or flattened; the
surface adjacent to the masses of carpospores pushed out and punctured by pores
through which the spores escape.
There is a single family Gelidiea [Gelidieae] Kiitzing ( Family Gelidiaceae Schmitz
and Hauptfleisch). Such economic importance as the red algae possess lies chiefly in
Fig. 7 — a, Thallus of Nemalion multifidum x 1. b, c, d^ production of sperms;
beginning of production of carpospores; and cluster of carpospores of Nemalion
multifidum after Bornet and Thuret (1867). e, Thallus of Chondrus crispus x 1.
{, Reproduction of Dudresnaya purpurifera (order Furcellariea or Cryptonemiales)
after Bornet and Thuret, op. cit. The trichogyne, whose free end with attached
sperms is seen above, is irregularly twisted below; it leads to the egg (carpogonium);
connecting filaments, growing from cells below the egg, make contact with auxiliary
cells at the summits of specialized filaments; each auxiliary cell gives rise to a cluster
of carpospores. g, Thallus of Delesseria sinuosa x 1. h^ Longitudinal section of
conceptacle of Polysiphonia nigrescens x 500, after KyUn (1923). The zygote z is
the fourth and terminal cell of the carpogonial filament whose connection with the
supporting cell b is not shown; the auxiliary cell a has grown from the supporting
cell after fertilization.
50 ] The Classification of Lower Organisms
this family, and particularly in the genus Gelidium. It is the chief source of agar agar.
This is the principal material of the cell walls of Gelidium. It is a jelly consisting
essentially of chains of galactose units, and has the property, that having been melted
by heat, it does not again become solid until cooled to a much lower temperature.
Algae containing it have long been used as foods in the orient. Brought into labora-
tory use by Koch, it has become a necessity in routine bacteriological work. The chief
source is Japan.
Kylin construed this order as relatively primitive; but its reproductive processes,
involving specialized nurse cells, appear less primitive than those of the Sphaerococ-
coidea. The production of elongate connecting filaments is shared with certain
examples both of the preceding order and of the following, and the Gelidialea are
probably derived by specialization from one or the other.
Order 4. Furcellariea [Furcellarieae] Greville Alg. Brit. 66 (1830).
Orders Spongocarpeae and Gastrocarpcae Greville op. cit. 68, 157 (1830).
Order Epiblasteae Kiitzing Phyc. Gen. 382 (1843).
Orders Cryptonemeae, Dumontieae, Squamarieae, and Corallineae J. Agardh Sp.
Alg. 2: 'l55, 346, 385 (1851), 506 (1852).
Cryptoneminae Schmitz in Flora 72: 452 (1889).
Order Cryptonemiales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 2: xi (1897).
The individuals are crustose or thallose, the thalli cylindrical or flattened, un-
branched or branched. On the two or three types of individuals of each species, the
reproductive structures may be scattered or clustered on the surfaces or gathered in
specialized pits called conceptacles. The eggs are as usual the terminal cells of spe-
cialized filaments; other filaments, homologous with these but abortive, bear the
auxiliary cells. After fertilization, the zygote may or may not establish connection
with a lower cell of the same filaments. Under either circumstance, it sends out fila-
ments which establish connection with the auxiliary cells, and these send out filaments
which bear the carpospores. In less specialized examples, the filaments growing from
the zygote may extend widely through the body; a single one, branching, may reach
many auxiliary cells.
Kylin ( 1932) placed nine families here.
The family Corallinea [Corallineae] Kiitzing (family Corallinaceae Hauck) is one
of the more specialized. The eggs, and subsequently the carpospores, are clustered in
conceptacles. In each conceptacle the zygotes, the filaments from them, and the
auxiliary cells, unite eventually in a single large multinucleate cell from whose mar-
gins grow the filaments which bear the carpospores. Members of this family have the
property of accumulating and depositing calcareous material, and were originally
classified as corals. In modern usage, the term coral means certain lower animals;
but the coralline algae are associated with them in coral reefs, being indeed, accord-
ing to Setchell (1926) and other authorities, responsible for the building of the reefs.
Fossil coralline algae are known from the Ordovician.
The parasite Callocolax and its host CallophylUs belong to this order; Coreocolax,
belonging to this order, attacks species of order Floridea.
The Furcellariea are a numerous group, rather unspecialized, varied almost to the
extent of a miscellany. They are related to the Sphaerococcoidea, and are believed
to represent the ancestry of the two following orders, and possibly also of the
Gelidialea.
Phylum Rho do phyta [51
Order 5. Coeloblastea [Coeloblasteae] Kutzing Phyc. Gen. 438 (1843).
Order Rhodymenieae J. Agardh Sp. Alg. 2: 337 (1851).
Rhodymeninae Schmitz in Flora 72: 442 (1889).
Order Rhodymeniales Engler in Engler and Prantl. Nat. Pfllanzenfam. I Teil,
Abt. 2: X (1897).
Heterocarpea producing auxiliary cells terminally on brief filaments which grow
from the supporting cells of the carpogonial filaments before fertilization; cystocarps
enclosed in cup- or vase-like pericarps; the thalli (cylindrical or flattened, branched
or unbranched) usually hollow. Champia may be regarded as the standard genus.
In various red algae, the germinating carpospore or tetraspore gives rise to a globe
of cells which grows to produce the thallus (Kylin, 1917). In the present group the
sporeling is particularly blastula-like. Its upper layer of cells becomes a ring of apical
cells, of definite number, distinguishing the group from others which grow by apical
cells either of a single filament or of a fascicle of indefinite number. The apical cells
are indeed homologous with the apical cells of filaments, but the cells derived from
them are arranged in a three-dimensional pattern as in the tissues of higher organisms;
it is only in the reproductive structures that the filamentous structure remains evident.
The order thus limited by Kylin (1932) is a specialized group including only the
two families Rhodymeniacea [Rhodymeniaceae] Hauck and Champiea [Champieae]
Kiitzing. The latter family is the more specialized; the hollow thalli are partitioned
by transverse septa and the supporting cells produce usually just two auxiliary cells.
In many examples of this family, after fertilization and the fusion of the zygote with
the auxiliary cells, the latter proceed to unite with further neighboring cells to pro-
duce a massive coenocyte from which the brief carpospore-bearing filaments arise.
The resulting structure is deceptively similar to that which occurs in the Corallinea.
The parasite Faucheocolax and its host Fauchea belong to this order.
Order 6. Floridea [Florideae] C. Agardh Syst. Alg. xxxiii (1824).
Order Floridees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 115 (1813).
Section Florideae C. Agardh Synops. Alg. Scand. xiii (1817).
Orders Trichoblasteae, Axonoblasteae, and Platynoblasteae Kiitzing Phyc. Gen.
370,413,442 (1843).
Orders Ceramieae, Spyridicae, Chondrieae, and Rhodomeleae J. Agardh Sp.
Alg. vol. 2 (1851-1863).
Ceramiales Oltmanns Morph. u. Biol. Alg. 1: 683 (1904).
Order Ceramiales Kylin in Kgl. Svensk. Vetensk.-Akad. Handl. 63, no. 11 : 132
(1923).
The Floridees of Lamouroux included the whole group of red algae organized as
four genera, Chondriis Stackhouse and the new genera Claudea, Delesseria, and
Gelidium. Lamouroux listed first Claudea and Delesseria, belonging to the present
order, to which the name is accordingly applied.
This order is characterized by specialized strict patterns in the development of the
feniale reproductive structures. The carpogonial filament is always of four cells. The
supporting cell initiates, in definite patterns, brief additional filaments. After fertili-
zation, the supporting cell cuts off one more cell adjacent to the zygote, and this be-
comes the auxiliary cell. The spore-bearing structures developed from it are naked
in the more primitive examples; in most, they are protected by pericarps, which, in
some examples, begin to develop before fertilization.
There are four families, all numerous in species: Ceramiea (Harvey) Kutzing,
52 ] The Classification of Lower Organisms
Dasyea Kiitzirxg, Delesseriea Kutzing, and Polysiphoniea Kiitzing [Rhodomelaceae
Hauck). The Ceramiea are mostly filaments, uniseriate or becoming pluriseriate by
lengthwise divisions. In many members of the other families the bodies are thallose,
though consisting essentially of filaments produced in definite patterns. In many
Delesseriea the branches of the thalli simulate leaves of higher plants.
Gonimophyllum is parasitic on Botryoglossum; both are Delesseriea. Various
species of Janczewskia, a genus of Polysiphoniea, attack Laurencia, Chondria, and
other members of the same family. This was the first genus of parasitic red algae to
be recognized as such, by Solms-Laubach (1877).
Such are the red algae. The Bangialea appear to represent the transition between
the organisms which lack nuclei and the generality of nucleate organisms. The
Heterocarpea appear to be a specialized offshoot, leading to no other group.
Chapter VI
PHYLUM PHAEOPHYTA
Phylum 2. PHAEOPHYTA Wettstein
FucoroEAE C. Agardh Synops Alg. Scand. ix (1817).
Orders Diatomeae and Fucoideae C. Agardh Syst. Alg. xii, xxxv (1824).
Stamme Diatomea and Fucoideae Haeckel Gen. Morph. 2: xxv, xxxv (1866).
Stamme Zygophyta in part and Phaeophyta Wettstein Handb. syst. Bot. 1: 71,
171 (1901).
Divisions Zygophyceae in part and Phaeophyceae Engler Syllab. ed. 3: 8, 15
(1903).
Chysophyta, with subordinate groups Chrysophyceae, Bacillariales, and Hetero-
kontae, Pascher in Ber. deutschen bot. Gess. 32: 158 (1914).
Stamm Chrysophyta Pascher in Siisswasserfl. Deutschland 11: 17 (1925).
Phyla Chrysophycophyta and Phaeophycophyta Papenfuss in Bull. Torrey Bot.
Club 73: 218 (1946).
Organisms typically living by photosynthesis, without chromoprotein pigments,
the plastids containing chlorophylls a and c, carotin, and various xanthophylls. Lutein
(the xanthophyll of typical plants) may be present but is usually exceeded in quantity
by flavoxanthin, violoxanthin, isofucoxanthin, or fucoxanthin, particularly the last.
The xanthophylls occur usually in quantity sufficient to give the organisms a yellow
or brown color. True starch is not produced. Many examples contain granules of a
white solid called leucosin, presumably a carbohydrate, which does not give a blue
color with iodine. The cells are usually enclosed in walls consisting of cellulose to-
gether with larger quantities of other carbohydrates or oxidized or esterized carbo-
hydrates. Silica or calcium carbonate may be deposited. Methanol extracts of the
cells contain fucosterol, a sterol distinct from the sitosterol of typical plants. Flagel-
late cells are usually produced; these bear one pantoneme or pantacroneme flagellum,
and usually, in addition, one acroneme or simple flagellum. Exceptional examples,
non-pigmented or without flagellate stages, are rather numerous. The obvious stand-
ard genus of the phylum is Fucus L.
The chemical characters are stated on the authority chiefly of Carter, Heilbron,
and Lythgoe (1939), Miwa (1940), and Tseng (1945). The character of the flagel-
lation, positively known of rather few examples, is stated by authority of Petersen
(1929), Vlk (1931, 1939), Couch (1938, 1941), Longest [1946), Manton (1952),
and Ferris (1954).
These characters bind together an assemblage of organisms which is in some re-
spects original herel. Engler (1897), West (1904), and Smith (1918, 1920) included
the chrysomonad flagellates in the group of brown algae. Pascher (1914) combined
as or!e group the chrysomonads, the diatoms, and the exceptional green algae called
Heierokontae. Later (1927, 1930), he included also the colorless flagellates of family
Moiiadina. He did not associate this group with the brown algae, and subsequent
authors have in general followed him. Kylin ( 1933 ) , however, considered the diatoms
to be the closest allies of the brown algae, both groups being descended from the
brown flagellates. Almost certainly, he was correct. Couch showed that the paired
unlike flagella of the typical Oomycetes are respectively pantoneme and acroneme,
iManton (1952) recognized this group, but omitted nomenclatural formalities.
54]
The Classification of Lower Organisms
Fig. 8. — Ochromonadalea: a, b, Chrysocapsa paludosa after West (1904); a, a
colony; b, zoospores. C-f, Phaeocystis globosa after ScherlTel (1900); c, a colony
X 50; d, a cell with two plastids, a mass of leucosin forming on a mound of proto-
plasm projecting into the central vacuole; e, production of zoospores; f, a zoospore.
g, h. Cell and statospore of Ochromonas granularis after Doflein (1922). i, Cell of
Monas sp. j. Two cells of Brehmiella Chrysohydra after Pascher ( 1928) . k, A very
young colony of Dendromonas virgaria after Stein (1878). 1, Colony of Ccphalo-
thamnium Cyclopum after Stein, op. cit. m. Cells of Epipyxis utriculus after Stein,
op. cit. n. Colony of Synura Uvella. x 1,000 except as noted.
Phylum Phaeophyta [ 55
and distinguished these fungi from practically all others by the presence of cellulose
in their walls.
The phylum thus assembled may be organized as four classes.
1. Miscellaneous groups, mostly small and rela-
atively unspecialized, of varied body type; not
of the characters of the following groups Class 1. Heterokonta.
1. Comparatively numerous and specialized
groups.
2. Unicellular brown organisms with shells
of silica consisting of two parts Class 2. Bacillariacea.
2. Organisms of fungal or chytrid body
types producing swimming spores with
paired unlike fiagella Class 3. Oomycetes.
2. Filamentous and thallose brown algae Class 4. Melanophycea.
Class 1 . HETEROKONTA Luther
Class Flagellata or Mastigophora Auctt., in part.
Class Heterokontae Luther in Bihang Svensk. Vetensk.-Akad. Handl. 24, part
3, no. 13: 19 (1899).
Subclass Chrysomonadineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: iv (1900).
Class Silicoflagellatae (Borgert) Lemmermann in Ber. deutschen bot. Gess. 19:
254 (1901).
Phylum Siphonophyceae and class Vaucherioideae Bessey in Univ. Nebraska
Studies?: 285, 286 (1907).
Chrysophyceae and Heterokontae Pascher in Ber. deutschen bot. Gess. 32: 158
(1914).
Divisions Chrysophyceae and Heterokontae, and classes Chrysomonadineae , Rhizo-
chrysidineae, Chrysocapsineae, Chrysosphaerineae, Chrysotrichineae, Hetero-
chloridineae, Rhizochloridineae, Heterocapsineae, Heterococcineae, Hetero-
trichineae, and Heterosiphoneae Pascher in Beih. bot. Centralbl. 42, Abt. 2:
323,324 (1931).
Classes Ebriaceae, Silico flagellata, and Coccolithophoridae Deflandre, and Chrys-
omonarfina Hollande in Grasse Traite Zool. 1, fasc. 1: 407,425,438, 471 (1952).
Class Phytomastigophorea Hall Protozoology 117 (1953), in part.
Phaeophyta which lack the distinctive characters of the remaining three classes.
Luther named the group on the occasion of his discovery of Chlorosaccus, and this
genus may be regarded as the type.
The chrysomonad flagellates are the core of this class and of the first two among
the five orders into which it is divided. In the classification of these two orders, three
novelties will be noted.
(a) Pascher (1913) made of the chrysomonad flagellates three orders character-
ized respectively by paired unequal flageila, paired equal flagella, and solitary
fiagella. Petersen (1929) found that the supposedly equal fiagella of Synura are
actually unlike, being respectively pantoneme and acroneme. Here, accordingly,
Pascher's first two orders are combined.
(b) Pascher made separate classes or orders of groups related to the chrysomonad
flagellates but of distinct body type, as palmelloid, chlorococcoid, filamentous, or
56]
The Classification of Lower Organisms
Fig. 9. — Ochromonadalea : a, Mallomonas roseola, based on Stein (1878) and
Conrad (1926). h, Syracosphaera Quadricornu; c, Calyptosphaera insignis; d, Cal-
ciconus vitreus; after Schiller (1925). Silicoflagei.lata: e, f, Colony and zoospore
of Epichrysis after Pascher (1925). g, Part of the thallose growth of Hydrurus
foetidus. h, Cell, and i, j, statosporos of Chromulina Pascheri after Hofeneder
(1913). k, 1, Skeletons of Dictyocha Fibula and Distephanus Speculum from di-
atomaceous earth at Lompoc, California, m, Rhizochrysis Scherffeli after Doflein
(1916). Mix 1,000.
Phylum Phaeophyta [57
amoeboid. By Pascher's own principle of the repeated evolution of body types,
these groups are surely artificial. Here most of them are broken up and their mem-
bers distributed between the two chrysomonad orders according to whether the
flagella of their motile stages are paired or single. It is not possible to divide by this
character ameboid forms not known to produce flagellate stages; these are lumped
in the second order.
fc) Since flagella appear to have evolved as a device for the dissemination of
unicellular pigmented organisms, examples whose vegetative state is that of clusters
of non-motile cells are placed in each order before those which are flagellate in
the vegetative condition.
The two chrysomonad orders are particularly characterized by production of
leucosin. They are further characterized by production of resting cells of a type
called statospores. This occurs by the deposition within the protoplast of a globular
shell impregnated with silica, punctured by a single pore, and often marked on the
outer surface by warts, spines, or ridges, of definite pattern. The external protoplasm
migrates through the pore to the interior of the shell, and the pore is then closed
by deposition of a silicified plug.
The group which is treated as the third order of the present class includes the
typical Heterokonta. Compared with typical green algae, these organisms give the
impression of a markedly distinct class; placed next to the chrysomonads, they
appear scarcely entitled to this rank. Their name is the oldest applicable to the
present class, and is accordingly so applied. If it appear expedient to maintain the
typical Heterokonta as a distinct class, the remainder of the present one will be
called Chrysomonadinea [Chrysomonadineae] (Engler) Pascher.
Of including the choanoflagellates and anisochytrids in the present class as addi-
tional orders, one may say that it is not contrary to current knowledge.
1. Mostly pigmented; non-pigmented examples
mostly producing motile cells with two
fiagella.
2. Brown or colorless.
3. Producing motile cells with two
flagella (exceptionally more) Order 1. Ochromonadalea.
3. Producing motile cells with one
flagellum; or without known flagel-
late stages Order 2. Silicoflagellata.
2. Green Order 3. Vaucheriacea.
1. Non-pigmented, producing motile cells with
one flagellum.
2. Predatory, flagellate in the vegetative
condition, each cell bearing a collar-like
protoplasmic ridge Order 4. Choanoflagellata.
2. Parasitic or saprophytic, the vegetative
cells non-motile, walled Order 5. Hyphochytrialea.
Order 1. Ochromonadalea [Ochromonadales] Pascher Siisswasserfl. Deutschland
2: 10,51 (1913).
Suborder Monadina Biitschli in Bronn KI. u. Ord. Thierreichs 1 : 810 (1884).
Order Isochrysidales Pascher op. cit. 10, 42.
Order Syracosphaerinae Schiller in Arch. Prot. 51: 108 (1925).
58] The Classification of Lower Organisms
Orders Heliolithae and Orthlithinae Deflandre in Grasse Traite Zool. 1, fasc.
1: 452, 457 (1952).
Brown or colorless Heterokonta, the swimming cells of typical examples with
two flagella which are respectively pantoneme and acroneme. In the exceptional
family Trimastigida there are a pair of equal flagella and a third flagellum shorter
or longer than these; the detailed structure of the flagella of this family is unknown.
Cells of pigmented types contain usually one or two lateral band-shaped plastids.
Details of nuclear division are known chiefly by the observations of Doflein (1918,
1922) on Ochromonas. The flagella spring from a granule which may be identified as
a blepharoplast, near which lies the nucleus. The blepharoplast is connected through
two stainable strands (rhizoplasts) to two granules, recognizable as centrosomes, on
the two sides of the nucleus. The spindle forms within the intact nuclear membrane
with its poles at the centrosomes. The chromosome number appears to be about 4.
The nuclear membrane presently disappears. At metaphase, the rhizoplasts are found
to lead to separate blepharoplasts, each bearing two flagella. Sexual processes are
scarcely known in this group. Schiller (1926) observed in Dinobryon the division of
calls into two which are released to swim and conjugate in pairs.
This order is believed to represent the direct ancestry of the two following, and
also of the typical brown algae.
1. Not filamentous.
2. Flagellate stages with a pair of equal
flagella and a third which is shorter or
longer Family 1. Trimastigida.
2. Flagellate stages with two unequal
flagella.
3. Without calcareous structures at-
tached to the cell walls.
4. Cells not enclosed in loricae,
i. e., open shells.
5. Not flagellate in the vege-
tative condition Family 2. Chrysocapsacea.
5. Flagellate in the vegeta-
tive condition, not forming
free-swimming circular or
globular colonies Family 3. Monadina.
5. Free-swimming circular or
globular colonies Family 4. Syncryptida.
4. Cells enclosed in loricae Family 5. Dinobryina.
3. With calcareous structures at-
tached to the cell walls Family 6. Hymenomonadacea.
1. Filamentous Family 7. Phaeothamnionacea.
Family 1. Trimastigida [Trimastigidae] Kent Man. Inf. 1: 307 (1880). Family
Trimastigaceae Senn in Engler and Prantl. Nat. Pflanzcnfam. I Teil, .\bt. la: 141
(1900). Family Prymncsiidae Hall Protozoology 127 (1953). Organisms producing
swimming cells with a pair of equal flagella and a third flagellum longer or shorter
than these. With a vegetative stage as globular non-motile colonies as large as pin-
heads, of pigmented cells; marine: Phacocystis. Motile solitary cells, pigmented:
Prymncsium, Chrysochromidina; Platychrysis with an amoeboid stage. Motile
solitary cells, not pigmented: Dallingeria, Trimastix, Macromastix.
Phylum Phaeophyta [ 59
Family 2. Chrysocapsacea [Chrysocapsaceae] Pascher in Siisswasserfl. Deutschland
2: 85 (1913). Family Chrysocapsidae Poche in Arch. Prot. 30: 156 (1913). Non-
motile cells with brown plastids (usually two), imbedded in gelatinous matter and
forming colonial aggregates, the protoplasts sometimes escaping as zoospores with
two flagella. Chrysocapsa Pascher, in fresh water, the colonies few-celled. Phaeo-
sphaera West and West, the colonies more extensive.
Family 3. Monadina Ehrenberg Infusionsthierrhen 1 (1838). Family Monades
Goldfuss ( 1818), the mere plural of a generic name. Family Dendromonadina Stein
Org. Inf. 3, I Halfte: x (1878). Family Monadidae Kent (1880). Family Hetero-
monadina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 815 (1884). Family Chryso-
monadaceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570
(1897), not family Chrysomonadina Stein. Family Ochromonadaceae Senn in
Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 163 (1900). Family Ochro-
monadidae Doflein. Pigmented or colorless Ochromonadalea, flagellate in the
vegetative condition, not forming circular or globular free-swimming colonies, nor
loricate, nor bearing calcareous structures on the cell walls (these being the distinc-
tions respectively of the three following families).
Ochromonas is considered to be in its normal condition when it occurs as solitary
swimming cells; it occurs also as gelatinous colonies like those of Chrysocapsa.
Stylochrysalis consists of O chromonas-like cells attached by a stalk at the end away
from the flagella. Chrysodendron is similar but colonial, the cells attached by branched
stalks. Brehmiella Pascher (1928) may occur as free-swimming Ochromonas-Vikt
cells, or these may become attached by the end away from tlie flagella and develop
a whorl of pseudopodia at the free end. Pseudopodia are a device for predatory nutri-
tion, here occurring in an organism which is capable also of photosynthesis. Hetero-
chromonas includes organisms of the structure of Ochromonas but without plastids, be-
ing presumably saprophytic, and containing only a pigmented speck by which it is sup-
posed that the direction of light is perceived. The historical generic name Monas
O. F. Miiller, as restricted in application by scholars up to Ehrenberg and as applied
ever since, designates totally non-pigmented cells, saprophytic or predatory, free-
swimming like Ochromonas or attached like Stylochrysalis [Physomonas Kent desig-
nates cells of Monas in the attached condition). There are believed to be several
species, but the group remains poorly known. It was in some member of it that Loeffler
(1889) first observed the pantoneme character of flagella. Dendromonas consists of
similar cells forming colonies like those of Chrysodendron. In Cephalothamnium
Stein, Monas-\ikc cells are gathered in capitate clusters on stout stalks. Anthophysis
Bory is an organism which Leeuwenhoeck had described as a microscopic water
plant: it consists of Monas-Vikt cells at the ends of branching stalks colored yellow
by deposits of iron. The comparatively unfamiliar original spellings of the two
generic names just mentioned were restored by Kudo ( 1946). The name Uvella Bory
appears to represent small clusters of cells of Cephalothamnium or Anthophysis
which have broken loose to swim free.
Family 3. Syncryptida [Syncryptidae] Poche in Arch. Prot. 30: 156 (1913). Family
Isochrysidaceae Pascher in Siisswasserfl. Deutschland 2: 43 (1913), not based on a
generic name. Family Isochrysidae Calkins Biol. Prot. 262 (1926). Families Synura-
ceae and Syncryptaceae Smith Freshw. Algae (1933). Ochromonas-Vike cells forming
circular or globular free-swimming colonies. Flagella markedly unequal, colonies
circular: Cyclonexis; colonies globular: Uroglena, Uroglenopsis. Flagella apparently
equal: Syncrypta, Synura.
60 ] The Classification of Lower Organisms
Family 4. Dinobryina Ehrenberg Infusionsthierchen 122 (1838). Family Dino-
hryaceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897).
Pigmented or colorless cells of the characters of Ochromonas or Monas, sheltered in
loricae, that is, in transparent open shells, solitary or colonial. The pigmented
examples have generally been referred to Ochromonadaceae (or whatever), the
colorless to Monadidae (or whatever). Pigmented, solitary, flagella markedly
unequal: Epipyxis, Stylo pyxis; flagella apparently equal: Chry so pyxis Stein {Dere-
pyxis Stokes). Pigmented, forming branching colonies: Dinobryon, Hyalobryon.
Poteriochromonas Scherffel resembles Stylopyxis, but the protoplast can project
pseudopodia from its lorica, thus supplementing photosynthesis by predatory nutri-
tion. Non-pigmented, solitary, flagella markedly unequal: Stokesiella; flagella ap-
prrently equal: Diplomita. Non-pigmented cells in colonies quite of the character
of those of Dinobryon: Stylobryon.
Family 5. Hymenomonadacea [Hymenomonadaceae] Senn in Engler and Prantl
Nat. Pflanzenfam. I Teil, Abt. la: 159 (1900). Family Coccolithophoridae Lohman
in Arch. Prot. 1: 127 (1902). Family Hymenomonadidae Doflein. Family Cocco-
lithidae Poche in Arch. Prot. 30: 157 (1913). Order Syracosphaerinae and family
Pontosphaeraceae Schiller in Arch. Prot. 51: 8 (1925). Families Syracosphaeraceae,
Halopappaceae, Deutschlandiaceae, and Coccolithaceae Kampter. Family Thora-
cosphaeracee Schiller in Rabenhorst Kryptog.-Fl. Deutschland ed. 2, 10, Abt. 2: 156
(1930). Y3imi\it5 Syracosphaeridae, Calcisolenidae, Thoracosphaeridae, and Braad-
rudosphaeridae Deflandre in Grasse Trait6 Zool. 1, fasc. 1: 452, 457, 458 (1952).
Family Discoasteridae Tan Sin Hok. Suborder Coccolithina Hall Protozoology 130
(1953). Solitary cells with one or two brown plastids, usually with two apparently
equal flagella, having a thin cell wall from which project bodies of calcium carbonate
(coccoliths) of definite form.
More than twenty genera and nearly 150 species have been described (Lohman;
Schiller; Kamptner, 1940). Neither the number of species nor the variety of form
appears to warrant making more than one family of the group. Nearly all examples
are marine. In Pontosphaera, Calyptosphaera, and allied genera, the coccoliths are
disks or hemispheres, sometimes umbonate and sometimes marked by one or more
pits. In Syracosphaera the coccoliths, or a few of them near the insertion of the
flagella, bear horn-like projections. In Najadea, Halopappus, and Calciconus, each
cell bears a whorl or elongate bristles. Cells of Calcisolenia are fusiform, without
flagella, with an armor of two layers of spiral bands of calcareous matter. In Hymen-
omonas and Coccolithus Swartz 1894 [Coccosphaera Wallich 1877, non Perty 1852;
Coccolithophora Lohman 1902) the coccoliths are punctured and accordingly ring-
shaped; Hymenomonas difi"ers from most of the group in occurring in fresh water. In
Discosphaera and Rhabdosphaera the punctured calcareous bodies are drawn out to
the form of tubes, spools, or trumpets.
These obscure organisms are not without importance. They occur in all oceans,
being most abundant in gulfs, such as the Adriatic, where the salinity is diminished
by rivers (Schiller, 1925). According to Bernard (1947) turbidity in the Mediter-
ranean depends chiefly on this group. Coccoliths are abundant in the ooze on the
bottoms of oceans. They occur as fossils as far back as the Cambrian, being par-
ticularly abundant in certain Cretaceous deposits.
Family 6. Phaeothamnionacea [Phaeothamnionaccae] Pascher in Siisswasserfl.
Deutschland2: 113 ( 1913). Family Chrysotrichaceae Fascher (1914). Family Nema-
tochrysidaceae Pascher (1925). Brown organisms, minute, marine, epiphytic, filamen-
Phylum Phaeophyta [ 61
tous, reproducing by zoospores bearing paired unequal flagella. Nematochrysis, the
filaments unbranched; Phaeothamnio7i, the filaments branched. These organisms are
believed to represent the transition between the Chrysocapsacea and the typical
brown algae.
There is a family Amphimonadidae or Amphimonadaceae of unwalled colorless
flagellates with paired supposedly equal flagella. They appear to belong to the
kingdom of plants, in the neighborhood of Chlamydomonas and Polytoma. If, how-
ever, future study shows their flagella actually to be respectively pantoneme and
acroneme, they are to be placed in the present order.
Order 2. Silicoflagellata Borgert in Zeit. wiss. Zool. 51: 661 (1891).
Chromomonadina Klebs in Zeit. wiss. Zool. 55: 394 (1893).
Order Chromomonadina Blochmann Mikr. Tierwelt ed. 2. Abt. I: 57 (1895).
Subclass Chrysomonadineae Engler in Engler and Prantl Nat. Pflanzenfam.
ITeil, Abt.'la: iv (1900).
Order Chrysomonadales Engler Syllab. ed. 3: 7 (1903).
Chrysomonadinae; Euchrysomonadinae , with order Chromulinales; Chryso-
capsinae; and Rhizochrysidinae Pascher in Siisswasserfl. Deutschland Heft 2
(1913).
Chrysomonadales, Chrysocapsales, Chrysosphaerales, and Chrysotrichales Pas-
cher in Ber. deutschen bot. Gess. 32: 158 (1914).
Order Chrysomonadina Doflein Lehrb. Prot. ed. 4: 401 (1916).
Order Chrysomonadida Calkins Biol. Prot. 258 (1926).
Classes Chrysomonadineae , Rhizochrysidineae, Chrysocapsineae, Chrysosphaeri-
neae, and Chrysotrichineae Pascher in Beih. bot. Centralbl. 48, Abt. 2: 323
(1931).
Suborders Euchrysomonadina, Silicoflagellina, Rhizochrysidina, and Chrysocap-
sina Hall Protozoology 125, 128, 130, 132 (1953).
Organisms of much the character of Ochromonadalea, but producing flagellate
stages with a single flagellum, or not producing flagellate stages. The detailed
structure of the flagella has seemingly never been determined. Statospores are known
to be produced by Chromulina, Mallonionas, and (of somewhat exceptional charac-
ter) by Hy drums. Sexual reproduction has not been observed. Mitosis, with an
intranuclear spindle and numerous chromosomes, was observed by Doflein (1916)
in Rhizochrysis.
This order is supposed to represent the direct ancestry of orders Choanoflagellata
and Hyphochytrialea.
1. Neither amoeboid nor truly filamentous.
2. Not flagellate in the vegetative condi-
tion.
3. Microscopic colonies Family 1. Chrysosphaeracea.
3. Macroscopic gelatinous colonies
simulating filaments Family 2. Hydruragea.
2. Flagellate in the vegetative condition.
3. Without prominent siliceous struc-
tures Family 3. Chrysomonadina.
3. With siliceous scales usually bearing
bristles Family 4. Mallomonadinea.
3. With siliceous internal skeletons Family 5. Actiniscea.
62 ] The Classification of Lower Organisms
1. Amoeboid Family 6. Chrysamoebida.
1. Filamentous Family 7. Thallochrysidacea.
Family 1. Chrysosphaeracea [Chr>'Sosphaeraceae] Pascher in Arch. Prot. 52: 562
(1925). Family Naegelliellaceae Pascher op. cit. 561. Family Nagelliellidae Hall
Protozoology 133 (1953). Non-motile brown cells, either capable of repeated division
into two, thus forming aggregates of indefinite number, or else undergoing multiple
division and producing colonies of definite number of cells; mostly known to produce
uniflagellate zoospores. Chrysosphaera, Epichrysis, Chrysospora, Gloeochrysis, Nae-
gelliella, and other genera.
Family 2. Hydruracea [Hydruraceae] West British Freshw. Algae 45 (1904).
Hydrurina Klebs in Zeit. wiss. Zool. 55: 420 (1893). Family Hydruridae Poche in
Arch. Prot. 30: 158 (1913). Like Chrysosphaeracea, but the colonies dendroid,
growing at the tips, becoming macroscopic; producing tetrahedral zoospores and
spheroidal resting cells bearing a unilateral crest. Hydrurus foetidus, in mountain
streams.
Family 3. Chrysomonadina Stein Org. Inf. 3, I Halfte: x (1878). Family
Chrysomonadidac Kent Man. Inf. (1880). Family Chromulinaceae Engler in Engler
and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). Family Chromulinidae
Doflein. Brown flagellates with a single anterior flagellum, sometimes producing
siliceous granules but without more extensive siliceous structures. Free-swimming,
walled: Chrysococcus, Microglena. Naked: Chromulina, the type genus of Chryso-
monadina, the generic name Chrysomonas being a synonym. Organisms of this genus
are rather freely capable of producing pseudopodia and supplementing photosynthetic
nutrition by predatism, or, alternatively, of producing gelatinous aggregates of
walled non-motile cells (Hofender, 1913; Gicklhom, 1922). Chrysapsis differs from
Chromulina in having in each cell a single plastid in the form of a network. Solitary
attached cells, producing pseudopodia only occasionally: Lepo chromulina. Bearing
whorls of permanent pseudopodia: Cyrtophora, Pedinella, Palatinella (Pascher,
1928).
Family 4. Mallomonadinea Diesing in Sitzber. Akad. Wiss. Wien Math.-Nat. CI.
52, Abt. 1: 304 (1866). Family Mallomonadidae Kent (1880). Brown uniflagellate
free-swimming cells with an armor of siliceous scales usually bearing bristles. Mallo-
m.onas, solitary cells, the bristle-bearing scales circular. Conradiella, the scales of the
form of rings about the body. Chrysosphaerella, spherical colonies, each cell with two
long bristles.
Family 5. Actiniscea [Actinisceae] Kiitzing Phyc. Germ 117 (1845). Family
Dictyochidae Wallich. Class Silicoflagellata (Borgert), orders Siphonotestales and
Stereotestales, and families Dictyochaceae and Ebriaccae Lemmermann in Ber.
deutschen bot. Gcss. 19: 254-268 ( 1901 ). Division (?) Silicoflagellatac Engler. Family
SiHcoflagellidae Calkins Biol. Prot. 263 (1926). Famihes Ebriopsidae, Ditripodiidae,
Ammodochidae, and Ebriidae Deflandre in Grasse Traite Zool. 1, fasc. 1: 421, 423,
424 (1952). Solitary brown uniflagellate cells with a continuous internal skeleton of
silica. Marine, commonest in colder oceans.
The skeletons are not subject to decay and are found as micro fossils in chalk
and diatomaceous earth. They have been reported from the Silurian and are com-
monest in certain Cretaceous deposits. Ehrcnbcrg described several fossil species,
classifying them as diatoms. The living forms, subsequently discovered, include
apparently the same species.
Gemeinhardt (in Rabcnhorst, 1930) accounted for the structure of the cells.
Phylum Phaeophyta [ 63
They are approximately of radial symmetry, the axis being shorter than the diameter.
The skeleton is completely imbedded in protoplasm. It may be a mere ring; or the
ring may bear radially projecting spines; or it may be the margin of a more or less
complicated basket-shaped network coaxial with the cell. Numerous brown plastids
lie near the surface of the protoplast. There is no cell wall. The double cells, like
two cells lying face to face, which have occasionally been seen, are not stages of
conjugation, but of cell division, in which one daughter cell retains the original
skeleton while the other develops a new skeleton in the position of a mirror image
of the original one.
Lemmermann and Gemeinhardt accounted for only six genera and twenty-four
species, but Gemeinhardt recognized numerous varieties, and it is probable that the
number of species has been underestimated. Mesocaena, the skeleton a mere ring,
smooth or spiny; Dictyocha, Distephanus, Cannopilus, the skeleton more or less
netted.
Family 6. Chrysamoebida [Chrysamoebidae] Poche in Arch. Prot. 30: 157 (1913).
Families Rhizochrysidaceae , Chrysarachniaceae, and Myxochrysidaceae Pascher in
Beih. Bot. Centralbl. 48, Abt. 2: 323 (1931). Family Rhizochrysididae Hollande in
Grasse Traite Zool. 1, fasc. 1: 547 (1952). Families Rhizochrysidae and Myxochry-
sidae Hall Protozoology 130, 132 (1953). Amoeboid organisms with brown plastids
of the form of one or two parietal films in each cell. Rhizaster, an attached organism
resembling Cyrtophora and Pedinella but lacking the flagellum. Chrysocrinus, at-
tached to algae, the protoplast covered by a dome-shaped shell punctured by many
pores through which project the slender psudopodia. Chrysamoeba, a freely moving
cell usually with one flagellum; Rhizochrysis, similar, without the flagellum. Myxo-
chrysis, a large multinucleate form. Chrysarachnion, the cells clustered and linked
together by strands of protoplasm. Lagynion, having an attached vase-shaped lorica
from which projects usually a single slender pseudopodium. Chrysothylakion, with
a retort-shaped lorica from which project many slender pseudopodia, branching
and anastomosing. Only the plastids distinguish these organisms from various genera
classified as Rhizopoda, Heliozoa, or Sarkodina.
Family 7. Thallochrysidacea [Thallochrysidaceae] Pascher (1925). Brown or-
ganisms producing definite filaments of walled cells and reproducing by anteriorly
uniflagellate zoospores. T hall ochry sis. Phaeodermatium.
Order 3. Vaucheriacea [Vaucheriaceae] Nageli Gatt. einzell. Alg. 40 (1849).
Class Heterokontae and orders Chloromonadales and Confervales Luther in
Bihang Svensk. Vetensk.-Akad. Handl. 24, part 3, no. 13: 19 (1899). Not
Chloromonadina Klebs (1893); not order Confervoidea C. Agardh (1824).
Vaucheriales Bohlin Grona Algernas 25 (1901).
Order Vaucheriales Clements Gen. Fung. 14 (1909).
Orders Heterochloridales, Heterocapsales, Heterococcales, Heterotrichales, and
H eter osiphonales Va.s.ch.tr mlitdwigizbZ: 10-21 (1912).
Division Heterokontae, Classes Heterochloridineae, Rhizochloridineae, Hetero-
capsineae, Heterococcineae, Heterotrichineae , and Heterosiphoneae, and or-
ders Rhizochloridales and Botrydiales Pascher in Beih. bot. Centralbl. 48,
Abt. 2: 324 (1931).
Class Xanthomonadina with orders Heterochloridea and Rhizo chloride a De-
flandre in Grasse Traite Zool. 1, fasc. 1: 212, 217, 220 (1952).
Order Heterochlorida Hall Protozoology 133 (1953).
64]
The Classification of Lower Organisms
Organisms producing motile cells with paired unequal flagella which Vlk (1931)
found to be respectively pantoneme and acroneme, differing from Ochromonadalea
in being of a green or yellow-green color, and in being mostly of algal body type,
i. e., walled and non-motile. The cell wall consists usually of two parts which become
separate when the cell divides; the two parts are believed to be distantly homologous
with the wall and pRig of the statospores of Ochromonadalea and Silicoflagellata
(Pascher, 1932). The storage products are oil and sometimes leucosin.
As this is the group to which the class name Heterokontae was first applied, it is
f-'^y-'v'^^^; ■■■• >i%'X°-^'^^
Fig. 10. — Vaucheriacea: a, b^ Chlorosaccus fluidus, cells of the colony and zoo-
spores, after Luther (1899). c, d^ Chlorarnoeba heteromorpha x 1,000 after Bohlin
(1897). e, f, g. Cell, empty cell, and zoospores of Characiopsis gibba x 1,000 after
Pascher (1912). h, Dioxys Incus after Pascher (1932). i, j, k, Cell, edge of cell,
and statospore of Pseudotetraedron neglectum x 1,000 after Pascher (1912). 1, Spi-
rodiscus fulvus x 1,000. m. End of a filament of Tribonema bombycina x 1,000.
n, Antheridium and oogonia of Vaucheria Gardneri x 100. o. Filament of Vaucheria
sessilis x 100.
Phylum Phaeophyta [ 65
the type group of the class. As established by Luther, the class consisted of the new
genus Chlorosaccus together with a few genera of flagellates ( Vacuolaria was included
in error) and a few transferred from the group of typical green algae. From time to
time, other green algae have been transferred, and it has become evident that the
group is a fairly extensive one. Green organisms can be recognized as belonging here
by a negative reaction to the iodine test for starch, and by the fact that they give a
b'uish color when heated with hydrochloric acid, instead of a yellow one, as typical
green algae do: the difference depends upon differences in the complement of
photosynthetic pigments. Bohlin (1901) placed Vaucheria here; most authors have
not followed him, but Smith (1950) has done so. This genus brings with itself the
oldest name for the group as an order.
Mitosis in Vaucheria was described by Hanatschek (1932) and Gross (1937). The
spindle is intranuclear; Hanatschek saw centrosomes at the poles. The conjugation of
equal free-swimming gametes was observed in Tribonema and several other genera by
ScherfFel (1901), and in Botrydium by Rosenberg (1930). Vaucheria was one of the
organisms by study of which the nature of fertilization was discovered (Pringsheim,
1855). Hanatschek and Gross found that the first two divisions of the nucleus of
the zygote are meiotic: the soma is haploid.
This order is believed to represent the direct ancestry of the two following classes,
Bacillariacea and Oomycetes.
Pascher (1912, 1925) arranged the green Heterokonta in subordinate groups
parallel to those of the typical green algae; and, as the main groups of green algae
are treated as orders, he treated these groups also as orders (in 1931 as classes).
They are scarcely entitled to such rank: too many of the classes or orders are of
single families, and too many of the families are of one or two genera. Here, then,
Pascher's classes and orders are suppressed and several of his families are reduced.
1. Not truly filamentous nor producing rhizoids.
2. The cells walled.
3. Cells regularly dividing into two,
forming gelatinous colonies; occa-
sionally producing small numbers
of zoospores.
4. The colonies globular or Irreg-
ular, becoming macroscopic Family 1. Chlorosagcacea.
4. The colonies dendroid, micro-
scopic Family 2. Mischococgacea.
3. Cells normally undergoing division
into several.
4. Producing zoospores Family 3. Chlorotheciacea.
4. Producing no motile cells Family 4. Botryococcagea.
2. The cells loricate Family 5. Stipitogogcacea.
2. The cells amoeboid Family 6. Chloramoebacea.
1. Filaments of uninucleate cells Family 7. Tribonematagea.
1. Cells becoming highly multinucleate, form-
ing filaments or at least producing rhizoids Family 8. Phyllosiphonacea.
Family 1. Chlorosaccacea [Chlorosaccaceae] Smith Freshw. Algae 145 (1933).
Family Heterocapsaceae Pascher in Hedwigia 53: 13 (1912); there is no correspond-
ing generic name. Gelatinous aggregates of cells which may divide, causing the
66] The Classification of Lower Organisms
aggregate to grow to macroscopic dimensions; or may produce one, two, or four
zoospores. Chlorosaccus Luther, the standard genus of class Heterokonta.
Family 2. Mischococcacea [Mischococcaceae] Pascher in Hedwigia 53 : 14 ( 1912).
Microscopic colonies of globular cells joined by dichotomously branching gelatinous
strands. Mischococcus.
Family 3. Chlorotheciacea [Chlorotheciaceae] Luther in Bihang Svensk. Vetensk-
Akad. Handl. 24, part 3, no. 13: 19 (1899). Families Chlorobotrydiaceae and Sci-
adiaccae Pascher in Hedwigia 53: 17 (1912). Family Halosphaeraceae Pascher
(1925). Family Ophiocytiaceae Auctt. Cells solitary, free or attached, capable of
reproduction by division to form multiple zoospores, in some examples capable
alternatively of producing multiple minute non-motile cells of the same form as
the parent. Large free multinucleate cells, more or less globular: Botrydiopsis, Leu-
venia. Smaller cells, elongate, curved or coiled: Characiopsis, Spirodiscus. Spirodiscus
fuluus Ehrenberg in Abh. Akad. Wiss. Berlin 1830: 65 (1832) {nomen nudum) and
Infusionsthierchen 86 (1838), whose identity has been a standing puzzle to bac-
teriological systematists, is an older name of Ophiocytium parvidum (Perty) A.
Braun (Copeland, 1954). It antedates the generic name Ophiocytium Nageli (1849);
new combinations are required for the dozen additional species of this genus. The
cells attached: some species of Characiopsis; Perionella; Dioxys.
Family 4. Botryococcacea [Botryococcaceae] Pascher in Hedwigia 53: 13 (1912).
Solitary or colonial cells reproducing strictly by production of non-motile cells.
Botryococcus. Pseudotetraedron.
Family 5. Stipitococcacea [Stipitococcaceae] Pascher in Beih. bot. Centralbl. 48,
Abt. 2: 324 (1931). Family Stipitochioridae Deflandre in Grasse Trate Zool. 1, fasc.
1: 221 (1952). Amoeboid cells with green plastids, partially enclosed in loricae at-
tached to objects in water. Stipitococcus.
Family 6. Chloramoebacea [Chloramoebaceae] Luther in Bihang Svensk. Vetensk.-
Akad. Handl. 24, part 3, no. 13: 19 (1899). Family Chloramoebidae Poche in Arch.
Prot. 30: 155 (1913). Families Heterochloridaceae and Rhizochloridaceae Pascher
Siisswasserfl. Deutschland 11: 22, 26 (1925). Y^.miWts Heterochloridae, Rhizochlori-
dae, Chlorarachnidae and Myxochloridae Deflandre in Grasse Traite Zool. 1, fasc. 1 :
217-222 (1952). Amoeboid organisms with green plastids, without loricae, some-
times swimming by means of paired unequal flagella. Chloramoeba, Chlorochromo-
nas, Rhizochloris.
Family 7. Tribonematacea [Tribonemataceae] Pascher in Hedwigia 53 : 19 ( 1912) .
Family Confervaceae Luther (1899). Family Monociliaceae Smith Freshw. Algae
160 (1933). Green Heterokonta producing filaments of uninucleate cells. The Lin-
naean genus Conferva included a great variety of growths in water. Definite groups
were separated from it, one after another, until the residue was a natural group; but
this residue cannot be assumed to be the type of Conferva L.; that name is to be
abandoned as a nomen confusum. The remnant in question has become two genera,
Tribonema Derbes and Solier, 1858, and Bumilleria Borzi, 1895. They are unbranched
filaments, common in freshwater pools. From typical green algae of similar appear-
ance they are distinguished in the first place by the presence in each cell of several
disk-shaped plastids without pyrenoids or with obscure ones. The cell walls, when
treated with sulfuric acid, can be seen to consist of two parts like a barrel sawed
across the middle. A broken filament ends always with a broken half wall. Monocilia,
an unfamiliar alga isolated from soil, difi^ers in producing branching filaments.
Phylum Phaeophyta [ 67
Family 8. Phyllosiphonacea [Phyllosiphonaceae] Wille in Engler and Prantl. Nat.
Pflanzenfam. I Teil, Abt. 2: 125 (1890). Family Vaucheriaceae (Nageli) Areschoug
(1850), preoccupied by order Vaucheriaceae Nageli. Family Botrydiaceae Luther
(1899). Heterokonta whose bodies are highly multinucleate single cells, filamentous
or anchored by filamentous rhizoids. Botrydium is found on damp soil as dark green
globes, sometimes as much as 2 mm. in diameter, anchored by much-branched color-
less rhizoids. Vaucheria is a familiar alga on damp earth or in fresh water. It consists
of irregularly branching filaments, green where exposed to light, colorless where
growing downward and serving as rhizoids. The reproductive cells are cut off by
walls. The end of an aerial filament, cut off in this fashion, may as a whole act as a
spore. In water, the protoplast of such a cell may escape as an exceptionally large
zoospore with as many pairs of flagella as the nuclei within it. Antheridia are brief
branches, each releasing many minute sperms each with two unequal flagella.
Oogonia are globular cells, multinucleate during development, but containing only
one functional nucleus when mature. Phyllosiphon is of much the same structure as
Vaucheria, but is parasitic in seed plants, particularly Araceae. It reproduces, ap-
parently, only by the breaking up of the protoplast to produce minute non-flagellate
spores.
Order 4. ChoanoflageUata [Choano-Flagellata] Kent Man. Inf. 1: 36 (1880).
Order Bicoecidea Grasse and Deflandre in Grasse Traite Zool. 1, fasc. 1: 599
(1952).
Non-pigmented flagellates, usually attached, each cell bearing a single flagellum
of the type called pantacroneme, with lateral appendages and a terminal whip-lash;
the cell bearing also a protoplasmic collar, usually surrounding the base of the flagel-
lum. The collar is a means of nutrition. Bacteria and other scraps of organic matter,
driven against it by the beating of the flagellum, adhere and are carried to the interior
of the cell by flow of the cytoplasm of which it consists.
It is probable that the pantacroneme flagellum is a variant of the pantoneme
flagellum, and that this order belongs naturally in class Heterokonta. It may have
evolved from Silicoflagellata; or it may be that the collar is a modified flagellum,
and that the group evolved from order Ochromonadalea.
Most authors have recognized more than one family of choanoflagellates, but
genera are not very numerous and one family seems sufficient to accommodate them.
Family Bicoekida Stein Org. Inf. 3, I Halfte: x (1878). Family Craspedornona-
dina Stein 1. c. Families Bikoecidae, Codonosigidae, Salpingoecidac, and Phalansteri-
idae Kent op. cit. Families Codonoecina and Bikoecina Biitschli in Bronn Kl. u. Ord.
Thierreichs 1: 814, 815 (1884). Families Bicoecaceae, Craspedoynonadaceae, and
Phalanasteriaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 121,
123, 129 (1900). Family Gymnocraspedidae Grasse Traite Zool. 1, fasc. 1: 590
(1952). Characters of the order. Cells naked, solitary: Monosiga; colonial: Codo-
siga James-Clark [Codonosiga Stein), Sphaeroeca. Cells imbedded in gelatinous
matter, the collars contracted: Phalanseterium. Loricate: Salpingoeca, Bicosoeca,
Poteriodendron.
The choanoflagellates were discovered by James-Clark (1866, 1868), who made at
the same time the discovery that certain internal cavities of sponges are lined by
minute cells (choanocytes) of the same structure as the choanoflagellates. From
these observations he drew the conclusion that sponges are a sort of flagellates dis-
tinguished by the production of exceptionally large and elaborate colonies. Kent
68]
The Classification of Lower Organisms
described Proterospongia Haeckeli as a colonial organism of amoeboid and choano-
flagellate cells in a common matrix; he regarded it as a transitional form, important
as evidence of the evolution of sponges from choanoflagellates. According to Duboscq
and Tuzet ( 1937) it is no organism, but a stage in the development of an individual
sponge from one which has been damaged. In spite of this, the hypothesis that the
choanoflagellates represent the evolutionary origin of the sponges, and accordingly
of the entire animal kingdom, continues to appear tenable.
Fig. 11. — Choanoflagellata : a, b, Monosiga spp.; c^ Phalanasterium digitatum;
A, Salpingoeca ampullacea; e, Salpingoeca Clarkii; i, Foteriodendron petiolatum.
c X 500, the remainder x 1,000. c-f after Stein (1878).
Phylum Phaeophyta [ 69
Order 5. Hyphochytrialea [Hyphochytriales] Bessey Morph. and Tax. Fungi 69
(1950).
Order Anisochytridiales Karling in American Jour. Bot. 30: 641 (1943), not
based on a generic name.
Non-pigmented organisms with walled cells, parasitic or saprophytic, the proto-
plasm with numerous granules not of a shining appearance, producing zoospores
with single anterior pantoneme flagella.
The naked zoospores come to rest upon appropriate hosts or substrata. Ordinarily,
in parasitic species, the protoplast of the zoospore makes its way to the interior of a
cell of the host. It swells and develops a thin wall. The resulting structure may be
called a center. In most members of the group, the center gives rise to a system of
slender rhizoids; in some species, these give rise to further centers like the original
one. Karling studied the cytology particularly in Anisolpidium. There are repeated
simultaneous mitoses in the growing centers. Resting nuclei contain conspicuous
karyosomes. Dividing ones show about five chromosomes in an intranuclear spindle
which ends sharply in centrosomes. Eventually, in the usual course of events, each
center produces an exit tube to the exterior. Its contents are released by delique-
scence of the tip of the exit tube. Either before this or afterward, the mass of proto-
plasm undergoes cleavage into uninucleate protoplasts which generate flagella. Some-
times, instead of discharging their contents, the centers are converted into resting
spores by the secretion of thick walls (this has been observed in only a few of the
species). The resting spores germinate by producing exit tubes and discharging
zoospores as ordinary centers do.
The body type which has just been described may be called the chytrid body
type; organisms of this body type were formerly assembled as a taxonomic group
typified by the genus Chytridium. Couch, however, showed that these organisms
form three groups distinguished by fundamental differences in type of flagellation.
The present group is here given a place implying relationship to order Silicoflagellata.
Karling (1943) accounted for fourteen species. He provided three families; only
one is here maintained.
Family Hyphochytriacea [Hyphochytriaceae] Fischer in Rabenhorst Kryptog.-Fl.
Deutschland 1, Abt. 4:131 (1892). Families Anisolpidiaceae and Rhizidiomycetaceae
Kailing in American Jour. Bot. 30: 641, 643 (1943). Characters of the order.
Without rhizoids: Anisolpidium on brown algae; Roesia on Lemna; Cystochytrium
on roots of Veronica. With rhizoids from a single center: Rhizidiomyces and Latr os-
tium on green algae, aquatic fungi, and the empty exoskeletons of insects. With
multiple centers: Hyphochytrium and Catenariopsis, on fungi and other hosts.
Class 2. BACILLARIACEA Engler and PrantI
Homalogonata Lyngbye Tent. Hydrog. Danicae 177 (1819).
Order Diatomeae C. Agardh Syst. Alg. xii (1824).
Division (of order ^/gae) Diatomaceae Harvey in Mackay Fl. Hibem. 166 (1836).
Family Bacillaria Ehrenberg Infusionsthierchen 136 (1838).
Series (of class Algae) Diatomaceae Harvey Man. British Alg. 15 (1841).
Abtheilung (of cldiS,?, Isocarpeae) Diatomaceae Kiitzing Phyc. Germ. 54 (1845).
Stamm Diatomea Haeckel Gen. Morph. 2: xxv (1866).
Division (of class Algae) Diatomaceae Rabenhorst Kryptog.-Fl. Sachsen 1: 1
(1863).
70]
The Classification of Lower Organisms
Fig. 12. — Hyphochytrialea: a-e^ Anisolpidium Ectocarpii; a-c, individuals de-
veloping in cells of Ectocarpus; d, mitotic figures x 2,000; e^ cell of Ectocarpus filled
by a mature individual discharging spores, f, g, Rhizidiomyccs apuphysatus; f, zoo-
spore; g, oogonium of Achlya parasitized by three individuals, h, i, j^ llyphochy-
trium catenoides; h, zoospore; i, young individual; j, mature individual with fila-
ments, sporangia, and zoospores in various stages of development. All after Karling
(1943, 1944, 1939). x 1,000 except as noted.
Phylum Phaeophyta [71
Class Bacillariaceae Engler and Prantl Nat. Pflanzenfam. II Teil; 1 (1889).
Subdivision and class Bacillariales Engler Syllab. 6 (1892).
Hauptclasse Diatomeae Haeckel Syst. Phylog. 1: 90 (1894).
Subclass Bacillariales Engler in Engler and Prantl Nat. Pflanzenfam. Teil I, Abt.
la: V (1900).
Class Bacillarieae Wettstein Handb. syst. Bot. 1: 74 (1901).
Class Bacillarioideae Bessey in Univ. Nebraska Studies 7: 283 (1907).
Class Diatomeae Schaffner in Ohio Naturalist 9: 447 (1909).
Abteilung Bacillariophyta Engler.
Ahteilung (of Stamm Chrysophyta) Diatomeae Pascher in Beih. bot. Centralbl.
48, Abt. 2: 324 (1931).
Class Bacillariophyceae Auctt.
Unicellular (occasionally filamentous or colonial) organisms without flagella in
the vegetative condition, each cell with one, two, or more plastids, brown, varying to
yellow or exceptionally to bluish or colorless, and bearing a siliceous shell of two
parts. Globules of oil and granules of something called volutin (the "red granules
of Biitschli," apparently protein) are present. Other granules in some examples are
said to be of leucosin.
These organisms, the diatoms, are very common. There are some 5300 species.
Microscopic examination of the bottoms of fresh water ponds reveals usually more of
diatoms than of any other kind of organisms. Diatoms are frequent prey of many kinds
of predators, from amoebas to whales. In using fish-liver oils as a source of vitamin
D, man adds himself to a long chain of predators of which it is believed that diatoms
are the usual ultimate prey.
The shells of diatoms are not subject to decay. In certain places which were in
the geologic past arms of the sea, there are enormous deposits of diatom shells in
the form of a white earth. The oldest deposits are of the Cretaceous age. Thus it ap-
pears that diatoms are a modern offshoot, no more ancient than the flowering plants.
Diatomaceous earth is mined for various uses. It is an effective insulating material,
and was the inert material first used in connection with nitroglycerine in the manu-
facture of dynamite.
The two parts of the shell of a diatom are called valves. They fit one over the
other "like the parts of a pill box" (ZoBell, 1941, objects to this traditional simile,
on the ground that in current language a pillbox is a concrete structure with loop-
holes). The shells consist basically of something of the nature of pectin heavily im-
pregnated with silica and characteristically sculptured. The cells appear markedly
different in different aspects: the aspect which is in effect top or bottom view is
called valve view, and that which is in effect side view is called girdle view. When a
cell divides, each of the daughter cells receives one of the valves and generates an
additional valve fitting within it. Diatoms in culture undergo a gradual diminution
in size; there is an old hypothesis that this is caused by the fact that one of each pair
of sister cells receives a slighly smaller valve than the other.
Lauterborn (1896) described mitosis in Surirella and other diatoms. He found a
centrosome, with radiating strands, near the nucleus. At the beginning of mitosis,
the centrosome generates a disk-shaped structure which enters the nucleus and grows
in such fashion as to become a cylinder extending through it. The cylinder is recog-
nizably a spindle, but the chromosomes, instead of appearing within it, form a ring-
shaped mass about its middle and divide into two ring-shaped masses which move
along it to its extremities. The nuclear membrane ceases to be recognizable early in
The Classification of Lower Organisms
Fig. 13. — Bacillariacea : a, Mclosira sp., a living cell and an empty one. b, c.
Girdle and valve views of cell of Cyclotella sp. d, e. Sections of a valve of Pinnu-
laria sp., highly magnified, after Otto Miillcr (1896); d, about half-way between the
middle and the end, e^ near the end. f, g, Girdle and valve views of Synedra sp.
h, i. Girdle and valve views of Rhoicosphenia curvata. j, k, Girdle and valve views
of Navicula sp. 1^ m, Girdle and valve views of Gomphonema sp. (the former show-
ing the gelatinous stalk by which the cell is attached), n, o. Girdle and valve views
of Cymbella sp. p, q, Surirella saxonica after Karsten (1900); p, two cells joined
before conjugation; q, zygote; x 250. r, s, Girdle and valve views of Cocconeis sp.
X 1,000 except as noted.
Phylum Phaeophyta [ 73
the process, but the nuclear cavity remains distinct until the chromosomes have
reached the ends of the spindle. The nuclear sap and the spindle are then absorbed
by the cytoplasm, but not until the spindle has budded off a new centrosome from
each end.
Subsequent authors, as Karsten (1900), Geitler (1927), Iyengar and Subrahman-
yan (1942, 1944), and Subrahmanyan (1947), have not seen as full a series of stages
as Lauterbom did. They have found centrosomes in at least some diatoms, and have
confirmed the point that the spindle is a cylinder which is surrounded by the
chromosomes instead of including them.
The same authors have described sexual processes in Surirella, Cymbella, Coc-
coneis, Cyclotella, and Navicula. In Surirella saxonica as described by Karsten, pairs
of the wedge-shaped cells become attached by little bodies of slime at the narrow
ends. Each nucleus divides twice, producing four, of which three are digested by the
cytoplasm. The two protoplasts then move in amoeboid fashion out of their shells
and they and their nuclei unite. The zygote protoplast grows to a size much greater
than that of the parent cells and secretes a membrane which becomes silicified. The
resulting cell is called an auxospore.
In most kinds of diatoms, each cell produces two gametes. In some, the cells pair
and proceed to produce auxospores individually, without conjugation. Karsten sup-
sposed the latter examples to represent a stage in the evolution of sexual reproduc-
tion under some zwingender Nothwendigkeit: much more probably, they are pro-
ducts of degeneration. In Cyclotella, Iyengar and Subrahmanyan found the produc-
tion of auxospores to involve autogamous karyogamy: the nucleus of a solitary cell
undergoes meiosis; two of the haploid nuclei are digested, and the two which remain
fuse with each other. It is evident that all diatoms are diploid in the vegetative
condition.
The filamentous green Heterokonta Tribonema and Bumilleria are closely similar
to the diatom Melosira, and it may reasonably be supposed that they represent the
evolutionary origin of the group.
Diatoms are preserved for study by violent methods which destroy the protoplasts,
and the classification is based strictly on characters of the shells. So uniform is the
group that Schiitt (in Engler and Prantl, 1896) treated it as a single family. He pro-
vided an elaborate subsidiary classification involving two main groups. Subsequent
scholars have found his system essentially sound as a representation of nature, but
have raised the main groups to the rank of orders and the minor ones in correspond-
ing degree.
Order 1. Disciformia [Disciformes] Kiitzing Phyc. Germ. 112 (1845).
Order Appendiculatae Kiitzing 1. c.
Centricae Schiitt in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 57
(1896).
Order Centricae Campbell Univ. Textb. Bot. 90 (1902).
Order Eupodiscales Bessey in Univ. Nebraska Studies 7: 284 (1907).
Diatoms basically of radial symmetry, which, however, is often distorted; not
motile in the vegetative condition; plastids numerous in the cells.
These are the more primitive diatoms. The majority are marine. Three types of
reproductive cells are known to be produced by them.
Occasionally, in mass catches of material from the ocean, diatoms are found
whose protoplasts have undergone repeated division within the shell and produced
74 ] The Classification of Lower Organisms
numerous little naked protoplasts. These protoplasts are said to bear flagella; whether
one or two, equal or unequal, is not certainly known. They are supposed to escape
and function as zoospores, but Karsten (1904), on rather scant evidence, supposed
them to be gametes.
A protoplast may contract and form a shell within its former shell. The new shell
consists like the old one of two parts, one fitting within the other. The outer shell is
usually more or less elaborately sculptured, while the inner is smooth. It is supposed
that the outer shell is deposited between outer and inner masses of protoplasm, and
that the entire protoplast then withdraws to the interior and deposits the inner shell
in the opening. It is in this manner that the statospores of chrysomonads are formed.
The resting cells of diatoms as just described are believed to be homologous with
them, and are called by the same term.
As a third manner of producing a reproductive cell, a protoplast may expand, force
apart the valves of its shell, and deposit an enlarged shell about itself. The resulting
spore is called an auxospore. As noted, Iyengar and Subrahmanyan found the pro-
duction of auxospores in Cyclotclla to involve sexual processes.
Schiitt divided the Centricae into three groups with names in -oideae (presum-
ably subfamilies) and these into nine groups with names in -eae (presumably tribes).
Subsequent authorities have made of Schiitt's groups a varying number of families.
The minimum tenable number of families is three, corresponding to Schutt's
subfamilies.
Family 1. Coscinodiscea [Coscinodisceae] Kiitzing Phyc. Germ. 112 (1845).
Family Melosireae Kiitzing op. cit. 66. Families Melosiraceae and Coscinodiscaceae
West British Freshw. Alg. 274, 276 (1904). Melosira, in fresh water, the shells feebly
silicified, the cells joined end to end in filaments. Cyclotclla, separate drum-shaped
cells in fresh water. Coscinodiscus, the cells disk-shaped. Triceratium, cells of the
form of 3-, 4-, or 5-sided prisms with abbreviated axes.
Family 2. Rhizosoleniacea [Rhizosoleniaceae] West British Freshw. Alg. 278
(1904). The cells, circular or elliptic in cross section, becoming elongate by inter-
calation of ring-shaped bands of wall between the valves. Rhizosolenia. Corethron.
Family 3. Biddulphiea [Biddulphieae] Kutzing Phyc. Germ. 115 (1845). Families
Biddulphiaceae and Chaetoceraceae Auctt. Cells laterally compressed, elliptic in
valve view, oblong or rhombic in girdle view. Cells of Biddulphia, solitary or colonial,
are familar as epiphytes on marine algae. Chaetoceros, the cells with a long spine at
each corner, frequently united valve to valve in filaments, abundant in subpolar
oceans.
Order 2. Diatomea [Diatomeae] C. Agardh Syst. Alg. xii (1824).
Tribe Striatae with orders Astomaticae and Stomaticae, and tribe Vittatae also
with orders Astomaticae and Stomaticae, Kutzing Phyc. Germ. ( 1845).
Pennatae Schiitt in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 101
(1896).
Order Pennatae Campbell Univ. Textb. Bot. 90 ( 1902).
Order Naviculales Bessey in Univ. Nebraska Studies 7: 284 ( 1907).
Diatoms basically of isobilateral symmetry, occasionally so skewed as to be dorsi-
ventral or asymmetric; valves usually punctured by a longitudinal cleft called the
raphe, or bearing a marking of some sort, called the pseudoraphe, in the same posi-
tion; exhibiting, when possessed of a true raphe, a gliding motion; cells usually with
two plastids.
Phylum Phaeophyta [ 75
The motion of the pennate diatoms is a gliding upon surfaces, with frequent re-
versal, in either direction of the long axis of the cell. It depends upon the flow of a
stream of exposed protoplasm. This is the opinion of Max Schultze (1865), Otto
Miiller (1889, 1896), and Lauterborn (1896); there have been other hypotheses.
Miiller showed that the true raphe, without which the motion does not occur, is an
actual opening. The raphe is not a simple crack; it enters the wall obliquely and
bends at a sharp angle to come from another oblique direction to the interior. Its
proportions vary along its length, and it is interrupted at the middle of the valve by
a knob, the central granule, projecting inward from the valve.
The pennate diatoms do not produce flagellate cells nor statospores, but they pro-
duce auxospores, usually by sexual processes. The majority inhabit fresh water.
Eleven families are currently recognized.
a. Without raphes.
Family 1. Fragilariea [Fragilarieae] (Harvey) Kutzing Phyc. Germ. 62 (1845).
Family Fragilariaceae West British Freshw. Alg. 285 (1904). Cells symmetrical with
respect to three planes, without internal partitions. Fragilaria. Synedra.
Family 2. Tabellariea [Tabellarieae] Kutzing op. cit. 110. Family Tahellariaceae
West op. cit. 281. Cells symmetrical with respect to three planes, with longitudinal
internal partitions. Tabellaria.
Family 3. Bacillaria Ehrenberg Infusionsthierchen 136 (1838). Family Diato-
maceae West op. cit. 284. Cells symmetrical with regard to three planes, with trans-
verse internal partitions, solitary, or joined valve to valve in ribbons, or corner to
comer in zig-zag chains. Diatoma.
Family 4. Meridiea [Meridieae] Kutzing op. cit. 61. Family Meridionaceae West
op. cit. 283. Cells symmetrical with regard to two planes, wedge-shaped both in valve
and in girdle view, with transverse internal partitions, often joined valve to valve
in fan-shaped colonies which are sometimes so extended as to produce spiral fila-
ments. Meridion.
b. With raphes, the valves of each cell alike.
Family 5. Naviculea [Naviculeae] Kiitzing op. cit. 90. Family Naviculaceae Rab-
enhorst Kryptog.-Fl. Sachsen 1: 33 (1863). This is the most numerous family of
diatoms. In most of the genera the cells are narrowly rectangular in girdle view,
narrowly elliptic in valve view, being of the shape of flat-bottomed boats. Navicula,
Pinnularia, etc. In other genera, as Gyrosigma and Pleurosigma, the cells are so
skewed as to be sigmoid in valve view.
Family 6. Gomphonemea [Gomphonemeae] Kiitzing op. cit. 87. Family Gom-
phonemaceae West op. cit. 297. Cells wedge-shaped. Gomphonema.
Family 7. Cymbellea [Cymbelleae] (Harvey) Kiitzing op. cit. 84. Family Cocco-
nemaceae West op. cit. 298. Cells with two planes of symmetry, in valve view crescent-
shaped or approximately so. Cymbella. Rhopalodia.
Family 8. Eunotiea [Eunotieae] Kiitzing op. cit. 57. Family Eunotiaceae West op.
cit. 287. Cells curved as in the preceding family, the raphes reduced to brief clefts
near the ends of the valves. Eunotia.
Family 9. Nitzschiacea [Nitzschiaceae] West op. cit. 301. Cells asymmetric in
valve view, the raphe along one margin. Nitzschia. Hantschia.
Family 10. Surirellea [Surirelleae] Kiitzing op. cit. 70. Family Surirellaceae West
op. cit. 303. Each cell with two marginal raphes. Surirella.
c. The two valves of each cell unlike, one with a raphe, one with a pseudoraphe.
76 ] The Classification of Lower Organisms
Family 11. Achnanthea [Achnantheae] Kiitzing op. cit. 81. Families Achnan-
thaceae and Cocconeidaceae West op. cit. 289, 290. Achnanthes, Rhoicosphenia,
Cocconeis.
Class 3. OOMYCETES Winter
Class OoMYCETEs Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1: 32
(1879).
Phycomyceten de Bary Vergl. Morph. Pilze 142 ( 1884), in part.
Class Phycojnycetes Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889), in
part.
Reihe Oomycetes Fischer in Rabenhorst Kryptog.-FI. Deutschland 1, Abt. 4: 310
(1892).
Stamm Phykomycophyta Pascher in Beih. bot. Centralbl. 48, Abt. 2: 330 (1931),
in part.
Biflagellatae Sparrow Aquatic Phycomycetes 487 (1943).
Organisms of fungal or chytrid body type, that is, non-pigmented saprophytes or
parasites whose bodies are walled filaments or cells with or without rhizoids; the
walls consisting partially of cellulose; reproducing asexually by zoospores with
paired unlike flagella which are, so far as is known, respectively pantoneme and
acroneme, and usually sexually by fertilization, the eggs being distinct cells within
the oogonia. The regularly cited example and evident standard genus of the group
is Saprolegnia.
Conventional botanical classification recognizes within the group of Fungi a sub-
ordinate group named Phycomycetes, which is in turn divided into Oomycetes and
Zygomycetes, the former including the chytrids. This arrangement suggests an evo-
lutionary series, originating perhaps among non-pigmented flagellates, and leading
through chytrids, typical Oomycetes, and Zygomycetes to the typical fungi. It does
not now appear tenable. Couch (1939) pointed out differences between Oomycetes
and Zygomycetes which make any direct connection between them appear quite
improbable; and his observations on flagella showed that only a small minority among
organisms of chytrid body type have anything to do with the proper Oomycetes.
There is an old hypothesis (Sachs, 1874) that Vauchcria may represent the direct
ancestry of Saprolegnia. This hypothesis could not be taken seriously while Sapro-
legnia and its allies were known to produce heterokont zoospores, while Vaucheria
was supposed to be a typical isokont green alga. Now it again appears probable. It
implies that in the present group the fungal body type is more primitive than the
chytrid.
The Oomycetes may be organized as three orders.
l.Of fungal body type, i.e., consisting of fila-
ments.
2. Essentially aquatic Order 1. Saprolegnina.
2. Mostly not aquatic, parasitic on higher
plants Order 2. Peronosporina.
1. Of chytrid body type, i.e., the cells not elong-
ated to filamentous form, though sometimes
proliferating or producing rhizoids Order 3. LAGENroiALEA.
Phylum Phaeophyta [ 77
Order 1. Saprolegnina [Saprolengninae] Fischer in Rabenhorst Kryptog.-Fl.
Deutschlandl,Abt.4: 311 (1892).
Order Eremospermeae and suborder Mycophyceae Kiitzing Phyc. Gen. 146
(1843), in part.
Order Oosporeae Cohn in Hedwigia 11: 18 (1872), in part.
Order Oomycetes and suborder Saprolegniineae Engler Syllab. 24 (1892).
Order Saprolegniineae Campbell Univ. Textb. Bot. 153 (1902).
Order Siphonomycetae Bessey in Univ. Nebraska Studies 7: 286 (1907).
Order Saprolegniales Auctt.
Order L^p^omzfa/^?^- Kanouse in American Jour. Bot. 14: 295 (1927).
Aquatic Oomycetes, filamentous, saprophytic or facultatively parasitic, the zoo-
spores diplanetic (exhibiting two periods of swimming) or giving evidence of an
ancestral diplanetic condition. The old ordinal names Eremospermeae and Oosporeae
designated miscellaneous collections of groups in which this one was listed at or near
the beginning. Either one, if taken up, would be applied here, but it seems better to
treat them as nomina confusa.
1. Filaments not constricted Family 1. Saprolegniea.
1. Filaments constricted at intervals.
2. Filaments not differentiated into basal
and reproductive parts Family 2. Leptomitea.
2. Filaments differentiated into basal and
reproductive parts Family 3. Rhipidiacea.
Family 1. Saprolegniea [Saprolegnieae] Kiitzing Phyc. Gen. 157 (1843). Family
Saprolegniaceae Cohn in Hedwigia 11 : 18 (1872). Aquatic Oomycetes consisting of
branching filaments of essentially uniform diameter without crosswalls other than
those which set apart differentiated reproductive structures.
These well-known organisms are called water molds. According to Coker (1923)
there are about eighty definitely recognizable species. They may be parasitic on
fishes or saprophytic on organic remains in water or soil. In almost any body of soil
or of fresh water they may be found by "baiting," in former practice with dead flies,
currently with hemp seeds.
Mitosis has rarely "been observed in the vegetative filaments, the nuclei being very
minute. Eggs are produced in large globular multinucleate oogonia borne at the ends
of filaments. The nuclei in the developing oogonia become enlarged and undergo a
single flare of concurrent mitoses (Davis, 1903; Couch, 1932). The sharp-pointed
spindles, ending in centrosomes, are formed within the nuclear membrane. The
membrane disappears toward the end of the mitotic process, and a nucleolus, which
has persisted to this stage, undergoes solution in the cytoplasm. The chromosome
numbers (Ziegler, 1953) are 3, 4, 5, 6, or 7.
Within each oogonium there appear one or a few minute bodies called coenocentra.
One nucleus becomes associated with each coenocentrum; all others break down and
disappear. Each surviving nucleus with the cytoplasm associated with it becomes
organized as an egg. When several eggs are produced, they share all of the cytoplasm
of the oogonium; when only one egg is produced, some of the cytoplasm is left out-
side of it.
Sperms are produced in small multinucleate antheridia borne at the tips of fila-
ments in contact with oogonia. Typically, each individual bears both oogonia and
antheridia. Some species are capable of self-fertilization; others exist as two kinds
of individuals, each capable of fertilizing the other; some occur as distinct male and
78]
The Classification of Lower Organisms
,^t*SJi
' •ITi''!' nVgi 'Villi
Fig. 14. — Oomycetes: a. Filaments and sporangia of Dictyuchus sp. x 50.
b, C, Zoospores of the second stage of swimming, of Achlya caroliniana and Sapro-
legnia ferax, after Couch (1941) x 1,000. d^ Oogonia and antheridia of Dictyuchus
X 400. e, f, g, Saprolegnia mixta after Davis (1903) : e, developing oogonium with
numerous nuclei x 500; f, metaphase of nuclear division x 2,000; g, developing
oogonium in which most of the nuclei have undergone degeneration; a few have
become associated with coenocentra, and the cytoplasm is undergoing cleavage to
produce eggs about these.
