The Classification of Lower Organisms Ernst Hkinrich Haickei, in 1874 From Rolschc (1906). By permission of Macrae Smith Company. C f3 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 I. II. III. Page Introduction An Essay on Nomenclature 1 6 Kingdom Mychota Phylum Archezoa Class 1. 12 17 Schizophyta 18 Order 1. Schizosporea Order 2. Actinomycetalea Order 3. Caulobacterialea Class 2. Myxoschizomycetes Order 1. Myxobactralea Order 2. Spirochaetalea Class 3. Archiplastidea Order 1. Order 2. Order 3. Order 4. IV. V. Rhodobacteria 31 Sphaerotilalea 33 33 Coccogonea Gloiophycea Kingdom Protoctista 33 37 Phylum Rhodophyta Class 1. 40 Bangialea 41 Order Bangiacea 41 Heterocarpea Order 1. Cryptospermea Order 2. Sphaerococcoidea Class 4. Furccllariea 44 47 47 49 50 5. Coeloblastea 51 6. Floridea 51 2. Order Order Order Order VI. 18 24 25 27 27 28 29 3. Gelidialea Phylum Phaeophyta Class 1. Order Order Order Order Order Class 2. 1. Ochromonadalea 53 55 57 2. Silicoflagellata 61 3. Vaucheriacea 63 4. Choanoflagellata 67 5. Hyphochytrialea 69 69 73 74 76 77 80 Heterokonta Bacillariacea Order 1. Disciformia Order 2. Diatomea Class 3. Oomycetes Order 1. Saprolegnina Order 2. Peronosporina Order 3. Lagenidialea Class 4. Melanophycea Order 1 Phaeozoosporea Order 2. Sphacelarialea Order 3. Dictyotea Order 4. Sporochnoidea 81 82 86 86 86 87 . V ly Chapter Page Orders. Cutlerialea Order 6. Laminariea Order 7. Fucoidea VII. 88 89 91 Phylum Pyrrhophyta 1. Cryptomonadalea 2. Adiniferidea 3. Cystoflagellata 94 95 96 98 99 4. Cilioflagellata 102 Class Mastigophora Order Order Order Order Order VIII. Astoma Phylum Opisthokonta 105 5. 110 Class Archimycetes Order Order IX. Monoblepharidalea Ill 2. Chytridinea 113 Phylum Inophyta 119 Zygomycetes Order 1. Mucorina Order 2. Entomophthorinea Class 2. Ascomycetes Order 1. Endomycetalea Order 2. Mucedines Order 3. Perisporiacea Order 4. Phacidialea Order 5. Cupulata Order 6. Exoascalea Order 7. Sclerocarpa Order 8. Laboulbenialea 121 3. Hyphomycetes Order 1. Phomatalea Order 2. Melanconialea Order 3. Nematothecia 140 Class 1. Class Class X. Ill 1. 4. .... 121 124 125 129 130 131 133 134 137 137 140 141 141 141 142 Basidiomycetes Order 1. Protobasidiomycetes Order 2. Hypodermia Order 3. Ustilaginea Order 4. Tremcllina Order 5. Dacryomycetalea Order 6. Fungi Order 7. Dermatocarpa Phylum Protoplasta Class 1. Zoomastigoda Order 1. Rhizoflagellata Order 2. Polymastigida Order 3. Trichomonadina Order 4. Hypcrmastiglna Class 2. Mycetozoa Order 1. Enteridiea Order 2. Exosporea vi 146 147 149 149 150 150 152 157 157 158 163 166 168 171 171 177 Chapter Page Order 3. Phytomyxida Class 3. Rhizopoda Order 1. Monosomatia Order 2. Miliolidea Order 3. Foraminifera Order 4. Globigerinidea Order 5. Nummulidnidea . Class 4. Heliozoa Order 1. Order 2. Order 3. Order 4. Orders. Class XI. 5. 1. XII. 2. 1. Class 2. . 185 187 188 Acantharia 195 Monopylaria 198 Phaeosphaeria Sarkodina 198 200 201 Sporozoa Neosporidia 205 206 207 209 211 211 214 215 215 216 218 219 219 221 222 223 228 228 229 230 233 233 235 235 237 238 Infusoria Order 1. Order 2. Order 3. Order 4. Order 5. 185 . 190 194 Order 1. Phaenocystes Order 2. Actinomyxida Order 3. Cryptocystes Phylum Ciliophora Class 183 Radiolaria Order 1. Oligosporea Order 2. Polysporea Order 3. Gymnosporidiida Order 4. Dolichocystida Orders. Schizogregarinida Order 6. Monocystidea Order 7. Polycystidea Order 8. Haplosporidiidea Class 179 189 Radioflagellata Order 1. Nuda Order 2. Lampramoebae Phylum Fungilli Class 177 Opalinalea Holotricha Heterotricha Hypotricha Stomatoda Tentaculifera Order Suctoria List of Nomenclatural Novelties Bibliography Index 271 VII LIST OF ILLUSTRATIONS Portrait of Ernst Heinrich Haeckel Frontispiece Figure Page 1. Structure of cells of blue-green algae 2. Photographs of Escherichia coli Caulobacterialea; Myxobactralea; Cristispira Veneris Coccogonea; Gloiophycea Bangialea Nuclear phenomena in Polysiphonia violacea Heterocarpea 3. 4. 5. 6. 7. 8. 9. 13 . Ochromonadalea Ochromonadalea; 10. Vaucheriacea 11. Choanoflagellata 12. Hyphochytrialea 13. Bacillariacea 14. Oomycetes . 15 . 26 32 42 45 48 54 56 64 68 70 72 78 84 90 92 97 104 106 108 114 116 122 Silicoflagellata 17. nuclear division in Stypocaulon Familiar kelps of Pacific North America Microscopic reproductive structures of Laminaria yezoensis 18. Cryptomonadalea 15. Stages of 16. 19. Cystoflagellata; Cilioflagellata 21. Astoma Astoma 22. Monoblepharidalea 23. Chytridinea 24. Zygomycetes Ascomycetes Ascomycetes Mycosphaerella personata 20. 25. 26. 27. ... 132 136 138 144 Basidiomycetes 29. Fruits of Agaricacea 153 30. Rhizoflagellata 160 28. 33. Trichomonadina Hypermastigina Mycetozoa 34. Ceratiomyxafruticulosa 31. Polymastigida; 164 32. 170 176 178 "Tretomphalus" Rhizopoda 35. Life cycle of 36. Shells of i. e., Discorbis or Cymbalo por a 39. Monopylaria; Phaeosphaeria Chaos Protheus 40. Sarkodina 41. Life cycle of Goussia 42. 43. cycle of 44. Infusoria, order 45. Schuhergi Plasmodium; Babesia bigemina Life cycle of Myxoceros Blennius LUe . . 180 184 37. Radioflagellata 38. Radiolaria; Acantharia; . Hypotricha Tokophrya Lemnarum ix 192 196 200 204 208 212 220 232 234 Chapter I INTRODUCTION The purpose of this work is to persuade the community of biologists that the accepted 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 and that the organisms excluded from them are to be organized as two other 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 limits, kingdoms. 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 organisms which may be called the taxowhich men have constructed it is to serve as an index to all that is known about organisms. This system is subject to certain 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 have developed a nomic system. Its classification of function —the purpose — for and Filicineae are classes, we are expressing a fact of human convenThe names assigned to groups are obviously conventional. Since the taxonomic system represents knowledge, and since knowledge is advancing, 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- say that Pisces ience, not a fact of nature. tions, a better representation of taxonomic system are related the natural system. Different presentations of the 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 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- the natural system is 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 kingdom from animals, it is because I believe that everyone will have a better understanding 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 reduction. 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 requiring perhaps hundreds of millions of years. The changes are conceived as accidents; 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 necessary 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 the fact that anaerobic energesis is in a sense the basic metabolic 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 process. original organisms 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 selection, brought photosynthesis into existence. The purple bacteria are believed to represent 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 the bibliography. listed in Introduction [ 3 was established in certain organisms, aerobic energesis became possible both 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 bluegreen 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 membrane 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 nucleus 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 complicated heredity. Beside mitosis, there are two other processes two only meiosis and karyogamy, by which nuclei may produce other normal and enduringly viable nuclei. synthesis to these — — — — — — In a sequence of generations of individuals sexually produced, these processes occur 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 chromosomes) 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 realternately, each lated to sexual reproduction, including The did: it Mendelian heredity, as structure to function. existence of organisms without nuclei shows that the nucleus evolved after life did not evolve at the same time as protoplasm. the nucleus and of its The essential uniformity of 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 accordingly 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 has made possible the precise transmission of a complicated heredity, the nucleus 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 many characters and gave rise to many it entered upon evolution in distinguishable groups. Among these groups, those which consist respectively of the typical plants and the typical animals are the however, neither any a priori reason, nor any evidence from nature, 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 presented. In his teaching, he treated the bacteria and blue-green algae as standing entirely 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 greatest. There is, for a belief that all groups of nucleate organisms 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). when the teaching of elementary botany was first fully my own came to the conclusion that the establishment of several kingdoms In the year 1926, responsibility, I of nucleate organisms in addition to plants and animals these organisms are to be treated as one kingdom. This is is not feasible; that all of 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 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" substitutes, 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 embryonic 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 of the kingdoms. Of kingdoms without exhave 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 course, the definitions are not warranted to describe the ception. For one thing, each is supposed to algae. Kingdom II. Protoctista. Nucleate organisms not of the characters of plants and animals; the protozoa, the red and Kingdom brown algae, and the fungi. 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 III. Plantae. Organisms in whose cells 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 Protoctista until work systems of their subordinate groups are available. consists of such systems. Complete systems The bulk of the present 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 known groups than in the larger or more obscure. 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 better The 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 attempts 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 are here applied to the groups. The is principles according to are set forth in the following chapter. I beg my colleagues, that of the which this in dealing names which has been done 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 a system of groups finds himself under the necessity of making The kingdoms have received more names than one and so have nearly all of the major groups within them: it has here been sets forth responsible decisions as to names. (Table 1 ), necessary to decide as to the validity and application of the names Flagellata Mastigophora, Rhodophyceae and Florideae, Rhizopoda and Sarcodina, and others. TABLE 1. Names Applied by Various Authors to the Kingdoms OF Systems of Four Kingdoms Authors Kingdoms and many An The Essay on Nomenclature [ 7 most part a condensation of an earlier edition 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 bacteriological code is for the of the botanical code. It includes the comprenne dispose a et soit les odd feature accepter." 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 It is the truth. of By common pret it, this law, statement, the legitimate powers of congresses are those of courts which avoid the and apply it. explicit making of law, but discover the law, Congresses and codes may legitimately (a) inter- 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. when One would not in theory deny a power (c) to validate breaches of principle on necessity; but its use by botanical con- these are of an expedience verging produce a roll of exceptions of twice the bulk of the text of the code leads doubt the expedience of this admission. It has been through failure to recognize 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 diffidence, this assumption is extended to the bacteria; it will be agreed that the nomenclatural practice applied to the bacteria must be the same as that which is applied gresses to one to — — 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-designations, 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 work (1735), Linnaeus took them first sig- 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 followed by one epithet, ordinarily an adjective, occasionally a noun in apposition or nificant in the genitive. for granted. 8 The ] A fifth Classification of Lower Organisms principle represents Linnaean practices as subsequently modified: Named taxonomic groups are necessarily of certain fixed ranks called categories, 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 kingdoms 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 fam5. i.e., lists. ilies is a separate category. Authority for the list of seven principal categories as given Agassiz (1857). Nothing prevents the assignment of groups to categories other than these, to suband the like. These may be called subordinate categories. The groups of any subordinate category embrace only fragments of the range of kinds of classes, tribes, 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 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 asis sociates (1843) When : 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 necessarily 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. 7. a group An Essay on Nomenclature [ 9 A type as specified in the original publication of a group, or as implied by the inis unchangeable. Linnaeus and his immediate successors had no conception of the device of types, and it is practically impossible clusion of a single subordinate group, 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 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, caution, declare types arbitrarily. 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 : organisms among which the selection of a standard would be purely arbitrary. Such names are called nomina confusa, and are to be abandoned. to altogether miscellaneous collections of from the principle of the binomial nomenclature of species that no genus one or more of its species are designated by binomial names. It follows 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 regulate 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 barbarisms 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. It follows is named until names in the nominative, in ap1 names of persons, places, or organisms in the 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 Specific epithets are capitalized if they are ( ) position with the generic names; (2) genitive; (3) adjectives derived from — originally published Names —has priority from the date of its original publication. 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 signifying "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 Coniferinae, applied by Engler to a class, is an adjective with an additional adjectival termiof groups higher than genera are in the plural. : nation superimposed. — 10 The ] Classification of Lower Organisms A name once applied in any principal category may not be transferred to another, 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. unless it 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 by the fact that groups in these catewhich they occur; one is not in reason responsible for being aware of their names in groups outside of ones own categories the full sanction of priority, is justified gories are of concern only to specialists in the groups in specialty. had names with the uniform ending -aceae from 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 expedient 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 Almost all families of plants have the point of time at — code designations rather than names. In attempting 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 termitheir designations be of the nature of nations for of priority, The in use neuter names is of phyla, classes, something of an terminations of ordinal and orders, beside involving wholesale violation insult to the intelligence. names in -ales and of family names in -aceae, currently among the Mychota, are here changed to -alea and -acea to name of the kingdom. A change of the gender of an adjective agree with the does not create 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 termination -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 a as orders. Such names do not supersede older ordinal names not based on names of genera. Mychota is formed of the stem of a generic name (not 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 The name of a family of necessarily a valid work. Accordingly: Each group of Protoctista is called by its oldest name of tenable form in the correct category, barring any previously used in other principal categories, irrespective An of termination. All names which Essay on Nomenclature [ 1 are adjectives are used in the neuter, but ascribed to the original authors. The practices described have resulted in the use of strange, producing lists many names which which are undeniably heterogeneous. A will seem friendly critic notes an example of these things the Hst of classes, Heterokonta, Bacillariacea, Oomyand 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. as cetes, 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 ( not sec( 1877 ) 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, ; : tion Protophyta nor cohors 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 chlorophyll together with other pigments which bacteria, but those make which contain 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 may is of the order of 0.02^ (Knasyi, 1944). The 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 walls contain cellulose, but consist chiefly of pectates, 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 blue-green algae (Gardner, 1906; Swellengrebel, 1910; Haupt, 1923) 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 discells of are divided into outer from that of higher organisms has been extracted (Gardner; Kylin, 1943). The some of which stain like chromatin, while others ("red granules of Biitschli") are stained red by methylene blue. Cell division tinct inner part contains rods and granules, is by constriction. Olive (1904) interpreted the inner part of the continually in process of mitosis, and accordingly without a nucleus It is true that one may recognize series of corresponding granules. and granules of the interior are divided random. Haupt expressed the impropriety of calling any part of these cells a in series of disk-shaped cells Where at cell as a membrane. the cells are nucleus. more elongate, the rods Kingdom Mychota [13 Recent studies of typical bacteria by conventional microtechnical methods (Robinow, 1942, 1949; Tulasne and Vendrely, 1947) and by the electron microscope (Hillier, Mudd, and Smith, 1949) have made it possible to recognize the essential identity The protoplast conouter part, considered as a substance, may be of the structure of their cells with those of the blue-green algae. of outer sists and inner parts. The called ectoplasm (Knasyi, 1930), the central body (Biitschli, 1890). than one fifth of the radius of the inner, considered as a body, may be called The ectoplasm is very thin, occupying usually less cell. The spiral bands which have often been seen and the % Bi^i Z# )ii-i'Ss;.- "ji;-^ Irx ••;i©\ ^-> 'l-i" V^ .?.. fw ^vWfe Fig. 1.- — Structure Gardner (1906). a slide .^,.'^» ; 4m of cells of blue-green algae, prepared by Dr. P. Maheshwari, x 1,000. Haupt (1923) x a, Symploca Muscorum after C, Lyngbya sp. from Anabacna circinnalis after b, Oscillatoria Princeps after Olive (1904). d, 2,000. and which Swellengrebel ( 1906) mistook for a nucleus, are thickenings 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 in cells of bacteria, 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 and may redivide, De Lamater and Hunter usually to divide by constriction before the cell begins to divide, so that the cell may contain two dumb-bell shaped bodies. (1951) succeeded in a partial de-staining of the dumb-bell shaped bodies and interpreted 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 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 Their positions and lengths were by the electron microscope, is of the order of 0.02 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 flagellum at one end; lophotrichous, with a tuft of flagella at one end; peritrichous, with bell [J.. 