Phylum Phaeophyta [ 79
female individuals. Parthenogenesis (reproduction by eggs which have not been
fertilized) is rather common in this group. There are no swimming sperms: nuclei
from the antheridia reach the eggs through fertilization tubes, or by migration through
the periplasm.
Ziegler found that the first nuclear divisions of the nucleus of the zygote are
meiotic: all cells except the zygotes are haploid.
The organs of asexual reproduction are cylindrical sporangia terminal on the fila-
ments. Within these the multinucleate protoplasts undergo cleavage into minute
uninucleate spores. It is chiefly by details of the behavior of the sporangia and spores
(the latter diplanetic, monoplanetic, or not swimming at all) that the dozen genera
are distinguished. Diplanetism is the character of zoospores which are not directly
infective; they undergo encystment, and the cysts release infective zoospores. During
the first stage of swimming, the spores are pear-shaped, with the nucleus drawn out
into a beak toward the narrow anterior end, where the flagella are attached. Spores re-
leased from cysts for a second period of swimming are bean-shaped, with the flagella
attached laterally, each connected through a separate rhizoplast to the nucleus, which
lies at some distance from the cell membrane (Cotner, 1930). No explanation of
this behavior, whether by phylogeny, genetics, physiology, or competitive advantage,
is known. The apparent trend of evolution is to eliminate it. Monoplanetic spores
in the present group are usually released from the sporangia as naked protoplasts
which undergo encystment and emerge subsequently as flagellate spores of the second
form.
Saprolegnia releases diplanetic spores through circular pores in the tips of sporangia
in which the spores are formed in several rows; new sporangia develop within empty
old ones. Organisms which differ from Saprolegnia only in producing new sporangia
beside, instead of within, the old ones, were formerly assigned to Achlya, but are now
called Isoachlya. Leptolegnia differs from Saprolegnia and Isoachlya in forming
spores in a single row. In Achlya proper, the spores are discharged without flagella,
to encyst and swim only once. In Thraustotheca the monoplanetic spores are re-
leased by irregular breakdown of the distal part of the sporangium. In Dictyuchus
the spores become encysted before discharge; their protoplasts escape in the form of
secondary swarmers through individual pores in the wall of the sporangium. Salvin
(1942) found that cultures while growing release into the medium substances which
affect the type of sporangium produced, so that a given culture may be while young
of the character of Achlya, and later of the character of Thraustotheca or Dictyuchus.
Family 2. Leptomitea [Leptomiteae] Kiitzing Phyc. Gen. 150 (1843). Family
Leptomitaceae Schroter in Engler and Prantl Nat. Pflanzenfam. I Tail, Abt. 1 : 101
(1893). Oomycetes consisting of filaments which are constricted at intervals, but
are not differentiated into a basal cell and reproductive branches. In sewage or on
organic matter decaying in water. Leptomitus, Apodachlya, Apodachlyella, with
some seven known species. The numbers of species and degree of distinction of this
family and the following do not appear to justify the proposed establishment of a
separate order for them.
Family 3. Rhipidiacea [Rhipidiaceae] Sparrow in Mycologia 34: 116 (1942).
Saprophytes resembling the Leptomitea, the body differentiated into a main part, the
basal cell, rhizoids of limited growth, and slender branches bearing the reproductive
structures. Sapromyces, Araiospora, Rhipidium, Mindeniella, with perhaps a dozen
known species.
80 ] The Classification of Lower Organisms
Order 2. Peronosporina [Peronosporinae] Fischer in Rabenhorst Kryptog.-Fl.
Deutschlandl,Abt.4: 383 (1892).
Suborder Peronosporineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1: iv (1897).
Order Peronosporineae Campbell Univ. Textb. Bot. 155 (1902).
Order Peronosporales Auctt.
Mostly parasites on terrestrial plants, but including also aquatic parasites and a
few saprophytes, the bodies filamentous, reproducing sexually by fertilization, the
eggs solitary in the oogonia, reproducing asexually chiefly by conidia, that is, by air-
born cells cut off from the ends of the filaments. The conidia are homologous with
the sporangia of the Saprolegnina : they germinate in most examples by release of
zoospores (which show no signs of diplanetism), but in the more highly evolved
examples they give rise to filaments. Ferris (1954) found the zoospores of Phytoph-
thora to bear the paired flagella, respectively pantoneme and acroneme, which are
typical of Phaeophyta.
In the multinucleate oogonia of most members of the group, single flares of mitoses
occur. The sharp-pointed spindles, described in some accounts as ending in centro-
somes, are formed within the persistent nuclear membrane, which undergoes con-
striction during the final stages of mitosis. A coenocentrum appears (this structure
was first described as occurring in Albugo, by Stevens, 1899); in general, one nucleus
becomes associated with it, and is thus selected as the egg nucleus, the remaining
nuclei being cast out to undergo disolution in a body of periplasm. The antheridium
develops in contact with the oogonium, and fertilization is accomplished by the
growth of a fertilization tube through the periplasm to the egg (Davis, 1900; Stevens,
1899,1901,1902).
In Albugo Bliti and A. Tragopogonis, Stevens observed two flares of simultaneous
mitoses in the oogonium and antheridium. If this phenomenon were general in the
group one would confidently identify it as meiosis. The single coenocentrum attracts
many nuclei; the fertilization tube delivers a large number of sperm nuclei; thus
multiple karyogamy occurs within a single cell. The further history of the resulting
peculiar zygote, containing many nuclei which are not by any evident necessity
genetically uniform, is unknown.
This order is evidently a specialized offshoot of the preceding. The family Pythiacea
is a good example of a transition group; many authorities have assigned it to the pre-
ceding order.
1. Producing solitary globular sporangia or
conidia at the ends of scarcely specialized
filaments; mostly aquatic Family 1. Pythiacea.
1. Producing conidia usually in clusters at the
ends of specialized filaments (conidio-
phores) ; parasites on land plants.
2. Conidiophores brief, unbranched, the
conidia in chains Family 2. Albuginacea.
2. Conidiophores elongate, usually branch-
ed, the conidia solitary or clustered, not
in chains Family 3. Peronosporacea.
Family 1. Pythiacea [Pythiaccae] Schroter in Engler and Prantl Nat. Pflanzenfam.
I Teil, Abt. 1: 104 (1893). Aquatic parasites and saprophytes releasing zoospores
from globular reproductive structures terminal on the filaments, together with para-
Phylum Phaeophyta [81
sites attacking land plants under moist conditions. The reproductive structures act
as sporangia if formed in water, as conidia if formed in air. Pythium, saprophytic on
plant remains in water or parasitic on algae or higher plants, includes some forty
species (Matthews, 1931). The few other genera include perhaps a dozen species.
Zoophagus produces specialized branches which serve as traps for rotifers which are
parasitized and killed.
Family 2. Albuginacea [Albuginaceae] Schroter op. cit. 110. Parasites of higher
plants, called white rusts, the masses of conidia which push up and burst through the
epidetmis being of a white color. Albugo.
Family 3. Peronosporacea [Peronosporaceae] Cohn in Hedwigia 11: 18 (1872).
Parasites of higher plants, called downy mildews. The ovoid conidia are produced
solitary or in clusters, not in chains, on elongate conidiophores, usually branched,
projecting through the stomata of the hosts. This numerous group includes the
agents of some of the most important diseases of cultivated plants. Plasmopara viti-
cola, causing downy mildew of grapes. Phytophthora injestans, the cause of the blight
of potatoes which produced the Irish famine of 1846. Peronospora, the many species
attacking many kinds of plants.
Order 3. Lagenidialea [Lagenidiales] Karling in American Jour. Bot. 26: 518
(1939).
Suborder Ancylistineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1 : iv ( 1897), for the most part, not as to the type genus Ancylistes.
Order Ancylistales Auctt., in part.
Oomycetes of chytrid body type, parasites consisting of walled cells which are
more or less isodiametric, sometimes proliferating or producing rhizoids, but not
forming extensive branched filaments. The cells become multinucleate. Mitotic
figures of Olpidiopsis as described by Barrett (1912) and McLarty (1941) are quite
as in the preceding orders, with sharp-pointed intranuclear spindles apparently with
centrosomes at the poles. In the usual course of events, each cell develops an exit
tube to the exterior of the host, and the protoplast becomes divided into uninucleate
cells which escape as unequally biflagellate zoospores. Fertilization, by the migration
of the protoplast of one cell into another, has been observed; the zygote becomes a
thick-walled resting spore.
1. Internal parasites without rhizoids.
2. The cells not proliferating.
3. The zoospores diplanetic Family 1. Ectrogellacea.
3. The zoospores not diplanetic Family 2. Olpidiopsidacea.
2. The cells proliferating.
3. Marine Family 3. Sirolpidiacea.
3. Fresh-water Family 4. Lagenidiacea.
1. External parasites with rhizoids Family 5. Thrau stock ytriacea.
Family 1. Ectrogellacea [Ectrogellaceae] Scherffel in Arch. Prot. 52: 6 (1925).
Ectrogella, Eurychasma, Eurychasmidium, Aphanomycopsis, with about a dozen
known species, attacking diatoms and red and brown algae.
Family 2. Olpidiopsidacea [Olpidiopsidaceae] Sparrow in Mycologia 34: 116
(1942). Olpidiopsis and a few other genera, with some thirty known species, attack-
ing water molds, green algae, red algae, and other aquatic organisms.
Family 3. Sirolpidiacea [Sirolpidiaceae] Sparrow 1. c. Sirolpidium and Pontisma,
each with one species, attacking marine algae, respectively green and red.
82 ] The Classification of Lower Organisms
Family 4. Lagenidiacea [Lagenidiaceae] Schroter in Engler and Prantl Nat.
Pfianzenfam. I Teil, Abt. 1 : 89 (1893). Lagenidium, Myzocytium, and Lagenocystis^,
with some twenty known species, attacking green algae, rotifers, pollen which has
fallen into water, and the roots of grasses.
Family 5. Thraustochytriacea [Thraustochytriaceae] Sparrow op. cit. 115. The
single species Thraustochytrium proliferum Sparrow was found as solitary cells ex-
ternal on certain marine green algae and red algae which are penetrated by means
of branching rhizoids. Reproduction is by release of naked protoplasts which become
laterally biflagellate after a period of rest.
Class 4. MELANOPHYCEA (Ruprecht) Rabenhorst
Order Fucacees Lamouroux in Ann. Mus. Hist. Nat. Paris 20: 28 (1813).
FucoiDEAE C. Agardh Synops. Alg. Scand. ix (1817).
Order Fucoideae C. Agardh Syst. Alg. xxxv (1824).
Division (of order Algae) Melanospermeae Harvey in Mackay Fl. Hibern. 157
(1836).
Series (of order Algae) Melanospermeae Harvey Man. British Alg. 1 (1841).
Order Pycnospermeae and tribe Angiospermeae Kiitzing Phyc. Gen. 333, 349
(1843).
Class Fucoideae J. Agardh Sp. Alg. 1 : 1 (1848).
Melanophyceae Ruprecht in Middendorff Sibir. Reise 1, part 2: 200 (1851).
Class Melanophyceae Rabenhorst Kryptog.-Fl. Sachsen 1: 275 (1863).
Stamm Fucoideae Haeckel Gen. Morph. 2: xxxv (1866).
Series {Reihe) Phaeophyceae Hauck in Rabenhorst Kryptog.-Fl. Deutschland 2:
282 (1885).
Class Phaeophyceae Engler and Prantl Nat. Pfianzenfam. H Teil: 1 (1889),
Class Dictyotales Engler in Engler and Prantl Nat. Pfianzenfam. I Teil, Abt. 2 :
ix(1897).
Classes Phaeosporeae, Tetrasporeae, and Cyclosporeae Bessey in Univ. Nebraska
Studies 7: 288, 290 (1907).
CXdiSS Dictyoteae Schaffner in Ohio Naturalist 9: 448 (1909).
Subclass Melanophyceae Setchell and Gardner in Univ. California Publ. Bot. 8:
387 (1925).
Classes Isogeneratae, Heterogeneratae (with subclasses Haplostichinae and Poly-
stichinae) and Cyclosporeae Kylin in Kungl. Fysiog. Sallsk. Handl. n. f. 44, no.
7: 91 (1933).
Filamentous or thallosc Phaeophyta, yellow to brown in color and living by photo-
synthesis, producing reproductive cells with paired unequal flagella.
These are the typical brown algae. They are almost exclusively marine, being
abundant along with red and green algae on most coasts, and particularly abundant
farther toward the poles than the red and green groups. The lower brown algae are
branched filaments of microscopic dimensions, commonly epiphytic on other algae.
More highly developed examples are thallosc and anchored to rocks. Some of these,
particularly the ones whose English name is kelp, reach great sizes and considerable
elaboration of structure. Papenfuss (in Smith, 1951) gives the number of genera as
about 240, and that of known species as about fifteen hundred.
^Lagenocystis nom. nov. Lagena Vanterpool and Ledingham in Canadian
Jour Res. 2: 192 (1930), non Parker and Jones 1859. L. radicicola (Vanter-
pool and Ledingham) comb. nov.
Phylum Phaeophyta [ 83
The cells are walled chiefly with readily hydrolyzable modified polysaccharides.
Algin, the soda extract of kelps, consists of chains of oxidized mannose units. A poly-
saccharide of the sugar fucose, with a sulfate radicle to each sugar unit, is also present.
A small percentage of cellulose is present, apparently as the immediate investment
of each protoplast. A glycogen- or dextrin-like dextrosan, laminarin, is stored (Miwa,
1940; Tseng, 1945). The plastids contain chlorophylls a and c (Strain, in Franck
and Loomis, 1949) and carotin; xanthophyll is also present in the more primitive
examples. In all examples, there is an additional carotinoid called fucoxanthin, which
produces the brown color. The analytic process of separating the pigments yields
also a sterol, fucosterol, not found in green plants; but this substance, and fucoxanthin,
are found in chrysomonads, green Heterokonta, and diatoms (Carter, Heilbron, and
Lythgoe, 1939).
Cytological study of a considerable variety of brown algae (Swingle, 1897; Farmer
and Williams, 1896; Mottier, 1898, 1900; Simons, 1906; Yamanouchi, 1909, 1912;
McKay, 1933) has shown that the spindle and chromosomes appear within an intact
nuclear membrane which disappears during the later stages of division. A centrosome,
usually with radiating rays, is present outside of the membrane at each pole of the
spindle. In Stypocaulon, a comparatively primitive brown alga, Swingle found the
centrosome to be a permanent structure, dividing as a preliminary to each division
of the nucleus. In the generality of brown algae, the centrosomes appear de novo as
division begins.
Swimming cells are produced by primitive brown algae as spores and as morpholo-
gically undifferentiated gametes; in the most advanced brown algae, such cells are
produced only as sperms. The flagella are attached laterally. The anterior flagellum
is the longer except in order Fucoidea (Kylin, 1916). Longest (1946) found in
Ectocarpus that the anterior flagellum is pantoneme, and the posterior one acroneme.
The swimming cells are without walls, and contain, beside the nucleus, usually one
plastid and a light-sentitive speck, the stigma or eyespot. They are quite small. No
system of structures linking the nuclei, centrosomes, and flagella has been discovered.
Thuret (1850) discovered that most brown algae produce swimming cells from
structures of two different sorts, which he named (1855) respectively plurilocular
sporangia and unilocular sporangia. The difference between them is this. In the
developing plurilocular structure, each division of the nucleus is followed by division
of the protoplast and deposition of a wall, with the result that the swimming cells
emerge from separate walled spaces. In the unilocular structure, the nucleus divides
repeatedly before the protoplast divides; the protoplast then undergoes cleavage to
produce swimming cells which emerge from a single walled space. A number of
studies (Clint. 1927; Higgins, 1931; Knight, 1923, 1929) have shown that the first
two nuclear divisions in the unilocular structure are normally meiotic. Unilocular
structures occur normally only on diploid individuals and release haploid swimming
cells. A few exceptional species, however, are known to bear unilocular structures
which produce swimming cells without the intervention of meiosis.
In Ectocarpus siliculosus as studied by Berthold (1881) at Naples, the swimming
cells from unilocular structures are spores which give rise to haploid individuals. In
the same species as studied in the Irish Sea by Knight (1929), they were found to
act as gametes, conjugating and giving rise to diploid individuals. Diploid and hap-
loid individuals of Ectocarpus are alike, and E. siliculosus may be said to have a
facultatively complete homologous life cycle. The haploid individuals produce pluri-
locular reproductive structures; the swarmers from these act either as spores, re-
84]
The Classification of Lower Organisms
Fig. 15. — Stages of nuclear division in Stypocaulon x 1,000 after Swingle (1897).
Phylum Phaeophyta [ 85
producing the haploid stage, or as gametes, initiating the diploid stage. The diploid
individuals produce both plurilocular and unilocular reproductive structures. The
swarmers from the former are spores, reproducing the diploid body. The swarmers
from the latter act either as spores, giving rise to haploid individuals, or as gametes,
reproducing the diploid body.
It is believed that the brown algae arose by evolution from order Ochromonadalea.
Filamentous organisms with a facultatively complete homolgous life cycle, as just
described, are believed to be primitive among them : such organisms appear to be the
starting point of evolution in many features. The filaments have become differentiated
and woven into thalli, and thalli of tridimensionally placed cells have been produced.
The haploid and diploid stages have become differentiated. The plurilocular and
unilocular structures have undergone specialization. Even in the most primitive
brown algae, there is a physiological differentiation of gametes; this has evolved into
extreme morphological differentiation. Every one of these evolutionary changes ap-
pears to have occurred in more than one line of descent; research is constantly reveal-
ing intermediate examples and rather free parallel evolution.
Conservative classification, such as that of Fritsch (1945), recognizes as orders a
comparatively primitive miscellany followed by a series of small derived groups
marked by distinctive specializations. Features of the life cycle, as applied to classi-
fication by Taylor (1922), Oltmanns (1922), Svedelius (1929) and Kylin (1933),
are not reliable as marks of natural groups. Kylin provided three classes (one of
them divided into two subclasses) and twelve orders. His system appears to provide
an excessive number of subdivisions of high category within a moderately small group
exhibiting no very profound evolutionary gaps. Tentatively, the seven orders dis-
tinguished as follows may be recognized.
1. Producing spores, that is, cells which germi-
nate without syngamy.
2. All spores bearing flagella.
3. Having an alternation of haploid
and diploid stages which are alike,
both being filamentous; or else com-
pletely lacking one of these stages.
4. The filaments uniseriate Order 1. Phaeozoosporea.
4. The filaments becoming pluri-
seriate Order 2. Sphacelarialea.
3. Not as above.
4. Haploid stage thallose, not dis-
tinctly less highly developed
than the diploid stage Order 5. Cutlerialea.
4. Haploid stage filamentous, dis-
tinctly less highly developed
than the diploid stage.
5. Diploid stage filamentous;
or, if partially or com-
pletely thallose, the thal-
lose part with apical growth Order 4. SpoROCHNoroEA.
5. Diploid stage thallose, its
growth intercalary Order 6. Laminariea.
2. Producing large non-motile spores Order 3. Dictyotea.
86 ] The Classification of Lower Organisms
1. Producing no spores; all individuals diploid
and reproducing exclusively sexually Order 7. FucoroEA.
Order 1. Phaeozoosporea [Phaeozoosporeae] Hauck in Rabenhorst Kryptog.-Fl.
Deutschland 2: 312 (1885).
Order Syntamiidae Areschoug in Act. Reg. Soc. Upsala 14: 387 ( 1850) , in part;
a nomen confusum.
Order Ectocarpeae J. Agardh Sp. Alg. 1: 6 (1848), preoccupied by family
EcTOCARPEAE KUtzing (1843).
Section (of Algae Zoosporeac) Phaeosporeae Thuret in Ann. Sci. Nat. Bot.
ser. 3, 14: 233 (1850).
Order Phaeosporeae Wettstein Handb. syst. Bot. 1: 173 (1901).
Order Ectocarpales Bessey in Univ. Nebraska Studies 7: 288 (1907).
Order Phaeosporales and suborder Ectocarpineae Taylor in Bot Gaz. 74: 435,
436 (1922).
Microscopic brown algae of the form of undifferentiated uniseriate branching fila-
ments, mostly with distinct haploid and diploid stages (exceptionally lacking the
former), the stages distinguishable only by the limitation of unilocular reproductive
structures to the diploid stage, the gametes morphologically uniform.
The order is typified by Ectocarpus, which is by coincidence also the theoretical
ancestral type of the brown algae, the living organism which supposedly represents
the evolutionary origin of the group. Recent systems of classification limit this order,
formerly construed as extensive, to this genus and a few others, as Pylaiella and Streb-
lonema, which make up the family Ectocarpea [Ectocarpeae] Kiitzing (family Ecto-
carpaceae Cohn).
Order 2. Sphacelarialea [Sphacelariales] (Oltmanns) Engler and Gilg Syllab. ed.
9 u. 10: 27 (1924).
Order Sphacelarieae J. Agardh Sp. Alg. 1: 27 (1848), preoccupied by family
Sphagelarieae Kiitzing (1843).
Sphacelariales Oltmanns Morph. u. Biol. Alg. ed. 2, 2: 2 (1922).
Brown algae distinguished from the Ectocarpea only by features of the vegetative
structure, namely that the filaments have large apical cells, and that the cells cut off
from them divide lengthwise without increasing considerably in thickness, with the
result that the filaments consist of tiers of cells. The life cycle is the same as in Ecto-
carpea. Family Sphacelariea [Sphacelarieae] Kiitzing (family Sphacelariaceae Cohn)
includes Sphacelaria and Stypocaulon. A few other families have been segregated.
Order 3. Dictyotea [Dictyoteae] Greville Alg. Brit. 46 (1830).
Tribe Dictyoteae Harvey in Mackay Fl. Hibern. 159 (1836).
Family Dictyoteae Kiitzing Phyc. Gen. 337 (1843).
Order Dictyotaceae Hauck in Rabenhorst Kryptog.-Fl. Deutschland 2: 302
(1885).
Class Dictyotales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2 :
ix (1897).
Akinetosporeae Oltmanns Morph u. Biol. Alg. 1: 473 (1904).
Order Tilopteridales and Class Tetrasporeae with order Dictyotales Bessey in
Univ. Nebraska Studies 7: 290 (1907).
Scries Aplanosporeae Setchell and Gardner in Univ. California Publ. Bot. 8:
649 (1925).
Phylum Phaeophyta [ 87
Filamentous or thallose brown algae with haploid and diploid stages equally de-
veloped, producing large spores without flagella, solitary or few in the sporangia.
Here are placed two families, Tilopteridea and Dictyotacea.
Family Tilopteridea [Tilopterideae] Cohn is a small group, apparently known
only from European coasts. They are evidently closely related to the Ectocarpea.
They consist of branching filaments which may become pluriseriate. In Haplospora
(poorly known; but Tilopteris and other genera are even more so), the haploid stage
bears both plurilocular structures, releasing minute swimming cells of the structure
usual in brown algae, and unilocular structures which release their contents as single
uninucleate protoplasts without flagella. The diploid stage bears only unilocular
structures which release their contents as single quadrinucleate non-motile spores. It
is inferred that the swimming cells from the plurilocular structures are sperms, and
that the protoplasts released from the unilocular structures on haploid bodies are
eggs, capable, however, of reproducing the haploid stage if not fertilized; further,
that the nuclei of the quadrinucleate spores released by diploid individuals are hap-
loid, and become on germination the nuclei of as many cells of the haploid body.
The Tilopteridea are believed to represent the evolutionary transition between
Ectocarpea and the following family.
Family Dictyotacea [Dictyotaceae] (Hauck) Kjellmann includes about twenty
genera, Dictyota, Zonaria, Padina, etc., with about one hundred species which are
commonest on the coasts of warmer oceans. They are thalli of moderate size, erect
and dichotomously branched or appressed and fan-shaped. They grow by the division
of a single apical cell or a row of apical cells in each branch. The cells multiplying
behind the apical cells become differentiated into two tissues, superficial small cells
rich in plastids and internal larger ones with fewer plastids, forming in different
species single or multiple layers of cells.
The Hfe cycle has been studied by Mottier (1898, 1900), Williams (1898), and
Haupt (1932). There are distinct male haploid individuals, female haploid indivi-
duals, and diploid individuals, all of the same vegetative structure. The males pro-
duce sperms from clusters of densely packed plurilocular antheridia. The females
produce eggs solitary in large oogonia solitary or clustered on the thalli. The eggs
are without flagella. The diploid individuals produce unilocular sporangia of much
the same structure as the oogonia. In Zonaria, each sporangium produces eight non-
motile spores; in Dictyota, each one produces four.
Order 4. Sporochnoidea [Sporochnoideae] Greville Alg. Brit. 36 (1830).
Order Chordarieae Greville op. cit. 44.
Order Chordariaceae Haeckel Gen. Morph. 2: xxxv (1866).
Orders Desmarestiales and Chordariales Setchell and Gardner in Univ. Califor-
nia Publ. Bot. 8: 554, 570 (1925).
Order Sporochnales Sauvageau in Compt. Rend. 182: 364 (1926).
Brown algae producing motile spores, the haploid stage reduced to scant undiffer-
entiated filaments, the diploid stage filamentous or thallose, when thallose with apical
growth. Ralfsia is an exception to the formal characters of the order: it has a haploid
stage of the same structure as the diploid. This is a rather miscellaneous assemblage,
rather arbitrarily separated from Phaeozoosporea on the one hand and from Lamin-
ariea on the other.
The haploid body of the form of a short-lived body of a few undifferentiated fila-
ments, like a reduced Ectocarpus, bearing gametangia reduced to single cells, has
88 ] The Classification of Lower Organisms
been demonstrated by Kylin ( 1933, 1934, 1937) in a wide variety ot genera, as Asco-
cyclus, Desmotrichum, Mesogloia, Eudesme, Leathesia, and Stilophora. In the more
primitive examples, the gametes are not visibly differentiated; in more advanced
ones, as Carpomitra and Desmarestia, different haploid bodies produce respectively
smaller sperms and larger eggs, the latter non-motile.
There is a series of families, Ralfsiacea, Myrionematacea, Myriogloiacea, Meso-
gloiacea, and others, in which the diploid body consists of filaments differentiated
into different types. In the simplest of these, the germinating zygote produces in the
first place a minute thallus-like plate, generally epiphytic on other algae, one cell
thick, and consisting obviously of branched filaments of limited growth. From this
plate grow erect filaments. Some of these are simply cylindrical and appear nutritive
in function; others are attenuate, and may function in protection or in absorbing
materials from the water; yet others bear the reproductive structures, unilocular or
plurilocular or both.
In the more advanced families, the diploid body, after passing through a Ralfsia-
or Myrionema-Vike stage, may produce a compacted column of filaments with a
terminal plate of apical cells. Besides adding cells to the column, the apical plate
gives rise to a fascicle of attenuate hairs projecting forward. Members of the families
Chordariacea, Sporochnea, and Desmarestiacea produce cylindrical or flattened
thallose bodies of tridimensionally placed cells differentiated into an outer layer of
small actively photosynthetic cells and an inner mass of nearly colorless cells. Super-
ficial hairs, growing in intercalary fashion, may become few, and growth may become
restricted to a single apical cell.
By differences in the detailed manner of growth, Setchell and Gardner distin-
guished two orders among the thalloid forms just mentioned. It is evident, however,
that the thallose structure (and, likewise, differentiation of gametes) has developed
repeatedly and independently in the present group. Knowledge which would make it
possible to divide it into several recognizably natural orders is not yet available.
Order 5. Cutlerialea [Cutlcriales] Bessey in Univ. Nebraska Studies 7: 289 ( 1907).
Brown algae producing motile spores, the haploid and diploid bodies being macro-
scopically visible thalli, alike or different.
This is a small group, of one family, Cutleriacea, with two genera, Zanardinia
and Cutlcria, known chiefly from the Mediterranean. In Zanardinia, both haploid
and diploid bodies are erect and rather freely branched. In Cutlcria, the haploid
bodies are of this description, while the diploid bodies are appressed and fan-shaped.
The distinct diploid bodies of Cutlcria were originally named as a different genus,
Aglaozonia. Falkenberg (1879) first showed that Cutlcria and Aglaozonia arc stages
of the same thing; Yamanouchi showed that they are respectively a haploid stage
with 24 chromosomes and a diploid stage with 48.
The growing margins of the thalli consist of laterally compacted filaments grow-
ing by the divisions of a band of mcristematic cells which produce free hairs in the
distal direction and a continuous body of cells in the proximal direction. The latter
cells are capable of further division, and produce a body several cells thick, with
small cells rich in plastids on the surface and larger ones with fewer plastids in the
interior.
Haploid individuals bear clusters of stalked plurilocular structures of two types,
almost always on different individuals, the larger ones consisting of fewer cells which
release eggs, the smaller of more numerous cells which release sperms. Both kinds of
Phylum Phaeophyta [ 89
gametes are flagellum-bearing cells of the type usual in brown algae. The eggs are
capable of germination without fertilization, reproducing the haploid stage. Diploid
individuals bear clusters of unilocular sporangia.
It is only in the life cycle that the Cutlerialea are decidedly different from higher
Sporochnoidea such as Desmarestia. Their evolutionary origin is explicable by the
hypothesis of a single mutation which enabled the haploid stage to exhibit the com-
paratively complicated morphology of the diploid stage, instead of being rudimentary
as in all Sporochnoidea except Ralfsia (and the exceptional life cycle of Ralfsia
would be explained by a similar mutation in some primitive example of Sporochnoi-
dea, such as Myrionema) .
Older 6. Laminariea [Laminarieae] Greville Alg. Brit. 24 (1830).
Order Pycnospermeae Kiitzing Phyc. Gen. 333 (1843).
Order Laminariaceae Haeckel Gen. Morph. 2: xxxv (1866).
Laminariales Oltmanns Morph. u. Biol. Alg. ed. 2, 2: 2 (1922).
Order Laminariales Engler and Gilg Syllab. ed. 9 u. 10: 27 ( 1924).
Order Dictyosiphonales Setchell and Gardner in Univ. California Publ. Bot.
8: 586 (1925).
Order Punctariales Kylin in Kungl. Fysiog, Sallsk. Hand!, n. f. 44, no. 7 : 93
(1933).
Brown algae with motile spores, the haploid stages reduced to microscopic dimen-
sions, the diploid stages thallose, growing in intercalary fashion.
This numerous group, like the preceding small one, is evidently a specialized off-
shoot from order Sporochnoidea. The familiar examples are the kelps, whose large
diploid bodies are differentiated into definite members. Kylin considered his order
Punctariales to represent the transition to the kelps. They are thallose, without dif-
ferentiation of members, but their microscopic and reproductive characters, as ob-
served in Soranthera by Angst (1926, 1927), tend to confirm Kylin's opinion, and
they are accordingly included in the same order with the kelps. Papenfuss (1947)
pointed it out that the Punctariales of Kylin are essentially the same group as the
Dictyosiphonales of Setchell and Gardner.
Sauvageau (1915) first showed that the reproduction of kelps is sexual. The
grossly visible individuals produce zoospores; these, on germination, produce micro-
scopic filamentous haploid individuals, generally of distinct sexes, releasing gametes
from unicellular gametangia. The eggs are without flagella, and it is characteristic
of them that in emerging from the oogonia they become attached at the opening
(Kylin, 1916, 1933; Myers, 1928; McKay, 1933; Kanda, 1936; Hollenberg, 1939).
The same things are true in Soranthera, except that the eggs, although much larger
than the sperms, are also flagellate.
The visible bodies of kelps consist of three kinds of members, holdfasts (hapteres),
being stout root-like growths by which the individuals are anchored to rocks, and
stalks and blades comparable to stems and leaves. Growth is most active at the sum-
mits of the stalks. The histology is the same in all members (A. I. Smith, 1939).
There is a superficial photosynthetic tissue of small cells rich in plastids; on the hold-
fasts and stalks, this tissue is meristematic, adding cells to the tissue within and in-
creasing the thickness. Internally there is a cortex of larger cells with fewer plastids.
In the center there is a medulla containing trumpet fibers, filaments whose cells are
expanded where they meet and marked by pit-pairs. In the trumpet fibers of Nereo-
cystis there are actual perforations from cell to cell. The trumpet fibers are not quite
90]
The Classification of Lower Organisms
Fig. 16. — Familiar kelps of Pacific North America: a, Egregia Menziesii; h, Nereo-
cystis Luetkeana; c, Macrocystis pyrifera; d, Postelsia palmaeformis. All approxi-
mately X /a-
Phylum Phaeophyta [91
perfectly analogous to the sieve tubes of higher plants; the nuclei remain alive. The
minute zoospores are produced in unilocular sporangia. These occur on the surface
of the body in dense masses, intermingled with, and protected while young by, spe-
cialized sterile hairs.
Individuals of Laminaria consist simply of hapteres, a stalk, and one or more
terminal blades. In various other genera, growth occurs in such fashion as to cause
the blades to split at the base. With further growth, the splits extend to the margins
of the blades and increase their number, while intercalary growth at the transitions
between the stalks and the blades produces elongation and branching of the stalks.
Early explorers described the stalks of Macrocystis pyrifera as reaching prodigious
lengths, matters of hundreds of meters, and these accounts have been repeated in
textbooks down to recent times. Frye, Rigg, and Crandall (1915) found a maximum
length of somewhat less than fifty meters. The stalks are dichotomously branched
to a moderate extent and bear series of blades, each with a pear-shaped pneumato-
cyst or float at the base. The stalks of Nereocystis Luetkeana also were said to be
extremely long, but the recent observers did not find them to attain fifty meters. They
are unbranched and bear a single large float from which spring several blades which
may exceed four meters in length. This great organism is an annual, growing and
dying within a year. Postelsia palmaeformis, called the sea palm, grows on rocks ex-
posed to surf. It has erect stalks some 30 cm. tall bearing many pendant linear blades.
Egregia Menziesii has flattened stalks many meters long with fringes of floats and
blades along the margins. Laminaria is widely distributed. Macrocystis occurs on the
northwest coast of North America and in southern oceans. The other kelps which
have bef:n mentioned are confined to the northwest coast of North America.
On coasts where they occur, kelps are used as fertilizer. They have been used com-
mercially as sources of potash, as much as 1-3% of the fresh weight being K as K2O
(Cameron, 1915); they have been used also as sources of iodine. These uses are not
economic at most times.
Setchell and Gardner divided the proper kelps, of which there are about one
hundred species, into four families. The groups of less elaborate structure which ap-
pear properly to be placed in the same order are treated by Papenfuss (under Dictyo-
siphonales) as six families.
Order 7. Fucoidea [Fucoideae] C. Agardh Syst. Alg. xxxv (1824).
FucoiDEAE C. Agardh Synops. Alg. Scand. ix (1817).
Tribe Angiospermeae Kiitzing Phyc. Gen. 349 (1843).
Order Cyclosporeae Areschoug in Act. Roy. Soc. Upsala 13: 248 (1847).
Order Fucaceae J. Agardh Sp. Alg. 1 : 180 ( 1848).
Order Sargassaceae Haeckel Gen. Morph. 2: xxxv (1866).
Order Fucales Bessey in Univ. Nebraska Studies 7: 290 (1907).
Order Cyclosporales and suborder Fucineae Taylor in Bot. Gaz. 74: 439 (1922).
Class Cyclosporeae Kylin in Kungl. Fysiog. Sallsk. Handl. n. f. 44, no. 7: 91
(1933).
Thallose brown algae, producing no spores, diploid in all stages except the gametes;
the latter being sperms, whose posterior flagellum is longer than the anterior one,
and non-motile eggs. The genus Fucus L. is to be construed as the type genus of order
Fucoidea, class Melanophycea, and phylum Phaeophyta.
Two families are usually recognized (others have been segregated). In family
Fucea [Fuceae] Kiitzing (family Fucaceae Cohn), called the rockweeds, the bodies
92]
The Classification of Lower Organisms
are flat dichotomously branching thalli. In family Sargassea [Sargasseae] Kiitzing
there is a differentiation of holdfasts, stalks, blades, and floats. Growth is by division
of a single apical cell in each branch or member. There are the usual two tissues, a
superficial photosynthetic tissue of small cells and an inner tissue of larger cells which
pull apart to produce a spongy or fibrous mass.
The gametangia are borne, mixed with sterile hairs, in pits called conceptacles.
These are clustered, in the Fucea near the tips of branches which have ceased to
grow (these tips are swollen, and are called receptacles), in the Sargassea on special
branches. Rarely, oogonia and antheridia occur in the same conceptacles; not infre-
quently, they occur in different conceptacles on the same individuals; commonly, they
occur on different individuals. Male and female conceptacles may be distinguished
by color, the male being orange-yellow, the female of the same dark color as the thalli.
Male conceptacles are full of branching hairs bearing minute antheridia. In each
antheridium, the original single nucleus undergoes six successive simultaneous divi-
sions, producing sixty-four nuclei. These become the nuclei of sperms. Female con-
ceptacles contain fewer, larger, oogonia, in which the nuclei divide three times, pro-
FiG. 17. — Microscopic reproductive structures of Laminaria yezoensis after Kanda
( 1938) : a, male haploid individual releasing sperms; b, sperm; C, zoospore; d, female
haploid individual of three cells; e, female individual with an egg extnided from the
oogonium and attached in the mouth f, female individual with two young diploid
individuals attached at the mouths of oogonia. All x 1,000.
Phylum Phaeophyta [ 93
ducing eight. In Fucus, these become the nuclei of as many eggs. In other genera,
the number of functional eggs is reduced by degeneration of some of them, or of some
of the nuclei before cell division. In Sargassum, Kunieda (1928) found each oogon-
ium to produce a single egg in which seven nuclei undergo dissolution while one re-
mains to function.
The first two nuclear divisions in each gametangium are meiotic. Farmer and
Williams (1896) and Strasburger (1897) showed that the bodies are diploid; Yama-
nouchi (1909) first gave a full account of the meiotic process. The haploid chromo-
some number of Fucus vesiculosus is 32. In Sargassum Horneri Kunieda found it to
be 16.
By a swelling of colloidal material in the conceptacles, the gametangia are forced
out into the water, where they burst and release the gametes. Fucus was one of the
first organisms in which syngamy was observed. Thuret (1855) saw multitudes of
sperms swarm about the eggs, and showed that without sperms the eggs would not
develop. This much had already been observed in frogs and certain fishes; the dis-
covery that the essential process is the union of just one sperm with the egg was not
made until later. The growing zygotes give rise directly to diploid thalli.
■ The gametangia of the Fucoidea appear to be homologous with the unilocular
sporangia of other brown algae. In the gametangia, as in unilocular sporangia, the
meiotic divisions are followed by a few divisions of the haploid nuclei: the Fucoidea
are not quite perfect examples of the reduction of the haplod stage to the gametes
only. As to which other brown algae may have provided their evolutionary origin,
there is no very satisfactory hypothesis; Sporochnus shows certain resemblances.
Chapter VII
PHYLUM PYRRHOPHYTA
Phylum 3. PYRRHOPHYTA Pascher
Order Astoma Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1: 10 (1848).
Order Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852).
Flagellata Cohn in Zeit. wiss. Zool. 4: 275 (1853).
Orders Flagellata and Cilio-flagellata Claparede and Lachmann Etudes Infus.
1: 73 (1858).
Suborder Mastigophora Diesing in Sitzber. Akad. Wiss. Wein Math. -Nat. CI.
52, Abt. 1: 294 (1866).
Stdmme Flagellata and Noctilucae Haeckel Gen. Morph. 2: xxv, xxvi (1866).
Class Flagellata Kent Man. Inf. 1: 27, 211 (1880).
Class Mastigophora and orders Flagellata, Dinoflagellata, and Cystoflagellata
Butschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 2, Inhalt (1887).
Class Peridineae Wettstein Handb. syst. Bot. 1: 71 (1901).
Divisions Flagellatae and Dinoflagellatae Engler Syllab. ed. 3: 6, 8 (1903).
Pyrrhophyta, Eugleninae, and Chloromonadinae Pascher in Ber. deutschen Bot.
Gess. 32: 158 (1914).
Stdmme Pyrrhophyta and Euglenophyta, and Abteilungen Cryptophyceae, Des-
mokontae, and Dinophyceae, Pascher in Beih. bot. Centralbl. 48, Abt. 2: 325,
326 (1931).
Division Pyrrhophyta G. M. Smith Freshw. Algae 10 (1933).
Protistes trichocystiferes ou progastreades Chadefaud in Ann. Protistol. 5: 323
(1936).
Phyla Pyrrhophycophyta and Euglenophycophyta Papenfuss in Bull. Torrey Bot.
Club 73: 218 (1946).
Unicellular or colonial organisms, typically with brown or green plastids, flagel-
late, the flagella solitary or more than one and unequal, the cells marked by grooves
or pits and sometimes containing trichocysts, i. e., minute structures which lie close
to the cell membrane and eject thread-like bodies when stimulated.
The organisms included here are the ones conventionally treated as four orders of
pigmented flagellates, cryptomonads, dinoflagellates, euglenids, and chloromonads.
These groups include organisms of the same varied body types, algal, amoeboid, and
chytrid, that occur in other groups in which the flagellate body type is construed
as typical. Peridinium may be considered to be the type of the phylum.
Deflandre (1934) designated as stichoneme [stichonemate) the type of flagcllum
which bears a single file of appendages, and which had been discovered by Fischer
(1894) in Euglena. Petersen (1929) reobserved the stichoneme flagellum of Etiglena,
and found it also in other euglenids, Phacus and Trachelomonas. Deflandre found
that one flagellum is stichoneme in various further euglenids (but not in all), and
also in the dinoflagcllate Glenodinium. This is the only report of a stichoneme flagel-
lum outside of the euglenid group. The fine structure of the flagella of cryptomonads
and chloromonads has not been determined.
In some cryptomonads, as Chilomonas, the cells contain granules which stain
blue with iodine; if these are not starch, one knows not what to call them. Dino-
flagellates produce a so-called starch which gives a reddish color with iodine, and
many of them have walls of a so-called cellulose which gives a reddish color with
Phylum Pyrrhophyta [ 95
zinc chlor-iodide. The euglenids store granules of a white solid believed not to be
starch and called paramylum.
The plastids of cryptomonads and dinoflagellates are of various colors, oflF-color
green, yellow, brown, bluish, or red. Those of dinoflagellates contain chlorophylls
a and e; the latter is an exceptional chlorophyll which occurs also in Tribonema.
Euglenids and chloromonads are typically of the same bright green color as typical
plants, and the euglenids are known to have the same chlorophylls, a and b, as
typical plants (Strain, in Franck and Loomis, 1949).
The groups here brought together exhibit family resemblances in details of the
mitotic process, so far as these are known. The nuclear membrane usually persists
through the process. In many examples the chromosomes appear to be present at all
times, and are quite numerous, elongate, and of the appearance of strings of beads.
In mitosis, quite as one would assume, they divide lengthwise; the point had been
disputed, and was established by Hall (1923, 1925, 1937) and Hall and Powell
(1928). There is a neuromotor apparatus consisting of a centrosome at or near the
nuclear membrane together with one or more rhizoplasts connecting it to as many
blepharoplasts at the bases of the flagella. No spindle has been seen, unless the
peculiar structure, seen in Noctiluca outside of and next to the dividing nucleus, is
such. The centrosomes may lie at the sides of the dividing nucleus instead of at its
ends. In the euglenids and some dinoflagellates the nucleus contains a nucleolus-like
body which does not disappear during mitosis, but divides as the chromosomes do.
There are few reports of sexual processes in this group.
Pascher (1914) united the crytomonads and dinoflagellates in a group which
he named Pyrrhophyta. He and those who follow him leave the euglenids as an iso-
lated group. Tilden (1933) placed the four groups of flagellates with which we are
here concerned in division Chrysophyceae, while leaving the Phaeophyceae as a
distinct division. Her arrangement does not appear to be contrary to nature: the
cryptomonads are apparently not very far removed from the chrysomonads. The
different arrangement here maintained, by which the brown algae instead of the
cryptomonads and so forth are placed in the same phylum with the chrysomonads, is
believed to have the advantage that that phylum as least is well marked by char-
acter.
Chadefaud (1936) proposed a group consisting of the four groups of flagellates
here under consideration together with the Infusoria: this on the ground that the
Infusoria also have deeply indented cells containing trichocysts. He did not give
to his proposed group a place in the taxonomic system by assigning it to a category
and giving it a Latin name: he called it by the French common names protistes
trichocystiferes and progastreades. He suggested two ideas: that if a cell marked
by a considerable indentation should become divided into many cells forming two
layers, respectively superficial and against the indentation, the resulting structure
would be a gastrula; and that the gastrula, and, in fact, the kingdom of animals,
might have come into existence in this fashion. Perhaps because of novelty, these
ideas seem far-fetched. So far as it concerns flagellates, Chadefaud's grouping appears
sound and has been followed in giving limits to the present phylum.
The phylum is treated as a single class.
Class MASTSGOPHORA (Diesing) Bu'tschli
Classes Cryptomonadineae , Rhizocryptineae, Cryptocapsineae, Cryptococcineae,
Desmomonadineae, Desmocapsineae, Dinoflagellatae, Rhizodininae, Dinocap-
96 ] The Classification of Lower Organisms
sineae, Dinococcineae, Dinotrichineae, Euglenineae, and Euglenocapsineae
Pascher in Beih. bot. Centralbl. 48, Abt. 2: 325, 326 (1931).
Classes Chloromonadina, Euglenoidina, and Cryptomonadina Hollande in Grasse
Traite Zool. 1, fasc. 1: 227, 238, 285 (1952).
Further synonymy as of the name of the phylum.
Characters of the phylum.
There are about one thousand known species. Clearly, thirteen classes for their
accommodation, as proposed by Pascher, are excessive; perhaps one goes too far
in the other direction in making the entire group a single class. The type of the
class is the euglenid Astasia. This is true because the family Astasiaea was listed
first in the earliest appearance of the traditional group Flagellata or Mastigophora
in due taxonomic form, as order Astoma Siebold, If the euglenids are set apart,
taking with them the class name Mastigophora, the remaining larger class will be
called Peridinea [Peridineae] Wettstein.
The traditional four orders are tenably natural; but that of dinoflagellates includes
about four-fifths of the species, while the chloromonad group is very inconsiderable.
The system will be more convenient if the former order is divided into three, and if
the latter is included in the euglenid order. The resulting five orders are distinguished
as follows:
1. Pigmentation if present brown, olive, or the
like; flagella normally two.
2. Flagella at the anterior end of the cell,
not moving in longitudinal and trans-
verse grooves.
3. Not walled in the flagellate con-
dition, flagella not markedly dif-
ferentiated, or not differentiated
as anterior and circumferential .Order 1 . Cryptomonadalea.
3. Usually walled in the flagellate
condition; flagella respectively an-
terior and circumferential Order 2. Adiniferidea.
2. Flagella attached laterally, respectively
longitudinal and circumferential, moving
in grooves impressed upon the cells.
3. Not walled in the flagellate con-
dition Order 3. Cystoflagellata.
3. Flagellate cells with a wall usually
of articulated plates Order 4. Cilioflagellata.
1. Pigmentation if present typically bright
green, flagella normally solitary, sometimes
two or more Order 5. Astoma.
Order 1. Cryptomonadalea [Cryptomonadales] Engler Syllab. ed. 3: 7 (1903).
Subclass Cryptomonadineae Engler in Engler and Prantl Nat. Pflanzenfam.
ITeil, Abt. la: iv (1900).
Cryptophyceae, including Phaeocapsales and Cryptococcales, Pascher in Ber.
deutschen bot. Gess. 32: 158 (1914).
Order Cryptomonadinae Pascher Siisswassei-fl. Deutschland 1: 28 (1914).
Order Cryptomonadina Doflein Lehrb. Prot. ed. 4: 417 (1916).
Phylum Pyrrhophyta
[97
Order Cryptomonadida Calkins Biol. Prot. 265 (1926).
Orders Cryptocapsales and Cryptococcales Pascher in Beih. bot. Centralbl 48,
Abt. 2: 325 (1931).
Solitary (exceptionally colonial) cells, usually with one or two plastids of various
colors, usually observed in the motile condition, then naked, of dorsiventral (excep-
tionally isobilateral) symmetry, with two anterior flagella which are not markedly
differentiated or not respectively anterior and circumferential.
The resting nucleus contains a karyosome, i. e., a globule which occupies most of
t 0
t>.>.
Fig. 18. — a, Cryptomonas sp. b, Rhodomonas baltica after Kylin ( 1935 ) . c, Chi-
lomonas Parmecium. d, Cyathomonas sp. e, Sennia sp. f. Vegetative cell, and
g, zoospore of Paradinium Pouchetii after Chatton (1920). All x 1,000.
its volume and contains most of the chromatin. Dangeard (1910) and Belar (1916)
have observed details of mitosis. The numerous chromosomes appear within an
intact nuclear membrane and form a disk- or drum-shaped figure with its axis at
right angles to the axis of the cell. No granule more massive than the chromosomes
persists and divides with them.
About thirty species are known. They may be treated as five families.
1. Flagellate cells elongate, with one plane
of symmetry.
2. Not parasitic, flagella not markedly dif-
ferentiated.
3. Non-motile in the vegetative con-
dition Family 1. Cryptococcacea.
3. Flagellate in the vegetative con-
dition Family 2. Cryptomonadina.
98 ] The Classification of Lower Organisms
3. Amoeboid in the vegetative con-
dition Family 3. Paramoebida.
2. Parasitic amoeboid organisms, the flag-
ella of swimming stages respectively
anterior and trailing Family 4. Paradinida.
1. Flagellate cells with two planes of symmetry Family 5. Nephroselmidacea.
Family 1. Cryptococcacea [Cryptococcaceae] Pascher in Beih. Bot. Centralbl. 48,
Abt. 2: 325 (1931). YdimWy Phaeocapsaceae West British Freshw. Alg. 48 (1904),
in part; Phaeocapsa is a chrysomonad. Family Phaeoplakaceae Pascher 1. c. Solitary
or clustered cells, non-motile in the vegetative condition, reproducing by flagellate
cells of cryptomonad type. Phaeococcus, Cryptococcus, Phaeoplax. Chrysidella in-
cludes yellowish cells called zooxanthellae, internally symbiotic in Radiolaria, Rhizo-
poda, sponges, coelenterates, and rotifers. It is believed that the supposed zoospores
of various amoeboid organisms are actually flagellate reproductive cells of Chrysi-
della escaping at certain stages of the life cycles of their hosts.
Family 2. Cryptomonadina Ehrenberg Infusionsthierchen 38 (1838). Family
Chilomonadidae Kent Man. Inf. 1: 423 (1880). Family Cryptomonadaceae Engler
Syllab. ed. 3: 7 (1903). Family Chilomonadaceae Lemmermann 1909. Family
Cryptomonadidae Poche in Arch. Prot. 30: 159 (1913). Flagellate in the vegetative
condition, the two flagella not markedly differentiated, springing from the anterior
end of the cells, usually from the mouth of a pit lined by granules of some sort.
Cryptomonas and Cryptochrysis have brown or yellow plastids; Chromomonas and
Cyanomonas have blue ones; Rhodomonas has red ones. Chilomonas is a colorless
saprophyte familiar in infusions. The colorless Cyathomonas, also from infusions,
was shown by tJlehla (1911) to be related to Chilomonas.
Family 3. Paramoebida [Paramoebidae] Poche in Arch. Prot. 30: 173 (1913).
Schaudinn (1896) discovered the sole known species, Paramoeba Eilhardi, in an
aquarium of sea water. It is an amoeboid organism with the peculiarity that each
cell contains beside the nucleus an additional body which divides when the nucleus
does. The cell may form about itself a shell of debris, and within this may undergo
division into many cells which escape as pigmented swarmers resembling cells of
Cryptomonas.
Family 4. Paradinida [Paradinidae] Chatton in Arch Zool. Exp. Gen. 59: 444
(1920). The sole known species, Paradinium Poucheti, is a parasite in the body
cavity of copepods. The amoeboid cells are linked together by slender pseudopodia
so as to form a network. The reproductive cells have a shorter anterior flagellum
and a longer trailing flagellum.
Family 5. Nephroselmidacea [Nephroselmidaceac] Pascher Siisswasserfl. Deutsch-
land 2: 'llO (1913). Family Nephroselmidae Calkins Biol. Prot. 267 (1926). Cells
isobilateral. Cells disk-shaped, the flagella on the margin: Scnnia. Cells laterally
extended, bean- or kidney-shaped, the indentation anterior and bearing the flagella:
Protochrysis, Nephroselmis.
Order 2. Adiniferidea Kofoid and Swczy in Mem. Univ. California 5: 108
(1921).
Suborder Adinida Blitschli in Bronn Kl. u. Ord. Thicrreichs 1: 1001 (1885).
Suborder Prorocentrinea Poche in Arch. Prot. 30: 160 (1913).
Desmokontae, including Desmomonadales and Desmocapsales, Pascher in Ber.
deutschen bot. Gess. 32: 158 (1914).
Phylum Pyrrhophyta [ 99
Division Desmokontae; classes Desmomonadineae and Desmocapsineae; and
orders Desmomonadales, Prorocentrales, and Desmocapsales Pascher in Beih.
bot. Centralbl. 48, Abt. 2: 325 (1931).
Suborder Prorocentrina Hall Protozoology 142 (1953).
Solitary cells, mostly flagellate in the vegetative condition, the flagellate stages
either naked or bearing a close wall of two valves, with two flagella at the anterior
end, one extending forward while the other is bent circumferentially and causes the
cell to whirl while swimming.
The few known organisms of this group may be treated as a single family.
Family Adinida Bergh in Morph. Jahrb. 7: 273 (1882). Family Prorocentrinen
Stein Org. Inf. 3, II Halfte: 8 (1883). Family Prorocentrina Butschli in Bronn Kl.
u. Ord. Thierreichs 1: 1002 (1885). Family Prorocentraceae Schiitt in Engler and
Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 6 (1896). Prorocentridae Kofoid in Bull.
Mas. Comp. Zool. Harvard 50: 164 (1907). Family Prorocentridae Poche in Arch.
Prot. 30: 160 (1913). Desmocapsa, Haplodiniuni, Desmomastix, Pleuromonas,
Exuviaella, Prorocentrum; minute brown organisms, mostly marine.
Order 3. CystoflageUata (Haeckel) Butschli in Bronn Kl. u. Ord. Thierreichs
1, Abt. 2, Inhalt (1887).
Tribe [group of families] Gymnodinioidae Poche in Arch. Prot. 30: 161 (1913).
Classes Rhizodininae, Dinocapsineae, Dinococcineae, and Dinotrichineae; orders
Gymnodiniales, Rhizodiniales, Dinocapsales, Dinococcales, and Dinotrichales
Pascher in Beih. bot. Centralbl. 48, Abt. 2 : 326 ( 193 1 ) .
Suborders Gymnodinina, Dinocapsina, and Dinococcina Hall Protozoology 143,
147, 149 (1953).
Haeckel ( 1866) made of Noctiluca alone a phylum under the name of Noctilucae.
He had the carelessness, as it appears, to publish in the same work the synonymous
phylar name Myxocystoda as a label in a phylogenetic diagram. In 1873 he used a
third name, CystoflageUata, and Biitschli took this up; in the text of the Klassen
und Ordnungen ambiguously as an Unterabtheilung or Ordnung, in the table of
contents definitely as an order. Allman (1872) had shown that Noctiluca belongs to
the present group. Biitschli did not agree with this opinion, but it is evidently correct,
and Haeckel's name becomes the valid one for the order to which Noctiluca belongs
Typical members of the present order are naked motile cells with brown plastids.
The two flagella are attached near the equator of the cell. One of them extends in a
posterior direction, in a groove called the sulcus. The other extends horizontally about
the cell (generally to the right, in the reversed image seen in the microscope), lying
in a groove called the girdle. The part of the cell anterior to the girdle is called the
epicone, the part posterior to it, the hypocone. From the typical structure as thus
described, there are, as will be seen, many deviations.
The species, of which more than three hundred are known, may be treated as nine
families.
1. Relatively unspecialized; having stages freely
propelled by two flagella, with a single girdle,
no tentacles, and unspecialized eyespots or
none; not parasitic; commonly pigmented.
2. Walled and non-motile in the vegeta-
tive condition Family 1. Phytodiniacea.
2. Flagellate in the vegetative condition Family 2. Gymnodiniacea.
100 ] The Classification of Lower Organisms
1. Not as above, always without photosynthetic
pigments.
2. Amoeboid Family 3. Dinamoebidina.
2. Flagellate or free-floating.
3. With multiplied girdles, without
tentacles or specialized light-sensi-
tive organelles Family 4. PoLYKRiKroA.
3. With one girdle or none.
4. Cells more or less isodiamet-
ric.
5. With prominent light-sen-
sitive organelles, some-
times with tentacles Family 5. Pouchetiida.
5. Without light-sensitive or-
ganelles, with tentacles.
6. Not exceptionally
large Family 6. Protodiniferida.
6. Reaching exceptional
sizes, to 1 mm. in di-
ameter Family 7. Noctilucida.
4. Cells dome-shaped Family 8. Lepodiscida.
2. Parasitic Family 9. Blastodinida.
Family 1 . Phytodiniacea [Phytodiniaceae] Schilling in Pascher Siisswasserfl.
Deutschland 3: 61 (1913). Family Phytodinidae Calkins Biol. Prot. 277 (1926).
Dinocapsales, Dinocapsaceae, Dinococcales, Dinotrichales, and Dinotrichaceae
Pascher in Ber. deutschen bot. Gess. 32: 158 (1914). Orders Dinocapsales, Dino-
coccales, and Dinotrichales, and families Gloeodiniaceae, Hypnodiniaceae, Dino-
trichaceae, and Dinocloniaceae Pascher in Beih. bot. Centralbl. 48, Abt. 2: 326
(1931). Organisms with numerous yellow to brown plastids, walled and non-motile
in the vegetative condition, reproducing by gymnodinioid zoospores. Some fifty
species are known; it is only recently that Thompson (1949) has found several of
these in America. Cells multiplying in a gelatinous matrix: Gloeodinium. Cells
solitary, dividing into several which escape usually in the flagellate condition; with
smooth ellipsoid walls: Phytodinium, Stylodinium; anvil-shaped, stalked and with
two horns: Racihorskya; tetrahedral, with horns at each comer: Tetradinium; with
a ring of about six horns: Dinastridium. Tending to produce filaments; marine:
Dinothrix, Dinoclonium.
Family 2. Gymnodiniacea [Gymnodiniaceae] Schiitt in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. lb: 2 (1896). Subfamily Gymnodinida Bergh in Morph.
Jahrb. 7: 274 (1882). Gymnodinidae Kofoid in Bull. Mus. Comp. Zool. Harvard
50: 164 (1907). Family Gymnodiniidae Poche in Arch. Prot. 30: 162 (1913). The
typical unarmored dinoflagellates, free-swimming, with sulcus and girdle, without
tentacles or a conspicuous light-sensitive organelle, commonly with photosynthetic
pigments.
The genus which is most numerous in species is Gymnodinium Stein. It includes
both pigmented and non-pigmcnted species, mostly marine, occasional in fresh water,
the girdles nearly equatorial and forming nearly complete circles. The cells readily
become encysted, and the cysts may grow to large sizes, reaching diameters of 0.5 mm.
These cysts have been taken for a distinct genus Pyrocystis. Observed in darkness,
Phylum Pyrrhophyta [ 101
the protoplasm in the cysts is seen to become luminous in response to disturbance of
the medium; they are among the agents of phosphorescence at sea. In Gymnodinium
Lunula the protoplast of each large globular cyst undergoes division into several
protoplasts which do not immediately become flagellate; each of them becomes
crescent-shaped, deposits a cell wall, and is released by dissolution of the wall of the
parent cyst. In the crescent-shaped cysts, the protoplasts divide into several which
develop flagella and escape as typical gymnodinioid cells.
In Hemidinium the girdle forms less than a complete circle; in Amphidinium, the
girdle is close to the anterior end of the cell; in Gyrodinium, it forms a steep left
spiral; in Cochliodinium it forms a left spiral of more than one and one half turns.
Family 3. Dinamoebidina nom. nov. Order Rhizodiniales and family Amoehodi-
niaceae Pascher (1931), not based on generic names. Non-pigmented amoeboid
organisms producing crescent-shaped cysts which germinate by releasing gymnodini-
oid zoospores. Dinamoebidium varians Pascher (1916; originally Dinamoeba, but
there is an earlier genus of this name, and the author changed it).
Family 4. Polykrikida [Polykrikidae] Kofoid and Swezy in Mem. Univ. California
5: 395 (1921). Family Polydinida Butschli (1885), not based on a generic name.
There is a single genus Polykrikos, of only three known species. They are colorless
predatory organisms of such a structure as might be produced if a cell of Gymnodi-
nium were repeatedly to enter upon division and fail to complete it. Each elongate
cell bears a single extended sulcus and a series of girdles; with each girdle are asso-
ciated the usual two differentiated flagella. Of nuclei there are usually half as many
as of girdles. The cells contain structures called nematocysts, whose development
and structure was studied by Chatton (1914). Each nematocyst consists of a conical
wall, with a peculiar operculum at the broad end, surrounding a minute cavity
containing fluid and a coiled thread. Nematocysts are supposed to be homologous
with trichocysts, and to contribute to protection, or to the capture of prey; the points
seem not fully established. They occur only in this family and the following.
Family 5. Pouchetiida [Pouchetiidae] Kofoid and Swezy in Mem. Univ. of Cali-
fcrnia 5: 414 (1921). Each of the gymnodinioid cells contains a light-sensitive ap-
paratus, the ocellus, consisting of a pigmented area and of one or more transparent
globes, of unknown composition, serving as lenses. Most species have nematocysts.
Protopsis, Pouchetia, etc.; Erythropsis, in warm seas, with a prominent tentacle.
Family 6. Protodiniferida [Protodiniferidae] Kofoid and Swezy in Mem. Univ.
California 5:111 (1921). Family Pronoctilucidae Lebour Dinofl. Northern Seas 10
(1925). Predatory organisms, the cells subglobular, without ocellus or nematocysts,
but with a tentacle. Pronoctiluca Fabre-Domergue 1889 {Protodinifer Kofoid and
Swezy 1921); 0.v}'rr/iw Dujardin.
Description of the neuromotor apparatus and process of division in Oxyrrhis
marina by Hall (1925) provides part of the authority for, and is in good conformity
to, the remarks on mitosis included above in the description of the phylum. The
nucleus contains a prominent internal body (endosome) which does not contain the
material of the chromosomes and does not disappear during mitosis. A centrosome,
close outside the nuclear membrane, is connected by two rhizoplasts to blepharo-
plasts at the bases of the flagella. When a cell is to divide, the centrosome divides;
the daughter centrosomes do not necessarily lie at the poles of the nucleus where
the chromosomes assemble. Each daughter centrosome appears to generate one
rhizoplast, blepharoplast, and flagellum to complete the neuromotor apparatus of a
eel]. In due course, the endosome, nucleus, and cell undergo constriction.
102 ] The Classification of Lower Organisms
Family 7. Noctilucida [Noctilucidae] Kent Man. Inf. 1: 396 (1880). The single
species Noctiluca scintillans (Mackartney) Kofoid and Swezy (1921; usually known
as A^. miliaris Suriray ) is a predatory marine organism, the subglobular cells reaching
dimensions exceeding 1 mm., luminescent when stimulated and accordingly contrib-
uting to phosphorescence at sea. Each cell is marked by an extensive depression
representing the sulcus; the girdle is obsolete. A part of the area of the sulcus func-
tions as a cytostome. A tooth in the sulcus represents the transverse flagellum. Present
are a longitudinal flagellum, minute in proportion to the cell, and a prominent
tentacle.
Mitosis in Noctiluca has been studied by Calkins (1899), van Goor (1918), and
Pratje (1921). Adjacent to the nucleus there is a body of differentiated cytoplasm,
as large as the nucleus, called by Calkins the attraction sphere. Before mitosis, the
tentacle and flagellum are absorbed. The attraction sphere becomes elongate and
its central part becomes converted into fibers. The nucleus becomes appressed to,
and curved about, the bundle of fibers, and the numerous elongate chromosomes
assemble against this. The two curved margins of the nucleus draw apart along the
bundle of fibers, appearing to draw the daughter chromosomes with them. Division is
completed by constriction of the nucleus and disappearance of the fibers, leaving a
daughter attraction sphere in association with each daughter nucleus. This peculiar
mitotic process is probably of no phylogenetic significance, being, like the organism
in which it occurs, an aberrant by-product of evolution.
Nuclear division may be followed by division of the cell into two, the entire
process requiring from twelve to twenty-four hours. Alternatively, the nucleus may
divide repeatedly, each division requiring from three to four hours; the numerous
nuclei produced are budded off from the cell in small uniflagellate spores. Ischikawa
( 1891 ) saw conjugation of pairs of cells, and van Goor stated that this is a preliminary
to the production of spores; Pratje, on the other hand, could find no evidence of
conjugation. The spores are believed to give rise by direct growth to cells like the
original one.
Family 8. Leptodiscida [Leptodiscidae] Kofoid 1905. Large dome-shaped preda-
tory marine organisms with small flagella or none. Leptodiscus R. Hertwig (1877)
was placed by Biitschli in order Cystoflagellata as the sole genus in addition to
Noctiluca; Craspedotella is a comparatively recent discovery of Kofoid.
Family 9. Blastodinida [Blastodinidae] Chatton in Arch. Zool. Exp. Gen. 59:
442 (1920). Ordre Blastodinides Chatton in Compt. Rend. 143: 981 (1906). Fam-
ilies Apodinidae, Haplozoonidae, Oodinidae, and Syndinidae Chatton op. cit (1920).
Dinoflagellates which are parasitic chiefly in copepods and tunicates, also in other
animals and in diatoms. As a general rule, after the parasite has grown to a certain
size, and a multiplication of nuclei has taken place, a part of the protoplast undergoes
division to form gymnodinioid zoospores, while the remainder resumes growth in
the host. Schizodinium, Blastodinium, Apodinium, Chytriodinium, etc.
Order 4. CiUoflagellata Claparede and Lachman Etudes Inf. 1: 394 (1858).
Family Peridinaea Ehrcnbcrg Infusionsthierchcn 249 (1838).
Family Dinifera Bergh in MoVph. Jahrb. 7: 273 (1882).
Order Dinoflagellata BiitschU in Bronn Kl. u. Ord. Thierreichs 1, Abt. 2: Inhalt
(1887).
Subclass Peridiniales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
lb: V (1896).
Phylum Pyrrhophyta [103
Class Peridineae Wettstein Handb. syst. Bot. 1: 71 (1901).
Division Dinoflagellata Engler Syllab. ed. 3: 8 (1903).
Dinophyceae and Dinoflagellatae Pascher in Ber. deutschen bot. Gess. 32: 158
(1914).
Order Diniferidea and tribe [group of families] Peridinioidae Kofoid and Swezy
in Mem. Univ. California 5 : 106, 107 ( 1921 ) .
Order Dinoflagellida Calkins Biol. Prot. 267 (1926).
Division Dinophyceae, Class Dinoflagellatae, and order Peridiniales Pascher in
Beih. bot. Centralbl. 48. Abt. 2: 326(1931).
Suborder Peridinina Hall Protozoology 144 (1953).
This order is very close to the preceding; its members are distinguished only by
the presence, while the cells are in the flagellate condition, of cell walls, consisting in
most examples of separable plates. The name Cilioflagellata is evidence of an early
error of observation: the circumferential flagellum was mistaken for a whorl of
cilia. This name and most of its synonyms were published as applying both to the
preceding order and this. For almost all of these names the type or obvious standard
example is Peridinium, with the effect that the names belong to the present order.
There are about five hundred species, prevalently marine. Five families may be
recognized.
Family 1. Peridinaea Ehrenberg Infusionsthierchen 249 (1838). Family Peridin-
idae Kent Man. Inf. 1: 441 (1880). Family Peridiniaceae Schiitt in Engler and
Prantl Nat. Pflanzenfam. I Teil, Abt. lb: 9 (1896). Ceratiidae Kofoid in Bull. Mus.
Comp. Zool. Harvard 50: 164 (1907). The typical dinoflagellates, of numerous
genera and species. The distinctions among them are largely matters of the detailed
arrangement of the plates making up the walls. Glenodinium, the plates scarcely
distinguishable. Peridinium, Goniodoma, Goniaulax, Ceratium, Oxytocum, etc. The
cells of certain species in various genera are ornamented with prominent horns; in
Ceratium especially the epitheca is drawn out into one long horn, and the hypotheca
into one, two, or three. Goniaulax becomes abundant at certain seasons, is eaten by
shellfish, and renders them poisonous.
The neuromotor apparatus (much as in Menoidium) and the process of nuclear
and cell division in Ceratium Hirundinella were described by Entz (1921) and Hall
(1925). Many nuclei lack the endosome; if present, it disappears during mitosis, as
does also the nuclear membrane. The daughter centrosomes lie at the sides of the
blunt-ended mitotic figure. When nuclear division is complete, the protoplast ex-
pands and then becomes constricted in such fashion that each daughter cell receives
certain plates of the wall; each daughter cell then secretes the plates which it lacks.
Zederbauer (1904) reported conjugation in Ceratium. He saw an elongate proto-
plast with each of its ends covered by a complete cell wall. Dividing cells are of
quite different appearance.
Families Ptychodiscida, Cladopyxida, and Amphilothida of Kofoid (1907, the
names in the feminine; explicitly made families by Poche, 1913) are minor segregates
from Peridinaea.
Family 5. Dinophysida (Bergh) Biitschli in Bronn Kl. u. Ord. Thierreichs 1:
1009 (1885). Subfamily Dinophysida Bergh in Morph. Jahrb. 7: 273 (1882). The
limits of the plates obscure; girdle near the anterior end; sulcus and girdle bordered
by prominent flanges. Strictly marine, mostly in warmer oceans. Dinophysis, Oxyphy-
sis, Amphisolenia, Triposolenia, etc.
104]
The Classification of Lower Organisms
Fig. 19, — a, Tetradinium javanicum x 1,000 after Thompson (1949). b, Gytnno-
dinium striatum x 500 after Kofoid & Swezy (1921). C, Gymnodiniian Lunula,
flagellate cells forming in a cyst x 500, after Kofoid & Swezy op. cit. d, e, f, Din-
amoehidium varians; amoeboid vegetative cell, cyst, and production of gymnodinioid
zoospores x 1,000 after Pascher (1916). g, Noctiluca scintillans x 100 after Allman
(1872). h, Peridinium cinctum x 1,000. i, Triposolcnia Ambulatrix x 500 after
Kofoid (1907). j, Amphisolcnia laticincta after Kofoid, op. cit.
Phylum Pyrrhophyta [ 105
Order 5. Astoma Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1 : 10 ( 1848) .
Order Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852), in part.
Order Flagellata Claparede and Lachmann Etudes Inf. 1: 73 (1858), in part.
Order Flagellato-Eustomata Kent Man. Inf. 1: 36 (1880).
Suborder Euglenoidina Biitschli in Bronn Kl. u. Ord. Thierreichs 1 : 818 ( 1884).
Abtheilung (suborder) Chloromonadina Klebs in Zeit. wiss. Zool. 55: 391
(1893).
Order Euglenoidina Blochmann Mikr. Tierwelt 1, ed. 2: 50 (1895).
Subclasses Chloromonadineae and Euglenineae Engler in Engler and Prantl
Nat. Pflanzenfam. I Teil, Abt. la: v, vi (1900).
Orders Euglenales and Chloromonadales Engler Syllab. ed. 3: 7 (1903).
Orders Eugleninae and Chloromonadinae Pascher Siisswasserfl. Deutschland 1 :
29 (1914).
Orders Euglenida and Chloromonadida Calkins Biol. Prot. 283, 285 (1926).
Mostly solitary flagellate cells of fresh water, unwalled and capable of contraction
and writhing movement; the anterior end of each cell (in the flagellate condition)
penetrated by a pit, the reservoir or cytopharynx, into which contractile vacuoles
open; having one flagellum, or two, usually unequal, or more, one flagellum of each
cell usually being stichoneme; mostly producing a solid storage product, not staining
blue with iodine, called paramylum.
Jahn (1946) reviewed this group. He recognized four families, to which one
more, to include the chloromonads, is to be added.
1. Producing paramylum.
2. Flagellum with a swelling near the base,
usually single but formed of two parts
which join below the swelling; cells
mostly pigmented.
3. Non-motile and walled in the vege-
tative condition Family 1 . Colaciacea.
3. Flagellate in the vegetative con-
dition Family 2. Euglenida.
2. Flagellum not swollen and usually not
forked near the base; cells not pig-
mented.
3. Cells without internal rod-shaped
structures; flagella stichoneme Family 3. Astasiaea.
3. Cells with internal rod-shaped struc-
tures; flagella acroneme or simple Family 4. ANisoNEMroA.
1. Not producing paramylum, storing oil Family 5. Coelomonadina.
Family 1. Colaciacea [Colaciaceae] Smith Freshw. Alg. 617 (1933). Family
Colaciidae Jahn in Quart. Rev. Biol. 21: 264 (1946). Euglenoid organisms which
are walled and non-motile in the vegetative condition. There is a single genus
Colacium, producing dendroid colonies.
Family 2. Euglenida Stein Org. Inf. 3, I Halfte: x (1878). Family Euglenina
Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 820 (1884). Family Euglenaceae
Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). Solitary
motile cells, mostly with abundant green plastids, the flagella with swellings near the
base, mostly solitary and forked below the swelling. Jahn recognized twelve genera.
Eutreptia has two flagella; Euglenamorpha has three. Members of the latter genus
106]
The Classification of Lower Organisms
Fig. 20. — a., Colacium Arbuscula after Stein (1878). b^ Euglena viridis. c, Eu-
glena Spirogyra. d, Euglena acus. e, Phacus sp. f, Trachelomonas sp. g, Kleb-
siella alligata after Pascher (1931). All x 1,000.
Phylum Pyrrhophyta [ 107
are entozoic in frog tadpoles; some of them are non-pigmented. Three genera having
the typical single flagella are among the most familiar of flagellates. Euglena has
fusiform to cylindrical cells freely capable of writhing changes in shape. Phacus has
flattened cells with a rigid membrane. In Trachelomonas, the protoplast lies loose
in a rigid lorica which is often ornamented with spines; variations in the form and
ornamentation of the lorica have made it possible to distinguish a large number of
species.
There are accounts of mitosis in Euglena by Keuten (1895), Baker (1926), Rat-
cliffe (1927) and Hall and Jahn (1929). All observers have seen within the nucleus
a large globule which divides as the nucleus does and appears to guide the separating
chromosomes. Keuten applied to it the term nucleolo-centrosome; the implications
of this term are not confidently to be accepted, and the body will better be called by
the neutral term endosome. RatclifTe's account of mitosis in Euglena Spirogyra is the
most detailed. It appears that division is initiated when the endosome buds oflE a
small granule which migrates to a position just within the nuclear membrane and
divides. The resulting granules may be regarded as centrosomes. The nucleus moves
forward within the cell and comes into contact with the cell membrane at the bottom
of the reservoir. Each centrosome appears to generate, just within the cell membrane,
a granule recognizable as a blepharoplast; the nucleus then withdraws from the cell
membrane, but the centrosomes remain connected to the blepharoplasts by rhizo-
plasts. The flagellum, already split at the base, divides throughout its length into two;
a new flagellum-base grows out from each blepharoplast and becomes fused to one
of the halves of the old one not far from the base of the latter. Meanwhile, withm
the intact nuclear membrane, the chromosomes and endosome are dividing. The
centrosomes are at the sides of the dividing nucleus. No spindle has been recognized.
Nuclear division is completed by constriction of the membrane. The cell divides by
constriction which proceeds longitudinally from the anterior end. The centrosomes
and rhizoplasts disappear, to be replaced during the next division by new ones.
Hall and Hall and Schoenborn (in several papers, 1938, 1939) have reported
experiments on nutrition in Euglena. All species are capable of photosynthesis. Some
of them, surprisingly, have lost the capacity to synthesize amino acids which usually
accompanies photosynthesis; and there are transitional species in which some in-
dividuals possess the capacity to make amino acids and others do not, evidently as
heritable characters.
Family 3. Astasiaea Ehrenberg Infusionsthierchen 100 (1838). Family Astasiidae
Kent Man. Inf. 1 : 375 ( 1880) . Family Astasiina Biitschli in Bronn. Kl. u. Ord. Thier-
reichs 1 : 826 ( 1884) . Family Astasiaceae Senn in Engler and Prantl Nat. Pflanzenfam.
I Teil, Abt. la: 177 (1900). Colorless organisms. Deflandre found the flagella sticho-
neme, as to the single flagella of Astasia and Menoidium, and as to one of the two
flagella of Distigma. Hall and Jahn (1929) found the flagella not swollen near the
base. The internal rod-shaped structures which characterize the following family are
absent.
Belar (1915) described mitosis in Astasia, and Hall (1923) described it in
Menoidium. There is a blepharoplast at the base of the flagellum, and some prepara-
tions show a rhizoplast connecting this to a centrosome immediately outside the
nuclear membrane. The blepharoplast divides during the early stages of mitosis, and
the flagellum appears to divide lengthwise. The daughter centrosomes mark the loci
toward which the dividing chromosomes move. The chromosome number appears to
be 12. A dividing endosome like that of Euglena is present.
108]
The Classification of Lower Organisms
Scytomonas pusilla Stein {Copromonas subtilis Dobell) occurs in the intestines
of frogs and toads. When cast out with the feces, it exhibits conjugation as a pre-
liminary to encystment (Dobell, 1908).
Family 4. Anisonemida [Anisonemidae] Kent Man. Inf. 1: 429 (1880). Families
Pernamina and Anisonemina Biitschli in Bronn Kl. u. Ord. Thierreichs 1 : 824, 828
(1884). Family Peranemaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: 178 (1900). Family Heteronemidae Calkins Biol. Prot. 285 (1926). Each
cell of these colorless organisms bears one conspicuous anterior flagellum; most of
them bear also a less conspicuous trailing flagellum. The trailing flagellum of Pera-
nema is grown fast to the cell membrane, and is detected only with difficulty (Hall,
Fig. 21.— a, Menoidium incurvum. b, c. Stages of mitosis in Menoidium incurvum
X 2,000 after Hall (1923). d, e, Peranema trichophorum. i, Stage of division in
Peranema trichophorum after Hall (1934). g, Anisoncma truncatum. h, Ento-
sipon sulcatum, i-m, Vacuolaria viridis: i, cell; j, neuromotor apparatus after Fott
(1935); k-m, stages of mitosis x 2,000 after Fott, op cit. x 1,000 except as noted.
Phylum Pyrrhophyta [ 109
1934). Deflandre was unable to find appendages on the flagella of members of this
family. As in other members of the order, the flagella spring from a deep anterior pit
in the cell; in this family, the pit is a functional cytopharynx (Hall, 1933). The cyto-
plasm of Peranema contains three brief rods, the pharyngeal rods or Staborgane,
lying near the cytopharynx; their function is unknown. Each cell of Urceolus, of
Anisonema, and of Heteronema contains a single conspicuous rod extending the length
of the body. Hall and Powell (1928) and Hall (1934) described the mitotic process
in Peranema, which is much as in Menoidium.
Family Coelomonadina Butschli in Bronn Kl. u. Ord. Thierreachs 1 : 819 (1884).
Family Vacuolariaceae Luther in Bihang Svensk. Vetensk-Akad. Handl. 24, part 3,
no. 13: 19 (1889). Family Chloromonadaceae Engler Syllab. ed 3: 7 (1903). Family
Thaumatonemidae Poche in Arch. Prot. 30: 155 (1913). Family Chloromonadidae
HoUande in Grasse Traite Zool. 1, fasc. 1: 235 (1952); family Thaumatomonadidae
Hollande op. cit. 686. Unicellular organisms, mostly green, with two diiTerentiated
flagella springing from a large reservoir, producing globules of oil but no solid storage
product. Klebs apologized for erecting the grossere Abtheilung Chloromonadina for
the single genus Vacuolaria, and in fact, this genus differs from other members of the
present order only in one conspicuous character, the failure to produce paramylum.
Fott (1935) studied the cytology of Vacuolaria. From the base of each flagellum, a
rhizoplast extends into the cytoplasm, but fails to come into contact with the nucleus.
Several granules or swellings, not definitely identifiable as blepharoplasts or centro-
somes, are distributed along the length of each rhizoplast. In mitosis, which takes
place within an intact nuclear membrane, the numerous subglobular chromosomes
form a blunt-ended figure much as in Chilomonas. Genera believed to be allied to
Vacuolaria include the green flagellate Goniostomum; Chysophaeum Lewis and
Bryan (1941), a marine organism forming non-motile yellow dendroid colonies of
m.acroscopic dimensions; and the colorless flagellate Thaumatomastix Lauterborn
(originally named Thaumatonema, but there is among plants an older genus of this
name).
Chapter VHI
PHYLUM OPISTHOKONTA
Phylum 4. OPISTHOKONTA, phylum novum
Chytridieae de Bary in Bot. Zeit. 16, Beil. 96 (1858).
Family Chytridieen de Bary and Woronin (1864).
Family Chytridiaceae Cohn in Hedwigia 11: 18 (1872).
Chytridineae Schroter in Engler and Frantl Nat. Pflanzenfam. I Teil, Abt. 1 :
62 (1892).
Series (Reihe) Archimycetes (Chytridinae) A. Fischer in Rabenhorst Kryp-
tog.-Fl. Deutschland 1, Abt. 4: 11 (1892).
Suborders Chrytidiineae and Monoblepharidineae Engler in Engler and Prantl
Nat. Pflanzenfam. I Teil, Abt. 1: iii, iv (1897).
Order Chytridineae Campbell Univ. Textb. Bot. 152 (1902).
Classes Archimycetae and Monoblepharideae Schaff'ner in Ohio Naturalist 9:
447,449 (1909).
Class Archimycetes Gaiimann Vergl. Morph. Pilze 15 (1926).
Uniflagellatae Sparrow Aq. Phyc. 21 (1943).
Parasites and saprophytes of simple structure (filamentous, of uniform diameter
or tapering; or unicellular, with or without rhizoids, i. e. tapering filamentous out-
growths), with cell walls of chitin, containing no cellulose; producing motile cells
with solitary posterior acroneme flagella. Type, Chytridium Olla Braun. From
6tt[o9ioc;, rearward, and KOVT6q^oar.
Chytrid is the English form of the generic name Chytridium, from Greek )(UTp(<;,
a jug. Braun (1856) applied this name to a colorless unicellular organism found
attached to green algae whose cells are penetrated by rhizoids which draw food from
them and kill them. By chytrids we mean organisms of body types of the general
nature of that of Chytridium. All such organisms were formerly treated as a single
taxonomic group. Couch (1938, 1941) showed that the organisms of chytrid body
type form three markedly distinct groups distinguished by types of flagellation. The
proper chytrids, those which legitimately constitute a taxonomic group, are marked
by swimming cells with solitary posterior acroneme flagella, and further by lack of
cellulose in the cell walls. The group thus marked includes, beside organisms of
chytrid body type, a few organisms of the filamentous body type of the typical fungi.
The cytoplasm of members of this group is described as peculiarly lustrous and
as containing shining globules. In mitosis (seen repeatedly, as by Dangeard, 1900,
Stevens and Stevens, 1903, Wager, 1913, and Karling, 1937), the sharp-pointed
spindle forms within the intact nuclear membrane. Some observers have seen centro-
somes at the poles. The nuclear membrane disappears toward the end of the process.
The formation of motile cells (zoospores and sometimes gametes) occurs in en-
larged cells. In these cells there are repeated simultaneous nuclear divisions. After
the last of these, uninucleate protoplasts, each one containing, ordinarily, one of the
above-mentioned shining globules, are separated by cleavage. On each of these
protoplasts a flagellum grows from the cell membrane at the point nearest that part
of the nucleus which represents a pole of the previous mitotic spindle. Among the
Blastocladiacea, the nucleus lies against the cell membrane and the flagellum appears
to spring from a granule within it (Cotner, 1930; Hatch, 1935). Similarly, in Clado-
Phylum Opisthokonta [111
chytrium, it appeared to Karling (1937) that the nucleolus generates the flagellum.
Within the developing swimming cell a body of granules assembles and produces a
"cap," prominent in stained material, on the anterior side of the nucleus, that is, on
the side away from the flagellum.
Nowakowski (1876) observed sexual processes in Polyphagus, and Scherffel
(1925) observed them in many other chytrids. Sexual processes were known in
Monoblepharis from the discovery of this genus, and have been studied in detail in
Allomyces by Emerson (1939, 1941) and Emerson and Wilson (1949).
The group thus characterized is of fewer than three hundred known species. One
takes no satisfaction in making it a phylum, but feels constrained to do so by its
isolation. Note has been taken that other groups including organisms of chytrid
body type, as Hyphochytrialea, Lagenidialea, and Phytomyxida, have nothing to do
with the proper chytrids. Furthermore, it will not do to thrust the proper chytrids
in with the groups of colorless flagellate and amoeboid organisms treated below as
phylum Protoplasta. One does not trust that group as natural, but it has a morpo-
logical continuity which would be defaced by the addition of this one.
Vischer, 1945, coined the name Opistokonten for organisms whose motile cells
have posterior flagella. Gams (1947) listed as such the green organisms Pedilomonas
and Chlorochytridion; the choanoflagellates; the proper chytrids; the Sporozoa (the
whole group by virtue of such examples as have flagellate stages); and the proper
animals. He inferred that these groups make up a major natural group derived
from the lowest green algae. This interesting hypothesis must as yet be treated as
far-fetched. Pedilomonas is scarcely known; it was described by Korschikoff, 1923,
as a green flagellate of somewhat the appearance of a Chlamydomonas lacking one of
its flagella. The flagella of the choanoflagellates are pantacroneme instead of acro-
neme. There remains a striking resemblance between the motile cells of the proper
chytrids and the sperms of animals. The nuclear cap of the former is quite similar,
in development and structure, to the beak of the latter.
The Opisthokonta are reasonably treated as a single class.
Class ARCHIMYCETES (A. Fischer) SchaflFner
Synonymy of the phylum.
Characters of the phylum.
Previous authors have arranged these organisms in a sequence from strictly uni-
cellular forms to typically filamentous forms. In the following treatment, this sequence
is reversed. The course of the evolution of the group is unknown, and it seems reason-
able to place the body types in the same sequence as among the Oomycetes. The class
is treated as two orders, Monoblepharidalea, essentially filamentous, and Chytridinea,
unicellular or producing filaments which taper or are swollen at intervals.
Order 1. Monoblepharidalea [Monoblepharidales] Sparrow in Mycologia 34: 115
(1942).
Suborder Monoblepharidineae Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
1: iv (1897).
Blastocladiincae Petersen in Bot. Tidsskr. 29: 357 (1909).
Order Blastocladiales Sparrow 1. c.
Opisthokonta whose bodies consist of filaments of uniform diameter, or are of
types apparently immediately derived from this. Saprophytes in fresh water or soil,
chiefly on vegetable remains. There are two families.
112 ] The Classification of Lower Organisms
Family 1. Monoblepharidacea [Monoblepharidaceae] A. Fischer in Rabenhorst
Kryptog.-Fl. Deutschland 1, Abt. 4: 378 (1892). Gonapodiineae and Gonapodiaceae
Petersen in Bot. Tidsskr. 29: 357 (1909). Producing extensive coenocytic filaments,
non-septate but with false septa of cytoplasm, anchored by rhizoids, reproducing
asexually by zoospores produced in sporangia which are usually terminal on the
filaments, the gametes produced in smaller antheridia and larger oogonia which are
in the more familiar forms terminal and subterminal on the filaments, the branches
commonly proliferating below them, the eggs without flagella.
The species, about a dozen, form three genera. In Monoblepharis, the zygote,
being the entire protoplast of the oogonium, moves out of the oogonium through a
terminal pore, becomes attached in the opening, and develops a thick wall. In
Monoblepharella the zygote, retaining the flagellum of the sperm, swims for a time
before becoming encysted. Gonapodya resembles Monoblepharella (Johns and Ben-
jamin, 1954). Myrioblepharis Thaxter is believed not to be an organism; it is de-
scribed as something which might be produced if sporangia of Monoblepharis were
parasitized by an infusorian.
Family 2. Blastocladiacea [Blastocladiaceae] Petersen in Bot. Tidsskr. 29: 357
(1909). Coenocytic filaments, in some examples of a false appearance of septation,
of the body type of the Rhipidiacea, i. e., differentiated into a basal cell anchored
by rhizoids and distal branches bearing reproductive structures, sometimes so re-
duced that the basal cell bears, or is itself, the reproductive structure; the reproduc-
tive structures including thin-walled zoosporangia, thick-walled resting spores which
germinate by releasing zoospores, and gametangia; the gametes morphologically
uniform or larger and smaller, all bearing flagella.
These organisms are not familiar, although they are readily isolated by baiting
pond water, or tap water to which soil has been added, with hemp seeds or pieces of
fruit. There are four genera, Allomyces, Blastocladia, Blastocladiella, and Sphaero-
cladia, with about twenty-five known species. Allomyces is of interest for varied life
cycles, and Blastocladia for a peculiar type of metabolism.
The first known species of Allomyces, A. Arbuscula, was discovered by Butler
(1911) on dead flies in water in India. The individuals are of the appearance of
minuscule shrubs, the branches divided by pseudosepta punctured in the middle and
ending in series of varicolored reproductive structures. Ordinary sporangia are
colorless, resting spores are brown, mature antheridia are pink, and mature oogonia
dull gray. Kniep (1929), in discovering the second species, A. javanicus, found that
the individuals are of two types, one bearing sporangia and resting spores, the other
oogonia and antheridia. Thus this organism has a complete life cycle of morpholo-
gically homologous haploid and diploid individuals. Kniep supposed that meiosis
occurs in the resting spores, and Emerson and Wilson (1949) established the point.
The chromosome number (n) oi A. Arbuscula is 7; that of A. javanicus var. macro-
gynus and of A. cystogenes is 14.
The life cycle of A. Arbuscula is the same as that of A. javanicus. In A. cystogenes,
the haploid stage consists merely of the zoospores from the resting spores; these
become encysted and germinate by releasing isogametes. Thus this species has a life
cycle essentially of the advanced type characteristic of animals. There are further
species of Allomyces in which a sexual cycle is believed not to occur.
In Blastocladia the basal cell bears directly multiple reproductive structures.
Organisms of this genus are less easily cultured than Allomyces; they require several
vitamins of the B group (Cantino, 1948). They tolerate oxygen, but do not require it.
Phylum Opisthokonta [113
They convert sugars to lactic and succinic acids, producing no CO2; the acids, if not
neutralized, check the growth of cultures (Emerson and Cantino, 1948; Cantino,
1949). Blastocladia appears to have lost the capacity to carry on the aerobic stages of
energesis, thus reverting to the type of metabolism characteristic of the supposedly
most primitive bacteria.
In Blastocladiella, the basal cell bears a single reproductive structure. DiflFerent
species have the same three types of life cycle which occur in Allomyces (Couch and
WhiflFen, 1942). In Sphacrodadia the vegetative body is reduced to the unicellular
condition which is characteristic of the following order rather than of this. The life
cycle is of the complete homologous type.
Order 2. Chytridinea [Chytridineae] (Schroter) Campbell Univ. Textb. Bot.
152 (1902).
Orders Myxochytridinae and My cocky tridinae A. Fischer in Rabenhorst Kryp-
tog. Fl. Deutschland 1, Abt. 4: 20, 72 (1892), not based on generic names.
Order Chytridiales Auctt.
Further synonymy as of the name of the phylum.
Opisthokonta which consist entirely or largely of more or less isodiametric bodies
called centers: the centers may send out filaments more slender than themselves,
generating at their ends further centers; or may be capable only of producing rhizoids,
i. e., tapering absorptive filaments; or may be by themselves complete individuals.
The chytrids are commonly thought of as prevalently parasitic on algae and
higher plants. They attack also rotifers, insects, nematodes, and other minute animals;
some parasitize other chytrids (Karling, 1942, 1948). It is probable, however, that
the majority of the group are saprophytic on organic remains. Some have been
cultured with no other organic food than cellulose (Haskins, 1939); new forms
have been discovered by baiting with, and culturing on, chitin (Karling, 1945;
Hanson, 1946) or keratin (Karling, 1946, 1947).
The following varieties of vegetative structure may be noted, (a) A zoospore,
settling upon the surface of an appropriate host or substratum, may penetrate this
by means of a walled filament which develops a terminal center; the center then
sends out rhizoids, and also filaments which generate further centers, (b) Develop-
ment may be as above except that only one center is formed. The body thus described
is of the Entophlyctis type of Sparrow (1943). (c) The zoospore may itself become
the single center, penetrating its host or substratum only by rhizoids. The resulting
body is of the Chytridium type if the center is in contact with the host or substratum,
of the Rhizidium type if it is not. (d) The protoplast of the zoospore may migrate
into the protoplast of the host and there become a center without rhizoids; the
resulting body is of the Olpidium type. To the varied bodies thus described, the
following terminology is applicable:
Pluricentric, with more centers than one; mono centric, with a single center.
Intramatrical, the center developing within the substratum or host; alternatively,
in a host, endobiotic.
Extramatrical or epibiotic, contrary to the foregoing.
Eucarpic, the center not constituting the entire body; holocarpic, the center con-
stituting the entire body.
The center regularly remains uninucleate during the vegetative phases and then
becomes the seat of successive simultaneous nuclear division, of cleavage, and of the
maturation of zoospores. Thus it is converted into a sporangium. In many forms, the
114]
The Classification of Lower Organisms
Fig. 22. — Monoblepharidalea: a-f, M o noble pharella Taylori x 1,000 after
Springer (1945); a, germinating spore producing a filament and a rhizoid; b, spor-
angium releasing spores; c, empty antheridium and sperm uniting with egg; d, sperms
escaping from antheridium and zygote escaping from oogonium; e, swimming zygote;
f. encysted zygote, g-i, Allomyces javanicus x 100 after Kniep (1929); g, asexual
(Continued bottom p. 115)
Phylum Opisthokonta [115
proximal part of the system of rhizoids develops a large swelling called the apo-
physis. In other forms, the center generates the sporangium as an outgrowth. In
these circumstances, the center is sometimes called an apophysis, but were better
called a presporangium. The sporangium discharges its spores, usually, through one
or more tubes which grow forth from it. The tube may open through a difTerentiated
cap, the operculum; the production of opercula appears to mark a natural subordi-
nate group.
Syngamy occurs in different chytrids in most of the possible fashions, by union of
like or unlike swimming cells, by the union of a swimming cell with a stationary one,
or by the establishment of contact by growth. The zygote regularly becomes a thick-
walled resting spore (asexual resting spores are also of frequent occurrence). Resting
spores germinate by producing zoospores. Meiosis has not been observed, but is be-
lieved to occur during the first nuclear divisions in the germinating zygote; the life
cycle is apparently of the primitive type, in which all cells except the zygote are
haploid {Phy so derma, or at least some of its species, is believed to be exceptional).
Sparrow (1943) recognized nine families. One of these does not appear tenable;
the remainder are distinguished as follows:
1. Sporangia not opening through opercula.
2. Eucarpic, i. e., producing rhizoids and
sometimes other filaments, the centers
not constituting the entire body.
3. Pluricentric Family 1. Cladochytriacea.
3. Monocentric.
4. Germinating spores generat-
ing the center as a distinct
body Family 2. Phlyctidiacea.
4. Zoospores themselves becom-
ing centers, and subsequently
sporangia or presporangia Family 3. Rhizidiacea.
2. Holocarpic, i. e., without rhizoids, the
individual consisting entirely of one or
more centers.
3. Centers becoming presporangia,
each one generating a cluster of
sporangia Family 4. Synchytriacea.
3. Centers proliferating, giving rise to
linear series of sporangia Family 5. Achlyogetonacea.
3. Each center becoming one spor-
angium Family 6. Olpidiacea.
individual with light sporangia and dark resting cells with pitted walls; h, branch of
sexual individual, the oogonia larger and darker than the antheridia; i, gametes.
j-m, Allomyces Arbuscula after Hatch (1935); j, k, gametes, x 1,000; 1, m, mitotic
figures in the gametangia, x 2,000. n-r, Blastocladiella cystogena, x 500, after Couch
and WhifFen (1942); n, individual producing a resting spore; O, resting spore germ-
inating by release of numerous naked protoplasts; these become flagellate zoospores,
p, which subsequently encyst; q, the protoplast of each cyst divides to produce four
gametes; r, young zygote with the flagella of both gametes.
116]
The Classification of Lower Organisms
' B^^
fiG. 23. — Chytridinea: a-c, Polyphagus Euglenae attacking cells of Euglena,
X 400, after Nowakowski (1876); in figure b, two individuals have made contact
and a zygote is developing at the point of junction; c, sporangium, d-i, Olpidium
Allomycetos attacking Alomyces anomalus, x 1,000, after Karling (1948); d, e, zoo-
spores; f, sporangium of the host beset with many parasites; g, h, resting cells of the
host containing respectively sporangia and resting cells of the parasite; i, germina-
tion of resting cell.
Phylum Opisthokonta [117
1. Sporangia opening through opercula.
2. Pluricentric Family 7. Nowakowskiellacea.
2. Monocentric Family 8. Chytridiacea.
Family 1. Cladochytriacea [Cladochytriaceae] Schroter in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1: 80 (1892). Family Hyphochytriaceae [Cladochytria-
ceae) A. Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 131 (1892), in
part. Family Physodermataceae Sparrow Aq. Phyc. 304 (1943). Pluricentric chytrids,
the sporangia not operculate. The members of this family are of the same body type
(designated by Karling, 1931, the rhizo mycelium) as the anisochytrid Hyphochy-
trium and the Nowakowskiellacea of the present order. In most Cladochytriacea the
rhizomycelium includes pairs of swollen cells ("turbinate organs") which give a false
appearance of conjugation. There are some forty known species, mostly of two
genera, Cladochytrium, saprophytic in vegetable remains, and Physoderma (including
Urophlyctis), parasitic in higher plants. Sparrow (1946, 1947) discovered in certain
species of Physoderma an alternation of morphologically distinguishable generations,
both on the same hosts; the generations are presumably haploid and diploid, but this
has not been established by observation of syngamy and meiosis. Polychytrium grows
well only on chitin (Ajello, 1948).
Family 2. Phlyctidiacea [Phlyctidiaceae] Sparrow in Mycologia 34: 114 (1942).
Family Sporochytriaceae [Rhizidiaceae, Polyphagaceae) subfamily Metasporeae A.
Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 85 (1892). Monocentric
eucarpic chytrids, the centers developed at the ends of filaments which grow from the
zoospores, sporangia without opercula.
These are the most familiar chytrids. There are more than one hundred species.
Many are parasitic, on blue-green and green algae, diatoms, pollen grains, nematodes,
and other minute fresh-water life; others are saprophytic, on cellulose, chitin, or
keratin. Rhizophidium, the most numerous genus; Phlyctidium, Phlyctorhiza, Ento-
phlyctis, Diplophlyctis, Loborhiza, etc.
Family 3. Rhizidiacea [Rhizidiaceae] Schroter in Engler and Prantl Nat. Pflanzen-
fam. I Teil, Abt. 1: 75 (1892). Family Sporochytriaceae [Rhizidiaceae, Polyphaga-
ceae) A. Fischer in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 4: 85 (1892)
and subfamily Orthosporeae op. cit. 124. Monocentric eucarpic chytrids, the zoospores
enlarging and becoming centers, which in turn become sporangia or presporangia; the
sporangia without opercula. A moderate number of species, parasitic on blue-green
or green algae, flagellates, or diatoms; or chitinophilous, saprophytic in the shed
exoskeletons of insects. Rhizidium, Siphonaria, Asterophlyctis, Polyphagus, etc.
Polyphargus Euglenae Nowakowski (1876) is a classic example. The centers lie free
in the water, parasitizing cysts of Euglena through freely branching and widely
spreading rhizoids. Most centers act as presporangia. Syngamy occurs when a rhizoid
from one center makes contact with another center. The protoplasm of the latter
migrates into the tip of the rhizoid, which swells and becomes a resting spore.
Family 4. Synchytriacea [Synchytriaceae] Schroter op. cit. 71. Family Merol-
pidiaceae [Synchytriaceae) A. Fischer op. cit. 45. Holocarpic chytrids, the intra-
matrical cell unwalled in the vegetative condition, becoming a presporangium or a
resting spore, either of which gives rise to a cluster of sporangia. Synchytrium,
parasitic on higher plants; Micromycopsis on Conjugatae.
Family 5. Achlyogetonacea [Achlyogetonaceae] Sparrow in Mycologia 34: 114
(1942). Chytrids without rhizoids, the intramatrical center proliferating and pro-
ducing a linear series of centers, each of which becomes a sporangium without an
118] The Classification of Lower Organisms
operculum. Achlyogeton, in green algae, diatoms, and nematodes; of very much the
appearance of certain Lagenidialea.
Family 6. Olpidiacea [Olpidiaceae] Schroter op. cit. 67. Family Monolpidiaceae
[Olpidiaceae) A. Fischer op. cit. 20. Holocarpic chytrids, each individual a single
intramatrical parasitic center, naked until the reproductive phase, when it becomes
a sporangium without an operculum. Olpidium, attacking blue-green and green algae,
diatoms, flagellates, Allomyces, Vampyrella, rotifers, and nematodes. Rozella, attack-
ing Oomycetes and producing spiny resting spores, has been confused with certain
Lagenidialea. The genera Sphaerita and Nucleophaga of Dangeard, including
intracellular parasites of amoebas and Infusoria, have been placed in this family;
it seems more probable that they should be placed among bacteria of family Rickett-
siacea.
Family 7. Nowakowskiellacea [Nowakowskiellaceae] Sparrow in Mycologia 34:
115 (1942). Family Megachytriaceae Sparrow Aq. Phyc. 378 (1943). Pluricentric
chytrids, the sporangia with opercula. A moderate number of saprophytes on material
of green algae and higher plants. Nowakowskiella, Megachytrium, etc. Zygochytrium
was described by Sorokin, 1874, as living on decaying insects, producing multiple
operculate sporangia, and exhibiting a conjugation of filaments to produce zygotes
much like those of Zygomycetes. It has apparently not been reobserved.
Family 8. Chytridiacea [Chytridiaceae] Cohn in Hedwigia 11: 18 (1872). Family
Chytridieen de Bary and Woronin in Berichte Verhandl. Naturf. Gess. Freiburg 3
(Heft 2) : 46 ( 1864). Monocentric eucarpic chytrids, the sporangia operculate. Some
fifty species, the majority parasitic on fresh water algae. Chytridium, etc. Catenochy-
tridium, saprophytic in cast-off exoskeletons of insects.
Chapter IX
PHYLUM INOPHYTA
Phylum 5. INOPHYTA Haeckel
Order Fungi L. Sp. PI. 1 1 7 1 (1 753 ) .
Hysterophyta Link, 1808.
Classes Fungi and Lichencs Bartling Ord. Nat. 4 (1830).
Regnum Mycetoideum Fries Syst. Myc. 1: Ivi (1832).
Class Lichenes and section Hysterophyta with class Fungi Endlicher Gen. PI. 11,
16 (1836).
Stamm Inophyta Haeckel Gen. Morph. 2: xxxvi (1866).
Subdivision Fungi Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889).
Division Eumycetes Engler Syllab. ed. 3: 25 (1903).
Phylum Carpomyceteae Bessey in Univ. Nebraska Studies 7: 249 (1907).
Stamm Mycophyta Pascher in Beih. bot. Centralbl. 48, Abt. 2: 330 ( 1931 ).
Kingdom Mycetalia Conard Plants of Iowa iv (1939).
Phylum Eumycophyta Tippo in Chron. Bot. 7: 205 (1942).
Parasites and saprophytes without flagellate stages, the bodies filamentous, the
w;,lls containing no cellulose.
This group represents the conventional division or subdivision Fungi of the
kingdom of plants, excluding, of course, the bacteria, Oomycetes, chytrids, and
Mycetozoa. The name Fungi, used as a scientific name, is properly to be applied,
by authority of Linnaeus, to an order. Agaricus campestris L. will be recognized
as the standard species of the phylum and of the order.
Those who study Inophyta are accustomed to use, for soma and filament respec-
tively, the terms mycelium and hypha. The walls of the hyphae are believed to consist
of pectic material. A small percentage of chitin is usually present (Schmidt, 1936);
cellulose is totally absent (Thomas, 1928; Nabel, 1939; Castle, 1945). The organism
Basidiobolus, having hyphae walled with cellulose, is tentatively retained among
Inophyta as an exception.
The multiplication and dissemination of those organisms is by spores, of various
types, scattered in the air. Most Inophyta produce two or more kinds of spores, some
of them asexually, others as features of a sexual cycle. Spores produced within cases
are called endospores, and the cases sporangia. Other spores are produced externally,
commonly by constriction of the ends of hyphae. Spores thus produced are called
conidia, and the hyphae or other structures which bear them, conidiophores. Spores
are commonly produced not directly on the mycelium but on macroscopic structures
of various types, all of which may be called by the familiar term fruit. The common
mushroom as we see it is a fruit; it is the temporary spore-producing structure of
an organism whose soma consists of filaments living saprophytically in the soil
below.
It is expedient to mention at this point the growths called lichens, which are
traditionally treated as a taxonomic group, either subordinate to Fungi or of the
same rank. Lichens are gelatinous or thallose growths, usually of an impure green
color, common everywhere, terrestrial or epiphytic, as on stones, trees, or fence
posts. The microscope, in the hands of de Bary and others, showed that they consist
of cells of two types, colorless filaments like those of Inophyta, and pigmented
120] The Classification of Lower Organisms
cells of quite the character of those of certain algae. De Bary (in Hofmeister, 1866)
concluded that some lichens are not organisms but combinations of totally diverse
organisms. Presently (1868) he was convinced by the work of Schwendener, soon
(1868) published under his own name, ". . . dass die Flechten sammt und senders
keine selbststiindigen Pflanzen seien, sondern Pilze aus der Abtheilung der Ascomy-
ceten, denen die fraglichen Algen — deren Selbststandigkeit ich also nicht bezweifle —
ah Nahrpflanzen dienen." In 1879 de Bary coined the term symbiosis to designate
the association of different kinds of organisms. In de Bary's usage the term included
parasitism; in general usage, it means association to mutual advantage. The lichens
are a classic example of symbiosis.
Clearly, the group of lichens is not to be maintained; the algal components are
known to have natural places among algae, and the inophyte components are to be
assigned to their natural places among Inophyta, almost all in various orders of
class Ascomycetes. This has already been done by Clements (1909) and Clements
and Shear ( 1931 ). The numerous names which students of lichens have given to them
are to be applied to the inophyte components.
Another common example of symbiosis involving inophytes is furnished by at least
some of those which live on or in the tissues of higher plants without killing them
(Kelley, 1950). They occur mostly on roots. Frank (1885) coined the term mycorhiza
to designate the combination of roots and inophytes; it will be more convenient to
hold that this term designates the inophyte component of the combination. Such
mycorhizae as cover the growing tips of roots are helpful to their hosts by serving as
agents of absorption.
Jones (1951) estimated the number of species of Inophyta as 40,000. This is
surely an extreme underestimate. Martin (1951) gives reason for believing the num-
ber to be about as great as that of flowering plants, of the order of 300,000.
The early classifications of "fungi," as by Persoon (1801) and Fries (1821-1832),
were based on gross characters. They presented, along with recognizable groups
whose names are to be applied in order of priority, others which were mere random
assemblages, and whose names are to be abandoned as nomina confusa. De Bary (in
Hofmeister, 1866; 1884), having applied comparatively modern methods, established
a dozen groups (under German names). These, so far as they are retained in the
present phylum, have been assembled as three classes distinguished by details of the
sexual cycle. A fourth class, acknowledgedly artificial, is maintained for the accomo-
dation of the numerous and important fungi whose sexual cycles are unknown. The
termination -mycetes, of the names of the classes and also of various subordinate
groups, is the Greek ^uKr]T£q, the plural of (auKT^c;, a mold or mildew. The termi-
nation -mycetae which some authors have used is a solecism.
1. Reproducing sexually, or by apomictic pro-
cesses clearly of sexual origin.
2. The zygote becoming a thick-walled
resting cell; fruits none or inconsiderable Class 1. Zygomycetes.
2. The zygote not becoming a thick-walled
resting cell; mostly producing fruits.
3. The zygotes giving rise, usually in-
directly, to sporangia called asci,
each typically containing eight
spores called ascospores Class 2. Ascomycetes.
Phylum Inophyta [121
3. The zygotes giving rise indirectly to
conidiophores called basidia, each
bearing typically four conidia
called basidiospores Class 4. Basidiomycetes.
1. Not known to reproduce sexually Class 3. Hyphomycetes.
Class 1. ZYGOMYCETES (Sachs ex Bennett and Thistleton-Dyer)
Winter
Zygomyceten Sachs Lehrb. Bot. ed. 4: 248 (1874).
Zygomycetes Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed. 847
(1875).
Class Zygomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1:
32 (1879).
Order Zygomycetes Engler Syllab. 23 (1892).
Class Zygomyceteae Schaffner in Ohio Naturalist 9: 449 ( 1909).
Inophyta whose zygotes are thick-walled resting cells, in germination giving rise to
spores indistinguishable from those produced asexually; hyphae usually without cross-
walls; mostly not producing fruits. The standard species is Mucor Mucedo L.
Among the Inophyta as here limited, the Zygomycetes appear to be primitive (an
alternative hypothesis, that certain Ascomycetes are primitive, will be discussed be-
low). Traditionally, the Zygomycetes are associated with the Oomycetes. The asso-
ciation is probably mistaken, being based merely on similarity of body form: the
Zygomycetes are terrestrial instead of aquatic, produce no flagellate cells, have no
cellulose in their cell walls (except in Basidiobolus) , and do not produce female
gametes by the cutting out of cells within a cell. In later editions of Engler's Syllabus
(1924), one finds most of the chytrids included among the Zygomycetes, instead of
in their conventional place among the Oomycetes. The hypothesis thus suggested,
that the Opisthokonta may represent the ancestry of the Inophyta, is attractive, but
not to present knowledge supported by convincing evidence. Class Zygomycetes and
phylum Inophyta must as yet be regarded as of unknown origin and treated as isolated.
There are some 500 known species of Zygomycetes. They form two orders. The
bulk of the group, and the typical examples, are order Mucorina. A minority,
distinguished by parasitism and by explosively discharged conidia, are order
Entomophthorinea.
Order 1. Mucorina [Mucorini] Fries Syst. Myc. 3: 296 (1832).
Suborder Mucorineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1: iv (1897).
Order Mucorineae Campbell Univ. Textb. Bot. 158 (1902).
Order Spirogyrales (presumably in part only) Clements Gen. Fung. 12 (1909).
Order Mucorales Smith Crypt. Bot. 1 : 405 (1938).
Order Zoopagales Bessey Morph. and Tax. Fungi 117 (1950).
The typical Zygomycetes, mostly saprophytic, not producing explosively dis-
charged conidia [Piloholus produces explosively discharged sporangia).
The asexual reproductive structures of the supposedly primitive Mucorina, as
Mucor and Rhizopus, are solitary globular sporangia terminal on erect hyphae. In
the developing sporangium, a dome-shaped basal sterile area, the columella, is set
apart by cleavage followed by deposition of a wall. The protoplasm above the
122]
'I'hc Classification of Lower Organisms
Fig. 24. — Zygomycetes: a-d, Rhizopus nigricans; a, sporangia x 50; b-d, prega-
mctes, suspensors and gametes, and zygote x 200. e. Zygote of Phycomyccs nitens
after Blakeslcc ( 1904). f, g, Conidiophore with young conidia, and mature conidia,
of Syncephalis pycnosperma after Thaxtcr (1897). h, i, Conjugation of Synce-
phalis nodosa after Thaxter, op. cit. j, Sporangium of Synccphalastrum raccmosum
after Thaxter, op. cit. k, Sporangium of Flaplospoiangium lignicola after Martin
(1937), x 1,000.
Phylum Inophyta [ 123
columella undergoes cleavage to form spores, which may remain plurinucleate
(Swingle, 1903). Other members of the order exhibit transitions (apparently two
distinct series of transitions) from sporangia as just described to typical conidia.
Syngamy occurs when the tips of pairs of hyphae meet and are cut off by crosswalls
to act as multinucleate gametes. The process is regarded as conjugation, although
the gametes of a pair are usually not of the same size. Conjugation does not occur
at random, but, in most Zygomycetes, between branches from hyphae of two mating
types, designated plus and minus (the distinction of mating types is not identical
with the differentiation of sexes). Zygomycetes were the first group reproducing by
conjugation in which a distinction of mating types was discovered; the discovery was
by Blakeslee (1904).
Syngamy is preceded by a flare of mitoses in the gametes. The mitotic figures are
sharp-pointed, as though centrosomes were present; the haploid chromosome number
appears to be 2. The process is not meiotic (Moreau, 1913). After these divisions,
the walls between the gametes break down and the nuclei unite in pairs. Unpaired
nuclei, presumably contributed in excess by one gamete or the other, undergo disso-
lution (Keene, 1914, 1919). Ordinarily, the zygote enlarges and becomes a thick-
walled resting spore; in some examples, the resting spore forms as an outgrowth on
what was one of the gametes. In Phycomyces, Absidia, and Syncephalis, the hyphae
which have produced the gametes, and to which the zygote remains attached, .send
out branches which form a layer about the zygote. These branches might be inter-
preted as making up fruits. Endogone produces definite fruits of considerable size.
A zygote germinates by production of a hypha bearing a sporangium (Blakeslee,
1906). Meiosis is believed to occur in the course of germination.
While Mucorina in general are saprophytic, some of them are parasitic on others,
Piptocephalis and Chaetocladhim on Mucor, and Parasitella on Absidia. Drechsler
(1935, 1937) discovered a number of organisms apparently of this group parasitizing
amoebas and nematodes in the soil.
The Mucorina may be treated as five families.
1. Not producing macroscopic fruits.
2. Not parasitic on amoebas or nematodes.
3. All spores produced in sporangia
with columellae Family 1. Mucoracea.
3. Not as above.
4. Producing sporangia or else
conidia as outgrowths from a
knob, homologous with a
sporangium, solitary on an un-
branched stalk Family 2. Piptocephalidacea.
4. Sporangia or conidia solitary
and terminal on branches of a
branched sporangiophore or
conidiophore; sporangia, if
produced, without columellae Family 3. Mortierellacea.
2. Parasitic on amoebas or nematodes Family 4. Zoopagacea.
1. Producing macroscopic fruits Family 5. Endogonacea.
Family 1. Mucoracea [Mucoraceae] Cohn in Hedwigia 11: 17 (1872). Mucorina
whose spores are produced exclusively in sporangia with columellae solitary on un-
branched sporangiophores. Mucor L., typified by M. Mucedo, is now limited to a
124] The Classification of Lower Organisms
small group mostly saprophytic on manure. Pilobolus, another coprophilous genus,
is distinguished by sporangiophores which become swollen at the summit, bend
toward the light, and discharge the sporangia violently to a distance of several meters.
Rhizopus nigricans, the common black bread mold; Phycomyces, Ahsidia, Sporodinia,
Zygorhynchus.
Family 2. Piptocephalidacea [Piptocephalidaceae] Schroter in Engler and Prantl
Nat. Pflanzenfam. I Teil, Abt. 1: 132 (1893). Family Choanephoraceae Schroter op.
cit. 131. Mucorina producing sporangia without columellae, or conidia, in compact
clusters terminal on unbranched stalks. Blakesleea, transitional between the preceding
family and this, may produce solitary sporangia with columellae, or else, as out-
growths from the primordia of sporangia, clusters of minuscule sporangia without
columellae. Cunninghamella, producing heads of globular conidia; Syncephalastrum,
with clustered cylindrical sporangia; Syncephalis and Piptocephalis, producuig
clustered chains of conidia.
Family 3. Mortierellacea [Mortierellaceae] Schroter op. cit. 130. Family Chaeto-
cladiaceae Schroter op. cit. 131. Mucorina whose sporangiophores or conidiophores
are branched, the sporangia (without columellae) or conidia solitary and terminal
on the branches. Thamnidium, Chaetocladium, Mortierclla, Haplosporangium.
Family 4. Zoopagacea [Zoopagaceae] Drechsler in Mycologia 27: 37 (1935). Mu-
corina parasitic in amoebas or nematodes, producing conidia. The hosts of Zoopaga-
cea inhabit the soil and are infected by contact with hyphae or conidia. From the point
of contact, a hypha grows into the host and gives rise to a mycelium; this is in some
examples reduced to a single coiled cell. The host being killed, the parasite sends
out hyphae which may produce conidia, usually in chains, or else may conjugate and
produce zygotes. Endocochlus, Cochlonema, Bdellospora, Zoopage, Acaulopage,
Stylopage.
Family 5. Endogonacea [Endogonaceae] Paoletti in Saccardo Sylloge Fungorum
8: 905 (1889). Endogonei Fries. Mucorina saprophytic in soil or wood, producing
macroscopic subterranean fruits. The fruits may reach a diameter of 2 cm. Within
them, the tips of hyphae are cut off by crosswalls, and develop either into sporangia
without columellae or into gametes.
Order 2. Entomophthorinea [Entomophthorineae] (Engler) Campbell Univ.
Textb. Bot 161 (1902).
Suborder Entomophthorineae Engler in Engler and Prantl Nat. Pflanzen-
fam. I Teil, Abt. 1: iv (1897).
Order Entomophthorales Smith Crypt. Bot. 1: 408 (1938).
Zygomycetes, mostly parasitic, producing explosively discharged conidia [Masso-
spora, while clearly belonging to the group, is an exception to the stated character).
These organisms, although of the general nature of ordinary Inophyta, exhibit
cytological characters markedly distinguishing the two families from the generality
of Inophyta and from each other. The position here given to them is the customary
one; it is doubtful that it is natural.
Family 1. Entomophthoracea [Entomophthoraceac] Berlese and de Toni in Sac-
cardo Sylloge 7: 280 (1888). Most species are parasitic in the bodies of insects,
whose tissues they replace. The hyphae become divided by crosswalls, and the multi-
nucleate cells thus produced tend to round up and become separate. A well-nourished
cell may send forth a hypha which reaches the outer air and whose tip is cut off and
discharged in the direction of the light. Martin (1925) and Couch (1939) described
Phylum Inophyta [ 125
the mechanism of discharge. The conidiophore ends in a columella projecting into
the base of the conidium. The columella develops a double wall. Increasing pressure
within the conidium causes a sudden eversion of the wall on the side of the conidium,
and this movement throws the conidium forth to a distance of perhaps 1 mm. Coni-
dia which come down on unfavorable substrata may form and discharge secondary
conidia.
Adjacent cells may conjugate, the thick-walled zygote forming either in one of
them or as an outgrowth from one of them. Many examples produce thick-walled
resting spores without conjugation.
Olive (1906) described the nuclei and the process of mitosis in Empusa. The
resting nuclei are fairly large, 7-9 [I in diameter. In the course of division, two stain-
resistant granules are seen, with strands of chromatin radiating from them. These
move apart, while the nucleus becomes dumb-bell shaped. The nuclear membrane
remains intact and division is completed by its constriction. As Olive remarked, the
process is much as in Euglena.
Entomophthora, Empusa, and Massospora attack insects; the first produces zygotes,
while the other two produce asexual resting spores; Massospora does not discharge
the conidia violently. Conidiobolus and Delacroixia are saprophytic. Completoria
attacks the prothallia of ferns. Ancylistes, a parasite in the green alga Closterium,
was formerly included among chytrids or Oomycetes. Berdan (1938) showed that it
belongs here; it produces conidia and zygotes quite of the character of the present
group, and does not produce zoospores.
Family 2. Basidiobolacea [Basidiobolaceae] Engler and Gilg Syllab. ed. 9 u. 10:
45 (1924). Basidiobolus ranarum Eidam (1886) occurs in the intestinal contents of
frogs and toads as uninucleate cells, solitary or in brief filaments, walled with cellu-
lose. In manure the filaments develop into a scant branching mycelium. The proto-
plasm gathers in the ends of erect hyphae which are cut off as conidia and discharged.
Conjugation occurs between adjacent cells of a filament. It is preceded by a single
nuclear division in each gamete (Fairchild, 1897). In this process, the nuclear mem-
brane disappears and the numerous minute chromosomes are found in a blunt-ended
spindle without centrosomes. Each gamete form a papilla; one of the two nuclei
enters the papilla, whose contents, after being cut oft by a wall, die and disappear.
The gametes and their nuclei unite and the zygote secretes a thick wall.
Class 2. ASCOMYCETES (Sachs ex Bennett and Thistleton-Dyer)
Winter
Order .4jco5porgag Cohn in Hedwigia 11: 17 (1872).
Ascomyceten Sachs Lehrb. Bot. ed. 4: 249 (1874).
AscoMYCETES Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed. 847
(1875).
Class AscoMYCETES Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1 : 32
(1879).
Class Ascosporeae Bessey in Univ. Nebraska Studies 7: 295 (1907).
Class Ascomycetae Schaffner in Ohio Naturalist 9: 449 (1909).
Inophyta which produce, as a feature of the sexual cycle, sporangia called asci, in
which the spores, called ascospores, typically eight in number, are delimited by the
manner of cell division called free cell formation, i.e., in such fashion as to exclude
a part of the cytoplasm.
126] The Classification of Lower Organisms
The hyphae of Ascomycetes are septate and the cells most often uninucleate.
Most Ascomycetes produce, beside the ascospores, conidia of one type or another.
A mycelium may produce a mass of densely woven hyphae with conidia on the sur-
face; such a mass is called an acervulus or sporodochium. Either a mycelium or an
acervulus or sporodochium may send up spore-bearing columns called coremia.
Many Ascomycetes produce, either directly from the mycelium or from special
structures consisting of interwoven hyphae, globular or flask-shaped structures which
produce conidia internally and release them through a pore. These structures are
called pycnidia, and the spores pycniospores. In many examples, the pycniospores
are capable of functioning as sperms; so far as this is true, the pycnidia may alter-
natively be called spermagonia, and the pycniospores spermatia.
Hyphae woven into a mass may go into a resting condition, becoming thick-walled,
hard, and usually dark in color. The resulting structure is a sclerotium. If a structure
of the general nature of an acervulus, sporodochium, or sclerotium gives rise either
to pycnidia or to fruits bearing asci, it is called a stroma.
As to asci and ascospores, Dangeard (1893, 1894, 1907) reached definitely the
conclusion that they are essentially sexual products. There had been earlier observa-
tions, beginning with de Bary, 1863, that there are meetings, coilings together, and
fusions of hyphae as a preliminary to the production of asci. Many ascomycetes are
of two mating types; this was first discovered of Glomerella, by Edgerton (1914). As
Dodge (1939) remarks, the mating types are not sexes; in forms producing recogniz-
able male and female reproductive structures, each mating type may produce both.
In Ascomycetes which may be regarded as primitive, difTerentiated male and fe-
male cells are produced. The male cell or antheridium is ordinarily terminal on a
hypha. The female cell (constituting, together with other differentiated cells of the
same hypha, if any are present, the ascogonium) may be terminal; more often it bears
an elongate cell, or a chain of cells, called the trichogyne, and having the function
of reaching the antheridium. In some Ascomycetes, antheridia are produced, but syn-
gamy does not take place; the egg is binucleate or multinucleate, and the nuclei
within it take the part of gamete nuclei in further development. There are others in
which no antheridia are produced. Hansen and Snyder (1943) found, in Hypornyces
Solani var. Cucurbitae, that "any part of the living thallus, ascospores, conidia or
bits of the mycelium could act as the male fertilizing agent." There are forms in
which fusions take place between undifferentiated hyphal cells; and yet others in
which it appears that the paired nuclei involved in sexual processes arise by divisions
of a single nucleus originally present in a spore.
In some Ascomycetes, syngamy is followed immediately by karyogamy, and the
zygote develops directly into a single ascus. In the overwhelming majority of the
group, asci are produced indirectly, and there is no fusion of nuclei until this takes
place. The zygote sends out hyphae called ascogenous hyphae, recognizably different
from the vegetative ones. The cells of the ascogenous hyphae arc binucleate; or,
arising from a multinucleate zygote, become binucleate by the establishment of
crosswalls. The two nuclei of each cell divide concurrently and the cell walls are so
placed that each cell receives nuclei of different origin. This effect is achieved in the
final cell division before ascus formation by a peculiar process called crozier forma-
tion. The terminal cell of the ascogenous hypha becomes bent to the form of a hook;
the nuclei divide concurrently, and cell walls appear between the daughter nuclei of
each pair; the middle cell of the row of three thus produced remains binucleate and
becomes an ascus. The uninucleate terminal and basal cells lie side by side, and may
Phylum Inophyta [127
fuse to form a binucleate cell which may become an additional ascus, or else may
grow forth and give rise to more asci than one.
The stage consisting of cells with two nuclei of different origin is called the
dikaryophase. It is characteristic of Ascomycetes ,and also of Basdiomycetes: among
Inophyta, it is a normal and familiar thing. To a concept of cytology founded on
studies overlooking the Inophyta, it would appear an extreme anomaly, almost an
impossibility. It has the appearance of a rather awkward device for making cells
genetically and physiologically diploid while the nuclei remain haploid. In most
Ascomycetes it is a brief stage, but there are some, as Taphrina, whose mycelium
consists prevalently of binucleate cells.
The detailed behavior of nuclei in the ascus was first described by Harper (1895,
1897, 1900) from studies of Peziza, Sphacrotheca, Erysiphe, and Pyronema. The two
nuclei in the primordium of the ascus unite into one. The fusion nucleus divides
three tim.es, each time in much the same manner. A centrosome with astral rays is
present at the nuclear membrane, apparently outside. It divides, and a spindle forms,
inside the intact nuclear membrane, between the daughter centrosomes. The chromo-
somes appear and divide. As they move toward the poles of the spindle, the nuclear
membrane collapses or dissolves, leaving the spindle free in the cytoplasm. The
mass of chromatin at each pole of the spindle shreds out into a nuclear network,
duly surrounded by a nuclear membrane and usually containing a nucleolus.
Haploid chromosome numbers of Ascomycetes (all of which have been observed in
the ascus) include the following:
Ascoidea rubescens, fide Walker (1935) 2
Eremascus alhus, fide De Lamater et al. (1953) 6
G/om^r(?//<z, fide Lucas (1946) 4
HypornycesSolaniv:xr.Cucurbitae,fi.dtY{.\r?,ch.{\9'^9) .... 4
Lachnea scutellata,^dt^ro\\'n {\9\\) 5
Neurospora crassa, fide McClintock (1945) 7
Peziza do miciliana,^dt ?)c\\n\iz {\921) 8
Phyllactinia corylea, fide Colson (1938) 10
Pyronema confiuens\?Lr. igiieum, ^dtV>ro\vn {\9\b) 5
Taphrina deformans, fide Martin (1940) 4
According to Harper, when the third division in the ascus is complete, each of
the eight nuclei produced by it thrusts forths its centrosome upon a beak. The astral
rays of the centrosomes become recurved in the cytoplasm about the nucleus, and
grow and multiply until they are converted into a smooth membrane, outside of
which a wall is deposited. Most observers have not seen so much detail. Brown (1911)
and Dodge (1937) describe the cell membrane of the ascus, apparently under the
influence of the centrosome of each nucleus, as cutting into the cytoplasm in an ellip-
soid pattern. In Taphrina (Martin, 1940), the cytoplasm of the spores is delimited
simply by accumulation about the nuclei. By whatever process the ascospores are cut
out. some of the cytoplasm of the ascus is excluded and left without nuclei. Harper
(1899) proposed to limit the older term free cell formation to processes which have
this effect; he observed that the occurrence of such processes distinguishes asci from
the sporangia of Oomycetes and Zygomycetes, in which spores are cut out by cleavage.
Harper believed that a fusion of nuclei follows immediately the fusion of gametes;
that the karyogamy observed in the ascus is a uniting of diploid nuclei, producing
tetraploid nuclei; and that the characteristic three nuclear divisions in the ascus are
necessary for reduction of the chromosome number from tetraploid to haploid. These
128] The Classification of Lower Organisms
hypotheses, long accepted as possible, were disproved by genetic studies by Betts and
Meyer (1939) and Keitt and Langford (1941). In the asci of many species, the
spores lie in a single series in which their order is determined by the divisions which
produce their nuclei. By refined technique, the spores from a single ascus may be
identified, separated, and cultivated. It is then observed that the mycelia grown from
the first four spores may differ in some particular character from those grown from
the second four spores; those from the first pair of spores may differ from those from
the second; but those from two members of any of the pairs, first, second, third or
fourth, are always alike. These observations mean that the first two divisions in the
ascus constitute the meiotic process, the third being mitotic. Lucas (1946) obtained
cytological evidence refined enough to confirm this conclusion.
Asci are almost always produced in fruits, which may be called ascocarps. The
ascocarp aside from the asci arises usually from vegetative hyphae; in the Ascomy-
cetes regarded as primitive, it does not begin to develop until after fertilization, but
in the higher ones it may develop in advance of fertilization and become the seat
of this process.
There are several types of ascocarps, among which three are most familiar. A
small ascocarp completely enclosing the asci is a cleistothecium. Cleistothecia were
formerly included under the term perithecium; that term will better be limited
to small fruits which are globular or vase-like, opening through a single pore, the
ostiole, and differing from the pycnidia already described in producing ascospores
instead of conidia. A fruit in which the asci form a broad layer which is typically
fully exposed at maturity, the whole being ordinarily of the form of a disk or cup,
larger than a cleistothecium or perithecium, is an apothecium.
Asci produced in perithecia or apothecia usually discharge the ascospores vio-
lently. The mechanism of discharge is apparently simply turgidity. Some asci show
no visible adaptations for the discharge of spores; others have lids (opercula)
whose position determines the direction of discharge. Certain large apothecia can
throw the spores to a distance of 10-20 cm.; the discharge is so governed by tempera-
ture and humidity as to occur in gently moving rather than in still air. By blowing
across these apothecia one can make them throw out a visible cloud of spores.
Heald and Walton (1914) reviewed many older observations of violent discharge by
perithecia, the oldest by Pringsheim on Sphaeria Scirpi, 1858. Rankin, 1913, found
that each ascus in turn breaks loose, comes up to the ostiole, projects through it,
throws out its spores, and collapses to make room for another. Weimer (1920) found
that the perithecia of Pleurage curvicolla bend toward the light and throw the spores
to a maximum distance of 45 cm., which is apparently the record.
There is a widely entertained hypothesis that the Ascomycetes evolved from the
red algae. It appears to have developed from a piece of classification by Sachs
(1874), who proposed a class Carposporeen, to consist of the red algae, certain higher
green algae, and the Ascomycetes and Basidiomycetes. A number of resemblances
support it. Both red algae and Ascomycetes include many parasites; both lack
flagellate cells; both have differentiated gametes, the egg bearing a trichogyne; in
both, fertilization leads to further development before spores are produced. In
addition to these genuine resemblances, an imaginary one was influential, namely
the double fertilization ascribed to the red algae by Schmitz and to the Ascomycetes
by Harper. Numerous as these resemblances are, they are not now believed to indicate
relationship. Atkinson (1915) formulated the counter-argument. The Ascomycetes
resemble the Mucorina in nutrition, in producing no flagellate cells, and in multi-
Phylum Inophyta [ 129
nucleate gametes. The germination of the zygote of the Mucorina, by the production
of a hypha bearing a sporangium, resembles the production of ascogenous hyphae by
the zygotes of Ascomycetes. Two principal changes would convert Mucorina into
Ascomycetes: the zygote should cease to be a resting spore, and cell division within
the sporangium should be by free cell formation. This could happen if the centro-
somes of the ultimate nuclei of the sporangia were in control of cleavage, and if
these nuclei were so far separated that considerable areas of cell membrane would
lie beyond the influence of the centrosomes, with the effect that the cell membrane,
furrowing in to delimit a spore around each nucleus, would leave some of the cyto-
plasm outside of all of the spores. The organisms listed below as the first order of
Ascomycetes, Endomycetalea, are but poorly known, yet seem genuinely to represent
the transition from Mucorina to typical Ascomycetes.
It is not yet possible to formulate a system of orders of Ascomycetes with the
expectation that it will not be found to require much amendment^. The following
will serve tentatively; excellent contemporary authority makes several orders each
of the ones listed fourth, fifth, and seventh.
l.Ascus developed directly from the zygote
(or apomictically from an unfertilized cell);
not producing fruits Order 1. Endomycetalea.
l.The zygote giving rise to filaments of cells
with more than one nucleus, these producing
the asci.
2. Producing fruits.
3. The fruits cleistothecia.
4. Asci scattered in the fruits;
mostly saprophytes with
branched conidiophores Order 2. Mucedines.
4. Asci in one cluster, or solitary,
in the fruits; mostly parasites
with unbranched conidiophores Order 3. Perisporiacea.
3. The fruits, originally closed, open-
ing by irregular pores or regular or
irregular clefts Order 4. Phacidialea.
3. The fruits apothecia Order 5. Cupulata.
3. The fruits perithecia.
4. Producing a normal mycelium Order 7. Sclerocarpa.
4. Parasitic on insects, the mycel-
ium reduced Order 8. Laboulbenialea.
2. Not producing fruits, the asci arising di-
rectly from the mycelium Order 6. Exoascalea.
Order 1. Endomycetalea [Endomycetales] Gaumann Vergl. Morph. Pilze 135
(1926).
Subclass Hemiasci Engler Syllab. 26 (1892).
ILuttrell (1951) has presented a complete reorganization of the class. He sets apart
as a major subordinate group Bitunicatae five orders in which the ripe ascus exudes
3 vesicle and discharges the spores from this.
130 ] The Classification of Lower Organisms
Subclass Hemiasci or Hemiasceae, with suborder (Unterreihe) Hemiascineae,
and suborder Protoascineae of subclass Euasci, Engler in Engler and Prantl
Nat. Pflanzelfam. I Teil, Abt. 1 : iv (1897), the names not based on those of
genera.
Order Protoascineae Campbell Univ. Textb. Bot. 165 (1902).
Order Hemiascalcs Engler Syllab. ed. 3: 28 (1903).
Ascomycetes whose asci develop directly from the zygotes. Two families may be
recognized.
Family 1. Endomycetacea [Endomycetaceae] Schroter in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1: 154 (1894). Family Ascoideaceae Schroter op. cit. 145.
Mostly saprophytes, the uninucleate or multinucleate cells of the filaments tending
to round up, become separate, and function as conidia; the zygotes, produced by
syngamy of scarcely differentiated cells, enlarging and becoming asci of 4, 8, or
many spores cut out by free cell formation. Dipodascus, Eremascus, Endomyces,
Ascoidea. The asci of the last are apparently produced asexually (Walker, 1935).
The genus Protomyces requires mention. It is a parasite on higher plants, producing
walled resting spores which germinate by producing a sporangium of many spores.
It is chytrid-like, but its spores are non-motile. Its proper place in classification has
for a long time been a puzzle.
Family 2. Saccharomycetacea [Saccharomycetaceae] (Rees) Schroter op. cit. 153.
CXa^?, znd iarm\y Saccharomycetes y<! inter m Rabenhorst Kryptog.-Fl. Deutschland 1,
Abt. 1: (1879). Unicellular, reproducing by budding, i.e., by production upon
the cells of outpocketings which are pinched off as additional cells, or by a sexual
cycle in which endospores are produced, usually by fours.
These are the organisms which are in English called yeasts. The common bread-
and beer-yeast called Sac char omyces cerevisiae has a good claim to be considered,
economically, the most important of all "fungi." Its metabolism, in which dextrose
is converted to alcohol and carbon dioxide, gives a superficial appearance of sim-
plicity, and has attracted much study, contributing much to an understanding of the
genuine intricacy of energesis.
In addition to agents of fermentation, this family includes pathogens causing
chronic infections of animals. These have been treated as a genus Torula, Torulopsis,
Blastodenna, or Cryptococcus. They have not been observed to produce endospores.
Order 2. Mucedines Fries Syst. Myc. 3: 380 (1832).
Order Gyjnnoascaceae Winter in Rabenhorst Kryptog-Fl. Deutschland 1, Abt.
2: 3 (1887).
Suborder Plectascineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1: V (1897).
Order Plectascineae Campbell Univ. Textb. Bot. 169 (1902).
Order Aspergilliales Bessey in Univ. Nebraska Studies 7: 304 (1907).
Order Gymnascales Clemens Gen. Fung. 93 (1909).
Order Plcctascales Gaumann Vergl. Morph. Pilze 164 (1926).
Ascomycetes producing cleistothecia in which the asci are scattered; mostly
saprophytic and producing branched conidiophores.
The name Mucedines means molds. Under this name Fries listed twelve genera,
with Aspergillus Link and Penicillium Link first. The former is the evident standard
genus of the order. Both genera are very common and numerous in species. They arc
readily recognized under the microscope by the forms of their clusters of conidia.
Phylum Inophyta [131
The conidiophore of Aspergillus ends in a globular swelling from which spring many
radiating rows of conidia, with the efTect that the entire mass, yellow, brown, black,
pink, or red in color, is globular. Penicillium has a branching conidiophore bearing
rows of conidia in a broom-like mass. The masses are usually blue or green, and are
familiar on cheese, jam, bread, cardboard, oranges, or almost any organic material.
Particular species of Penicilliuni are involved in the making of genuine Camem-
bert and Roquefort cheeses. The genus has become best known for the production by
P. notatum of the drug penicillin. In 1929, Dr. Alexander Fleming of London noticed
that a mycelium of this species, growing as a contaminant on a plate of bacteria,
interfered with the growth of the latter. This observation led to the discovery of a
substance clinically useful against actinomycetes, spheres, and Gram positive rods,
but not against Gram negative rods. Production was for several years very scant, and
the drug expensive accordingly; in the early 1940's, as a war measure, the United
States financed large scale production along with the appropriate scientific study
(Elder, 1944; Committee on Medical Research, Washington, and the Medical Re-
search Council, London, 1945). Several forms of penicillin have been recognized;
they differ in the radicle R in the formula CgHnOiSNiR. The structural formula is
believed to be as follows (Editorial Board of the Monograph on the Chemistry of
Penicillin, 1947):
RCONH — CH — CH — S — C (CH3)2
I I I
OC N CH COOH.
The sexual reproduction of Aspergillus and Penicillium involves the syngamy
of differentiated cells. The zygote sends out ascogenous hyphae which bud oflF
scattered asci; the neighboring cells send out hyphae which become woven into a
minute firm-walled cleistothecium enclosing them.
Link, who named Aspergillus and Penicillium., gave to the ascocarp-producing
stage of Aspergillus the name Eurotium. There is a rule of botanical nomenclature
which allows only a tentative status to names given to the conidium-producing
stages of inophytes. Thom and his associates (1926, 1945), in presenting a workable
system of the species of Aspergillus, remarked that "It is better to forget Eurotium
along with the technicality."
This order includes a variety of other molds: Gymnoascus, producing only a
loose weft of hyphae about the asci; Ctenomyces, on feathers, recognized by comb-
like outgrowths from the loosely woven ascocarps; Monascus, its name a misnomer,
the minute fruit containing many asci; Onygena, saprophytic on horns and hoofs,
producing puffball-like fruits as much as 1 cm. high; Elaphomyces, forming a
mycorrhiza on roots of conifers and producing hypogaeous fruits as large as walnuts.
Order 3. Perisporiacea [Perisporiaceae] Fries Syst. Myc. 3: 220 (1829).
Order Perisporia Fries op. cit. 1: xlviii (1832).
Suborder Perisporiaceae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1,
Abt. 2: 21 (1887).
Subsuborder [Underordnung) Perisporiales Engler in Engler and Prantl Nat.
Pflanzenfam. I Teil, 1: v'(1897).
Order Perisporiales Bessey in Univ. Nebraska Studies 7: 295 (1907).
Ascomycetes producing cleistothecia containing a compact cluster of asci or a
solitary ascus; mostly parasites producing unbranched conidiophores.
132]
The Classification of Lower Organisms
Fig. 25. — Ascomycetes: a-e, Dipodascus albidus after Juel (1902), x 1,000;
a, gametes; b, syngamy; c, development of ascus; d, e, lower and upper parts of a
mature ascus. f, Erysiphe graminis, haustorium penetrating an epidermal cell of
a grass and conidiophore bearing a chain of conidia x 500. g-k, Cleistothecia of
Perisporiacea x 100: g, of Erysiphe sp.; h, of Microsphaera sp.; i, of Podosphaera sp.;
j, of Uncinula sp.; k, of Phyllactinia sp.
Phylum Inophyta [ 133
The more familiar Perisporiacea are those of family Erysiphea [Erysipheae]
Winter. They are parasites on plants, mostly producing a white mycelium on the
surface and sending brief haustoria into the epidermal cells. They produce abundant
conidia in erect unbranched chains; this habit explains the common name of powdery
mildews. Harper's important studies of the morphology of Ascomycetes were in large
part made on powdery mildews. The gametes are uninucleate and unite directly, the
egg bearing no trichogyne; the ascogenous hyphae are brief; each minute black
globular cleistothecium bears an equatorial whorl of appendages of a form charac-
teristic of the genus. In Erysiphe and Sphaerotheca iS. pannosa is the common rose
mildew), the fruits bear unbranched sinuous appendages like vegetative hyphae; the
fruit of Erysiphe contains several asci, while that of Sphaerotheca contains one. In
Microsphaera {M. Alni is the powdery mildew of lilac) and Podosphaera, the ap-
pendages are dichotomously forked near the tip; the fruit of Microsphaera contains
several asci, that of Podosphaera only one. The appendages of Uncinula are hooked
at the tip. Those of Phyllactinia are like sharp spikes with bulbous bases.
Other Perisporiacea, parasitic or saprophytic on plant material, are compara-
tively poorly known. The fruits may bear appendages of other characters than those
of the Erysiphea, or none, and may be characteristically clustered or borne in
stromata. In some examples the fruits have no definite dehiscence mechanism; in
others they open by deliquescence or by a separation of plates. Some open by a
single pore, and appear transitional to those of order Sclerocarpa; some open by a
cleft, or by lobes separated by radiating clefts, and appear transitional to those of
order Phacidialea.
Order 4. Phacidialea [Phacidiales] Bessey in Univ. Nebraska Studies 7: 298
(1907).
Phacidiacei Fries Syst. Myc. 1: li (1832).
Order Hysteriaceae and suborders (of order Discomycetes) Phacidiaceae,
Stictideae, and Tryblidieae Rehm in Rabenhorst Kryptog.-Fl. Deutsch-
land 1, Abt. 3: 1, 60, 112, 191 (1896); the ordinal name preoccupied by
family Hysteriaceae Saccardo.
Suborders Phacidiineae and Hysteriineae Engier in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1 : v ( 1897) .
Orders Graphidiales and Hysteriales Bessey op. cit. 298, 303.
Order Hemisphaeriales Theissen in Ann. Myc. 11: 468 (1913).
Order Microthyriales Clements and Shear Gen. Fung. ed. 2: 94 (1931),
Ascomycetes producing fruits which are not typical cleistothecia, apothecia, or
perithecia.
This group is here used as a catch-all for three or more distinct groups, which
appear to form cross-connections among orders Perisporiacea, Cupulata, and Sclero-
carpa. This appearance suggests the probability that the present group, and the
usually accepted orders assembled under it, are not natural, but represent parallel
developments from several sources. The present groups include moderately numerous
ordinary parasites and saprophytes, together with great numbers of lichen-formers.
Only the latter are common and familiar in temperate countries. There has been little
study of the morphology.
The families which appear tenable are distinguished as follows:
a. Fruits minute and flattened, usually releasing the spores through one or more
pores or clefts (Order Hemisphaeriales Theissen, Microthyriales Clements
and Shear).
134] The Classification of Lower Organisms
Family Microthyriacea [Microthyriaceae] Lindau (in Engler and Prantl, 1897).
Parasitic on plants, surfaces of the fruits marked by radiating ridges.
Family Micropeltidacea [Micropeltidaceae] Clements and Shear (1931). Family
Hemisphaeriaceae Theissen (1913), not based on a generic name. Like the fore-
going, but the surface of the fruit not radiate or radiate only at the margin.
Family Trichothyriacea [Trichothyriaceae] Theissen and Sydow. Parasitic on
inophytes, the mycelium a pseudoparenchymatous layer, asci pendant within the
fruits from the apparent summit.
b. Fruits elongate, hard, dark, opening by a narrow cleft (suborder Hysteruneae
Engler).
Family Hysteriacea [Hysteriaceae] Saccardo Sylloge 2: 721 (1883). Parasitic on
higher plants or saprophytic.
Family Graphidiacea [Graphidiaceae] Clements (1909). An enormous group of
lichens or parasites on lichens, largely tropical and chiefly crustose, the openings
of the fruits forming dark lines.
c. Fruits not as above, mostly with a roundish area of asci exposed by the
irregular or stellate shattering of a superficial layer; if long and narrow, not
hard and dark (Suborder PHACiDnNEAE Engler).
Family Phacidiea [Phacidieae] Saccardo Sylloge 8: 705 (1889). Phacidiaceae
Saccardo (1889). Family Phacidiaceae Lindau (in Engler and Prantl, 1896). The
dark fruits thin and weak laterally and below.
Family Tryblidacea [Tryblidaceae] Rehm (in Rabenhorst, 1896). The dark fruits
hard and thick laterally and below.
Family Stictea [Sticteae] Saccardo Sylloge 8: 647 (1889). Stictaceae Saccardo
(1889). Family Stictidaceae Lindau (1896). Fruits light-colored or white. Higgins
(1914) found that the agents of the shot-hole disease of plums and cherries, which,
on the basis of non-fruiting stages, have been called Cylindrosporium Pruni, produce
on fallen leaves ascocarps distinguishable as three species of the genus Coccomyces
of the present family.
Order 5. Cupulata [Cupulati] Fries Syst. Myc. 1 : 2 (1821 ).
Order Mitrati Fries 1. c; order Uterini Fries op. cit. 1 : liii (1832).
Family Discomycetes Fries Epicrisis 1 (1836).
Orders Discomycetes and Tuberaceae Winter in Rabenhorst Kryptog.-Fl.
Deutschland 1, Abt. 2: 3 (1887).
Suborders Helevellincae, Pczizineae, and Tuherineae Engler in Engler and
Prantl Nat. Pflanzenfam. I Teil, Abt. 1: v (1897).
Orders Helevellincae, Pezizineae, and Tuberineae Campbell Univ. Textb. Bot.
166, 167, 168 (1902).
Orders Pezizales, Discolichcnes, Helvellales, and Tuberales Bessey in Univ.
Nebraska Studies 7: 299, 300, 303, 304 (1907).
This order includes primarily the cup fungi, the inophytes which produce cup-
or disk-shaped fruits bearing a single hiyer of closely packed asci on the inner or
upper surface. There has been much study of some of them, notably of Pyronema, by
Harper, Dangeard, Claussen, and Brown. The disk-shaped flesh-colored apothecia of
Pyronema, 1-3 mm. in diameter, are found particularly on damp charcoal. The
mycelium produces difTcrentiatcd multinucleate antheridia and ascogonia, the latter
bearing one-celled multinucleate trichogynes. After syngamy, or sometimes without
it, but always to the best of our knowledge without any fusion of nuclei, the ascogonia
Phylum Inophyta [ 135
send out branching filaments which become septate in such fashion that the ultimate
cells are binucleate. These cells form croziers and produce asci. During the develop-
ment of the ascogenous hyphae, other hyphae, more slender, grow up from the
vegetative mycelium; these produce a disk of undifferentiated cells below the layer
of asci, and send up sterile hairs (paraphyses) among them.
Gaumann (1926) divided the families of this group into two series by the
presence or absence of a differentiated operculum at the summit of the ascus. The
names being put into neuter form, and family Tuberacea being added, the lists are
as follows:
Inoperculata : Patellariacea, Dermateacea, Bulgariacea, Cyttariacea, Mollisi-
acea, Helotiacea, Geoglossacea, Tuberacea.
Operculata: Rhizinacea, Pyronemacea, Ascobolacea, Fezizacea, Helvellacea.
Along with these, Clements and Shear (1931) list eight families of lichen-formers,
some of them very numerous.
Families Pezizacea and Ascobolacea include the ordinary cup fungi. They are
mostly saprophytes in soil or on manure, and do not usually produce conidia. Peziza
was listed by Fries first in order Cupulata; it is the evident standard genus of the order.
Families Dermateacea and Helotiacea include many parasites on plants. One of
the Helotiacea is Sclerotinia cinerea, the agent of the brown rot of stone fruits.
As an active parasite it produces conidia of a type which, if the fruits were unknown,
would place it in the genus Monilia. These spread the disease rapidly. The killed
fruits fall and the organism lives in them as a saprophyte, replacing their tissues
with a hard black mass of hyphae, a sclerotium. This survives the winter and in
spring sends up stalked white apothecia.
The Helvellacea have been treated as a separate order, but are not sufficiently
numerous and distinct to justify this treatment. They are saprophytes in soil, pro-
ducing large stalked apothecia bearing an extensive layer of asci which is everted
and wrinkled. The most familiar genera are Elvella and Morchella. The fruits are
edible, indeed delicious; they should be boiled briefly, then creamed and served on
toast. When found in abundance they should be preserved by drying for use through-
out the year.
The Tuberacea, the truffles, also usually treated as a distinct order, produce
underground fruits which appear to be apothecia distorted and rolled into balls.
They are associated with particular species of trees on which the mycelia are be-
lieved to live as mycorhizae (Dangeard, 1894). The asci commonly contain reduced
numbers of spores. The fruits are prized by gourmands.
The relationships of the Cupulata are a puzzle. Pyronema could be interpreted
as representing an evolutionary transition from the order Mucedines to this. Certain
parasitic cup fungi produce minute apothecia, hard, dark, and nearly closed, sug-
gesting a transition to order Sclerocarpa. Some species, particularly among the
parasites and lichen-formers, seem to intergrade with order Phacidialea, and thence
again both to Mucedines and Sclerocarpa. The operculate asci which mark a part of
the group occur also in other orders. Thus there is among Ascomycetes an appearance
of reticulate relationships, such as reputable naturalists of the past supposed to
exist in many groups. The appearance is of course illusory; sufficient study of other
groups has made it possible to distinguish the resemblances among them which indi-
cate relationship from those which are results of parallel evolution. The study of the
Ascomycetes has not yet been carried this far.
136]
The Classification of Lower Organisms
Fig. 26. — Ascomycetes: a-k, Lachnca scutellata after Brown (1911) x 1,000;
a, b, formation of crozicr; C, karyogamy; d, fusion nucleus; e-i, stages of mciosis;
j, k, early stages of free cell formation. 1, Apothecia x 2, and m, ascus x 250, of
Lamprospora leiocarpa. n, Apothecia x 2, and O, ascus x 250, of Aleuria rutilans.
V, Apothecia of Sclcrotinia cinerea x 2. q, Fruit x 1, and r, ascus x 250, of Mor-
chella conica. s-x, Taphrina deformans after Martin (1940) x 1,000; s, growth on
surface of an infected leaf; t, karyogamy; u, mitosis; V, homeotypic anaphase in the
ascus; w^ development of ascospores; x^ germination.
Phylum Inophyta [137
Order 6. Exoascalea [Exoascales] Bessey in Univ. Nebraska Studies 7 : 305 ( 1907) .
Suborder Protodiscineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1: V (1897), not based on a generic name.
Order Protodiscineae Campbell Univ. Textb. Bot. 166 (1902).
Order Agyriales Clements and Shear Gen. Fung. ed. 2: 141 (1931), in part.
Ascomycetes parasitic on plants, producing no fruits but a broad layer of asci
directly on the mycelium.
The leaves of the hosts of these parasites become swollen and distorted; the
diseases recognized by these symptoms are called curly-leaf diseases. The most
familiar is the curly-leaf of peaches, caused by Taphrina {Exoascus) deformans.
Many others are known. The agents of all of these diseases may be regarded as a
single family Exoascacea [Exoascaceae] Schroter (in Engler and Prantl, 1894), and
all are commonly treated as a single genus, Taphrina Fries, typified by T. aurea on
poplar trees; there are differences among them which might well be treated as of
generic rank.
Clements and Shear associated the curly-leaf parasites with a collection of sapro-
phytes producing small and undifferentiated disk-like or indefinite fruits, as Pyro-
nema, Ascocorticium, and Agyrium; and offended against the principles of nomen-
clature by re-naming the order Agyriales. It is probable that something of the nature
of Agyrium may represent the transition from order Cupulata to this one.
Martin (1940) described the cytology of Taphrina deformans. The mycelium
grows between the cells of the host, not penetrating them. It is a dikaryophase
mycelium, the cells binucleate, the nuclei dividing concurrently, cell division occur-
ring in such fashion as to separate the daughter nuclei of each pair. In preparation for
reproduction, hyphae of short round cells form a single layer between the epidermis
and the cuticle of the host. In each cell of these hyphae, the nuclei unite and then
divide. The division is mitotic, the fusion nuclei and the daughter nuclei having
each eight chromosomes. The cell divides, by a wall parallel to the surface of the
leaf, into two. The daughter cell which lies against the tissues of the host dies,
and its wall becomes empty; the other cell grows and bursts through the cuticle of
the host and becomes an ascus. Its nucleus divides three times; the first two divisions
are the meiotic process, and the chromosome number is reduced to four. Cytoplasm
accumulates around each of the resulting eight nuclei and is presently cut out by a
membrane and a wall. No centrosome is evident at any stage of the process. The
spores germinate by sending out buds, as yeasts form buds; sometimes they do this
before being discharged from the ascus. So far as Martin could determine, the
binucleate condition of the mycelium is established by division of the nucleus of
the spore from which it grows.