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 matter 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 uncertain. The appearance of the movement is as though it were caused by local secretion of substances affecting surface tension. by constriction of the cells, each into two 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. Subsequently 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- The normal reproduction equal daughter turies, cells; of whence Mychota is the various names doubtfully of millenia. Lohnis and Smith (1916, 1923) observed of Azotobactcr that numbers of protoplasts 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 authorities. 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 derived 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.^ — Photographs Robinow, reproduced by show the ectoplasm, in which there are thickenings which tend to be spiral; right, stained to show the large repeatedly dividing granule in the central body. About x 2,000. By courtesy of Dr. Robinow and of the Society of .\merican Bacteriologists. Fig. 2. Hillier, of Escherichia coli by Dr. C. F. Mudd, and Smith (1949); left, stained to Kingdom Mychota The metabolic ficial list [17 Mychota are remarkably diverse. The most superwould include the following: (a) anaerobic parasites facultatively aerobic parasites and saprophytes; (c) the vinegar systems of the of physiological types and saprophytes; (b) known organisms which, while bacteria, being apparently the only 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 and certain other formations Archeozoic rocks estimated as more are ancient. Iron deposits believed to have been produced by them occur in 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- Manual, 6th ed., 1948). than one thousand. gey's The species of blue-green algae are probably fewer The classification of this group is inescapably highly tentative. The morphology 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 is Mychota makes it probable that many groups which appear to belong 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 antiquity of the or phyla; but it is not expedient to make many would produce too many divisions of The kingdom is accordingly treated as a this divisions of a group of 2500 species: a single class or classes of a single order. single phylum, and its main divisions as classes. Phylum ARCHEZOA Haeckel 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 yhylB. life as animal. The conventional division of the group into two classes, bacteria and blue-green 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. algae, 1. is Cells without internal pigment, heterotrophic or living by chemosynthesis; not usually pro- ducing filaments with prominent sheaths. 18 The ] 2. Cells with firm Classification of walls, Lower Organisms non-motile or motile by means of flagella 2. means of changes of shape, also 1. Schizophyta. Class 2. Myxoschizomycetes. Class 3. some- times by flagella 1. Class Cells with thin walls or none, motile by Cells mostly with internal pigment, living by photosynthesis or chemosynthesis, exceptionally heterotrophic; often producing filaments with prominent sheaths Class 1. SCHIZOPHYTA (Cohn) Archiplastidea. 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 1. into cells 1. Order 1. Schizosporea. Order 2. Actinomycetalea. Order 3. Caulobacterialea. Consisting of branched filaments not divided Cells attached by stalks, free ends differentiated the attached and Schizosporea [Schizosporeae] Cohn in Hedwigia 11: 17 (1872). Schizomycetes (Nageli) McNab in Jour, of Bot. 15: 340 (1877). Eubacteria Schroter 1886. Haplobacteriacei Fischer in Jahrb. wiss. Bot. 27: 139 (1895). Orders Cephalotrichinae and Peritrichinae Orla-Jensen in Centralbl. Bkt. Abt. Order 1. Order Order Order 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 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. small 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 and the peculiar character called Gram reaction. Gram, 1884, consists of staining successively with gentian violet and iodine. It gives an intense blue-black color. From some bacteria, 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 of endospores; metabolic products; The method of staining invented by is believed to be lipoid, such as occurs in all cells. The Gram positive believed to consist in a relatively low isoelectric point, a capacity, that is, plus iodine quality to is combine with anions in a relatively acid medium. This quality lies in the ectoplasm of the cells and disappears in aging cultures. The is Manual (1923, 1925, 1930, 1934, 1939, 1948) among Americans) as standard. The following system of thirteen classification given in Bergey's accepted (at least families is a moderate rearrangement of the Bergeyan system, with certain ideas or and Stanier ( 1925), Pribram (1929) names from Enderlein (1917, 1925), Buchanan andvanNiel (1941). 1. Gram positive, which are with exceptions many of intracellular parasites; atrichous or peritrichous. 2. Spheres dividing in more planes than one. 3. 3. 2. Gram Gram positive gens in animals Rods, or spheres dividing in one plane. 3. Gram 2. Micrococcacea. Family 2. Neisseriacea. Atrichous. 5. Not 5. Intracellular parasites intracellular parasites. 4. Peritrichous 1. 1. Not producing endospores. 4. 3. Family negative; intracellular patho- . Family 3. Corynebacteriacea. Family 4. Rickettsiacea. Family 5. Kurthiacea Family 6. Bacillacea. . Producing endospores negative. Atrichous or peritrichous, requiring comparatively complicated organic food. 3. Not plant pathogens. 4. Not fixing nitrogen. Capable of growth on ordinary media Family 7. Achromobacteriacea. 5. Requiring special media; minute atrichous pathogens. Family 8. Pasteurellacea. 4. Fixing nitrogen Family 10. Azotobacteriacea. Plant pathogens Family 9. Rhizobiacea. 5. 3. 2. Atrichous, monotrichous, or lophotrichous; the atrichous representatives, many and can survive with organic foods simpler than carbohydrates, or with none. 3. Mostly requiring at least carboothers, hydrates Family 11. Spirillacea. : 20 The ] 3. Not requiring carbohydrates. 4. Oxidizing acid, 4. and 1. alcohol to acetic acetic acid to and H2O Not as above; strictly Family Lower Organisms Classification of CO2 Family 12. Acetobacteriacea. Family 13. Nitrobagteriacea. many examples autotrophic 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^. ]Veichselbaumii 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 and serious pathogens causing, for example, and pneumonia. Diplococcus, spheres usually in pairs, encapsulated. D. pneumoniae occurs in many immunologically distinct races which are the usual causes of pneumonia. butter and cheese; and commensals abscesses, septicemia, erysipelas, Lactobacillus, rods, microaerophilic, producing lactic acid. In milk, involved in making and cheese; in the oral cavity, being the usual agent of dental 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 the of butter caries (Rosebury, Linton, 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 intracellular 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 satisthem is not yet possible. Howard Taylor Ricketts showed that Rocky Mountain spotted fever is transmitted by the tick Dcrmocentor, and factory classification of observed, in the cells of diseased tissues, minute irregularly staining bodies; in 1910, Kingdo7n Mychota [21 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, in the course of further studies of the disease, 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. construed Bartonella as a sporozoan. Good authority has Students of flagellates, Sarkodina, and Infusoria have occasionally observed in the cytoplasm or nuclei of these organisms minute bodies multiplying to form considerable masses. These parasites have generally been construed as chytrids, but have little in common with proper chytrids. The genus Caryococcus Dangeard includes at them. Kurthiacea, fam. nov. least a part of Family 5. Gram positive peritrichous rods, not producing endospores. Kurthia, harmless; Listeria Pirie ex 408 (1948), pathogenic Family 6. BaciUacea in sheep Murray in Bergey's Manual 6th ed. and man. [Bacillacei] Fischer in Jahrb. wiss. Bot. 27: 139 (1895). 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 The which can be can be definitely applied because the type species 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. Exdefinitely applied: it B. subtilis was so described as to be recognizable. 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 PrazmowAs 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 ski. (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 capacity by Winogradsky forms of gangrene, occasion for the study and distinction of which was found during World War I. Achromobacteriacea [Achromobacteriaceae] Breed 1945. Family Bacin 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 Family teriaceae 7. McNab 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. standard method of testing waters for contamination presence of these organisms. Klebsiella also produces acid tract. The cells are and gas from is essentially a test for sugars. It inhabits heavily capsulated and non-motile. The The the the respiratory type species K. pneumo- 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. Immunological study of cultures of Salmonella from cases of disease and from waters have resulted 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, niae is S. dystenteriae, is the cause of dystentery. numerous group of acid-producing gut bacteria, motile or nonfrom the foregoing as strictly anaerobic. Alcaligenes fecalis, an apparently harmless organism isolated from intestinal contents, does not produce acid from sugars; grown in milk, it produces an alkaline Bacteroides is a motile, generally harmless.^ distinguished reaction. and are capable which produce an extracellular red pigment are Serratia (one of the oldest generic names for bacteria); those which produce yellow pigment are Flavobacterium; those which produce blue, black, or violet growths are Chromobacterium. Cultures which lack the distinc- Numerous races of bacteria which have been isolated from soil of attacking cellulose are assigned to the genus Cellulomonas. Bacteria above named genera (most such cultures have been 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. Pfeiftive characters of all of the isolated ferella mallei is the cause of glanders. 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. Family 9. Rhizobiacea [Rhizobiaceae] Conn Erwinia commemorates Erwin F. Smith, the discoverer of many bacteria pathogenic 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 to plants. 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 Agrobacterium 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 Radicicola Beijerinck 1888) is that which attacks plants of the pea tribe. Bewley and Hutchinson (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 monotrichous 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 remaining families of the present order are more definitely distinct, being marked by monotrichous or lophotrichous flagella. Migula 1894. Family Pseudomonadaceae 555 (1917). Rods and spirals, Gram negative, monotrichous or lophotrichous; not producing much acetic acid, and mostly heterotrophic. Pseudomonas is a numerous genus of rods which may or may not produce a fluorescent pigment soluble in water; they do not produce a yellow pigment which is inFamily Winslow 11. Spirillacea [Spirillaceae] et al. in Jour. Bact. 2: soluble in water. The original species, P. aeruginosa, was isolated from pus, in which 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. it 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 intolerable 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 to acetic acid, or acetic acid, which processes are strictly aerobic: to it, one is make is alternatively alcohol, which is oxidized oxidized to carbon dioxide and water. These vinegar, one exposes wine to air; to preserve seals the vessels. Family 13. Nitrobacteriacea [Nitrobacteriaceae] Buchanan in Jour. Bact. 2: 349 (1917). Organisms oxidizing the simplest organic compounds; or facultatively capable of chemosynthesis; or living strictly by chemosynthesis and strictly aerobic: mostly Gram negative monotrichous or atrichous rods. is capable of oxidizing methane; Carboxidomonas of oxidizing carbon monoxide; Hydrogenomonas, of oxidizing elemental hydrogen. Thiobacillus Methanomonas 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 contained 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 and Nitrobacter, non-motile rods oxidizing nitrites to nitrates. Subsequent authors have validated Winogradsky's names by creating the combinations Nitrosococcus nitrosus and Nitrobacter VVinogradskyi. Subsequently, Winograd- as Nitrosomonas; sky discovered yet other bacteria capable of the The presence of nitrifying bacteria is same oxidations. 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 Order 2. in fallow fields. 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 The order may be treated as a single family. fast. Kingdom Mychota [ 25 Family Mycobacteriacea [Mycobacteriaceae] Chester 1907. Family ActinomyceBuchanan 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 Actinomyces. 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 penetrate 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 immunity 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. taceae 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 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. manner The 26] Classification of Lower Organisms m — Caulobacterialea after Henrici and Johnson (1935) x 2,000: a, Caulobacter vibrioides; c, Caulobacter sp.; d, Pasteuria sp.; e, Blastocaulis sp. f. Various stages of Cytophaga Hutchinsonii [Spirochaeta cytophaga) after Hutchinson and Clayton ( 1919). g-k, Myxobactralea after Thaxtcr (1892), the cells 3 Fig. a-e, Nevskia sp.; b, X 1,000, in the fruits x 200. des. I, m, Dividing and fruit of Chondromyccs crocatus; i, fruit and spores, and fruit, of Myxococcus coralloi g, h, Cells of C. aurantiacus; j,k, vegetative cells cells of Cristispira Veneris after Dobell (1911) x 2,000. Kingdom Mychota Family 1. [ Leptotrichacea [Leptotrichacei] Schroter 1886. The cells 27 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 believed 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 likewise 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, PasBlastocaulis, colorless saprophytes in waters or parasites in aquatic and teuria, animacules. Class 2. MYXOSCHIZOMYCETES Myxoschizomycetae Class Class Polyyangidae Jahn Schaffner Schaffner in Ohio Naturalist 9: 447 (1909). Beitr. bot. Protistol. 1: 65 (1924). Class Spirochaetae Stanier and van Niel in Jour. Bact. 42 459 ( 1941 ) Mychota, the elongate cells with thin walls or none, capable of bending movements and sluggishly or actively motile. In many examples : Parasitic or saprophytic there is a resting stage: the cell contracts generally, so as to diminish the surface, 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). and deposits 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 ] cells Classification of Lower Organisms 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 As a digesting fairly easy. and absorbing food as they proceed. Labratory culture is 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 Winogradsky [Spirochaeta cytophaga Hutchinson and Clayton) is one of several species discovered as active fermenters of cellulose. The slenderly spindle-shaped cells are and produce ellipsoid chlamydospores resembling yeasts. Family 2. Archangiacea [Archangiacae] Jahn op. cit. 66. Spores elongate in irregularly 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. sluggishly motile, 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 recurrentis (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 Schaudinn, 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 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 material, under more or less anaerobic conditions. Each species requires its peculiar variant of the conditions, to which it is quite sensitive. success Kingdom Mychota Spirochaeta plicatilis and other saprophytic [ 29 with certain species species, together 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 chromatin, 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 parasitic in mollusks, are fairly large. The species 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 Treponema pallidum. Reproduction is normally by transverse diviDuring 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 ends of cells of sion into two. sui generis. The larger Family 1. and smaller spirochaets are reasonably treated as separate families. Spirochaetacea [Spirochaetaceae] Swellengrebel 1907. The cells com- paratively large, 80-500(1 long. Spirochaeta, Saprospira, Cristispira. Family 2. Treponematacea [Treponemataceae] Robinson (1948). Family Treponemidae Schaudinn 1905. The cells in Bergey Man. 6th ed. 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. microdentium, 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, supposedly from cases of yellow fever, yellow fever fever that is caused by a virus. Noguchi lost his life Class 3. by is It perhaps the same thing: it is now known that in pursuing in Africa his study of yellow was this disease (Flexner, 1929; Eckstein, 1931). ARCHSPLASTIDEA Bessey Myxophykea Wallroth 1853. Myxophyceae Stizenberger 1860. Division (of Class Algen) Pkycochromaceae and order Krytog.-Fl. Sachsen 1: 56' (1863). Cyanophyceae Sachs Lehrb. Bot. ed. 4: 248 (1874). Gloiophyceae Rabenhorst OrAtx Cyanophyceae or Pkycochromaceae yicNdLhm]o\iT. Schizophyceae Kryptog.-Fl. Cohn 2, oi'Qot. 15: 340 (1877). 1879, not suborder Schizophyceae Rabenhorst Deutschland's Abt. 2: 16 (1847). — . 30 The ] Classificatio7i of Order Schizophyceae Schenck Lower Organisms 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). Myxophyceae Setchell and Gardner Subclass part 1: in Univ. California Publ. Bot. 8, 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) appears 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 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 bluegreen algae. Only the genus Sphaerotilus remains at loose ends. It is credibly reported and blue-green algae belong to produce A cells together. swimming by means variety of purple bacteria do this. which contain a red pigment of flagella; no proper blue-green algae —bacteria, that is, 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 photosynthesis; 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 and ZoBell, 1951; Williams and Burris, 1952). Four orders may be distinguished: 1. fixing nitrogen (Sisler Possessing a red ("purple") intracellular pigment, or a green pigment not masked by others l.With green pigment masked by Order 1 Order 2. . Rhodobacteria. others, or colorless. 2. Producing cells with flagella; non-pig- mented sheathed filaments not accumulating ferrugineous matter Sphaerottlalea. Kingdom Mychota 2. Never producing 3. cells Cells dividing in with [ 31 flagella. more planes than one, growing to full size before redividing; unicellular or colonial, not filamentous 3. Cells dividing in one plane, accordingly producing Order 3. Coccogonea. Order 4. Gloiophycea. and filaments; exceptional examples reproducing by budding (unequal division) or by repeated division into minute spores 1. Rhodobaeteria Molisch Purourbakterien 27 (1907). Rods, spheres, and spirals, solitary or colonial, with red or green pigment, performing 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, Thiorhodaceae, aerobic, accumulating granules of sulfur, and Athiorhodaceae, microaerophilic 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 ChroMATiACEAE Migula. Family Rhodobacteriaceae Migula; Family Thiorhodaceae Molisch; the family does not include genera with corresponding names. Purple bac- Order accumulating granules of sulfur. Chromatium Perty includes the organism 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 Winoteria, aerobic, gradsky. Rhodobacillacea nom. nov. Family Athiorhodaceae Molisch. Molisch 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 Family named 2. in this family a — The 32] Fig. 4. Coccogonea: phycea: d, Oscillatoria a, Classification of Chroococcus splendida; e, sp.; b, C, m, Clonothrix and Gardner 1919); after Setchell X 1,000. 1, ( Achromatiuni oxalijerum. Phormidium sp.; f, Beggiatoa sp.; g, \, fusca after o, Gloio- Chamae- Cylidrospcrmum majus; j, ChlarnydoKolk (1938); n, Dermocarpa protea Crenothrix polyspora after Kolk 1938). All siphon incrustans; h, Anabaena inacqualis; thrix ochracea; k, Lower Organisms ( 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 corresponding 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 The cells may escape from the filaments, become lophotri- laterally out of the sheath. 