Order 7. Sclerocarpa [SclerocarpiJ Persoon Syst. Meth. Fung, xii (1801).
Order Pyrenomycetes Fries Syst. Myc. 2:312 (1822); order Uterini, suborder
Pyrenomycetes Fries op. cit. 1: li (1832).
Family Pyrenomycetes Fries Epicrisis 1 (1836).
Order Pyrenomycetes, suborders Hypocreaceae, Sphaeriaceae, and Dothideaceae,
Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 2: 18, 82, 152, 893
(1887).
Suborder {Unterreihe) Pyrenomycetineae, sub-suborders [Unterordnungen)
Hypocreales, Dothideales, and Sphaeriales Engler in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1: v, vi (1897).
138
The Classification of Lower Organis?ns
Order Pyrcnomycetales Bessey in Univ. Nebraska Studies 7: 295 (1907).
Orders Hypocreales, Sphaeriales, and Dothideales Gaumann Vergl. Morph.
Pilze 222, 253, 284 (1926).
Ascomycetes producing, from a normal mycelium, perithecia, i. e., small fruits
of the shape of a small globe or flask opening through a single pore, the ostiole.
Sphaeria, which Persoon and Fries listed first under the names which they respec-
tively used, is the evident standard genus; but this genus has become broken up and
lost in the work of subsequent scholars.
This order includes very many species and is by the generality of authority
divided into three. Forms whose perithecia are borne directly on the m.ycelium,
together with those whose perithecia are borne in or on but distinct from a dark
Fig. 27. — Mycosphaerella personata after Higgins (1929), x 1,000; a, conidio-
phorcs and conidia of Ccrcospora type, b, longitudinal section of pycnidium;
C, primordium of perithccium with ascogonoium bearing a trichogyne; d, e, ascogen-
ous hyphae; f, crozier formation; g, longitudinal section of mature peridiecium.
Phylum Iiiophyta [ 139
stroma, are assigned to order Sphaeriales. Forms with perithecia in or on a brightly
colored stroma are Hypocreales. Those whose perithecia are cavities with a wall
indistinguishable from a dark stroma are Dothideales. These groups are not confi-
dently acceptable as natural: the stromatic Sphaeriales (Wehmeyer, 1926), the
Hypocreales, and the Dothideales appear each to include more than one line of
descent from Sphaeriales with solitary perithecia.
As a general rule, each perithecium develops in consequence of a separate act of
fertilization, of a differentiated ascogonium, either by an antheridium, a spermatium,
or otherwise.
Gaumann recognized fourteen families in the present group or groups. To these
are to be added a great number of lichen-formers, properly Sphaeriales and Hypocre-
ales,. but construed as a single family Verrucariacea; and a smaller number, repre-
senting the Dothideales, and called Mycoporacea.
Exmples include the following:
Among Sphaeriales with solitary perithecia, Mycosphaerella is a genus of more
than one thousand parasites on plants, mostly inconspicuous, causing leaf spots.
Their conidia are of various types, Septoria, Phleospora, Ramularia, Cercospora.
Venturia, another numerous genus, includes V. inaequalis, causing apple scab; its
conidia are of a type called Fusicladium.
Four species of Neurospora were discovered by Shear and Dodge (1927) as the
fruiting stages of a red mold on bread called Monilia sitophila. Genetic study of
this genus particularly by Tatum, Beadle, and their associates (Ryan, Beadle, and
Tatum, 1943; McClintock, 1945; Beadle and Tatum, 1945; Tatum and Bell, 1946;
Mitchell and Houlahan, 1946; Tatum, Barratt, Fries, and Bonner, 1950) has yielded
results of the highest theoretical significance. Normal cultures require no other
food than minerals, carbohydrate, and a single vitamin, biotin (Butler, Robbins,
and Dodge, 1941). Either spontaneously or under violent treatment (with x-rays,
ultra-violet radiations, or mustard gas) the cultures give rise to many mutations,
behaving as Mendelian recessives, each consisting of the inability to synthesize
some one vitamin or amino acid. These observations mean that life in its aspect of
metabolism consists of unit chemical processes, each controlled by a specific enzyme,
each enzyme being dependent upon a specific area in a specific chromosome.
Among stromatic Sphaeriales, Glomerella, with conidial stages identified as
Gloeosporium or Colletotrichum, attacks many plants; G. cingulata causes the bitter
rot of apples. Valsa, Diatrype, and Diaporthe are numerous in species. Endothia
parasitica causes the chestnut blight, destructive in the eastern United States.
Xylaria, Daldinia, and other genera are saprophytic on wood; the former produces
black fruits, club-shaped or branched; the latter, fruits of the form of black knobs
which may reach the size of golf balls.
Among Hypocreales, Nectria cinnabarina is common as a saprophyte on dead
twigs of poplar. It produces small wart-like red stromata which bear first conidia,
then perithecia. Claviceps purpurea causes a disease of rye; it produces conidia of
various types, and converts the grains of rye into sclerotia. These bodies are called
ergot; they are extremely poisonous, sometimes dangerously so, because they may
be ground with the grain. They are used in medicine. After lying in the earth through
the winter, the sclerotia send up fruits of the form of a stalk bearing a knob consisting
of radiating perithecia. Cordyceps kills subterranean larvae or pupae of insects and
then sends up a stalk bearing an elongate head of many perithecia.
140 ] The Classification of Lower Organisms
The Dothideales include Plowrightia morhosa, the agent of the black knot of
plums. Diseased twigs become swollen and covered with a black stroma which bears,
according to the season, conidia of various types or else perithecia.
Order 8. Laboulbenialea [Laboulbeniales] Engler Syllab. ed. 3: 42 (1903).
Order Laboulbeniaceae Thaxter, the name (ascribed to Peyritsch) preoccupied
by family Laboulbeniaceae Berlese in Saccardo Sylloge 8: 909 (1889).
Suborder Laboulbeniineae Engler in Engler and Prantl Nat. Pflanzenfam.
I Teil, Abt. 1: vi (1897).
Class LabouJbeniomycctes Engler Syllab. 1. c.
Class Laboulbenieae Schaffner in Ohio Naturalist 9: 450 (1909).
Parasites on insects, the mycelium scant or reduced to a single cell, producing
antheridia which discharge spermatia into the air and small numbers of perithecia.
These organisms have the appearance of excep^^ional setae on their hosts, which
are not usually seriously injured by them. They were first mentioned in a note by
the entomologist Rouget, 1850; Montagne and Robin, in Robin's book on parasitic
plants, 1853, gave the first names, Laboulbenia Rougetii and L. Guerinii, the generic
name honoring the entomologist Laboulbene. Only a few scholars, notably Thaxter
(1896, 1908, 1924, 1926, 1931) have given much attention to this group; they have
distinguished well over a thousand species, forming three families and about fifty
genera.
Many Laboulbenialea occur as two forms, male and hermaphrodite. A male indi-
vidual produces a series of flask-shaped antheridia, each of which discharges into
the air, one at a time, a series of globular naked sperms. A hermaphrodite individual
produces first a series of antheridia as described and then one or more perithecia.
A perithecium consists of a wall, of a definite number of cells produced in definite
order and pattern, surrounding an egg which bears a trichogyne; the trichogyne
protrudes from the perithecium and receives the sperms. The zygote gives rise to a
fascicle of asci which crowd aside and destroy the inner cells of the wall and dis-
charge the ascospores (usually eight in the ascus, and divided into two cells) through
the ostiole.
Those who would link the Ascomycetes with the red algae entertain the hypothesis
that the Laboulbenialea represent the transition. This hypothesis is surely mistaken.
The Laboulbenialea are a highly specialized group, not a link between others. They
appear to have evolved from Sphaeriales with solitary perithecia.
Class 3. HYPHOMYCETES Fries
Classes Hyphomycetes and Coniomycctes Fries Syst. Myc. 3: 261, 455 (1832).
Families Hyphomycetes and Coniomycctes Fries Epicrisis 1 (1836).
Fungi imperfecti or Deuteromycetes Auctt.
Inophyta of which the structures involved in sexual reproduction are unknown.
It has been noted that a particular genus of Ascomycetes may produce conidia
of more types than one, as Sclcrotinia produces types called Monilia and Botrytis,
and Glomerella produces types called Gloeosporium. and Colletotrichum. The same
type may be produced by many genera; the Monilia type recurs in Neurospora,
which does not belong to the same order as Sclerotinia. Collecting naturalists, and
plant pathologists in the pursuit of their duties, are constantly encountering conidial
stages whose assignment to an order of Ascomycetes is impossible. It is an obvious
Phylum Inophyta [ 141
practical necessity that a register of these observations be kept. The register is pro-
vided by the present group, one which is named, defined, and assigned to the category
of classes, and divided into named orders, families, and genera under which specimens
may be identified as of species old or new. Class, orders, families, and genera are
known not to be valid taxonomic groups; many of the ostensible species are known,
and most of the rest are believed, to be stages of organisms which would in other
stages have other names. Almost all of them are Ascomycetes; Zygomycetes and
Basidiomycetes do not usually occur in unidentifiable stages.
The ascus-bearing stages are constantly being discovered. When this happens, the
species is re-named in its proper place among Ascomycetes. Theoretically, it loses
its place in the list of imperfect fungi; practically, it retains it, because the next
collector or plant pathologst is most likely to try to find it there.
The system of Hyphomycetes is as follows:
Order 1. Phomatalea [Phomatales] Clements Gen. Fung. 121 (1909).
Sphaeropsideae Saccardo Sylloge 8: xvi (1889).
Order Sphaeropsidales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil
Abt. 1**: v (1900), not based on a generic name.
Order Phomales Clements and Shear Gen. Fung. ed. 2: 175 (1931).
Producing pycnidia. The four families correspond with as many groups of Ascomy-
cetes.
Family 1. Phomatacea [Phomataceae] Clements Gen. Fung. 121 (1909). Family
Sphaerioideae or Sphaerioidaceae Saccardo; but Sphaeria belongs to order Sclero-
carpa. Family Phomaceae Clements and Shear (1931). Pycnidia hard and black as
in Sphaeriales and Dothideales. Phoma, Ascochyta, Diplodia, Septoria, each of
many species.
Family 2. Zythiacea [Zythiaceae] Clements Gen. Fung. 128 (1909). Family
Nectrioideae or Nectrioidaceae Saccardo; but Nectria belongs to order Sclerocarpa.
Pycnidia in brightly colored stromata as of Hysteriales.
Family 3. Leptostromatacea [Leptostromataceae] Saccardo Sylloge 3: 625 (1884).
Pycnidia in shield-like stromata, like the fruits of Microthyriacea.
Family 4. Discellacea [Discellaceae] Clements and Shear Gen. Fung. ed. 2: 192
(1931). Family Excipulaceae Saccardo; but Excipula is a cup fungus. Pycnidia wide
open like the fruits of Phacidiea.
Order 2. Melanconialea [Melanconiales] Engler in Engler and Prantl Nat. Pflan-
zenfam. I Teil, Abt. 1**: v (1900).
The conidia borne on a stroma but not in pycnidia.
Family Melanconiacea [Melanconiaceae] (Saccardo, without category) Lindau
in Engler and Prantl op. cit. 398, the single very numerous family: Gloeosporium;
Coryneum, C. Beijerinckii, the shot-hole of almonds; Pestallozia.
Order 3. Nematothecia [Nematothecii] Persoon Synops. Meth. Fung, xix (1801).
Orders Dematiei, Sepedoniei, Tubercularini, and Stilhosporei Fries Syst. Myc. Order
Hyphomycetes (Fries) Auctt. Order Moniliales Clements Gen. Fung. 138 (1909).
Conidia directly on the mycelium, or none.
Family 1. Tuberculariea [Tubercularieae] Saccardo Sylloge 4: 635 (1886).
Tuberculariaceae Saccardo (1889). Family Tuherculariaceae Lindau (1900).
Scarcely distinct from Melanconiacea, the conidia on a mass of interwoven hyphae
142 ] The Classification of Lower Organisms
less compact than a stroma. Fusarium, an enormous number of species producing as
conidia crescent-shaped rows of cells. Snyder and Hansen (1941, 1945) find that
the fruiting stages are species of Hypomyces, Nectria, Gibberella, or Calonectria, all
Hypocreales.
Family 2. Stilbellacea [Stilbellaceae] Bessey Morph. and Tax. Fungi 584 (1950).
Family .Siz/^e'a^ Saccardo Sylloge 4: 563 ( 1886). .S^z/foacfflP Saccardo ( 1889). Family
Stilbaceae Lindau (1900); Bessey observed that the type of the genus Stilbum does
not belong to this family. Mostly molds producing coremia.
Family 3. Dematiea [Dematieae] Saccardo Sylloge 4: 235 (1886). Dematiaceae
Saccardo (1889). Family Dematiaceae Lindau (1900). Dark-colored parasites, as
Helminthosporium, Cladosporium, and Cercospora, or molds, as Alternaria.
Family 4. Moniliacea [Moniliaceae] Clements Gen. Fung. 138 (1909). Mucedineae
Persoon, family Mucedineae or Mucedinaceae Saccardo, not based on a generic
name. White or brightly colored parasites or molds, as Oidium, with colorless spores
in chains, Monilia, Botrytis, etc. The parasites on animals which have been referred
to Monilia are currently called Candida.
Family (?') 5. Sterile mycelia. Many mycorhizae must be left here. Rhizoctonia,
dark net-like masses of hyphae occurring as parasites or saprophytes. Trichophyton,
parasitic on the skins of man and animals, causing ringworm, athlete's foot, etc.
Class 4. BASSDIOMYCETES (Sachs ex Bennett and Thistleton-Dyer)
Winter
Order Basidiosporeae and subordinate group Basidiomycetae Cohn in Hedwigia
11: 17 (1872).
Basidiomyceten Sachs Lehrb. Bot. ed. 4: 249 (1874).
Basidiomycetes Bennett and Thistleton-Dyer in Sachs Textb. Bot. English ed.
847 (1875).
Class Basidiomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt.
1: 72 (1884).
Classes Teliosporeae and Basidiosporeae Bessey in Univ. Nebraska Studies 7 : 305,
306 (1907).
Classes Teliosporeae and Basidiomycetae Schaffner in Ohio Naturalist 9 : 450
(1909).
Inophyta which produce, as a feature of the sexual cycle, conidiophores called
basidia, each producing typically four conidia called basidiospores.
Germinating basidiospores give rise to mycelia of cells with solitary haploid
nuclei. Syngamy occurs among cells of these mycelia, usually simply by contact of
vmdifTerontiated cells; the rusts produce differentiated sperms in spermagonia re-
sembling the pycnidia of Ascomycetes. In some species any haploid hypha may
conjugate with any; in some there are two mating types, and in some four. Raper
(1953) has studied the interesting genetics of the mating types.
The cell produced by .syngamy remains undifferentiated, but gives rise, by con-
current division of its nuclei, to a dikaryote mycelium. The nuclei are minute, and
mitosis has rarely been seen. The nuclear divisions are often followed by a peculiar
manner of cell division, comparable to the crozier formation of Ascomycetes, and
producing structures called clamp connections.
Either the original haploid mycelium or the dikaryophase may produce conidia
without nuclear change. Such reproduction is familiar among the rusts, rather un-
familiar among other Basidiomycetes.
Phylum Inophyta [ 143
Only the dikaryophase produces the specialized conidiophores called basidia,
which are regularly the seat of karyogamy and meiosis. There is a considerable
variety of types of basidia. Van Tieghem (1893) originated the terminology ap-
plicable to these; Martin (1938) has attempted to refine it, and Linder (1940) to
simplify it.
Frequently, the seat of meiosis is a thick-walled resting spore or an otherwise
difTerentiated cell called a probasidium, upon which the proper basidium develops,
after meiosis, as an outgrowth. A basidium arising in this fashion is commonly
elongate and divided into four cells each of which produces a basidiospore. Such a
hypha-like basidium may be called a promycelium or a phragmobasidium; the latter
term is applicable also to an elongate four-celled basidium which does not arise
from a probasidium. In a few Basidiomycetes, the basidium is divided into four cells
by longitudinal walls; such basidia are called cruciate basidia. In the familiar
Basidiomycetes the basidium does not become divided by walls and is called a holo-
basidium or autobasidium. Gaumann ( 1926) distinguished two types of holobasidia:
the stichobasidium, in which the spindles of the dividing nuclei lie at various levels
and in various directions, and which frequently produces more than four nuclei;
and the chiastobasidium, in which the spindles lie transversely near the summit,
and which regularly produces just four nuclei. Dodge, translating Gaumann (1928),
denies much importance to this distinction.
The meiotic divisions have repeatedly been studied. Apparent centrosomes have
been seen at the poles of the spindles (Lewis, 1906; Lander, 1933), but not by most
microtechnical methods (Savile, 1939; Ritchie, 1941). The chromosomes gather as
usual at the middle of the spindle and divide. The nuclear membrane becomes in-
distinct, but the nuclear sap remains distinct from the cytoplasm nearly until the
completion of division; it then disappears, leaving the groups of daughter chromo-
somes connected by a spindle of the appearance of a dark streak in the cytoplasm.
Ob:;erved haploid chromosome numbers include the following:
Coleosporium, fideMoxczM (1914) 2
Coleosporium Vernoniae, fide Olive (1949) 8
Coj&rmuj, fide Yokes (1931) 4
Eocronartium, fide Fitzpatrick (1918) 4
&;frffa, fide Whelden (1935) 4
Gymnosporangium, fide Stevens (1930) 2
Melampsora, fide Savile (1939) 4
Myxomycidium flavum, fide Martin (1938) 8
Puccinia, fide Savile (1939) 4
Transchelia, fide Savile (1939) 4
Russula, fide Ritchie (1941) 4
Scleroderma, fide Lander (1933) 2
f/romycg'^, fide Savile (1939) 4
Savile suggests that some at least of the reports of a chromosome number of 2
may have resulted from misinterpreted observations of one pair of choromosomes
behind another.
Normally, only the two meiotic divisions, producing four nuclei, occur in the
basidium; exceptionally, there are further, mitotic, divisions, resulting in more than
four spores on the basidium. The basidiospores are usually borne on slender stalks
called sterigmata. Sterigmata and spores are formed by evagination of the wall of
the basidium; the nuclei migrate through the sterigmata into the spores.
144]
The Classification of Lower Organisms
Fig. 28. — Basidiomygetes : a. Two germinating basidiosporcs of Agariciis campcs-
tris produce mycelia which anastomose freely, the cells becoming plurinucleate,
after Hein ( 1930) , x 500. b, c. Young and older basidia of Cystobasidium sebaceum,
after Martin (1939). d-g, Eocronartium muscicola after P'itzpatrick (1918); d, fus-
ion nucleus; e, homeotypic division in the basidium; f, four-celled basidium; g, pro-
duction of basidiospore. h^ i^ i, Basidia of Ustilago Heujlcri, U. Hurdei, and Tille-
(Continued bottom p. 145)
Phylum Inophyta [ 145
Most basidia discharge the spores actively, to a distance of a fraction of a milli-
meter. Buller (1929) observed that just before a spore is cast off a minute droplet
of liquid appears at the summit of the sterigma. This occurs in precisely the same
fashion in mushrooms, rusts, certain smuts, and the yeast-like organism Sporoholo-
myces. Buller inferred that the force which discharges the spore is surface tension
in the droplet. The fruits of Basidiomycetes are evidently adapted to the feebleness
of the mechanism by which the spores are discharged. If the fruits are cup-like,
they open laterally or downward. The basidia of mushrooms stand horizontally on
gills which are commonly less than one millimeter apart, allowing the spores to
fall from between them without touching them.
The groups of Ascomycetes and Basidiomycetes are evidently related. Morels and
mushrooms, truffles and puffballs, taste alike. The technical scholar will be con-
vinced that the groups are related by the occurrence in both of a dikaryophase
stage, a character too strongly in contrast with those of the generality of organisms
to be a probable product of parallel evolution. Gaumann quotes an old opinion of
Vuillemin (1893), "qu'une baside est un asque dont chaque cellule-fille avant de
passer a I'etat de spore, fait saillie au dehors et se transforme en une sorte de conidie
pour mieux s'adapter au transport par la vent." In dealing with the Zygomycetes,
Gaumann emphasized the apparent evolution of conidia from endospores by evagina-
tion of the walls of the sporangia. Largely, as it seems, by Gaumann's influence,
Vuillemin's hypothesis has become generally accepted.
Gaumann was disposed to derive the Basidiomycetes from something like Asco-
cortkiiim, and began his account of several of the groups of Basidiomycetes with
forms having scant flat fruits, or having basidia which spring directly from the
substratum or host. Linder (1940) suggested a derivation from Cupulata or Sclero-
carpa having operculate asci. He took note that many such asci open by producing a
vescicle, bounded by the stretched inner wall of the ascus, into which the asco-
spores pass. This led to the conclusion that the Basidiomycetes producing probasidia
are the lowest, and to this extent his reasoning appears cogent. He went on to identify
the rusts as the lowest Basidiomycetes, which seems far-fetched, the rusts being
distinctly a specialized group.
The generally accepted groups of Basidiomycetes are those which were set forth
by Engler (1897, 1900), as follows:
Subclass HEMiBAsron, having basidia bearing indefinite numbers of spores; the
smuts.
Subclass EuBAsron, the basidia bearing definite numbers of spores.
Order {Reihe) Protobasidigmycetes, the basidia divided into cells.
Suborder {Unterreihe or Ordnung) ^uricularhneae, the basidia divided
by transverse walls.
Sub-suborder [Unter ordnung) Uredinales, the rusts.
tia Tritici, after Sartoris ( 1924) . k, 1, Basidia of Patouillardina cinerea after Martin
(1935). m, Basidium of Sebacina sublilacina after Martin (1934). n, Basidium
of Protodontia Uda after Martin ( 1932). o, p, younger and older basidia of Tulas-
nella phaerospora, after Martin (1939). q-t, Development of the basidium of
Guepinia Spathularia, after Bodman (1938). u-x, Russula emetica after Ritchie
(1941); binucleate primordium of basidium, fusion nucleus, homeotypic division,
development of basidiospores. y, z, Basidia of Lycogalopsis Solmsii after Martin
(1939). X 1,000 except as noted.
146 ] The Classification of Lower Organisms
Sub-suborder Auriculariales.
Suborder Tremellineae, the basidia divided by longitudinal walls.
(At this point should appear Reihe Autobasidiomycetes, to include eight
Unterreihen of ordinary Basidiomycetes. The name Autobasidiomycetes
does not appear in the table of contents, the text, or the index of the
Natilrlichen Pflanzenfamilien; it was published in Engler's Syllabus, 1892).
Rearranging these groups according to current opinion, and suppressing the sub-
sidiary categories, one arrives at the following system of orders:
1. Producing probasidia or transversely divided
basidia, usually both.
2. Probasidia, if formed, terminal on the
hyphae.
3. Mostly saprophytic and producing
gelatinous fruits Order 1. Protobasidiomycetes,
3. Parasitic, mostly not producing
fruits; the rusts Order 2. Hypodermia.
2. Probasidia produced by rounding up and
deposition of thiclc walls by the gener-
ality of the cells of the mycelium Order 3. Ustilaginea.
1. Without probasidia, the basidia divided lon-
gitudinally Order 4. Tremellina.
1. Without probasidia, the basidia undivided.
2. Fruits gelatinous, basidia producing only
two spores on stout sterigmata Order 5. Dacryomygetalea.
2. Not as above.
3. Basidia in a layer which forms with-
out protection or becomes exposed. . .Order 6. Fungi.
3. Basidia formed in closed fruits
which do not open to expose them
as a single layer Order 7. Dermatocarpa.
Order 1. Protobasidiomycetes Engler in Engler and Prantl Nat. Pflanzenfam. I
Teil, Abt 1**: iii (1900).
Suborder Auriculariineae and sub-suborder Auriculariales Engler 1. c.
Order Auricularineac Campbell Univ. Textb. Bot. 175 (1902).
Order Auriculariales Bcssey in Univ. Nebraska Studies 7: 309 (1907).
Basidiomycetes mostly producing probasidia, th.^ basidia divided by transverse
walls, mostly saprophytic and producing gelatinous fruits.
This order includes the family Auriculariacea [Auriculariaceae] Lindau in Engler
and Prantl Nat. Pflanzenfam. I Teil, Abt. 1**: 83 (1900), from which two or three
others have been segregated; about fifteen genera and about 125 species.
Martin (1943) has discussed the name of the genus Auricularia and of its type
species. The organism in question is surely the Jew's ear, Tretnella Auricula L.; the
genus Auricularia Bulliard 1795 can have nothing else as a type. The right name
of the species is Auricularia Auricula (L. ) Underwood 1902. It is a saprophyte on
logs and sticks, producing flattened brown gelatinous fruits a few centimeters in
diameter, vaguely resembling human ears. There are no probasidia. Hyphae growing
toward the surfaces of the fruits produce a palisade of elongate basidia. Each basi-
dium becomes divided by transverse walls into four cells, and each of these sends out
Phylum Inophyta [ 147
to the surface an elongate sterigma which bears a curved basidiospore. The organism
produces also conidia, either from the mycelium, the fruits, or directly from the
basidiospores.
A series of unfamiliar other genera, Platygloca, Cystobasidium, Septobasidium,
etc., have been studied notably by Martin (1934, 1937, 1939, 1942). Jola and
Eocronartium are parasites on mosses. All of these genera produce probasidia, from
which four-celled phragmobasidia arise, as a layer near the surfaces of the fruits.
Most of them produce also conidia.
Order 2. Hypodermia [Hypodermii] Fries Syst. Myc. 3: 460 (1832).
Uredinees Brongniart in Bory de Saint Vincent Diet. Class. Hist. Nat. 16: 471
(1830).
Order Uredineae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1:
74 (1884).
Sub-suborder Uredinales Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1**: iii (1900).
Order Uredinales Bessey in Univ. Nebraska Studies 7: 306 (1907).
Order Pucciniales Clements and Shear Gen. Fung. ed. 2: 147 (1931).
The rusts: parasitic Basidiomycetes, the haploid and dikaryote mycelia usually
attacking different hosts; the dikaryote mycelium producing probasidia, these not
usually compacted into fruits, usually heavily walled and serving as resting spores,
becoming or giving rise to four-celled phragmobasidia.
The typical reproductive structure of the haploid stage is the aecium, a cup-shaped
structure which releases spores called aeciospores; this stage is accordingly called the
aecial stage, and its host the aecial host. In addition to aecia, this stage usually pro-
duces pycnidia or spermagonia. The typical reproductive structures of the dikaryote
mycelium are clusters (telia) of spores called teliospores or teleutospores; this stage,
then, is the telial stage, and its host the telial host. The telial stage usually produces,
beside the teliospores, others called uredospores. The teliospore, or rather (since the
teliospore commonly consists of two or more cells) each cell of the teliospore, is a
probasidium, producing a promycelium which bears four basidiospores. These state-
ments mean that a normal rust produces spores of five kinds. Rusts producing differ-
ent kinds of spores were formerly supposed to be different genera; such were the
Aecidium, Uredo, and Puccinia of Persoon, who, however, remarked of Uredo line-
aris, "vereor, ne junior plantula Pucciniae graminis modo sit." De Bary first proved
that Aecidium Berberis is yet another stage of Puccinia graminis.
The dikaryophase is initiated, of course, by syngamy among cells of the aecial
stage. In Phragmidium violaceum, Blackman observed this to take place between
different cells of the same hypha. Christman (1905) and Moreau (1914), studying
other species of Phragmidium, observed fusion to take place between tips of different
hyphae. Craigie, 1927, showed that Puccinia graminis occurs in two mating types,
and that the fertilizing elements are pycniospores or spermatia. De Bary (1884) had
suggested that this is the truth; his suggestion waited some forty years to be confirmed.
Allen (1930) has described much of the detail. The pycniospores are carried out of
the pycnidium in exuding fluid, and are carried by insects; they make protoplasmic
connection with paraphyses growing from pycnidia of the opposte mating type. The
binucleate uredospores arise from a dikaryote mycelium, but the cup-shaped wall of
the aecium is produced by the haploid mycelium.
148 ] The Classification of Lower Organisms
The first-formed reproductive structures of the dikaryote mycelium on the telial
host are usually uredospores, which remain binucleate and have the function of
spreading the infection of the telial host.
Teliospores may be compacted into palisade-like masses which break through the
epidermis of the host; the masses may be gelatinous and yellow, like fruits of
Auriculariacea. In other genera, the teliospores are gathered into hard, microscopic-
ally stout columns, and in yet others they break through the epidermis in masses not
compacted, each teliospore on a separate stalk. The teliospores of Phragynidium are
chains of several probasidia; those of the many species of Pucciriia are chains reduced
to two probasidia; those of Ravcnelia are globular clusters of probasidia. Almost
always, the teliospores are thick-walled; outside of the tropics, they have the function
of overwintering. Each probasidium contains two nuclei. These unite as a preliminary
to germination: this was first observed by Sappin-Troufi^y (in Dangeard and Sappin-
Troufi'y, 1893). Thereafter the probasidium gives rise to the four-celled promycelium.
The life cycle thus described is not perfectly stable. Aeciospores, uredospores, and
young teliospores are alike dikaryote, and are genetically identical. Spores of the
structure and behavior of any of these types may be produced by processes which
normally lead to another. Thus in Puccinia Malvaccaruvi, the hollyhock rust, syng-
amy leads directly to the production of teliospores on the host of the haploid mycel-
ium; spermagonia, aecia, and uredosori are not produced.
Four families of rusts may be recognized (various authorities make fewer or more).
There are about five thousand species.
Family 1. Melampsoracea [Melampsoraceae] Dietel in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1**: 38 (1900). Teliospores forming a single compact
layer and germinating by producing promycelia. The aecial stages are mostly on
conifers. Some have telial stages on ferns, and FauU (1929) regards these as most
primitive; others attack a variety of flowering plants.
Family 2. Coleosporiacea [Coleosporiaceae] Auctt. The teliospores themselves
becoming basidia by transverse division. In some examples, as Gallowaya, they are
thin-walled.
Family 3. Cronartiacea [Cronartiaceae] Auctt. The teliospores compacted into
columns. Cronartium, with aecial stages on pines; C. ribicola, the important white
pine blister rust, its telial stage on gooseberries and currants.
Family 4. Uredinacea [Uredinaceae] Cohn in Hedwigia 11: 17 (1872). Family
Pucciniaceae Dietel op. cit. 48. The bulk of the rusts, producing teliospores on indi-
vidual stalks. Hemileia vastatrix, the coffee rust; Phragmidium spp., autoecious (at-
tacking a single host) on Rosaceae; Gymnosporangium, the aecial stage on junipers,
the telial (with no uredospores) on plants of the apple tribe; Puccinia, a great num-
ber of species. The races which attack barberry and grasses are all called Puccinia
graminis; but there are morphologically distinguishable strains on wheat, rye, oats,
timothy, Agrostis, and blue grass. Leading an active sexual life and capable of muta-
tion, these strains are subdivisible into large numbers of races distinguished by capa-
city to attack different races of hosts. Given a specimen of rust on wheat, one deter-
mines by trial upon seedlings of ten varieties of wheat to which of 189 numbered races
it belongs. The races occur characteristically in different wheat-growing areas. If one
breeds wheat for resistance to rust, there is good probability of success against the
races occurring locally; but some other race is likely to move into the area (Stakman,
1947).
Phylum Inophyla [ 149
Order 3. Ustilaginea [Ustilagineae] (Tulasne and Tulasne) Winter in Rabenhorst
Kryptog.-Fl. Deutschland 1, Abt. 1: 73 (1884).
Ustilagineae Tulasne and Tulasne in Ann. Sci. Nat. Bot. ser. 3, 7: 73 (1847).
Subclass Hemibasidii Engler Syllab. 26 (1892).
Order Ustilaginales Bessey in Univ. Nebraska Studies 7: 306 (1907).
The smuts: parasitic Basidiomycetes completing their development on a single
host, the dikaryophase mycelium breaking up into thick-walled black spores, these
functioning as probasidia, the basidia usually bearing more than four basidiospores.
In the apparently more primitive smuts, the promycelia are four-celled phragmo-
basidia. The haploid nuclei divide before passing into the basidiospores, with the
effect that each cell of the promycelium buds off a series of basidiospores. In other
examples the promycelia do not become divided by walls, but are of the character of
holobasidia. The basidiospores of some species are capable of budding like yeasts.
In some species, they are capable of syngamy with each other, and in some they send
out hyphae which bear conidia of characteristic form. In many species, syngamy has
not been observed, but is beheved to take place between vegetative hyphae. Hybridi-
zation, and mutation, particularly in the capacity to attack particular races of hosts,
take place freely in smuts, which are accordingly well fitted to cope with the efforts
of plant breeders.
The smuts are believed to be somewhat degenerate descendants of the rusts.
There are two families, about thirty genera, about six hundred species.
Family 1. Ustilaginacea [Ustilaginaceae] Cohn in Hedwigia 11: 17 (1872). The
basidia divided by transverse walls. Ustilago, on grasses and other plants.
Family 2. Tilletiacea [Tilletiaceae] Dietel in Engler and Prantl Nat. Pflanzenfam.
I Teil, Abt. 1** : 15 ( 1900) . The basidia not divided by walls. Tilletia, on grains, etc.
Tuburcinia, Doassansia, the resting spores produced in globular masses.
Order 4. Tremellina [Tremellinae] Fries Syst. Myc. 1: 2 (1821); 2: 207 (1822).
Order Tremellinei Fries Hymen. Eur, 1 (1874).
Order Tremellineae Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1 :
74 (1884).
Suborder Tremellineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1**: iii (1900).
Order Tremellales Bessey in Univ. Nebraska Studies 7: 309 (1907).
Order Tulasnellales Gaumann Vergl. Morph. Pilze 487 (1926).
Saprophytic Basidiomycetes producing gelatinous fruits bearing a layer of basidia
which typically become divided into four cells by longitudinal walls. Each cell pro-
duces a long stout sterigma which reaches the surface of the fruit and bears a spore.
The mycelia, the young fruits, or the basidiospores may bear conidia.
The number of species is perhaps one hundred. Nearly all belong to family Tre-
mellacea [Tremellaceae] Cohn in Hedwigia 11: 17 (1872). Martin (1935, 1937,
1939) has given much study to this group. It is clearly related to the Protobasidi-
omycetes; Patouillardina, having basidia divided by oblique walls, is clearly transi-
tional. Tremella, Sebacina, Tremellodendron, Hyaloria.
Tulasnella differs from the generality of Tremellina in producing holobasidia of
a peculiar type, with bulbous sterigmata (Lindau interpreted the sterigmata as basi-
diospores borne without sterigmata and not released, but producing conidia; it may
be that this interpretation is more sound than the obvious one). It is supposed that
the holobasidia of this genus are derived from the cruciate basidia of proper Tremel-
150 ] The Classification of Lower Organisms
lina by a line of descent separate from those which have produced the holobasidia of
other groups. By leaving Tulasnella in order Tremellina, we spare ourselves the recog-
nition of one more insignificant order.
Order 5. Dacryomycetalea [Dacryomycetales] Gaumann Vergl. Morph. Pilze 490
(1926).
Suborder Dacryomycetineae Engler in Engler and PrantI Nat. Pflanzenfam.
ITeil, Abt. 1**: iv (1900).
Saprophytic Basidiomycetes producing small gelatinous fruits bearing holobasidia
in which two of the nuclei produced by meiosis undergo degeneration, while two
pass into the basidiospores by way of stout sterigmata which give the basidium the
form of a Y. Conidia are produced either from the mycelium, from the young fruits,
or from the basidiospores.
There is a single family Dacryomycetacea [Dacryomycetaceae] Hennings in Engler
and PrantI Nat. Pflanzenfam. I Teil, Abt. 1**: 96 (1900). Dacryomyces, Dacryomi-
tra, Guepinia. Bodman ( 1938) observed the details of the cytological processes in the
basidia.
This insignificant order, like Tulasnella and the two great orders next to be con-
sidered, is evidently derived from Protobasidiomycetes, through Tremellina, by loss
of septa in the basidia; the peculiarities of its basidia suggest an independent origin.
Order 6. Fungi L. Sp. PI. 1171 (1753).
Order Hynienothecii Persoon Syst. Meth. Fung, xvi (1801).
Class Hymenomycetes and orders Pilcati and Clavati Fries Syst. Myc. 1: 1, 2
(1821').
Yzmily Hymenomycetes Fries Espicrisis 1 (1836).
Family Agaricaceae Cohn in Hedwigia 11: 17 (1872).
Order Hymenomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt.
1: 74 (1884).
Suborders Exobasidiineae and Hymenomycetineae Engler in Engler and PrantI
Nat. Pflanzenfam. I Teil, Abt.'l**: iv (1900).
Orders Hymenomycetales and Exohasidiales Bessey in Univ. Nebraska Studies
7: 307, 308 (1907).
Order A^aricalcs Clements Gen. Fung. 102 (1909).
Orders Cantharellales, Polyporales, and Agaricales Gaumann Vergl. Morph.
Pilze495, 503, 519 (1926).
Basidiomycetes producing holobasidia in a layer which is or becomes exposed to
the air, usually on fruits which are woody, leathery, or fleshy, rather than waxy or
gelatinous.
The layer of basidia is called the hymenium. In the lowest members of the group,
the hymenium is formed directly on the mycelium, on the surface of the host or
substratum; in higher examples, it is formed on the surface of more or less compli-
cated fruits; in the highest, it is formed in closed fruits which open to expose it. The
area of the hymenium, and the number of basidia it can bear, is increased when it is
not smooth, but thrown into teeth, ridges, plates, or other projections. Families have
been distinguished chiefly on the basis of the form of the hymenium. The system is not
reliably entirely natural; Overholts (1929) pointed out various microscopic details
which promise to contribute to a more natural system. Among these are cystidia,
swollen cells imbedded in the hymenium and projecting from it; in some examples at
Phylum Inophyta [151
least, they are sterile basidia and serve to hold apart the ridges bearing the hymenium.
Other microscopic features are setae, similar to cystidia but hard, dark, and pointed;
slender hairs called paraphyses; latex ducts; and crystalline inclusions.
There are some fifteen thousand species. The following famiHes are for the most
part the conventionally accepted ones.
Family 1. Exobasidiacea [Exobasidiaceae] Hennings in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. 1**: 103 (1900). The basidia directly on the mycelium. A
sniall group, mostly parasitic on plants. Exohasidium.
Family 2. Thelephoracea [Thelephoraceae] (Saccardo) Hennings (1900). Order
Thelephorei Fries Hymen. Eur. 1 ( 1874) . FamHy Thelephorei Winter ( 1884) . Thele-
phoraceae Saccardo Sylloge 8: xiii (1889). Fruits of various form, gelatinous, fleshy
or leathery, the hymenium covering the surface generally except where it faces up-
ward. Corticium, saprophytic, the fruit a mere appressed layer; Stereum, leathery
shelf-like extensions from decaying sticks and logs: these genera seem to lead into
Lnmily Polyporacea. Cora, a tropical variant of Stereum, is the only lichen-forming
basidiomycete. Thelephora, Craterellus, the fruits club-, funnel-, or cup-like.
Family 3. Clavariacea [Clavariaceae] (Saccardo) Hennings (1900). Order
Clavariei Fries (1874). Family Clavariei Winter (1884). Clavariaceae Saccardo
(1889). Fruits fleshy, club-like or branched; stag-horn fungi. Clavaria, generally
edible.
Family 4. Hydnacea [Hydnaceae] (Saccardo) Hennings (1900). Order Hydnei
Fries (1874). Family //yi/n^i Winter (1884). Hydnaceae Saccardo (1889). Hymen-
ium on the surface of downward-pointing teeth. Fruits assigned to the genus Hydnum
may be massive or variously branched or mushroom-shaped, leathery or fleshy; the
fleshy examples are edible. Fruits of Irpex are little leathery brackets projecting from
sticks and logs, distinguished from Stereum or Polystictus by the masses of fine teeth
projecting below.
Family 5. Polyporacea [Polyporaceae] (Saccardo) Hennings (1900). Order Poly-
poreiYries (1874). Family Polyporei Winter (1884). Polyporaceae Saccardo (1889).
The hymenium lining vertical tubes open below. These are mostly woody or leathery
shelf fungi, mostly saprophytic on wood, numerous and varied in detail. Cooke ( 1940)
recognized forty-six genera in North America. Polyporus, Fames, Polystictus. In Dac-
dalea, the pores are not cylinders but slits; this genus leads into Lenzites, in which the
hymenium is borne on radiating plates, and which is conventionally stationed in
Agaricacea. Boletus has stout fleshy mushroom-shaped fruits, yellow to brown, turn-
ing green when bruised. These fruits are unattractive, but some species are eaten;
others are supposed to be poisonous.
Family 6. Agaricacea [Agaricaceae] Cohn in Hedwigia 11: 17 (1872). Order
Agaricini Fries (1874). Family Agaricini Winter (1884). The hymenium on vertical
plates, radiating from a center, called gills.
These are the Fungi whose fruits are called mushrooms or toadstools. The fruits
are mostly mushroom-shaped, sometimes shelf-like; the texture is usually fleshy, vary-
ing to leathery on the one hand, and on the other to deliquescent, i.e., becoming
converted after maturity into black fluid. There has been much study of the develop-
ment of the fruits (Levine (1922) and Hein (1930) give extensive bibliographies).
This occurs in any of several different fashions, leading to recognizable differences in
the mature structure. For the identification of agarics, many mushroom books are
available. Any interested person, noting the details of structure which result from
the different courses of development, together with the color of the spores (of one
152 ] The Classification of Lower Organisms
of five classes, white, pink to red, light brown to rust color, dark brown or purple, or
black), will find identification reasonably easy. Popular interest in agarics is con-
cerned, of course, with the edible and poisonous. Many amateur mycophagists need to
be convinced that there is no single test for poisonous agarics except the final one.
One who encounters an unfamiliar species may chew and eat a small scrap of it; if
it is tasty and without bad after-effects, one may collect and eat the same species
when one again recognizes it by its technical characters. At the present point, it is
expedient to mention only a few examples.
Deliquescent agarics with black spores are called inky caps and constitute the genus
Coprinus. All are edible; they should be fried in butter and served on toast.
Fruits of Agaricus campestris, the field mushroom, are rather large, white or gray
on top, the stalk marked by a ring but no cup, the gills pink when young, dark brown
to nearly black when mature. Anything of this character is safely edible.
Fruits of Pleurotus have an excentric or lateral stalk, or none, being shelf- or
bracket-like, fleshy, with white spores. All species are edible. The most familiar is
the oyster mushroom, P. ostreatus, producing large white to gray fruits on dead
trees, commonly on poplars.
Fruits of Amanita are marked by cup and ring, and bear white spores. Some species
are known to be edible; others, as the fly agaric, A. muscaria, recognized by a red cap
flecked with white, are extremely poisonous.
Family 7. Podaxacea [Podaxaceae] Fischer in Engler and Prantl Nat. Pflanzenfam.
I Teil, Abt. 1**: 332 (1900). Gyrophragmium produces fruits much like those of
Agaricus, but coming up only to ground level, and drying and shattering irregularly
instead of opening like mushrooms. The gills are quite evident in immature fruits.
Podaxon is similar, but does not form definite gills. These organisms are convention-
ally stationed in the next order, but their obvious natural position is next to
Agaricacea.
Order 7. Dermatocarpa [Dermatocarpi] Persoon Syst. Meth. Fung, xiii (1801).
Order Lytothccii Persoon op. cit. xv.
Class Gasteromycetes and orders Angiogastres and Trichospermi Fries Syst. Myc.
2: 275, 276 (1822).
Family Gasteromycetes Fries Epicrisis 1 (1836)
Order Gasteromycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt.
1:864(1884).
Suborders Phallineae, Hymenogastrineae, Lycoperdineae, Nidulariineae, and
Plectobasidiineae Engler in Engler and Prantl Nat. Pflanznfam. I Teil, Abt.
1**: iv (1900).
Orders Phallineae, Lycoperdineae, and Nidularineae Campbell Univ. Textb.
Bot. 186, 187, 188 (1902).
Orders Hymenogastrales, Phallales, Lycoperdales, Nidulariales, and Sclcroder-
matales Bessey in Univ. Nebraska Studies 7: 306-307 (1907).
Orders Plectobasidiales and Gasteromycetes Gaumann Vergl. Morph. Pilze 537,
544(1926).
Basidiomycetes producing holobasidia enclosed in fruits, not forming a continuous
layer or not exposed as such, not discharging the spores directly into the air, sterig-
mata more or less suppressed.
Distinguished by negative characters, this order may be suspected of being artificial;
but Engler's attempt to correct this produced orders which were small and numerous
I'lixUnu Innfihytd
[ 153
Fig. 29. — Fruits of Agaricacea: upper left, Coprinus atramcntarius; upper right,
Galera tenera; below, Agaricus campestris. Photographs by the late Dr. J. J. McCabe,
by courtesy of the Department of Botany, University of California.
Phylum Inophyta [ 155
to an unsatisfactory degree, and to some of which the suspicion of artificiality con-
tinued to attach.
Dodge, translating Gaumann (1928), took account of the course of development
of the fruits in rearranging those families whose fruits are characteristically pro-
duced underground. The roll of families which appear tenable is as follows.
A. Fruits typically formed underground.
Family 1. Rhizopogonacea [Rhizopogonaceae] Dodge in Gaumann Comp. Morph.
Fungi 469 (1928).
Family 2. Sclerodermea [Sclerodermei] Winter in Rabenhorst Kryptog.-Fl.
Deutschland 1, Abt. 1: 865 (1884). Family Sclerodermataceae Fischer in Engler
and Prantl Nat. Pflanzenfam. I Teil, Abt. 1**: 334 (1900).
Family 3. Hydnangiacea [Hydnangiaceae] Dodge in Gaumann op. cit. 485.
Family 4. Hymenogastrea [Hymenogastrei] Winter in Rabenhorst op. cit. 865.
Family Hymenogastraccae de Toni in Saccardo Sylloge 7: 154 (1888).
Family 5. Hysterangiacea [Hysterangiaceae] Fischer in Engler and Prantl op. cit.
304.
B. Fruits appearing on the surface of the ground.
Family 6. Lycoperdacea [Lycoperdaceae] Cohn in Hedwigia 11: 17 ( 1872) . These
are the common puffballs, Lycoperdon, Bovista, Calvatia, Lycogalopsis, etc. The con-
tents of the more or less globular fruits become disorganized, leaving a mass of spores
m.ixed with fibers (modified hyphae constituting a capillitium), enclosed in one or
more continuous layers of tissue (peridia) which open usually through one stellate
pore at the summit. Geaster has a double peridium. The outer peridium becomes
split by meridional clefts from the apex nearly to the base, and the lobes curl back
in damp weather, exposing the inner peridium with its terminal pore. The appearance
of the fruit in the damp condition explains the common name, earth star, and the
scentific name of the same meaning.
Family 7. Tulostomea [Tulostomei] Winter in Rabenhorst op. cit. 866. Family
Tulostomataceae Fischer in Engler and Prantl op. cit. 342. Tulostoma produces at
ground level puffball-like fruits which are found to stand upon buried stalks some
centimeters long. The basidia bear the spores scattered along the sides instead of in
a crown at the summit. This is probably a minor deviation from the condition in
ordinary puffballs, and not a token of independent origin.
Family 8. Nidulariea [Nidulariei] Winter in Rabenhorst 1. c. Family Nidulariaceae
de Toni in Saccardo Sylloge 7: 28 (1888). The bird's nest fungi, Nidularia, Cyathus,
etc., with small fruits growing on sticks or earth, the outer peridium opening and
exposing several peridioles.
Family 9. Sphaerobolacea [Sphaerobolaceae] Fischer in Engler and Prantl op.
cit. 346. Sphaerobolus, a saprophyte on wood, produces minute puffball-like fruits
which discharge mechanically a globular mass of spores.
Family 10. Clathracea [Clathraceae] Fischer in Engler and Prantl op. cit. 280.
Closely related and transitional to the following family.
Family 11. Phalloidea [Phalloidei] Winter in Rabenhorst 1. c. Family Phallaceae
Fischer in Engler and Prantl op. cit. 289. The stinkhorns. Phallus, Dictyophora,
Mutinus, etc. These organisms produce highly specialized fruits. A fruit is first seen
as a white globe, as large as a marble or a golf-ball, at ground level. It has a leathery
peridium containing certain structures imbedded in gelatinous matter: there is a
firm thimble-shaped structure upon whose surface the basidia develop; below or
within this there is a body of the form of a hollow cylinder of spongy structure. When
156 ] The Classification of Lower Organisms
the spores are ripe, the spongy body grows, so to speak, by unfolding, and becomes,
it may be within an hour, a stalk as much as 15 cm. tall. This happens usually during
the night or at dawn, and is not commonly observed. The growing stalk carries the
basidium-bearing structure into the air, bursting the peridium, which remains as a
cup about the base, and exposing the spores in a mass of jelly which is of an odor
repulsive to man but attractive to carrion-seeking insects. The latter are used as
agents of dissemination.
Chapter X
PHYLUM PROTOPLASTA
Phylum 6. PROTOPLASTA Haeckel
Stdmme Protoplasta and Myxomycetes Haeckel Gen. Morph. 2: xxiv, xxvi
(1866).
Subphylum Plasmodroma Doflein Protozoen 13 (1901), in part.
Subphylum Rhizoflagellata Grasse Traite Zool. 1, fasc. 1: 133 (1952), not order
Rhizoflagellata Kent (1880).
Further names for the myxomycetes as a phylum are cited below under class
Mycetozoa.
Organisms without photosynthetic pigments, mostly with flagellate stages, the
flagella simple or acroneme, not paired and equal nor solitary and posterior; com-
monly occurring also in amoeboid stages. By Haeckel's original publication, the type
or standard is Amoeba, i.e., Amiba diffluens.
Amoeboid organisms are those whose protoplasts lack walls or shells, or are only
incompletely covered by them, and which thrust forth temporary bodies of proto-
plasm, called pseudopodia, functional in motion and in predatory nutrition. Pseudo-
podia are of several types. If massive and blunt they are lobopodia. If fine and
straight, not anastomosing and usually not branching, they are filopodia; or, if they
contain inner filaments, axopodia. If fine, branching, and anastomosing, they are
rhizopodia.
The characters of the pseudopodia distinguish the accepted primary groups of
amoeboid organisms. Variations in this character tend to run parallel to variations
in the structure and composition of shells and skeletons: to a considerable extent,
the accepted groups appear natural. This applies to the second, third, and fourth
among the classes treated below. The phylum, on the other hand, is acknowledgedly
artificial. Some of its groups appear to have had their origins (presumably more
origins than one) among the chrysomonads; others are of unguessed origin.
1. Flagellate in the vegetative condition Class 1. Zoomastigoda.
1. Amoeboid in the vegetative condition.
2. Producing rhizopodia; with shells, these
usually calcareous Class 3. Rhizopoda.
2. Producing filopodia or axopodia; mostly
with skeletons, these usually siliceous Class 4. Heliozoa.
2. Producing lobopodia.
3. Producing flagellate reproductive
cells; mostly macroscopic, subaerial Class 2. Mycetozoa.
3. Not as above; without flagellate
stages Glass 5. Sarkodina.
Class 1 . ZOOMASTIGODA Calkins
Subclass Zoomastigina Doflein Lehrb. Prot. ed. 4: 462 (1916).
Class Zoomastigoda Calkins Biol. Prot. 285 (1926).
Class Zooflagellata Grasse Traite Zool. 1, fasc. 1: 574 (1952).
Class Zoomastigophorea Hall Protozoology 170 (1953).
158] The Classification of Lower Organisms
Non-pigmented flagellates having acroneme or simple flagella; amoeboid stages,
if they occur, having lobopodia. The standard is Bodo. Four orders are to be recog-
nized.
1. Flagella one or two Order 1. Rhizoflagellata.
1. Flagella four to eight (in each neuromotor
system, if these are more than one).
2. Axostyles, if present, homologous with
flagella; parabasal body commonly ab-
sent Order 2. PoLYMASXiGroA.
2. Axostyles present, not homologous with
flagella; parabasal body present, disap-
pearing during mitosis Order 3. Trichomonadina.
1. Flagella of indefinite large numbers Order 4. Hypermastigina.
Order 1. Rhizoflagellata [Rhizo-Flagellata] Kent Man. Inf. 1: 220 (1880).
Orders Trypanosomata (the mere plural of a generic name) and Flagellato-
Pantostomata in part Kent op. cit. 218, 229.
Suborders Monadina in part and Heteromastigoda Biitschli in Bronn Kl. u. Ord.
Thierreichs 1: 810, 827 (1884).
Protomastigina Klebs in Zeit. wiss. Zool. 55: 293 (1893).
Order Protomonadina Blochmann Mikr. Tierwelt ed. 2, 1 : 39 (1895).
Subclasses Pantostomatineae and Protomastigineae Engler in Engler and Prantl
Nat. Pflanzenfam. I Teil, Abt. la: iv (1900).
Orders Pantostomatales and Protomastigales Engler Syllab. ed. 3: 7 (1903).
Orders Cercomonadinea and Monadidea in part Poche in Arch. Prot. 30: 139,
140 (1913).
Orders Pantostomatineae and Protomastigineae Lemmermann in Pascher Siiss-
wasserfl. Deutschland 1: 30, 52 (1914).
Order Rhizomastigina Doflein Lehrb. Prot. ed. 4: 704 (1916).
Orders Pantostomatida and Protomastigida Calkins Biol. Prot. 286, 288 (1926).
Orders Trypanosomidea Grasse, Bodonidea Hollande, and Proteromonadina
Grasse in Grasse Traite Zool. 1, fasc. 1: 602, 669, 694 (1952).
Orders Rhizomastigida and Protomastigida Hall Protozoology 171, 173 (1953).
Non-pigmented flagellates with one flagellum or two unequal flagella, these
simple or acroneme; commonly with amoeboid stages, or amoeboid while bearing
flagella. The type, being the sole genus of Rhizo-Flagellata as originally published, is
Mastigamoeba, i. e., Chaetoproteus Stein.
As the synonymy shows, most authorities have made these organisms two orders,
Pantostomatales (or some such name), amoeboid in the vegetative condition, and
Protomastigina (or the like), not definitely so. Monas, and the choanoflagellates
and Amphimonadaceae, usually included in the latter order, have in the present work
been given places elsewhere. The residue of the Protomastigina are not sharply
different in character from the original Rhizoflagellata, and are accordingly placed
in the same order. The resulting group is not a very numerous one. Some examples
appear to occur naturally as predators in uncontaminatcd waters; the majority have
been found in foul or contaminated waters, or in feces, and are believed to be
naturally cntozoic, cither commensal or parasitic. Further examples are parasites in
blood. A cytological character marking the majority of the goup, but not confined
to it, is the parabasal body (better, perhaps, the kinetoplast; Kirby, 1944). This is a
Phylum Protoplasta [ 159
rather massive extranuclear body regularly present in the cell and distinct both
from the centrosome and the blepharoplast. In the present group, it divides when
the nucleus does. Thus this group, although marked chiefly by characters which are
negative or derived, appears possibly to be natural.
1. Flagella two.
2. Cells not notably slender Family 1. CERCOMONADroA.
2. Cells notably slender Family 2. TRYPANOPLASMroA.
1. Flagellum one.
2. Not regularly markedly amoeboid Family 3. Oicomonadacea.
2. Conspicuously amoeboid Family 4. CHAETOPROXEroA.
Family 1. Cercomonadida [Cercomonadidae] Kent Man. Inf. 1: 249 (1880).
Family Bodonina Butschli in Bronn Kl. u. Ord. Thierreichs 1: 827 (1884). Family
5oc?onflccag Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. la: 133 (1900).
Family Bodonidae Doflein Protozoen 73 (1901). Family Cercobodonidae Hollande
1942. Family Proteromonadidae Grasse Traite Zool. 1, fasc. 1: 694 (1952). Non-
pigmented flagellates, the bodies not notably slender, with two flagella, one directed
anteriorly, the other trailing. Fischer (1894) found both of the flagella of Bodo to
be acroneme.
In Bodo both flagella are free of the body. There are numerous species, in infusions
or foul or polluted waters, or entozoic in a wide variety of animals, from insects to
men. Prowazekia, Proteromonas, and Pleuromonas are doubtfully c'istinct. Rhyncho-
monas, from fresh or foul waters, is distingished by a protoplasmic beak in which
the anterior flagellum is imbedded. Cercomonas, of like habitats, has the trailing
flagellum grown fast to the cell membrane; the eel! : exhiljit a considerable capacity
to send out lobopodia.
Biflagellate organisms which can lose their flagella and take on the appearance
of ordinary amoebas have repeatedly been discovered and variously named. So far as
the pseudopodia are lobopodia and the flagella are unequal, these organisms belong
in this family; but many accounts fail to establish the equality or inequality of the
flagella, with the result that the names used in them cannot be applied with confidence.
This is true of various organisms originally named under Pseudospora, Dimastiga-
moeba, and Naegleria. The earliest generic name definitely applicaple to organisms
as described in Cercobodo Senn, 1910.
Belar (1914, 1916, 1920, 1921), Kuhn (1915), and others have described mitosis
in various examples of this family; the most detailed account is of Bodo Lacertae in
Belar's paper of 1921. The flagella spring from a blepharoplast from which a rhizo-
plast extends into the nucleus. The chromatin is reticulate, not massed in a karyo-
some, but no centrosome has been recognized in it when it is not dividing. The
rhizoplast, where it passes through the cytoplasm, is surrounded by stainable Ring-
korper. The parabasal body, located on the posterior side of the nucleus, is massive
and often irregular. In division, the blepharoplast divides, each part retaining one
flagellum and generating an additional one. The rhizoplast appears to begin to split,
but presently it and the Ringkorper become invisible. Within the intact nuclear mem-
brane there appears a spindle with evident centrosomes at the poles. The centrosomes
come presently to the inner surface of the nuclear membrane, while the blepharo-
plasts move to adjacent positions on the outside. Chromosomes duly assemble at
the equator of the spindle and undergo division. Division of the nucleus is com-
pleted by constriction of the nuclear membrane; the parabasal body undergoes
constriction; the cell divides by constriction lengthwise. The Ringkorper and the
rhizoplast are apparently regenerated by the blepharoplast.
160]
The Classification of Lower Organisms
5.
Fig. 30. — Rhizoflagellata : a, Bodo sp. x 1,000. b, c, Cercomonas longicauda
as identified by Wenyon (1910) in material from a cholera patient; d, the same as
identified by Hovasse (1937) in swamp water, e-h, Cryptobia spp.; e-g, cell and
division stages of a species from the conger eel after Martin (1910); h, a species
from siphonophores after Keysselitz (1904) x 1,000. i, Phytomonas Donovani after
Franga (1914). j-p, Trypanosoma Lewisi; j, k, forms from the rat after Minchin
(1909); 1-p, forms from the flea Ceratophyllus fasciatus after Minchin & Thomp-
son (1915). q. Division stage of Trypanosoma Brucii after Kiihn & Schuckmann
(1911). r, Chaetoproteus [Mastigamoeba aspera) after Schulze (1875) x 100.
X 2,000 except as noted.
Phylum Protoplasta [161
AlexeiefF (1924) described fusions of pairs of cells of Bodo edax.
Family 2. Trypanoplasmida [Trypanoplasmidae] Hartmann and Jollos 1910. Fam-
ily Cryptobiidae Poche in Arch. Prot. 30: 148 (1913). Family Trypanophidae Hol-
lande in Grasse Traite Zool. 1, fasc. 1: 680 (1952). Organisms of essentially the
structure of Cercomonas, but notably slender in adaptation to parasitic life, the
trailing flagellum forming the margin of an undulating membrane on the body.
Parasitic in various invertebrates and in the gut and blood of fishes.
The numerous species may be included in a single genus Cryptobia Leidy [Try-
panoplasma Laveran and Mesnil; Trypanophis Keysselitz).
According to Martin's (1910) description of a species from the eel Conger niger,
both flagella spring from a blepharoplast ("basal granule") at the anterior end.
As preliminary to division, the blepharoplast and flagella divide, and one blepharo-
plast migrates to the posterior end of the cell. The nucleus divides by constriction of
the nuclear membrane. There is a prominent parabasal body ("kinetonucleus") which
divides by constriction, as does the cell, transversely.
Belar (1916) described sexual fusions of differentiated individuals of a species
parasitic in snails.
Family 3. Oicomonadacea [Oicomonadaceae] Senn in Engler and Prantl Nat.
Pflanzenfam. I Teil, Abt. la: 118 (1900). Family Trypanosomidae Doflein Proto-
zoen 55 (1901). Family Trypanosomatidae Grobben 1904. Family Oicomonadidae
Hartog. Non-pigmented anteriorly uniflagellate organisms, not markedly amoeboid
while in the flagellate condition.
Oikomonas includes organisms of the character of the family without particular
specialization, occurring in contaminated water or soil, and as commensals in the
intestine of animals.
The bulk of the family consists of the slender-celled parasites which may be
celled trypanosomes in the broad sense of the word. From the viewpoint of man, these
are the most important flagellates, and they have been the most intensely studied.
Some are known only from the guts of insects; some occur alternatively in insects
and plants; some in insects and vertebrates; and some in vertebrates and in inverte-
brates other than insects, as ticks and leeches. The range of parasitization is as
though the group had evolved as parasites in insects, and had been carried to
other hosts by the activity of insects and other biting or sucking invertebrates.
Most trypanosomes occur in varied forms. The forms are designated by words
which originated as names of genera and remain in use as such. ( 1 ) The leptomonas
form has an anterior flagellum but no undulating membrane; it resembles a cell of
Oikomonas but is notably slender. (2) The leishmania form has no flagellum; the cell
is rounded up and lives attached to, or inside of, cells of the host. (3) In the crithidia
form, the base of the flagellum is continued as an undulating membrane more or less
to the middle of the cell. (4) In the trypanosoma form, the base of the flagellum is
continued as an undulating membrane to the posterior end of the cell.
The accepted genera are distinguished (artificially, as one may suspect) by stages
produced and groups of hosts attacked, as follows:
l.With leptomonas stages in insects and in
Euphorbiaceae, Ascelepiadaceae, and other
plants with milky juice Phytomonas.
1. Confined to invertebrate animals.
2. Trypanosoma stage known Herpetomonas.
2. Trypanosoma stage unknown; crithidia
162 ] The Classification of Lower Organisms
stage known Crithidia.
2. Trypanosoma and crithidia stages un-
known Leptomonas.
1. Attacking vertebrate animals.
2. Trypanosoma stage known Trypanosoma.
2. Trypanosoma stage unknown Leishmania.
Man has been concerned particularly with Trypanosoma gambiense, the agent of
African sleeping sickness; T. Cruzi, the cause of Chagas' disease; T. Brucii, T. Evansi,
T. equinum, and T. equiperdum, which cause in domestic animals the diseases,
respectively, nagana, surra, mal de caderas, and dourine; Leishmania Donovani and L.
tropica, causing kala azar and oriental sore; and L. brasiliensis, causing espundia,
ferida brava, or chicleros' ulcer, usually appearing as a grievous disfigurement of
the features.
Schaudinn (1903), having studied a trypanosome occurring in mosquitoes and in
the owl Athene noctua, described the nucleus as undergoing repeated unequal divi-
sions. It appeared to him that when a cell is to produce a flagellum, one of the
minor nuclei produced by unequal division generates it. Prowazek (1903) described
similar phenomena in a Herpetomonas occurring in flies. These reports led Woodcock
(1906) to apply to the proper nucleus of trj^panosomes the term trophonucleus, and
to the large granule near the base of the flagellum the term kinetonucleus.
There has been much other study of the cytology of trypanosomes (as by Minchin,
1908, 1909; Robertson, 1909; Woodcock, 1910; Minchin and Woodcock, 1910, 1911;
Kiihn and Schuikmann, 1911; Minchin and Thomson, 1915; Schuurmans Stekhoven,
1919). This has not confirmed the foregoing accounts and conclusions, but appears
to have established the following points.
The base of the flagellum is slightly swollen and may be construed as a blepharo-
plast. Separated from the blepharoplast by a distance of one or two microns there
is a conspicuous parabasal body (the kinetonucleus of Woodcock). Fine strands con-
necting the blepharoplast, parabasal body, and nucleus, have been observed. Most of
the stainable material in the resting nucleus is aggregated in a globular karyosome.
In mitosis, the karyosome breaks up to form a moderate number of chromosomes and
a central granule, evidently a centrosome, which stains more heavily than the chromo-
somes. It divides before the chromosomes, the daughter centrosomes remaining con-
nected by a fine fiber, the centrodesmose. An obscure spindle forms about the centro-
desmose; thi' chromosomes undergo division within the spindle, and the daughter
chromosome > assemble about the centrosomes. Mitosis is completed by constriction
of the nuclear membrane.
The blepharoplast divides at the same time as the nucleus. The flagellum splits
to a short distance and one of the branches breaks loose; one daughter blepharoplast
retains essentially the whole of the original flagellum while the other generates one
which is almost entirely new. The parabasal body undergoes constriction. The cell
membrane cuts in in such fashion as to divide the cell longitudinally. The blepharo-
plast and the parabasal body persist through the non-flagellate leishmania stage.
Reports that the nucleus may generate these structures, or that one of them may
generate another, were apparently mistaken.
Schaudinn described complicated processes by which a trypanosome generates
differentiated male and female gametes which duly undergo syngamy. His account is
believed to have resulted from mistaking stages of a sporozoan for those of a trypa-
nosome. Still, the occurrence of syngamy among trypanosomes is inherently probable.
Phylum Protoplasta [ 163
Family 4. Cliaetoproteida [Chaetoproteidae] Poche in Arch. Prot. 30: 172 (1913).
Family Rhizomastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 810 (1884).
Family Rhizomastigaceae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt.
la: 113 (1900). Family Mastigamoebidae Kudo Protozoology ed. 3: 263 (1946).
Amoeboid organisms bearing one anterior flagellum, either permanently or tempor-
arily. In polluted soil or water, or commensal or pathogenic in animals.
The oldest genus, Chaetoproteus Stein {Mastigamoeba F. E. Schulze, 1875; Din-
amoeba Leidy ?) remains poorly known. This organism and Mastigella are described
as fairly large; Craigia is much smaller. Rhizomastix is doubtfully distinct from
Craigia. Early names of this family appear to refer to Rhizomastix as the type, but
the family is much older than the genus, and the names are not valid.
Order 2. Polymastigida Calkins Biol. Prot. 292 (1926).
Family Polymastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 842 (1884).
Order Polymastigina Blochmann Mikr. Tierwelt ed. 2, 1: 47 (1895).
Subclass Distomatineae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: iv (1900).
Order Distomatinales Engler Syllab. ed. 3: 7 (1903), not based on a generic
name.
Orders Pyrsonymphina, Oxymonadina, Retortomonadina, and Distomata Grasse
Traite Zool. 1: fasc. 1: 788, 801, 824, 963 (1952).
Non-pigmented flagellates with simple or acroneme flagella of definite number,
from four to eight (two in Retortomonas), in the individual neuromotor system, and
accordingly on the individual cell, except when the neuromotor systems are multi-
plied; not of the definite characters of the following order. Free-living, chiefly in
foul waters, or commensal or parasitic in animals. Polymastix is presumably the type
of the group. It was listed with a query in Biitschli's original publication of family
Polymastigina.
In the generality of Polymastigida, the cells are dorsiventral and have single nuclei
and neuromotor systems. There are derived examples in which the cells are spirally
twisted. There is a group in which the cells are double, having two nuclei and neuro-
motor systems. In another group there are two or more neuromotor systems, usually
with more than one nucleus; the cells consist of units in a whorled or spiral arrange-
ment, so that as wholes they are of radial symmetry.
The neuromotor system consists primarily of ( 1 ) the flagella; (2) one or more
blepharoplasts from which the flagella spring; (3) one or more rhizoplasts linking
together the parts of the system; and (4) a centrosome located just outside the nuclear
membrane. Furthermore, (5) a parabasal body may be present. (6) An axostyle is a
rod imbedded in the cytoplasm. In Hexamita the axostyles are the proximal ends of
backwardly directed flagella; axostyles occurring in various other genera of the order
appear also to be homologous with flagella.
Nuclear and cell division have been observed in various genera, as in Hexamita by
Swezy (1915); in Streblomastix by Kidder (1929); in Giardia by Kofoid and Chris-
tianson (1915) and Kofoid and Swezy (1922); and in O.V);mona^ by Connell (1930).
Cleveland (1947) observed in Saccinobaculus a multiplication of nuclei followed
by their fusion in pairs, and by meiosis in the fusion nuclei: thus there is a .sexual
cycle without fusion of cells. It is not probable that sexual reproduction does not
occur in the generality of the group, but it has not been observed in any others.
164]
The Classification of Lower Organisms
Fig. 31. — PoLYMASTAcroA : a, Polymastix Mclolonthae after Swezy (1916). b,
Streblomastix Strix x 1,000 after Kidder ( 1929) . c, d, Giardia cnterica after Kofoid
& Swezy (1922). Trichomonadina: e, Hcxamastix Tcrmopsidis after Kirby
(1930). i' Tricercomitus Termopsidis 2ihtv YJirhy (1930). g, Macrotrichomonas
pulchra after Kirby (1938). h. Trichomonas tenax x 4,000 after Hinshaw (1926).
i, Pentatrichomonas obliqua after Kirby (1943). j, Snydcrella Tabogae x 500 after
Kirby ( 1929) . x 2,000 except as noted.
Phylum Protoplasta [165
In making the clearly natural group of trichomonads a separate order, Kirby ( 1947 )
removed the majority of the species formerly assigned to this order, and left a mis-
cellany of small isolated families. It seems not expedient to make them several small
orders, as Grasse has done; rather they are to be held together until their respective
relationships become evident. A hint of Hall has led in the present work to the trans-
fer of family Trimastigida to order Ochromonadalea.
1. With a single nucleus and neuromotor system.
2. Cells not spirally twisted, at least not as
wholes and not conspicuously Family 1. TEXRAMiTroA.
2. Entire cells conspicuously spirally
twisted.
3. With four free flagella Family 2. Streblomastigida.
3. With four or eight flagella whose
proximal ends are grown fast to the
cell membrane Family 3. Dinenymphida.
1. With one or several nuclei and two or more
neuromotor systems Family 4. Oxymonadida.
1. With two nuclei and neuromotor systems Family 5. Trepomonadida.
Family 1. Tetramitida [Tetramitidae] Kent Man. Inf. 1: 312 (1880). Families
Tetramitina and Polymastigina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 841,
842 (1884). Family Tetramitaceae Senn in Engler and Prantl Nat. Pflanzenfam. I
Teil, Abt. la: 143 ( 1900) . Family Polymastigidae Doflein Protozoen 83 ( 1901 ) . Fam-
ily Chilomastigidae Wenyon (1926). Family Costiidae Kudo Handb. Prot. 153
(1931). Family Retortomonadidae Wenrich 1932. Cells mostly dorsiventral and
with four flagella; these uniform or differentiated; when differentiated, one or two
may trail behind the cell. Axostyles present or absent, parabasal bodies not reported.
Like the order, the family is a miscellany; good authority has made as many as four
families of the few genera. Tetramitus, free-living, unfamiliar. Costia, occurring
usually as sessile parasites on fishes. Polymastix, in insects. Monocercomonoides, in
insects and vertebrates. Chilomastix, in insects and vertebrates, cells marked by a
cytostomal groove into which one of the flagella, shorter than the others, is recurved.
The species which occurs in man (usually, as it appears, as a harmless commensal)
is in most works called C. Mesnili; the correct name is apparently Chilomastix Hom-
inis (Davaine) n. combl. Current authority places next to Chilomonas the biflagellate
Retortomonas, also in insects and vertebrates, and having cells of essentially the
same structure.
^Kofoid (1920) gave the history involved in this combination. Davaine, 1860, de-
scribed the flagellates Cercomonas Hominis var. A and var. B. The two forms are
not of the same species, and Moquin-Tandon, in the same year, re-named them
respectively C. Davainei and C. obliqua. They are not of the same genus, being re-
spectively a Chilomastix and a Pentatrichomonas, under which genera they have
various names. Kofoid named them respectively Chilomastix davainei and Tricho-
monas hominis. In so doing, he may be held to have exercised his right to choose a
type in a group in which no type has been designated; but it is arguable on the con-
trary that an author who designates a var. A designates the type in doing so. It is
on the basis of this argument that the new combination here published is applied
to the Cercomonas Hominis var. A of Davaine.
166] The Classification of Lower Organisms
Family 2. Streblomastigida [Streblomastigidae] Kofoid and Swezy in Univ. Cali-
fornia Publ. Zool. 20: 15 (1919). The only known species is Strehlornastix Strix, a
slender spirally twisted organism with four anterior flagella, free-swimming or at-
tached in the gut of the termite Termopsis. The significance of the epithet Strix (a
Greek noun meaning screech owl) as applied to this species is not clear.
Family 3. Dinenymphida [Dinenymphidae] Grassi in Atti Accad. Lincei ser. 5.
Rendiconti CI. Sci. 20, 1° Semestre: 730 (1911). Elongate flagellates, the four or
eight anterior flagella adherent to the body and spirally twisted with it, free at their
distal ends. Often beset with spirochaets, which have been mistaken for additional
flagella; the family has been misplaced in order Hypermastigina. Dinenympha and
Pyrsonympha in termites; Saccinohaculus in the wood roach Cryptocercus.
Family 4. Oxymonadida [Oxymonadidae] Kirby in Quart Jour. Micr. Sci. n. s. 72:
380 ( 1928) . Flagellates with radially symmetrical bodies including two or more neuro-
motor systems, entozoic in termites of subfamily Kalotermitinae. Each pear-shaped
cell of Oxymonas has one nucleus and two neuromotor systems (Kofoid and Swezy,
1926). In Microrhopalodina {Proboscoidella) each cell contains a whorl of nuclei,
each with its separate neuromotor system (Kofoid and Swezy, 1926; Kirby, 1928).
These organisms are superficially closely similar to the Calonymphida, from which
Kirby distinguished them.
Family 5. Trepomonadida [Trepomonadidae] Kent Man. Inf. 1: 300 (1880).
Family Hexamitidae Kent op. cit. 318. Distomata Klebs in Zeit. wiss. Zool. 55: 329
(1893). Family Distomataccae Senn in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. la: 148 (1900). Flagellates each with two nuclei and two neuromotor systems.
In most examples, each half-cell is dorsiventral, and the whole isobilateral, with two
cytostomes. Most of the genera, Trepomonas, Gyromonas, Trigonomonas, are free-
living in fresh or foul waters and have been little studied. Hexamita occurs both free-
living and entozoic, in roaches and in all classes of vertebrates; the cells have eight
flagella {Octomitus Prowazek and Urophagus Moroff are synonyms). In Giardia
the half-cells are asymmetric, and the whole cells dorsiventral, with one cytostome.
There are several species, serious pathogens in mammals. The valid name of the
species in man, usually known as G. Lamblia, appears to be G. enterica (Grassi)
Kofoid (1920).
Order 3. Trichomonadina Grasse Traite Zool. 1, fasc. 1: 704 (1952).
Order Trichomonadida Kirby in Jour. Parasitol. 33: 215, 224 (1947), preoc-
cupied by family TRiCHOMONADroAE Wenyon (1926).
Flagellates of the general nature of the Polymastigida having in each neuromotor
system one trailing flagellum; axostyle present, rigid, apparently not homologous
with the flagella; parabasal body present, disappearing during mitosis. Entozoic, the
majority of the species, to the number of fully 150, occurring in termites.
The base of the trailing flagellum may be underlain by a cresta, a more or less
prominent body distinct both from parabasal body and from axostyle. The trailing
flagellum may be grown fast to the cell membrane and converted into an undulating
membrane; in this case it is underlain by a rod called the costa, apparently homolo-
gous with the cresta (Kirby, 1931).
Nuclear and cell division have been described in Trichomonas by Kuczynski
(1914), Kofoid and Swezy (1915, 1919; the Trichomitiis described in the latter
year is a Trichomonas) and Hinshaw (1926). The centrosome (or a combined cen-
trosome and blcpharoplast, the centroblcpharoplast of Kofoid and Swezy, 1919) lies
Phylum Protoplasta [167
outside the nuclear membrane. This structure divides and the daughter structures
move apart along the nuclear membrane. They remain connected, usually until mito-
sis is complete, by a stainable strand, the paradesmose. Definite chromosomes, usually
few in number, and an intranuclear spindle, are formed. Mitosis is completed by con-
striction of the nuclear membrane. In what appears to be the typical course of cell
division, the rhizoplast and blepharoplast divide when the centrosome does. Of other
parts of the neuromotor system, some may remain connected to one blepharoplast
and some to the other; some may disappear. The parts needed to complete a neuro-
motor system are regenerated in each daughter cell.
1. With a single nucleus and neuromotor system.
2. Lacking a cresta, costa, or undulating
membrane Family 1. MoNOCERCOMONADroA.
2. With a trailing flagellum whose base is
underlain by a cresta Family 2. DEVEScoviNroA.