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. chous, and function as Order 3. Coccogonea [Coccogoneae] (Thuret) Campbell Univ. Textb. Bot. 84 (1902). Tribe Chroococcaceac [Coccogoneae) Thuret in Ann. Sci. Nat. Bot. 1: 377 (1875). Subclass Coccogoneae Engler in Engler and Prantl Nat. Pfianzenfam. Abt. la: (1900). Order Coccogonales Atkinson 1903. Orders Chroococcales and Entophysalidales Geitler in Pascher et ser. 6, I Teil, iii al. Siisswasserfl. Deutschland 12: 52, 120 (1925). Cells solitary or colonial, not filamentous, never flagellate; mostly of blue-green 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 color parasites within the cells of the green algae Glaucocystis sulting bodies, having the color and Gloeochaete. The of blue-green algae with the structure re- of green algae, resisted classification until Geitler (1923) explained their nature. Family 2. Achromatiacea [Achromatiaceae] Buchanan. Cells solitary, large, ellipsoidal, 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 (1863). Tribe Nostochineae {Hormonogoneae) Thuret in Ann. Sci. Nat. Bot. 1: 377 (1875). 1 : ser. 56 6, 34 The ] Classification of Lower Organisms Family Hormogoneae Bomet and Flahault in Ann. Sci. Nat. Bot. ser. 7, 3 (1886). Subclass Hormogoneae Engler in Engler and Prantl Nat. Pflanzenfam. 337 : 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 the type called "false": it one plane, any branching of the filaments occurs by breaks in the continuity of the series of followed by the outgrowth, beside the original some members series, of the newly formed is of cells, tips. In of the group, however, the cells are not strictly confined to division in is possible. There are a few apparwhich 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 differentiated cells called heterocysts (the word means "different cells"). These are enlarged thick-walled cells with colorless contents; their most obvious function is to one plane, with the result that "true" branching ently derived examples in furnish breaking points for the filaments. They arthrospores; they have been seen to germinate Ten 1. families may be are believed to be variants of the and give rise to normal filaments. distinguished as follows: 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 organisms accumu4. Colorless lating sulfur Filaments reduced to single cells which reproduce by budding Filaments branching or tapering, or producing spores or heterocysts, or showing Family 1. Oscillatoriacea. Family 2. Beggiatoacea. Family 3. Chamaesiphonacea. Family 4. Nostocacea. Family 5. Scytonematacea. 3. 2. several of these characters. 3. Filaments not tapering. 4. Filaments not branching 4. Filaments branching. 5. Blue-green algae mostly producing heterocysts 5. Minute without heterocysts Filaments tapering Cells dividing in more planes than one, usually after a preliminary filamentous phase. 2. Pigmented, blue-green. 3. Producing extensive filaments with 3. 1. colorless filaments heterocysts Family 6. Chlamydotrichacea. Family 7. Rivulariacea. Family 8. Sirosiphonacea. Kingdom Mychota 3. Filaments more or less reduced, reproducing by minute spores (gonidia) formed by repeated division Family in all planes 2. [35 Colorless; filamentous, 9. Pleurocapsacea. reproducing by Family 10. Crenotrichacea. gonidia 1. Oscillatoriacea [Oscillatoriaceae] Harvey 1858. Blue-green algae consisting of unbranched filaments, not tapering, without spores or heterocysts; mostly actively motile by mechanisms as yet unknown. In the commonest genus, Oscillatoria, Family 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 filaments are coiled; those of Spirulina are not visibly septate, and are said to be unicellular. 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 was the natural one. Family 3. Chamaesiphonacea [Chamaesiphonaceae] Borzi 1882. Order Chamaesipkonales 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 position originally assigned to the species of Beggiatoa 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. 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 If in difference that heterocysts are usually present. filament may produce The multiplication of the cells of a 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 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. Chlamydothrix 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 filaments dividing repeatedly (always in one plane) to produce spherical non-motile laterally gonidia (Kolk, 1.938). Family Fdvulariacea [Rivulariaceae] Rabenhorst op. 7. 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 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. SiisswasserFl. Deutschland 12: 124 (1925). This group was formerly included in Chamaesi- become attenuate and 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 become 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. Crenotrichacea [Crenotrichaceae] Hansgirg. This family includes the Cohn, one of the traditional iron bacteria. There is every appearance that it is a colorless variant of the Pleurocapsacea. A germinating 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 released by its decay. Such are the Mychota, the organisms which may properly be characterized as Family single 10. known species Crenothrix polyspora lacking nuclei. Chapter IV KINGDOM PROTOCTISTA Kingdom PROTOCTISTA Hogg !l. 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 Kingdom Primalia Wilson and Cassin in Proc. Acad. 2: 6 (1861). Nat. Sci. Philadelphia 1863: 117 (1864). Kingdom Kingdom Morph. 2: xix (1866). 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. Protista Haeckel Gen. The name Protista, of Haeckel, is the most familiar among those which have been not the earliest. Among followers of Cuvier, the animal kingdom consisted necessarily of four branches. Presumapplied to the kingdom here to be discussed, but ably, it was this tradition that induced Owen it is to refer the Infusoria and Amorphozoa A year later, Owen puban alternative name for this kingdom; but Hogg had already published modifications 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 (sponges) to a separate kingdom, which he called Protozoa. lished author (1938, 1947). They have been accepted, with exception in a single significant and Rothmaler (1948). point, by Barkley (1939, 1949) It is assumed that the evolutionary origin of the Protoctista consisted of the evoluall nuclei are essentially the same thing. Kofoid tionary origin of the nucleus, and that (1923) insisted that enduringly viable nuclei originate among protozoa, as among and animals, regularly by mitosis, never by binary or multipe fragmentation, nor by aggregation of stainable granules. He did not recognize the nucleus as essentially a device for sexual reproduction. Several considerable groups of protozoa, howplants 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 nonnucleate and nucleate organisms, the nucleus is believed to have evolved in organisms living by this function. The closest approach between non-nucleate and nucleate organisms 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 (Besever, 38 The ] The Classification of Lower Organisms organisms are not homologous with those 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 chlorophylls 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 sey, 1905). flagella of nucleate bacteria; they are much larger and of earth. Organisms of the body type of solitary tinoid pigments but not chromoproteins, walled cells, having chlorophylls and caro- and producing flagellate reproductive cells, 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 occurring in nucleate organisms are here particularly to be noted. appear to Loeffler (1889), in the original publication of the standard method of staining the remarked that he had applied this method also to certain larger found that the flagellum of Manas bears numerous lateral appendages, flagella of bacteria, organisms. He and that the method cilia of a certain infusorian bear solitary terminal appendages. Loeffler's is difficult, and has not been much used. Fischer (1894) used it and coined and Peitschengeisseln, designating structures of the respective terms, Flimmergeisseln types seen by Loeffler. Petersen (1929), having applied Loeffler's method to a reasonable variety of flagellates, introduced refinements of terminology. Flagella of the type of the larger flagellum of Monas (the organism bears also a minute simple flagellum) 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 appears 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. ( The stichoneme flagellum bears a single file of appendages. The point in which Barkley and Rothmaler take exception to (4) kingdom the limits here given 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 pigments; 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 to Protoctista is this, that they include in this plant kingdom. Rothmaler up a system of only four phyla, being the red organisms, basically which are typically yellow to brown, having pantoneme flagella; 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. without set flagella; those 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 characters, 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 1. Phylum 1. Rhodophyta. Phylum 2. Phaeophyta. Phylum 3. Pyrrhophyta. Phylum 4. Opisthokonta. Phylum 5. Inophyta. Phylum 6. Protoplasta. Phylum 7. Fungilli. .Phylum 8. Ciuophora. Without chromoprotein pigments. 2. photosynthesis, living by brown, yellow, or green in color. 3. Producing flagellate cells each with one pantoneme or pantacroneme flagellum, often with additional Typically acroneme flagella Producing flagellate cells whose flagella are never pantoneme or pantacroneme, often stichoneme Dependent; motile cells with acroneme flagella or cilia, or amoeboid, or none. 3. Not producing cilia, i. e., structures 3. 2. acroneme flagella, numerous and widely distributed on the surfaces of the cells. of the nature of walled in the vegetative 4. Cells condition. 5. Producing motile cells with single posterior flagella; bodies mostly with tapering rhizoids 5. Producing no motile cells; bodies filamentous not walled in the vegeta- 4. Cells tive condition. 5. Mostly predatory, flagel- amoeboid or with flagellate or amoeboid stages late or 5. Parasitic in animals, pro- ducing flagellate cells as rare exceptions 3. With cilia only . V PHYLUM RHODOPHYTA Chapter 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). Heterocarpeae Kiitzing Phyc. Gen. 369 (1843). 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 Class Class Florideae 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 primitive 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- 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 thrin, are applied; they are not, (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 1. classes: 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 1. with Cells protoplasmic [41 interconnections, containing except in the lowest examples several parietal plastids, organized as filaments with apical growth, the filaments usually massed as thalloid bodies; zygotes giving rise Class to spores indirectly Class 1. BANGIALEA 2. Heterocarpea. (Engler) Wettstein Algarum 4: 4 (1897). Subclass Bangiales Engler in Engler and Prantl Nat. Pflanzenfam. Subclass Bangioideae de Toni Sylloge 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. Nebraska Studies 7: 291 (1907). Class Bangieae SchaflFner in Ohio Naturalist Protoflorideae Rosenvinge in Mem. 9: 448 (1909). 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 interconnections, 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, known species is about eighty. five families, about fifteen genera; the number of Order Bangiacea [Bangiaceae] Nageli 1847. Characters of the class. 1. Cells forming irregular aggregates 1. Cells forming filaments or thalli. 2. 2. Vegetative cells becoming spores without dividing Vegetative cells undergoing division to Family 1. Porphyridiacea. Family 2. Rhodochaetacea. produce spores. Family 3. Porphyrea. 3. Organisms red, purplish, etc Family 4. Schizogoniacea. 3. Organisms green Family 5. Compsopogonacea. 2. Spores formed solitary in special cells 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 distributed 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 The 42] — Classification of Lower Organisms x 1/2. b-g, Porphyra tenera after Ishikawa produce sperms; d, sperms; e, fertilization; 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. Fig. 5. a, Porphyra (1921); b, cells; f, "carpospores," c, laciniata, thallus cell dividing to 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 considrable 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 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. observations Algae 121 (1933). Branching filaments, sometimes becoming multiseriate by lengthcells capable of escaping and functioning as spores. Sexual reproduction unknown. Asterocystis, uncommon, in fresh water; the remaining genera marine, epiphytic on other algae. Goniotrichum. Rhodochaete and Goniotrichopsis, the cells with numerous plastids. Family 3. Porphyrea [Porphyreae] Kutzing (1843). Family Porphyraceae Rabenhorst 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 release their protoplasts as wholes or may undergo division into many. Rosenvinge wise division, the vegetative (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 fuscopurpurea, 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 centrosomes. 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 gaNaked 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 metes. 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 in which all cells is except the zygotes are haploid. a life cycle of the primitive type, 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 {Schizogonium 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 pluriseriate, 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 140[i per minute. Sexual reproduction Erythrocladia. Compsopogon, is much as in in fresh water, the cells Class 2. HETEROCARPEA move as far as Porphyrea. Erythrotrichia. with numerous parietal plastids. Kutzing Heterocarpeae Kiitzing Phyc. Gen. 369 (1843). v ( 1848). J. Agardh Sp. Alg. 1 Subclass Florideae Engler in Engler and Prantl Nat. Pflanzenfam. Class Class Florideae (C. Agardh) : 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 examples) 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- 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 filament, the carpogonial filament, and bears a filiform terminal extension, the trichogyne (Bornet and Thuret, 1867), whose function is to receive the sperms. The cell, often diflferentiatcd, from which the carpogonial filament grows, is the support- uals. ing cell [Trugzelle). In the more primitive members of the mass of cells called the cystocarp. as spores called carpospores. the original ones. first The The the zygote gives rise by budding to a cystocarp release their protoplasts These on germination produce haploid individuals zygote nucleus divisions arc meiotic. class, cells of the is the only diploid nucleus in the like life cycle; its Phylum Rhodophyta [45 In more advanced examples, the first step of development after fertilization conof the establishment of protoplasmic contact between the zygote and other cells. These may be adjacent cells, reached directly, or distant cells, reached by the outgrowth 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, sists The give rise to diploid individuals. diploid individuals are of the same vegetative 'ymW''^- :^^>^^ ^^y i^ •«!» —Nuclear phenomena Fig. 6 a, b, c. Stages of mitosis, in Polysiphonia violacea after d, e. Stages of homeotypic Yamanouchi (1906). division. structure as the haploid individuals, but do not produce spermatia, carpogonia, or cystocarps. Certain cells, sets of four spores commonly scattered which are accordingly called and imbedded in the body, produce 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, multiplicative phase of the diploid stage (Janet named this stage the carposporophyte; Drew, 1954); and free-living diploid individuals, reproducing by tetraspores. The production 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 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 much 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 Oltmanns (1898) disproved this: he showed that the nuclei of auxiliary 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. Experfertilization. cells are inert, 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 classificaEngler ( 1897) made these groups ( 1889) Subsequent scholars have found this system sound in principle, but necessary, on the basis of studies of additional examples (for example, tion of the typical red algae as four groups ; definitely orders. have found it by Kylin, 1923, 1924, 1925, 1928, 1932; Papenfuss, 1930, number The lished currently recognized to following key to the orders six. is a rather considerable modification of those pub- by Kylin (1932) and Smith (1944). l.All free-living individuals haploid; tetra- spores not produced, or produced as carpospores. 1. 1926; least four orders have been proposed but reductions have decreased in addition to those of Engler the 1944; Sjostedt, and genera. At Svedelius, 1942) radically to rearrange the families . Order 1. Cryptospermea. Order 2. Sphaerococcoidea. Order 3. Gelidialea. Order 4. Furcellariea. Order 5. Coeloblastea. Free-living individuals of two types, the one producing gametes (the zygotes giving rise to carpospores), the other producing tetraspores. 2. Without specialized auxiliary nurse cells, cells or the lower cells of the carpo- gonial filaments, or normal vegetative serving as auxiliary cells cells, 2. With specialized nurse cells, the carpo- spores produced from filaments which have made contact with these 2. With which 3. specialized auxiliary cells from the carpogonia develop. The auxiliary cells being intercalary cells in specialized filaments homol- ogous with the carpogonial filaments. 3. The auxiliary cells terminal in . . . fila- ments which grow from the supporting cells of ments before the carpogonial fertilization fila- Phylum Rhodophyta 3. The [ 47 auxiliary cells originating after fertilization as porting cells branches of the supthe carpogonial of Order filaments 6. FLORroEA. Order 1. Cryptospermea [Cryptospermeae] Kiitzing Phyc. Gen. 321 (1843). Order Periblasteae Kutzing op. cit. 387, in part. Orders 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. H 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 producing 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 The forms is mere called Nemalionales. Eight families are rec- Rhodochorton, and (Family Chantransiaceae 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. ognized. others, consisting of filaments, Acrochaetium, are family Acrochaetiacea [Acrochaetiaceae] Fritsch 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 classified as corals. They have certained their and tetrasporic stages. Svedelius (1942) asCarpospore-bearing filaments arise both from the zygote distinct sexual life cycle. 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 exhibiting 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. 75 (1926). f. 37, no. 4: 48] The Classification of Lower Organisms (Legend on bottom of page 49) Phylum Rhodophyta This order, in one. The all recent literature called Gigartinales, bodies are generally erect; they may be [ is a 49 numerous and varied unbranched cylindrical or flattened, 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 or branched. In cells with which contact is made are auxiliary produce carpospores, and the carpospores produce 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 established 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 carpogonial filament. In either case, the cells and give rise to cystocarps; these (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 Plocamium; Gracilariophila and its host Gracilaria (Wilson, 1910). Order 3. Gelidialea [Gelidiales] Kylin in Kgl. Svensk. Vetensk.-Akad. Handl. 132 (1923). 63, no. 11: 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 b, c, d^ production of sperms; a, Thallus of Nemalion multifidum x 1. 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 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 of carpospores. g, cell after fertilization. 50 The ] this family, This Classification of and particularly Lower Organisms in the genus Gelidium. It is the chief source of agar agar. the principal material of the cell walls of Gelidium. It is essentially of chains of galactose units, is a jelly consisting 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 laboratory 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 involving specialized nurse cells, appear less its reproductive processes, primitive than those of the Sphaerococ- 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. coidea. 