2. With a trailing flagellum grown fast to
the cell membrane, forming an undula-
ting membrane underlain by a costa Family 3. Trichomonadida.
1. With several nuclei and neuromotor systems. . Family 4. CALONYMPHroA.
Family 1. Monocercomonadida [Monocercomonadidae] Kirby in Jour. Parasitol.
33: 225 (1947). Minute flagellates of the appearance of certain Tetramitida, but
having a firm axostyle, the parabasal body disappearing and a paradesmose forming
between the daughter centrosomes during mitosis; lacking a cresta, costa, or undulat-
ing membrane; entozoic in termites and other insects, and in all classes of vertebrates.
Monocercomonas, Hexamastix, Tricercomitus.
Family 2. Devescovinida [Devescovinidae] Doflein Lehrb. Prot. ed. 3: 537 (1911).
Subfamily Devescovininae Kirby in Univ. California Publ. Zool. 36: 215 (1931).
Organisms with three anterior flagella and a larger trailing flagellum underlain by a
cresta; confined to termites of the families Mastotermitidae, Hodotermitidae, and
Kalotermitidae, being most abundant in the last. The cells, usually fairly large, ingest
scraps of wood and are presumed to contribute to the lives of their hosts by digesting
it. Devescovina, Gigantomonas, Macrotrichomonas, Foaina, Parajoenia, Metadeves-
covina. Spirochaets which share the habitat of these organisms are commonly found
adhering to their cell membranes, and were mistaken for additional flagella in the
original descriptions of some of the genera.
Family 3. Trichomonadida [Trichomonadidae] Wenyon Protozoology 1 : 646
(1926). Flagellates with three or more flagella directed forward and one trailing, the
proximal part of the latter grown fast to the cell membrane and forming an undula-
ting membrane underlain by a costa. Entozoic in a wide variety of animals. Tricho-
monas, normally with four anterior flagella, is the most numerous genus. It occurs
in termites, including those of the advanced family Termitidae, in which scarcely
any other flagellates occur; it does not ingest wood, and is not believed to be benefi-
cial to its hosts. It occurs also in all classes of vertebrates. Man harbors Trichomonas
tenax as a commensal in the mouth. T. vaginalis may be a serious pathogen. Penta-
trichomonas obliqua (Moquin-Tandon) comb. nov.,l commensal (or pathogenic?)
in the gut has at the anterior end a fifth flagellum separate from the other four
(Kirby,^1943).
icf. footnote, p. 165.
168] The Classification of Lower Organisms
Family 4. Calonymphida [Calonymphidae] Grass! in Atti Accad. Lincei ser. 5,
Rendiconti CI. Sci. 20, 1° Semestre: 730 (1911). Flagellates with radially symmetri-
cal bodies including more than two nuclei and neuromotor systems, the latter of
trichomonad type; entozoic in termites of subfamily Kalotermitinae. These flagellates
ingest scraps of wood and are believed to contribute to the nutrition of their hosts.
In Coronympha each cell contains one whorl of nuclei each with its separate neuro-
motor system (Kirby, 1929). In Stephanonympha, the nuclei and neuromotor systems
are so numerous as to form a spiral band of several cycles in the anterior part of the
cell. In Calonympha, besides numerous neuromotor systems associated with nuclei,
there are others free of any nucleus; in Snyderella, the two types of structures are
independently multiplied.
Order 4. Hypermastigina Grassi in Atti Accad. Lincei ser. 5, Rendiconti CI. Sci.
20, 1° Semestre: 727 (1911).
Order Trichonyynphidea Poche in Arch. Prot. 30: 149 (1913).
Order Hypermastigida Calkins Biol. Prot. 29"5 (1926).
Order Lophomonadida Light in Univ. California Publ. Zool. 29: 486 (1927).
Orders Joeniidca, Lophomonadina, Trichonymphina, and Spiratrichonym-
phina, Grasse Traite Zool. 1, fasc. 1: 837, 851, 862, 916 (1952).
Flagellates, mostly large and of radial symmetry, with single nuclei and indefi-
nitely numerous flagella. Entozoic in roaches and in termites excluding those of
family Termitidae. Lophomonas is to be regarded as the type.
Cleveland (1925, 1926) found it possible, by starvation or by exposure to high
pressures of oxygen or high temperatures, to rid insects of all of their intestinal
flagellates or of some of the kinds. When completely freed of flagellates, wood roaches
and termites of the lower families are able to remain alive only for a few weeks. The
life of Termopsis is not prolonged by the presence of Streblomastix, and it is pro-
longed only moderately by the presence of Trichomonas Termopsidis. But if infested
with either Trichonympha Campanula or T. sphaerica, it can survive indefinitely on
a diet of pure cellulose. Both species ingest the ground scraps of wood which reach
the part of the intestine in which they occur; it is evident that they serve their hosts
as agents of digestion. Cleveland's observations raise unanswered questions as to the
occurrence of fixation of nitrogen; it is known only that termites are quite economical
in their use of nitrogenous compounds available to them.
The Hypermastigina have elaborate neuromotor systems. There is regularly a large
centroblepharoplast. In what appears to be the relatively primitive type of cell divi-
sion, as in Trichonympha (Kofoid and Swezy, 1919), the neuromotor system of the
mother cell is divided between the daughter cells. In Spirotrichonympha (Cupp,
1930), only the centroblepharoplast divides; the neuromotor system of the mother
cell remains attached to one of the daughter centroblcpharoplasts, while the other
generates the remaining parts of a complete system. In Lophomonas (Kudo, 1926),
and Kofoidia (Light, 1927), the neuromotor system of a dividing cell is absorbed
or discarded, with the exception of the centroblcpharoplasts, from which new systems
develop.
In Trichonympha and Spirotrichonympha the details of nuclear division have
much the appearance of meiosis. A double set of chromosomes appears, and the
chromosomes form pairs which are divided in the spindle. It is supposed that this
appearance is produced by a precocious splitting of the chromosomes.
Phylum Protoplasta [ 169
In species of Trichonympha, Leptospironympha, and Eucomonympha from the
wood roach Cryptocercus, Cleveland (1947, 1948) observed the syngamy of undiffer-
entiated or diflFerentiated gametes; the appearance of the process is as though the egg
ingested the sperm. Syngamy is followed immediately by meiosis. This means that
vegetative individuals are haploid. Barhulanympha achieves without syngamy an al-
ternation of haploid and diploid stages. Diploid cells are produced when a centro-
blepharoplast fails to divide, with the result that the nucleus remains intact, while
chromosomes appear and divide. Reduction division, by the separation of undivided
chromosomes, occurs when a centroblepharoplast divides at an exceptionally early
stage. Cleveland concluded that the early division of the central body is the event
which primarily distinguishes meiosis from mitosis. It is possible that he has recog-
nized an essential feature of the evolution of the sexual cycle. His words suggest the
idea that the sexual cycle may have originated within the present group. This is an
impossibility; the sexual cycle is a normal character of nucleate organisms, and is
fully established in nucleate organisms far more primitive than these.
There are fewer than one hundred known species of Hypermastigina. They are
treated as seven families.
1. Body without segmented appearance.
2. Flagella distributed generally over the
surface of the body or its anterior part. . . . Family 1. TRiCHONYMPHroA.
2. Flagella in spiral bands Family 2. HoLOMASTiGOTororoA.
2. Flagella in tufts.
3. Flagella in a single tuft Family 3. LoPHOMONAoroA.
3. Flagella in two tufts Family 4. HoPLONYMPHroA.
3. Flagella in four tufts Family 5. SxAUROjOENnDA.
3. Flagella in many tufts Family 6. KoForonoA.
1. Body with segmented appearance Family 7. Teratonymphida.
Family 1. Trichonymphida [Trichonymphidae] Leidy ex Doflein Lehrb. Prot. ed.
3: 537 (1911). The numerous flagella distributed generally over the surface of the
body or its anterior part. Trichonympha {Leidy opsis), Eucomonympha, etc.
Family 2. Holomastigotoidida [Holomastigotoididae] Janicki in Zeit. wiss. Zool.
112: 644 (1915). Family S pirotrichonymphidae Grassi in Mem. Accad. Lincei CI.
Sci. ser. 5, 12: 333 (1917). The numerous flagella arranged in spiral bands. Holo-
m.astigotoides, S pirotrichonympha, etc.
Family 3. Lophomonadida [Lophomonadidae] Kent Man. Inf. 1: 321 (1880).
Family Joeniidae Janicki in Zeit wiss. Zool. 112: 644 (1915). The numerous flagella
assembled in a single anterior tuft. Lophomonas, in cockroaches, all of the flagella
directed forward. Joenia, Joenina, Joenopsis, etc., in termites, the outer flagella
directed backward.
Family 4. Hoplonymphida [Hoplonymphidae] Light in Univ. California Publ.
Zool. 29: 138 (1926). The flagella assembled in two anterior tufts. Hoplonympha,
Barhulanympha, etc.
Family 5. Staurojoeninda [Staurojoenindae] Grassi in Mem. Accad. Lincei CI. Sci.
ser. 5, 12: 333 (1917). The flagella assembled in four anterior tufts. Staurojoenina.
Family 6. Kofoidiida [Kofoidiidae] Light in Univ. California Publ. Zool. 29: 485
(1927). The flagella fused at their bases into several bundles. Kofoidia, a single
known species in Kalotermes.
Family 7. Teratonymphida [Teratonymphidae] Koidzumi in Parasitology 13: 303
(1921). Family Cyclonymphidae Reichenow. Elongate and segmented, with a single
170]
The Classification of Lower Organisms
Fig. 32. — Hypermastigina : a-d, Trichonympha Campanula after Kofoid &
Swezy (1919); a, cell x 250; b, division of centroblcpharoplast and formation of
paradesmose, and c and d, earlier and later stages of mitosis x 500. e, f, g, Sperm,
egg, and fertilization of Trichonympha sp. from the roach Cryptocercus after Cleve-
land (1948). h, Hoplonympha Natator x 250 after Light (1926). i, Staurojoenina
assimilis x 250 after Kirby (1926). j, Tcratonympha mirabilis after Koidzumi
(1921).
Phylum Protoplasta [171
nucleus in the anterior segment; flagella distributed generally on the surface, most
abundant on an anterior beak. Teratonympha Koidzumi {Cyclonympha Dogiel), a
single known species in Reticulitermes.
Class 2. MYCETOZOA de Bary
Order Dermatocarpi Persoon Syst. Meth. Fung, xiii (1801), in part.
Suborder Myxogastres Fries Syst. Myc. 3: 3 (1829); suborder Trichospermi Fries
op. cit. 1 : xlix (1832), in part.
Suborder MyATomyce^^j Link 1833.
Mycetozoen de Bary in Bot. Zeit. 16: 369 (1858); Zeit. wiss. Zool. 10: 88 (1859).
Stamm Myxomycetes Ylatcktl Gen. Morph. 2: xxvi (1866).
Class Mycetozoa de Bary ex Rostafinski Versuch Systems Mycetozoen 1 (1873).
Division Mycetozoa and classes Myxogasteres and Phytomyxini Engler and Prantl
Nat. Pflanzenfam. IITeil: 1 (1888).
Division Myxothallophyta Engler in Engler and Prantl Nat. Pflanzenfam. I Teil,
Abt. 1: iii (1897).
Stamm Myxophyta Wettstein Handb. syst. Bot. 1: 49 (1901).
Division Phytosarcodina, Myxothallophyta, or Myxomycetes Engler Syllab. ed. 3:
1 (1903).
Division Myxomycophyta Tippo in Chron. Bot. 7: 205 (1942).
Order Mycetozoida Hall Protozoology 227 (1953).
Organisms whose walled resting cells produce in germination anteriorly unequally
biflagellate cells; these giving rise to bodies called plasmodia, being multinucleate
bodies of amoeboid character.
1. Predatory, subaerial, producing macroscopic
spore-bearing fruits.
2. Spores produced within the fruits Order 1. Enteridiea.
2. Spores produced on the surfaces of the
fruits Order 2. Exosporea
1. Parasitic, not producing definite fruits Order 3. Phytomy.xii>a.
Order 1. Enteridiea [Enteridieae] Rostafinski Vers. 3 (1873).
Cohort Endosporeae and orders Anemeae, Heterodermeae, Reticularieae, Ain-
aurochaeteae, Calcareae, and Calonemeae Rostafinski op. cit.
Order Endosporea Lankester in Enc. Brit. ed. 9, 19: 840 (1885).
Orders Protodermieae and Columniferae Rostafinski ex Berlese in Saccard)
Sylloge7: 328,417 (1888).
Cohorts Amaurosporales and Lamprosporales, with numerous orders with names
in -aceae. Lister Monog. Mycetozoa 21-23 (1894).
Subclass Myxogastres and orders Physaraceae, Stemonitaceae, Cribrariaceae,
Lycogalaceae, and Trichiaceae Macbride North American Slime Molds 20
(1899).
Subsuborder (!) Endosporinei Poche in Arch. Prot. 30: 200 (1913).
Orders Physarales, Stemcnitales, Cribrariales, Lycogalales, and Trichiales Mac-
bride op. cit. cd. 2 (1922).
Order Liceales Ma. bride and Martin (1934).
Suborder Eumycetozoina Hall Protozoology 230 (1953).
172 ] The Classification of Lower Organisms
Predatory Mycetozoa producing macroscopic fruits, these producing internal uni-
nucleate spores. The type is Lycogala, the sole genus of the order as originally
published.
The fruits of many examples are of the appearance of minute puffballs, and Per-
soon and Fries classified them as puffballs; Fries took note that they are primitus
mucilaginosi and made them a suborder distinct from the proper pufTballs. De Bary
studied the non-reproductive stages; concluded "dass die Myxomyceten nicht dem
Pflanzenreiche angehoren, sondern dass sie Thiere, und zwar der Abtheilung der
Rhizopoden angehorig, sind"; and renamed the group Mycetozoen. This name was
apparently first published in Latin form, in the category of classes, by de Bary's stu-
dent Rostafinski. Conventional botany continues to list Myxomycetes as a class of
Fungi; conventional zoology makes the group an order of Rhizopoda or Sarcodina.
The spores germinate readily in water or appropriate solutions (Jahn, 1905; Gil-
bert, 1929; Smith, 1929). Their nuclei usually divide once or twice, during or just
after germination; thus each spore produces from one to four naked cells.
It is in germinating spores that mitosis is most easily observed. Mitosis takes place
in a clear area, about which some observers have found a persistent nuclear membrane.
The spindle is sharp-pointed. Only a few observers (as Skupienski, 1927) have dis-
cerned definite centrosomes. When the one or two divisions associated with germina-
tion are complete, the flagella grow forth from the areas of the poles of the mitotic
spindle. All earlier observers described the spores as uniflagellate, but Ellison (1945)
and Elliott (1949) found them biflagellate. The flagella may be apparently equal or
moderately unequal; or one of them may be very brief. Each nucleus remains con-
nected to the base of the flagella by a conical body of clear protoplasm, the Geissel-
glocke of Jahn (Jahn, 1904; Howard, 1931).
The flagellate cells are not spores, but gametes; they fuse with each other. Skupien-
ski (1917) affirms that they are of two mating types. Fusion is at first by pairs, and
Howard (1931) found that each zygote develops into a plasmodium by itself. All
other observers (de Bary, 1858, 1859; Cienkowski, 1863; Skupienski, 1917, 1927;
Schiinemann, 1930) have found the zygotes to fuse with each other and with further
gametes. The flagella are lost. The nuclei fuse in pairs; those which fail to find
partners are digested.
The cell formed by the fusion of zygotes and gametes is a young plasmodium. The
term was coined by Cienkowski ( 1863, p. 326) : "Das Protoplasmanetz der Myxomy-
ceten werde ich mit den Namen Plasmodium bezeichnen." The plasmodium nour-
ishes itself in predatory fashion, on fungus spores, bacteria, and other digestible ob-
jects, and grows accordingly. Mitosis occurs simultaneously in all nuclei of the plas-
modium, and takes 20 to 40 minutes; it has accordingly only rarely been observed
(Lister, 1893; Howard, 1932). Plasmodia do not ordinarily divide, but grow to great
sizes. They are not very familiar objects because during most of their life they keep to
dark and moist places, chiefly among vegetable remains. Drouth does not kill them;
they can become dry and hard while retaining the capacity to resume activity upon
the return of moisture. When an active plasmodium reaches a certain stage, its re-
actions change; it moves out into the light and to dry places. A plasmodium in this
stage is conspicuous, being of the form of a network which may be many centimeters
in diameter, in some species brilliantly colored. The whole is a single naked protoplast.
Each Plasmodium proceeds to produce a fruit or fruits. The entire mass may heap
itself up, or it may break up into portions, large or minute. In species whose plas-
modia break up into small fragments, each of these may secrete a column of lifeless
Phylum Protoplasta [173
material, a millimeter or more in height, and ascend upon it. Each separate body of
protoplasm secretes an external wall and begins to undergo cleavage within it. Har-
per (1900) described the details of the process. All authorities agree that the nuclei
undergo a flare of divisions at this time (Strasburger, 1884; Harper, 1900, 1914;
Bisby, 1914). It is almost certain that there are two flares of division, constituting the
meiotic process, but few authorities have positively affirmed this (Schiinemann,
1930)1. Cleavage is carried to the point of producing uninucleate protoplasts. While
this is taking place, many species secrete a network of hollow tubes or a system of
hollow fibers, called the capillitium, by deposition of lifeless material outside the
cell membranes. In species which produce a true capillitium, all of the uninucleate
protoplasts secrete walls and become spores. Strasburger found the capillitium and
the walls of the spores to consist of impure cellulose; others have found no cellulose.
In many species which do not produce a true capillitium, an analogous structure
called a pseudocapillitium, consisting of solid bodies of various forms, is modelled
from a part of the nucleate protoplasm which is deprived of its reproductive function
and killed. In many species, much calcium carbonate is deposited in the wall, or
both in the wall and in the capillitium or pseudocapillitium.
A small separate fruit is called a sporangium. A fruit of the form of a large mass,
or of many sporangia not completely separate, is an aethalium. The spores are re-
leased by collapse of the outer wall.
These organisms are of no known economic importance. There are some forty
genera, between four and five hundred species. As Lister remarked, the same species
occur everywhere: collections from Colombia (Martin, 1938) and from Mount
Shasta (Cooke, 1949) consist entirely of familiar species.
Rostafinski (1873) arranged the genera in two cohorts, seven orders, and nineteen
tribes, the last with names in -aceae. His subsequent monograph of the group ( 1875)
was regrettably published in a barbarous language, and is for nomenclatural purposes
a nullity. All later systems are based on Rostafinski's original system. The group being
essentially uniform, it is properly treated as a single order.
Definite families were first established by Lankester, mostly under names which
Rostafinski had applied to tribes. Berlese (in Saccardo, 1888) provided a complete
set of names in -aceae, valid under botanical rules; Poche provided a complete set in
-idae, valid under zoological rules. Authorities have differed moderately as to the
list of families; here, somewhat arbitrarily, fourteen are maintained.
1. Capillitium none (order Cribrariales Mac-
bride).
2. Producing separate sporangia, pseudo-
capillitium none.
3. Sporangia shattering irregularly or
opening through a terminal oper-
culum Family 1 . Liceacea.
3. Sporangia opening through numer-
ous pores, the walls becoming sieve-
like Family 2. Cribrariacea.
2. Fruits aethalioid, pseudocapillitium
present.
iWhile the present work was in proof, Wilson and Ross (1955) established the point
that meiosis occurs immediately before the formation of spores.
174 ] The Classification of Lower Organisms
3. Aethalia consisting of more or less
separate sporangia.
4. Sporangia tubular, opening
through terminal pores Family 3, Tubiferida.
4. Sporangia indistinct, their walls
becoming freely punctured and
converted into a reticulate
pseudocapillitium Family 4. Retigulariacea.
3. Aethalia not consisting of distin-
guishable sporangia Family 5. Lycogalacttoa.
1. Capillitium present.
2. Fruits without considerable deposits of
calcium carbonate.
3. Spores black or dark, capillitial
hairs smooth (order Stemonitales
Macbride).
4. Fruits aethalioid, capilHtium
poorly defined, without a cen-
tral axis Family 6. Amaurochaetacea.
4. Fruits of separate sporangia
with a definite capillitium in-
cluding a central axis (colu-
mella).
5. Capillitium spreading hor-
izontally from the colu-
mella Family 7. Stemonitea.
5. Capillitium spreading
chiefly from the summit of
the columella Family 8. Enerthenemea.
3. Spores pallid or yellow (order Tri-
chiales Macbride).
4. Capillitial hairs smooth, un-
branched or sparsely branched.
5. Capillitial threads hori-
zontal, attached at both
ends Family 9. Margaritida.
5. Capillitial threads run-
ning at random, not at-
tached at the ends Family 10. Perichaenacea.
4. Capillitium reticulate, sculp-
tured, but not with spiral bands. . .Family 11. Arcyriagea.
4. Capillitial threads unbranched
or sparsely branched, sculp-
tured with spiral bands Family 12. Trighiagea.
2. Fruits containing considerable deposits
of calcium carbonate (order Physarales
Macbride).
3. Calcium carbonate both in walls
and in capillitium Family 13. Physarea.
Phylum Protoplasta [175
3. Calcium carbonate in walls but not
in capillitium Family 14. DrovMiACEA.
Family 1. Liceacea [Liceaceae] (Rostafinski) Lankester in Enc. Brit. ed. 9, 19:
841 (1885). Tribe Liceaceae Rostafinski Vers. 4 (1873). Order Liceaceae Lister
Monog. Mycetozoa 149 (1894). Family Liceidae Doflein 1909. Family Orcadel-
lidae Poche in Arch. Prot. 30: 200 (1913). Family Orcadellaceae Macbride N. Am.
Slime Molds ed. 2: 203 (1922). Sporangia separate, sessile or stalked, without capil-
litium or pseudocapillitium, the walls shattering irregularly or opening by means of
a terminal operculum. Licea, Orcadella.
Family 2. Cribrariacea [Cribrariaceae] (Rostafinski) Lankester 1. c. Tribe Crib-
rariaccac Rostafinski op. cit. 5. Order Cribrariaceae Macbride N. Am. Slime Molds
20 (1899). Order Heterodermaceae Lister op. cit. 136. Family Cribrariidae Poche
1. c. The wall of the stalked fruit becoming sieve-like. Cribraria. Dictydium.
Family 3. Tubiferida [Tubiferidae] Poche in Arch. Prot. 30: 200 (1913). Order
Tubulinaceae Lister op. cit. 152 (1894). Family Tubulinidae Doflein 1909. Family
Tubiferaceae Macbride in N. Am. Slime Molds ed. 2: 203 (1922). Aethalia consist-
ing of tubular sporangia opening through terminal pores. Tubifer (its older name
Tubulina preoccupied), Lindbladia, Alwisia.
Family 4. Reticulariacea [Reticulariaceae] (Rostafinski) Lankester 1. c. Tribes
Dictydiaethaliaceae and Reticulariaceae Rostafinski op. cit. 5, 6. Order Reticularia-
ceae Lister op. cit. 156. Family Dictydiaethaliidae Poche I.e. Aethalia of indistinct
sporangia whose walls become porous and are converted into a reticulate pseudo-
capillitium. Reticularia, Dictydiaethallium, etc.
Family 5. Lycogalactida [Lycogalactidae] Poche in Arch. Prot. 30: 201 (1913).
Tribe Lycogalaceae de Bary. Order Lycogalaceae Macbride N. Am. Slime Molds 20
(1899). Y di.m.i\y Lycogalaceae Macbride and Martin Myxomycetes (1934). Aethalia
with a pseudocapillitium, not divided into sporangia. Lycogala, the brownish fruits a
few millimeters in diameter clustered on wood, of much the appearance of small
puffballs.
Family 6. Amaurochaetacea [Amaurochaetaceae] (Rostafinski) Berlese in Sac-
cardo Sylloge 7: 401 (1888). Tribe Amaurochaetaceae Rostafinski op. cit. 8. Order
Amaurochaetaceae Lister op. cit. 134. Family Amaurochaetidae Doflein 1909.
Fruits aethalioid with dark spores and a poorly defined capillitium without a central
axis. Amaurochaete.
Family 7. Stemonitea Lankester in Enc. Brit. ed. 9, 19: 841 (1885). Tribes Stemo-
nitaceae and Brefeldiaceae Rostafinski op. cit. 6, 8. Families Stemonitaceae and
Brefeldiaceae Berlese in Saccardo op. cit. 390, 402. Order Stemonitaceae Macbride
N. Am. Slime Molds 20 (1899). Family Stemonitidae Doflein 1909. Families Bre-
feldiidae and Stemonitidae Poche op. cit. 202. Sporangia with dark spores and a
capillitium of smooth threads spreading from a central axis, the columella. Stemo-
nitis, comm.on, the clustered stalked fruits of the appearance of minuscule dark
bottle-brushes. Brefeldia, Comatricha; Diachea, exceptional in containing much lime
in the stalk and wall.
Family 8. Enerthenemea Lankester 1. c. Tribes Echinosteliaceae and Enerthene-
maceae Rostafinski op. cit. 7, 8. Families Echinosteliaceae and Enerthenemaceae
Berlese in Saccardo op. cit. 389, 402. Family Lamprodermaceae Macbride N. Am.
Slime Molds ed. 2: 189 (1922). Like Stemonitea, in which this family has usually
been included, but the capillitium attached chiefly at the summit of the columella.
Enerthenema, Clastoderma, Lamproderma, Echinostelium.
176]
The Classification of Lower Organisms
Family 9. Margaritida [Margaritidae] Doflein 1909. Order Margaritaceae Lister
op. cit. 202. Family Dianemaceae Macbride N. Am. Slime Molds ed. 2: 237 (1922).
Sporangia with pale or yellow spores and a capillitium of smooth threads attached at
both ends. Dianema, Margarita.
Family 10. Perichaenacea [Perichaenaceae] (Rostafinski) Lankester 1. c. Tribe
Perichaenaceae Rostafinski op. cit. 15. Sporangia with pale or yellow spores and a
capillitium of unattached smooth threads. Perichaena, Ophiotheca.
Family 11. Arcyriacea [Arcyriaceae] (Rostafinski) Lankester 1. c. Tribe Arcyri-
aceae Rostafinski op. cit. 15. Order Arcyriaceae Lister op. cit. 182. Family Arcyriidae
Doflein 1909. Sporangia with pale or yellow spores and a reticulate capillitium,
usually sculptured, but not with spiral bands. Arcyria, Lachnobolus.
Family 12. Trichiacea [Trichiaceae] (Rostafinski) Berlese in Saccardo Sylloge 7:
437 (1888). Tribe Trichiaceae Rostafinski op. cit. 14. Family Trichinaceae Lankes-
Fig. 33. — Mycetozoa. a-f, Spore, germination, gametes, syngamy, and zygote of
Physarum polycephalum after Howard (1931) x 1,000. g-1, Stages of mitosis in the
Plasmodium of Physarum polycephalum after Howard ( 1932) x 2,000. m-o. Stages
of mitosis in the plasmodium of Trichia after Lister (1893) x 1,000. p, Cleavage
in the developing fruit of Physarum polycephalum after Howard (1931) x 1,000.
q, Capillitium and spores of Lepidoderma Chailletii x 1,000. r-W, fruits of Myce-
toza X 5; r, Sternonitis splcndens; s, Lycogala cpidcndrum; i, Lcocarpus fragilis;
U, Lepidoderma Chaillettii; v, Physarum notabile; w, Hemitrichia intorta.
Phylum Protoplasta [177
ter 1. c; the genus Trichina does not belong to this family! Order Trichiaceae Mac-
bride N. Am. Slime Molds 20 (1899). Family Trichiidae Doflein 1909. Sporangia
with pale or yellow spores, the capillitium of free threads, unbranched or sparsely
branched, marked with spiral bands. Trichia, Hemitrichia, Oligonema, Calonema.
Family 13. Physarea Lankester 1. c. Tribes Cienkowskiaceae, Physaraceae, and
Spumariaceae Rostafinski op. cit. 9, 13. Families Cienkowskiaceae , Physaraceae, and
Spumariaceae Berlese in Saccardo op. cit. 328, 329, 387. Order Physaraceae Macbride
N. Am. Slime Molds 20 (1899). Family Physaridae Doflein 1909. Fruits sporangial
or aethalioid, with capillitium, both wall and capillitium containing considerable
deposits of calcium carbonate. Physarum, with some seventy-five species, is the most
numerous genus of Mycetozoa; the little gray sporangia may be spherical or irregular,
sessile or stalked. Fuligo septica produces dirty yellow aethalia reaching several cen-
timeters in diameter on vegetable trash; observed on spent tan bark, it has the com-
mon name of flowers of tan. Badhamia, Craterium, Leocarpus, Chondrioderma,
Spumaria, etc.
Family 14. Didymiacea [Didymiaceae] (Rostafinski) Lankester 1. c. Tribe Didy-
miaceae Rostafinski op cit. 12. Order Didymiaceae Lister op. cit. 93. Family Didy-
midae Doflein 1909. Family Didymiidae Poche op. cit. 202. Family Collodermata-
ceae Macbride and Martin Myxomycetes 145 (1934). Sporangia with deposits of
calcium carbonate in the wall and a simple capillitium free of mineral deposits.
Didymium, Leangium, Lepidoderrna, Colloderma.
Order 2. Exosporea (Rostafinski) Lankester in Enc. Brit. ed. 9, 19: 841 (1885).
Cohors Exosporeae Rostafinski Vers. 2 (1873).
OrAtr Ectosporeae Y.ng\tr ?>y\\2ih. 2 (1892).
Order Ceratiomyxaceae (Schroter) Lister Monog. Mycetozoa 25 (1894).
Subsuborder (!) Exosporinei Poche in Arch. Prot. 30: 200 (1913).
Organisms of much the character of the Enteridiea, but the spores forming a single
layer on the surface of the fruits. There is a single family with only one well-marked
species.
Family Ceratiomyxacea [Ceratiomyxaceae] Schroter (in Engler and Prantl, 1889).
Ceratiomyxa Schroter [Ceratium Albertini and Schweinitz, 1805, non Schrank, 1793);
C. fruticulosa (O. F. Miiller) Macbride. The fruits are white pillars, sometimes
branched, 1-2 mm. tall, of secreted material. Each spore of the single superficial
layer generates a microscopic stalk and ascends upon it before becoming walled.
Meiosis then takes place, making the spores 4-nucleate; the chromosome number is
cut from 16 to 8 (Gilbert, 1935). In germination, the contents of the spore are re-
leased as a single amoeboid protoplast, whose nuclei divide once; the cell then divides
into eight, and these generate flagella (Rostafinski, 1873; Jahn, 1905; Gilbert, 1935).
Order 3. Phytomyxida Calkins Biol. Prot. 330 (1926).
Class Phytomyxini Engler and Prantl Nat. Pflanzenfam. II Teil: 1 (1889); class
Phytomyxinae op. cit. I Teil, Abt. 1: iii (1897).
Order Phytomyxinae Campbell Univ. Textb. Bot. 71 (1902).
Class Plasmodiophorales Engler Syllab. ed. 3: 1 (1903).
Order Plasmodiophorales Sparrow in Mycologia 34: 115 (1942).
Suborder Plasmodia phorina Hall Protozoology 228 (1953).
Intracellular parasites chiefly of higher plants, attacking also algae, Oomycetes,
and beetles, being naked multicellular plasmodia producing walled resting cells,
1781
The Classification of Lower Organisms
the walls containing no cellulose; these releasing naked infective colls with paired
unequal simple flagella.
This inconsiderable group was made known by the discovery of Plasmodiophora
Brassicae, the agent of the clubroot disease of cabbage, by Woronin (1878). The
proper place of the group in classification has been a puzzle; some students treat it
as a class of myxomycetes, others as an order of chytrids. The known characters —
paired unequal simple flagella; cells naked in the vegetative condition; and non-pro-
duction of cellulose — assure us that this group has nothing to do with proper chytrids,
nor with Oomycetes of chytrid body type. The traditional association with myxomy-
cetes is tenable. Alternatively, the group would not be out of place next to order
Rhizoflagellata (anyone who chooses to put it there should take note that the class
name Phytomyxini is older than Zoomastigoda).
The Plasmodium causes often much hypertrophy of the host tissue. In some forms
the mature plasmodium becomes walled; the protoplast undergoes cleavage into uni-
nucleate portions; these become swimming cells and are released through a discharge
tube. These forms are of much the appearance of Lagenidialea. In the majority of
the group the naked plasmodium undergoes cleavage; the resulting protoplasts be-
come walled; the resulting spores or cysts, released by decay of the host, discharge
their contents as one or two swimming cells. Ledingham ( 1939) and Sparrow ( 1947)
report both types of development as occurring in Polyniyxa. Karling (1944) found
the walls to contain no cellulose. Ellison (1945) found the flagella to be simple.
iKyuiU/Tnuiwuium/^iiiMiivnirm-
Fig. 34. — Ceratiomyxa jruticulosa. a. Fruits x 5. b-q, reproductive processes
after Gilbert (1935); b, young spores on the surfaces of the fruit; c, d, the same
raised on stalks; e^ f, heterotypic division; g, homeotypic division; h, the mature spore
ou its stalk; i-n, germination and subsequent processes: the amoeboid protoplast
passes through a "thread stage" before rounding up and dividing into four and then
into eight; o, production of flagcllum; p, "zoospore" (gamete); q, gametes fusing to
initiate the plasmodium. All x 1,000 except Fig. a.
Phylum Protoplasta [179
In the growing plasmodium, a nucleus which is not dividing contains an endosome
("nucleolus"). During mitosis, which occurs within the intact nuclear membrane,
the endosome becomes elongate, and a ring of chromatin, within which separate
chromosomes have not been distinguished, forms about its middle. The resulting "cru-
ciform" figure resembles some which have been seen in trypanosomes. The nuclear
divisions which occur immediately before cleavage are of a different character: no en-
dosome is seen, but there is a spindle with centrosomes at the poles, and definite
chromosomes are present. The occurrence of these two types of nuclear division has
been noted by every careful observer, Schwartz (1914), Home (1930), Cook (1933),
Ledingham (1939), and Karling (1944). Home was probably correct in supposing
the divisions which precede cleavage to be meiotic. Conjugation of the flagellate cells
of Spongospora has been observed.
There are monographic accounts of the Phytomyxida by Cook ( 1933) and Karling
(1942). The group may be treated as a single family with a dozen genera and about
twenty-five species.
Family Plasmodiophorea [Plasmodiophoreae] Berlese in Saccardo Sylloge 7 : 464
(1888). Family Plasmodiophoreen Zopf Pilzthiere 129 (1885). Family Plasmodio-
phoraceae Engler Syllab. 1 (1892). Family Woroninaceae Minden 1911. Families
Phytomyxidae and Woroninidae Poche in Arch Prot. 30: 198 (1913). Plasmodio-
phora, Polymyxa, Spongospora, and Sorosphaera attack land plants; Tetramyxa,
Ligniera, and Sorodiscus, chiefly aquatic seed plants; Woronina and Octomyxa,
Oomycetes; Phagomyxa, brown algae; Sporomyxa (Leger, 1908) and Mycetosporid-
ium, beetles.
Class 3. RHSZOPODA Siebold
Order Foraminiferes d' Orbigny in Ann. Sci. Nat. 7: 128, 245 (1826).
Order Foraminifera Zborewski 1834.
Rhizopodes Dujardin in Compt. Rend. 1: 338 (1835).
Class Foraminifera d'Orbigny in de la Sagra Hist. Cuba vol. 8 (1839).
Order Polythalamia Ehrenberg in Abh. Akad. Wiss. Berlin (1838) : table 1 ( 1839) .
Class Rhizopoda and orders Monosomatia and Polysomata Siebold in Siebold
and Stannius Lehrb. vergl. Anat. 1 : 3, 11 (1848).
Reticulosa Carpenter 1862.
Stamm Rhizopoda and Class Acyttaria Haeckel Gen. Morph. 2: xxvii (1866).
Thalamophora R. Hertwig Hist.Radiolar. 82 (1876).
Class Reticidaria Lankester in Enc. Brit. ed. 9, 19: 845 (1885).
Order Reticulosa Poche in Arch. Prot. 30: 203 (1913).
Order Granuloreticulosa de Sacdeleer in Mem. Mus. Roy. Hist. Nat. Belgique 60:
7 (1934).
Order Foraminiferida Hall Protozoology 250 (1953).
Amoeboid organisms, the pseudopodia of the character of rhizopodia, i.e., fine,
freely branching and anastomosing; producing shells, these usually calcareous; com-
monly reaching macroscopic dimensions; mostly marine.
The first examples of rhizopodes mentioned by Dujardin were milioles, vorticiales,
and le gromia: the genus Miliola is to be construed as the type. These organisms, the
proper rhizopods, are in general usage called Foraminifera, but that name was orig-
inally applied in the categoiy of orders.
180]
The Classification of Lower Organisms
Fig. 35. — Life cycle of "Tretomphalus," i.e., Discorbis or Cymbalopora, from
Myers (1943); 1-3, microspheric individuals, in 3 releasing young megalospheric
individuals; 4-8 megalospheric individuals; 9-12, gametes and syngamy.
Phylum Protoplast a [181
The individual rhizopod originates as a minute amoeboid cell which secretes a
shell from which the pseudopodia project. In the fresh-water forms, each protoplast,
after moderate growth, divides into two, one of which retains the original shell while
the other secretes a new one. In some of the marine forms, the original protoplast,
having a cylindrical or irregular shell, enlarges this as it grows. In the great majority
of the group, the original shell, called the proloculus, is of definite size and form and
has a constricted orifice. When the protoplast reaches a certain stage, it expands, pro-
trudes from the orifice, and secretes an extension of the shell in the form of a second
chamber. In some few examples, the second chamber is the final one, being capable
of indefinite extension. But again in the great majority, the second chamber, although
diff'erent from the proloculus, resembles it in being definite in form and in having a
constricted orifice. After further development, the protoplast again protrudes through
the orifice and secretes a third chamber, generally of the same form as the second,
though often larger. Repetition of this process produces macroscopically visible
bodies. Even though becoming a centimeter or more in diameter, the individuals
continue to be single cells.
As a result of different patterns of growth, the developed shells are of highly varied
forms, linear, globular, or coiled in one plane; trochoid or rotaloid, that is, helical,
of the form of a low cone; of the form of high cones; or screw-like, with the chambers
in fixed longitudinal rows. The grov/th pattern may change during the life of the
individual. There are apparently degenerate forms, simple or irregular. It is highly
probable that some of the forms have evolved repeatedly.
The shells may be of gelatinous material or of chitin, without or with imbedded
grains of sand. Exceptionally, they are siliceous. They are sometimes of crystallized
calcium carbonate with imbedded grains of sand. In the bulk of the group they consist
of crystallized calcium carbonate without foreign matter, and are of either of two
t>'pes of texture: vitreous, that is, hyaline, and punctured by numerous pores a few
microns in diameter; or porcellanous, white by reflected light and amber by trans-
mitted light, and with no perforations except the proper orifices. In fossil shells, other
textures than these may occur; it is supposed that these are products of modification
during preservation. Some of the textures, like some of the forms, are believed to
have evolved repeatedly.
Most rhizopods occur in two forms which are most readily distinguished by the
size of the proloculi. This was first pointed out by Munier-Calmas, 1880; who,
jointly with Schlumberger, 1885, designated the smaller and larger proloculi re-
spectively microsperes and megaspheres. Lister (1895), by study in culture of Elphi-
dium crispiirn [Polystomella crispa Lamarck), showed that the two forms are alter-
nate generations. He observed that the microspheric cells become multinucleate
during growth, while the megalospheric cells remain uninucleate until just before
reproduction. The reproduction of the megalospheric cells is by release of numerous
minute biflagellate cells.
Schaudinn (1902) confirmed much of what Lister had observed. He was mistaken
in describing nuclear division (except just before the production of the swimming
cells) as non-mitotic; and correct in identifying the swimming cells as gametes.
Winter (1907) observed a similar life cycle in Peneroplis, but described the gametes
as having solitary flagella.
Myers" (1934, 1935, 1936), dealing with Patellina and Spirillina, described the
details of mitosis. This takes place within an intact nuclear membrane, and is com-
pleted by its constriction. The spindle is blunt-ended; there is no evidence of centre-
182 ] The Classification of Lower Organisms
somes. The chromosomes are numerous, long, and slender; the mitotic figures re-
semble those of Pyrrhophyta. Reduction of the chromosome number is said to be
effected by a single nuclear division, the last one before the formation of gametes,
which cuts the chromosome number of Patellina from 24 to 12, and that of Spirillina
from 12 to 6. Before they reach this stage, the megalospheric individuals have
gathered themselves in clusters of two or more within cyst walls consisting of secreted
gelatinous matter and scraps from the neighborhood. Gametes from one individual
are unable to unite with each other. The gametes are amoeboid, positively without
flagella. In Discorbis and Cymbalopora, however, Myers (1943) observed the produc-
tion of biflagellate gametes.
Le Calvez (1950) has cleared up various questions raised by earlier studies. Some
forms, as Discorbis orbicularis, appear to lack a sexual cycle. Patellina and Spirillina
produce amoeboid gametes 40-50[.i in diameter. Most rhizopods produce biflagellate
gametes 1.5-4[i long. Le Calvez found the flagella definitely unequal. In Discorbis
mediterranensis he showed that the megalospheric individuals are of two mating
types. Earlier zoologists, apparently misled by familiarity with the normal life cycle
cf animals, had identified meiosis as occurring at the time of gametogenesis; it is the
fact, on the contrary, that it occurs in the last two nuclear divisions in the micro-
spheric individuals. The megalospheric and microspheric stages of rhizopods are
respectively haploid and diploid, like the gametophytes and sporophytes of plants.
With the possible exception of some of the one-chambered fresh water forms, the
rhizopods are clearly a natural group. The fresh water forms appear to intergrade
with organisms which Pascher identified as chrysomonads.
The shells of dead rhizopods may under appropriate conditions be preserved
through geologic ages. Natural chalk consists of shells of Textularia mixed with coc-
coliths. Certain forms of limestone consist chiefly of shells of Miliola. Certain fossil
rhizopods have long been known as indicators of division of geologic time. Since about
1917, it has been found that the whole group offers one of the beautiful illustrations
of evolution as related to geologic time: the shells of rhizopods found under magnifi-
cation in a particular stratum serve promptly and precisely to identify it. The services
of experts on "Foraminifera" have acquired a high economic value in the petroleum
industry: these experts have found themselves promoted from the status of pure
biologists to that of economic geologists.
Among some eleven hundred genera which have been published, Galloway (1933)
maintains 542. Of the number of species one can only say that it is a matter of
thousands, but probably not many tens of thousands. Economic micropaleontologists
find themselves dealing with great numbers of forms which are slightly, yet signifi-
cantly, distinct. They find it expedient not to name these, but to identify them by
comparison with available collections.
Some of the marine and fossil forms are similar, on a small scale, to the animal
Nautilus, and Linnaeus placed some of them in that genus. Montfort and Lamarck
treated them as several genera of mollusks. In first distinguishing these organisms as
the order Foraminiferes of class Cephalopodes, d'Orbigny intended to contrast them
with Nautilus, in whose shells a series of chambers arc connected, not by holes (fora-
mina) but by cylindrical tubes. Dujardin ( 1835) found that his Rhizopodes are with-
out definite organs. Their shells enclose a clear semiliquid substance; their apparent
tentacles are merely temporary structures, formed of this substance, thrust forward
in the direction of the movement of the shell and withdrawn as it advances. Dujardin
named this substance sarcode; it is, of course, the same which has since been called
Phylum Protoplasta [ 183
protoplasm. The effect of his discoveries was to show that the rhizopods or Foramini-
fera are not mollusks, but one-celled organisms.
Very much taxonomic study has been given to this interesting group. The standard
system, in the modern period of practical concern with the group, has been that of
Cushman (1928).
Galloway (1933), attempting to recognize phylogeny and concluding that certain
types of form and texture of shells have evolved repeatedly, has radically revised
Cushman's system and set up a system of thirty-five families. The following survey
of the group is based on Galloway's system. The names applied to the families are
those which he has cited as the oldest, and the groups treated as orders are the blocks
of families which to him appeared natural.
1. Shell one-chambered, or of a proloculus fol-
lowed by one other chamber, not of a series
of similar chambers Order 1. Monosomatia.
1. Shell a series of similar chambers.
2. Shell porcellanous, imperforate Order 2. Miliolidea.
2. Not as above.
3. Not specialized as in the following
orders Order 3. Foraminifera.
3. Shell hyaline, perforate, typically
trochoid, i.e., having the succes-
sively larger chambers helically ar-
ranged so that all may be seen from
one side and only the last whorl
from the other Order 4. GLOBiGERiNroEA.
3. Chambers of the fundamentally
planispiral shell with specialized
walls containing channels or pro-
ducing chamberlets Order 5. Nummulitinidea.
Order 1. Monosomatia (Ehrenberg) Siebold in Siebold and Stannius Lehrb. vergl.
Anat. 1: 11 (1848).
Monosomatia Ehrenberg in Abh. Akad. Wiss. Berhn (1838) : table 1 (1839).
Order Astrorhizidea Lankester in Enc. Brit. ed. 9. 19: 846 (1885).
Order Imperforida Delage and Herouard Traite Zool. 1: 107 (1896).
Order Archi-Monothalamia Calkins Biol. Prot. 354 ( 1926) .
Rhizopoda consisting of a single chamber, or of a proloculus followed by one other
chamber; exceptionally, after passing through a stage of this character, producing
a series of similar chambers.
Family 1. Allogromiida [Allogromiidae] Cash and Wailes. Minute, with one-
chambered chitinous or gelatinous shells, usually subglobular; large in fresh water.
Allogromia Rhumbler; Mikrogromia Hertwig, the pseudopods of sister cells retaining
contact so that small colonies are formed; etc.
Family 2. Astrorhizida [Astorhizidae] Brady (1881). Family Astrorhizina Lankes-
ter (1885). Family Astrorhizidaceae Lister. Families Rhizamminidae , Saccammini-
dae, and Hyperamminidae Cushman. Shell of agglutinated foreign material, usually
elongate, often branched, but not coiled. In Astrorhiza there is a central chamber
from which grow elongate arms. In Rhizammina, the shell is tubular, open at both
ends; in Bathysiphon it is a tube closed at one end; in Hyperammina a proloculus is
formed before the extended tube.
184]
The Classification of Lower Organisms
- ^ / m\^■^V^//'":^^^^^.^V^\\■•\\
Fig. 36 — Shells of Rhizopoda. a, Ophthalimidium. h, c, Triloculina. d, Verte-
bralina. e, Peneroplis. f, Archaias x 25. g, Nodosaria. h, Dcntilina. i, Flabel-
lina. j, Lagena. k, 1, Nonion. m, n, Rotalia. o, Globigerina. x 50 except as
noted.
Phylum Protoplasta [ 185
Family 3. Spirillinidea Reuss 1861. Family Spirillinina Lankester (1885). Family
Silicinidae Cushman. In Spirillina, the perforate hyaline one-chambered shell is
planispirally coiled; the family is distinguished by a shell of this form in the young
stages if not throughout life. Silicina is a Jurassic fossil whose shell is silicified. In
Patellina the spirally coiled first chamber is followed by others arranged in a heHx.
Family 4. Ammodiscida [Ammodiscidae] Rhumbler 1895. Like the preceding
family, but the shell consisting of agglutinated foreign matter. Ammodiscus etc.
Order 2. Miliolidea Lankester in Enc. Brit. ed. 9, 19: 846 (1885).
Order Flexostylida Calkins Biol. Prot. 355 (1926).
Rhizopoda with imperforate porcellanous shells, a numerous and important group.
Family 1. Miliolida [Miliolidae] d'Orbigny (1839). Families N uhecularina, Milio-
lina, and Hauerinina Lankester (1885). Fisherinidae Cushman. The genus Cornu-
spira, known from the carboniferous, differs from Spirillina only in the texture. Evi-
dently evolved from this are genera of planispirally coiled tubes divided into chambers,
and from these others in which the series of chambers becomes straight or irregular,
as in Vertebralina and Tubinella. There is an important block of genera in which each
cycle of chambers is of two members, the second opening at the opposite end of the
body from the first. In O phthalmidium and Pyrgo alternate chambers lie regularly
on opposite sides of a body whose form is that of an elliptic flake. In other genera
of this group successive chambers are not opposite each other, but separated by less
than 180°, so that more than two appear on the outside. In Triloculina three cham-
bers are externally visible. In Miliola Lamarck [Miliolina Lamarck, the latter name
applied to fossil representatives of the same genus) the chambers are 144° apart, so
that five appear on the outside. In many members of the family the apertures are
partially blocked by teeth, single, double, or multiple, or extended as bars clear across.
Family 2. Soritina Ehrenberg (1839). Family Helicosorina Ehrenberg op. cit.
Family Peneroplidea Reuss 1861. Family Peneroplidina Lankester (1885). Family
Peneroplidae Cushman. Family Soritidae Galloway (1933). Specialized derivatives
of the lower Miliolida: planispiral shells in which the chambers become successively
larger, as in Peneroplis, and, by a further development, divided into large numbers
of secondary chambers, as in Archaias, Sorites, and Orbitolites. Spirolina, the shell
coiled in the oldest part, straight in the remainder.
Family 3. Alveolinea Ehrenberg (1839). Family Alveolinida Schultze 1854.
Families Alveolinina and Keramosphaerina Lankester, Alveolinellidae and Keramo-
sphaeridae Cushman. Another group of specialized derivatives, the planispirally coil-
ing chambers broadened and divided into many chamberlets with separate apertures,
the entire body more or less globular. Borelis; Fasciolites Parkinson 1811 [Alveo-
lina d'Orbigny 1826); Alveolinella; Keramosphaera Brody, a rare antarctic form.
The organisms of these last two families resemble, as a parallel development, those
of order Nummulitinidea, from which they are distinguished by the texture.
Order 3. Foraminifera Zborewski 1834.
Order Polythalamia and subordinate group Polysomatia Ehrenberg in Abh.
Akad. Wiss. Berlin (1838): table 1 (1839).
Order Polysomatia Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1 : 11
(1848).
Orders Lituolidea, Textularidea, and Lagenidea Lankester in Enc. Brit. ed. 9,
19: 847 (1885).
186] The Classification of Lower Organisms
Order Perforida Delage and Herouard Traite Zool. 1 : 107 (1896).
Orders Nodosalida and Textulinida Calkins Biol. Prot. 355, 356 (1926).
Comparatively unspecialized Rhizopoda, the shells of various textures, not porcel-
lanous; not usually of trochoid form, and if so, usually not vitreous.
Early students, Montfort, Lamarck, and d'Orbigny, were much concerned with
organisms which they called Geophonus, Vorticialis, or Polystornella. These names
represent organisms of much the appearance of Nautilus; all are synonyms of Elphi-
dium Montfort, which is to be considered the type or standard genus of Foraminifera.
Family 1. Endothyrina Lankester (1885). Family Endothyridae Rhumbler 1895.
Fossils, pre-Cambrian to Carboniferous, the calcareous shells granular or fibrous, not
porcellanous or vitreous. Cayeuxina Galloway ( 1933) includes minute globular shells
solitary or irregularly clustered, described by Cayeux, 1894, from the pre-Cambrian
of Brittany; Matthewina Galloway includes Cambrian fossils of similar character.
Endothyra and Cribrospira are Carboniferous forms, planispirally coiled; Tetrataxis
produced trochoid shells.
Family 2. Nodosinellida [Nodosinellidae] Rhumbler 1895. Shells like those of
the Endothyrina or containing imbedded grains of sand, one-chambered or forming
straight or curved, not coiled, rows. Mostly Carboniferous, rare as late as the Eocene.
Archaelagena, Nodosinclla, Nodosaroum, Pedangia, etc.
Family 3. Reophacida [Reophacidae] Cushman 1827. A small group of forms ap-
parently degenerate from the foregoing, the chambers in straight, curved, or irregular
series, walls chitinous or sandy; sometimes parasitic in other rhizopods. Reophax, etc.,
surviving to the present in cold deep water.
Family 4. Trochamminida [Trochamminidae] Schwager 1877. Family Trocham-
minina Lankester (1885). Family Plocapsilinidae Cushman. Cells planispiral or
trochoid, becoming evolute or irregular; walls with imbedded grains of sand. Penn-
sylvanian to recent, abundant only in the Cretaceous. Trochamniina, Plocapsilina, etc.
Family 5. Lituolidea Reuss 1861. Family Lituolidae Brady (1881). Families
Lituolina and Loftusiina Lankester. Family Lituolidaceae Lister. Families Loftu-
siidae and Neusinidae Cushman. Shells spiral or becoming evolute or irregular, with
walls of agglutinated siliceous or calcareous matter, the chambers subdivided as in
order Nummulitinidea. Cyclammina, Lituola, Loftusia, Neusina, etc.; Mississippian
to recent, most abundant in the Cretaceous and at present.
Family 6. Orbitolinida [Orbitolinidae] Martin 1890. Specialized derivatives of
the preceding family, walls agglutinated as in that group, the numerous chambers
forming a conical or nearly circular body. Dictyoconus, Orbitolina, etc. Mesozoic and
Eocene.
Family 7. Ataxophragmidea Schwager 1877. Families Valvulinidae and Vcr-
neulinidae Cushman 1927. Family Ataxophragmidae Galloway (1933). Having
walls of agglutinated material and allied to the preceding families; chambers of the
shell tending to form an elongate, screw-like spiral. Valvulina, Ataxophragmium,
Verneulina, etc.; since early Mesozoic, abundant in the present.
Family 8. Textularina Ehrcnberg (1839). Family Textularidac d'Orbigny (1839).
Family Textulariaccac Lister. Walls more or less agglutinated, the chambers usually
in an elongate spiral with two members to a cycle, so that they form two series, the
body as a whole tending to be wedge-shaped. Textularia, Cuneolina, Vulvulina, etc.;
Ordovician to the present.
Family 9. Nodosarina Ehrcnberg (1839). Family Nodosarida Schultzc 1854.
Family Lagenidae Brady (1881). Family Lagenina Lankester (1885). Family Nodo-
Phylum Protoplasta [ 187
saridae Rhumbler. Family Lagenaceae Lister. Walls calcareous, hyaline, perforate;
chambers planispiral in the earliest forms, becoming curved or straight in the major-
ity; orifice ordinarily of radiating slits, becoming reduced to a single slit. A numerous
group, Triassic to the present. Lenticulina Lamarck {Lenticulites Lamarck and
Crist ellaria Lamarck are synonyms) is Naiitilus-Vikt. Hemicristellaria and Vaginulina
resemble the sheath of a dagger; Flabellina and Frondicularia resemble fans; Glandu-
lina is shaped like a jug. Nodosaria is like a row of enlarging beads. Lagena is a one-
chambered form connected to Nodosaria by transitions, and evidently reduced, not
primitive.
Family 10. Polymorphinida [Polymorphinidae] d'Orbigny. Families Polymorphin-
ina and Ramulinina Lankester ( 1885). Specialized irregular forms related to the pre-
ceding, as indicated by orifices of the same character. Polymorphina, etc., present in
the Mesozoic, abundant in the Cenozoic to the present.
Family 11. Nonionidea [Noninideae] Reuss 1860. Family Polystomellina Lankester
(1885). Family Hantkeninidae Cushman. Shells mostly nautiloid, that is, plani-
spiral with successively larger chambers, a few of the highest trochoid; walls hyaline,
perforate; aperture generally a transverse slit. N onion Montfort {Nonionina d'Or-
bigny) and Elphidium Montfort [Geophonus Montfort, Vorticialis Lamarck, Poly-
stomella Lamarck, the apparent type of Foraminifera) are simply nautiloid; Hant-
kenina is ornamented with spines. Jurassic to the present.
Order 4. Globigerinidea Lankester in Enc. Brit. ed. 9, 19: 847 (1885).
Orders Rotalidea and Chilostomellida Lankester 1. c, both names having prev-
ious use in the category of families.
Order Rotalida Calkins Biol. Prot. 356 (1926).
The main body of Rhizopoda with perforate hyaline shells, many-chambered, the
chambers primitively of the trochoid arrangement.
Family 1. Rotalina Ehrenberg (1839). Family Rotalidea Reuss 1861. Family
Rotalidae Brady (1861). Family Rotalina Lankester. Family Rotaliaceae Lister.
Families Globorotaliidae, Anomalinidae , and Planorbidinidae Cushman. A numerous
family, including unspecialized forms, Globorotalia, Rotalia, etc., as well as degen-
erate and irregular forms, Piano pulvinulina, etc., and moderately specialized ones
with conical or disk-shaped bodies of numerous chambers, Cymbalopora, Planorbu-
lina, etc. Triassic, rare; Jurassic to the present, common.
Family 2. Acervulinida Schultze 1854. Family Rupertiidae Cushman. A small
group of degenerate derivatives of the foregoing, the bodies attached, irregular, some-
times reduced to one chamber. Rupertia, Acervulina, etc.. Cretaceous to the present.
Family 3. Tinoporidea Schwager 1877. Family Calcarinidae Cushman. Another
small group derived from Rotalina, the disk-shaped cells with a whorl of prominent
spines. Calcarina, Tinoporus, etc. Cenozoic, to the present.
Family 4. Asterigerinida [Asterigerinidae] d'Orbigny (1839). Two genera, Asteri-
gerina and Amphistegina, diverging from Rotalina in having each chamber divided
into two by an oblique wall. Doubtfully in the Cretaceous; Eocene to the present.
Family 5. Chapmaniida [Chapmaniidae] Galloway (1933). The numerous cham-
bers arranged in a low cone whose inside is filled with deposited solid material. Chap-
mania, Halkyardia, Dictyoconoides. Eocene and Oligocene.
Family 6. Chilostomellida [Chilostomellidae] Brady (1881). Family Chilostomell-
aceae Lister. A few genera of reduced derivatives of Rotalina with few chambers.
Allomorphina, Chilostomella, Sphaeroidina, etc. Jurassic to the present.
188 ] The Classification of Lower Organisms
Family 7. Orbulinida Schultze 1854. Family Globigerinida Carpenter 1862.
A few genera with the chambers mostly few, subglobular, clustered rather than ar-
ranged in a definite pattern. Orbulina. Globigerina, abundant, pelagic in all oceans,
the shells abundant in the ooze on the bottom. Pennsylvanian, doubtful; Jurassic, rare;
Cretaceous to the present, common.
Family 8. Pegidiida [Pegidiidae] Heron-Allen and Earland 1928. A few genera
much like the Orbulinida but with thinner walls. Pegidia, etc. Oligocene to the
present.
Family 9. Heterohelicida [Heterohelicidae] Cushman 1927. A numerous group,
the shells screw-like, biseriate, uniseriate, sheath-like or fan-like, the walls often with
exterior ornamentation; paralleling the Nodosarina, but without the radiate orifices.
Heterohelix, Sagrina, Eouvigerina, Pavonina, Plectojrondicidaria, Bolivina, Mucron-
ina. Common, Jurassic to the present.
Family 10. Buliminida Jones 1876. Family Uvellina Ehrenberg (1839), not
based on a generic name. Family Buliminina Lankester (1885). Shells mostly high
spirals, screw-like, often with spines or other external ornamentation, the orifices
various, commonly comma-shaped. Turrilina, Bulimina, Virgulina, etc. Triassic to
the present.
Family 11. Cassidulinida [Cassidulinidae] d'Orbigny (1839). A small group with
high-spiralled shells and comma-shaped orifices, evidently derived from the forego-
ing family. Cassidulina, etc. Eocene to the present.
Family 12 Uvigerinida [Uvigerinidae] Galloway and Wissler, 1927. Further vari-
ants from Heterohelicida, the high-spiralled shells with chambers in three rows at
first, varying to biseriate and uniseriate. Uvigerina, Siphonogenerina, etc. Jurassic
to the present, common since the Miocene.
Family 13. Pleurostomellida [Pleurostomellidae] Reuss 1860. An additional
rather small family of the same general character as the few preceding. Pleurosto-
mella, Nodosarella, Daucina, Ellipsoidina, etc. Cretaceous to the present, commonest
in upper Cretaceous and Eocene.
Order 5. Nummulitinidea Lankester in Enc. Brit. ed. 9, 19: 848 (1885).
Rhizopoda with large specialized shells, the walls hyaline, perforate, generally
thickened and traversed by channels and thrown into internal ridges which subdivide
the chambers.
Family 1. Fusulinida [Fusulinidae] Moller 1878. Carboniferous fossils, the
chambers short and broad, numerous, in a planispiral coil, forming bodies which are
usually fusiform or globular. Orobias, Fusidina, Triticina, Verbeekina, etc.
Family 2. Nummulitida [Nummulitidae] Reuss 1861. Family Camerinidae
Meek and Hayden 1865. Family Nummulinidae Brady (1881). Family Nummuli-
tina Lankester ( 1885). Family Nummulitaceae Lister. Mostly disk-shaped, planispiral,
the walls not highly specialized. Camerina Bruguiere 1792 {Nxunrnulites Lamarck
1801), Operculina, Heterostegina, etc. Jurassic to the present, most abundant in the
Eocene.
Family 3. Orbitoidida [Orbitoididae] Schubert 1920. Similar to the foregoing,
the numerous chambers divided into numerous chamberlcts. A considerable group of
Mcsozoic and Ccnozoic fossils. Orbitoides, Cyclosiphon, etc.
Family 4. Cycloclypeina Lankester (1885). Family Cycloclypeidae Galloway
(1933). Similar to the preceding. A number of Mcsozoic and Ccnozoic genera, most
numerous in the Eocene. Asterocydina. The only living species is Cycloclypeiis Car-
penteri Brady.
Phylum Protoplasta [ 189
Class 4. HEUOZOA Haeckel
Family Polycystina Ehrenberg in Abh. Akad. Wiss. Berlin 1838: 128 (1839).
Rhizopoda radiaria seu Radiolaria J. Miiller in Abh. Akad. Wiss. Berlin (1858) :
16 (1859).
Echinocystida Claparede.
Order Radiolaria Haeckel Radiolarien 243 (1862).
Stamm Moneres for the most part, and classes Heliozoa and Radiolaria, Haeckel
Gen. Morph. 2: xxii, xxviii, xxix (1866).
Subclasses Heliozoa and Radiolaria Biitschli in Brown Kl. u. Ord. Thierreichs 1, 1
Teil: Inhalt (1882).
Class Proteomyxa Lankester in Enc. Brit. ed. 9, 19: 839 (1885).
Subclasses Proteomyxiae, Heliozoariae, and Radiolariae Delage and Herouard
Traite Zool. 1: 66, 156, 169 (1896).
Class Actinopoda Calkins Biol. Prot. 318 (1926).
Class Actinopodea and orders Helioflagellida, Heliozoida, Radiolarida, and Proteo-
myxida HaU Protozoology 202, 203, 212, 220 (1953).
Subphylum Actinopoda Grasse and Deflandre, and classes Acantharia, Radiolaria,
and Heliozoa Tregouboff in Grasse Traite Zool. 1, fasc. 2: 267, 270, 321, 437
(1953).
Organisms having pseudopodia of the character of filopodia, stiffly radiating, or of
axopodia, stiffly radiating and having inner fibers; often with siliceous skeletons.
Here, not without authority, one combines in one class the three groups which have
been treated as the classes Proteomyxa, Heliozoa, and Radiolaria; and adds further
two families of shelled amoebas.
Cienkowski (1865) listed as "Monaden" the new species or genera Monas amyli,
Colpodella (apparently a chytrid), Pseudospora, and Vampyrella. They are minute
fresh-water amoeboid organisms, in part having flagellate stages. Haeckel (1866)
placed most of them (the Monas under the new generic name Protomonas), together
with his own discoveries Protamoeba and Protogenes, and also the bacteria, in his
Stamm Moneres, i.e., his group of Protista without nuclei. Later (1868) he omitted
the bacteria. Zopf (1885) found several of Haeckel's Moneres to possess nuclei, and
Lankester renamed the group Proteomyxa. Publication of subsequent original obser-
vations of these organisms has been scant and scattered; they remain poorly known.
The Heliozoa as conventionally construed are also mostly inhabitants of fresh
water. Ehrenberg observed some of them and took them for Infusoria with immobile
cilia. There are only a few dozen species of Heliozoa sensu strict o (Schaudinn, 1896) :
the whole group is no more than a reasonable order.
The Radiolaria (this name also used at this point in its conventional sense) are
marine. Examples were first observed as floating gelatinous bodies. These were taken
for fragments and remained unnamed until 1834, when Mayen named Physematium
and Sphaerozoum. Fossil skeletons of many examples were described by Ehrenberg
(1839). Huxley (1851) named Thalassicolla and gave an accurate account of its
structure. It was by work on organisms of this group that Haeckel first distinguished
himself (1862).
Haeckel dealt further with this group in four important papers (1879, 1882, 1887,
1887-1888). In the last of these, the Radiolaria are a class of four legions, eight sub-
legions, twenty orders, 85 families, 739 genera, and more than four thousand species.
The categories, Haeckel explained, are purely relative: Radiolaria would as well be
190] The Classification of Lower Organisms
a phylum, the legions classes, and so forth. This idea served him as license for con-
founding the application of many names, by shifting them among the categories, or
by substituting new names for old. All subsequent authors have followed Haeckel's
system of Radiolaria, applying names as best they might.
The class Heliozoa in the extended sense here proposed may be organized as five
orders distinguished as follows:
1. Cells without a central capsule, i.e., without
a firm membrane surrounding the inner part
of the protoplast Order 1. Radioflagellata.
1. Cells with a central capsule.
2. Central capsule of spherical symmetry
or with three planes of symmetry at
right angles, punctured by many pores.
3. Pores of the central capsule evenly
distributed; skeleton absent or pres-
ent, without spicules which cross the
central capsule or meet in its center Order 2. Radiolaria.
3. Pores of the central capsule clust-
ered; skeleton including spicules
which cross the central capsule or
meet in its middle Order 3. Acantharia.
2. Central capsule of radial symmetry, with
one opening Order 4. Monopylaria.
2. Central capsule of isobilateral symmetry,
with one main opening and two minor
ones Order 5. Ph.'SlEosphaeria.
Order 1. Radioflagellata Kent Man. Inf. 1 : 225 ( 1880) .
Subdivision or subclass Heliozoa (Haeckel), and orders Aphrothoraca (Hert-
wig), Chlamydophora (Archer), Desmothoraca (Hertwig and Lesser), and
Chalarothoraca (Hertwig and Lesser) Biischli in Bronn Kl. u. Ord. Thier-
reichs 1: 261, 320, et seq. (1881, 1882).
Suborder Protoplasta Filosa Leidy in Rept. U.S. Geol. Survey Territories 12:
189(1879).
Class Proteomyxa Lankester ( 1885) .
Subclass Proteomyxiae and orders Acystosporidia, Azoosporidia, and Zoospori-
dia; subclass Heliozoariae and orders Aphrothoracida, Chlamydophorida,
Chalarothoracida, and Desmothoracida Delage and Herouard Traite Zool.
1: 66-72, 156-168 (1896).
Order Heliozoa Doflcin Protozoen 13 ( 1901 ) .
Orders Vampyrellidea and Chlamydomyxidea Poche in Arch. Prot. 30: 182,
193 (1913).
The proper Heliozoa together with the Proteomyxa: organisms of the character
of the class, lacking central capsules, that is, firm membranes about the inner part of
the protoplasts. Mostly fresh water organisms of spherical .symmetry, commonly with-
out skeletons. The type, being the only genus assigned to the order by Kent, is
Actinomonas.
1. Pscudopodia unspecialized; amoeboid organ-
isms with or without flagellate stages.
Phylum Protoplasta [191
2. Without shells Family 1. Pseudosporea.
2. With shells; without known flagellate
stages.
3. Shells chitinous, without siliceous
scales Family 2. Lagyntoa.
3. Shells bearing circular siliceous
scales Family 3. Euglyphida.
1. Pseudopodia slender, with apical knobs Family 4. Vampyrellacea.
1. Pseudopodia of the character of typical axo-
podia, without apical knobs; the cells or their
main bodies usually regularly spherical.
2. Bearing flagella as well as axopodia in
the vegetative condition Family 5. AcTiNOMONADroA.
2. Without flagella in the vegetative
condition.
3. Cells without a lifeless outer coat Family 6. AcTiNOPHRYroA.
3. Cells having a gelatinous outer coat
without siliceous spicules Family 7. Heterophryida.
3. Cells having a gelatinous outer coat
with siliceous spicules Family 8. Acanthocystida.
3. Cells with a hard shell punctured
by pores Family 9. Clathrulinida.
Family 1. Pseudosporea [Pseudosporeae] Berlese in Saccardo Sylloge 7: 460
fl888). Monadineae Zoosporcae Cienkowski in Arch. mikr. Anat. 1: 213 (1865).
Family Pseudosporeen Zopf Pilzthiere 115 (1885). Orders Azoosporidea for the
most part and Zoosporidca Delage and Herouard (1896). Azoosporidae for the most
part and Zoosporidae Doflein Protozoen 40, 41 ( 1901 ) . Family Pseudosporidae Poche
in Arch. Prot. 30: 197 (1913). Amoeboid organisms without shells or skeletons, the
pseudopodia tapering from a broad base to a filamentous termination. Flagellate
stages (with one flagellum or two unequal flagella) occur in Protovionas, Pseudo-
spora. and Diplophysalis. In other genera, as Arachnula and Chlamydomyxa, no
flagellate stages are known.
Family 2. Lagynida Schultze 1854. Order Gromida Claparede and Lachmann
1859. Family Gromida Carpenter 1862. Family Gromiidae Brady (1881). Families
Monostomina and A.mphistomina Lankester (1885). Amoeboid organisms having
chitinous shells without siliceous scales with a broad orifice through which project
pseudopodia of the character of filopodia. Grom.ia, Lagynis, etc.
Family 3. Euglyphida [Euglyphidae] Wallich 1874. Amoeboid organisms with a
chitinous shell beset with circular siliceous scales, the filopodia projecting through a
broad orifice. Euglypha, Cyphoderia, Campuscus, Trinema, etc.
Family 4. Vampyrellacea [Vampyrellaceae] Zopf Pilzthiere 99 (1885). Monadin-
eae Tetraplasteae Cienkowski op. cit. 218. Family Vampyrelleae Berlese in Saccardo
Sylloge 7: 454 (1888). Family Vampyrellidae Poche in Arch. Prot. 30: 182 (1913).
Cells subglobular, slowly creeping, with slender pseudopodia, numerous, densely
packed and stiffly radiating on mature individuals, bearing terminal knobs. Vampy-
rclla, the cells colored faintly pink by some metabolic by-product, is not unfamiliar
as a predator on freshwater algae cultured under unfavorable conditions.
Family 5. Actinomonadlda [Actinomonadidae] Kent Man. Inf. 1: 226 (1880).
Family Ciliophryidae Poche in Arch Prot. 30: 187 (1913) Family Helioflagellidae
192]
The Classification of Lower Organisms
I
%j^
t)v:
h \
Fig. 37. — Radioflagellata : a-f, Diplophysalis stagnalis after Karling (1930);
a, b, young cells with one or two flagella; c, active amoeboid form; d, walled cell;
e, iame releasing flagellate cells; f, resting cell, g. Young cell of Vampyrella x 1,000.
h, Actinosphaerium Eichhornii x 1,000.
Phylum Protoplasta [ 193
Doflein. Organisms bearing at the same time flagella and typical axopodia. Dimor-
pha, free-swimming, with two unequal flagella. Actinomonas, Pteridomonas, Cilio-
phrys, with one flagellum, either free-swimming or attached by a protoplasmic stalk.
Family 6. Actinophryida [Actinophryidae] Glaus 1874. Askeleta Hertwig and
Lesser in Arch. mikr. Anat. 10 Suppl. 164. (1874). Aphrothoraca seu Actinophryidae
Hertwig Org. Radiolar. 142 (1879). Order Aphrothoraca Butschli (1881). Suborder
Aphrothoraca Minchin (1912). Family Camptonematidae Poche in Arch. Prot. 30:
187 (1913). Cells typically spherical, with typical axopodia, having no flagella nor
shells nor skeletons. Actinophrys Sol Ehrenberg and Actinosphaerium Eichhornii
(Ehrenberg) Stein are common in fresh water among algae, living as predators
largely on diatoms; Actinophrys is uninucleate, the cells to 50^ in diameter; Actono-
spaerium is multinucleate, the cells to lOOOjl in diameter. Camptonema is marine.
Mitosis in Actinophrys as described by Schaudinn (1896) occurs within an intact
nuclear membrane which undergoes constriction; the dividing nucleus lies within a
spindle-like body of cytoplasm. Schaudinn and Belar (1923) observed conjugation.
Pairs of gametes, which are usually sister cells but may sometimes be random pairs,
lie within a cyst wall of secreted material. The nucleus of each gamete undergoes
meiosis; at the end of each meiotic division, one of the daughter nuclei is digested;
thus each gamete comes to possess a single haploid nucleus. Syngamy and karyogamy
follow in due course and the zygote becomes walled. An old account of the cytology
of Actinosphaerium by Hertwig is defaced by descriptions of the origin of nuclei from
fragments of nuclei (chromidia), and of nuclear fusions at two separate stages of
development.
Family 7. Heterophryida [Heterophryidae] Poche in Arch. Pr«it. 30: 189 (1913).
Heliozoa Chtamydophora Archer in Quart. Jour. Micr. Sci. n.s. 16: 348 (1876).
Order Chlamydophora Butschli (1882). Suborder Chlamydophora Minchin (1912).
Family Lithocollidae Poche I.e. The cells or their main bodies spherical with axopodia
projecting through a gelatinous envelope. Heterophrys and Astrodisculus are simply
globular cells. Elaeorhanis and Lithocolla are similar but with grains of sand or dia-
tom shells imbedded in the envelope. Actinolophus is stalked. Sphaerastrum becomes
colonial by incomplete division of the cells.
Family 8. Acanthocystida [Acanthocystidae] Glaus 1874. Chalarothoraca Hert-
wig and Lesser in Arch. mikr. Anat. 10 Suppl. 193 ( 1874). Chalarothoraca seu Acan-
thocystidae Hertwig Org. Radiolar. 142. (1879). Order Chalarothoraca Butschli in
Bronn Kl. u. Ord. Thierreichs 1: 325 (1882). Suborder Chalarothoraca Minchin
(1912). Resembling the preceding family, but the gelatinous envelope containing
hard bodies, supposedly usually siliceous, of definite form. In Raphidophrys, these
bodies are curved needles; in Pinacocystis, small plates; in Acanthocystis and Pinacio-
phora, disks bearing a central spine which is in some species forked. The cell of Wag-
nerella (a marine form, on rocks in bays) consists of a globular head with spines and
axopodia, borne on a protoplasmic stalk attached by a foot; the nucleus lies in the foot.
In these forms the axial filaments of the pseudopodia radiate from a central gran-
ule located outside the nucleus (in Wagnerella, in the head). Schaudinn (1896) re-
ported nuclear division in Acanthocystis as being either amitotic or mitotic: the
report of amitosis is of course not to be taken seriously. In the mitotic process, the
central granule acts as a centrosome; the chromosomes are numerous and minute;
the nuclear membrane disappears during the middle stages. Nuclear division may be
followed by division of the cell into two, or may be repeated and followed by produc-
tion of buds. The buds may lose their pseudopodia and develop paired flagella. It is
194 ] The Classification of Lower Organisms
suspected that the flagellate cells may be gametes. The central granule is said to
originate by extrusion from the nucleus of a bud.
Zuelzer (1909) found in Wagner ella two types of individuals, slender and stout,
supposedly respectively haploid and diploid. In either type the nuclei may become
numerous (and it is said that they sometimes develop from chromidia). The nuclei
may migrate to the head and be released in buds, or they may become distributed
throughout the protoplast, which then breaks up into biflagellate cells. It is supposed
that these may be gametes, but a fusion of the heads of individuals of the slender
type was observed.
Family 9. Clathrulinida [Clathrulinidae] Glaus 1874. Desmothoraca Hertwig
and Lesser op. cit. 225. Desmothoraca seu Clathrulinidae Hertwig Org. Radiolar.
142 (1879). Order Desmothoraca Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 328
(1882). Family Choanocystidae Poche in Arch. Prot. 30: 192 (1913). Protoplasts ly-
ing within globular shells, apparently of chitin, usually stalked, punctured by numer-
ous pores through which the axopodia project. In reproduction, the protoplast may
divide into two, one of which escapes from the shell and secretes a new one; or it may
divide into many which become unequally biflagellate. Clathrulina, Hedriocystis,
Choanocystis.
Order 2. Radiolaria (J. Miiller) Haeckel Radiolarien 243 (1862).
Rhizopoda radiaria sen Radiolaria J. Miiller in Abh. Akad. Wiss. Berlin 1858:
16 (1859).