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, Order Abt. 2: xi (1897). individuals are crustose or thallose, the thalli cylindrical or flattened, unbranched or branched. On the two or three types of individuals of each species, the The reproductive structures may be scattered or clustered on the surfaces or gathered in specialized pits called conceptacles. The cialized filaments; other filaments, homologous with these but abortive, bear the eggs are as usual the terminal cells of spe- auxiliary cells. After fertilization, the zygote with a lower cell of the same filaments. may Under or may not establish connection 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. ( 1932) placed nine families here. 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 Kylin The whose margrow 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, according 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, auxiliary cells, unite eventually in a single large multinucleate cell from gins belonging to The this order, attacks species of order Floridea. 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 Furcellariea are a 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 produce 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 Lamouroux included the whole group of red algae organized as 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 fertiliFloridees of four genera, one more cell adjacent to the zygote, and this becomes 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, zation, the supporting cell cuts off 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 though consisting many members of the other families the bodies are thallose, 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 Heterokontae, 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 together with larger quantities of other carbohydrates or oxidized or esterized carbo- hydrates. Silica or calcium carbonate may be deposited. Methanol extracts of the contain fucosterol, a sterol distinct from the sitosterol of typical plants. Flagellate cells are usually produced; these bear one pantoneme or pantacroneme flagellum, and usually, in addition, one acroneme or simple flagellum. Exceptional examples, cells non-pigmented or without flagellate stages, are rather ard genus of the phylum Fucus L. The chemical numerous. The obvious stand- characters are stated on the authority chiefly of Carter, Heilbron, and Lythgoe (1939), lation, positively is Miwa known (1940), and Tseng (1945). The character of the flagelfew examples, is stated by authority of Petersen of rather (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 respects 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. — The 54] Classification of Ochromonadalea: Fig. 8. colony; b, zoospores. C-f, a, b, Lower Organisms Chrysocapsa paludosa after West (1904); Phaeocystis globosa after ScherlTel (1900); c, a, a a colony X 50; d, a cell with two plastids, a mass of leucosin forming on a mound of protoplasm projecting into the central vacuole; e, production of zoospores; f, a zoospore. i, Cell of g, h. Cell and statospore of Ochromonas granularis after Doflein (1922). k, A very Monas sp. j. Two cells of Brehmiella Chrysohydra after Pascher ( 1928) young colony of Dendromonas virgaria after Stein (1878). 1, Colony of Ccphalo. m. Cells of Epipyxis utriculus after Stein, op. cit. Colony of Synura Uvella. x 1,000 except as noted. thamnium Cyclopum op. cit. n. after Stein, Phylum Phaeophyta and distinguished these fungi from practically all [ 55 others by the presence of cellulose in their walls. The phylum 1. thus assembled may be organized as four Miscellaneous groups, mostly small and relaatively unspecialized, of varied body type; not of the characters of the following groups 1. Comparatively numerous classes. and Class 1. Heterokonta. Class 2. Bacillariacea. Class 3. Oomycetes. Melanophycea. specialized groups. 2. Unicellular brown organisms with shells of silica consisting of two parts 2. Organisms of fungal or chytrid body types producing swimming spores with paired unlike 2. fiagella Filamentous and thallose brown algae Class 1 . Class 4. HETEROKONTA Luther Class Flagellata or Mastigophora Auctt., in part. Class Heterokontae Luther 3, no. 13: in Bihang Svensk. Vetensk.-Akad. Handl. 24, part 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). classes Chrysomonadineae , RhizoChrysosphaerineae, Chrysotrichineae, Heterochloridineae, Rhizochloridineae, Heterocapsineae, Heterococcineae, Heterotrichineae, and Heterosiphoneae Pascher in Beih. bot. Centralbl. 42, Abt. 2: Divisions Chrysophyceae chrysidineae, and Heterokontae, and Chrysocapsineae, 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 genus may be regarded as the type. The chrysomonad his discovery of Chlorosaccus, flagellates are the core of this class the five orders into which it is and of the first and this two among divided. In the classification of these two orders, three novelties will be noted. (a) Pascher (1913) made of the chrysomonad flagellates three orders character- 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 ized respectively by paired — The 56] Fig. 9. Classification of Ochromonadalea Conrad (1926). h, : a, Lower Organisms Mallomonas roseola, based Syracosphaera Quadricornu; c, on Stein (1878) and Calyptosphaera insignis; d, Cal- Colony and zoospore g, Part of the thallose growth of Hydrurus h, Cell, and i, j, statosporos of Chromulina Pascheri after Hofeneder foetidus. (1913). k, 1, Skeletons of Dictyocha Fibula and Distephanus Speculum from diatomaceous earth at Lompoc, California, m, Rhizochrysis Scherffeli after Doflein ciconus vitreus; after Schiller (1925). of Epichrysis after Pascher (1925). (1916). Mix 1,000. Silicoflagei.lata: e, f, 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 members 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 pigmented organisms, examples whose vegetative state is that of clusters unicellular 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 They are further characterized by production of resting cells of a type leucosin. called statospores. This occurs by the deposition within the protoplast of a globular 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. shell The group which typical Heterokonta. is treated as the third order of the present class includes the Compared with impression of a markedly distinct typical green algae, these organisms give the 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 additional orders, one may say that it is not contrary to current knowledge. 1. Mostly pigmented; non-pigmented examples mostly producing motile cells with two class; fiagella. 2. Brown or 3. colorless. Producing motile cells flagella (exceptionally 3. Producing motile cells flagellum; or without late with two Order more) with known 2. 1. Ochromonadalea. flagel- stages Green Non-pigmented, producing motile 1. one cells Order Order 2. Silicoflagellata. 3. Vaucheriacea. Order 4. Choanoflagellata. Order 5. Hyphochytrialea. with one flagellum. 2. Predatory, flagellate in the vegetative condition, each cell bearing a collar-like protoplasmic ridge 2. Parasitic or saprophytic, the vegetative cells Order 1. non-motile, walled 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). : The 58] 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. with The spindle forms within the intact nuclear The chromosome number appears membrane 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 its poles at the centrosomes. to two which are released to swim and conjugate in pairs. This order is believed to represent the direct ancestry of the two following, and calls into brown Not filamentous. also of the typical 1. 2. algae. Flagellate stages with a pair of equal and a third which flagella is shorter or longer 2. Flagellate two with stages Family 1. Trimastigida. Family 2. Chrysocapsacea. Family 3. Monadina. Family Family 4. Syncryptida. Dinobryina. unequal flagella. 3. Without calcareous structures at- tached to the cell walls. 4. Cells not enclosed in loricae, i. e., 5. open Not shells. flagellate in the vege- tative condition 5. Flagellate in the vegeta- forming free-swimming circular or tive condition, not globular colonies 5. Free-swimming circular or globular colonies 4. 3. Cells enclosed in loricae With calcareous structures 5. at- Family 6. Hymenomonadacea. tached to the cell walls Family 7. Phaeothamnionacea. 1. Filamentous 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 pinheads, 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 Family 2. Chrysocapsacea [Chrysocapsaceae] Pascher [ in Siisswasserfl. 59 Deutschland 85 (1913). Family Chrysocapsidae Poche in Arch. Prot. 30: 156 (1913). Nonmotile cells with brown plastids (usually two), imbedded in gelatinous matter and 2: forming colonial aggregates, the protoplasts sometimes escaping as zoospores with two flagella. Chrysocapsa Pascher, in fresh water, the colonies few-celled. Phaeosphaera 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 Heteromonadina Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 815 (1884). Family Chrysomonadaceae 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 Ochromonadidae 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 distinctions 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 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 nutrition, here occurring in an organism which is capable also of photosynthesis. Heterochromonas includes organisms of the structure of Ochromonas but without plastids, being presumably saprophytic, and containing only a pigmented speck by which it is supposed 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, freestalks. swimming Ochromonas like nates cells of Monas Kent desigThere are believed to be several was in some member of it that Loeffler or attached like Stylochrysalis [Physomonas in the attached condition). group remains poorly known. It 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 species, but the (1889) first by deposits of generic names iron. just The comparatively unfamiliar original spellings of the two ( 1946). The name Uvella Bory mentioned were restored by Kudo 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 Synuraceae 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 Dinohryaceae Engler in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 2: 570 (1897). Pigmented or loricae, that Ochromonas or Monas, sheltered in transparent open shells, solitary or colonial. The pigmented colorless cells of the characters of is, in examples have generally been referred to Ochromonadaceae (or whatever), the to Monadidae (or whatever). Pigmented, solitary, flagella markedly unequal: Epipyxis, Stylo pyxis; flagella apparently equal: Chry so pyxis Stein {Derepyxis 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 nutrition. Non-pigmented, solitary, flagella markedly unequal: Stokesiella; flagella apprrently 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 Coccolithidae 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 Thoracosphaeracee Schiller in Rabenhorst Kryptog.-Fl. Deutschland ed. 2, 10, Abt. 2: 156 (1930). Y3imi\it5 Syracosphaeridae, Calcisolenidae, Thoracosphaeridae, and Braadrudosphaeridae 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 colorless (coccoliths) of definite form. More than twenty genera and nearly 150 species have been described (Lohman; 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 Schiller; cell bears a whorl or elongate bristles. Cells of Calcisolenia are fusiform, without with an armor of two layers of spiral bands of calcareous matter. In Hymenomonas and Coccolithus Swartz 1894 [Coccosphaera Wallich 1877, non Perty 1852; flagella, Coccolithophora Lohman 1902) the coccoliths are punctured and accordingly ringshaped; 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 Mediterranean 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 particularly abundant in certain Cretaceous deposits. Family 6. Phaeothamnionacea [Phaeothamnionaccae] Pascher in Siisswasserfl. Deutschland2: 113 ( 1913). Family Chrysotrichaceae Fascher (1914). Family Nematochrysidaceae Pascher (1925). Brown organisms, minute, marine, epiphytic, filamen- Phylum Phaeophyta [ reproducing by zoospores bearing paired unequal tous, flagella. 61 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, however, future study shows their flagella actually to be respectively pantoneme and acroneme, they are to be placed Order 2. in the present order. 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. Subclass Chrysomonadineae Engler in Engler and ITeil, Abt.'la: iv (1900). I: 57 (1895). Prantl Nat. Pflanzenfam. Order Chrysomonadales Engler Syllab. ed. 3: 7 (1903). Chrysomonadinae; Euchrysomonadinae with order Chromulinales; Chrysocapsinae; and Rhizochrysidinae Pascher in Siisswasserfl. Deutschland Heft 2 , (1913). Chrysomonadales, Chrysocapsales, Chrysosphaerales, and Chrysotrichales Pascher 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, Chrysosphaerineae, and Chrysotrichineae Pascher in Beih. bot. Centralbl. 48, Abt. 2: 323 , (1931). Suborders Euchrysomonadina, Silicoflagellina, Rhizochrysidina, and Chrysocapsina 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 produced by Chromulina, Mallonionas, and (of somewhat exceptional character) by Hy drums. Sexual reproduction has not been observed. Mitosis, with an intranuclear spindle and numerous chromosomes, was observed by Doflein (1916) to be in Rhizochrysis. This order is supposed to represent the direct ancestry of orders Choanoflagellata and Hyphochytrialea. amoeboid 1. Neither 2. Not flagellate nor in truly filamentous. the vegetative condi- tion. 3. 3. Microscopic colonies Macroscopic gelatinous simulating filaments 2. Family 1. Chrysosphaeracea. Family 2. Hydruragea. Family 3. Chrysomonadina. colonies Flagellate in the vegetative condition. 3. Without prominent siliceous struc- tures 3. With siliceous scales usually bearing bristles 3. With siliceous internal skeletons Family Family 4. 5. Mallomonadinea. Actiniscea. 62 The ] Classification of Lower Organisms Amoeboid Family 6. Chrysamoebida. 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, Nae1. 1. 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 gelliella, Arch. Prot. 30: 158 (1913). Like Chrysosphaeracea, but the colonies dendroid, becoming macroscopic; producing tetrahedral zoospores and spheroidal resting cells bearing a unilateral crest. Hydrurus foetidus, in mountain growing at the tips, 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 Chrysomonadina, the generic name Chrysomonas being a synonym. Organisms of this genus are rather freely capable of producing pseudopodia and supplementing photosynthetic by predatism, or, alternatively, of producing gelatinous aggregates of walled non-motile cells (Hofender, 1913; Gicklhom, 1922). Chrysapsis differs from nutrition Chromulina having in each cell a single plastid in the form of a network. Solitary producing pseudopodia only occasionally: Lepo chromulina. Bearing whorls of permanent pseudopodia: Cyrtophora, Pedinella, Palatinella (Pascher, attached in cells, 1928). Mallomonadinea Diesing in Sitzber. Akad. Wiss. Wien Math.-Nat. CI. 304 (1866). Family Mallomonadidae Kent (1880). Brown uniflagellate free-swimming cells with an armor of siliceous scales usually bearing bristles. MalloFamily 4. 52, Abt. 1: m.onas, solitary cells, the bristle-bearing scales circular. Conradiella, the scales of the form of long rings about the body. Chrysosphaerella, spherical colonies, each cell with two 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 commonest 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 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 two of the original one. Lemmermann and Gemeinhardt accounted for only six genera and twenty-four 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 species, but 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 547 (1952). Families Rhizochrysidae and MyxochryAmoeboid 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, attached 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. Myxochrysis, 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 organisms producing definite filaments of walled cells and reproducing by anteriorly Grasse Traite Zool. 1, fasc. 1: sidae Hall Protozoology 130, 132 (1953). uniflagellate zoospores. T hall ochry sis. Phaeodermatium. Vaucheriacea [Vaucheriaceae] Nageli Gatt. einzell. Alg. 40 (1849). 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 Order 3. Class Heterokontae and H eter osiphonales Va.s.ch.tr mlitdwigizbZ: 10-21 (1912). Division Heterokontae, Classes Heterochloridineae, Rhizochloridineae, Hetero- capsineae, Heterococcineae, Heterotrichineae , and Heterosiphoneae, ders Rhizochloridales and Botrydiales Pascher in Beih. bot. and or- 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). — The 64] 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°-^'^^ Chlorosaccus fluidus, cells of the colony and zooChlorarnoeba 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, Spim. End of a filament of Tribonema bombycina x 1,000. rodiscus fulvus x 1,000. o. Filament of Vaucheria n, Antheridium and oogonia of Vaucheria Gardneri x 100. Fig. 10. Vaucheriacea: spores, after Luther (1899). sessilis x 100. a, b^ c, d^ 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 This order is soma is haploid. believed to represent the direct ancestry of the two following classes, 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. Bacillariacea 2. The 3. cells walled. Cells regularly dividing into two, forming gelatinous colonies; occasionally producing small numbers of zoospores. 4. 4. The colonies globular or Irreg- ular, becoming macroscopic The colonies dendroid, micro- scopic 3. Family 1. Chlorosagcacea. Family 2. Mischococgacea. Family Family Family Family Family 3. 4. Chlorotheciacea. Botryococcagea. 5. Stipitogogcacea. 6. Chloramoebacea. Cells normally undergoing division into several. 4. 4. 2. 2. The The Producing zoospores Producing no motile cells cells loricate cells amoeboid 7. Tribonematagea. Filaments of uninucleate cells 1. Cells becoming highly multinucleate, formFamily 8. Phyllosiphonacea. ing filaments or at least producing rhizoids Family 1. Chlorosaccacea [Chlorosaccaceae] Smith Freshw. Algae 145 (1933). Family Heterocapsaceae Pascher in Hedwigia 53: 13 (1912); there is no correspond1. ing generic name. Gelatinous aggregates of cells which may divide, causing the :. The 66] 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 Sciadiaccae 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 the parent. Large free multinucleate cells, more cells of the same form as 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- an older name of Ophiocytium parvidum (Perty) A. antedates the generic name Ophiocytium Nageli (1849); new combinations are required for the dozen additional species of this genus. The teriological systematists, is Braun (Copeland, 1954). It cells attached: some species of Characiopsis; Perionella; Dioxys. Family 4. Botryococcacea [Botryococcaceae] Pascher in Hedwigia 53: 13 (1912). by production of non-motile cells. Solitary or colonial cells reproducing strictly 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). tached Amoeboid cells with green plastids, partially enclosed in loricae at- 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, Rhizochloridae, Chlorarachnidae and Myxochloridae Deflandre in Grasse Traite Zool. 1, fasc. 1 217-222 (1952). Amoeboid organisms with green plastids, without loricae, sometimes swimming by means of paired unequal flagella. Chloramoeba, Chlorochromonas, 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 Linnaean 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 : common in freshwater pools. From typical green algae of similar appearance 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. 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 sometimes as much as 2 mm. in diameter, anchored by much-branched colorVaucheria 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 globes, less rhizoids. 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 of the type called pantacroneme, with lateral appendages bearing a single flagellum and a terminal whip-lash; the cell bearing also a protoplasmic collar, usually surrounding the base of the flagellum. 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. 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 CraspedornonaIt is dina Stein idae Kent 1. c. op. Families Bikoecidae, Codonosigidae, Salpingoecidac, and PhalansteriFamilies Codonoecina and Bikoecina Biitschli in Bronn Kl. u. Ord. cit. 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 Thierreichs (1952). Characters of the order. Cells naked, solitary: Monosiga; colonial: CodoJames-Clark [Codonosiga Stein), Sphaeroeca. Cells imbedded in gelatinous matter, the collars contracted: Phalanseterium. Loricate: Salpingoeca, Bicosoeca, siga Poteriodendron. choanoflagellates were discovered by James-Clark (1866, 1868), who made at 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 distinguished by the production of exceptionally large and elaborate colonies. Kent The the — The 68] Classification of Lower Organisms described Proterospongia Haeckeli as a colonial organism of amoeboid and choanoflagellate 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 c X 500, the remainder x 1,000. Phalanasterium digitatum; Foteriodendron petiolatum. spp.; c^ A, Salpingoeca ampullacea; e, Salpingoeca Clarkii; i, c-f after Stein (1878). Phylum Phaeophyta Order 5. [ 69 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 protoplasm 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 deliquescence of the tip of the exit tube. Either before this or afterward, the mass of protoplasm undergoes cleavage into uninucleate protoplasts which generate flagella. Sometimes, 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 The resting spores germinate by producing exit tubes and discharging zoospores as ordinary centers do. species). may be called the chytrid body 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. The body type which has just been described type; organisms of this 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 ostium 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). — The 70] Classification of Lower Organisms Hyphochytrialea: a-e^ Anisolpidium Ectocarpii; a-c, individuals deEctocarpus; d, mitotic figures x 2,000; e^ cell of Ectocarpus filled by a mature individual discharging spores, f, g, Rhizidiomyccs apuphysatus; f, zooFig. 12. veloping in cells of spore; g, oogonium of Achlya parasitized by three individuals, h, i, j^ llyphochytrium catenoides; h, zoospore; i, young individual; j, mature individual with filaments, 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; Subdivision and class Bacillariales Engler Syllab. 6 (1892). 1 (1889). Hauptclasse Diatomeae Haeckel Syst. Phylog. 1: 90 (1894). Subclass Bacillariales Engler in Engler and Prantl Nat. Pflanzenfam. Teil la: I, Abt. 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 oldest deposits are of the Cretaceous age. Thus it apmodern 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- the form of a white earth. pears that diatoms are a 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 loopholes). The shells consist basically of something of the nature of pectin heavily imThe appear markedly 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 recognizably a spindle, but the chromosomes, instead of appearing within it, form a ringshaped mass about its middle and divide into two ring-shaped masses which move pregnated with silica and characteristically sculptured. different in different aspects: along it to its extremities. The the aspect which nuclear is membrane cells in effect top or ceases to be recognizable early in — The Fig. 13. Bacillariacea Classification of : a, Mclosira Lower Organisms sp., a living cell and an empty one. b, c. d, e. Sections of a valve of PinnuGirdle and valve views of cell of Cyclotella sp. 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. j, k, Girdle and valve views m, Girdle and valve views of Gomphonema sp. (the former shown, o. Girdle and valve views ing the gelatinous stalk by which the cell is attached), 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. h, i. Girdle and valve views of Rhoicosphenia curvata. of Navicula sp. 1^ 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 Subrahmanyan (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, Cocconeis, 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 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 supsposed the latter examples to represent a stage in the evolution of sexual reproduction under some zwingender Nothwendigkeit: much more probably, they are products of degeneration. In Cyclotella, Iyengar and Subrahmanyan found the production 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 resulting cell condition. The to the filamentous green Heterokonta Tribonema and Bumilleria are closely similar 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 provided 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 corresponding 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 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- numerous duction of auxospores in Cyclotclla to involve sexual processes. Schiitt divided the Centricae into three groups with names in -oideae (presumably subfamilies) and these into nine groups with names in -eae (presumably tribes). Subsequent authorities have made of Schiitt's The minimum families tenable number of groups a varying number of families. corresponding to Schutt's is three, 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 intercalation 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. 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 Order (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 position; exhibiting, when possessed of a true raphe, a gliding motion; cells usually with two plastids. Phylum Phaeophyta The motion of the pennate diatoms [ 75 upon surfaces, with frequent redepends 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 produce auxospores, usually by sexual processes. The majority inhabit fresh water. Eleven families are currently recognized. is a gliding versal, in either direction of the long axis of the cell. It 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 West op. 2. cit. Tabellariea [Tabellarieae] Kutzing op. cit. 110. Family Tahellariaceae 281. Cells symmetrical with respect to three planes, with longitudinal internal partitions. Tabellaria. Family 3. Bacillaria Ehrenberg Infusionsthierchen 136 (1838). Family Diatomaceae West op. cit. 284. Cells symmetrical with regard to three planes, with transverse internal partitions, solitary, or joined valve to valve in ribbons, or corner to comer Diatoma. Meridiea [Meridieae] Kutzing op. cit. 61. Family Meridionaceae West 283. Cells symmetrical with regard to two planes, wedge-shaped both in valve in zig-zag chains. Family op. cit. and 4. 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. Naviculea [Naviculeae] Kiitzing op. cit. 90. Family Naviculaceae Rabenhorst 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 Gomphonemaceae West op. cit. 297. Cells wedge-shaped. Gomphonema. Family 7. Cymbellea [Cymbelleae] (Harvey) Kiitzing op. cit. 84. Family Cocconemaceae West op. cit. 298. Cells with two planes of symmetry, in valve view crescentshaped 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 Family 5. 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. c. 303. Each The two cell with two marginal raphes. Surirella. valves of each cell unlike, one with a raphe, one with a pseudoraphe. 76 The ] Family Classification of Achnanthea [Achnantheae] 11. thaceae and Cocconeidaceae West op. Lower Organisms Kiitzing op. cit. cit. 81. Families Achnan- 289, 290. Achnanthes, Rhoicosphenia, Cocconeis. Class 3. Class OoMYCETEs Winter in OOMYCETES Winter Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. 1: 32 1 (1889), in (1879). Phycomyceten de Bary Vergl. Morph. Pilze 142 1884), in part. Class Phycojnycetes Engler and Prantl Nat. Pflanzenfam. II Teil: ( 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. 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 Biflagellatae is Saprolegnia. Conventional botanical classification recognizes within the group of Fungi a subnamed Phycomycetes, which is in turn divided into Oomycetes and Zygomycetes, the former including the chytrids. This arrangement suggests an evolutionary 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. ordinate group There is an old hypothesis (Sachs, 1874) that Vauchcria may represent the direct ancestry of Saprolegnia. This hypothesis could not be taken seriously while Saprolegnia and its allies were known to produce heterokont zoospores, while Vaucheria was supposed to be a typical isokont green alga. Now it implies that in the present group the fungal body type again appears probable. It more primitive than the is chytrid. The Oomycetes may be organized l.Of fungal body type, i.e., as three orders. consisting of fila- ments. 2. Essentially aquatic 2. Mostly not aquatic, parasitic on higher plants 1. Of chytrid body type, i.e., Order 1. Saprolegnina. Order 2. Peronosporina. Order 3. LAGENroiALEA. the cells not elong- ated to filamentous form, though sometimes proliferating or producing rhizoids 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- swimming) or giving evidence of an 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. spores diplanetic (exhibiting two periods of ancestral diplanetic condition. The old ordinal 1. 1. Filaments not constricted Filaments constricted at intervals. 2. Filaments not differentiated into basal and reproductive parts 2. Family 1. Saprolegniea. Family 2. Leptomitea. Filaments differentiated into basal and Family 3. Rhipidiacea. Saprolegniea [Saprolegnieae] Kiitzing Phyc. Gen. 157 (1843). Family 18 (1872). Aquatic Oomycetes consisting of Saprolegniaceae Cohn in Hedwigia 11 branching filaments of essentially uniform diameter without crosswalls other than reproductive parts Family 1. : those which set apart differentiated reproductive structures. Coker (1923) 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, These well-known organisms are called water molds. According there are about eighty definitely recognizable species. currently with hemp to They may be 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 Couch, 1932). The sharp-pointed formed within the nuclear membrane. The membrane disappears toward the end of the mitotic process, and a nucleolus, which single flare of concurrent mitoses (Davis, 1903; spindles, ending in centrosomes, are 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 organized as an egg. of the oogonium; side of When when it becomes several eggs are produced, they share all of the cytoplasm only one egg is produced, some of the cytoplasm is left out- it. Sperms are produced in small multinucleate antheridia borne at the tips of filaments 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 — The 78] ,^t*SJi Classification of ' •ITi''!' Lower Organisms nVgi 'Villi Oomycetes: a. Filaments and sporangia of Dictyuchus sp. x 50. Zoospores of the second stage of swimming, of Achlya caroliniana and Saprod^ Oogonia and antheridia of Dictyuchus legnia ferax, after Couch (1941) x 1,000. 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. Fig. 14. b, C, : 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 except the zygotes are haploid. The organs of asexual reproduction are cylindrical sporangia terminal on the filaments. Within these the multinucleate protoplasts undergo cleavage into minute uninucleate spores. It is chiefly by details of the behavior of the sporangia and spores meiotic: all cells (the latter diplanetic, monoplanetic, or not swimming at all) that the dozen genera 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 are distinguished. Diplanetism is into a beak toward the narrow anterior end, where the flagella are attached. Spores released 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 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 lies at 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 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 in spores in a single row. In Achlya proper, the spores are discharged without flagella, and swim only once. In Thraustotheca the monoplanetic spores are rebreakdown 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 to encyst leased by irregular : 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. (1897). 1: iv 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 airfrom 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 Phytophthora to bear the paired flagella, respectively pantoneme and acroneme, which are born cells cut off : typical of Phaeophyta. In the multinucleate oogonia of most members of the group, single flares of mitoses The sharp-pointed spindles, described in some accounts as ending in centrosomes, are formed within the persistent nuclear membrane, which undergoes constriction during the final stages of mitosis. A coenocentrum appears (this structure occur. 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 peculiar zygote, containing genetically uniform, This order is a is is many cell. The further history of the resulting nuclei which are not by any evident necessity unknown. evidently a specialized offshoot of the preceding. good example of a transition group; many The family Pythiacea 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 1. Family 1. Pythiacea. Family 2. Albuginacea. Producing conidia usually in clusters at the (conidioends of specialized filaments phores) parasites on land plants. the 2. Conidiophores brief, unbranched, ; conidia in chains 2. Conidiophores elongate, usually branched, the conidia solitary or clustered, not Family 3. Peronosporacea. 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 parain chains Family 1. Phylum Phaeophyta sites attacking land plants under moist conditions. as sporangia if formed in water, as conidia if [81 The formed reproductive structures act 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 Peronosporacea [Peronosporaceae] Cohn 3. Parasites of higher plants, called in Hedwigia downy mildews. The ovoid solitary or in clusters, not in chains, 11: 18 (1872). conidia are produced 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 downy mildew cola, causing of potatoes Order which produced the many attacking 3. viti- of grapes. Phytophthora injestans, the cause of the blight Irish famine of 1846. Peronospora, the many species kinds of plants. 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 3. 3. 2. The 3. cells not proliferating. The zoospores diplanetic The zoospores not diplanetic Family Family 1. Ectrogellacea. 2. Olpidiopsidacea. Family 3. Sirolpidiacea. cells proliferating. Marine 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 3. known diatoms and red and brown algae. Olpidiopsidacea [Olpidiopsidaceae] Sparrow in Mycologia 34: 116 (1942). Olpidiopsis and a few other genera, with some thirty known species, attacking 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. species, attacking Family 2. : 82 The ] Classification of Lower Organisms Lagenidiacea [Lagenidiaceae] Schroter in Engler and Prantl Nat. 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 external 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 Family 4. Pfianzenfam. I Teil, : 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). Melanospermeae Harvey Man. British Alg. 1 (1841). Order Pycnospermeae and tribe Angiospermeae Kiitzing Phyc. Gen. 333, 349 Series (of order Algae) (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). Phaeophyceae Hauck in Rabenhorst Kryptog.-Fl. Deutschland Series {Reihe) 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). Dictyoteae Schaffner in Ohio Naturalist 9: 448 (1909). CXdiSS Subclass Melanophyceae Setchell and Gardner in Univ. California Publ. Bot. 8: 387 (1925). Classes Isogeneratae, Heterogeneratae (with subclasses Haplostichinae stichinae) 7: 91 and Cyclosporeae Kylin in Kungl. Fysiog. Sallsk. Handl. n. and Polyf. 44, no. (1933). Filamentous or thallosc Phaeophyta, yellow to brown in color and living by photoproducing 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. synthesis, ^Lagenocystis nom. nov. Lagena Vanterpool and Ledingham in Canadian Jour Res. 2: 192 (1930), non Parker and Jones 1859. L. radicicola (Vanterpool and Ledingham) comb. nov. Phylum Phaeophyta The cells are [ 83 walled chiefly with readily hydrolyzable modified polysaccharides. mannose units. A polysaccharide 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 Algin, the soda extract of kelps, consists of chains of oxidized of each protoplast. A glycogen- or dextrin-like 1940; Tseng, 1945). The dextrosan, laminarin, and plastids contain chlorophylls a is stored c (Strain, in (Miwa, 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, membrane each pole of the 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 is present outside of the spindle. In Stypocaulon, a comparatively primitive brown at alga, division begins. Swimming cells are produced by primitive brown algae as spores and as morpholomost advanced brown algae, such cells are gically undifferentiated gametes; in the produced only as sperms. The flagella are attached laterally. the longer except in order Fucoidea (Kylin, 1916). The anterior flagellum (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. is Longest Thuret (1850) discovered that most brown algae produce swimming cells from 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 haploid individuals of Ectocarpus are alike, and E. siliculosus may be said to have a facultatively complete homologous life cycle. The haploid individuals produce plurilocular reproductive structures; the swarmers from these act either as spores, restructures of The 84] Fig. 15. — Stages Classification of Lower Organisms 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 ing intermediate examples and rather free parallel evolution. is constantly reveal- 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 classification 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- may be recognized. Producing spores, that is, cells which germinate without syngamy. tinguished as follows 1. 2. All spores bearing flagella. 3. Having an alternation of haploid and diploid stages which are alike, both being filamentous; or else completely lacking 4. 4. The The one of these stages. filaments uniseriate seriate 3. Not 4. 1. Phaeozoosporea. Order 2. Sphacelarialea. Order 5. Cutlerialea. Order 4. SpoROCHNoroEA. Order Order 6. 3. Laminariea. Dictyotea. as above. Haploid stage thallose, not dis- developed than the diploid stage tinctly 4. Order filaments becoming pluri- less highly Haploid stage filamentous, distinctly less highly developed than the diploid stage. 5. Diploid stage filamentous; or, if partially or com- pletely thallose, the thal- 5. 2. lose part with apical growth Diploid stage thallose, its growth intercalary Producing large non-motile spores : 86 The ] 1. Classification of Lower Organisms Producing no spores; all individuals diploid and reproducing exclusively sexually Order 7. FucoroEA. 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. Order , 14: 233 (1850). Phaeosporeae Wettstein Handb. syst. Bot. 1: 173 (1901). Ectocarpales Bessey in Univ. Nebraska Studies 7: 288 (1907). Phaeosporales and suborder Ectocarpineae Taylor in Bot Gaz. 74: 435, ser. 3, Order Order Order 436 (1922). Microscopic brown algae of the form of undifferentiated uniseriate branching filaments, 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 Streblonema, which make up the family Ectocarpea [Ectocarpeae] Kiitzing (family Ectocarpaceae Cohn). Order 9 2. Sphacelarialea [Sphacelariales] (Oltmanns) Engler and Gilg Syllab. ed. 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 Ectocarpea. 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: (1885). Class Dictyotales Engler in Engler and Prantl Nat. Pflanzenfam. (1897). Akinetosporeae Oltmanns I Teil, 302 Abt. 2 ix Morph u. Biol. Alg. 1: 473 (1904). Order Tilopteridales and Class Tetrasporeae with order Dictyotales Bessey Univ. Nebraska Studies 7: 290 (1907). Scries Aplanosporeae Setchell and Gardner in Univ. California Publ. Bot. 649 (1925). in 8: Phylum Phaeophyta [ 87 Filamentous or thallose brown algae with haploid and diploid stages equally developed, producing large spores without flagella, solitary or few in the sporangia. Here are placed two families, Tilopteridea and Dictyotacea. Family Tilopteridea [Tilopterideae] Cohn only from European coasts. They They is a small group, apparently known are evidently closely related to the Ectocarpea. which may become consist of branching filaments pluriseriate. In Haplospora more so), the haploid stage minute swimming cells of the structure (poorly known; but Tilopteris and other genera are even bears both plurilocular structures, releasing usual in brown algae, and unilocular structures uninucleate protoplasts without flagella. structures is which release inferred that the their contents as single diploid stage bears only unilocular their contents as single quadrinucleate non-motile spores. It swimming that the protoplasts released eggs, capable, which release The cells from the plurilocular structures are sperms, and from the unilocular structures on haploid bodies are however, of reproducing the haploid stage if not fertilized; further, that the nuclei of the quadrinucleate spores released by diploid individuals are hap- and become on germination the nuclei of loid, The as many cells of the haploid body. Tilopteridea are believed to represent the evolutionary transition between Ectocarpea and the following family. Family Dictyotacea [Dictyotaceae] (Hauck) genera, Dictyota, Zonaria, Padina, etc., Kjellmann includes about twenty 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 Haupt (1932). There are studied by Mottier (1898, 1900), Williams (1898), and distinct male haploid individuals, female haploid indivi- and diploid individuals, all of the same vegetative structure. The males produce 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 nonmotile spores; in Dictyota, each one produces four. duals, 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. California Publ. Bot. 8: 554, 570 (1925). Order Sporochnales Sauvageau Brown in Compt. Rend. 182: 364 (1926). algae producing motile spores, the haploid stage reduced to scant undiffer- when thallose with apical 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 Laminentiated filaments, the diploid stage filamentous or thallose, growth. Ralfsia ariea on the is an exception to the other. The haploid body of the form of a short-lived body of a few undifferentiated ments, like a reduced Ectocarpus, bearing gametangia reduced to single cells, fila- has 88 The ] Classification of Lower Organisms been demonstrated by Kylin ( 1933, 1934, 1937) in a wide variety ot genera, as Ascocyclus, 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, Mesogloiacea, 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 Ralfsiaor 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. Superficial hairs, growing in intercalary fashion, may become few, and growth may become restricted to a single apical cell. By differences in the detailed guished two orders among manner of growth, Setchell and Gardner the thalloid forms just mentioned. It is distin- evident, however, that the thallose structure (and, likewise, differentiation of gametes) has developed repeatedly and independently in the present group. possible to divide it Knowledge which would make into several recognizably natural orders is it not yet available. Cutlerialea [Cutlcriales] Bessey in Univ. Nebraska Studies 7: 289 Order 5. Brown algae producing motile spores, the haploid and diploid bodies being macro- ( 1907). 