Orders Thalassicollen, Sphacrozoen, and Peripyleen Hertwig Org. Radiolar. 133
(1879). *
Orders Pcripylaria, Collodaria, Symbelaria, and Syncollaria Haeckel in Jen-
aische Zeitschr. 15: 447, 469, 471, 472 (1882).
Legion S pumellaria or Peripylea, with orders Collodaria and Sphaerellaria and
seven suborders, Haeckel in Rept. Voy. Challenger Zool. 18: 5, 9 ( 1887) .
Legion Spiimellaria, sublegions Collodaria and Sphaerellaria, and six orders,
Haeckel Radiolarien 2: 87 (1887).
Order Peripylida Delage and Herouard Traite Zool. 1 : 176 ( 1896).
Suborder Peripylaria Minchin Protozoa 225 (1912).
Order Sphaeridca Poche in Arch. Prot. 30: 206 (1913).
Order Peripylea Calkins Biol. Prot. 343 (1926).
Suborder Peripylea Kudo Handb. Protozool. 259 (1931).
Suborder Pen'py/ma Hall Protozoology 218 (1953).
This order and the three which follow, being the Radiolaria as conventionally con-
strued, are unicellular marine organisms with axopodia, having within the protoplast
a layer of organic material, variously punctured and of various types of symmetry,
which separates the inner protoplasm from the outer. The central capsule consists of
this layer (the central capsule membrane) and its contents, including the one or more
nuclei of the cell. Imbedded in the protoplasm there is usually a skeleton, usually sili-
ceous, various in structure and sometimes highly complicated. The outer cytoplasm
is commonly inhabited by symbiotic cryptomonads in the resting condition (yellow
cells, zooxanthellae), and sometimes contains masses of dark material, apparently
debris extruded from the central capsule. The type of Radiolaria is evidently Thalas-
sicolla; this genus was the first one described from living material, and was listed
first by J. Miiller in the original publication of the name.
The order which includes Thalassicolla, and to which the name Radiolaria is here
restricted, is distinguished by uniformly distributed small punctures in the central
Phylum Protoplasta [ 195
capsule membrane and by the absence of skeletal spicules extending across the central
capsule or meeting in its middle. Except during reproduction, each central capsule
contains a single nucleus, but the cells of many examples are coenocytic, containing
several or many central capsules.
Brandt (1885, 1905) observed reproduction particularly as it occurs by the produc-
tion of swimming cells by some of the coenocytic forms. The nucleus divides to pro-
duce very many, and the intracapsular cytoplasm divides to produce uninucleate
flagellate cells. In Collosphaera, all the nuclei are included in these cells. The cells
are of two sizes, produced by different individuals, and are supposed to be gametes. In
Sphaerozoum and its allies, some of the nuclei degenerate instead of being included
in the swimming cells, of which two sizes are produced by single individuals. It ap-
pears that the swimming cells have characteristically two unequal flagella, though
many are found to have only one, and some produce a third appendage by which
they can attach themselves.
Haeckel listed thirty-two families in his legion Spumellaria. Other authors recog-
nize about a dozen, including the following.
Family ThalassicoUida Haeckel (1882). Thalassicollen J. Miiller (1859). Family
Collida Haeckel (1862); there is no corresponding generic name. Order Collida
Haeckel (1887). Globular forms with a single central capsule, skeleton none or of
numerous small spicules. Thalassicolla, Physematium, Lampoxanthium, etc.
Family Sphaerozoida Haeckel (1882). Family Collozoida Haeckel op. cit. Family
Sphaeroidina Haeckel (1862); there is no corresponding generic name. Coenocytic,
each cell with several nuclei in separate central capsules; skeleton none or of numer-
ous small spicules. Sphaerozoum, the cells globular, to 1 mm. in diameter; Raphido-
zoum, the cells elongate.
Family CoIIosphaerida Haeckel (1862). Coenocytic, the spherical cell to 1 mm. in
diameter, with several central capsules, each with an individual lattice-like skeleton.
Collosphaera.
Family Haliommatina Ehrenberg 1847. Families Ethmosphaerida, Ommatida,
and Cladococcida Haeckel ( 1862) . Family Sphaerida Haeckel ( 1882) . Order Sphaer-
oidea, with six families, Haeckel '(1887). Globular, with small numbers of radiating
main spicules, the main spicules bearing tangential branches which form a globular
network of definite pattern, or, often, two or more concentric networks. Haliomma,
Actinomma, Hexacontium, Cladococcus, and many other genera.
Further families are of the character of the Haliommatina, but with the spherical
symmetry modified by abbreviation or elongation of one or more axes:
Family Spongurida Haeckel (1862). Order Prunoidea, with seven families,
Haeckel (1887). Having one axis elongate. Spongurus, Pipetta, etc.
Family Lithocyclidina Ehrenberg 1847. Family Discida Haeckel (1862). Order
Discoidea, with six families, Haeckel ( 1887) . Having one axis shorter than the others.
Lithocyclia, Staurocyclia, Heliodiscus, etc.
Family Larcarida Haeckel (1887). Order Larcoidea, with this and seven other
families, Haeckel (1887). The skeleton with three unequal axes, or spiral. Cenolar-
cus, etc.
Order 3. Acantharia Haeckel in Jenaische Zeitschr. 15: 465 (1882).
Order Actipyleen Hertwig Org. Radiolar. 133 (1879).
Legion Acantharia or Actipylea, orders Acanthometra and Acanthophracta, and
seven suborders, Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887).
196]
The Classification of Lower Organisms
'^^'
0-
f
*lty
-S^'^v"^
^-'v-^if**^^^^
^^
>o,
::Q)t^^a
^/-T
Fig. 38. — Radiolaria: a, motile cells of Collosphaera Huxleyi after Brandt ( 1885).
b. Skeleton of Haliomma capillaris after Haeckel ( 1862) . c, Skeleton of Actinomma
Asteracanthion after Haeckel (1862). d. Skeleton of //f/zorfz,?cu5 P/zaco^wcu^ after
Haeckel (1887). Acantharia: e, Skeleton of Dorataspis costata after Haeckel
(1387). Monopylaria: f, Central capsule of Tridictyopus elegans after R. Hertwig
f 1879) . g, Skeleton of Lithocircus productus after R. Hertwig, op. cit. h. Skeleton
of Eucyrtidium carinatum after Haeckel (1862). Phaeosphaerl^: i. Typical cen-
tral capsule after R. Hertwig, op. cit.
Phylum Protoplasta [ 197
Legion Actipyea or Acantharia, sublegions Acanthometra and Acanthophracta,
and orders Actinellida, Acanthonida, Sphaerophracta, and Prunophracta
Haeckel Radiolarien II Teil (1887).
Order Actipylida Delage and Herouard Traite Zool. 1 : 204 ( 1896) .
Suborder Acantharia Minchin Protozoa 256 (1912).
Order Acanthometrida Poche in Arch. Prot. 30: 212 (1913).
Order Actipylea Calkins Biol. Prot. 345 (1926).
Suborder Actipylea Kudo Handb. Protozool. 216 (1931).
Suborder Actipylina Hall Protozoology 216 (1953).
In this group the central capsule membrane has many punctures arranged in
clusters. The skeleton includes radiating spicules; in some examples these extend
through the cell from side to side, passing through the central capsule; in the majority,
their proximal ends meet in the center of the central capsule. In the latter forms,
the number of radiating spicules is twenty, and they are arranged according to a pat-
tern discovered by Johannes Miiller and called Miiller's law; they form five parallel
whorls of four. Usually they bear tangential branches of definite, often highly elab-
orate, patterns: these may form a globular frame, or two or more concentric frames.
Haeckel, Hertwig, and Brandt found the spicules not to consist of silica. They are
soluble in acids and alkalis, and were reported to be destroyed by heat. They were
supposed, accordingly, to consist of some organic substance; Haeckel named it acan-
thin. Schewiakoff found them resistant to heat, and Biitschli (1906) analyzed them
and found them to consist of strontium sulfate. This surprising fact has recently been
confirmed by Odum (1951).
The cytoplasm at each point where a spicule passes through the surface is attached
to the spicule by a whorl of minute fibers called myophrisks. The myophrisks are
believed to be contractile, and to have the function of changing the volume, and
hence the density, of the cells, enabling them to sink or float.
Young cells contain a single nucleus, eccentric in the central capsule; older ones
have several to many nuclei.
Haeckel Hsted twenty families in his legion Acantharia. Other authors recognize
about a half dozen, including the following.
Family Litholophida Haeckel (1882). Family Astrolophida Haeckel (1887). Spi-
cules numerous, radiating, not arranged according to Miiller's law. Litholophus, As-
trolophus, Actinelius, etc.
Family Chiastolida Haeckel (1887). Spicules ten to twenty, extending clear
through the body. Chiastolus, Acanthochiasma.
Family Acanthometrida Haeckel (1862). Acanthometren J. MuUer (1859). Fam-
ily Acanthonida Haeckel ( 1882). With twenty spicules arranged according to Miiller's
law; they may be branched, but do not form a continuous network. In most examples,
as Acanthometron, Xiphacantha, etc., they are equal; in others, as Amphilonche, two
of the spicules of the equatorial whorl are much longer than the others.
Family Sphaerocapsida Haeckel (1882). Family Dorataspida Haeckel I.e. Order
Sphaerophracta Haeckel (1887). Like the foregoing, but the branches of the radiat-
ing spicules forming a globular network, or two or more concentric networks. Dora-
taspis, Sphaerocapsa, Lychnaspis.
Family Diploconida Haeckel (1862). Order Prunophracta Haeckel (1887). Again
hke the foregoing, but with the eight spicules of the two polar whorls either extended
or abbreviated. Diploconus, Hexaconus.
198] The Classification of Lower Organisms
Order 4. Monopylaria Haeckel in Jenaische Zeitschr. 15: 422 (1882) .
Order Afowopy/^^n Hertwig Org. Radiolar. 133 (1879).
Legion Nassellaria with orders Plectellaria and Cyrtellaria, and six suborders,
Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887).
Legion Nassellaria with sublegions Plectellaria and Cyrtellaria and orders Nas-
soidea, Plectoidea, Stephoidea, and Cyrtoidea, Haeckel Radiolarien H Teil
(1887).
Order Monopylida Delage and Herouard Traite Zool. 1: 215 (1896).
Suborder Nassellaria Doflein.
Suborder Monopylaria Minchin Protozoa 256 (1912).
Order Mo7zop};/ga Poche in Arch. Prot. 30: 218 (1913).
Suborder Monopylea Kudo Handb. Protozool. 261 (1931).
Suborder Monopylina Hall Protozoology 218 (1953).
This order is distinguished by a central membrane with one opening, or with a
single circular field of pores. From this opening or field as a base, there extends into
the central capsule a large conical body (apparently a bundle of protoplasmic fibers)
called the porocone. The skeleton varies from none to highly elaborate; it does not
in any form consist of separate spicules. Its symmetry is dorsiventral, not radial.
These skeletons are well known as microfossils.
In this group, under the name of legion Nassellaria, Haeckel placed twenty-six
families. Other authors recognize about a half dozen.
Family Nassellida Haeckel (1887). Skeleton none. Cystidium.
Family Plectonida Haeckel (1887). Family PlectidaYiztcktl (1882), not based
on a generic name. Skeleton consisting of three arms radiating from a point opposite
the mouth of the central capsule; sometimes with a fourth forming a caltrop.
Triplagia.
Family Stephanida Haeckel (1887). Family Stephida Haeckel (1882), not based
on a generic name. Skeleton including a ring in the sagittal plane, often with a basal
tiipod and with branches and crossbars. Lithocircus, Zygostephanus. This family is
well represented by microfossils as far back as the Cambrian.
Family Eucyrtidina Ehrenberg 1847. Family Polycystina Ehrenberg in Abh.
Akad. Wiss. Berlin ( 1838) : 128 ( 1839), not based on a generic name. Families Hali-
cryptina and Lithochytridina Ehrenberg 1847. Family Cyrtida Haeckel (1862).
Order Cyrtoidea, with twelve families, Haeckel (1887). Skeleton a more or less bas-
ket-shaped network; the root cyrt- in many of the names is Greek KupTr|, a fishing
basket. Lithocampe, Cryptocalpis, Eucyrtidium, Theoconus, Dictyoconus, and many
other genera. This group is common as Mesozoic and Cenozoic microfossils, occurring
mixed with diatoms and silicoflagellates.
Family Spyridina Ehrenberg 1847. Family Spyrida Haeckel (1882). Order
Spyroidea, with four families, Haeckel (1887). The skeleton divided by sagittal
grooves into two lobes.
Family Cannobotryida Haeckel (1887). Family Botrida Haeckel (1882), not
based on a generic name. Order Botryoidea, with three families, Haeckel ( 1887) . The
skeleton divided by three or more longitudinal grooves into as many lobes.
Order 5. Phaeosphaeria Haeckel in Sitzber. Jenaische Gess. Med. Naturw. 1879:
156(1879).
Phaeodariae, with orders Phaeocystia, Phaeogromia, Phaeosphaeria. and
Phaeoconchia Haeckel op. cit.
Phylum Protoplasta [199
Order Tripyleen Hertwig Org. Radiolar, 133 (1879).
Order Phaeodaria Haeckel in Jenaische Zeitschr. 15: 470 (1882).
Legion Phaeodaria and orders Phaeocystina, Phaeosphaeria, Phaeogromia, and
Phaeoconchia Haeckel in Rept. Voy. Challenger Zool. vol. 18 (1887).
Order Tripylea Doflein.
Suborder Tripylaria Minchin Protozoa 256 (1912).
Suborder Tripylea Kudo Handb. Protozool. 263 (1931).
Suborder Tripylina Hall Protozoology 218 (1953).
In this order, the central capsule is of isobilateral symmetry, having a rather
small main opening (astropyle) at one end and two smaller openings (parapyles)
at the other. The openings are located on projections of the central capsule mem-
brane; inside of each, the protoplasm is so differentiated as to appear to be a conical
bundle of fibers with the apex at the opening (in contrast to the preceding order, in
which the base of the conical structure is at the opening) . A mass of variously colored
bodies, supposedly excreted from the central capsule, lies in the extracapsular cyto-
plasm about the astroplyle. The skeletons consist in part of organic matter and are
not well preserved as fossils.
Borgert (1896, 1900) described nuclear division in Aulacantha. A very large
number of chromosomes, a matter of several hundred, form a plate which splits into
two; the two plates move apart in a body of differentiated cytoplasm, but no definite
spindle, and no centrosomes, were seen. The margins of the plates draw apart faster
than the middles, with the effect that the plates become saucer-shaped, then bowl-
shaped, and finally globular, after which nuclear membranes form about them. While
the nucleus divides, the central capsule membrane becomes constricted by longi-
tudinal grooves so placed that each daughter central capsule membrane receives
one parapyle and an astropyle formed from half of the original astropyle. The rudi-
ments of additional parapyles are first seen as granules in the intracapsular cyto-
plasm. Each granule grows slightly and becomes "hat-shaped," and migrates so as
to come into contact with the central capsule membrane at the point appropriate for
the development of its second parapyle.
Later, Borgert (1909) described a process in which the nucleus divides repeatedly,
producing many. The divisions are mitotic, with small numbers of chromosomes,
perhaps twenty; the eventual products become the nuclei of gametes. There are re-
ports, in part illustrated with photographs, of similar processes in family Thalassi-
collida (Hacker, 1907; Huth, 1913). According to Hollande (in Grasse, 1953) the
small nuclei are those of a parasitic dinoflagellate, Solenodinium. Le Calvez (1935)
found Coelodendrurn to produce zoospores with a pair of unequal simple flagella.
They resemble cells of Cryptomonas or of Bodo.
Haeckel's legion Phaeodaria was of fifteen families. These have been maintained
by the generality of authors.
A. Skeleton none or of distinct spicules; cells usually nearly spherical.
Family Aulacanthida [Aulacanthidae] Haeckel (1879). Aulacantha.
Family Astracanthida [Astracanthidae] Hacker. Spicules more or less thorny at
the distal ends. Aulactinium.
B. Skeleton spherical or of two concentric spheres, with no main opening.
Family Aulosphaerida Haeckel (1862). Aulosphaera.
Family Cannosphaerida [Cannosphaeridae] Haeckel (1879). Cromodromys.
Family Sagosphaerida Haeckel (1887).
C. Skeleton with a distinct main opening, either nearly spherical, radially sym-
metrical, or distinctly dorsiventral.
200]
The Classification of Lower Organisms
Family Castanellida [Castanellidae] Haeckel (1879). Skeleton nearly globular
with numerous pores. Castanidium.
Family Circoporida [Circoporidae] Haeckel (1879). Like the foregoing, but with
the pores gathered about the bases of radiating spines. Circoporus.
Family Tuscarorida Haeckel (1887). The main pore on an extended neck, the
skeleton accordingly flask-shaped. Tuscarora. Tuscarilla.
D. Shell strongly dorsiventral.
Family Challengerida [Challengeridae] J. Murray. Shell finely pitted. Chal-
lengeron.
Family Medusettida Haeckel (1887). Shell smooth or with small spines. Medu-
setta.
E. Shell bilaterally divided into two parts.
Family Concharida [Concharidae] Haeckel (1879).
Family Coelodendrida Haeckel (1862). The shell bearing extensive branched
appendages. Coelodendrum.
Class 5. SARKODINA (Hertwig and Lesser) Biitschli
Sarkodina Hertwig and Lesser in Arch. mikr. Anat. 10 Suppl. 43 (1874).
Class Sarkodina Biitschli in Bronn Kl. u. Ord. Thierreichs 1, 1 Teil: Inhalt ( 1882).
Class, subclass, etc., Rhizopoda Auctt.; class, subclass, etc. Sarcodina Auctt.
Amoeboid organisms without flagellate stages, the pseudopodia of the character
of lobopodia; without skeletons, without or with shells of various materials.
Fig. 39 — Chaos Protheus: a, b, cells x 25, after the original figures of Pelomyxa
carolinensis by Wilson (1900); c, mitotic figure x 2,000 after Short (1945).
Phylum Protoplasta [201
Hertwig and Lesser took note that the name Rhizopoda was originally applied to
organisms such as Miliola, which have rhizopodia; they proposed the name Sarko-
dina for all amoeboid organisms, with Rhizopoda as a subordinate group. Among
examples of Sarkodina which are not Rhizopoda, they listed first Difflugia, which
may accordingly be considered the standard genus.
The Sarkodina as here presented are not assumed to be a natural group. Their
common characters are probably the outcome of degeneration, by which organisms of
diverse evolutionary origins have lost their distinctions.
This assemblage is obviously and superficially divisible into two by the absence or
presence of shells. The resulting groups are construed as orders.
Order 1. Nuda Schultze 1854.
Family Amoebaea Ehrenberg Infusionthierchen 125 (1838).
Order Lofco^a Carpenter 1861.
Order Gymnamoebae Haeckel Gen. Morph. 2: xxiv (1866).
Order AmoebinaKtnt Man. Inf. 1: 27 (1880).
Suborder Amoebaea Biitschli in Bronn. Kl. u. Ord. Thierreichs 1: 176 (1880).
Order Gymnamoebida Delage and Herouard (1896).
Order Chaidea Poche in Arch. Prot. 30: 170 (1913).
Subclass Amoebaea and order Amoebida Calkins Biol. Prot. 335, 337 (1926).
Order Amoebaea Kudo Handb. Protozool. 204 (1931).
Sarkodina without shells. The type is the common amoeba, Amiba diffluens.
1 Protoplasts not tending to form pseudoplas-
modial communities.
2. Free-living Family 1. Amoebaea.
Family 2. MAYORELLroA.
Family 3. TnECAMOEBroA.
Family 4. Hyalodiscida.
2. Entozoic Family 5. ENDAMOEBroA.
1. Protoplasts assembling and acting in unison
in pseudoplasmodial communities.
2. Parasitic in plants Family 6. Labyrinthulida.
2. Predatory on bacteria, in air on moist
surfaces; mostly producing complicated
fructifications Family 7. Guttulinacea.
Family 1. Amoebaea Ehrenberg Infusionsthierchen 125 (1838). Family Amoe-
bidae Bronn 1859. Family Chaidae Poche in Arch. Prot. 30: 171 (1913). Family
Chaosidae Chatton in Grasse Traite Zool. 1, fasc. 2: 58 (1953). The ordinary free-
living amoebas. SchaeflFer (1926) limited the family to forms which produce numer-
ous indefinite granular pseudopodia. There has been much confusion as to the iden-
tity of the species. There are apparently two species of common large amoebas:
1. Chaos Protheus L. Syst. Nat. ed. 12: 1326 (1767) {Volvox Chaos L. Syst. Nat.
ed. 10: 821. 1758. Vibrio Protheus O. F. Muller Verm. Terr, et Fluv. 1: 45. 1773.
Pelomyxa carolinensis Wilson in American Nat. 34: 535. 1900. Chaos chaos Stiles).
Schaeffcr identified Pelomyxa carolinensis as the original Chaos Protheus L. It is
exceptionally large, being macroscopically visible, and is multinucleate. Surely,
sound nomenclature will apply to this species the name which Linnaeus gave it.
2. Amiba [Amoeba] diffluens (O. F. Muller) Ehrenberg Infusionsthierchen 127
(1838) [Proteus diffluens O. F. Muller Animac. Infus. 9. 1786; there is an older genus
202 ] The Classification of Lower Organisms
Proteus; Amiba divergens Bory Diet. Class. Hist. Nat. 1: 261. 1822; Amoeba Proteui
Leidy). It appears that MiiUer intended to rename the Chaos Protheus of Linnaeus;
that in 1773 he actually did so; but that in 1786 he applied another new name to a
different organism. Ehrenberg's amended spelling Amoeba, although in general use,
is not valid as that of a generic name; as Schaeffer suggests, the word may be used as
a common noun. Amiba diffluens is uninucleate; large, but not visible to the naked
eye.
In nuclear division in the common amoebas the nuclear membrane disappears.
There are many chromosomes in a blunt-ended spindle. Short (1945) noted a pecu-
liar twisting of the spindle of Chaos Protheus.
Schaeffer included in the present family three further genera, Trichamoeba Fro-
mentel, Polychaos Schaeffer, and Metachaos Schaeffer. Here, in ignorance of its
relationships, another well-known genus is assigned to this family.
Pelomyxa, typified by P. palustris Greeff ( 1874) , resembles Chaos Protheus in being
exceptionally large, macroscopically visible, and multinucleate. It is definitely dif-
ferent from Chaos Protheus in manner of movement (King and Jahn, 1948) and in
chemical characters (Andressen and Holter, 1949).
Minute amoebas moving by means of a single pseudopodium are called Vahlkamp-
fia. They are believed to have swimming stages with paired equal flagella. If so, they
do not belong to the present group, but perhaps to the plant kingdom.
Family 2. Mayorellida [Mayorellidae] Schaeffer in Publ. Carnegie Inst. 345: 47
(1926). Producing numerous brief conical pseudopodia, but moving by a single large
clear one. Mayorella, Pontifex, and several other genera proposed by Schaeffer;
Dactylosphaerium Hertwig and Lesser; Dinamoeba Leidy? The last may be the non-
flagellate stage of Chactoproteus Stein.
Family 3. Thecamoebida [Thecamoebidae] Schaeffer op. cit. 83. Amoebas with a
tough pellicle simulating a shell, moving by the outflow of clear protoplasm at the
anterior margin. Thccamocba Fromentel. Rugipes Schaeffer.
Family 4. ''Hyalodiscida [Hyalodiscidae] Poche in Arch. Prot. 30: 182 (1913).
Family Cochliopodiidae de Saedeleer in Mein. Mus. Roy. Hist. Nat. Belgique 60: 5
(1934). Similar to the foregoing but without the tough pellicle. Commonly dome-
shaped, with a row of small pseudopodia projecting from the margin. Hyalodiscus
and Cochliopodium of Hertwig and Lesser, together with certain genera of Schaeffer.
Family 5. Endamoebida [Endamoebidae] Calkins. Entozoic amoebas.
Endamoeba Leidy is found in cockroaches and termites. The nucleus contains no
karyosome, but many separate granules; in mitosis, definite chromosomes arc formed
(twelve in E. disparita), but there is apparently no centrosome; at least, no intra-
desmose is seen (Kirby, 1927).
Entamoeba Casagrandi and Barbagello, named at nearly the same time as the fore-
going and regrettably similarly, is widely distributed in invertebrate and vertebrate
hosts. E. dysenteriae (Councilman and Lafleur) Craig {Endamoeba histolytica Schau-
dinn) is a serious pathogen to man, the cause of amoebic dysentery. E. coli and E.
gingivalis are believed to be harmless commensals. The fully mitotic character of
nuclear division in these organisms was established by Kofoid and Swezy ( 1921, 1922,
1925). The nucleus contains a small karyosome and an intranuclear centrosome. Mi-
tosis begins with division of the centrosome into two, which remain connected, as they
draw apart, by a stainable strand, the intradesmose (the term is of Kofoid and Swczy,
1921). The karyosome breaks up into chromosomes, six in the species mentioned.
Spindle fibers connecting these to the centrosomes have been seen; Child (1926)
Phylum Protoplasta [ 203
found that the two halves of the spindle swing apart as the centrosomes move apart
like the legs of a compass being extended. There is no doubt that Endamoeba and
Entamoeba are generically distinct.
Endolimax, lodamoeba, and Councilmania occur chiefly in vertebrates and include
species commensal in man. A refined technique is required to discern the characters
by which they are distinguished from Entamoeba. Karyamoebina Kofoid and Swezy
(1924, 1925,) another commensal in man, resembles Vahlkampfia in details of the
mitotic process, and probably does not belong to the present group. Hydramoeba,
usually listed in the present family, is not an entozoic organism, but a predator on
Hydra.
Family 6. Labyrinthulida [Labyrinthuhdae] Haeckel ex Doflein Protozoen 47
(1901). There is a single genus Labyrinthula Cienkowski, and probably only one
species, L. macrocystis, parasitic in green and brown algae and in the marine seed
plant Zostera. The uninucleate cells are spindle-shaped. These cells send out from
one or both ends fine filaments which writhe in the water. The filaments from differ-
ent cells coil together and produce "tracks" along which the cells glide. The tracks
form a network on which the cells may be scattered or gathered into clusters; or the
cells may abandon their tracks and generate new ones. The nature of the tracks is
not clear. Possibly they are pseudopodia, on which the cells move by absorbing them
at one end while generating them at the other. Young (1943) found Labyrinthula
remarkably indifferent to variations in temperature, reaction, and salinity.
Family 8. Guttnlinacea [Guttulinaceae] Berlese in Saccardo Sylloge 7: 325 (1888)
Tribe Dictyosteliaceae Rostafinski Vers. 4 (1873). Sorophoreen with families Gut-
tulineen and Dictyostcliaceen Zopf Pilzthiere 131-134 (1885). Families Guttulineae
and Dictyosteliaceae Berlese op. cit. 451. Sappiniaceae Olive in Proc. American Acad.
37: 334 (1901). Families Sappiniidae, Guttulinidae , and Dictyostelidae Doflein
1909. Family Acrasidae Poche in Arch. Prot. 30: 177 (1913). Suborder Acrasina
Hall Protozoology 228 ( 1953 ) . Amoeboid cells predatory on bacteria and other scraps
of organic matter, in air on moist surfaces, commonly on dung. The cells are capable
of assembling and moving and going into a resting stage in unison. These organisms
have generally been included among the Mycetozoa; the resemblance is superficial.
More recently than Olive, Raper (1940) and Bonner (1944) have surveyed the
group and studied the behavior. Three families have been maintained, but one appears
sufiicient to accommodate the seven genera and approximately twenty species.
Cells of Sappinia are binucleate. They do not necessarily assemble in clusters; a
single cell may secrete a stalk, by which it is raised into the air, where it rounds up
and becomes dry. Alternatively, small numbers of cells may assemble and secrete a
common stalk. The dry cells are "pseudospores" : they are capable of resuming ac-
tivity without casting off a wall. Hartmann and Nagler (1908) described a peculiar
sexual process in Sappinia diploidea.
Guttulina and Guttulinopsis produce larger clusters of resting cells than Sappinia
does; in Guttulina the resting cells are said to be walled spores.
Acrasis produces fruits, solitary or clustered, of the form of uniseriate rows of spores
terminal on stalks consisting of rows of dead cells.
Distyostelium produces fruits consisting of a column of dead cells bearing a globu-
lar cluster of spores; Polys phondylium and Coenenia produce slightly more elaborate
fruits of the same general nature. In Dictyostelium, Raper and Bonner saw that the
amoeboid active cells, having devoured the available food, gather into a disk-shaped
mass which may exceed a millimeter in diameter. Wilson (1953) found syngamy,
204]
The Classification of Lower Organisms
Fig. 40. — a, Labyrinthula as a parasite in cells of Ectocarpus Mitchelliae x 1,000
after Karling (1944). b. Cell of Labyrinthula x 2,000 after Young (1943).
c, d^ Sappinia pedata, active cell and cyst, x 1,000 after Dangeard ( 1896) . e, f, Sap-
pinia pedata, cluster of pseudospores x 100 and single pseudospore x 1,000 after
Olive (1902). g, h, Guttulina scssilis, cluster of pseudospores x 100 and individual
pseudospores x 1,000 after Olive (1902). i-n, Dictyostelium discoideum x 10 after
Bonner (1944) : i, the pseudoplasmodium; j, it heaps itself up; k, falls toward the
light and creeps; 1, m, again heaps itself up and becomes a fruit, n. o, p. Fruits of
Dictyostelium mucoroides; q, of Polysphondylium violaccum; x 10, after Bonner,
op. cit.
Phylum Protoplasta [ 205
karyogamy, and meiosis to occur at this point; the chromosome number (n) is 7. The
disk changes into a column which bends, and then falls, toward the light, and after-
ward creeps some distance in the same direction. When this has happened, the fore-
most cells, being those which were originally in the middle of the disk-shaped mass,
pile up again to form a sterile stalk perhaps one millimeter tall; the cells behind them
crawl up the stalk to form the globular mass of spores; the hindmost, being those
which were last to arrive at the disk-shaped mass, remain behind to form a flange
about the base of the stalk.
Order 2. Lampramoebae Haeckel Gen. Morph. 2: xxiv (1866).
Order Testacea Schultze 1854, non L. (1758).
Order Thecamoebae Haeckel.
Order Conchulina Cash and Hopkinson British Freshw. Rhizop. 1: 37 (1905).
Suborder Testaceolobosa de Saedeleer in Mem. Mus. Roy. Hist. Nat. Belgique
60: 5 (1934).
Order Testacida Hall Protozoology 241 (1953).
Order Testaceolobosa Deflandre in Grasse Traite Zool. 1, fasc. 2: 97 (1953).
Amoeboid organisms without known flagellate stages, bearing shells and producing
lobopodia. Various organisms producing rhizopodia or filopodia, traditionally asso-
ciated with these, have here been placed among Rhizopoda or Heliozoa, as suggested
by de Saedeleer (1934) and Grasse (1953). Deflandre (in Grasse, op. cit.) distin-
guishes several families beside the following:
1. Shell without secreted scales of silica.
2. Shell of uniform secreted material Family 1. Arcellina.
2. Shell with imbedded grains of sand Family 2. Difflugiida.
1. Shell with secreted scales of silica Family 3. NEBELroA.
Family 1. Arcellina Ehrenberg Infusionsthierchen 129 (1838). Family Arcellidae
Schultze 1876. Arcella, etc.
Family 2. Difflugiida [Difflugiidae] Taranek 1881. Difflugia, etc.
Family 3. Nebelida [Nebelidae] Schouteden 1906. Nebela, the shell beset with
circular siliceous scales; Quadrula, the scales square; etc.
Chapter XI
PHYLUM FUNGILLI
Phylum 7. FUNGILLI Haeckel
Order Gregarinae Haeckel Gen. Morph. 2: xxv (1866).
Class Sporozoa Leuckart Parasiten der Menschen 1, part 1: 241 (1879).
Phylum FuNGiLLi Haeckel Syst. Phylog. 1: 90 (1894).
Class Sporozoaria Delage and Herouard Traite Zool. 1 : 254 (1896).
Subphylum Sporozoa Calkins Biol. Prot. 249 (1926).
Essentially unicellular organisms (the cells sometimes becoming multinucleate or
multiple, but remaining undifferentiated except in connection with reproduction);
commonly with a writhing motion; reproduction usually involving complicated sexual
processes and the production of walled cysts (spores); flagella absent except some-
times on the sperms; parasitic in animals.
The class Sporozoa as originally published by Leuckart included the following
groups: (a) the gregarines, first described by Dufour (1826) as worms parasitic in
beetles: the generic name Gregarina Dufour (1828) refers to their occurrence in
crowds; (b, c) coccidians and psorosperms, different sorts of parasites discovered in
fishes by J. Miiller and Retzius (1842); and, doubtfully, (d) Miescher's tubes
{Mieschersche Schlduche), being certain abnormal growths in muscles. The cause of
the pebrine disease of silkworms, which Nageli (in Caspary, 1857) had named A^o-
sema Bomhycis, belongs to this group but was not originally included, presumably
because Nageli had considered it to be a schizomycete.
It has subsequently become known that almost every species of the animal kingdom
is parasitized by one or more species of Fungilli. Not all of these parasites, but many,
are serious pathogens. Thus the Fungilli are a very important group and very num-
erous. The number of species duly registered by name and description is apparently
some two or three thousand; this is surely a small fraction of the number which exist.
The transmission of disease by biting arthropods was first demonstrated when
Theobald Smith (1893) showed that the Texas fever of cattle, caused by Babesia
bige?7iina, is transmitted by ticks.
All who have classified the Sporozoa or Fungilli have recognized two prime sub-
ordinate groups, the first including the gregarines and coccidians, the second includ-
ing the organisms which were formerly called psorosperms (Myxosporidia or Neo-
sporidia). In addition to the main bodies of these groups, there are certain organisms
which have resisted definite placement and have been assigned sometimes to one of
the main groups, sometimes to the other, and sometimes to additional main groups.
In the present work the two main groups are treated as classes and the groups of
uncertain relationship are included in the first. Clearly, this class is to bear the name
of Sporozoa Leuckart. Schaudinn's famous paper on parasites in the owl (1903) is
apparently authority for the widely entertained opinion that this class is artificial,
representing at least two lines of descent. In fact, the class appears natural with the
possible exception of some of the poorly known groups. The second class is marked
by positive specialized characters and is clearly natural; it is not clearly certain that
the second class is related to the first, and it is accordingly not certain that the phylum
is natural. The classes are distinguished as follows:
Phylum Fungilli [ 207
1. Producing resting cells protected by cell walls
and not containing polar capsules; or not
producing resting cells Class 1. Sporozoa.
1. Producing resting cells whose walls consist
(at least usually) of modified cells, and
which contain "polar capsules" enclosing
coiled threads Class 2. Neosporidia.
Class 1. SPOROZOA Leuckart
Class Sporozoa Leuckart Parasiten der Menschen 1, Abt. 1: 241 (1879).
Subclass Gregarinida Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1: Inhalt
(1882).
Class Sporozoaria and subclass Rhabdogeniae Delage and Herouard Traite Zool.
1: 254, 255 (1896).
Legion Cytosporidia Labbe in Thierreich 5: 3 (1899).
Subclass Telosporidia Schaudinn in Zool. Jahrb. Anat. 13: 281 (1900).
Class Telosporidia Calkins Biol. Prot. 421 (1926).
Class Telosporidca with subclasses Gregarinidia, Coccidia, and Haemosporidia;
and class Acnidosporidea Hall Protozoology 270, 271, 290, 301, 323 (1953).
Sous-emhranchement des Sporozoaires, with classes Gregarinomorpha, Coccidio-
morpha, and Sarcosporidia Grasse Traite Zool. 1, fasc. 2: 545 et seq. (1953).
Fungilli which produce resting cells protected by cell walls and not containing
polar capsules; or else do not produce resting cells.
The nature of the organisms included in this class may be made clear by an
example, Goussia Schubergi (Schaudinn) comb. nov. [Coccidium Schubergi Schau-
dinn, 1900).
Goussia is parasitic in centipedes. Infection is by certain spindle-shaped cells which
have a certain power of movement, and which make their way to the interior of cells
of the epithelium of the gut of the host. Each parasitic cell grows and becomes
globular; it becomes multinucleate; when the host cell dies and breaks up, the para-
sitic cell divides into many spindle-shaped cells which infect other cells of the gut
epithelium.
Alternatively, a sexual process takes place. Some of the parasites emerge into the
gut and do not divide but function as eggs. Others produce numerous cells which
are more slender than the usual infective cells. These become flagellate, each pro-
ducing two unequal flagella, and function as sperms.
The zygote becomes walled. Its nucleus divides twice. Each of the four resulting
nuclei becomes the nucleus of a walled cyst. The cysts are apparently formed by a
process of free cell formation: not all of the cytoplasm of the zygote is included in
them. In each cyst, two of the spindle-shaped infective cells are produced, again ap-
parently by free cell formation, excluding a part of the cytoplasm. The zygotes, with
their included cysts and infective cells, pass out with the feces of the host. If a centi-
pede eats one of them with its food, the infective cells are released to perform their
function.
No feature of the life cycle described is peculiar to the Sporozoa as contracted
with other nucleate organisms. Nevertheless, largely by the authority of Schaudinn,
specialists in Sporozoa use an extensive system of special terms. A familiarity with
these is necessary to anyone reading about Sporozoa. They include the following:
208]
The Classification of Lower Organisms
Sporozoite, the original infective cell.
Trophozoite, the vegetative individual.
Nucleogony, the multiplication of nuclei.
Plasmotomy, multiplication of cells.
Meront or schizont, the individual in process of dividing to produce further infec-
tive cells.
Schizogony or agamogony, the process of dividing to produce infective cells.
Merozoite or agamete, the infective cell produced from a trophozoite.
Gamogony, the production of gametes.
Macrogamete and microgamete mean, of course, egg and sperm; macrogametocyte
and microgametocyte mean the cells which produce them.
Sporoblast or sporont, the zygote or other cell inside of which walled cysts are
produced.
Sporogony, the sexual cycle which produces walled cysts.
Sporulation, the production of walled cysts by asexual processes.
Spore, the walled cyst.
Trophozoite (or schizont) and sporont are regarded as the alternating main stages
in the life cycle of Sporozoa. The point at which meiosis occurs is uncertain. In the
Fig. 41. — Life cycle of Goussia Schubergi after Schaudinn (1900) : a, sporozoites;
b-d, developing trophozoites; e, schizogony; f, merozoites; g, young gamctocytes;
h. i, development of egg; j-m, development of sperms; n, fertilization; o, zygote
(sporoblast); p-t, development of spores; u, germination of spores.
Phylum Fungilli [ 209
monocystid gregarines, Muslow (1911) and Calkins and Bowling (1926) described
a reduction of the chromosome number immediately before gametogenesis, quite as
in typical animals. They described reduction as accomplished by a single process of
nuclear division; to current cytological theory, this is an impossibility. Dobell and
Jameson (1915), Jameson (1920), and Dobell ( 1925), dealing with organisms of the
same group and also with the coccodian Aggregata, found meiosis to occur immed-
iately after karyogamy. They conclude that all nuclei except those of zygotes are hap-
loid, as among most of the lower plants.
The coccidian group, to which Goussia belongs, is here treated as primitive among
Sporozoa because the sperms of this group are flagellate. The detailed structure of
the flagella is unknown; they appear to resemble those of Bodo and Cryptobia. This
fact conveys the best available hint as to what may have been the evolutionary origin
of the Sporozoa. The majority of Sporozoa, having gametes which are alike or
scarcely differentiated, appear to be derived from forms with markedly differentiated
gametes.
The Sporozoa are classified primarily by whether or not the trophozoites are intra-
cellular; by the occurrence or non-occurrence of asexual reproduction; and by the
production or non-production of spores in the sense in which the term is used in
deaHng with this group, that is, of walled cysts.
1. Sexual reproduction, so far as it is known,
involving oocytes which produce single large
eggs and spermatocytes which produce from
few to many sperms; the organisms multiply-
ing also asexually.
2. The gametocytes not attached in pairs.
3. Producing walled spores Order 1. Oligosporea.
3. Not producing walled spores.
4. Intracellular in erythrocytes Order 3. GYMNOSPORiDnoA.
4. Producing macroscopic bodies
in muscle Order 4. Dolichocystida.
2. The gametocytes pairing before gameto-
genesis; sperms few; with or without
walled spores Order 2. Polysporea.
1. Gametes slightly differentiated or undifferen-
tiated, produced by the gametocytes in more
or less equal, usually large, numbers.
2. The organisms multiplying also asex-
ually.
3. Spores producing several sporozoites. . . . Order 5. Schizogregarinida.
3. Each spore producing one sporozoite. . . . Order 8. Haplosporidhdea.
2. The organisms not multiplying asexually.
3. Cells not elongate and divided into
two parts Order 6. Monogystidea.
3. Cells elongate and divided into two
parts Order 7. Polycystidea.
Order 1. OUgosporea Lankester in Enc. Brit. ed. 9, 19: 855 (1885).
Tribe Monosporees and groups Disporees and Tetrasporees Schneider in Arch.
Zool. Exp. Gen. 9: 387 (1881).
210] The Classification of Loivcr Organisms
Coccididae, with tribes Monosporea and Oligosporea, Biitschli in Bronn Kl. u.
Ord. Thierreichs 1: 574, 575 (1882).
Order Monosporea Lankester op. cit. 854.
Suborder Coccididae Delage and Herouard Traits Zool. 1 : 278 ( 1896).
Order Coccidiidia Lahhe in Thierreich 5: 51 (1899).
Order Coccidiomorpha Doflein Protozoen 95 (1901).
Order, suborder, or tribe Eimeridea Leger in Arch. Prot. 22 : 80 (1911).
Order Eimeriidea, suborders Selenococcidinea and Eimeriinea, and tribe Eimer-
ioidae, Poche in Arch. Prot. 30: 237, 238 (1913).
Subclass Coccidiomorpha and order Coccidia Calkins Biol. Prot. 435, 436
(1926).
Suborder Eimeridea Reichenow in Doflein Lehrb. Prot. ed. 5, 3: 921 (1929).
Order Eimeriida Hall Protozoology 297 (1953).
Sporozoa living mostly within epithelial cells of their hosts, multiplying asexually,
the gametocytes not pairing before gametogenesis, the macrogametocytes producing
single eggs and the microgametocytes numerous flagellate sperms, the zygotes usually
producing definite walled spores.
The organisms of the present order and the following are called coccidians.
Schneider ( 1881 ) classified them by the number of spores produced in each sporoblast
(i.e., zygote), either one, two, four, or many. Biitschli and Lankester gave due form
to Schneider's system. As between their names Monosporea and Oligosporea, the one
which included the typical example Eimeria is here chosen in preference to the one
which had page priority. Leger classified these organisms primarily by the number of
sporozoites per zygote, and distinguished eight families. Here, with the authority,
for example, of Reichenow (1929) and Kudo (1931), these are reassembled as one
family to which are appended three others including markedly exceptional or poorly
known forms.
Family 1. Eimerida [Eimeridae] Minchin 1903. The typical coccidians. In
Eimeria Schneider {Coccidium Leuckart) the zygote produces four firmly-walled
spores each with two sporozoites. The spores are symmetrically ellipsoid and release
the sporozoites through a terminal pore. Species of this genus parasitize many verte-
brate hosts, rabbits, sheep, goats, swine, dogs, cats, chickens, turkeys, frogs, and
fishes. Some of the other genera differ from this as follows: Jarrina, attacking birds,
is distinguished by spores bearing the pore at the end of a brief neck. Goussia, in cen-
tipedes, has spores whose walls split lengthwise into two valves. The zygote of Iso-
spora, in mammals, including man, produces two spores each with four sporozoites;
that of Caryospora, in snakes, produces one spore with eight sporozoites. Barrouxia,
in various invertebrates, produces from each zygote numerous bivalved spores each
containing one sporozoite.
Family 2. Dobelliida [Dobelliidae] Ikeda. The single known species, Dohcllia bi-
nuclcata, occurs in a siphuncuHd worm. It exhibits an exception to the characters
of the order: the male and female gametocytes become attached to each other; the
male gametocyte, however, produces many sperms, as in the generality of the order.
Family 3. Aggregatida [Aggrcgatidae] Labbe in Thierreich 5: 6 ( 1899). This fam-
ily is distinguished by hetcroocism. In Aggrcgata Ebcrthi, vegetative growth and
multiplication take place in crabs. When these are eaten by squids, the cells either
develop into single eggs or else divide to produce many sperms. The zygote produces
about twenty bivalved spores which pass out with the feces and infect crabs. The
number of sporozoites per spore is variable. There are several other species of Ag-
Phylum Fungilli [211
gregata. Various other genera, Merocystis, Hyaloklossia, Myriospora, Caryotropha,
etc., attacking mussels, polychaet worms, and other marine invertebrates, are as-
signed to this family although their life cycles are not fully known.
Family 4. Selenococcidiida [Selenococcidiidae] Poche in Arch. Prot. 30: 238
(1913) includes the single species Selenococcidium intermedium Leger and Dubosq
(1910) in the lobster. The vegetative cell is long and slender, and asexual reproduc-
tion is regularly by transverse division into eight.
Order 2. Polysporea Lankester in Enc. Brit. ed. 9, 19: 855 (1885).
Tribe Polysporea Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 576 (1882).
Suborder Haemosporidae Delage and Herouard Traite Zool. 1 : 284 ( 1896) .
Order Haemosporidiida Labbe in Thierreich 5: 73 (1899).
Order, suborder, or tribe Adeleidea Leger in Arch. Prot. 22: 81 (1911).
Tribe Adeleoidae Poche in Arch. Prot. 30: 239 (1913).
Order Adeleida with suborders Adeleina and Haemogregarinina Hall Proto-
zoology 296 (1953).
It is characteristic of this order that pairs of reproductive cells, essentially mero-
zoites, which are to become gametocytes, become attached to each other. The mac-
rogametocyte becomes converted into a single egg; the microgametocyte produces,
at least usually, four sperms.
Family 1. Adeleida [Adeleidae] Mesnil in Bull. Inst. Pasteur 1: 480 (1903).
Chiefly in invertebrates, either in the gut epithelium or in the kidneys, testes, or other
organs. Zygote usually producing definite spores, these numerous (commonly twenty
or more), thin-walled, without definite dehiscence mechanism, with two or four
sporozoites. Adelea and Adelina chiefly in centipedes; Klossia and Orcheobius in
snails; Klossiella in the kidney of the mouse; Legerella in various arthropods, the zy-
gote not producing spores but numerous sporozoites.
Family 2. Haemogregarinida [Haemogregarinidae] Liihe in Mense Handb. Tro-
penkrankheiten 3: 205 (1906). Heteroecious, with vegetative multiplication in the
tissues of a vertebrate host. The infection spreads to the erythrocytes of the host, and
blood-sucking invertebrates are infected by these. Sexual reproduction occurs in the
invertebrate host. Production of spores is suppressed; the zygote produces numerous
sporozoites. Haemogregarina Danilewski (1885; Drepanidium Lankester 1882, non
Ehrenberg 1861) in turtles, frogs, fishes, transmitted by leeches; Hepatozoon in ro-
dents, Karyolysus in lizards, transmitted by mites.
Order 3. Gymnosporidiida Labbe in Thierreich 5: 77 (1899).
Suborder Gymnosporidae Delage and Herouard Traite Zool. 1: 284 (1896).
Suborder Haemosporidia Doflein Protozoen 121 (1901).
Order Haemosporidia Calkins Biol. Prot. 441 (1926).
Subclass Haemosporidia with orders Plasmodiida and Babesiida Hall Proto-
zoology 301, 302, 306 (1953).
In this order the vegetative cells occur in vertebrates and infect the erythrocytes.
Sexual reproduction, so far as it has been discovered, occurs in blood-sucking arthro-
pods. The gametocytes do not become associated in pairs; the male gametocytes pro-
duce numerous spirochaet-like sperms by a process of budding. In the zygote, the
nucleus undergoes a series of divisions, after which numerous naked uninucleate
sporozoites are budded off from the surface. There are no walled spores.
212]
The Classification of Lower Organisms
The name Haemosporidia, commonly applied to this order, appears to belong by
priority to the preceding.
Schaudinn (1903) was disposed to connect this order with the trypanosomes, while
connecting the coccidians with Bodo and Cryptobia. This view has been entertained
by Liihe (inMense, 1906), Woodcock (1909), and Leger (1910). In spite of authority
thus good, it appears far-fetched. The Gymnosporidiida are of the same general na-
ture as the Aggregatida, Adeleida, and Haemogregarinida.
Fig. 42. — a-m. Life cycle of Plasmodium compiled from various sources: a, infec-
tion of an erythrocyte by a sporozoite; b-e, trophozoites, plasmotomy, and mero-
zoites; f, spermatocyte; g, oocyte; h, production of sperms; i, fertilization; j, k, pro-
duction of sporozoites in cells of the gut epithelium of the mosquito; 1, sporozoites;
m sporozoites entering the salivary gland of the mosquito, n-q. Stages of division
of cells of Babesia bigemina in erythrocytes of cattle x 2,000 after Dennis (1930).
The Gymnosporidiida are organized, somewhat arbitrarily, as three families.
Family 1. Halteridiida [Halteridiidae]Hartmann and Jollos 1910. Family Leu-
cocytozoidae Hartmann and Jollos. Family Hacmoproteidae Doflcin. Hacmoproteus
Kruse {Haltcridium Labbc) occurs in reptiles and birds. Vegetative growth and re-
production occur in tissue cells. Some of the merozoites infect erythrocytes, and are
believed to become gametocytes, and to develop no further unless swallowed by some
blood-sucking arthropod. In the best known example, H. Columbae of pigeons
Phylum Fungilli [ 213
(Argao, 1908), the alternate host is a fly. In the gut of the fly, the spermatocytes pro-
duce the elongate sperms as outgrowths. The zygotes make their way into the wall of
the gut of the fly, grow, and produce very numerous sporozoites. These migrate to the
salivary gland, from which they are injected into pigeons.
Leucocytozoon attacks birds; its cells become fairly large in certain blood cells
which become colorless and spindle-shaped.
Family 2. Plasmodida [Plasmodidae] Mesnil in Bull. Inst. Pasteur 1: 480 (1903).
The malaria organisms, differing from Haemoproteus in that they multiply in the
erythrocytes of their hosts. With a few obscure exceptions, the species are construed
as a single genus Plasmodium. Three species attack man; they have perhaps done
mankind more injury than any comparable group of living creatures. Several com-
paratively poorly known species attack apes and monkeys. The alternate hosts of all
species are mosquitoes of the genus Anopheles.
The vegetative individuals complete their growth within erythrocytes of their hosts
in more or less definite periods of time, and undergo multiple division; the erythro-
cytes then break up and release the merozoites. The chill and fever of malaria are as-
sociated with the destruction of erythrocytes. In the ordinary form of malaria, called
tertian malaria, development requires forty-eight hours, and the chill and fever occur
every other day. Another form, called malignant tertian or tropical malaria, exhibits
the same rhythm; it is distinguished by details of the appearance of the infected
erythrocytes. In the third form of malaria in man, called quartan, development re-
quires 72 hours, and the chill and fever occur every third day.
The course of development in the mosquito is quite like that of Haemoproteus
Columbae in the fly. Some of the parasites inside the erythrocytes are gametocytes;
each female gametocyte in an erythrocyte swallowed by a mosquito develops into a
single egg, while each male gametocyte buds off several spirochaet-like sperms. The
fertilized eggs are able to move. They break into the epithelium of the gut of the
mosquito, grow into large globes, and become multinucleate; their protoplasts divide
into numerous masses of protoplasm each of which buds off large numbers of sporozo-
ites. The sporozoites are released into the body cavity of the mosquito, migrate to
the salivary gland, and are injected into whatever animal the mosquito may bite.
The scientific names usually applied to the three species which cause human
malaria are not valid by priority. Extensive synonymy is given by Sabrosky and
Usinger, in their application to the International Commission on Zoological Nomen-
clature for action arbitrarily maintaining the current names (1944), and in the
report by Hemming (1950) of the action of the Commission.
Certain structures in the erythrocytes of malaria patients were first recognized as
parasites by Laveran, 1880, who, in 1881, named them Oscillaria malariae. The
organism is believed to have been that of malignant tertian or tropical malaria. The
word Plasmodium, properly designating a certain type of body, was applied by Mar-
chiafava and Celli 1885, in the combination Plasmodium malariae, believed also
originally to have designated the agent of malignant tertian malaria. Feletti and
Grassi, 1889, introduced the generic name Haemamoeba, with two species, H. vi-
vax, the agent of tertian malaria, and H. malariae, that of quartan malaria; it is be-
lieved that the latter epithet was applied under the misapprehension that this was the
organism which Marchiafava and Celli had named. It appears that Liihe, 1900, is
responsible for the currently used names:
Plasmodium vivax, the organism of tertian malaria;
P. malariae, that of quartan malaria;
214] The Classification of Lower Organisms
P. falciparum, that of malignant tertian.
In order that a great mass of literature may be read without confusion, it is ex-
pedient that these names be arbitrarily maintained. The International Commission
of Zoological Nomenclature has duly taken action to this effect.
Family 3. Babesiida [Babesiidae] Poche in Arch. Prot. 30: 241 (1913). Family
Theileridae du Toit in Arch. Prot. 39: 94 (1918). Minute intracellular parasites
transmitted by arthropods; sexual reproduction unknown. Theileria Bettencourt et al.
causes a fever of cattle in Africa; the parasites multiply in the tissue cells and spread
to the ery'throcytes, by which ticks are infected. Babesia Stercovici [Piro plasma Pat-
ton) is similar, but the parasites multiply in the erythrocytes. B. bigemina causes
the Texas fever of cattle.
The minute nucleus of Babesia bigemina is largely filled by a single granule, a
karyosome. This is connected by a rhizoplast to an extranuclear granule which has
been identified as a blepharoplast, although no flagellum is present. In nuclear divi-
sion, as described by Dennis (1930), the blepharoplast divides; the rhizoplast splits;
the nucleus widens, the karyosome becoming a rod; karyosome, nucleus, and cell
undergo constriction. No chromosomes are seen.
If Bartonella bacilliformis, the agent of the disease variously known as verruga
peruana, Oroya fever, or Carrion's disease, is not a bacterium, perhaps it may be
placed in or near this family.
Order 4. Dolichocystida Delage and Herouard Traite Zool. 1 : 289 (1896).
Sarcosporidia Balbiani 1882.
Class Sarcosporidia Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1 : Inhalt
(1882).
Subclass Sarcocystidca Lankester in Enc. Brit. ed. 9, 19: 855 (1885).
Order Sarcosporidia Doflein Protozoen 214 (1901).
Order Sarcocystidca Poche in Arch. Prot. 30: 245 (1913).
Subclass Sarcosporidia Calkins Biol. Prot. 461 (1926).
The characters are those of the single family and genus:
Family Sarcocystida [Sarcocystidae] Poche in Arch. Prot. 30: 245 (1913).
Sarcocystis Lankester produces the Mieschersche Schlauche, macroscopically visible
bodies, globular, fusiform, or filiform, of dimensions up to several millimeters, in
muscles of animals. The several supposed species, from mice, sheep, swine, deer, etc.,
are not morphologically distinguishable. Miescher observed these things in mice, in
which they are called Sarcocystis Muris; material from swine is called S. Miescher-
iana.
The visible body is a mass of cells, the whole walled by modified muscle of the
host. The mass originates as a single cell which divides repeatedly; the ultimate
division products are crescent-shaped uninucleate reproductive cells. Erdmann (1910)
observed the infection of epithelial cells of the gut of mice. Each infective cell grew
and divided into several, v\'hich made their way, or were carried, to the muscles, where
they gave rise to the Mieschersche Schlauche. Crawley (1914, 1916), on the other
hand, found the infective cells to be gametocytes. In cells of the gut epithelium of
the host, they may be converted as wholes into eggs, or else may give rise to numerous
elongate sperms. These conflicting observations could be explained by an alternation
of sexual and asexual generations, but the point is not established.
Phylum Fungilli [ 215
Order 5. Schizogregarinida Calkins Biol. Prot. 433 (1926).
Amoebosporidies Schneider.
Amoebosporidia Labbe in Thierreich 5: 120 (1899).
Suborder ^mo^feo^/^oncfm Doflein Protozoan 171 (1901).
^iuhordev Schizocystinea Poche in Arch. Prot. 30: 233 (1913).
Suborder Schizogregarinaria Reichenow in Doflein Lehrb. Prot. ed. 5, 3 : 872
(1929).
Orders Archegregarina and Neogregarina Grasse Traite Zool. 1, fasc. 2: 622,
665 (1953).
The Sporozoa previously considered, particularly those of the first two orders, are
called coccidians; those of the present order and the two which follow are called
gregarines. The latter are characterized (not without exceptions) by inter- instead
of intra-cellular active stages, and by the production of numerous gametes, alike or
not strongly differentiated, from paired scarcely differentiated gametocytes. The
present order includes the gregarines which exhibit asexual reproduction. They are
a rather miscellaneous assemblage.
Family 1. Schizocystida [Schizocystidae] Leger and Duboscq in Arch. Prot. 12:
102 (1908). Family Monoschizae V^eiser in ]our. Protozool. 2: 10 (1955), including
the two following families. In marine worms and other invertebrates. The sporozoites
enlarge in the host and become multinucleate individuals which reproduce freely by
producing uninucleate buds. Some of these buds continue the infection directly;
others become attached in pairs, each pair secreting a common cyst wall. Each of the
individuals in the cyst become multinucleate and buds off numerous uninucleate
gametes. The zygotes become walled spores which are cast out with the feces of the
host, to infect others which ingest them. Each produces eight sporozoites. Schizo-
cystis, Siedleckia.
Family 2. Seleniida [Seleniidae] Brasil in Arch. Prot. 8: 394 (1907). In marine
worms. Vegetative individuals notably long and slender; spores spiny, with four
sporozoites. Selenidiu7n, Meroselenidium.
Family 3. Merogregarinida [Merogregarinidae] Fantham 1908. Family Caul-
leryellidae Keilin. Merogregarina, Caulleryella, Tipulocystis.
Family 4. Spirocystida [Spirocystidae] Calkins Biol. Prot. 435 (1925). Family
Spirocystidees Leger and Duboscq in Arch. Prot. 35: 210 (1915). In earthworms.
Spores containing a solitary sporozoite which escapes through a pore. Spirocystis.
Family 5. Ophryocystida [Ophryocystidae] Leger and Duboscq in Arch. Prot. 12:
102 (1908). Family Amoebosporidiidae Brasil (1907), not based on a generic name.
Family Dischizac Weiser in Jour. Protozool 2: 10 (1955). In Ophryocystis Schneider
(Leger, 1907), the vegetative individuals, attached to the walls of the Malpighian
tubules of beetles, grow and become multinucleate and send out branches whose ends
develop into additional individuals. Eventually, different individuals become at-
tached in pairs. Each of these individuals buds off a single uninucleate gamete. The
remaining protoplasm of the gametocytes forms a protective sheath around the zygote,
which becomes a single spore with eight sporozoites.
Order 6. Monocystidea Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 574 (1882).
Order Haplocyta Lankester in Enc. Brit. ed. 9, 19: 853 (1885).
Suborder Acephalina Labbe in Thierreich 5 : 37 ( 1899) .
Organisms of the character of gregarines, not multiplying asexually, the vegetative
individuals not elongate and divided into serial parts.
216] The Classification of Lower Organisms
The genus which is best known is Monocystis Stein, including several species which
are common in earthworms. The cells grow within epithelial cells of the seminal fun-
nels; they and their nuclei reach considerable sizes without dividing. At maturity,
they escape into the seminal vescicles, where they form pairs, each pair secreting a
common cyst wall. The pairing and encystment were observed, more definitely of the
related genus Zygocystis than of Monocystis, by Stein (1848). The nuclei of the
paired cells divide. Several observers, as Brasil (1905) and Mulsow (1911); also, as
to related genera, Jameson (1920) and Noble (1938); have observed peculiarities in
the first nuclear division. The peculiarities amount to this, that the large nucleus
breaks up and, for the most part, undergoes dissolution, leaving a small number of
definite chromosomes to undergo normal mitosis in a spindle. Repeated subsequent
divisions are of normal character. The numerous nuclei thus produced become those
of gametes which are budded oflF from the surfaces of the gametocytes. This was first
observed by Wolters (1891). The gametes from the respective paired cells are pre-
sumably always of different mating types, and are usually visibly differentiated,
larger and smaller. Each zygote becomes a spindle-shaped walled spore; the enucleate
remainder of the gametocytes provides nourishment during their development. Each
spore produces eight sporozoites.
The number of known species of Monocystidea is of the order of 150. The majority
occur in annelid worms; others attack flatworms, echinoderms, insects, tunicates, and
other invertebrates. Bhatia (1930) distinguished twelve famiHes which are here
merely listed.
A. The two ends of the spore alike.
Family 1. Monocystida [Monocystidae] (Biitschli) Poche in Arch. Prot. 30: 236
(1913). Family Monocystiden Stein in Arch. Anat. Phys. 1848: 187 (1848). Mono-
cystidae Biitschli ( 1882). Monocystis, etc.
Family 2. Rhynchocystida [Rhynchocystidae] Bhatia in Parasitology 22: 158
(1930). Rhynchocystis.
Family 3. Stomatophorida [Stomatophoridae] Bhatia op. cit. 159. Stomatophora,
Choanocystis, etc.
Family 4. Zygocystida [Zygocystidae] Bhatia op. cit. 160. Zygocystis, Pleurocystis.
Family 5. Akinetocystida [Akinetocystidae] Bhatia op. cit. 160. Akinetocystis.
Family 6. Syncystida [Syncystidae] Bhatia op. cit. 161. Syncystis.
Family 7. Diplocystida [Diplocystidae] Bhatia op. cit. 161. Diplocystis, Lankcsteria.
Family 8. Schaudinellida [Schaudinellidae] Poche in Arch. Prot.' 30: 236 (1913).
Schaudinella.
B. The ends of the spores differentiated.
Family 9. Doliocystida [Doliocystidae] Labbe in Thierreich 5: 33 (1899). Family
Lecudinidae Kamm. Lxcndina Mingazzini {Doliocystis Legcr).
Family 10. Urosporida [Urosporidae] Woodcock 1906. Family Choanosporidae
Dogiel. Gonospora; Lithocystis; Urospora, the spores with long tails; Ceratospora;
Pterospora, the spores with longitudinal flanges.
Family 11. Ganymedida [Ganymcdidae] J. S. Iluxlcy in Quart. Jour. Micr. Sci.
n..s. 55: 169 (1910). Ganymcdcs.
Family 12. Allantocystidae [Allantocystidae] Bhatia op. cit. 163. Allantocystis.
Order 7. Polycystidea Biitschli in Bronn Kl. u. Ord. Thicrreichs 1: 578 (1882).
Order Grcgarinae Haeckel Gen. Morph. 2: xxv (1866), the mere plural of a
generic name.
Phylum Fungilli [217
Subclass Gregarinida Biitschli op. cit. Inhalt (1882).
Order Septata Lankester in Enc. Brit. ed. 9, 19: 853 (1885).
Order Brachycystida, suborder Gregarinidae, and tribe Cephalina or Folycystina
Delage and Herouard Traite Zool. 1: 255, 256, 269 (1896).
Order Gregarinida Labbe in Thierreich 5: 4 (1899).
^uhordtr Eugregarinaria Doflein Protozoen 160 (1901).
Order Gregarinoidca Minchin (1912).
Suborder Gregarininea and tribe Gregarinoidae Poche in Arch. Prot. 30: 234
(1913).
Subclass Gregarinida, order Eugregarinida, and suborder Cephalina Calkins Biol.
Prot. 422, 428 (1926).
The typical gregarines, the vegetative cells elongate and divided by more or less
definite constrictions into two (or, occasionally, more than two) parts; not repro-
ducing asexually.
Typical gregarines occur chiefly in insects. The vegetative cell consists of an an-
terior portion (protomerite) serving for attachment and a posterior portion (deuto-
merite), containing the nucleus, lying in the gut cavity of the host. Both parts have
a thick outer layer, commonly differentiated upon the protomerite into a more or
less elaborate knob, the epimerite. Longitudinal fibrils, presumably contractile, are
present. The cells writhe actively.
The individuals are commonly found in pairs, one member attached to the epi-
thelium of the gut, the other to the posterior end of the first. This arrangement is
produced by active self-placement on the part of the second member. When both are
mature, they take common action to produce a globular cyst. The protoplasts remain
distinct until both have become multinucleate, after which they produce numerous
gametes. In some forms, as Nina, studied by Goodrich (1938), all of the gametes
migrate from one cell, recognizably male, into the other, the female cell; the male
cell is left empty and is compressed or crushed by the growth of the zygotes in the
female cell. The zygotes are spores, usually fusiform, and usually producing sporo-
zoites by eights. In Gregarina and Gamocystis, an inner layer of the cyst wall is so
modelled as to form tubes (sporoducts) running from the surface to the interior.
When the spores are ripe, the sporoducts become extroverted and the spores are ex-
truded through them in uniseriate rows. In connection with this behavior, the spores
have flat ends like barrels.
Family 1. Stenophorida [Stenophoridae] Crawley 1903. Protomerite a mere
knob. Stenophora.
Family 2. Gregarinida [Gregarinidae] Greene 1859. Family Gregarinarien Stein
in Arch. Anat. Phys. 1948: 187 (1848). Gregarines which are without epimerites
and are not notably elongate. There are about a dozen genera. Cysts without sporo-
ducts: Hirmocystis, Hyalospora, Cnemidospora. Cysts with sporoducts, the spores
barrel-shaped: Gregarina, Gamocystis. The earliest observations of Sporozoa were by
Dufour ( 1826) , who, studying the anatomy of insects, found them in the gut of beetles.
He took them for worms and illustrated an individual with an epimerite, which he
took for a sucker. Later (1828) he applied names, Gregarina conica to the form first
seen, G. ovata to a form without an epimerite found in the forficule, i.e., in an ortho-
pteran. The former does not belong to the genus Gregarina as subsequently construed;
it appears to be a member of the family Actinocephalida. Gregarina ovata should be
regarded as the type of Gregarina, but the genus has usually been interpreted by G.
cuneata, which Stein observed in cockroaches.
218] The Classification of Lower Organisms
Family 3. Didymophyida [Didymophyidae] Wasilewski 1896. Family Didymo-
phyiden Stein (1848). Like the foregoing, but the cells extremely elongate. Didymo-
phyes.
Family 4. Acanthosporida [Acanthosporidae] Labbe in Thierreich 5: 27 (1899).
The spores with polar or equatorial bristles. Acanthospora.
Family 5. Stylocephalida [Stylocephalidae] Ellis 1912. Family Stylorhynchidae
Labbe op. cit. 30, based on a generic name which is a later homonym. Epimerite
elongate with a small terminal knob. Stylocephalus.
Family 6. Actinocephalida [Actinocephalidae] Wasilewski 1896. Epimerite with
thorns. Numerous genera, Sciadophora, Acanthorhynchus, Actinocephalus, Hoplo-
rhynchus, Pileoccphalus, etc.
Family 7. Menosporida [Menosporidae] Labbe op. cit. 29. Epimerite with a long
stalk, distally branched and bearing appendages. Menospora.
Family 8. Dactylophorida [Dactylophoridae] Wasilewski 1896. Epimerite dis-
tally broadened, clinging to the host epithelium by means of numerous filiform pro-
cesses. Dactylophorus, Nina [Pterocephalus), etc.
Family 9. Porosporida [Porosporidae] Labbe op. cit. 7. Heteroecious: in Porospora,
the gregarinoid stage occurs in crabs and the production of spores occurs in mussels.
The spores contain a single sporozoite and open through a pore.
Order 8. Haplosporidiidea Poche in Arch. Prot. 30: 178 (1913).
Order Aplosporidies Caullery and Mesnil 1899.
Order H aplosporidies Caullery and Mesnil in Arch. Zool. Exp. Gen. ser. 4, 4:
104 (1905).
Order Haplosporidia Auctt., the mere plural of a generic name.
Subclass Haplosporidia Hall Protozoology 326 (1953).
Unicellular intracellular parasites, the cells becoming multinucleate and multiply-
ing by fragmentation, producing walled spores which germinate by releasing the
protoplasts as single sporozoites.
The vegetative body is of the type properly called a plasmodium. The nuclei and
the process of division, described by Granata (1914) are characteristic. The resting
nucleus contains an "axial rod" as well as a nucleolus-like body. In mitosis the axial
rod becomes converted into an intranuclear spindle. Individual chromosomes have
not been seen; the chromatin gathers in a mass about the middle of the spindle (the
figures are curiously diatom-like). The mass of chromatin, the nucleolus-like body,
and the entire nucleus, divide by constriction; the ends of the spindle persist as the
axial rods of the daughter nuclei. Eventually, the plasmodium secretes a thin wall
and the protoplast divides into uninucleate naked cells. Granata found that these
cells are gametes, and that conjugation takes place among gametes produced by the
same plasmodium. The zygotes become walled spores which germinate by casting
off a circular operculum and releasing the contents. If the life cycle is correctly
understood, we may suppose that these organisms are degenerate gregarinos.
In the present state of knowledge, it will be as well to treat the typical haplo-
sporidians as a single family:
Family Haplosporidiida [Haplosporidiidae] Caullery and Mesnil in Arch. Zool.
Exp. Gen. ser. 4, 4: 106 (1905). Families Bartramiidae and Coelosporidiidae Caul-
lery and Mesnil op. cit. 107. Characters of the order. Haplosporidium (spores with
appendages at both ends) and Urosporidium (spores with a single appendage) attack
Phylum Fungilli [219
chiefly annelid worms. Bartramia attacks rotifers; Ichthyosporidium is a serious
parasite of fishes; Coelosporidium attacks cockroaches.
The following family, of uncertain position, may tentatively be associated with the
Haplosporidiidea :
Family Metchnikovellida [Metchnikovellidae] Caullery and Mesnil in Compt.
Rend. Soc. Biol. 77: 527 (1914), Ann. Inst. Pasteur 33: 214 (1919). Secondary
parasites, intracellular in gregarines; cells naked at first, with very minute nuclei,
which become numerous, later converted into walled cysts of characteristic form, the
protoplasts undergoing division into uninucleate infective cells. Mctchnikovella,
Amphiamblys, Amphiacantha.
Class 2. NEOSPOR!D!A (Schaudinn) Calkins
Myxosporidia Biitschli in Zool. Jahresber. 1880: 162 (1881).
Subclass Myxosporidia Biitschli in Bronn Kl. u. Ord. Thierreichs 1, Abt. 1 : Inhalt
(1882).
Subclass Amoebogeniae Delage and Herouard Traite Zool. 1: 291 (1896).
Subclass Neosporidia Schaudinn in Zool. Jahrb. Anat. 13: 281 (1900).
Order Cnidosporidia Doflein Protozoen 177 (1901).
Class Cnidosporidia Poche in Arch. Prot. 30: 224 (1913).
Class Neosporidia and subclass Cnidosporidia Calkins Biol. Prot. 445, 448 (1926).
Subphylum Cnidosporidia Grasse Traite Zool. 1, fasc. 1: 129 (1952).
Class Cnidosporidea Hall Protozoology 311 (1953).
Fungilli whose resting cells contain polar capsules; are walled, at least usually,
by a layer of modified cells; and, in most examples, release a single infective cell.
As a general rule, the vegetative bodies of Neosporidia are plasmodia, i.e., naked
multinucleate bodies, usually freely capable of asexual reproduction by internal or
external budding. An entire small plasmodium may become converted into one or two
spores, or the spores may be cut out internally and produced continually. The spores,
at least in the two better-known orders, are structures formed from several cells; they
are not homologous with the spores of the proper Sporozoa. In most examples, only
one of the cells involved in the formation of a spore is fertile, and only one infective
protoplast is released on germination. Of the sterile cells, one or more become con-
verted into the structures called polar capsules. These resemble the nematocysts of
coelenterates : they contain a coiled hollow thread capable of swift extroversion.
Extroversion occurs during germination. Its significance is unknown. The presence
of polar capsules marks the class as a natural group.
Three orders are recognized:
1. Spores covered by two valves formed from
accessory cells Order 1. Phaenocystes.
1. Spores covered by three valves formed from
accessory cells Order 2. Actinomyxida,
1. Spores very minute, with a continuous mem-
brane Order 3. Cryptocystes.
Order 1. Phaenocytes Gurley in Bull. U. S. Fish Comm. 11 : 410 (1893).
Order N^maiocj^^ffrfa Delage and Herouard Traite Zool. 1: 291 (1896).
Order Phaenocystida Labbe in Thierreich 5 : 85 (1899).
220]
The Classification of Lower Organisms
Order Cnidosporidia Doflein Protozoen 177 (1901).
Order Myxosporidia Calkins Biol. Prot. 449 (1926).
Most species of this order parasitize fishes, living either in internal cavities or in
the tissue cells; fewer than a dozen species are known from miscellaneous other ani-
mals, amphibia, reptiles, insects, and worms. Most of these parasites are not
extremely injurious.
The infective protoplast which issues from a spore is, at least usually, binucleate.
The nuclei fuse and the fusion nucleus divides repeatedly as the plasmodium grows.
Fig. 43.— Diagram of the life cycle of Myxoceros Blennius after E. Noble (1941).
In the examples which are believed to be more primitive, the plasmodia are freely
capable of budding, and the mature plasmodia are rather small and are converted as
wholes into single or paired spores. In the remaining examples, the plasmodia do not
multiply by budding, but produce spores continually.
Noble (1941) described the mitotic process. There is a rather large intranuclear
centrosome, which divides, the daughter centrosomes moving to opposite sides of
the nuclear cavity. Four chromosomes appear; this is apparently constant throughout
the order. The nuclear membrane and the centrosomes disappear. No spindle has
been seen. The chromosomes divide, and the daughter chromosomes move apart and
melt into two masses. The masses swell, a nuclear membrane appears about each,
and a centrosome appears inside of each.
Phylum Fungilli [221
The spore-forming structure (sporoblast) is a protoplast with several nuclei; it is
either a whole small plasmodium, or half of one, or a protoplast cut out endogen-
ously within a plasmodium. Two of the nuclei are set apart in cells which become
converted into the valves of the spore. Two or four are set apart in cells which become
converted into polar capsules. Two, of which it is established that they have two
chromosomes each, are the nuclei of the infective protoplast.
In a review of the literature as to Hfe cycles, Noble (1944) remarks as follows. "A
survey of the literature reveals that there is little agreement on the details of nuclear
changes in the Myxosporidia. Some authors maintain that the cycle is mainly haploid,
others have described a diploid cycle. Some reports indicate that there are two reduc-
tion divisions and two zygotes in one cycle. When only one zygote is reported the
reduction division in one case occurs just before fertilization, in another case it occurs
just after fertilization. Some authors have maintained that there is no sexual process."
Noble's own conclusions include the following. The organisms are diploid at most
stages. The meiotic divisions are among those by which the sporoblast becomes multi-
nucleate. The two haploid nuclei of the spore, which unite after germination, are
derived from a single diploid nucleus. Authors who have described fusions of proto-
plasts, or transfers of nuclei from one protoplast to another, have had no evidence
beyond an understandable unwillingness to accept fusions of sister nuclei.
Nearly two hundred species of the present order are listed in the monograph of
Kudo (1920), who established three suborders.
A. Valves conical, spores biconic (suborder Eurysporea Kudo).
Family 1. Myxoceratida nom. nov. Family Ceratomyxidae Doflein Protozoen 182
(1901), based on a generic name which is a later homonym. Characters of the sub-
order. Myxoceros nom. nov. [Ccratomyxa Thelohan 1892, non Ceratiomyxa Schroter
1889; if ever names are homonymous without being absolutely identical, these are.)
Some thirty-five species; the type is M. sphaerulosa (Thelohan) comb, nov.; Noble
studied mitosis in M. Blennius (Noble) comb. nov. Leptotheca, Myxoproteus, War-
dia, Mitraspora.
B. Valves hemispherical, spores spherical (suborder Sphaerosporea Kudo) .
Family 2. Chloromyxida [Chloromyxidae] Gurley in Bull. U. S. Fish Comm. 1 1 :
418 (1893). Chloromyxees Thelohan in Bull. Soc. Philomath. Paris ser. 8, 4: 176
(1892). Chloromyxea Braun in Centralbl. Bakt. 14: 739 (1893). With four polar
capsules. Chloromyxum.
Family S.Sphaerosporida [Sphaerosporidae] Davis 1917. With two polar cap-
sules. Sphaerospora, Sinuolinea.
C. Valves saucer- or boat-shaped, spores disk-shaped or fusiform (suborder
Platysporea Kudo ) .
Family 4. Myxidiida [Myxidiidae] Gurley op. cit. 420. Myxidiees Thelohan op.
cit. 175. Myxidiea Braun I.e. Myxidium, Sphaeromyxa, Zschokkella.
Family 5. Coccomyxida [Coccomyxidae] Leger and Hesse 1907. Coccomyxa.
Family 6. Myxosomatida [Myxosomatidae] Poche in Arch. Prot. 30: 230 (1913).