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 growing 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 comparatively 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 Sporochnoidea, 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 dimengrowing in intercalary fashion. sions, the diploid stages thallose, This numerous group, like the preceding small one, is evidently a specialized offshoot 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- members, but their microscopic and reproductive characters, as observed 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 microscopic 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 ferentiation of than the sperms, are also The flagellate. 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 summits 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 fasts and stalks, this tissue is meristematic, adding cells to the plastids; on the hold- within and inwith fewer plastids. tissue creasing the thickness. Internally there is a cortex of larger cells 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 Nereocystis there are actual perforations from cell to cell. The trumpet fibers are not quite The 90] — Classification of Lower Organisms Familiar kelps of Pacific North America: a, Egregia Menziesii; h, NereoLuetkeana; c, Macrocystis pyrifera; d, Postelsia palmaeformis. All approximately X /aFig. 16. cystis 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, specialized 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 hundreds of meters, and these accounts have been repeated in 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 pneumatocyst or float at the base. The stalks of Nereocystis Luetkeana also were said to be lengths, matters of textbooks down to recent times. Frye, Rigg, 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 exposed 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 On mentioned are confined to the northwest coast of North America. where they occur, kelps are used as fertilizer. They have been used com- coasts mercially as sources of potash, as much 1-3% as 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 appear properly to be placed in the same order are treated by Papenfuss (under Dictyosiphonales) 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 Fucea [Fuceae] (others have been segregated). In family Kiitzing (family Fucaceae Cohn), called the rockweeds, the bodies The 92] are dichotomously branching flat there Classification of thalli. Lower Organisms In family Sargassea [Sargasseae] Kiitzing Growth is by division a differentiation of holdfasts, stalks, blades, and floats. is of a single apical cell in each branch or member. There are the usual two tissues, a and an inner tissue of larger cells which superficial photosynthetic tissue of small cells 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 infrequently, 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 divisions, producing sixty-four nuclei. These become the nuclei of sperms. Female conceptacles 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 individuals attached at the mouth f, female individual with two young diploid All x 1,000. mouths of oogonia. Phylum Phaeophyta [ 93 In Fucus, these become the nuclei of as many eggs. In other genera, is reduced by degeneration of some of them, or of some of the nuclei before cell division. In Sargassum, Kunieda (1928) found each oogonium to produce a single egg in which seven nuclei undergo dissolution while one re- ducing the eight. number mains of functional eggs 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; Yamanouchi (1909) first gave a full account of the meiotic process. The haploid chromosome 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 discovery 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. Order Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852). 1: 10 (1848). Cohn in Zeit. wiss. Zool. 4: 275 (1853). Orders Flagellata and Cilio-flagellata Claparede and Lachmann Etudes Infus. Flagellata 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). Man. Inf. 1: 27, 211 (1880). 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). Class Flagellata Kent Class 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, Desmokontae, and Dinophyceae, Pascher in Beih. bot. Centralbl. 48, Abt. 2: 325, 326 (1931). Division Pyrrhophyta G. Protistes trichocystiferes M. Smith Freshw. Algae 10 (1933). 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, flagellate, 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 membrane and The organisms included to the cell eject thread-like bodies when stimulated. 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 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- which bears a 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. Dinoflagellates 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 zinc chlor-iodide. The [ 95 euglenids store granules of a white solid believed not to be and called paramylum. The plastids of cryptomonads and starch green, yellow, brown, bluish, or red. a and dinoflagellates are of various colors, oflF-color Those of dinoflagellates contain chlorophylls an exceptional chlorophyll which occurs also in Tribonema. Euglenids and chloromonads are typically of the same bright green color as typical plants, e; the latter is and the euglenids are known to have the same chlorophylls, a and Franck and Loomis, 1949). b, as typical plants (Strain, in 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 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. times, 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 isolated 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 character. 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 name: he called it by the French common names protistes 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. and giving it a Latin trichocystiferes The phylum is treated as a single class. Class MASTSGOPHORA (Diesing) Bu'tschli Classes Cryptomonadineae , Rhizocryptineae, Cryptocapsineae, Cryptococcineae, Desmomonadineae, Desmocapsineae, Dinoflagellatae, Rhizodininae, Dinocap- The 96 ] Lower Organisms Classification of 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. sineae, 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. if present brown, olive, or the normally two. Flagella at the anterior end of the cell, not moving in longitudinal and trans- Pigmentation like; 2. flagella verse grooves. 3. Not walled in the flagellate con- not markedly dif- flagella dition, or not differentiated as anterior and circumferential Usually walled in the flagellate ferentiated, 3. .Order 1 . Cryptomonadalea. condition; flagella respectively anterior 2. and circumferential longitudinal and circumferential, in 2. Adiniferidea. Order 3. Cystoflagellata. Order 4. Cilioflagellata. Order 5. Astoma. moving impressed upon the cells. Not walled in the flagellate con- grooves 3. dition 3. Flagellate cells with a wall usually of articulated plates 1. Order Flagella attached laterally, respectively Pigmentation if present typically bright green, flagella normally solitary, sometimes two or more 7 (1903). 1. Cryptomonadalea [Cryptomonadales] Engler Syllab. ed. 3: Subclass Cryptomonadineae Engler in Engler and Prantl Nat. Pflanzenfam. Order 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 observed in the motile condition, then naked, of dorsiventral (exceptionally 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 colors, usually t t>.>. — c, Chib, Rhodomonas baltica after Kylin ( 1935 ) Fig. 18. a, Cryptomonas sp. lomonas Parmecium. d, Cyathomonas sp. e, Sennia sp. f. Vegetative cell, and All x 1,000. g, zoospore of Paradinium Pouchetii after Chatton (1920). its . volume and contains most have observed of the chromatin. details of mitosis. membrane and form intact nuclear right angles to the axis of the cell. Dangeard (1910) and Belar (1916) The numerous chromosomes appear a disk- or No drum-shaped figure with within an its axis at granule more massive than the chromosomes and divides with them. About thirty species are known. They may be treated persists 1. Flagellate cells elongate, with one as five families. plane of symmetry. 2. Not parasitic, flagella not markedly dif- ferentiated. 3. Non-motile in the vegetative con- dition 3. Flagellate dition in the vegetative Family 1. Cryptococcacea. Family 2. Cryptomonadina. con- 98 The ] 3. Amoeboid Classification of Lower Organisms vegetative con- amoeboid organisms, the flag- in the Family dition 2. Parasitic of ella swimming stages 3. Paramoebida. respectively anterior and trailing Family 4. Paradinida. 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 includes yellowish cells called zooxanthellae, internally symbiotic in Radiolaria, Rhizopoda, sponges, coelenterates, and rotifers. It is believed that the supposed zoospores of various amoeboid organisms are actually flagellate reproductive cells of Chrysi1. Flagellate cells with two planes of symmetry 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. Deutschland 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 Division Desmokontae; classes [ 99 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 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. either The few known organisms of this group Family Adinida Bergh Morph. Jahrb. may be treated as a single family. 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. in 7: 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. Abt. u. Ord. Thierreichs Inhalt (1887). Tribe [group of families] Gymnodinioidae Poche in Arch. Prot. 30: 161 (1913). Classes Rhizodininae, Dinocapsineae, Dinococcineae, and Dinotrichineae; orders 1, 2, 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 phylar the carelessness, as name Myxocystoda it appears, to publish in the same work the synonymous 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 as a label in a phylogenetic families. 1. Relatively unspecialized; having stages freely propelled by two flagella, with a single girdle, tentacles, and unspecialized eyespots or no none; not parasitic; 2. commonly pigmented. Walled and non-motile in the vegeta- tive condition 2. Flagellate in the vegetative condition Family Family 1. Phytodiniacea. 2. Gymnodiniacea. 100 1. The ] Not as above, Classification of Lower Organisms always without photosynthetic pigments. 2. Amoeboid 2. Flagellate or free-floating. 3. With multiplied Family 3. Dinamoebidina. Family 4. PoLYKRiKroA. Family 5. Pouchetiida. Family 6. Protodiniferida. without girdles, tentacles or specialized light-sensitive organelles 3. With one 4. girdle or none. Cells more or less isodiamet- ric. 5. With prominent sitive light-sen- organelles, some- times with tentacles 5. Without light-sensitive or- ganelles, with tentacles. 6. Not exceptionally large 6. Reaching exceptional sizes, to 1 mm. in di- ameter Cells dome-shaped Family 7. Noctilucida. 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, Dinococcales, and Dinotrichales, and families Gloeodiniaceae, Hypnodiniaceae, Dinotrichaceae, 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 4. . 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 the protoplasm in the cysts is seen to [ become luminous 101 in response to disturbance of the agents of phosphorescence at sea. In Gymnodinium Lunula the protoplast of each large globular cyst undergoes division into several the medium; they are among which do not immediately become protoplasts crescent-shaped, deposits a cell wall, and is flagellate; each of them becomes released by dissolution of the wall of the 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- parent niaceae Pascher (1931), not based on generic names. Non-pigmented amoeboid organisms producing crescent-shaped cysts which germinate by releasing gymnodinioid 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. 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 Gymnodinium 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- There ciated the usual two differentiated flagella. The Of nuclei there are usually half as many 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 as of girdles. cells 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 They occur only in this family and the following. Pouchetiida [Pouchetiidae] Kofoid and Swezy in Mem. Univ. of Califcrnia 5: 414 (1921). Each of the gymnodinioid cells contains a light-sensitive apparatus, 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 seem not Family fully established. 5. 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 the daughter centrosomes do not necessarily the to divide, the centrosome divides; the poles of the nucleus where chromosomes assemble. Each daughter centrosome appears to generate one and flagellum to complete the neuromotor apparatus of a In due course, the endosome, nucleus, and cell undergo constriction. rhizoplast, blepharoplast, eel]. is lie at 102 The ] Family 7. Classification of Lower Organisms Inf. 1: 396 (1880). The single and Swezy (1921; usually known Noctilucida [Noctilucidae] Kent Man. species Noctiluca scintillans (Mackartney) Kofoid as A^. miliaris Suriray ) is a predatory marine organism, the subglobular cells reaching dimensions exceeding 1 mm., luminescent when stimulated and accordingly contributing to phosphorescence at sea. Each cell is marked by an extensive depression representing the sulcus; the girdle tions as a cytostome. is obsolete. A tooth in the sulcus A part of the area of the sulcus func- 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 and flagellum are absorbed. The attraction sphere becomes elongate and 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 tentacle its central part 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 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 nuclei produced are ( 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 a comparatively recent discovery of Kofoid. Blastodinida [Blastodinidae] Chatton in Arch. Zool. Exp. Gen. 59: Noctiluca; Craspedotella is Family 9. 442 (1920). Ordre Blastodinides Chatton in Compt. Rend. 143: 981 (1906). Families 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. Family Peridinaea Ehrcnbcrg Infusionsthierchcn 249 (1838). Family Dinifera Bergh in MoVph. Jahrb. 7: 273 (1882). Order Dinoflagellata BiitschU in Bronn Kl. u. Ord. Thierreichs 394 1: 1, (1858). Abt. 2: Inhalt (1887). Subclass Peridiniales Engler in Engler and Prantl Nat. Pflanzenfam. lb: V (1896). I Teil, Abt. Phylum Pyrrhophyta Class Peridineae Wettstein Handb. syst. [103 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 for a whorl of 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 error of observation: the circumferential flagellum was mistaken cilia. recognized. Peridinaea Ehrenberg Infusionsthierchen 249 (1838). Family PeridinInf. 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 expands 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 protoplast with each of its ends covered by a complete cell wall. Dividing cells are of Family idae 1. Kent Man. quite different appearance. Families Ptychodiscida, Cladopyxida, and Amphilothida of Kofoid (1907, the in the feminine; explicitly made families by Poche, 1913) are minor segregates from Peridinaea. names 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, Oxyphysis, Amphisolenia, Triposolenia, etc. The 104] — Classification of Lower Organisms Fig. 19, b, Gytnnoa, Tetradinium javanicum x 1,000 after Thompson (1949). 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, Dinamoehidium 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 Order 5. Order Order Order [ Astoma 105 Siebold in Siebold and Stannius Lehrb. vergl. Anat. 1 10 ( 1848) Phytozoidea Perty Kennt. kleinst. Lebensf. 161 (1852), in part. Flagellata Claparede and Lachmann Etudes Inf. 1: 73 (1858), in part. 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 being stichoneme; mostly producing a solid storage product, not staining blue with iodine, called paramylum. cell usually 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- Family tative condition 3. Flagellate in the vegetative 1 . Colaciacea. con- Family 2. Euglenida. dition Flagellum not swollen and usually not forked near the base; cells not pigmented. 3. Cells without internal rod-shaped Family 3. Astasiaea. structures; flagella stichoneme 3. Cells with internal rod-shaped structures; 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 2. — The 106] Fig. 20. a., Classification of Colacium Arbuscula d, Euglena acus. glena Spirogyra. siella alligata after Pascher (1931). Lower Organisms after Stein (1878). Phacus sp. All x 1,000. e, f, b^ Euglena viridis. Trachelomonas sp. EuKleb- c, g, 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(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 rhizocliffe plasts. a new The flagellum, already split at the base, divides throughout its length into two; 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. is completed by constriction of the membrane. The cell divides by 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 individuals possess the capacity to make amino acids and others do not, evidently as Nuclear division constriction heritable characters. Astasiaea Ehrenberg Infusionsthierchen 100 (1838). Family Astasiidae 375 ( 1880) Family Astasiina Biitschli in Bronn. Kl. u. Ord. Thier1 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 stichoneme, 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 Family 3. Kent Man. : base. The Inf. . : . 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 preparations show a rhizoplast connecting this to a centrosome immediately outside the nuclear membrane. The blepharoplast divides during the early stages of mitosis, and 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. the flagellum appears to divide lengthwise. 108] The Classification of Lower Organisms Scytomonas pusilla Stein {Copromonas subtilis Dobell) occurs in the and toads. When cast out with the feces, it exhibits conjugation of frogs liminary to encystment (Dobell, 1908). Family 4. Anisonemida [Anisonemidae] Kent Man. Inf. 1: intestines as a pre- 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 Peranema is grown fast to the cell membrane, and is detected only with difficulty (Hall, : b, c. Stages of mitosis in Menoidium incurvum Fig. 21.— a, 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, Entosipon 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 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 cytofamily. 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. Hollande op. cit. flagella springing 1, fasc. 1: 235 (1952); family Thaumatomonadidae 686. Unicellular organisms, mostly green, with two diiTerentiated 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. rhizoplast extends into the cytoplasm, but fails to From come the base of each flagellum, a into contact with the nucleus. Several granules or swellings, not definitely identifiable as blepharoplasts or centrosomes, are distributed along the length of each rhizoplast. In mitosis, which takes place within an intact nuclear membrane, the numerous subglobular chromosomes 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 form a blunt-ended (originally name). named Thaumatonema, but there is among plants an older genus of this : 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 Kryptog.-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, growths), with cell walls of chitin, i. e. tapering filamentous out- containing no cellulose; producing motile cells with solitary posterior acroneme flagella. Type, Chytridium Olla Braun. From KOVT6q^oar. 6tt[o9ioc;, rearward, and 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 centrosomes at the poles. The nuclear membrane disappears toward the end of the process. The formation of motile cells (zoospores and sometimes gametes) occurs in enlarged 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 morpological 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 acroneme. 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 Synonymy ARCHIMYCETES (A. Fischer) SchaflFner of the phylum. Characters of the phylum. Previous authors have arranged these organisms in a sequence from strictly unicellular forms to typically filamentous forms. In the following treatment, this sequence seems reasonis reversed. The course of the evolution of the group is unknown, and it Oomycetes. The class among the as sequence same the in types the body able to place and Chytridinea, is treated as two orders, Monoblepharidalea, essentially filamentous, unicellular or producing filaments Order 1. which taper or are swollen Monoblepharidalea [Monoblepharidales] Sparrow at intervals. in Mycologia 34: 115 (1942). Suborder Monoblepharidineae Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. (1897). Blastocladiincae Petersen in Bot. Tidsskr. 29: 357 (1909). 1: iv 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 ] Family 1. Classification of Lower Organisms Monoblepharidacea [Monoblepharidaceae] A. Fischer in Rabenhorst 1, Abt. 4: 378 (1892). Gonapodiineae and Gonapodiaceae Kryptog.-Fl. Deutschland 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 produced in smaller antheridia and larger oogonia which are more familiar forms terminal and subterminal on the filaments, the branches commonly proliferating below them, the eggs without flagella. filaments, the gametes in the 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 Benjamin, 1954). Myrioblepharis Thaxter is believed not to be an organism; it is described 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 Sphaerocladia, 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 morphologically 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. macrogynus 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 They [113 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 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. species have the Order 2. Chytridinea [Chytridineae] (Schroter) Campbell Univ. Textb. Bot. 152 (1902). Orders Myxochytridinae and My cocky tridinae A. Fischer in Rabenhorst Kryptog. 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 The higher plants. They may be by commonly thought chytrids are themselves complete individuals. of as prevalently parasitic on algae and 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 settling following varieties of vegetative structure may be noted, (a) A zoospore, 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) Development may be as above except that only one center is formed. The body thus described is of the Entophlyctis type of Sparrow (1943). the single center, penetrating body is its of the Chytridium type of the Rhizidium type if it is if (c) The zoospore may host or substratum only by rhizoids. the center not. (d) The is itself The become resulting in contact with the host or substratum, protoplast of the zoospore may migrate and there become a center without rhizoids; the of the Olpidium type. To the varied bodies thus described, the into the protoplast of the host resulting body is following terminology Pluricentric, with is more applicable: 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 constituting 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 — The 114] Classification of Lower Organisms M a-f, o noble pharella Taylori x 1,000 after germinating spore producing a filament and a rhizoid; b, sporangium releasing spores; c, empty antheridium and sperm uniting with egg; d, sperms escaping from antheridium and zygote escaping from oogonium; e, swimming zygote; g-i, Allomyces javanicus x 100 after Kniep (1929); g, asexual f. encysted zygote, Fig. 22. Monoblepharidalea: Springer (1945); a, (Continued bottom p. 115) Phylum Opisthokonta [115 proximal part of the system of rhizoids develops a large swelling called the aposporangium 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- physis. In other forms, the center generates the nate group. Syngamy occurs in different chytrids in most of the possible fashions, by union of 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- like or unlike walled resting spore (asexual resting spores are also of frequent occurrence). Resting spores germinate by producing zoospores. Meiosis has not been observed, but is believed 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 3. Monocentric. 4. Germinating ing the as a Family 2. Phlyctidiacea. Family 3. Rhizidiacea. Family 4. Synchytriacea. Family 5. Achlyogetonacea. Family 6. Olpidiacea. Zoospores themselves becoming centers, and subsequently sporangia or presporangia 2. Cladochytriacea. distinct body 4. 1. generat- spores center Family without rhizoids, the individual consisting entirely of one or Holocarpic, more i. e., centers. becoming presporangia, 3. Centers 3. each one generating a cluster of sporangia Centers proliferating, giving rise to linear series of sporangia Each center becoming one spor- 3. angium 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 n-r, Blastocladiella cystogena, x 500, after Couch figures in the gametangia, x 2,000. and WhifFen (1942); n, individual producing a resting spore; O, resting spore germ- numerous naked protoplasts; these become flagellate zoospores, which subsequently encyst; q, the protoplast of each cyst divides to produce four gametes; r, young zygote with the flagella of both gametes. inating by release of p, — The 116] Classification of Lower Organisms ' B^^ 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, zoofiG. spores; 23. f, Chytridinea: a-c, sporangium of the host beset with many parasites; g, h, resting and resting cells of the parasite; host containing respectively sporangia tion of resting cell. cells of the i, germina- Phylum Opisthokonta 1. [117 Sporangia opening through opercula. Family 7. Nowakowskiellacea. Pluricentric Family 8. Chytridiacea. Monocentric Family 1. Cladochytriacea [Cladochytriaceae] Schroter in Engler and Prantl Nat. Pflanzenfam. I Teil, Abt. 1: 80 (1892). Family Hyphochytriaceae [Cladochytriaceae) 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 2. 2. (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. 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- Many phlyctis, Diplophlyctis, Loborhiza, etc. Rhizidiacea [Rhizidiaceae] Schroter in Engler and Prantl Nat. PflanzenAbt. 1: 75 (1892). Family Sporochytriaceae [Rhizidiaceae, Polyphagaceae) 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 Family fam. 3. I Teil, sporangia without opercula. A moderate number of species, parasitic on blue-green or green algae, flagellates, or diatoms; or chitinophilous, saprophytic in the shed 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 exoskeletons of migrates into the tip of the rhizoid, which swells and becomes a resting spore. Family 4. Synchytriacea [Synchytriaceae] Schroter op. cit. 71. Family Merolpidiaceae [Synchytriaceae) A. Fischer op. cit. 45. Holocarpic chytrids, the intramatrical cell unwalled in the vegetative condition, becoming a presporangium or a resting spore, either of which gives rise to a cluster of sporangia. Synchytrium, on higher plants; Micromycopsis on Conjugatae. 5. Achlyogetonacea [Achlyogetonaceae] Sparrow in Mycologia 34: 114 (1942). Chytrids without rhizoids, the intramatrical center proliferating and producing a linear series of centers, each of which becomes a sporangium without an parasitic Family The 118] 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, attacking 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 Rickettsiacea. 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 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- of green algae : ( 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). 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 respectively, the terms mycelium and hypha. The walls of the hyphae are believed to consist Parasites of pectic material. A small percentage of chitin is usually present (Schmidt, 1936); (Thomas, 1928; Nabel, 1939; Castle, 1945). The organism Basidiobolus, having hyphae walled with cellulose, is tentatively retained among cellulose is totally absent Inophyta as an exception. The multiplication and dissemination of those organisms types, scattered in the air. them Most Inophyta produce two or is by spores, of various more kinds of spores, some 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 of asexually, others as features of a sexual cycle. Spores below. expedient to mention at this point the growths called lichens, which are taxonomic group, either subordinate to Fungi or of the It is traditionally treated as a same rank. Lichens are gelatinous or color, posts. common an impure green trees, or fence microscope, in the hands of de Bary and others, showed that they consist of two types, colorless filaments like those of Inophyta, and pigmented The of cells thallose growths, usually of everywhere, terrestrial or epiphytic, as on stones, — The 120] Lower Organisms Classification of 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 Ascomyceten, 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 cells of quite the . . — are a classic example of symbiosis. Clearly, the group of lichens known to is not to be maintained; the algal components are have natural places among algae, and the inophyte components are to be among Inophyta, almost all in various orders of 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 assigned to their natural places class are to be applied to the inophyte components. Another common example furnished by at least 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 some of those which live of symbiosis involving inophytes on or is in the tissues of higher plants agents of absorption. Jones (1951) estimated the number of species of Inophyta as 40,000. This surely an extreme underestimate. is 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 accomodation 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 1. is a solecism. Reproducing sexually, or by apomictic processes clearly of sexual origin. 2. The becoming a thick-walled none or inconsiderable The zygote not becoming a thick-walled zygote resting cell; fruits 2. Class 1. Zygomycetes. Class 2. Ascomycetes. resting cell; mostly producing fruits. 3. The zygotes giving directly, each to rise, usually in- sporangia called typically containing spores called ascospores asci, eight Phylum Inophyta 3. The [121 zygotes giving rise indirectly to conidiophores called basidia, each bearing typically four conidia called basidiospores 1. Not known Class 1. to reproduce sexually ZYGOMYCETES Class 4. Basidiomycetes. Class 3. Hyphomycetes. (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 (1875). Class Zygomycetes Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, ed. Abt. 847 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 crosswalls; 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 below). Traditionally, the Zygomycetes are associated with the Oomycetes. The association 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. , 1. Mucorina [Mucorini] Fries Syst. Myc. 3: 296 (1832). Suborder Mucorineae Engler in Engler and Prantl Nat. Pflanzenfam. Order Abt. Order Order Order Order 1: iv I Teil, (1897). Mucorineae Campbell Univ. Textb. Bot. 158 (1902). Spirogyrales (presumably in part only) Clements Gen. Fung. 12 (1909). Mucorales Smith Crypt. Bot. 1 405 (1938). Zoopagales Bessey Morph. and Tax. Fungi 117 (1950). : The typical Zygomycetes, mostly saprophytic, not producing explosively discharged 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 Zygomycetes: Lower Organisms sporangia x 50; b-d, pregaZygote of Phycomyccs nitens f, g, Conidiophore with young conidia, and mature conidia, after Blakeslcc 1904). h, i, Conjugation of Synceof Syncephalis pycnosperma after Thaxtcr (1897). phalis nodosa after Thaxter, op. cit. j, Sporangium of Synccphalastrum raccmosum k, Sporangium of Flaplospoiangium lignicola after Martin after Thaxter, op. cit. (1937), x 1,000. Fig. 24. a-d, Rhizopus nigricans; mctes, suspensors and gametes, and zygote x 200. ( e. a, 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 most Zygomycetes, between branches from hyphae of two mating 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 dissolution (Keene, 1914, 1919). Ordinarily, the zygote enlarges and becomes a thickwalled 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 interpreted as making up fruits. Endogone produces definite fruits of considerable size. A zygote germinates by production of a hypha bearing a sporangium (Blakeslee, at random, but, in types, designated plus 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 3. Not 4. Family 1. Mucoracea. Family 2. Piptocephalidacea. as above. Producing sporangia or else conidia as outgrowths from a homologous with a sporangium, solitary on an unbranched stalk Sporangia or conidia solitary and terminal on branches of a branched sporangiophore or knob, 4. if sporangia, conidiophore; produced, without columellae Family 3. Mortierellacea. Family 4. Zoopagacea. Family 5. Endogonacea. 1. Producing macroscopic fruits Family 1. Mucoracea [Mucoraceae] Cohn in Hedwigia 11: 17 (1872). Mucorina whose spores are produced exclusively in sporangia with columellae solitary on unbranched sporangiophores. Mucor L., typified by M. Mucedo, is now limited to a 2. Parasitic on amoebas or nematodes The 124] 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 outgrowths 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. Mortierellacea [Mortierellaceae] Schroter op. cit. 130. Family Chaetocit. 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). Mucorina parasitic in amoebas or nematodes, producing conidia. The hosts of Zoopagacea inhabit the soil and are infected by contact with hyphae or conidia. From the point Family 3. cladiaceae Schroter op. 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, of contact, a Stylopage. Family 5. Endogonacea [Endogonaceae] Paoletti in Saccardo Sylloge Fungorum Endogonei Fries. Mucorina saprophytic in soil or wood, producing 8: 905 (1889). 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 Abt. in Engler and Prantl Nat. Pflanzen- (1897). Order Entomophthorales Smith Crypt. Bot. 1: 408 (1938). Zygomycetes, mostly parasitic, producing explosively discharged conidia [Massospora, while clearly belonging to the group, is an exception to the stated character). fam. I Teil, 1: iv These organisms, although of the general nature of ordinary Inophyta, exhibit markedly distinguishing the two families from the generality of Inophyta and from each other. The position here given to them is the customary cytological characters one; it is doubtful that it is natural. Entomophthoracea [Entomophthoraceac] Berlese and de Toni in Saccardo 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 multinucleate 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 Family 1. Phylum Inophyta the mechanism [ 125 The conidiophore ends in a columella projecting into The columella develops a double wall. Increasing pressure of discharge. the base of the conidium. 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. Conidia which come down on unfavorable substrata may form and discharge secondary conidia. Adjacent them or as cells may conjugate, the thick-walled zygote forming either in one of Many examples produce thick-walled an outgrowth from one of them. resting spores without conjugation. The Olive (1906) described the nuclei and the process of mitosis in Empusa. resting nuclei are fairly large, 7-9 [I in diameter. In the course of division, two stain- with strands of chromatin radiating from them. These 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, resistant granules are seen, move apart, while the nucleus 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 cellulose. In manure the filaments develop into a scant branching mycelium. The protoplasm 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 membrane 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 11: 17 (1872). Order .4jco5porgag Cohn in Ascomyceten Sachs Lehrb. Bot. ed. 4: 249 (1874). AscoMYCETES Bennett and Thistleton-Dyer in Sachs Textb. Bot. English Hedwigia (1875). Class AscoMYCETES Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. (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 which the manner spores, called ascospores, typically eight in of cell division called free cell formation, a part of the cytoplasm. i.e., 847 ed. 1 : 32 asci, in number, are delimited by the in such fashion as to exclude The 126] The hyphae of Classification of Lower Organisms Ascomycetes are septate and the most often uninucleate. cells 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 surface; 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- and the pycniospores spermatia. a mass may go into a resting condition, becoming thick-walled, natively be called spermagonia, Hyphae woven into 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, As it is called a stroma. and ascospores, Dangeard (1893, 1894, 1907) reached definitely the conclusion that they are essentially sexual products. There had been earlier observations, 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 recognizable male and female reproductive structures, each mating type may produce both. In Ascomycetes which may be regarded as primitive, difTerentiated male and female cells are produced. The male cell or antheridium is ordinarily terminal on a to asci 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 syngamy 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 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 place. This effect is achieved in the formation by a peculiar process called crozier formation. 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 placed that each cell receives nuclei of different origin. final cell division before ascus Phylum Inophyta [127 fuse to form a binucleate cell grow forth The and give stage rise to which may become an additional more asci than one. ascus, or else may two nuclei of different origin is called the 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 consisting of cells with dikaryophase. It is characteristic of consists prevalently of binucleate cells. The was first described by Harper (1895, 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 chromosomes appear and divide. As they move toward the poles of the spindle, the nuclear detailed behavior of nuclei in the ascus 1897, 1900) 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, Eremascus fide alhus, fide G/om^r(?//<z, fide Walker (1935) De Lamater et al. 2 6 (1953) Lucas (1946) HypornycesSolaniv:xr.Cucurbitae,fi.dtY{.\r?,ch.{\9'^9) .... 4 4 Lachnea scutellata,^dt^ro\\'n {\9\\) Neurospora crassa, fide McClintock (1945) 5 Peziza do miciliana,^dt 8 {\921) Phyllactinia corylea, fide Colson (1938) Pyronema confiuens\?Lr. igiieum, ^dtV>ro\vn {\9\b) Taphrina deformans, fide Martin (1940) ?)c\\n\iz 7 10 5 4 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) According to 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- 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. soid pattern. In Harper believed that a fusion of nuclei follows immediately the fusion of gametes; 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 that the The 128] 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 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 Ascomycetes 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 spores 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 which are globular or vase-like, opening through a single pore, the 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. to small fruits ostiole, and differing Asci produced in perithecia or apothecia usually discharge the ascospores vioThe mechanism of discharge is apparently simply turgidity. Some asci show no visible adaptations for the discharge of spores; others have lids (opercula) lently. 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 temperature 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 formulated the counter-argument. The Ascomycetes resemble the Mucorina in nutrition, in producing no flagellate cells, and in multi- relationship. Atkinson (1915) 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 centrosomes 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 cytoplasm 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); Order 1. Endomycetalea. not producing fruits l.The zygote giving rise to filaments of cells with more than one nucleus, these producing the asci. 2. Producing 3. The 4. fruits. fruits cleistothecia. Asci scattered in the fruits; mostly saprophytes with 4. 