Myxosoma, Lentospora.
Family 7. Myxobolida [Myxobolidae] Gurley op. cit. 413. Myxobolees Thelohan
op. cit. 176. Myxobolea Braun I.e. Myxoboliis, Henneguya, Hoferellus.
Order 2. Actinomyxida Stole 1911.
This order includes about a dozen parasites in annelid worms. A plasmodial stage
and asexual reproduction are believed not to occur; the infective protoplast grows
222 ] The Classification of Lower Organisms
into an individual whose one or two nuclei remain undivided until the commence-
ment of the ill-understood process by which the complicated spores, with three valves
and three polar capsules, are produced.
Family 1. Tetractinomyxida [Tetractinomyxidae] Poche in Arch. Prot. 30: 231
(1913). Family Haploactinomyxidae Granata in Arch. Prot. 50: 205 (1925). Spores
subglobular, with a single binucleate sporozoite. Tetractinomyxon.
Family 2. Synactinomyxida [Synactinomyxidae] Poche I.e. Family Euactinomyxi-
dae Granata I.e. Family Triactinomyxidae Kudo Handb. Protozool. 314 (1931).
Spores producing eight or more sporozoites. S phaeractinomyxon and Neactinomyxon,
the spores subglobular. Synactinomyxon, with two of the valves protruding as con-
siderable horns, the whole horse-shoe shaped. Triactinomyxon and Hexactinomyxon,
all three valves drawn out into long horns, the whole caltrop- or anchor-shaped.
Order 3. Cryptocystes Gurley in Bull. U. S. Fish Comm. 11 : 409 ( 1893).
Microsporidies Balbiani 1882.
Order Microsporidiida Labbe in Thierreich 5: 104 (1899).
The parasites of this order attack chiefly arthropods and fishes. They multiply
asexually and produce serious epizootics. The spores are very minute, and the details
of the processes by which they are formed are unknown. A polar capsule is present in
each spore (those of Telornyxa have two polar capsules). The polar capsules are not
visible in living material, but are revealed by treatment with alkali.
In Kudo's monograph of this order ( 1924) , more than 150 species are treated. They
form four families.
Family 1. Glugeida [Glugeidae] Gurley op. cit. 409. Glugeidees Thelohan op. cit.
174. Glugeidea Braun I.e. Family Nosematidae Labbe in Thierreich 5: 104 (1899).
Family Plistophoridae Doflein Protozoen 205 ( 1901 ) . Spores oval, ovoid, or pyriform.
Nosema Bombycis Nageli causes the pebrine disease of silkworms; another species of
Nosema causes an epizootic of honeybees. Gliigea attacks several species of fishes.
Gurleya, Thelohania, Duboscquia, Plistophora, etc.
Family 2. Coccosporida [Coccosporidae] Kudo Handb. Protozool. 323 (1931).
Family Cocconemidae Leger and Hesse 1922, based on a generic name which is a
later homonym. Spores globular. Coccospora Slavinae (Leger and Hesse) Kudo, in
the oligochaet worm Slavina.
Family 3. Mrazekiida [Mrazekiidae] Leger and Hesse 1922. Spores elongate,
exceedingly minute, resembling bacteria. Mrazekia, Octospora, Spironema,
Toxonema.
Family 4. Telomyxida [Telomyxidae] Leger and Hesse 1910. Telornyxa glugei-
formis, in the fat body of the larva of Ephemera vulgata, producing ellipsoid spores
with a polar capsule at each end.
Chapter XII
PHYLUM CILIOPHORA
Phylum 8. CIUOPHORA (Doflein) nomen phylare novum
Class Infusoires Lamarck Phil. Zool. 1: 127 (1809).
Class Infusoria Lamarck Anim. sans Verteb. 1: 392 (1815).
Class Protozoa Goldfuss in Isis 1818: 1008 (1818).
Class Polygastrica Ehrenberg Infusionsthierchen p.* (1838).
Hauptgruppe Protozoa, class Infusoria, and order Stomatoda Siebold in Siebold
and Stannius Lehrb. vergl. Anat. 1 : 3, 10 ( 1848).
Subkingdorn Archezoa Perty Kennt. kl. Lebensf. 22 (1852), not phylum Archezoa
Haeckel (1894).
Order Ciliata Perty op. cit. 137.
Subphylum Infusoria Haeckel Gen. Morph. 2: Ixxviii (1866).
Phylum Infusoria Haeckel Syst. Phylog. 1: 90 (1894).
Subphylum Ciliophora Doflein Protozoen 227 (1901).
Dependent organisms, mostly predatory, unicellular but mostly of complicated
structure; swimming by means of cilia at least at some stage of life; mostly with
nuclei of two types in each cell. Vorticella, the only genus named by Linnaeus, is to
be considered the type.
These organisms are the typical examples of the accepted groups Infusoria and
Protozoa. The name Infusoria, referring to creatures which appear in infusions, is
said to have been introduced by Ledermiiller, 1763, or Wrisberg, 1764. As a scien-
tific name it has status from its application to a class by Lamarck (1815). The name
Protozoa, applied to a class in its original publication by Goldfuss, is a later synonym
of Infusoria. In treating the group as a phylum, one finds it necessary to apply a new
name, and takes up as such the name which Doflein applied to it as a subphylum.
The essential point in the definition of the phylum is the word cilia. Cilia are cell-
organs of the same nature as flagella, differing in being smaller in proportion to the
cell which bears them, more numerous, and distributed generally on the surface. In
Loeffler's classic investigation (1889), they were found to bear solitary terminal ap-
pendages; by subsequent terminology, they are acroneme. Doflein appears to have
been mistaken in emphasizing the difference between flagella and cilia; there is no
fundamental difference. A verbal distinction, nevertheless, is expedient: the applica-
tion of the term ciHum is to be restricted to two things, (a) the swimming organelles
of the Ciliophora, and (b) moving fibrils protruding abundantly from certain epithe-
Hal cells of animals. Botanical usage, which treats cilium and flagellum as synonyms,
is unsound. The structures which in botany have been called cilia are definitely
flagella.
The cells of Ciliophora reach moderately large sizes; those of the classroom
example Paramaecium attain a length of 0.25 mm. and are perceptible to the naked
eye. The cells of some of the Ciliophora are the most highly compHcated of all indi-
vidual cells. In addition to the cilia, the cell organs which require discussion are the
pellicle, neuromotor fibrils, trichocysts, structures involved in nutrition, contractile
vacuoles, and nuclei.
The cell has a firm ectoplasm or pellicle which gives it a definite form. The cilia
spring from basal granules imbedded in the pellicle. In simpler examples, the cilia
224 ] The Classification of Lower Organisms
are essentially uniform and uniformly distributed on the surface. Other examples
are without separate cilia upon part or all of the surface, but bear a variety of struc-
tures which consist of coalescent cilia. Membranelles are triangular appendages con-
sisting of brief rows of ciHa; undulating membranes represent long rows; cirri represent
tufts. The organisms of class Tentaculifera bear cilia only in the juvenile condition.
At maturity they bear extensible tubular structures called tentacles, by means of
which they capture free-swimming ciUates and absorb their contents.
The basal granules of the cilia are linked together by a system of fibrils; the cilia
and fibrils make up the neuromotor apparatus. This term was coined by Sharp, in his
study of Diplodinium (1914). The neuromotor fibrils form a highly elaborate net-
work, not connected with the nucleus, as in flagellates, but to a central structure,
apparently regulative, called the motorium. The motorium of Diplodinium is a mas-
sive body near the anterior end; that of the tintinnids is fairly large in proportion to
the cells (Campbell, 1926, 1927); that of Paramaecium, presumably a comparatively
primitive organism, is a minute body lying near the dorsal side of the cytopharynx
(Lund, 1933).
Imbedded in the pellicle, in addition to the neuromotor fibrils, there are certain
minute ellipsoid bodies called trichocysts. These, when the cell is irritated, discharge
their contents in the form of elongate rods or threads. Their mechanism and effect
are not understood.
In most Ciliophora, each cell has a mouth and gullet; or better, since these struc-
tures are not homologous with those of animals, a cytostome and cytopharynx. The
cytopharynx is a more or less funnel-shaped impression in the cell. It is bounded
laterally by ciliate pellicle; its outer opening is the cytostome; it is closed at the inner
end by a layer of cell membrane directly over fluid cytoplasm. Prey, chiefly bacteria
and small algae, encountered by the organism as it swims, is swept into the cyto-
pharynx by the action of the cilia. When a certain mass of prey has accumulated,
the cell membrane at the inner end of the cytopharynx becomes impressed and under-
goes constriction, enclosing the prey in a food vacuole. The material in the food
vacuole undergoes digestion; while this is taking place, movement of the cytoplasm
carries the vacuole along a definite circuitous course through the interior of the cell.
After some time, the vacuole arrives at a certain point on the pellicle, the anus or
cytoproct, where it discharges its contents and disappears by bursting through the
pellicle.
In freshwater species, each cell contains one or more contractile vacuoles which
appear at definite points and disappear periodically by discharge of their contents
to the exterior. Associated with the proper contractile vacuoles, there may be systems
of "canals" which are in fact additional contractile vacuoles. These structures have
been much studied; there are notable accounts by Day (1930) and Mac Lennan
(1933). When a vacuole has disappeared by discharge, it reappears as one or more
minute vacuoles in the same area: minute bodies of gelled protoplasm turn into sol,
and then become lifeless liquid. The discharge of a "canal" into the proper con-
tractile vacuole occurs by dissolution of the bounding membranes of gelled proto-
plasm where the two are in contact, followed by contraction of the membrane of the
canal. The proper contractile vacuole discharges by essentially the same mechan-
ism. Its membrane meets and becomes fused with the bounding membrane of the
cell, generally at the end of a brief channel through the pellicle; the combined mem-
brane breaks, and the membrane of the vacuole contracts.
Phylum Ciliophora [ 225
Earlier biologists supposed tliat the contractile vacuole is an excretory mechanism.
More probably, its function is purely hydrostatic, to rid the cell of the water which
is constantly entering by osmosis. Marine Ciliophora have no contractile vacuoles.
In many members of the family Opalinoea each cell has many similar nuclei. These
divide, from time to time, by mitosis. Cell division takes place independently of
nuclear division, by transverse constriction, when a certain size has been reached.
In the generaHty of Ciliophora, each cell has one or more nuclei of each of two
types, macronuclei, which are conspicuous, and micronuclei, discerned with difficulty.
Cell division occurs by transverse constriction and is necessarily associated with
nuclear division. The macronucleus becomes elongate and divides by constriction
without any formation of chromosomes; in other words, amitotically. The micro-
nucleus also becomes elongate and divides by constriction. Early observers supposed
this process also to be amitotic. Actually, there appear within the intact nuclear
membrane a spindle and a definite number of chromosomes. Reichenow (editing
Doflein, 1927) compiled the following diploid counts:
Stentor coeruleus 28
Didinium nasutum 16
Chilodon uncinatus 4
C arche Slum poly pinum 16
Turner (1930) found 8 in Euplotes Patella. Thus the chromosome numbers of
Cihophora appear usually to be small powers of 2.
The chromosomes duly undergo division, the daughter chromosomes going to dif-
ferent ends of the nuclear cavity. The nucleus becomes greatly elongate and its mem-
brane presses in from the sides and cuts it in two. Turner observed in the axis of the
spindle of Euplotes a rather small endosome which becomes elongate and undergoes
constriction while the chromosomes are forming.
Opalina has a sexual process in which the multinucleate cells divide into many
uninucleate gametes. These are sexually differentiated, larger and smaller; they
duly unite in pairs and the zygotes grow and become ordinary multinucleate
individuals.
In the generality of Ciliophora, early observers discovered a sexual process in
which the cells, apparently undifferentiated, join in pairs but maintain their individ-
uality. The uniting cells become attached to each other in definite positions: in
Paramaecium, by their ventral or mouth-bearing surfaces; in Euplotes, by the left
halves of their broad ventral surfaces; in the ophryoscolecids and various other groups,
by their anterior ends. They remain attached, while continuing to swim, for several
hours, during which an exchange of nuclei takes place, and then resume their separate
life. Calkins (1926) was disposed, contrary to historical usage, to confine application
of the term conjugation to this exceptional form of syngamy.
The nuclear details of conjugation were described by Maupas (1889) and Richard
Hertwig (1889), whose observations have repeatedly been confirmed. When a pair
have joined, their macronuclei divide several times; the ultimate fragments are
digested and disappear. The micronuclei also divide, concurrently in both conju-
gants, a fixed number of times, in Paramaecium three, in Euplotes four. These divi-
sions include a meiotic process. Most of the haploid nuclei produced are digested;
as a general rule, only one survives to undergo the final division, which is mitotic,
producing in each conjugant two genetically identical haploid nuclei. By this time
a cytoplasmic connection has been established between the conjugants. In Paramae-
cium, the spindles of the mitotic final nuclear divisions extend through this connection,
226 ] The Classification of Lower Organisms
so that when mitosis is complete each protoplast contains two haploid nuclei of dif-
ferent origin. In other ciliates the same result is attained, apparently, by the migration
of one nucleus of each pair. Karyogamy takes place in each conjugant. The cyto-
plasmic connection is broken and the conjugants separate from each other. During
several subsequent hours, the zygote nucleus undergoes a characteristic number of
divisions, three in Paramaecium. Among the nuclei produced, one usually enlarges
and becomes a macronucleus; others, of the number characteristic of the form, survive
as micronuclei; the remainder are digested.
In Vorticella and its allies, syngamy consists of the complete fusion of a smaller
swimming individual with a larger one attached by a stalk. The nuclear processes
are believed to be essentially as in other ciliates. The reproduction of the Tentaculi-
fera has not been much studied, but here also the nuclear changes are as in the
generahty of ciliates (Noble, 1932).
The possibility of conjugation is limited by the occurrence of mating types. Certain
early observations had suggested the existence of these; the definite discovery was by
Sonneborn, in Paramaecium Aurelia (1937). Results of further study are available
in a symposium edited by Jennings (1940) and in a review by Kimball (1943). To
current knowledge, then:
Paramaecium caudatum includes four mating types divided into two groups; types
I and II conjugate with each other, and types III and IV with each other, but the
two groups are mutually sterile.
Paramaecium Aurelia includes eight mutually sterile groups, each of two mutually
fertile mating types.
Paraviaecium Bursaria includes three mutually sterile groups. The first group is
of four types, each self-sterile but able to conjugate with any other; the second group
is of eight such types, and the third again of four.
Paramaecium. multimicronucleatum is without mating types; any race can conju-
gate with any other
Euplotes Patella includes six mating types all in one group; each can conjugate
with any other.
The heredity of mating types is not understood. It is not a matter of simple Men-
dclian heredity. In Paramaecium Bursaria group I, the progeny of a cell of a given
mating type may include after conjugation either two or all four of the mating types.
The mating type of a line becomes fixed in connection with the first or second cell
division after conjugation, at the time that macronuclei are being differentiated; it
is accordingly believed that something in the macronuclei fixes the mating types.
So far as mating types are present, pure lines of ciliates cannot conjugate. Early
attempts to maintain pure cultures failed by death after intervals of some months.
These observations led to speculations that the vitality of protoplasm is limited, and
that sexual reproduction restores it. Woodruff, however, proved it possible to maintain
Parflmagau?n ^urc/m indefinitely without conjugation: he reported (1926) a culture
so maintained for sixteen years, an estimated eleven thousand generations.
The cultures are not thus persistent without nuclear change. At intervals, the macro-
nuclei break up and dissolve, and are replaced by new ones formed by division of the
micronuclei. Woodruff and Erdmann (1914) applied to this process of replacement
of nuclei the term cndomixis. It is not possible that this process is the genetic equiva-
lent of karyogamy. It is, presumably, the physiological equivalent of conjugation in
its feature of providing new macronuclei.
Phylum Ciliophora [ 227
Diller (1936) observed in P. Aurelia a different manner of replacement of nuclei,
by autogamy. In this process, the nuclei of a solitary cell go through the preliminaries
of conjugation; two haploid nuclei, sister products of one act of mitosis, unite to form
a zygote nucleus; and this divides in the usual manner to produce micronuclei and
macronuclei. Wichtermann (1939, 1940) observed that two cells, joined as in conju-
gation, may simultaneously undergo autogamy instead of exchanging nuclei.
In the normal conjugation of ciliates, the gamete nuclei produced in each cell,
being sister products of mitosis, are genetically identical; and the zygote nuclei pro-
duced after interchange are also genetically identical with each other. Autogamy is
believed to produce diploid nuclei which are completely homozygous. Thus the sexual
processes of the ciliates tend strongly to limit the variability of the progeny. This is a
peculiar and surprising feature of the group.
The ciliates have attracted experimental study, beyond what has already been
implied, of various functions, including nutrition, inheritance of acquired characters,
and regeneration after injury.
Hall and his associates (1940-1945) have shown that Colpidiinn campylum and
Tetraphymena Geleii (the latter is in their earlier papers called Glaucoma piriformis)
require thiamin and probably riboflavin. Nutritional requirements, rather than such
an entity as vitality, are presumably responsible for the limited life of early attempted
pure cultures. As to minerals, the same scholars demonstrated the necessity of Ca and
Fe: others have demonstrated the necessity of K, Mg, and P.
It has been observed of certain cultures in which the rate of division has been in-
creased by exposure to high temperature that they would continue to divide abnor-
mally rapidly when returned to normal temperatures. The peculiarity disappeared in
individuals which conjugated. By refrigeration or by application of chemicals, there
have been produced "monsters," individuals of abnormal structure, which have repro-
duced themselves through many generations, and have proved capable eventually of
giving rise to normal individuals. Jollos (1913) designated as Dauermodifikationen,
that is, enduring changes, modifications of the type described. They are actually
acquired characters which can be inherited within limits. It is evident that they are
determined by macronuclei or by cytoplasm, and that they are not in conflict with
the principle that the truly enduring heredity of nucleate organisms Hes in nuclei
which divide mitotically.
Balamuth (1940) reviewed the literature of experimental mutilation of Protozoa
and gave a bibliography of 173 titles. Most of the experiments have been performed
on ciliates. The conclusions from them include these, that regeneration of parts arti-
ficially cut away takes place with different degrees of facility in different groups,
and that it is effected, if at all, by the same mechanism by which the parts are pro-
duced after division or excystment. The less elaborate ciliates, as Opalina and
Paramaecimn, are usually killed by mutilation, since this allows the fluid inner
cytoplasm to escape. In Stentor, injury to the crown of membranelles results in the
appearance of a new crown of membranelles on the side of the body, followed by its
migration to the injured area. In Stylonychia and Euplotes, destruction of one cirrus
is followed by the appearance, in a certain area of the surface, of the primordia of a
complete set of cirri; the original cirri are absorbed, and the new ones migrate along
the surface to their proper stations. The regulation of regeneration is explained, as
are various other phenomena, in a review by Weisz (1954).
Micronuclei are necessary for unlimited hfe and for sexual reproduction, but not
for regeneration and a long period of Hfe. Schwartz kept a culture of Stentor alive
228 ] The Classification of Lower Organisms
without micronuclei for more than a year. Macronuclear material is necessary for
regeneration, but any fragment of a macronucleus is sufficient. This is a very signifi-
cant observation. It means that all the factors controlling the vegetative structure and
behavior of a cell can be spread out and intermingled in all parts of a body of con-
siderable size; it furnishes an analogy to the state of affairs which may be supposed
to exist in bacteria.
The Ciliophora are treated as two classes, Infusoria and Tentaculifera. Hartog
(1909) estimated the number of known species of the former as about five hundred.
This number would have included practically all of the fresh-water species known up
to the present. Entozoic and marine species were known, but hundreds of species of
these ecological groups have subsequently been discovered. Including some two
hundred species of Tentaculifera, the phylum Ciliophora appears to be of about
twelve hundred known species.
Class 1. INFUSORIA Lamarck
Class Ciliata Haeckel Gen. Morph. 2: Ixxviii (1866).
Class Ciliatea Hall Protozoology 333 (1953).
Further synonymy essentially as of the name of the phylum.
Ciliophora lacking tentacles, bearing cilia or modified cilia in the mature condition.
Stein (1867) provided four orders of Infusoria. These orders are surely natural.
Subsequent authors have proposed many modifications of Stein's system, and many
of these are surely sound; but among groups proposed as additional orders, only the
opahnids are positively entitled to this status.
1. Nuclei all alike, commonly numerous. Order 1. Opalinalea.
1. Nuclei diflferentiated into macronuclei and
micronuclei.
2. Without a spiral band of membranelles
or cilia about the cytostome Order 2. Holotricha.
2. With a spiral band of membranelles or
cilia about the cytostome.
3. The spiral sinistrorse.
4. Not of the character of the fol-
lowing order Order 3. Heterotricha.
4. Flattened, cirri and most cilia
confined to the ventral surface Order 4. Hypotricha.
3. The spiral dextrorse Order 5. Stomatoda.
Order 1. Opalinalea nom. nov.
Suborder Opalininea Poche in Arch. Prot. 30: 250 (1913).
Protociliata Metcalf in Anat. Record 14: 89 (1918) and Jour. Washington Acad.
Sci. 8: 431 (1918).
Subclass Protociliata Kudo Handb. Protozool. 335 (1931).
Order Opalinida Hall Protozoology 113 (1953), preoccupied by family Opalini-
dae Claus.
Nuclei not differentiated into two types; cilia abundant, undifferentiated; sexual
reproduction by the complete union of differentiated minute uninucleate gametes.
Commensal in the gut of amphibia and fishes.
The group has been treated monographically by Metcalf (1923). A single family
is usually recognized.
Phylum Ciliophora [ 229
Family Opalinoea Pritchard 1842. Family Opalinaea Siebold in Siebold and
Stannius Lehrb. vergl. Anat 1: 10 (1848). Family Opalinina Stein Org. Inf. 2: 169
(1867). Family Opalinidae Glaus 1874. Family Protoopalinidae Metcalf 1940.
There are about 150 known species of four approximately equally numerous genera:
Protoopalina Metcalf, cylindrical, with one or two nuclei which are always found in
a stage of mitosis; Zelleriella Metcalf, similar, the cells flattened; Cepedia Metcalf,
cylindrical, with many nuclei; Opalina Purkinje and Valentin, flattened and
m.ultinucleate.
Order 2. Holotricha Stein Org. Inf. 2 : 169 ( 1867) .
Orders Gymnostomata and Trichostomata, and suborder Aspirotricha Biitschli
in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889).
Suborder Hymenostomata Hickson 1903.
Orders Gymnostomataceae and Aspirotrichaceae Hartog in Cambridge Nat.
Hist. 1: 137 (1909).
Order Holotricha with suborders Anoplophryinea, Gymnostomata, and Hymeno-
stomata Poche in Arch. Prot. 30: 250-255 (1913).
Order Holotrichida Calkins Biol. Prot. 376 (1926).
Infusoria with differentiated macronuclei and micronuclei, with simple cilia dis-
tributed generally over the surface of the body, not having membranelles in a spiral
band about the cytostome.
This group is the mass of the more primitive typical Infusoria, of numerous fami-
lies, not all of which are to be listed here. Arrangements of the families in other
groups than the three here maintained have been proposed and are presumably more
nearly natural.
a. Cytostome anterior. Suborder Gymnostomata (Biitschli) Poche. Suborder
Gym.no stom,ina Hall.
Family Enchelia Ehrenberg Infusionsthierchen 298 (1838). Family Enchelina
Stein Org. Inf. 2: 169 (1867). Family Enchelyidae Kent. Families Holophryidae and
Cyclodinidae Schouteden. Family Didiniidae Poche. Comparatively unspecialized
forms, radially symmetrical or nearly so. Enchelis O. F. Miiller; Holophrya, Chaenia,
Prorodon; Ichthiophthirius, becoming parasitic in the skins of fishes; Lacryviaria,
the cytostome at the end of an extensible proboscis; Didinium, barrel-shaped, with
the cilia confined to two belts, having an extensible proboscis by means of which it
seizes other Infusoria and through which it swallows them.
Family Colepina Ehrenberg op. cit. 316 includes the single genus Coleps. The cells
look like hand grenades of World War I: they are approximately barrel-shaped (the
axis more or less curved), the pellicle forming hardened quadrangular plates between
which the cilia project. The anterior cytostome can be opened widely to ingest other
Infusoria.
b. Cytostome lateral. Suborder Aspirotricha Biitschli.
Family Parameciina Perty (1852). Family Paramoecidae Grobben. Paramaecium
[Hill] O. F. Miiller Verm. Terr. Fluv. 1: 54 (1773). The name is variously spelled;
the spelling here used is Miiller's in what is believed to be the first publication under
binomial nomenclature.
Family Colpodaea Ehrenberg Infusionsthierchen 345 (1838). Family Colpodidae
Glaus 1879. Family Ophryoglenidae Kent 1882. Small forms, oval, bean-shaped,
or flattened. Ophryoglena, Glaucoma, Colpoda, Tetrahymena, and many others.
230 ] The Classification of Lower Organisms
Family Cyclidina Ehrenberg op. cit. 244. Family Pleuronemidae Kent. Family
Pleuronemina Biitschli (1889). Similar, with a conspicuous undulating membrane
along one side. Cyclidium and many other genera.
Family Urocentrina Claparede and Lachmann Etudes Inf. 1 : 134 ( 1858) . Family
Urocentridae Schouteden. Urocentrum, the single genus, top-shaped, with cilia con-
fined to two belts and a tail-like tuft, constantly whirling in the water.
Family Trachelina Ehrenberg op. cit. 319. Family Tracheliidae Kent. Having an
anterior proboscis, the mouth at the base of this. Trachelius, Dileptus, Lionotus,
Loxodes, etc.
Family Chlamydodontida [Chlamydodontidae] Glaus 1874. Family Chlamydo-
donta Stein, the mere plural of a generic name. Family Chilodontida Biitschli. Fam-
ily Nassulidae Schouteden. Flattened. The cytopharynx surrounded by longitudinal
rods, apparently of hardened protein, collectively forming a conical basket, enclosed
except when the cytostome is open for ingestion. Chilodon, Chlamydodon, Nassula.
c. Cytostome lacking; parasitic, mostly in invertebrates. Suborder Anoplophry-
INEA Poche; suborder Astomina Hall.
Family Anoplophryida [Anoplophryidae] and seven other families, all named by
Cepede, 1910.
Order 3. Heterotricha Stein Org. Inf. 2: 169 (1867).
Suborder Spirotricka, sections Heterotricha and Oligotricha Biitschli in Bronn
Kl. u. Ord. Thierreichs 1 : 1674 ( 1889).
Section Chonotricha Wallengren in Acta Univ. Lund 31, part 2, no. 7 : 48 ( 1895) .
Order Oligotricha Doflein Protozoen 240 (1901).
Orders Heterotrichaceae and Oligotrichaceae Hartog in Cambridge Nat. Hist.
1: 137 (1909).
Orders Heterotrichida and Oligotrichida Calkins Biol. Prot. 386, 388 (1926).
Suborder Entodiniomorpha Reichenow in Doflein Lehrb. Prot. ed. 5, 3: 1195
(1929); Order Chonotricha Reichenow op. cit. 1211; suborder Ctenostomata
Kahl ex Reichenow op. cit. 1024.
Orders Spirotrichida and Chonotrichida Hall Protozoology 380, 411 (1953).
Infusoria having a sinistrorsc spiral band of cilia about the cytostome, these cilia
united (except in family Spirochonina) in triangular-attenuate membranellcs; not
having the body flattened and the cilia or cirri confined to the ventral surface.
The peristomal apparatus of this order is an evidently derived character, so pecu-
liar as to appear to have evolved only once: in short, the order appears natural. There
are numerous subordinate groups. Several of these, of many species or of exceptional
character, have been segregated as additional orders; it is by an arbitrary decision
that they are here treated as suborders.
a. Comparatively unspecialized examples. Suborder Spirotricha Biitschli.
Suborders Hctcrolrichina and Oligolrichina Flail.
Family Plagiotomina Biitschli op. cit. 1719 (1889). Family Plagiotomidae Poche
(1913). Peristomal area narrow and elongate, extending from the anterior end to a
cyto.^tome located near the middle of one side. Blepharisyna. Spirostomum.
Family Bursarina Stein Org. Inf. 2: 169, 295 (1867). Family Bursariidae Kent.
Cytostome seated in a deep pit in one side of the body. Bursaria. Balantidium, para-
sitic in the gut of Amphibia and mammals; B. coli, a serious pathogen in man.
Family Stentorina Stein op. cit. 169, 217. Family Stentoridae Claus. Peristomal
area anterior, more or less transverse. Stcntor, sessile and obconic, familiar. Follicu-
Phylum alio phor a [231
Una, the posterior end seated in a chitinous lorica, the peristomal area broadly ex-
panded as two wings.
Family Halterina Claparede and Lachmann Etudes Inf. 1: 367 (1858). Family
Halteriidae Claus. Halteria, subglobular, with a single whorl of long cilia; familiar in
infusions, recognizable by the motion of the cells, alternately revolving slowly and
snapping violently from place to place.
b. Loricate, free-swimming. Suborder Tintinnoinea Kofoid and Campbell.
Suborder Tintinnina Hall.
Family Tintinnodea Claparede and Lachmann Etudes Inf. 1: (1858). Family
Tintinnidae Claus. Peristomal membranelles elongate and ciliate, the cylindrical or
conical body attached in and retractile into the lorica; characteristically with two
macronuclei and two micronuclei. Mostly marine. Kofoid and Campbell, who mono-
graphed the group ( 1929) , found it possible to distinguish the natural and subordinate
groups entirely by the structure of the lorica. They divided the former single family
into twelve and recognized more than three hundred species.
c. Laterally flattened, with a tough membrane and few cilia and membranelles.
Suborder Ctenostomata Kahl. Suborder Ctenostomina Hall.
Family Ctenostomida [Ctenostomidae] Lauterborn in Zeit. wiss. Zool. 90: 665
(1908). Kahl (1932) monographed the group and found twenty-five species, which
he arranged in six genera and three families.
d. Cylindrical, entozoic, with no ciliation except the membranelles. Suborder
Entodiniomorpha Reichenow. Suborder Entodiniornorphina Hall.
Family Ophryoscolecina Stein Org. Inf. 2: 168 (1867). Family Ophryoscolecidae
Claus. Becker (1932) reviewed previous studies of this group, examples of which
were first mentioned by Gruby and Delafond, 1843. He noted 71 species, of the
genera Entodinium, Diplodinium, Ophryoscolex, Epidinium, etc. (the genera were
first named by Stein) in the domestic ox; and 52 {Didesmis, Paraisotricha, Spirodin-
ium, Cycloposthium, etc.) in the horse. Dogiel (1927) monographed the family, but
it is certain that large numbers of species remain to be discovered in wild animals,
oxen and others.
The barrel-shaped cells are about 0.1-0.25 mm. long. The cytostome is anterior,
surrounded by the usual spiral band of membranelles; this may be broken up into
several partial files, and there may be belts or clusters of membranelles on other parts
of the body. The posterior end is drawn out into processes, one, few, or many, ob-
scure or prominent, horn-like or fringe-like. Internally, beside contractile vacuoles
and a neuromotor apparatus including a large motorium, there are characteristic
skeletal plates. These consist of minute cylindrical bodies imbedded in an amorphous
matrix, the whole staining with iodine and consisting supposedly of some polysac-
charide carbohydrate.
Animals are infected by eating food contaminated with the saliva of others. The
ciliates may be present in the rumen in numbers from one thousand to three million
per cc. It has been supposed that they are symbiotic, benefitting their hosts by carry-
ing on useful syntheses, or perhaps merely by controlling numbers of bacteria in the
rumen. There is no good evidence for these ideas: the probability is, that they are
harmless commensals.
e. Cylindrical or obconic, sessile, cilia of the peristomal band separate, body
otherwise naked. Suborder Chonotricha (Wallengren) subordo novus.
Family Spirochonina Stein Org. Inf. 2: 168 (1867). Family Spirochonidae Grob-
232
The Classification of Lower Organisms
Fig. 44. — Infusoria, order Hypotricha: a, Aspidisca x 800. b, Stylonychia
X 400. C, Euplotes x 400. d-n, Euplotes Patella after Turner (1930); d-h, stages
of mitosis x 2,000, i, conjugating cells x 400; j, k, polar and equatorial views of the
heterotypic division in a conjugant x 2,000; 1, early anaphase of the homeotypic
division x 2,000; m, first division of the zygote nucleus x 2,000; n, a cell after con-
jugation X 400, the macronucleus breaking up, the zygote nucleus divided into four,
of which one is to become a macronucleus, one a micronucleus, and two are to
undergo dissolution.
Phylum Ciliophora [ 233
ben. Spirochona and a few other genera, attached to aquatic animals, fresh-water or
marine, best known from the crustacean Gammarus.
Order 4. Hypotricha Stein Org. Inf. 2 : 168 ( 1867) .
Section Hypotricha Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889).
Order Hypotrichaceae Hartog in Cambridge Nat. Hist. 1: 137 (1909).
Order Hypotrichida Calkins Biol. Prot. 389 (1926).
Suborder Hypotricha Kudo Man. Protozool. ed. 3: 668 (1946).
Suborder Hypotrichina Hall Protozoology 381 ( 1953 ) .
Flattened Infusoria bearing a band of membranelles crossing the upper surface
near the anterior end from right to left and continued rearward on the lower surface
beside the cytostome, along which lie also undulating membranes; mostly bearing
cirri, which are confined to the lower surface, as are most free cilia, if these are
present.
This group is evidently natural, and evidently a specialized offshoot from the pre-
ceding order. It might reasonably be treated as a subordinate group of the preceding
order; Biitschli, Kudo, and Hall have done so. There are comparatively few species.
Several are familiar in infusions and have been much studied.
Family 1. Peritromina Stein Org. Inf. 2: 168 (1867). Family Peritromidae Kent.
Cilia abundant on the lower surface, cirri none. Peritromus.
Family 2. UrostyUda [Urostylidae] Calkins Biol. Prot. 390 (1926). As above, but
with frontal and sometimes also anal cirri. Numerous genera, Urostyla, Uroleptus,
Epiclintes, Stilotricha; Kerona O. F. Miiller, an ectoparasite on the animal Hydra.
Family 3. Oxytrichina Ehrenberg Infusionsthierchen 362 (1838). Family Oxytri-
chidae Kent. Family Pleurotrichidae Biitschli. Cirri present; cilia in one or two mar-
ginal rows, few or absent on the ventral surface. Oxtricha, Stylonychia, Pleurotricha,
Euplotes, etc.
Order 5. Stomatoda Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1: 10
(1848).
Order Ciliata Perty Kennt. kl. Lebensf. 137 (1852).
Order Peritricha Stein Org. Inf. 2: 168 (1867),
Section Peritricha Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889).
Order Peritrichaceae Hartog in Cambridge Nat. Hist. 1: 138 (1909).
Order Peritrichida Calkins Biol. Prot. 395 (1926).
Infusoria having a dextrorse spiral band of membranelles about the cytostome,
which can in most examples be concealed and protected by contraction of the body;
free-swimming only in the immature condition, at maturity attached and without
separate cilia; syngamy occurring by the complete union of a smaller swimming indi-
vidual with a larger attached one. Vorticella is the apparent type of the old ordinal
names Stomatoda and Ciliata, which are accordingly held to belong to this order.
Family Vorticellina Ehrenberg Infusionsthierchen 259 (1838). Family Vaginifera
Perty (1852). Family Vorticellidae Fromentel 1874. Vorticella L., a familiar mic-
roscopic organism in material from ponds and ditches, consists of solitary bell-shaped
cells on contractile stalks. Carchesium and Zoothamnium are similar organisms in
colonies. Ophrydium, Epistylis, etc., consist of similar colonies of non-contractile
cells. Cothurnia and Vaginicola are solitary stalkless cells having conical loricae into
which they can withdraw themselves.
234]
The Classification of Lower Organisms
Fig. 45 — Tokophyra Lemnarum after A. Noble (1932) : a, representative individ-
ual; b, budding; c, swimming bud; d, conjugation; e, feeding on a cell of Euplotes;
t, cyst; g, tentacles, feeding, expanded, and contracted, g x 2,000, all others x 400.
Phylum Ciliophora [ 235
Family Urceolarina Perty (1852). Family Trichodinidae Glaus. Family Urceol-
aridae Kudo. Urceolaria, Trichodina, etc., disk- or barrel-shaped cells attached on
or in aquatic animals by means of a whorl of hard hooks.
Class 2. TENTACULIFERA (Huxley) Kent
Order lufusoires suceurs and group Acinetina Claparede and Lachmann Etudes
Inf. 1: 377,381 (1858).
Class Acinetae Haeckel Gen. Morph. 2: Ixxix (1866), the mere plural of a generic
name.
Tentaculifera Huxley Man. Anat. Invert. 100 (1877).
Glass Tentaculifera with orders Suctoria and Acinetaria Kent Man. Inf. 1 : 36
(1880).
Class Acinetaria and order Suctoria Lankester in Enc. Brit. ed. 9, 19: 865 ( 1885).
Subclass Suctoria Butschli in Bronn Kl. u. Ord. Thierreichs 1: 1842 (1889).
Class Acinetoidea Poche in Arch. Prot. 30: 263 (1913).
Class Sudor ea Hall Protozoology 413 (1953).
Organisms swimming by means of cilia while immature, at maturity lacking cilia
and usually attached, provided with tentacles by which they capture and paralyze
their prey and absorb food. Acineta is the type genus.
These organisms are rather unfamiliar. They occur both in fresh water and in salt,
and prey chiefly upon Infusoria. There are differentiated macronuclei and micro-
nuclei; in branching or colonial individuals, a single macronucleus may extend to all
parts. Asexual reproduction is by budding, often endogenous. Conjugation occurs
either between attached individuals or between an attached individual and a swim-
ming bud. The fact that one individual may bend past another to conjugate with a
third indicates the presence of mating types. Conjugating individuals exhibit pregamic
and postgamic nuclear divisions quite as among Infusoria (Noble, 1932). The group
is undoubtedly derived from Infusoria; whether from something of the nature of
Didinium, Vorticella, or Spirochona remains uncertain.
Collin (1912) accounted for about 170 species and recognized eight families. One
of these families has subsequently been transferred to order Holotricha. The re-
mainder may be construed as a single order:
Order Suctoria Kent (1880). Lankester chose this as between two ordinal names
which Kent published at the same time.
a. Individuals subglobular, usually stalked, their tentacles essentially uniform.
Family 1. Podophryina Butschli in Bronn Kl. u. Ord. Thierreichs 1 : 1926 (1889).
Family Podophryidae, Rousseau and Schouteden 1907. Buds produced exogenously.
Podophrya, Sphaerophrya, naked; Urnula, loricate.
Family 2. Acinetida [Acinetidae] Glaus 1874. Acinetina Claparede and Lach-
mann (1858). Family Acinetina Biitschli (1889). Bodies with a thin pellicle, with or
without loricae; budding endogenous. Acineta, Tokophrya, etc.
Family 3. Discophryida [Discophryidae] Collin in Arch. Zool. Exp. Gen. 51: 364
(1912). Body with a firm pellicle, budding endogenous. Discophrya, etc.
b. Individuals branching or colonial.
Family 4. Dendrosomida [Dendrosomidae] Kent Man. Inf. 2: 215 (1882). Family
Dendrosomina BiitschU (1889). Family Dendrosomatidae Poche (1913). Dendro-
soma, etc.
236 ] The Classification of Lower Organisms
Family 5. Ophryodendrida [Ophryodendridae] Kent I.e. Family Ophryodendrina
Biitschli (1889). Ophryodendron, etc.
Family 6. Dendrocometida [Dendrocometidae] Kent I.e. Family Dendrocometina
Biitschli (1889). Dendrocometes, Stylocometes.
c. With differentiated tentacles for piercing and sucking.
Family 7. Ephelotida [Ephelotidae] Kent. I.e. Family Ephelotina Sand 1899.
Marine, individuals subglobular, stalked. Ephelota, naked; Podocyathus, loricate.
With this peculiar and highly evolved group, the here-proposed classification of
organisms which lack the distinctive characters both of plants and of animals is
concluded.
List of N omenclatural Novelties [ 237
LIST OF NOMENCLATURAL NOVELTIES
Page
P'amily Kurthiacea fam. nov 21
Family Pasteurellacea nom. nov 22
Family Chromatiacea nomen familiare novum 31
Family Rhodobacillacea nom. nov 31
Family Chlorobiacea nom. nov 31
Order Sphaerotilalea nom nov 33
Lagenocystis, nom. nov., and L. radicicola, comb, nov 82
Family Dinamoebidina nom nov 101
Phylum Opisthokonta phylum novum 110
Chilomastix hominis comb, nov 165
Pentatrichomonas obliqua comb, nov 167
Goussia Schubergi comb, nov 207
Family Myxoceratida and Myxoceros, nomina nova; M. sphaerulosa and
M. Blennius, combinationes novae 221
Phylum Ciliophora nomen phylare novum 223
Order Opalinalea nom. nov 228
Suborder Chonotricha subordo novus 231
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INDEX
OF NAMES OF ORGANISMS AND GROUPS
Absidia, 123, 124
Acantharia, 189, 190, 195, 196, 197
Acanthochiasma, 197
Acanthocystida, 191, 193
Acanthocystidae, 193
Acanthocystis, 193
Acanthometra, 195, 197
Acanthometren, 197
Acanthometrida, 197
Acanthometron, 197
Acanthonida, 197
Acanthophracta, 195, 197
Acanthorhynchus, 218
Acanthospora, 218
Acanthosporida, 218
Acanthosporidae, 218
Acaulopage, 124
Acephalina, 215
Acervulina, 187
Acervulinida, 187
Acetobacter aceti, 24
Acetobacteriacea, 20, 24
Acetobacteriaceae, 24
Achlya, 70, 79
Achlya caroliniana, 78
Achlyogeton, 118
Achlyogetonacea, 115, 117
Achlyogetonaceae, 117
Achnanthea, 76
Achnantheae, 76
Achnanthes, 76
Achnanthaceac, 76
Achromatiacea, 33
Achromatiaceae, 33
Achromatium, 33
Achromatium oxaliferum, 32, 33
Achromobacter, 22
Achromobacteriacea, 19, 21
Achromobacteriaceae, 2 1
Acineta, 235
Acinetae, 235
Acinetaria, 235
Acinetida, 235
Acinetidae, 235
Acinetina, 235
Acinetoidea, 235
Acnidosporidea, 207
Acrasidae, 203
Acrasina, 203
Acrasis, 203
Acrita, 37
Acrochaetiacea, 47
Acrochaetiaceae, 47
Acrochaetium, 47
Actinelius, 197
Actinollida, 197
Actiniscea, 61, 62
Actinisceae, 62
Actinocephalida, 217, 218
Actinocephalidae, 218
Actinocephalus, 218
Actinolophus, 193
Actinomma, 195
Actinomma Asteracanthion, 196
Actinomonadida, J 91
Actinomonadidae, 191
Actinomonas, 190, 193
Actinomyces Bovis, 25
Actinomycetaceae, 25
Actinomycetalea, 18, 24
Actinomycetales, 24
Actinomyxida, 219, 221
Actinophryida, 191, 193
Actinophryidae, 193
Actinophrys, 193
Actinophrys Sol, 193
Actinopoda, 189
Actinopodea, 189
Actinosphaerium, 193
Actinosphaerium Eichhornii, 192, 193
Actipylea, 195, 197
Actipyleen, 195
Actipylida, 197
Actipylina, 197 '
Acystosporidia, 190
Acyttaria, 179
Adelea, 211
Adeleida, 211,212
Adeleidae, 211
Adeleidea, 211
Adeleina, 21 1
Adeleoidae, 21 1
Adelina, 211
Adinida,_98, 99
Adiniferidea, 96, 98
Aecidium, 147
Aerobacter aerogenes, 22
Agaricacea, 151, 153
Agaricaceae, 150, 151
Agaricales, 150
Agaricini, 151
Agaricus campestris, 119, 145, 152, 153
Agarics, 152
Aggregata, 209
Ae;gregata Eberthi, 210
Aggregatida, 210, 212
Aggregatidae, 210
Aglaozonia, 88
Agrobacterium, 23
Agrobactcrium tumefaciens, 23
Agrostis, 148
Agyriales, 137
272]
The Classification of Lower Organisms
Agyrium, 137
Ahnfeldtia, 49
Akinetocystida, 216
Akinetocystidae, 216
Akinetocystis, 216
Akinetosporeae, 86
Albuginacea, 80, 81
Albuginaccae, 81
Albugo, 80, 81
Albugo Bliti, 80
Albugo Tragopogonis, 80
Alcaligenes fecalis, 22
Aleuria rutilans, 136
Algae, 9, 10, 69, 113, 118, 120, 177, 224
Algae, blue-green, 2, 3, 12, 13, 14, 17, 30,
37,41, 117, 118
Algae, brown, 39, 69, 53, 179, 203
Algae, green, 38, 41, 53, 69, 82, 117, 118,
128, 203
Algae, red, 37, 39, 41, 82, 128, 140
Algae Zoosporeae, 86
Algen, 29, 120
Allantocystida, 216
Allantocystidae, 216
Allan tocystis, 216
Allogromia, 183
Allogromiida, 183
Allogromiidae, 183
Allomorphina, 187
Allomyces, 111, 112, 113, 118
Allomyces anomalus, 116
Allomyces Arbuscula, 112, 115
Allomyces cystogenes, 112
Allomyces javanicus, 112, 114
Almond, 141
Alveolina, 185
Alveolinea, 185
Alveolinella, 185
Alveolinellidae, 185
Alveolinida, 185
Alveolinina, 185
Alternaria, 142
Alwisia, 175
Amanita, 152
Amanita muscaria, 152
Amaurochaetacea, 174, 175
Amaurochaetaceae, 175
Amaurochaete, 175
Amaurochacteae, 171
Amaurochaetidae, 175
Amaurosporales, 171
Amiba diffluens, 37, 157, 201
Amiba divergcns, 202
Ammodiscida, 185
Ammodiscidae, 185
Ammodiscus, 185
Ammodochidae, 62
Amoeba, 71, 118, 124, 157, 189, 201
Amoeba Proteus, 202
Amorbaca, 201
Amocbida, 10,201
Amocbidac, 10, 201
Amoebina, 201
Amoebodiniaceae, 101
Amoebogeniae, 219
Amoebosporidia, 215
Amoebosporidies, 215
Amoebosporidiidae, 215
Amorphoctista, 37
Amorphozoa, 37
Amphibia, 220
Amphiacantha, 219
Amphiamblys, 219
Amphidinium, 101
Amphilonche, 197
Amphilothida, 103
Amphimonadaceae, 61, 158
Amphimonadidae, 61
Amphisolenia, 103
Amphisolenia laticincta, 104
Amphistegina, 187
Amphistomina, 191
Anabaena, 35
Anabaena circinnalis, 13
Anabaena inaequalis, 32
Ancylistales, 81
Ancylistes, 125
Ancylistinaeae, 81
Anemeae, 171
Angiococcus, 28
Angiogastres, 152
Angiospermeae, 82, 91
Animacule, 18
Animal kingdom, Animalia. Animals, 1, 2,
4, 6, 10,' 68, 95, 111, 113, 159, 163,
167, 206, 214, 220, 223, 231, 233, 235
Anisochytridiales, 69
Anisochytrids, 57
Anisolpidiaceae, 69
Anisoplidium, 69
Anisolpidium Ectocarpii, 70
Anisonema, 109
Anisonema truncatun, 108
Anisonemida, 105, 108
Anisonemidae, 108
Anisonemina, 108
Anomalinidae, 187
Anopheles, 213
Anoplophryida, 230
Anoplophryinca, 229, 230
Anthophysis, 59
Anucleobionta, 6, 12
Ape, 213
Aphanizomenon, 35
Aphanomycopsis, 81
Aphrothoraca, 190, 193
Aphrothoracida, 190
Aplanosporeao, 86
Aplosporidics, 218
Apodachlya, 79
Apodachlyclla, 79
Apodinidae, 102
Apodinium, 102
Appcndiculatae, 73
Index
[273
Apple, 139, 148
Araceae, 67
Arachnula, 191
Araiospora, 79
Arcella, 205
Arcellidae, 205
ArcelHna, 205
Archaelagena, 186
Archaias, 184, 185
Archangiacea, 28
Archangiaceae, 28
Archangium, 28
Archegregarina, 215
Archephyta, 17
Archezoa (of Haeckel), 17
Archezoa (of Perty}, 223
Archi-Monothalamia, 183
Archimycetae, 110
Archimycetes, 110, 111
Archiplastidea, 18, 29
Archiplastideae, 30
Arcyria, 176
Arcyriacea, 174, 176
Arcyriaceae, 176
Arcyriidae, 176
Arthropods, 211, 212, 222
Arthrospira, 35
Asclepiadaceae, 161
Ascobolacea, 135
Ascochyta, 141
Ascocorticium, 137, 145
Ascocyclus, 88
Ascoidea, 130
Ascoidea rubescens, 127
Ascoideaceae, 130
Ascomvcetae, 125
Ascomycetes, 120, 125, 140, 142, 145
Ascomyceten, 125
Ascosporeae, 125
Askcleta, 193
Aspergillus, 130, 131
Aspergilliales, 130
Aspidisca, 232
Asplrotricha, 229
Aspirotrichaceae, 229
Astasia, 96, 107
Astasiaceae, 107
Astasiaca, 96, 105, 107
Astasiidae, 107
Astasiina, 107
Asterigerina, 187
Asterigerinida, 187
Asterigerinidae, 187
Asterocyclina, 188
Asterocystis, 43
Asterophlyctis, 1 1 7
Astoma, 94, 96, 105
Astoinaticae, 74
Astomina, 230
Astracanthida, 199
Astracanthidae, 199
Astrodisculus, 193
Astrolophida, 197
Astrolophus, 197
Astrorhiza, 183
Astrorhizida, 183
Astrorhizidaceae, 183
Astrorhizidae, 183
Astrorhizidea, 183
Astrorhizina, 183
Ataxophragmidae, 186
Ataxophragmidea, 186
Ataxophragmium, 186
Athene noctua, 162
Aulacantha, 199
Aulacanthida, 199
Aulacanthidae, 199
Aulactinium, 199
Aulosphaera, 199
Aulosphaerida, 199
Auricularia, l46
Auricularia Auricula, 146
Auriculariacea, 146, 148
Auriculariaceae, 146
Auriculariales, 146
Auriculariineae, 145, 146
Auricularineae, 146
Autobasidiomycetes, 146
Aves, 6
Axonoblasteae, 51
Azoosporidae, 191
Azoosporidca, 191
Azoosporidia, 190
Azotobacter, 14
Azotobacter Chroococcum, 23
Azotobacteriacca, 19, 23
Azotobacteriaceae, 23
Babesia, 214
Babesia bigemina, 206, 212, 214
Babesiida, 211, 214
Babesiidae, 214
Bacillacea, 19, 21
Bacillacei, 21
Bacillaria, 69, 75
Bacillariacea, 1 1, 55, 65, 69, 72
Eacillariaceae, 71
Bacillariales, 53, 71
Bacillarieae, 71
Bacillarioideae, 71
Bacillariophyceae, 71
Bacillariophyta, 71
Bacillus, 21
Bacillus alvei, 21
Bacillus Amylobacter, 21
Bacillus anthracis, 21
Bacillus, colon, 22
Bacillus, gas, 22
Bacillus Radicicola, 23
Bacillus, Shiga, 22
Bacillus subtilis, 18, 21
Bacteria, 2, 3, 4, 6, 7, 12, 13, 14, 17, 18,
30, 38, 118, 119, 189, 222, 224, 231
274]
The Classification of Lower Organisms
Bacteriaceae, 21
Bacteriophyta, 17
Bacteroides, 22
Badhamia, 177
Balantidium, 230
Balantidium coli, 230
Bangia, 43
Bangia fuscopurpurea, 43
Bangiacea, 41
Bangiaceae, 41, 43
Bangialea, 40, 41, 52
Bangiales, 41
Bangieae, 41
Bangiineae, 41
Bangioideae, 41
Barbulanympha, 169
Barley, 6
Barrouxia, 210
Bartonella bacilliformis, 21, 214
Bartonellaceae, 20
Bartramia, 219
Bartramiidae, 218
Basidiobolacea, 125
Basidiobolaceae, 125
Basidiobolus, 119, 121
Basidiobolus ranarum, 125
Basidiomycetae, 142
Basidiomyceten, 142
Basidiomycetes, 121, 127, 128, 141, 142,
145
Basidiosporeae, 142
Bathysiphon, 183
Batrachospermaceae, 47
Batrachospermum, 47
Bdellospora, 124
Beetles, 177, 215, 217
Beggiatoa, 24, 30, 31, 32, 35
Beggiatoacea, 34, 35
Beggiatoaceae, 35
Bicoecaceae, 67
Bicoecidea, 67
Bicoekida, 67
Bicosoeca, 67
Biddulphia, 74
Biddulphiaceae, 74
Biddulphica, 74
Biddulphicac, 74
Biflagcllatae, 76
Bikoecidae, 67
Bikoccina, 67
Birds, 6, 210. 212, 213
Bitunicatae, 129
Blakeslcca, 124
Blastocaulis, 26, 27
Blastocladia, 112, 113
Blastocladiarca, 110, 112
Blastocladiaceae, 112
Blastocladialcs, 1 1 1
Blastocladiclla, 112, 113
Blastocladiclla cystogena, 115
Blastocladiineae, 111
Blastodcrma, 130
Blastodinida, 100, 102
Blastodinidae, 102
Blastodinides, 102
Blastodinium, 102
Blastosporaceae, 44
Blepharisma, 230
Blue grass, 148
Blue-green algae, see Algae, Blue-green
Bodo, 159, 160, 199, 209, 212
Bodo edax, 161
Bodo Lacertae, 159
Bodonaceae, 159
Bodonidac, 159
Bodonidca, 158
Bodonina, 159
Boletus, 151
Bolivina, 188
Borelis, 185
Borrelia, 29
Borrelia recurrentis, 28, 29
Borrelia Vincenti, 29
Botrida, 198
Botrydiaceae, 67
Botrydiales, 63
Botrydiopsis, 66
Botrydium, 65, 67
Botryococcacea, 65, 66
Botryococcaceae, 66
Botryococcus, 66
Botryoglossum, 52
Botryoidca, 198
Botrytis, 140, 142
Bovista, 155
Braadrudosphaeridae, 60
Brachycystida, 217
Brefcldia, 175
Brefeldiaceae. 175
Brefeldiidae, 175
Brehmiella, 59
Brehmiella chrysohydra, 54
Brown algae, see Algae, Brown
Brucella, 22
Bulgariacea, 135
Bulimina, 188
Buliminida, 188
Buliniinina, 188
Bumilleria, 66, 73
Bursaria, 230
Bursariidae, 230
Bursarina, 230
Cabbage, 1 78
Calcaroae, 1 71
Calcarina, 187
Calcarinidae, 187
Calciconus, 60
Galciconus vitrcus, 56
Calrisolcnia. 60
Calcisolonidae, 60
Callocolax, 50
Callophyllis, 50
Index
[275
Calonectria, 142
Calonema, 177
Calonemeae, 171
Calonympha, 168
Calonymphida, 166, 167, 168
Calonymphidae, 168
Calothrix, 36
Calvatia, 155
Calyptosphaera, 60
Calyptosphaera insignis, 56
Camerina, 188
Camerinidae, 188
Camptonema, 193
Camptonematidae, 193
Campuscus, 191
Candida, 142
Cannobotryida, 198
Cannopilus, 63
Cannosphaerida, 199
Cannosphaeridae, 199
Cantharellales, 150
Carageen, 49
Carboxidomonas, 24
Carchesium, 233
Carcheslum polypinum, 225
Carpomitra, 88
Carpomycetae, 119
Carpophyceae, 40
Carposporeen, 128
Caryococcus, 21
Caryospora, 210
Caryotropha, 211
Cassidulina, 188
Cassidulinida, 188
Cassidulinidae, 188
Castanellida, 200
Castanellidae, 200
Castanidium, 200
Cat, 6, 210
Catenariopsis, 69
Catenochytridium, 118
Cattle, 206, 214
Caulleryella, 215
CauUeryellidae, 215
Caulobacter, 26, 27
Caulobacter vibrioides, 26
Caulobacteriacea, 27
Caulobacteriaceae, 27
Caulobacterialea, 18, 25, 26
Caulobacteriales, 25
Cayeuxina, 186
Cellulomonas, 22
Cenolarcus, 195
Centipedes, 207, 210, 211
Centricae, 73, 74
Cepedia, 229
Cephalina, 217
Cephalopodes, 182
Cephalothamnium, 59
Cephalothamnium Cyclopum, 54
Cephalotrichinae, 18
Ceramiales, 51
Ceramiea, 51, 52
Ceramieae, 51
Ceratiidae, 103
Ceratiomyxa, 177, 221
Ceratiomyxa fruticulosa, 177, 178
Ceratiomyxacea, 177
Ceratiomyxaceae, 177
Ceratium (dinoflagellate), 103
Ceratium ( myxomycete ) , 177
Ceratium Hirundinella, 103
Ceratomyxa, 221
Ceratomyxidae, 221
Ceratophyllus fasciatus, 160
Ceratospora, 216
Cercobodo, 159
Cercobodonidae, 159
Cercomonadida, 159
Cercomonadidae, 159
Cercomonadinea, 158
Cercomonas, 159, 161
Cercomonas Davainei, 165
Cercomonas Hominis, 165
Cercomonas longicauda, 160
Cercomonas obliqua, 165
Cercospora, 138, 139, 142
Chaenia, 229
Chaetangieae, 47
Chaetoceraceae, 74
Chaetoceros, 74
Chaetocladiaceae, 124
Chaetocladium, 123, 124
Chaetoproteida, 159, 163
Chaetoproteidae, 163
Chaetoproteus, 158, 160, 163, 202
Chaidae, 201
Chaidea, 201
Chalarothoraca, 190, 193
Chalarothoracida, 190
Challengerida, 200
Challengeridae, 200
Challengeron, 200
Chamaesiphon, 35, 36
Chamaesiphon incrustans, 32
Chamaesiphonacea, 34, 35
Chamaesiphonaceae, 33, 35
Champia, 51
Champiea, 51
Champieae, 51
Chantransia, 47
Chantransiaceae, 47
Chaos Protheus, 200, 201, 202
Chaosidae, 201
Chapmania, 187
Chapmaniida, 187
Chapmaniidae, 187
Characiopsis, 66
Characiopsis gibba, 64
Chestnut, 139
Chiastolida, 197
Chiastolus, 197
Chicken, 210
Chilodon, 230
276
The Classification of Lower Organisms
Chilodon uncinatus, 225
Chilodontida, 230
Chilomastigidae, 165
Chilomastix, 165
Chilomastix davainei, 165
Chilomastix Hominis, 165, 237
Chilomastix Mesnili, 165
Chilomonadaceac, 98
Chilomonas, 94, 109
Chilomonas Paramaecium, 97
Chilostomella, 187
Chilostomellida, 187
Chilostomellidae, 187
Chlamydodon, 230
Chlamydodonta, 230
Chlamydodontida, 230
Chlamydodontidae, 230
Chlamydomonas, 61, 111
Chlamydomyxa, 191
Chlamydomyxidea, 190
Chlamydophora, 190, 193
Chlamydophorida, 190
Chlamydothrix ochracea, 32, 36
Chlamydotrichacea, 34
Chlamydotrichaceae, 36
Chlamydozoaceae, 20
Chloramoeba, 66
Chloramoeba heteromorpha, 64
Chloramoebacca, 65, 66
Chloramoebaceae, 66
Chloramoebidac, 66
Chlorarachnidae, 66
Chlorobacteriaceae, 31
Chlorobacterium, 33
Chlorobiacea, 31, 237
Chlorobium, 31
Chlorobotrydiaceae, 66
Chlorochromonas, 66
Chlorochytridion, 1 1 1
Chloromonadaceae, 109
Chloromonadales, 63, 105
Chloromonadida, 105
Chloromonadidae, 109
Chloromonadina, 63, 96, 105
Chloromonadinae, 94, 105
Chloromonadineae, 105
Chloromonads, 94
Chloromyxea, 221
Chloromyxees, 221
Chloromyxida, 221
(^hloromyxidac, 221
Chloromyxum, 221
Chlorosaccacca, 65
Chlorosaccaceac, 65
Chlorosaccus, 55, 65, 66
Chlorosaccus fluidus, 64
Chlorotheciacea, 65, 66
Chlorothcciaceae, 66
Choancphoraccac, 124
Choano-Flagellata, 67
Choanocystidac, 194
Choanocystis, 194, 216
Choanoflagellata, 57, 61, 67, 68
Choanoflagcllates, 57, 158
Choanosporidae, 216
Chondria, 52
Chondrieae, 51
Chondrioderma, 177
Chondrococcus, 28
Chondromyces, 28
Chondromyces aurantiacus, 26
Chondromyces crocatus, 26
Chondrus, 51
Chondrus crispus, 49
Chonotricha, 230, 231, 237
Chonotrichida, 230
Chordariacea, 88
Chordariaceae, 87
Chordariales, 87
Chordarieae, 87
Chromatiacea, 31, 237
Chromatiaceae, 31
Chromatium, 31
Chromobacterium, 22
Chromomonas, 98
Chromulina, 61, 62
Chromulina Pascheri, 56
Chromulinaceae, 62
Chromulinales, 61
Chromulinidae, 62
Chroococcacea, 33
Chroococcaceae, 33
Chroococcales, 33
Chroococcus, 32, 33
Chrysamoeba, 63
Chrysamoebida, 62, 63
Chrysamoebidae, 63
Chrysapsis, 62
Chrysarachniaceae, 63
Chrysarachnion, 63
Chrysidella, 98
Chrysocapsa, 59
Chrysocapsa paludosa, 54
Chrysocapsacea, 58, 59
Chrysocapsaceae, 59
Chrysocapsales, 61
Chrysocapsidae, 59
Chrysocapsina, 61
Chrysocapsinae, 61
Chrysocapsineae, 55, 61
Chrysochromulina, 58
Chrysococcus, 62
Chrysocrinus, 63
Chrysodcndron, 59
Chrysomonadaceae, 59
Chrysomonadales, 61
Chrysomonadida, 61
Chrysoinonadidao, 62
Chrysomonadina, 59, 61, 62
Chrysomonadinae, 61
Chrysomonadinca, 57
Chrysoinonadincac, 55, 57, 61
Chrysonionads, 53, 83
Chrysomonas, 62
Index
[277
Chrysophaeum, 109
Chrysophyceae, 53, 55, 95
Chrysophycophyta, 53
Chrysophyta, 53
Chrysopyxis, 60
Chrysosphaera, 62
Chrysosphaeracea, 61, 62
Chrysosphaeraceae, 62
Chrysosphaerales, 61
Chrysosphaerella, 62
Chrysosphaerineae, 55, 61
Chrysospora, 62
Chrysothylakion, 63
Chrysotrichaceae, 60
Chrysotrichales, 61
Chrysotrichineae, 55, 61
Chytridiacea, 117, 118
Chytridiaceae, 110, 118
Chytridiales, 113
Chytridieae, 110
Chytridieen, 110, 118
Chytridiineae, 110
Chytridinae, 110
Chytridinea, 111, 113, 116
Chytridineae, 110, 113
Chytridium, 69, 110, 113, 118
Chytridium Olla, 110
Chytrids, 76, 110, 111, 119, 121, 125, 130,
178
Chytriodinium, 102
Cienkowskiaccae, 177
Ciliata, 223, 228, 233
Ciliatea, 228
Cilio-flasrellata, 94
Cilioflagellata, 96, 102
Giliophora, 39, 223, 237
Ciliophryidae, 191
Ciliophrys, 193
Circoporida, 200
Circoporidae, 200
Circoporus, 200
Cladochytriacea, 115, 117
Cladochytriaceae, 117
Cladochytrium, 110, 117
Cladococcida, 195
Cladococcus, 195
Cladopyxida, 103
Cladosporium, 142
Cladothrix dichotoma, 33
Clastoderma, 175
Clathracea, 155
Clathraceae, 155
Clathrochloris, 31
Clathrulina, 194
Clathrulinida, 191, 194
Clathrulinidae, 194
Claudea, 51
Clavaria, 151
Clavariacea, 151
Clavariaceae, 151
Clavariei, 151
Clavati, 150
Claviceps purpurea, 139
Clonothrix fusca, 32, 36
Closterium, 125
Clostridium, 21
Clostridium botulinum, 21
Clostridium butyricum, 21
Clostridium Pastorianum, 21
Clostridium septicum, 21
Clostridium tetani, 21
Cnemidospora, 21 7
Cnidosporidea, 219
Cnidosporidia, 219, 220
Coccaceac, 20
Coccidia, 207, 210
Coccidians, 260, 209, 210, 212, 215
Coccididae, 210
Coccidiidea, 210
Coccidiomorpha, 207, 210
Coccidium, 210
Coccidium Schubergi, 207
Coccogonales, 33
Coccogonea, 31, 32, 33
Coccogoneae, 33
Coccolithaceae, 60
Coccolithidae, 60
Cocclithina, 60
Coccolithophora, 60
Coccolithophoridae, 55, 60
Coccolithus, 60
Coccomyces, 134
Coccomyxa, 221
Coccomyxida, 221
Coccomyxidae, 221
Cocconeidaceae, 76
Cocconeis, 72, 73, 76
Cocconemaceae, 75
Cocconemidae, 222
Coccosphaera, 60
Coccospora Slavinae, 222
Coccosporida, 222
Coccosporidae, 222
Coccus, 20
Cochliodinium, 101
Cochliopodiidae, 202
Cochliopodium, 202
Cochlonema, 124
Cockroach, 169,217,219
Codonoecina, 67
Codonosiga, 67
Codonosigidae, 67
Codosiga, 67
Coeloblastca, 46
Coeloblasteae, 51
Coclodendrida, 200
Coelodendrum, 199, 200
Coelomonadina, 105, 109
Coelosphaerium, 33
Coelosporidiidae, 218
Coelosporidium, 219
Coenenia, 203
Coffee, 148
Colaciacea, 105
278]
The Classification of Lower Organisms
Colaciaceae, 105
Colaciidae, 105
Colacium, 105
Colacium Arbuscula, 106
Coleosporiacea, 148
Coleosporiaceae, 148
Coleosporium, 143
Coleosporium Vernoniae, 143
Colepina, 229
Coleps, 229
Colletotrichum, 139, 140
Collida, 195
Collodaria, 194
Colloderma, 177
Collodermataceae, 177
CoUosphaera, 195
Collosphaera Huxleyi, 196
Collosphaerida, 195
Collozoida, 195
Colpidium campylum, 227
Colpoda, 229
Colpodaea, 229
Colpodella, 189
Colpodidae, 229
Columniferae, 171
Comatricha, 175
Completoria, 125
Compsopogon, 44
Compsopogonacea, 41, 44
Compsopogonaceae, 44
Concharida, 200
Concharidae, 200
Conchulina, 205
Conferva, 66
Confervaceae, 66
Confervales, 63
Confervoidea, 63
Conger niger, 161
Conidiobolus, 125
Coniferinae, 9
Conifers, 148
Coniomycetes, 140
Conjugatae, 117
Conradiella, 62
Coprinus, 143, 152
Coprinus atramentarias, 153
Copromonas subtilis, 108
Cora, 151
Corallinaceae, 50
Corallinea, 50
Corallineae, 50
Cordyceps, 139
Coreocolax, 50
Corethron, 74
Cormobionta, 6
Cornuspira, 185
Coronympha, 168
Corticiiun, 151
Corynebacteriacea, 19, 20
Coryncbacteriaceae, 20
Corynebacteriidae, 20
Coryncbactcrium, 20, 21
Corynebacterium diphtheriae, 20
Coryneum, 141
Coryneum Beijerinckii, 141
Coscinodiscaceae, 74
Coscinodiscea, 74
Coscinodiscus, 74
Costia, 165
Costiidae, 165
Cothurnia, 233
Councilmania, 203
Crab, 218
Craigia, 163
Craspedomonadaceae, 67
Craspedomonadina, 67
Craspedotella, 102
Craterellus, 151
Craterium, 177
Crenothrix polyspora, 32, 36
Crenotrichacca, 35, 36
Crenotrichaceae, 36
Cribraria, 175
Cribrariacea, 173, 175
Cribrariaceae, 171, 175
Cribrariales, 171, 173
Cribrariidae, 175
Cribrospira, 186
Cristellaria, 187
Cristispira, 29
Cristispira Veneris, 26
Crithidia, 162
Cromodromys, 199
Cronartiacea, 148
Cronartiaceae, 148
Cronartium, 148
Cronartium ribicola, 148
Cryptobia, 160, 161,209,212
Cryptobiidae, 161
Cryptocalpis, 198
Cryptocapsales, 97
Cryptocapsineae, 95
Cryptocercus, 166, 169, 170
Cryptochrysis, 98
Cryptococcacea, 97, 98
Cryptococcaceae, 98
Cryptococcalcs, 96, 97
Cryptococcineae, 95
Cryptococcus, 98, 130
Cryptocystes, 219, 222
Cryptomonadaceae, 98
Cryptomonadalca, 96
Cryptomonadalcs, 96
Cryptomonadida, 97
Cryptomonadidae, 98
Cryptomonadina, 96, 97, 98
Cryptonionadinae, 96
Cryptonionadincae, 95, 96
Crvptomonads, 94, 194
Cryptomonas, 97,98, 199
Cryptoncnicac, 50
Cryptonemiales, 50
Cryptoncniinae, 50
Cryptophyceae, 94, 96
Index
[279
Cryptospermea, 46, 47
Cryptospermeae, 47
Ctenomyces, 131
Ctenostomata, 230, 231
Ctenostomida, 231
Ctenostomina, 231
Cumagloia, 47
Cuneolina, 186
Cunninghamella, 124
Cup fungi, 134
Cupulata, 129, 134, 137, 145
Cupulati, 134
Currants, 148
Cutleria, 88
Cutlcriacea, 88
Cutlerialea, 85, 88
Cutleriales, 88
Cyanomonas, 98
Cyanophyceae, 29
Cyanophyta, 17, 30
Cyathoxnonas, 97, 98
Cyathus, 155
Cyclammina, 186
Gyclidina, 230
Cyclidium, 230
Cycloclypeidae, 188
Cycloclypeina, 188
Cycloclypeus Carpenter!, 188
Cyclodinidae, 229
Cyclonexis, 59
Cyclonympha, 171
Cyclonymphidae, 169
Cycloposthium, 231
Gyclosiphon, 188
Cyclosporales, 91
Cyclosporeae, 82, 91
Cyclotella, 72, 73, 74
Cylindrospermum, 35
Cylindrospermum majus, 32
Cylindrosporium Pruni, 134
Cymbalopora, 180, 182, 187
Cymbella, 72, 73, 75
Cymbellea, 75
Cymbelleae, 75
Cyphoderia, 191
Cyrtellaria, 198
Cyrtida, 198
Cyrtoidea, 198
Cyrtophora, 62, 63
Cystidium, 198
Cystobasidium, 147
Cystobasidium sebaceum, 145
Cystochytrium, 69
Cystoflagellata, 94, 96, 99
Cytophaga Hutchinsonii, 26, 28
Cytophagacea, 28
Cytophagaceae, 28
Cytosporidia, 207
Cyttariacea, 135
Dacryomitra, 150
Dacryomyces, 150
Dacryomycetacea, 150
Dacryomycetaceae, 150
Dacryomycetalea, 146, 150
Dacryomycetales, 150
Dacryomycetineae, 150
Dactylophorida, 218
Dactylophoridae, 218
Dactylophorus, 218
Dactylosphaerium, 202
Daedalea, 151
Daldinia, 139
Dallingeria, 58
Dasyea, 51
Daucina, 188
Deer, 214
Delacroixia, 125
Delesseria, 51
Delesseria sinuosa, 49
Delesseriea, 51
Dematiaceae, 142
Dematiea, 142
Dematieae, 142
Dematiei, 141
Dendrocometes, 236
Dendrocometida, 236
Dendrocometidae, 236
Dendrocometina, 236
Dendromonadina, 59
Dendromonas, 59
Dendromonas virgaria, 54
Dendrosoma, 235
Dendrosomatidae, 235
Dendrosomida, 235
Dendrosomidae, 235
Dendrosomina, 235
Dentilina, 184
Derepyxis, 60
Dermateacea, 135
Dermatocarpa, 146, 152
Dermatocarpi, 152, 171
Dermocarpa, 36
Dermocarpa protea, 32
Dermocentor, 20
Desmarestia, 88, 89
Desmarestiacea, 88
Desmarestales, 87
Desmobacteriales, 33
Desmocapsa, 99
Desmocapsales, 98, 99
Desmocapsineae, 95, 99
Desmokontae, 94, 98, 99
Desmomastix, 99
Desmomonadales, 98, 99
Desmomonadineae, 95, 99
Desmothoraca, 190, 194
Desmothoracida, 190
Desmotrichum, 88
Deuteromycetes, 140
Deutschlandiaceae, 60
Devescovina, 167
280
The Classification of Lower Organisms
Devescovinida, 167
Devescovinidae, 167
Devescovininae, 167
Diachea, 175
Dianema, 176
Dianemaceae, 176
Diaporthe, 139
Diatoma, 75
Diatomaceae, 69, 75
Diatomea, 53, 69, 71, 74
Diatomeae, 53, 69, 71, 74
Diatoms, 53, 71,83, 117, 118
Diatrype, 139
Dictydiaethaliaceae, 175
Dictydiaethaliidae, 1 75
Dictydiaethalium, 175
Dictydium, 175
Dictyocha, 63
Dictyocha Fibula, 56
Dictyochaceae, 62
Dictyochidae, 62
Dictyoconoides, 187
Dictyoconus, 186, 198
Dictyophora, 155
Dictyosiphonales, 89, 91
Dictyosteliaceae, 203
Dictyosteliaceen, 203
Dictyostelidae, 203
Dictyostelium, 203
Dictyostclium discoideum, 204
Dictyostelium mucoroides, 204
Dictyota, 87
Dictyotacea, 87
Dictyotaceae, 86, 87
Dictyotales, 82, 86
Dictyotea, 85, 86
Dictyotcae, 82, 86
Dictyuchus, 78, 79
Didesmis, 231
Didiniidae, 229
Didinium, 229, 235'
Didinium nasutum, 225
Didymiacea, 175, 177
Didymiaceae, 177
Didymidae, 177
Didymiidae, 177
Didymium, 177
Didymohelix ferruglnea, 27
Didymophyes, 218
Didymophyida, 218
Didymophyidac, 218
Difflugia, 201, 205
DIfflugiida, 205
Difflugiidae, 205
Dilcptus, 230
Dimastigamocba, 159
Dimorpha, 193
Dimychota, 17
Dinamocba (dinoflagellate), 101
Dinamocba (amoeba), 16, 202
Dinamocbidina, 100, 101, 237
Dinamocbidium varians, 101, 104
Dinastridium, 100
Dinenympha, 166
Dinenymphida, 165, 166
Dinenymphidae, 166
Dinifera, 102
Diniferidea, 103
Dinobryaceae, 60
Dinobryina, 58, 60
Dinobryon, 58, 60
Dinocapsaceae, 100
Dinocapsales, 99, 100
Dinocapsina, 99
Dinocapsineae, 95, 99
Dinococcales, 99, 100
Dinococcina, 99
Dinococcineae, 96, 100
Dinoclonium, 100
Dinocloniaceae, 100
Dinoflagellata, 94, 102
Dinoflagellatae, 94, 95
Dinoflagellates, 94, 199
Dinoflagellida, 103
Dinophyceae, 94, 103
Dinophysida, 103
Dinophysis, 103
Dinothrix, 100
Dinotrichales, 99, 100
Dinotrichineae, 96, 99
Dioxys, 66
Dioxys Incus, 64
Diplococcus, 20
Diplococcus pneumoniae, 20
Diploconida, 197
Diploconus, 197
Diplocystida, 216
Diplocystidae, 216
Diplocystis, 216
Diplodia, 141
Diplodinium, 224, 231
Diplomita, 60
Diplophlyctis, 117
Diplophysalis, 191
Diplophysalis stagnalis, 192
Dipodascus, 130
Dipodascus albidus, 132
Discellacea, 141
Discellaceae, 141
Dischizae, 21 5
Discida, 195
Disciformia, 73
Discoasteridac, 60
Dificoidca, 195
Discolichenes, 134
Discomycetes, 133, 134
Discophrya, 235
Discophryida, 235
Discophryidae, 235
Discorbis, 180, 182
Discorbis mcditerrancnsis, 182
Discorbis orbicularis, 182
Discosphacra, 60
Disporees, 209
Index
[281
Distephanus, 63
Distephanus Speculum, 56
Distigma, 107
Distomata, 163
Distomataceae, 166
Distomatinales, 163
Distomatineae, 163
Ditripodiidae, 62
Doassansia, 149
Dobellia binucleata, 210
Dobeliida, 210
Dobelliidae, 210
Do?, 210
Dolichocystida, 209, 214
Doliocystida, 216
Doliocystidae, 216
Doliocystis, 216
Dorataspida, 197
Dorataspis, 197
Dorataspis costata, 196
Dothideaceae, 137
Dothideales, 137, 138, 139, 140, 141
Drepanidium, 211
Duboscqia, 222
Dudresnaya purpurifera, 49
Dumontieae, 50
Earth star, 155
Earthworm, 215, 216
Eberthella, 22
Eberthella typhi, 22
Ebriaceae, 55, 62
Ebriidae, 62
Ebriopsidae, 62
Echinocystida, 189
Echinoderms, 216
Echinosteliaceae, 175
Echinostelium, 175
Ectocarpales, 86
Ectocarpea, 86
Ectocarpeae, 86
Ectocarpineae, 86
Ectocarpus, 70, 83, 86, 87
Ectocarpus Mitchelliae, 204
Ectocarpus siliculosus, 83
Ectosporeae, 177
Ectrogella, 81
Ectrogellacea, 81
Ectrogellaceae, 81
Eel, 161
Egregia Menziesii, 90, 91
Eimeria, 210
Eimerida, 210
Eimeridae, 210
Eimeridea, 210
Eimcriidea, 210
Eimeriinea, 210
Eimerioidae, 210
Elaeorhanis, 193
Elaphomyces, 131
Ellipsoidina, 188
Elphidium, 186, 187
Elphidium crispum, 181
Elvella, 135
Empusa, 125
Enchelia, 229
Enchelina, 229
Enchelis, 229
Enchelyidae, 229
Endamoeba, 202, 203
Endamoeba disparita, 202
Endamoeba histolytica, 202
Endamoebida, 201, 202
Endamoebidae, 202
Endocochlus, 124
Endogonacea, 123, 124
Endogonaceae, 124
Endogone, 123, 124
Endogonei, 124
Endoiimax, 203
Endomyces, 130
Endomycetacea, 130
Endomycetaceae, 130
Endomycetalea, 129
Endomycetalcs, 129
Endosporea, 171
Endosporeae, 171
Endosporinei, 171
Endothia parasitica, 139
Endothyra, 186
Endothyridae, 186
Endothyrina, 186
Enerthenema, 175
Enerthenemaceae, 175
Enerthenemea, 174, 175
Entamoeba, 202, 203
Entamoeba coli, 202
Entamoeba dystenteriae, 202
Entamoeba gingivalis, 202
Enteridiea, 171
Enteridieae, 171
Enterobacteriaceae, 21
Entodiniomorpha, 230, 231
Entodiniomorphina, 231
Entodinium, 231
Entomophthora, 125
Entomophthoracea, 124
Entomophthoraceae, 124
Entomophthorales, 124
Entomophthorinea, 121, 124
Entomophthorineae, 124
Entophlycis, 113, 117
Entophysalidales, 33
Entosiphon sulcatum, 108
Eocronartium, 143, 147
Eocronartium muscicola, 145
Eouvigerina, 188
Ephelota, 236
Ephelotida, 236
Ephelotidae, 236
Ephelotina, 236
Ephemera vulgata, 222
Epiblasteae, 50
282]
The Classification of Lower Organisms
Epichrysis, 56, 62
Epiclintes, 233
Epidinium, 231
Epipyxis, 60
Epipyxis utriculus, 54
Epistylis, 233
Eremascus, 130
Eremascus albidus, 127
Eremospermeae, 77
Erica, 9
Ericae, 9
Erysiphe, 127, 132, 133
Erysiphe graminis, 132
Erysiphea, 133
Erysipheae, 133
Erythrocladia, 44
Erythropsis, 101
Erythrotrichia, 44
Erythrotrichia carnea, 44
Erythrotrichiaceae, 44
Erwina, 22
Erwinia amylovora, 22
Escherichia coli, 14, 15, 22
Ethmosphaerida, 195
Euactinomyxidae, 222
Euasci, 130
Eubacteria, 18, 25
Eubacteriales, 18
Eubasidii, 145
Euchrysomonadina, 61
Euchrysomonadinae, 61
Eucomonympha, 169
Eucyrtidina, 198
Eucyrtidium, 198
Eucyrtidium carinatum, 196
Eudesme, 88
Euflorideae, 44
Euglena, 38, 94, 107, 116, 117, 125
Euglena acus, 106
Euglena Spirogyra, 106, 107
Euglena viridis, 106
Euglenaceae, 105
Englenales, 105
Euglenamorpha, 105
Euglenida, 105
Euglenids, 94, 106
Euglenina, 105
Euglcninae, 94, 105
Euglenineae, 96, 105
Euglenocapsineae, 96
Euglcnoidina, 96, 105
Euglenophycophyta, 94
Euglenophyta, 94
Euglypha, 191
Euglyphida, 191
Euglyphidae, 191
Eugregarinaria, 217
Eugregarinida, 217
Eumycetes, 119
Eumycctozoina, 171
Eumycophyta, 119
Eunotia, 75
Eunotiaceae, 75
Eunotiea, 75
Eunotieae, 75
Euphorbiaceae, 161
Euplotes, 227, 232, 233, 234
Euplotes Patella, 225, 226, 232
Eupodiscales, 73
Eurotium, 131
Eurychasma, 81
Eur)'chasmidium, 81
Eurysporea, 221
Eutreptia, 105
Excipula, 141
Excipulaceae, 141
Exidia, 143
Exoascalea, 129, 137
Exoascales, 137
Exoascus, 137
Exobasidiacea, 151
Exobasidiaceae, 151
Exobasidiales, 1 50
Exobasidiineae, 150
Exobasidium, 151
Exosporea, 171, 177
Exosporeae, 177
Exosporinei, 177
Exuviaella, 99
Fasciolites, 185
Fauchea, 51
Faucheocolax, 51
Felis Catus, 6
Ferns, 125, 148
Filicineae, 1
Fisherinidae, 185
Fishes, 165, 210, 211, 219. 220, 222
Flabellina, 184, 187
Flagellata, 6, 55, 94, 96, 105
Flagellatae, 94
Flagellates, 10, 53, 55, 76, 94, 118
Flagellato-Eustomata, 105
Flagellato-Pantostomata, 158
Flatworms, 216
Flavobacterium, 22
Flea, 160
Flexostylida, 185
Floridea, 47, 50,51
Florideae, 6, 40, 44, 51
Floridees, 40, 51
Floridineae, 44
Flowers of tan, 177
Fly, 213
Foaina, 167
Folliculina, 230
Fomcs, 151
Foraminifera, 179, 182, 183, 185
Foraminiferes, 179, 182
Foraminiferida, 179
Forficule, 217
Fragilaria, 75
Fragilariaceae, 75
Index
[283
Fragilariea, 75
Fragilarieae, 75
Frogs, 125, 210, 211
Frondicularia, 187
Fucaceae, 91
Fucales, 91
Fucea, 91
Fuceae, 91
Fucineae, 91
Fucacees, 82
Fucoidea, 83, 86, 91
Fucoideae, 53, 82
Fucus, 53, 91, 93
Fucus vesciculosus, 93
Fuligo septica, 177
Fungi, 39, 69, 76, 110, 119, 146, 150, 172
Fungi, bird's-nest, 155
Fungi imperfecti, 140
Fungilli, 39, 206
Furcellariea, 46, 50
Furcellarieae, 50
Fusarium, 142
Fusiformis, 29
Fusobacterium, 29
Fusulina, 188
Fusulinida, 188
Fusulinidae, 188
Galaxaura, 47
Galera tenera, 153
Gallionella, 27
Gallowaya, 148
Gammarus, 233
Gamocystis, 217
Ganymedes, 216
Ganymedida, 216
Ganymedidae, 216
Gasteromycetes, 152
Gastrobionta, 6
Gastrocarpeae, 50
Geaster, 155
Gelidiaceae, 49
Gelidialea, 46, 49, 50
Gelidiales, 49
Gelidieae, 49
Gelidium, 50, 51
Geophonus, 186, 187
Geoglossacea, 135
Giardia, 163, 166
Giardia enterica, 164, 166
Giardia Lamblia, 166
Gibberella, 142
Gigantomonas, 167
Gigartina mammilosa, 49
Gigartinales, 47
Gigartineae, 47
Gigartininae, 47
Glandulina, 187
Glaucocystis, 33
Glaucoma, 229
Glaucoma pyriformis, 227
Glenodinium, 94, 103
Globigerina, 184, 188
Globigerinida, 188
Globigerinidea, 183, 187
Globorotalia, 187
Globorotaliidae, 187
Gloeocapsa, 33
Gloeochaete, 33
Gloeochrysis, 62
Gloeodiniaceae, 100
Gloeodinium, 100
Gloeosporium, 139, 140, 141
Gloeotrichia, 36
Gloiophycea, 31, 32, 33
Gloiophyceae, 29, 33
Glomerella, 126, 127, 139, 140
Glomerella cingulata, 139
Glugea, 222
Glugeida, 222
Glugeidae, 222
Glugeidea, 222
Glugeidees, 222
Goat, 210
Gomphonema, 72, 75
Gomphonemaceae, 75
Gomphonemea, 75
Gomphonemeae, 75
Gomphosphaeria, 33
Gonapodiaceae, 112
Gonapodiineae, 112
Gonapodya, 112
Goniaulax, 103
Gonimophyllum, 52
Goniodoma, 103
Goniostomum, 109
Goniotrichaceae, 43
Goniotrichopsis, 43
Goniotrichum, 43
Gonococcus, 20
Gonospora, 216
Gooseberries, 146
Goussia, 209, 210
Goussia Schubergi, 207, 208, 237
Gracilaria, 49
Grains, 149
Granuloreticulosa, 179
Graphidiacea, 134
Graphidiaceae, 134
Graphidiales, 133
Grasses, 149
Green algae, see Algae, Green
Gregarina, 206, 217
Gregarina conica, 217
Gregarina cuneata, 217
Gregarina ovata, 217
Gregarinae, 206, 216
Gregarinarien, 217
Gregarines 206, 209, 215, 219
Gregarinida, 207, 217
Gregarinidae, 217
Gregarinidia, 207
Gregarininea, 217
284]
The Classification of Lower Organisms
Gregarinoidae, 217
Gregarinoidea, 217
Gregarinomorpha, 207
Gromia, 179, 191
Gromida, 191
Guepinia, 150
Guepinia apathularia, 145
Gurleya, 222
Guttulina, 203
Guttulina sessilis, 204
GuttuHnacea, 201, 203
Guttulinaceae, 203
GuttuHneae, 203
Guttulineen, 203
Guttulinidae, 203
Guttulinopsis, 203
Gymnamoebae, 201
Gymnamoebida, 201
Gymnascales, 130
Gymnoascaceae, 130
Gymnoascus, 131
Gymnocraspedidae, 67
Gymnodiniacea, 99, 100
Gymnodiniaceae, 100
Gymnodlniales, 99
Gymnodinida, 100
Gymnodinidae, 100
Gymnodiniidae, 100
Gymnodinina, 99
Gymnodinioidae, 99
Gymnodinium, 100
Gymnodinium Lunula, 101, 104
Gymnodinium striatum, 104
Gymnogongrus, 49
Gymnosporangium, 143, 148
Gymnosporidae, 211
Gymnosporidiida, 209, 211
Gymnostomata, 229
Gymnostomataceae, 229
Gymnostomina, 229
Gyrodinium, 101
Gyromonas, 166
Gyrophragmium, 152
Gyrosigma, 75
Haemamoeba, 213
Hacmamoeba malariac, 213
Haemamoeba vivax, 213
Hacmogregarina, 2 1 1
Haemogregarinida, 211, 212
Haemogregarinidac, 211
Haemogrcgarinina, 2 1 1
Haemoproteidae, 212
Haemoproteus, 213
Haemoproteus Columbae, 212, 213
Hacmosporidae, 211
Hacmosporidia, 207, 211, 212
Hacmosporidiida, 211
Haliarchnion, 49
Halicryptina, 198
Haliomma, 195
Haliomma capillaris, 196
Haliommatina, 195
Halkyardia, 187
Halopappaceae, 60
Halopappus, 60
Halosphaeraceae, 66
Halteria, 231
Halteridiida, 212
Halteridiidae, 212
Halteridium, 212
Halteriidae, 231
Halterina, 231
Hantkenina, 187
Hantkeninidae, 187
Hantschia, 75
Haploactinomyxidae, 222
Haplobacteriacei, 18
Haplocyta, 215
Haplodinium, 99
Haplospora, 87
Haplosporangium, 124
Haplosporangium lignicola, 122
Haplosporidia, 218
Haplosporidies, 218
Haplosporidiida, 218
Haplosporidiidae, 218
Haplosporidiidea, 209, 218
Haplosporidium, 218
Haplostichinae, 82
Haplozoonidae, 102
Hauerinina, 185
Hedriocystis, 194
Helicosorina, 185
Heliodiscus, 195
Heliodiscus Phacodiscus, 196
Heliolithae, 58
Helioflagellida, 189
Helioflagellidae, 191
Heliozoa, 63, 157, 189, 190, 205
Heliozoariae, 189, 190
Heliozoida, 189
Helminthocladeae, 47
Helminthosporium, 142
Helotiacea, 135
Helvellacea, 135
Helvellales, 134
Helvellineae, 134
Hemiascales, 130
Hemiasceae, 130
Hemiasci, 129
Hemiascineae, 130
Hemibasidii, 145, 149
Hcmicristcllaria, 187
Hemicyclomorpha, 18
Hcmidinium, 101
Hemileia vastatrix, 148
Hemisphaeriaceae, 134
Hcmisphaeriales, 133
Hemitrichia, 177
Hemitrichia intorta, 176
Hemophilus, 22
Henneguya, 221
Index
[285
Hepatozoon, 211
Herpetomonas, 161, 162
Heterocapsaceae, 65
Heterocapsales, 63
Heterocapsineae, 55, 63
Heterocarpea, 41, 44, 52
Heterocarpeae, 40, 44
Heterochlorida, 63
Heterochloridaceae, 66
Heterochloridae, 66
Heterochloridales, 63
Heterochloridea, 63
Heterochloridineae, 55, 63
Heterochromonas, 59
Heterococcales, 63
Heterococcineae, 55, 63
Heterodermaceae, 175
Heterodermeae, 171
Heterogeneratae, 82
Heterohelicida, 188
Heterohelicidae, 188
Heterohelix, 188
Heterokonta, 11, 55, 83
Heterokontae, 53, 55, 63
Heteromastigoda, 158
Heteromonadina, 59
Heteronema, 109
Heteronemidae, 108
Heterophryida, 191, 193
Heterophryidae, 193
Heterophrys, 193
Heterosiphonales, 63
Heterosiphoneae, 55, 63
Heterostegina, 188
Heterotricha, 228, 230
Heterotrichaceae, 230
Heterotrichales, 63
Heterotrichida, 230
Heterotrichina, 230
Heterotrichineae, 55, 63
Hexacontium, 195
Hexaconus, 197
Hexactinomyxon, 222
Hexamastix, 167
Hexamastix Termopsidis, 164
Hexamita, 163, 166
Hexamitidae, 166
Hirmocystis, 217
Hodotermitidae, 167
Hoferellus, 221
Holocyclomorpha, 18
Holomastigotoides, 169
Holomastigotoidida, 169
Holomastigotoididae, 169
Holophrya, 229
Holophryidae, 229
Holotricha, 228, 229
Holotrichida, 229
Homalogonata, 69
Homo sapiens, 6
Honey bees, 222
Hoplonympha, 169
Hoplonympha natator, 170
Hoplonymphida, 169
Hoplonymphidae, 169
Hoplorhynchus, 218
Hordeum vulgare, 6
Hormogonales, 34
Hormogoneae, 34
Horse, 231
Hyalobryon, 60
Hyalodiscida, 201, 202
Hyalodiscidae, 202
Hyalodiscus, 202
Hyaloklossia, 211
Hyaloria, 149
Hyalospora, 217
Hydnacea, 151
Hydnaceae, 151
Hydnangiacea, 155
Hydnangiaceae, 155
Hydnei, 151
Hydnum, 151
Hydra, 203, 233
Hydramoeba, 203
Hydrocoleum, 35
Hydrogenomonas, 24
Hydruracea, 61, 62
Hydruraceae, 62
Hydruridae, 62
Hydrurina, 62
Hydrurus, 61
Hydrurus foetidus, 56, 62
Hyella, 36
Hymenogastraceae, 155
Hymenogastrales, 152
Hymenogastrea, 155
Hymenogastrei, 155
Hymenogastrineae, 152
Hymenomonadacea, 58, 60
Hymenomonadaceae, 60
Hymenomonadidae, 60
Hymenornonas, 60
Hymenomycetales, 150
Hymenomycetes, 150
Hymenomycetineae, 150
Hymenothecii, 150
Hymenostomata, 229
Hyperammina, 183
Hyperamminidae, 183
Hypermastigida, 168
Hypermastigina, 158, 166, 168
Hyphochytriacea, 69
Hyphochytriaceae, 69, 117
Hyphochytrialea, 57, 61, 69, 70, 1 11
Hyphochytriales, 69
Hyphochytrium, 69, 117
Hyphochytrium catenoides, 70
Hyphomycetes, 121, 140, 141
Hypnodiniaceae, 100
Hypocreaceae, 137
Hypocreales, 137, 138, 139, 142
Hypodermia, 146, 147
Hypodermii, 147
286]
The Classification of Lower Organisms
Hypomyces, 142
Hypomyces Solani var. Cucurbitae, 126,
127
Hypotricha, 228, 232, 233
Hypotrichaceae, 233
Hypotrichida, 233
Hypotrichina, 233
Hysterangiacea, 155
Hysterangiaceae, 155
Hysteriacea, 134
Hysteriaceae, 133, 134
Hysteriales, 133, 141
Hysteriineae, 133, 134
Hysterophyta, 119
Ichthyophthirius, 229
Ichthyosporidium, 219
Imperforida, 183
Infusoires, 223
Infusoires suceurs, 235
Infusoria, 2, 37, 95, 118, 223, 228, 232,
235
Inoperculata, 135
Inophyta, 39, 119
Insects, 69, 113, 117, 118, 124, 125, 155,
159, 161, 165, 167, 216, 217, 220
Invertebrates, 161, 210, 211, 215, 216
lodamoeba, 203
Irish moss, 49
Irpex, 151
Isoachlya, 79
Isocarpeae, 69
Isochrysidaceae, 59
Isochrysidae, 59
Isochrysidales, 57
Isogeneratae, 82
Isospora, 210
Janczewskia, 52
Jarrina, 210
Joenia, 169
Joeniidae, 169
Joeniidea, 168
Jocnina, 169
Joenopsis, 169
Jola, 147
Junipers, 148
Kalotermes, 169
Kalotermitidae, 167
Kalotcrmitinac, 166, 168
Karyamocbina, 203
Karyolysus, 2 1 1
Kelps, 82, 83, 89, 90
Keramosphaera, 185
Keramosphacridac, 185
Keramosphaerina, 185
Kerona, 233
Klebsiella (bacterium), 7, 22
Klebsiella pneumoniae, 22
Klebsiella (flagellate), 7
Klebsiella alligata, 106
Klossia, 211
Klossiella, 211
Kofoidia, 168, 169
Kofoidiida, 169
Kofoidiidae, 169
Kurthia, 21
Kurthiacea, 19, 21, 237
Laboulbenia, 140
Laboulbenia Guerinii, 140
Laboulbenia Rougetii, 140
Laboulbeniaceae, 140
Laboulbenialea, 129, 140
Laboulbeniales, 140
Laboulbenieae, 140
Laboulbeniineae, 140
Laboulbeniomycetes, 140
Labyrinthula, 203, 204
Labyrinthula macrocystis, 203
Labyrinthulida, 201, 203
Labyrinthulidae, 203
Lachnea scutellata, 127, 136
Lachnobolus, 176
Lacrymaria, 229
Lactobacillaceae, 20
Lactobacillus, 20
Lactobacteriaceae, 20
Lagena (oomycete), 82
Lagena (rhizopod), 82, 184, 187
Lagenaceae, 187
Lagenidae, 186
Lagenidea, 185
Lagenidiacca, 81, 82
Lagenidiaceae, 82
Lagenidialea, 76, 81, 111, 118
Lagenidiales, 81
Lagenidium, 82
Lagenina, 186
Lagenocystis, 82, 237
Lagenocystis radicicola, 82, 237
Lagynida, 191
Lagynion, 63
Lagynis, 191
Laminaria, 91
Laminaria yczoensis, 92
Laminariaccae, 89
Laminariales, 89
Laminariea, 85, 89
Laminarieae, 89
Lampoxanthium, 195
Lampramocbac, 205
Ivamprodernia, 1 75
Lamprodermaceae, 1 75
Lamprospora Iciocarpa, 136
Lamprosporalcs, 171
Lankestcria, 216
Larcarida, 195
Larcoidea, 195
Index
[287
Latrostium, 69
Laurencia, 52
Leangium, 177
Leathesia, 88
Lecudina, 216
Lecudinidae, 216
Leeches, 161, 211
Legerella, 211
Leidyopsis, 169
Leishmania, 162
Leishmania brasiliensis, 162
Leishmania Donovani, 162
Leishmania tropica, 162
Lemanea, 47
Lemna, 69
Lenticulina, 187
Lenticulites, 187
Lentospora, 221
Lenzites, 151
Leocarpus, 177
Leocarpus fragilis, 176
Lepidoderma, 177
Lepidoderma Chailletii, 176
Lepochromulina, 62
Leptodiscida, 100, 102
Leptodiscidae, 102
Leptodiscus, 102
Leptolegnia, 79
Leptomitaceae, 79
Leptomi tales, 77
Leptomitea, 77, 79
Leptomiteae, 79
Leptomitus, 79
Leptomonas, 162
Leptospira, 29
Leptospira icteroides, 29
Leptospira icterohaemorrhagiae, 29
Leptospironympha, 169
Leptostromatacea, 141
Leptostromataceae, 141
Leptotheca, 221
Leptothrix, 27
Leptothrix ochracea, 27, 36
Leptotrichacea, 27
Leptotrichaceae, 20
Leptotrichacei, 20, 27
Leptotrichia, 20
Leucocytozoidae, 212
Leucocytozoon, 213
Leuvenia, 66
Liagora tetrasporifera, 47, 49
Licea, 175
Liceacea, 173, 175
Liceaceae, 175
Liceales, 171
Liceidae, 175
Lichenes, 119
Lichens, 119, 120
Ligniera, 179
Lindbladia, 175
Lionotus, 230
Listeria, 21
Lithocampe, 198
Lithochytridina, 198
Lithocircus, 198
Lithocircus productus, 196
Lithocolla, 193
Lithocollidae, 193
Lithocyclia, 195
Lithocyclidina, 195
Lithocystis, 216
Litholophida, 197
Litholophus, 197
Lituola, 186
Lituolidaceae, 186
Lituolidae, 186
Lituolidea, 185, 186
Lituolina, 186
Liverworts, 10
Lizards, 211
Lobosa, 201
Loborhiza, 117
Lobster, 211
Loftusia, 186
Loftusiidae, 186
Loftusiina, 186
Lophomonadidae, 169
Lophomonadida, 168, 169
Lophomonadina, 168
Lophomonas, 168, 169
Loxodes, 230
Lychnaspis, 197
Lycogala, 172, 175
Lycogala epidendrum, 1 76
Lycogalaceae, 171, 175
Lycogactida, 174, 175
Lycogalactidae, 175
Lycogalales, 171
Lycogalopsis, 155
Lycogalopsis Solmsii, 145
Lycoperdacea, 155
Lycoperdaceae, 155
Lycoperdales, 152
Lycoperdineae, 152
Lycoperdon, 155
Lyngbya, 13, 35
Lytothecii, 152
Macrocystis pyrifera, 90, 91
Macromastix, 58
Macrotrichomonas, 167
Macrotrichomonas pulchra, 164
Maize, 6
Mallomonadidae, 62
Mallomonadinea, 61, 62
Mallomonas, 61, 62
Mallomonas roseola, 56
Mammals, 166, 210
Man, Mankind, Men, 6, 159, 165, 210,
213,230
Margarita, 176
Margaritaceae, 176
Margaritida, 174, 176
288]
The Classification of Lower Organisms
Margaritidae, 176
Massospora, 124, 125
Mastigamoeba, 158, 163
Mastigamoeba aspera, 160
Mastigamoebidae, 163
Mastigella, 163
Mastigophora, 6, 55, 94, 95
Mastotermitidae, 167
Matthewina, 186
Mayorella, 202
Mayorellida, 201, 202
Mayorellidae, 202
Medusetta, 200
Medusettida, 200
Megachytriaceae, 118
Megachytrium, 118
Melampsora, 143
Melampsoracea, 148
Melampsoraceae, 148
Melanconiacea, 141
Melanconiaceae, 141
Melanconialea, 141
Melanconiales, 141
Melanophycea, 11, 55, 82
Melanophyceae, 82
Melanospermeae, 82
Melitangium, 28
Melosira, 72, 73, 74
Melosiraceae, 74
Melosireae, 74
Meningococcus, 20
Menoidium, 103, 107, 109
Menoidium incurvum, 108
Menospora, 218
Menosporida, 218
Menosporidae, 218
Meridiea, 75
Meridieae, 75
Meridion, 75
Meridionaceae, 75
Merismopedia, 33
Merocystis, 211
Merogregarina, 215
Merogregarinida, 215
Merogregarinidae, 2 1 5
Merolpidiaceae, 1 1 7
Meroselenidium, 215
Mesocaena, 63
Mesogloia, 88
Mesogloiacca, 88
Metachaos, 202
Metadevescovina, 167
Metaphyta, 6
Metasporeae, 1 1 7
Metazoa, 6
Metchnikovclla, 219
Metchnikovellida, 219
Mctchnikovellidae, 219
Methanomonas, 24
Micrococcacea, 19, 20
Micrococcaceae, 20
Micrococcus, 20
Microcoleus, 35
Microglena, 62
Micromycopsis, 117
Micropeltidacea, 134
Micropeltidaceae, 134
Microrhopalodina, 166
Microsphaera, 132, 133
Microsphaera alni, 133
Microsporidia, 222
Microsporidies, 222
Microthyriacea, 134, 141
Microthyriaceae, 134
Microthytriales, 133
Miescher's tubes, 206
Mieschersche Schlauche, 206, 214
Mikrogromia, 183
Miliola, 182, 185, 201
Milioles, 179
Miliolida, 185
Miliolidae, 185
Miliolidea, 183, 185
Miliolina, 185
Mindeniella, 79
Mischococcacea, 65, 66
Mischococcaceae, 66
Mischococcus, 66
Mites, 211
Mitraspora, 221
Mitrati, 134
Molds, 142
Molds, water, 77
Mollisiacea, 135
Monaden, 189
Monades, 59
Monadidae, 59, 60
Monadidea, 158
Monadina, 57, 58, 59, 158
Monadineae Tetraplasteae, 191
Monadineae Zoosporeae, 191
Monads, collared, 38
Monas, 38, 54, 59, 60, 158
Monas amyli, 189
Monas Okenii, 31
Monascus, 131
Monera, 6, 12
Moneres, 12, 189
Monilia, 135, 140, 142
Monilia sitophila, 139
Moniliacea, 142
Moniliaceae, 142
Moniliales, 141
Monkeys, 213
Monoblepharella, 112
Monoblepharella Taylori, 114
Monoblcpharidacca, 112
Monoblcpharidaceae, 112
Monoblepharidalea, 111, 114
Monoblepharidales, 1 1 1
Monoblcpharideae, 110
Monoblcpharidineae, 110, 111
Monoblepharis, 111, 112
Monocercomonadida, 167
Index
[289
Monocercomonadidae, 167
Monocercomonas, 167
Monocercomonoides, 165
Monocilia, 66
Monociliaceae, 66
Monocystid gregarines, 209
Monocystida, 216
Monocystidae, 216
Monocystidea, 209, 215'
Monocystiden, 216
Monolpidiaceae, 118
Monomychota, 17
Monopylaria, 190, 196, 198
Monopylea, 198
Monopyleen, 198
Monopylida, 198
Monopylina, 198
Monoschizae. 215
Monosiga, 67, 68
Monosomatia, 179, 183
Monosporea, 210
Monosporees, 209
Monostomina, 191
Morchella, 135
Morchella conica, 136
Mortierella, 124
Mortierellacea, 123, 124
Mortierellaceae, 124
Mosquitoes, 162, 213
Moss, Irish, 49
Mosses, 10
Mouse, Mice, 211,214
Mrazekia, 222
Mrazekiida, 222
Mrazekiidae, 222
Mucedinaceae, 142
Mucedineae, 142
Mucedines, 129, 130, 135
Mucor, 121, 123
Mucor Mucedo, 121, 123
Mucoracea, 123
Mucoraceae, 123
Mucorales, 121
Mucorina, 121, 128
Mucorineae, 121
Mucorini, 121
Mucronina, 188
Mushrooms, 145, 151
Mussels, 211,218
Mutinus, 155
Mycetalia, 119
Mycetoideum, Regnum, 119
Mycetosporidium, 179
Mycetozoa, 119, 157, 171, 176, 203
Mycetozoen, 171, 172
Mycetozoida, 171
Mychota, 1,4,6,8, 10, 12
Mycobacteriacea, 25
Mycobacteriaceae, 25
Mycobacterium, 25
Mycobacterium leprae, 25
Mycobacterium tuberculosis, 25
Mycochytridinae, 113
Mycoderma mesentericum, 24
Mycophyceae, 77
Mycophyta, 119
Mycoporacea, 139
Mycosphaerella, 139
Mycosphaerella personata, 138
Myrioblepharis, 112
Myriogloiacea, 88
Myrionema, 89
Myrionematacea, 88
Myriospora, 2 1 1
Myxidiea, 221
Myxidiees, 221
Myxidiida, 221
Myxidiidae, 221
M>Tcidium, 221
Myxobacter, 28
Myxobacteria, 12, 14
Myxobacteriacea, 28
Myxobacteriaceae, 27, 28
Myxobacter iales, 27
Myxobactralea, 26, 27
Myxobactrales, 27
Myxobolea, 221
Myxobolees, 221
Myxobolida, 221
Myxobolidae, 221
Myxobolus, 221
Myxoceratida, 221, 237
Myxoceros, 221, 237
Myxoceros Blennius, 220, 221, 237
Myxoceros sphaerulosa, 221, 237
Myxochloridae, 66
Myxochrysidaceae, 63
Myxochrysidae, 63
Myxochrysis, 63
Myxochytridinae, 113
Myxococcacea, 28
Myxococcaceae, 28
Myxococcus, 28
Myxococcus coralloides, 26
Myxocystoda, 99
Myxogastres, 171
Myxomycetes, 10, 157, 171, 172, 178
Myxomyceten, 172
Myxomycidium flavum, 143
Myxomycophyta, 171
Myxophyceae, 17, 29, 30
Myxophykea, 29
Myxophyta, 171
Myxoproteus, 221
Myxoschizomycetae, 27
Myxoschizomycetes, 18, 27
Myxosoma, 221
Myxosomatida, 221
Myxosomatidae, 221
Myxosporidia, 206, 219, 220
Myxothallophyta, 171
Myzocytium, 82
290]
The Classification of Lower Organisms
Naegelliella, 62
Naegelliellaceae, 62
Naegelliellidae, 62
Naegleria. 159
Najadea, 60
Nassellaria, 198
Nassellida, 198
Nassoidea, 198
Nassula, 230
Nassulidae, 230
Nautilus, 182, 186, 187
Navicula, 72, 73, 75
Naviculaceae, 75
Naviculales, 74
Naviculea, 75
Naviculeae, 75
Neactinomvxon, 222
Nebela, 205
Nebelida, 205
Nebelidae, 205
Nectria, 141, 142
Nectria cinnabarina, 139
Nectrioidaceac, 141
Nectrioideae, 141
Neisseria gonorrhoeae, 20
Neisseria intracellularis, 20
Neiseria meningitidis, 20
Neisseria Weichselbaumii, 20
Neisseriacea, 19, 20
Neisseriaceae, 20
Neisseriacees, 20
Nemalion, 47
Nemalion multifidum, 49
Nemalionales, 47
Nemalioninae, 47
Nemastomatales, 47
Nematochrysidaceae, 60
Nematochrysis, 61
Nematocystida, 219
Nematodes, 113, 118, 124
Nematothecia, 141
Nematothecii, 141
Neogregarina, 215
Neosporidia, 206, 207, 219
Nephroselmidacea, 98
Nephrosclmidaceae, 98
Nephrosclmidae, 98
Nephroselmis, 98
Nereocystis, 89
Nereocystis Luetkeana, 90, 91
Neurospora, 139, 140
Neurospora crassa, 127
Neusinidae, 186
Neusina, 186
Nevskia, 27
Nidularia, 155
Nidulariaceae, 155
Nidularialcs, 152
Nidularica, 155
Nidularici, 155
Nidulariincae, 152
Nina, 217,218
Nitrobacter, 24
Nitrobacter Winogradskyi, 24
Nitrobacteriacea, 20, 24
Nitrobacteriaceae, 24
Nitromonas, 24
Nitrosococcus, 24
Nitrosococcus nitrosus, 24
Nitrosomonas europaea, 24
Nitrosomonas javanensis, 24
Nitzschia, 75
Nitzschiacea, 75
Nitzschiaceae, 75
Noctiluca, 95, 99, 102
Noctiluca miliaris, 102
Nectiluca scintillans, 102, 104
Noctilucae, 94, 99
Noctilucida, 100, 102
Noctilucidae, 102
Nodosalida, 186
Nodosarella, 188
Nodosaria, 184, 187
Nodosarida, 186
Nodosaridae, 186
Nodosarina, 186, 188
Nodosaroum, 186
Nodosinella, 186
Nodosinellida, 186
Nodosinellidae, 186
Nonion, 184, 187
Nonionidea, 187
Nonionideae, 187
Nonionina, 187
Nosema, 222
Noscma bombycis, 206, 222
Nosematidae, 222
Nostoc, 35
Nostocacea, 34, 35
Nostocaceae, 35
Nostochineae, 33
Nowakowskiella, 118
Nowakowskiellacea, 117, 118
Nowakowskiellaceae, 118
Nubecularina, 185
Nucleophaga, 118
Nuda, 201
Nummulitaceae, 188
Nummulites, 188
Nummulitida, 188
Nummulinidae, 188
Nunmiulitina, 188
Nummulitinidea, 183, 185, 188
Oats, 148
Ochromonadaceae, 59, 60
Ochromonadalea, 54, 56, 57, 61, 64, 67,
85, 165
Ochromonadales, 57
Ochromonadidae, 59
Ochromonas, 58, 59, 60
Ochromonas granulans, 54
Octomitus, 166
Index
[291
Octomyxa, 179
Octospora, 222
Oicomonadacea, 159, 161
Oicomonadaceae, 161
Oicomonadidae, 161
Oidium, 142
Oikomonas, 161
Oligochaet worms, 222
Oligonema, 177
Oligosporea, 209, 210
Oligotricha, 230
Oligotrichaceae, 230
Oligotrichida, 230
Oligotrichina, 230
Olpidiacea, 115, 118
Olpidiaceae, 118
Olpidopsidacea, 81
Olpidiopsidaceae, 81
Olpidiopsis, 81
Olpidium, 113, 118
Olpidium Allomycetos, 116
Ommatida, 195
Onygena, l31
Oodinidae, 102
Oomycetes, 11, 53, 55, 65, 76, 78,
118, 119, 121, 125, 127, 177, 178,
Oosporeae, 77
Opalina, 225, 227, 229
Opalinalea, 228, 237
Opalinida, 228
Opalinidae, 228, 229
Opalinina, 229
Opalininea, 228
Opalinoea, 225, 229
Operculata, 135
Operculina, 188
Ophiocytiaceae, 66
Ophiocytium, 66
Ophiocytium parvulum, 66
Ophiotheca, 176
Ophrydium, 233
Ophryocystis, 215
Ophryocystida, 2 1 5
Ophryocystidae, 215
Ophryodendrida, 236
Ophryodendridae, 236
Ophryodendrina, 236
Ophryodendron, 236
Ophryoglena, 229
Ophryoglenidae, 229
Ophryoscolecidae, 231
Ophryoscolecids, 225
Ophryoscolccina, 231
Ophryoscolex, 231
Ophthalmidium, 184, 185
Opisthokonta, 39, 110, 121, 237
Opistokonten, 111
Orbitoides, 188
Orbitoidida, 188
Orbitoididae, 188
Orbitolina, 186
Orbitolinida, 186
Orbitolinidae, 186
Orbitolites, 185
Orbulina, 188
Orbulinida, 188
Orcadella, 175
Orcadellaceae, 175
Orcadellidae, 175
Orcheobius, 211
Orobias, 188
Ortholithinae, 58
Orthopteran, 217
Orthosporeae, 117
Oscillaria malariae, 213
Oscillatoria, 30, 35, 36
Oscillatoria Princeps, 13
Oscillatoria splendida, 32
Oscillatoriacea, 34, 35
Oscillatoriaceae, 35
Owl, 162
Ox, Oxen, 162, 231
Oxymonadida, 165, 166
Oxymonadidae, 166
Oxymonadina, 163
Oxymonas, 163, 166
111, Oxyphysis, 103
179 Oxyrrhis, 101_
Oxyrrhis marina, 101
Oxytocum, 103
Oxytricha, 233
Oxytrichidae, 233
Oxytrichina, 233
Pacinia, 23
Pacinia cholerae-asiaticae, 23
Padina, 87
Palatinella, 62
Pantostomatales, 158
Pantostomatida, 158
Pantostomatineae, 158
Paradinida, 98
Pardinidae, 98
Paradinium Pouchetii, 97, 98
Paraisotricha, 231
Parajoenia, 167
Paramaecium, 223, 224, 225, 226, 227,
229
Paramaecium Aurelia, 226, 227
Paramaecium Bursaria, 226
Paramaecium caudatum, 226
Paramaecium multimicronucleatum, 226
Parameciina, 229
Paramoeba Eilhardi, 98
Paramoebida, 98
Paramoebidae, 98
Paramoecidae, 229
Parasitella, 123
Parvobacteriaceae, 22
Pasteurella avicida, 22
Pasteurella pestis, 22
Pasteurellacea, 19, 22, 23, 237
Pasteuria, 26, 27
292]
The Classification of Lower Organisms
Patellariacea, 135
Patellina, 181, 182, 185
Patouillardina, 149
Patouillardina cinerea, 145
Pavonina, 188
Peach, 137 _
Pectobacterium, 23
Pectobacterium carotovorum, 23
Pedangia, 186
Pedilomonas, 111
Pedinella, 62, 63
Pegidia, 188
Pegidiida, 188
Pegidiidae, 188
Pelodictyon, 31
Pelomyxa, 202
Pelomyxa carolinensis, 200, 201
Pelomyxa palustris, 202
Peneroplidae, 185
Peneroplidea, 185
PeneropHdina, 185
Peneroplis, 181, 184, 185
Penicillium, 130, 131
Penicillium notatum, 25, 131
Pennatae, 74
Penta trichomonas, 165
Pentatrichomonas obliqua, 164, 167, 237
Peranema, 108, 109
Peranema trichophorum, 108
Peranemaceae, 108
Peranemina, 108
Perforida, 186
Periblasteae, 47
Perichaena, 176
Perichaenacea, 174, 176
Perichaonaceae, 176
Peridinaca, 102, 103
Peridinea, 96
Peridineae, 94, 96, 103
Peridiniaceae, 103
Peridiniales, 102
Peridinidae, 103
Peridiriina, 103
Peridinioidae, 103
Peridinium, 94, 103
Peridinium cinctum, 104
Perionella, 66
Pcripylaria, 194
Pcripylea, 194
Peripyleen, 194
Peripylida, 194
Peripylina, 194
Perisporia, 131
Porisporiacea, 129, 131
Perisporiaceae, 131
Perisporialcs, 131
Peritricha, 233
Peritrichaceae, 233
Peritrichida, 233
Peritrichinae, 18
Pcritromidae, 233
Peritromina, 233
Peritromus, 233
Peronospora, 81
Peronosporacea, 80, 81
Peronosporaceae, 81
Peronosporales, 80
Peronosporina, 76, 80
Peronosporinae, 80
Peronosporineae, 80
Peziza, 127, 135
Peziza domiciliana, 127
Pezizacea, 135
Pezizales, 134
Pezizineae, 134
Pestallozia, 141
Pfeiflferella mallei, 22
Phacidiaceae, 133, 134
Phacidiacei, 133
Phacidialea, 129, 133, 135
Phacidiales, 133
Phacidiea, 134, 141
Phacidieae, 134
Phacidiineae, 133, 134
Phacus, 94, 106, 107
Phaenocystes, 219
Phaenocystida, 219
Phaeocapsa, 98
Phaeocapsaceae, 98
Phaeocapsales, 96
Phaeococcus, 98
Phaeoconchia, 198, 199
Phaeocystia, 198
Phaeocystina, 199
Phaeocystis, 58
Phaeocystis globosa, 54
Phaeodaria, 199
Phacodariae, 198
Phaeodermatium, 63
Phaeogromia, 198, 199
Phaeophyceae, 53, 82, 95
Phacophycophyta, 53
Phaeophyta, 39, 53
Phaeoplakaceae, 98
Phaeoplax, 98
Phaeosphaera, 59
Phacosphaeria, 190, 196, 198, 199
Phaeosporales, 86
Phaeosporcae, 82, 86
Phaeothamnion, 61
Phaeothamnionacea, 58, 60
Phaeothamnionaccae, 60
Phaeozoosporea, 85, 86, 87
Phaeozoosporeac, 86
Phagomyxa, 179
Phalanastrriaccae, 67
Phalanastcriidae, 67
Phalanasterium, 67
Phalanasterium digitatum, 68
Phallaccae, 155
Phallales, 152
Phallineac, 152
Phalloidca, 155
Phalloidei, 155
Index
[293
Phallus, 155
Phlebotomus, 21
Phleospora, 139
Phlyctidiacea, 115, 117
Phlyctidiaceae, 117
Phlyctidium, 117
Phlyctorhiza, 117
Phoma, 141
Phomaceae, 141
Phomales, 141
Phomatacea, 141
Phomataceae, 141
Phomatalea, 141
Phomatales, 141
Phormidium, 32, 35
Phraginidium, 147, 148
Phragmidium violaceum, 147
Phycochromaceae, 29
Phycomyces, 123, 124
Phycomyces nitens, 122
Phycomyceten, 76
Phycomycetes, 76
Phycomycophyta, 76
Phyllactinia, 132, 133
Phyllactinia corylea, 127
Phyliophora, 49
Phyllosiphon, 67
Phyllosiphonacea, 67
Phyllosiphinaceae, 67
Physaraceae, 171, 177
Fhysaraks, 171, 174
Physarea, 174, 177
Physaridae, 177
Physarum, 177
Physarum notabile, 176
Physarum polycephalum, 176
Physematium, 189, 195
Physoderma, 115, 117
Physodermataceae, 117
Physomonas, 59
Phytodiniacea, 99, 100
Phytodiniaceae, 100
Phytodinidae, 100
Phytodinium, 100
Phytomastigophorea, 55
Phytomonas (bacterium), 7, 23
Phytomonas (flagellate), 7, 161
Phytomonas Donovani, 160
Phytomyxida, 111, 171, 177
Phytomyxidae, 179
Phytomyxinae, 177
Phytomyxini, 177
Phytophthora, 80
Phytophthora infestans, 81
Phytosarcodina, 171
Phytozoidea, 94, 105
Pigeon, 212
Pileati, 150
Pileocephalus, 218
Pilobolus, 121, 124
Pinaciophora, 193
Pinacocystis, 193
Pines, 148
Pinnularia, 72, 75
Pipetta, 195
Piptocephalidacea, 123, 124
Piptocephalidaceae, 124
Piptocephalis, 123, 124
Piroplasma, 214
Pisces, 1
Plagiotomidae, 230
Plagiotomina, 230
Planopulvinulina, 187
Planorbulina, 187
Planorbulinidae, 187
Plant kingdom, Plantae, Plants, 1,2, 4, 6,
8, 10, 24, 38, 61, 67, 95, 113, 117, 118,
130, 137, 148, 151, 161, 177, 179, 202
Plasmodida, 213
Plasmodidae, 213
Plasmodiida, 211
Plasmodiophora, 179
Plasmodiophora Brassicae, 178
Plasmodiophoraceae, 1 79
Plasmodiophorales, 177
Plasmodiophorea, 179
Plasmodiophoreae, 179
Plasmodiophoreen, 179
Plasmodiophorina, 177
Plasmodium, 212, 213
Plasmodium falciparum, 214
Plasmodium malariae, 213
Plasmodium vivax, 213
Plasmodroma, 157
Plasmopara viticola, 81
Platychrysis, 58
Platygloea, 147
Platynoblasteae, 51
Platysporea, 221
Plectascales, 130
Plectascineae, 130
Plectellaria, 198
Plectida, 198
Plectobasidiales, 152
Plectobasidiineae, 152
Plectofrondicularia, 188
Plectoidea, 198
Plectonema, 36
Plectonida, 198
Pleurage curvicolla, 128
Pleurocapsa, 36
Pleurocapsacea, 35, 36
Pleurocapsaceae, 36
Pleuromonas (dinoflagellate), 99
Pleuromonas (zoomastigote), 159
Pleuronemidae, 230
Pleurosigma, 75
Pleurostomella, 188
Pleurostomellida, 188
Pleurostomellidae, 188
Pleurotricha, 233
Pleurotrichidae, 233
Pleurotus, 152
Pleurotus ostreatus, 152
294]
The Classification of Lower Organisms
Plistophora, 222
Plistophoridae, 222
Plocapsilina, 186
Plocapsilinidae, 186
Plowrightia morbosa, 140
Pneumobacilliis, 22
Podangium, 28
Podaxacea, 152
Podaxaceae, 152
Podaxon, 152
Podocyathus, 236
Podophrya, 235
Podophyridae, 235
Podophryina, 235
Podosphaera, 132, 133
Polyangiaceae, 28
Polyangidae, 27
Polyangium, 28
Polychaos, 202
Polychytrium, 117
Polycystidea, 209, 216
Polycystina (of Ehrenberg), 189, 198
Polycystina (of Delage and Herouard),
217
Polydinida, 101
Polygastrica, 223
Polykrikida, 100, 101
Polykrikos, 101
Polymastigida, 158, 163, 164
Polymastigidae, 165
Polymastigina, 158, 163, 165
Polymastix, 163, 165
Polymastix melolonthae, 164
Polymorphina, 187
Polymorphinida, 187
Polymorphinidae, 187
Polymorphinina, 187
Polymyxa, 178, 179
Polyphagaceae, 117
Polyphagus, 111, 117
Polyphagus Euglcnae, 116, 117
Polyporacea, 151
Polyporaceae, 151
Polyporales, 150
Polyporei, 151
Polyporus, 151
Polysiphonia nigrescens, 49
Polysiphonia violacea, 45, 46
Polysiphonieae, 51
Polysomatia, 179, 185
Polysphondylium, 203
Polysphondylium violaceum, 204
Polysporea, 209,211
Polystichinae, 82
Polystictus, 151
Polystomclla, 186, 187
Polystomella crispa, 181
Polystomellina, 187
Polythalamia, 179, 185
Polytoma, 61
Pontifex, 202
Pontisma, 81
Pontosphaera, 60
Pontosphaeraceae, 60
Porospora, 218
Porosporida, 218
Porosporidae, 218
Porphyra, 43
Porphyra laciniata, 42
Porphyra tenera, 42, 43
Porphyra umbilicaris, 42, 43
Porphyraceae, 43
Porphyrea, 41, 43
Porphyreae, 43
Porphyridiacea, 41
Porphyridiaceae, 41
Porphyridiales, 4l
Porphyridium, 3, 40
Porphyridium cruentum, 41
Postelsia palmaeformis, 90, 91
Poteriochromonas, 60
Poteriodendron, 67
Poteriodendron petiolatum, 68
Pouchetia, 101
Pouchetiida, 100, 101
Pouchetiidae, 101
Prasiola, 3, 40, 44
Prasiolaceae, 44
Primalia, 37
Primigenium, Regnum, 37
Proboscoidella, 166
Progastreades, 94, 95
Pronoctiluca, 101
Prorocentraceae, 99
Prorocentrales, 99
Proroccntridae, 99
Proroccntrina, 99
Prorocentrinea, 98
Prorocentrinen, 99
Prorocentrum, 99
Prorodon, 229
Protamoeba, 189
Proteomyxa, 189, 190
Proteomyxiae, 189, 190
Proteomyxida, 189
Proteromonadidae, 159
Protcromonadina, 158
Proteromonas, 159
Proterospongia Haeckcli, 68
Proteus diffluens, 201
Proteus vulgaris, 22
Protista, 4, 6, 37, 189
Protistcs trichocystiferes, 94, 95
Protoascineac, 130
Protobasidiomycetes, 145, 146, 150
Protobionta, 6, 37
Protochrysis, 98
Protociliata, 228
Protoctista, 1, 4,6,8, 10,37
Protodennieae, 171
Protodinifcr, 101
Protodiniferida, 100, 101
Protodiniferidae, 101
Protodiscineae, 137
Index
[295
Protodontia Uda, 145
Protoflorideae, 41
Protogenes, 189
Protomastigales, 158
Protomastigida, 158
Protomastigina, 158
Protomastigineae, 158
Protomonas, 189, 191
Protomonadina, 158
Protomyces, 130
Protoopalina, 229
Protoopalinidae, 229
Protophyta, 6, 12, 18
Protoplasta, 39, 111, 157
Protoplasta filosa, l90
Protopsis, 101
Protozoa, 6, 12, 29, 37, 39, 223
Prowazekia, 159
Prunoidea, 195
Prunophracta, 197
Prymnesiidae, 58
Prymneslum, 58
Pseudomonas, 23
Pseudomonas aeruginosa, 23
Pseudospora, 159, 189, 191
Pseudosporea, 19l
Pseudosporeae, 191
Pseudosporeen, 191
Pseudosporidae, 191
Pseudotetraedron, 66
Pseudotetraedron neglectum, 64
Psorosperms, 206
Psychodiere, Regne, 37
Psychodies, 37
Pteridomonas, 193
Pterocephalus, 218
Pterospora, 216
Ptychodiscida, 103
Puccinia, 143, 147, 148
Puccinia graminis, 147, 148
Puccinia Malvacearum, 148
Pucciniaceae, 148
Pucciniales, 147
Puffballs, 155, 172
Punctariales, 89
Pycnospermeae, 82, 89
Pylaiella, 86
Pyrenomycetales, 138
Pyrenomycetes, 137
Pyrenomycetineae, 137
Pyrgo, 185
Pyrocystis, 100
Pyronema, 127, 134, 135,_ 137
Pyronema confluens var. igneum, 127
Pyronemacea, 135
Pyrrhophycophyta, 94
Pyrrhophyta, 39, 94, 182
Pyrsonympha, 166
Pyrsonymphina, 163
Pythiacea, 80
Pythiaceae, 80
Quadrula, 205
Rabbit, 210
Raciborskya, 100
Radaisia, 36
Radioflagellata, 190
Radiolaria, 189, 190, 194, 196
Radiolariae, 189
Radiolarida, 189
Ralfsia, 87, 89
Ralfsiacea, 88
Ramularia, 139
Ramulinina, 187
Raphidophrys, 193
Raphidozoum, 195
Rat, 160
Ravenelia, 148
Red algae, see Algae, Red
Regne Psychodiere, 37
Regnum Mycetoideum, 119
Regnum Primigenium, 37
Reophacida, 186
Reophacidae, 186
Reophax, 186
Reptiles, 212, 220
Reticularia, 175, 179
Reticulariacea, 174, 175
Reticulariaceae, 175
Reticularieae, 171
Reticulitermes, 171
Reticulosa, 179
Retortomonadidae, 165
Retortomonadina, 163
Retortomonas, 163, 165
Rhabdogeniae, 207
Rhabdosphaera, 60
Rhipidiacea, 77, 79
Rhipidiaceae, 79
Rhipidium, 79
Rhizammina, 183
Rhizamminidae, 183
Rhizaster, 63
Rhizidiacea, 115, 117
Rhizidiaceae, 117
Rhizidiomyces, 69
Rhizidiomyces apophysatus, 70
Rhizidiomycetaceae, 69
Rhizidium, 113, 117
Rhizinacea, 135
Rhizo-Flagellata, 158
Rhizobiacea, 19, 22, 23
Rhizobiaceae, 22
Rhizobium, 23
Rhizobium Leguminosarum, 23
Rhizochloridaceae, 66
Rhizochloridae, 66
Rliizochloridales, 63
Rhizochloridea, 63
Rhizochloridineae, 55, 63
Rhizochloris, 66
Rhizochrysidaceae, 63
296]
The Classification of Lower Organisms
Rhizochrysidae, 63
Rhizochrysidina, 61
Rhizochrysidinae, 61
Rhizochrysidineae, 55
Rhizochrysis, 61, 63
Rhizochrysis Scherffeli, 56
Rhizocryptineae, 95
Rhizoctonia, 142
Rhizodiniales, 99, 101
Rhizodininae, 95, 99
Rhizoflagellata, 157, 158, 160, 178, 192
Rhizomastigaceae, 163
Rhizomastigida, 158
Rhizomastigina, 158, 163
Rhizomastix, 163
Rhizopoda, 6, 63, 157, 172, 179, 184, 200,
205
Rhizopoda radiaria, 189, 194
Rhizopods, 179
Rhizopodes, 179
Rhizopogonacea, 155
Rhizopogonaceae, 155
Rhizopus, 121
Rhizopus nigricans, 122, 124
Rhizosolenia, 74
Rhizosoleniacea, 74
Rhizosoleniaceae, 74
Rhodobacillacea, 31, 237
Rhodobacillus, 31
Rhodobacteria, 30, 31
Rhodobacteriaceae, 31
Rhodochaetacea, 41, 43
Rhodochaetaceae, 43
Rhodochaete, 43
Rhodochorton, 47
Rhodomelaceae, 51
Rhodomeleae, 51
Rhodomonas, 98
Rhodomonas baltica, 97
Rhodophyceac, 6, 40
Rhodophycophyta, 40
Rhodophyllis, 49
Rhodophyta, 39, 40, 44
Rhodopseudomonas, 31
Rhodospermeae, 40
Rhodospirillum, 31
Rhodymcniacea, 51
Rhodymeniaceae, 51
Rhodymeniales, 51
Rhodymcnieae, 51
Rhodymeninae, 51
Rhoicosphenia, 76
Rhoicosphenia curvata, 72
Rhopalodia, 75
Rhynchocystida, 216
Rhynchocystidae, 216
Rhynchocystis, 216
Rhynchomonas, 159
Rickettsia Mclophagi, 21
Rickettsia Prowazekii, 21
Rickettsia Rickettsii, 21
Rickettsiacea, 19, 20, 118
Rickettsiaceae, 20
Rivularia, 36
Rivulariacea, 34, 36
Rivulariaceae, 36
Roach, 166, 168,170
Rodents, 211
Roesia, 69
Rosaceae, 148
Rotalia, 184, 187
Rotaliaceae, 187
Rotalida, 187
Rotalidae, 187
Rotalidea, 187
Rotalina, 187
Rotifers, 113, 118, 219
Rozella, 118
Rugipes, 202
Rupertia, 187
Rupertiidae, 187
Russula, 143
Russula emetica, 145
Rusts, 145, 147
Rye, 148
Saccamminidae, 183
Saccharomyces cerevisiae, 130
Saccharomycetacea, 130
Saccharomycetaceae, 130
Saccharomycetes, 130
Saccinobaculus, 163, 166
Sagosphaerida, 199
Sagrina, 188
Salmonella, 22
Salpingoeca, 67
Salpingoeca ampullacea, 68
Salpingoeca Clarkii, 68
Salpingoecidae, 67
Sappinia, 203
Sappinia diploidea, 203
Sappinia pedata, 204
Sappiniaceae, 203
Sappiniidae, 203
Saprolegnia, 76, 79
Saprolegnia ferax, 78
Saprolegnia mixta, 78
Saprolegniaceae, 77
Saprolegniales, 77
Saprolegniea, 77
Saprolegnicae, 77
Saprolcgniineae, 77
Saprolegnina, 77
Saprolegninae, 77
Sapromyces, 79
Saprospira, 29
Sarcina, 20
Sarcocystida, 214
Sarcocystidae, 214
Sarcocystidca, 214
Sarcocystis, 214
Sarcocystis Miescheriana, 214
Sarcocystis Muris, 214
Index
[297
Sarcodina, 6, 172, 200
Sarcosporidia, 207, 214
Sargassaceae, 91
Sargassea, 92
Sargasseae, 92
Sargassum, 93
Sargassum Horneri, 93
Saricodina, 63, 157,200
Schaudinella, 216
Schaudinellida, 216
SchaudinelHdae, 216
Schinzia Leguminosarum, 23
Schizocystida, 215
Schizocystidae, 215
Schizocystinea, 215
Schizocystis, 215
Schizodinium, 102
Schizogoniacea, 41, 44
Schizogoniaceae, 44
Schizogonium, 44
Schizogregarinaria, 215
Schizogregarinida. 209, 215
Schizomycetae, 17, 18
Schizomycetes, 18, 206
Schizomycophyta, 17
Schizophyta, 12, 18
Schizophytae, 12
Schizosporea, 18
Schlauche, Mieschersche, 206, 214
Sciadiaceae, 66
Sciadophora, 218
Sclerocarpa, 129, 133, 135, 137, 145
Sclerocarpi, 137
Scleroderma, 143
Sclerodermataceae, 155
Sclerodermatales, 152
Sclerodermea, 155
Sclerodermei, 155
Sclerotinia, 140
Sclerotinia cinerea, 135, 136
Scytomonas pusilla, 108
Scytonema, 36
Scytonematacea, 34, 35
Scytonemataceae, 35
Sebacina, 149
Sebacina sublilacina, 145
Sebdenia, 49
Selenidium, 215
Seleniida, 215
Seleniidae, 215
Selenococcidiida, 211
Selenococcidiidae, 211
Sclenococcidinea, 210
Selenococcidium intermedium, 211
Sennia, 97, 98
Sepedonei, 141
Septata, 217
Septobasidium, 147
Septoria, 139, 141
Sheep, 210, 214
Shigella, 22
Shigella dysenteriae, 22
Serratia, 22
Siderocapsa, 27
Sideromonas, 27
Siedleckia, 215
Silicina, 185
Silicoflagellata, 55, 56, 57, 61, 62, 64, 67,
69
Silicoflagellatae, 55, 62
Silicoflagellidae, 62
Silicoflagellina, 61
Silkworms, 206, 222
Sinuolinea, 221
Siphonaria, 117
Siphonogenerina, 188
Siphonomycetae, 77
Siphonophyceae, 55
Siphonotestales, 62
Sirolpidiacea, 81
Sirolpidiaceae, 81
Sirolpidium, 81
Sirosiphon, 36
Sirosiphonacea, 34, 36
Sirosiphonaceae, 36
Slavina, 222
Smuts, 145, 149
Snails, 161,211
Snakes, 210
Snyderella, 168
Snyderella Tabogae, 164
Solenodinium, l99
Sorangiacea, 28
Sorangiaceae, 28
Sorangium, 28
Soranthera, 89
Sorites, 185
Soritidae, 185
Soritina, 185
Sorodiscus, 179
Sorophoreen, 203
Sorosphaera, 179
Sphacelaria, 86
Sphacelarialea, 85, 86
Sphacelariales, 86
Sphacelariea, 86
Sphacelarieae, 86
Sphaeractinomyxon, 222
Sphaerastrum, 193
Sphaerellaria, 194
Sphaeria, 138, 141
Sphaeria Scirpi, 128
Sphaeriaceae, 137
Sphaeriales, 137, 138, 139, 141
Sphaerida, 195
Sphaeridea, 194
Sphaerioidaceae, 141
Sphaerioideae, 141
Sphaerita, 118
Sphaerobolacea, 155
Sphaerobolaceae, 155
Sphaerobolus, 155
Sphaerocapsa, 197
Sphaerocapsida, 197
298]
The Classification of Lower Organisms
Sphaerocladia, 112, 113
Sphaerococcales, 47
Sphaerococcoidea, 46, 47, 50
Sphaerococcoideae, 47
Sphaeroeca, 67
Sphaeroidea, 195
Sphaeroidina (genus of Rhizopoda), 187
Sphaeroidina (family of Radiolaria), 195
Sphaeromyxa, 221
Sphaerophracta, 197
Sphaerophrya, 235
Sphaeropsidales, 141
Sphaeropsideae, 141
Sphaerospora, 221
Sphaerosporida, 221
Sphaerosporidae, 221
Sphaerosporea, 221
Sphaerotheca, 127, 133
Sphaerotheca pannosa, 133
Sphaerotilacea, 33
Sphaerotilaceae, 33
Sphaerotilalea, 30, 33, 237
Sphaerotilus, 30
Sphaerotilus natans, 33
Sphaerozoen, 194
Sphaerozoida, 195
Sphaerozoum, 189, 195
Spirillacea, 19, 23
Spirillaceae, 23
SpirilHna, 181, 182, 185
Spirillinidea, 185
Spirillinina, 185
Spirillum, 24
Spirochaeta, 29
Spirochaeta cytophaga, 26, 27
Spirochaeta plicatilis, 28, 29
Spirochaetacea, 29
Spirochaetaceae, 29
Spirochaetae, 27
Spirochactalea, 28
Spirochaetales, 28
Spirochaets, 12, 14, 166, 167
Spirochona, 233, 235
Spirochonidae, 231
Spirochonina, 230, 231
Spirocystida, 215
Spirocystidae, 215
Spirocystidces, 215
Spirocystis, 215
Spirodinium, 231
Spirodiscus, 66
Spirodiscus fulvus, 64, 66
Spirogyrales, 121
Spirolina, 185
Spironema, 222
Spirophyllum, 27
Spirostomum, 230
Spirotricha, 230
Spirotrichida, 230
Spirotrichonympha, 168, 169
Spirotrichonymphidae, 169
Spirotrichonymyjhina, 168
Spirulina, 35
Sponges, 37, 67
Spongocarpeae, 50
Spongospora, 179
Spongurida, 195
Spongurus, 195
Sporobolomyces, 145
Sporochnales, 87
Sporochnea, 88
Sporochnoidea, 85, 87, 89
Sporochnoideae, 87
Sporochnus, 93
Sporodinia, 124
Sporochytriaceae, 117
Sporomyxa, 179
Sporozoa, 111, 206, 207, 219
Sporozoans, 21, 162
Sporozoaires, 207
Sporozoaria, 206, 207
Spumaria, 177
Spumariaceae, 177
Spumellaria, 194, 195
Spyrida, 198
Spyridieae, 51
Spyridina, 198
Spyroidea, 198
Squamarieae, 50
Squids, 210
Staphylococcus, 20
Staurocyclia, 195
Staurojoenina, 169
Staurojoenina assimilis, 170
Staurojoeninida, 169
Staurojoeninidae, 169
Stelangium, 28
Stemonitaceae, 171, 175
Stemonitales, 171, 174
Stemonitea, 174, 175
Stemonitidae, 175
Stemonitis, 175
Stemonitis splendens, 176
Stenophora, 217
Stenophorida, 21 7
Stenophoridae, 217
Stentor, 227, 230
Stentor coeruleus,
Stentoridae, 230
Stcntorina, 230
Stephanida, 198
Stephanonympha,
Stephida, 198
Stephoidea, 198
Stereotestales, 62
Stereum, 151
Stictaceac, 134
Stictea, 134
Sticteac, 134
Stictidaceac, 134
Stictideae, 133
Stigonema, 36
Stigoncmataceae, 36
Stiibaceae, 142
225
168
Index
[299
Stilbeae, 142
Stilbellacea, 142
Stilbellaceae, 142
Stilbosporei, 141
Stilbum, 142
Stilophora, 88
Stilotricha, 233
Stipitochloridae, 66
Stipitococcacea, 65, 66
Stipitococcaceae, 66
Stipitococcus, 66
Stokesiella, 60
Stomaticae, 74
Stomatoda, 223, 228, 233
Stomatophora, 216
Stomatophorida, 216
Stomatophoridae, 216
Streblomastigida, 165, 166
Streblomastigidae, 166
Streblomastix, 163, 168
Streblomastix Strix, 164, 166
Streblonema, 86
Streptococcus, 20
Streptomyces, 25
Streptomycetaceae, 25
Streptothrix, 25
Striatae, 74
Stylobryon, 60
Stylocephalida, 218
Stylocephalidae, 218
Stylocephalus, 218
Stylochrysalis, 59
Stylocometes, 236
Stylodinium, 100
Stylonychia, 227, 232, 233
Stylopage, 124
Stylopyxis, 60
Stylorhynchidae, 218
Stypocaulon, 83, 84, 86
Suctorea, 235
Suctoria, 235
Surirella, 71, 73, 75
Surirella saxonica, 72, 73
Surirellaceae, 75
Surirellea, 75
Surirelleae, 75
Swine, 210, 214
Symbelaria, 194
Symploca Muscorum, 13
Synactinomyxida, 222
Synactinomyxidae, 222
Synactinomyxon, 222
Synchytriacea, 115, 117
Synchytriaceae, 117
Synchtrium, 117
Syncephalastrum, 124
Syncephalastrum racemosum, 122
Syncephalis, 123, 124
Syncephalis nodosa, 122
Syncephalis pycnosperma, 122
Syncollaria, 194
Syncrypta, 59
Syncryptaceae, 59
Syncryptida, 58, 59
Syncryptidae, 59
Syncystida, 216
Syncystidae, 216
Syncystis, 216
Syndinidae, 102
Synedra, 72, 75
Syntamiidae, 86
Synura, 55, 59
Synura Uvella, 54
Synuraceae, 59
Syracosphaera, 60
Syracosphaera Quadricornu, 56
Syracosphaeraceae, 60
Syracosphaeridae, 60
Syracosphaerinae, 57, 60
Tabellaria, 75
Tabellariaceae, 75
Tabellariea, 75
Tabellarieae, 75
Taphrina, 127, 137
Taphrina aurea, 137
Taphrina deformans, 127, 136, 137
Teliosporeae, 142
Telomyxa, 222
Telomyxa glugeiformis, 222
Telomyxlda, 222
Telomyxidae, 222
Telosporidea, 207
Telosporidia, 207
Tentaculifera, 224, 228, 235
Teratonympha, 171
Teratonympha mirabilis, 170
Teratonymphida, 169
Teratonymphidae, 169
Termites, 166, 167, 168, 169
Termitidae, 168
Termopsis, 166, 168
Testacea, 205
Testacida, 205
Testaceolobosa, 205
Tetractinomyxida, 222
Tetractinomyxidae, 222
Tetractinomyxon, 222
Tetradinium, 100
Tetradinium javanicum, 104
Tetrahymena, 229
Tetrahymena Geleii, 227
Tetramitaceae, 165
Tetramitida, 165
Tetramitidae, 165
Tetramitina, 165
Tetramitus, 165
Tetramyxa, 179
Tetrasporeae, 82, 86
Tetrasporees, 209
Tetrataxis, 186
Textularia, 182, 186
Textulariaceae, 186
300]
The Classification of Lower Organisms
Textularidae, 186
Textularidea, 185
Textularlna, 186
Textulinida, 186
Thalamophora, 179
Thalassicolla, 189, 194, 195
Thalassicollen, 194, 195
Thalassicollida, 195, 199
Thallochrysidacea, 62, 63
Thallochrysidaceae, 63
Thallochrysis, 63
Thamnidium, 124
Thaumatomastix, 109
Thaumatomonadidae, 109
Thaumatonema, 109
Thaumatonemidae, 109
Thecamoeba, 202
Thecamoebae, 205
Thecamoebida, 201, 202
Thecamoebidae, 202
Theileria, 214
Theileridae, 214
Thelephora, 151
Thelephoracea, 151
Thelephoraceae, 151
Thelephorei, 151
Thelohania, 222
Theoconus, 198
Thiere, 172
Thiobacillus, 24
Thiobacteria, 30, 31, 35
Thiorhodaceae, 31
Thioploca, 35
Thiospira, 24, 31
Thiospirillum, 31
Thiothrix, 35
Thoracosphaeraceae, 60
Thoracosphaeridae, 60
Thorea, 47
Thraustochytriacea, 81, 82
Thraustochytriaceae, 82
Thraustochytrium proliferum, 82
Thraustotheca, 79
Ticks, 161, 206
Tilletia, 149
Tilletia Tritici, 145
Tilletiacea, 149
Tilletiaceae, 149
Tilopteridales, 86
Tilopteridca, 87
Tilopterideae, 87
Tilopteris, 87
Timothy, 148
Tinoporidea, 187
Tinoporus, 187
Tintinnidac, 231
Tintinnids, 224
Tintinnina, 231
Tintinnodea, 231
Tintinnoinea, 231
Tipulocystis, 215
Toads, 125
Toadstools, 151
Tokophrya, 235
Tokophrya Lemnarum, 234
Tolypothrix, 35, 36
Torula, 130
Torulopsis, 130
Toxonema, 222
Tracheliidae, 230
Trachelina, 230
Trachelius, 230
Trachelomonas, 94, 106, 107
Transchelia, 143
Tremella, 149
Tremella Auricula, 146
Tremellacea, 149
Tremellaceae, 149
Tremellales, 149
Tremellina, 146, 149, 150
Tremellineae, 146, 149
Tremellinei, 149
Tremellini, 149
Tremellodendron, 149
Trepomonadida, 165, 166
Trepomonadidae, 166
Trepomonas, 166
Treponema, 29
Treponema macrodentium, 29
Treponema microdentium, 29
Treponema pallidum, 28, 29
Treponema pertenue, 29
Treponematacea, 29
Treponemataceae, 29
Tretomphalus, 180
Triactinomyxon, 222
Triactinomyxidae, 222
Tribonema, 65, 66, 73, 95
Tribonema bombycina, 64
Tribonematacea, 65, 66
Tribonemataceae, 66
Triceratium, 74
Tricercomitus, 167
Tricercomitus Termopsidis, 164
Trichamoeba, 202
Trichia, 176, 177
Trichiacea, 174, 176
Trichiaceae, 171, 176
Trichiales, 171, 174
Trichiidae, 177
Trichina, 177
Trichinaceae, 1 76
Trichoblasteae, 51
Trichocystiferes, Protistes, 94, 95
Trichodina, 235
Trichodinidae, 235
Trichomitus, 166
Trichomonadida, 166, 167
Trichomonadidae, 166, 167
Trichomonadina, 158, 164, 166
Trichomonads, 165
Trichomonas, 166, 167
Trichomonas hominis, 165
Trichomonas tenax, 164, 167
Index
[301
Trichomonas Termopsidis, 168
Trichomonas vaginalis, 167
Trichonympha, 168, 169, 170
Trichonympha Campanula, 168, 170
Trichonympha sphaerica, l68
Trichonymphida, 169
Trichonymphidae, 169
Trichonymphidea, 168
Trichonymphina, 168
Trichophyton, 142
Trichospermi, 152, 171
Trichostomata, 229
Tridictyopus elegans, 196
Trigonomonas, 166
Triioculina, 184, 185
Trimastigaceae, 58
Trimastigida, 58, 165
Trimastigidae, 58
Trimastix, 58
Trinema, 191
Triplagia, 198
Triposolenia, 103
Triposolenia Ambulatrix, 104
Tripylaria, 199
Tripylea, l99
Tripyleen, 199
Tripylina, 199
Triticina, 188
Trochammina, 186
Trochamminida, 186
Trochamminidae, 186
Trochamminina, 186
Truffles, 135
Tryblidacea, 134
Tryblidaceae, 134
Tryblidieae, l33
Trypanophidae, 161
Trypanophis, 161
Trypanoplasma, 161
Trypanoplasmida, 159, 161
Trypanoplasmidae, 161
Trypanosoma, 162
Trypanosoma Brucii, 160, 162
Trypanosoma Cruzi, 162
Trypanosoma equinum, 162
Trypanosoma equiperdum, 162
Trypanosoma Evansi, 162
Trypanosoma gambiense, 162
Trypanosoma Lewisi, 160
Trypanosomata, 158
Trypanosomatidae, 161
Trypanosomes, 161, 212
Trypanosomidae, 161
Trypanosomidea, 158
Tuberacea, 135
Tuberaceae, 134
Tuberales, 134
Tuberculariaceae, 141
Tuberculariea, l4l
Tubercularieae, 141
Tubercularini, 141
Tuberineae, 134
Tubifer, 175
Tubiferaceae, 175
Tubiferida, 174, 175
Tubiferidae, 175
Tubinella, 185
Tubulina, 175
Tubulinaceae, 175
Tubulinidae, 175
Tuburcinia, 149
Tulasnella, 149, 150
Tulasnella sphaerospora,
Tulasnellales, 149
Tulostoma, 155
Tulostomataceae, 155
Tulostomea, 155
Tulostomei, 155
Tunicates, 216
Turillina, 188
Turkeys, 210
Turtles, 211
Tuscarilla, 200
Tuscarora, 200
Tuscarorida, 200
Ulvina aceti, 24
Umbina aceti, 24
Uncinula, 132, 133
Uniflagellatae, 110
Urceolaria, 235
Urceolaridae, 235
Urceolarina, 235
Urceolus, 109
Uredinacea, 148
Uredinaceae, 148
Uredinales, 145, 147
Uredineae, 147
Uredinees, 147
Uredo, 147
Uredo linearis, 147
Urnula, 235
Urocentridae, 230
Urocentrina, 230
Urocentrum, 230
Uroglena, 59
Uroglenopsis, 59
Uroieptus, 233
Uromyces, 143
Urophagus, 166
Urophlyctis, 117
Urospora, 216
Urosporida, 216
Urosporidae, 216
Urosporidium, 218
Urostyla, 233
Urostylida, 233
Urostylidae, 233
Ustilaginacea, 149
Ustilaginaceae, 149
Ustilaginales, 149
Ustilaginea, 146, 149
Ustilagineae, 149
145
302]
The Classification of Lower Organisms
Ustilago, 149
Ustilago Heufleri, 145
Ustilago Hordei, 145
Uterini, 134, 137
Uvella, 59
Uvellina, 188
Uvigerina, 188
Uvigerinlda, 188
Uvigerinidae, 188
Vacuolaria, 65, 109
Vacuolaria viridis, 108
Vacuolariaceae, 109
Vaginicola, 233
Vaginifera, 233
Vaginulina, 187
Vahlkampfia, 202, 203
Valsa, 139
Valvulina, 186
Valvulinidae, 186
Vampyrella, 118, 189, 191, 192
Vampyrellacea, 191
Vampyrellaceae, 191
Vampyrelleae, 191
Vampyrellidae, 191
Vampyrellidea, 190
Vaucheria, 67, 76
Vaucheria Gardner!, 64
Vaucheria sessilis, 64
Vaucheriacea, 57, 63, 64
Vaucheriaceae, 63, 67
Vaucheriales, 63
Vaucherioideae, 55
Venturia, 139
Venturia inaequalis, 139
Verbeekina, 188
Vermes, 9
Verneulina, 186
Verneulinidae, 186
Veronica, 69
Verrucariacea, 139
Vertebralina, 184, 185
Vertebrates, 161, 165, 166, 167, 210, 211
Vibrio, 23
Vibrio Protheus, 201
Virgulina, 188
Volvox Chaos, 201
Vorticclla, 223, 226, 233, 235
Vorticellidae, 233
Vorticellina, 233
Vorticialcs, 179
Vorticialis, 186, 187
Vulvulina, 186
Wagnerella, 193, 194
Wardia, 221
Water molds, 77
Whales, 71
Wheat, 148
Wood roach, 166, 169
Worms, 215, 217, 220
Worms, annelid, 216, 219, 221
Worms, oligochaet, 222
Worms, polychaet, 211
Worms, siphunculid, 210
Woronina, 179
Woroninaceae, 179
Woroninidae, 179
Wrangelieae, 47
Xanthomonadina, 63
Xanthomonas, 23
Xenococcus, 36
Xiphacantha, 197
Xylaria, 139
Zanardinia, 88
Zea Mays, 6
Zelleriella, 229
Zonaria, 87
Zooflagellata, 157
Zoomastigina, 157
Zoomastigoda, 157, 178
Zoomastigophorea, 157
Zoopagacea, 123, 124
Zoopagaceae, 124
Zoopagales, 121
Zoopage, 124
Zoophagus, 81
Zoosporidae, 191
Zoosporidea, 191
Zoosporidia, 190
Zoothamnium, 233
Zooxanthellae, 194
Zostera, 203
Zschokkella, 221
Zygochytrium, 118
Zygocystis, 216
Zygocystida, 216
Zygocystidae, 216
Zygomyceteae, 121
Zygomyceten, 121
Zygomycetes, 76, 1 18, 120, 121, 122, 127,
141
Zygophyceac, 53
Zygophyta, 53
Zygorhynchus, 124
Zygostephanus, 198
Zythiacea, 141
Zythiaceac, 141
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