3. branched conidiophores Asci in one cluster, or solitary, in the fruits; mostly parasites with unbranched conidiophores The fruits, originally closed, open- Order 2. Mucedines. Order 3. Perisporiacea. ing by irregular pores or regular or Order 4. Phacidialea. Order 5. Cupulata. irregular clefts The fruits apothecia 3. The fruits perithecia. 3. 4. Producing a normal mycelium on insects, the mycel- Order 7. Sclerocarpa. Order 8. Laboulbenialea. Order 6. Exoascalea. 4. Parasitic 2. Order ium reduced Not producing fruits, the asci rectly from the mycelium 1. arising di- Endomycetalea [Endomycetales] Gaumann Vergl. Morph. Pilze 135 (1926). Subclass Hemiasci Engler Syllab. 26 (1892). ILuttrell (1951) has presented a 3 complete reorganization of the class. He sets apart in which the ripe ascus exudes major subordinate group Bitunicatae five orders vesicle and discharges the spores from this. as a 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. 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 Pflanzenfam. many I Teil, Abt. 1: spores cut out by free cell formation. Dipodascus, Eremascus, The Endomyces, 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 Ascoidea. asci of the last are 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 breadand 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 simplicity, and has attracted much study, contributing much to an understanding of the for a long time 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. I Teil, 2: 3 (1887). Suborder Plectascineae Engler Abt. 1: V (1897). in Engler and Prantl Nat. Pflanzenfam. 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 The conidiophore [131 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. making of genuine Camemand 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 Particular species of Penicilliuni are involved in the bert 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 in the formula CgHnOiSNiR. The structural formula is R believed to be as follows (Editorial Board of the Penicillin, Monograph on RCONH — CH — CH — S — C I OC The of the Chemistry of 1947): I N (CH3)2 I CH COOH. sexual reproduction of Aspergillus and Penicillium involves the syngamy differentiated cells. The zygote sends out ascogenous hyphae which bud scattered asci; the neighboring cells send out hyphae which become woven oflF 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 hyphae about the asci; Ctenomyces, on feathers, recognized by comblike 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. loose weft of 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 25. Ascomycetes: a-e, Dipodascus albidus after Juel (1902), x 1,000; gametes; b, syngamy; c, development of ascus; d, e, lower and upper parts of a f, Erysiphe graminis, haustorium penetrating an epidermal cell of mature ascus. g-k, Cleistothecia of a grass and conidiophore bearing a chain of conidia x 500. Fig. a, Perisporiacea x 100: g, of Erysiphe sp.; h, of Microsphaera of Uncinula sp.; k, of Phyllactinia sp. j, sp.; i, of Podosphaera sp.; Phylum Inophyta The more They familiar Perisporiacea are those of family Erysiphea [ 133 [Erysipheae] 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 characteristic of the genus. In Erysiphe and Sphaerotheca iS. pannosa is the common rose mildew), the fruits bear unbranched sinuous appendages like vegetative hyphae; the Winter. are parasites on plants, mostly producing a white 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 appendages 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 comparatively 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 Sclerocarpa. 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 a. which appear tenable are distinguished as follows: and flattened, usually releasing the spores through one or more pores or clefts (Order Hemisphaeriales Theissen, Microthyriales Clements and Shear). families Fruits minute The 134] Classification of Lower Organisms Family Microthyriacea [Microthyriaceae] Lindau (in Engler and Prantl, 1897). 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 foreParasitic going, but the surface of the fruit not radiate or radiate only at the margin. Family Trichothyriacea [Trichothyriaceae] Theissen and Sydow. Parasitic on 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 inophytes, the Engler). Family Hysteriacea [Hysteriaceae] Saccardo Sylloge 721 (1883). Parasitic on 2: higher plants or saprophytic. Family Graphidiacea [Graphidiaceae] Clements (1909). lichens or parasites on lichens, largely tropical and An enormous group of 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 cupor 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 send out branching filaments which become septate [ in 135 such fashion that the ultimate are binucleate. These cells form croziers and produce asci. During the development of the ascogenous hyphae, other hyphae, more slender, grow up from the cells vegetative mycelium; these produce a disk of undifferentiated cells below the layer and send up sterile hairs (paraphyses) among them. (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 of asci, Gaumann 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, producing 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 throughout 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 believed 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, suggesting a transition to order Sclerocarpa. Some species, particularly among the 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 parasites of reticulate relationships, such as reputable naturalists of the past supposed to many groups. The appearance is of course illusory; sufficient study of other groups has made it possible to distinguish the resemblances among them which indicate relationship from those which are results of parallel evolution. The study of the exist in Ascomycetes has not yet been carried this far. — The 136] Classification of Lower Organisms Lachnca scutellata after Brown (1911) x 1,000; karyogamy; d, fusion nucleus; e-i, stages of mciosis; 1, Apothecia x 2, and m, ascus x 250, of j, k, early stages of free cell formation. Lamprospora leiocarpa. n, Apothecia x 2, and O, ascus x 250, of Aleuria rutilans. q, Fruit x 1, and r, ascus x 250, of MorV, Apothecia of Sclcrotinia cinerea x 2. s-x, Taphrina deformans after Martin (1940) x 1,000; s, growth on chella conica. surface of an infected leaf; t, karyogamy; u, mitosis; V, homeotypic anaphase in the ascus; w^ development of ascospores; x^ germination. Fig. a, b, 26. Ascomycetes: formation of crozicr; a-k, C, Phylum Inophyta [137 305 ( 1907) 6. Exoascalea [Exoascales] Bessey in Univ. Nebraska Studies 7 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 Order : generic rank. Clements and Shear associated the curly-leaf parasites with a collection of saprophytes producing small and undifferentiated disk-like or indefinite fruits, as Pyronema, Ascocorticium, and Agyrium; and offended against the principles of nomenclature by re-naming the order Agyriales. It is probable that something of the nature 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 occurring 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. of 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). {Unterreihe) Pyrenomycetineae, sub-suborders [Unterordnungen) Hypocreales, Dothideales, and Sphaeriales Engler in Engler and Prantl Nat. Suborder 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 respectively 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, conidiophorcs and conidia of Ccrcospora type, b, longitudinal section of pycnidium; C, primordium of perithccium with ascogonoium bearing a trichogyne; d, e, ascogenous 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 confidently 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 are to be ales,. recognized fourteen families in the present group or groups. To these a great number of lichen-formers, properly Sphaeriales and Hypocre- added 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, 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 ultra-violet radiations, or which may reach the Among size of golf balls. Hypocreales, Nectria cinnabarina is common as a saprophyte twigs of poplar. It produces small wart-like red stromata which bear first on dead 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. Ohio Naturalist 9: 450 (1909). 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 Class Laboulbenieae Schaffner in Parasites genera. Many Laboulbenialea occur as two forms, male and hermaphrodite. A male individual 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 A perithecium consists of a wall, of order and pattern, surrounding an protrudes from the perithecium and fascicle of asci as described a definite and then one or more of cells produced number perithecia. in definite egg which bears a trichogyne; the trichogyne receives the sperms. The zygote gives rise to a which crowd aside and destroy the inner cells of the charge the ascospores (usually eight in the ascus, and divided into two wall and discells) through the ostiole. Those who would link the Ascomycetes with the red algae entertain the hypothesis that the Laboulbenialea represent the transition. This hypothesis The Laboulbenialea appear to have evolved from Sphaeriales with Class Classes is surely mistaken. are a highly specialized group, not a link between others. 3. HYPHOMYCETES Hyphomycetes and Coniomycctes They solitary perithecia. Fries Fries Syst. Myc. Families Hyphomycetes and Coniomycctes Fries Epicrisis 1 3: 261, 455 (1832). (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 provided 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 is most likely to try Hyphomycetes is as follows: collector or plant pathologst The system of to find it there. 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- Order cetes. Family 1. Phomatacea [Phomataceae] Clements Gen. Fung. 121 (1909). Family Sphaerioideae or Sphaerioidaceae Saccardo; but Sphaeria belongs to order Sclerocarpa. Family Phomaceae Clements and Shear (1931). Pycnidia hard and black as in Sphaeriales and Dothideales. Phoma, Ascochyta, Diplodia, Septoria, each of many species. Clements Gen. Fung. 128 (1909). Family 2. Zythiacea [Zythiaceae] 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). Family 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. Melanconialea [Melanconiales] Engler in Engler and Prantl Nat. PflanI 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 2. zenfam. 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 The 142 ] less compact than Classification of Lower Organisms enormous number of species producing as Snyder and Hansen (1941, 1945) find that a stroma. Fusarium, an conidia crescent-shaped rows of cells. 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 17 (1872). 11: 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, : 450 306 (1907). Classes Teliosporeae and Basidiomycetae Schaffner in Ohio Naturalist 9 (1909). Inophyta which produce, as a feature of the sexual cycle, conidiophores called basidia, each producing typically four conidia called basidiospores. Germinating basidiospores give nuclei. Syngamy occurs among rise to mycelia of cells of these cells with solitary haploid mycelia, usually simply by contact of cells; the rusts produce differentiated sperms in spermagonia resembling 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 concurrent 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- vmdifTerontiated 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 apto it, and Linder (1940) plicable to these; Martin (1938) has attempted to refine simplify it. Frequently, the seat of meiosis is a thick-walled resting spore or difTerentiated cell called a probasidium, an otherwise upon which the proper basidium develops, 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 after meiosis, as 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 holobasidium or autobasidium. Gaumann 1926) distinguished two types of holobasidia: ( 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 indistinct, but the nuclear sap remains distinct from the cytoplasm nearly until the completion of division; it then disappears, leaving the groups of daughter chromothe stichobasidium, in 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) Coleosporium Vernoniae, fide Olive (1949) Coj&rmuj, fide Yokes (1931) Eocronartium, fide Fitzpatrick (1918) &;frffa, fide Whelden (1935) Gymnosporangium, fide Stevens (1930) Melampsora, fide Savile (1939) Myxomycidium flavum, fide Martin (1938) Puccinia, fide Savile (1939) Transchelia, fide Savile (1939) Russula, fide Ritchie (1941) Scleroderma, fide Lander (1933) f/romycg'^, fide Savile (1939) 2 8 4 4 4 2 4 8 4 4 4 2 4 some at least of the reports of a chromosome number of 2 misinterpreted observations of one pair of choromosomes from may have resulted behind another. Normally, only the two meiotic divisions, producing four nuclei, occur in the Savile suggests that 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. — The 144] Classification of Lower Organisms Basidiomygetes a. Two germinating basidiosporcs of Agariciis campcsproduce mycelia which anastomose freely, the cells becoming plurinucleate, after Hein ( 1930) x 500. b, c. Young and older basidia of Cystobasidium sebaceum, d-g, Eocronartium muscicola after P'itzpatrick (1918); d, fusafter Martin (1939). ion nucleus; e, homeotypic division in the basidium; f, four-celled basidium; g, production of basidiospore. h^ i^ i, Basidia of Ustilago Heujlcri, U. Hurdei, and Tille(Continued bottom p. 145) Fig. 28. : tris , Phylum Inophyta [ 145 Most basidia discharge the spores actively, to a distance of a fraction of a millimeter. 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 Sporoholomyces. 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 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- of the 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 evagination of the walls of the sporangia. Largely, as Vuillemin's hypothesis has it seems, by Gaumann's influence, become generally accepted. Gaumann was disposed to derive the Basidiomycetes from something like Ascocortkiiim, 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 Sclerocarpa 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 ascospores pass. This led to the conclusion that the Basidiomycetes producing probasidia are the lowest, and to this extent his reasoning appears cogent. the rusts as the lowest Basidiomycetes, which seems He went on to identify 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. k, 1, Basidia of Patouillardina cinerea after Martin ( 1924) (1935). m, Basidium of Sebacina sublilacina after Martin (1934). n, Basidium o, p, younger and older basidia of Tulasof Protodontia Uda after Martin ( 1932). q-t, Development of the basidium of nella phaerospora, after Martin (1939). tia Tritici, after Sartoris . 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 Guepinia Spathularia, after (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 subsidiary 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 Order 1. Protobasidiomycetes, Order 2. Hypodermia. Order 3. Ustilaginea. gitudinally Order 4. Tremellina. Without probasidia, the basidia undivided. 2. Fruits gelatinous, basidia producing only two spores on stout sterigmata Order 5. Dacryomygetalea. .Order 6. Fungi. Order 7. Dermatocarpa. gelatinous fruits 3. Parasitic, mostly not producing fruits; the rusts 2. Probasidia produced by rounding up and deposition of thiclc walls by the gener- mycelium Without probasidia, the basidia divided ality of the cells of the 1. 1. 2. Not 3. lon- as above. Basidia in a layer which forms with- out protection or becomes exposed. 3. . formed in closed fruits which do not open to expose them Basidia as a single layer Protobasidiomycetes Engler in Engler and Prantl Nat. Pflanzenfam. I 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 basidium becomes divided by transverse walls into four cells, and each of these sends out Order 1. Teil, Phylum Inophyta [ 147 an elongate sterigma which bears a curved basidiospore. The organism produces also conidia, either from the mycelium, the fruits, or directly from the to the surface basidiospores. A series of unfamiliar other genera, Platygloca, Cystobasidium, Septobasidium, 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. etc., 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, Order 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 which releases spores called aeciospores; this stage is accordingly called the stage, and its host the aecial host. In addition to aecia, this stage usually pro- structure aecial duces pycnidia or spermagonia. mycelium are then, is The typical reproductive structures of the dikaryote clusters (telia) of spores called teliospores or teleutospores; this stage, the telial stage, and its The telial stage usually The teliospore, or rather host the telial host. beside the teliospores, others called uredospores. produces, (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 statements mean that a normal rust produces spores of five kinds. Rusts producing different kinds of spores were formerly supposed to be different genera; such were the Aecidium, Uredo, and Puccinia of Persoon, who, however, remarked of Uredo linearis, "vereor, ne junior plantula Pucciniae graminis modo sit." De Bary first proved that Aecidium Berberis is yet another stage of Puccinia graminis. by syngamy among cells of the aecial 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. The dikaryophase is initiated, of course, stage. In much The pycniospores are carried out of 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 Allen (1930) has described of the detail. the pycnidium in exuding fluid, and are carried by insects; they 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- 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 ally stout chains of several probasidia; those of the many species of Pucciriia are chains reduced 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 was first observed by Sappin-Troufi^y (in Dangeard and SappinThereafter the probasidium gives rise to the four-celled promycelium. to germination: this Troufi'y, 1893). The young life cycle thus described structure not perfectly stable. Aeciospores, uredospores, and and are genetically identical. Spores of the and behavior of any of these types may be produced by processes which normally lead amy is teliospores are alike dikaryote, to another. Thus in Puccinia Malvaccaruvi, the hollyhock rust, syng- 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 individual stalks. Hemileia vastatrix, the coffee rust; Phragmidium spp., autoecious (attacking 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 number 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 mutation, these strains are subdivisible into large numbers of races distinguished by capacity to attack different races of hosts. Given a specimen of rust on wheat, one determines by it trial belongs. upon The seedlings of ten varieties of wheat to which of 189 numbered races 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 Order 3. [ 149 Ustilaginea [Ustilagineae] (Tulasne and Tulasne) Winter in Rabenhorst Kryptog.-Fl. Deutschland 1, Abt. Ustilagineae Tulasne and Tulasne 1: in 73 (1884). 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 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 phragmobasidia. 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. Hybridization, and mutation, particularly in the capacity to attack particular races of hosts, host, the 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 Abt. 1**: in Engler and Prantl Nat. Pflanzenfam. I Teil, (1900). Order Tremellales Bessey in Univ. Nebraska Studies 7: 309 (1907). Order Tulasnellales Gaumann Vergl. Morph. Pilze 487 (1926). iii Saprophytic Basidiomycetes producing gelatinous fruits bearing a layer of basidia which typically become divided into four cells by longitudinal walls. Each cell produces 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 Tremellacea [Tremellaceae] Cohn in Hedwigia 11: 17 (1872). Martin (1935, 1937, 1939) has given much study to this group. It is clearly related to the Protobasidiomycetes; Patouillardina, having basidia divided by oblique walls, is clearly transitional. 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; be that the holobasidia of this more sound than it may supposed that genus are derived from the cruciate basidia of proper Tremel- this interpretation is the obvious one). It is 150 lina The ] by a Lower Organisms Classification of line of descent separate from those which have produced the holobasidia of By leaving Tulasnella in order Tremellina, we spare one more insignificant order. other groups. nition of Order 5. ourselves the recog- Dacryomycetalea [Dacryomycetales] Gaumann Vergl. Morph. Pilze 490 (1926). Suborder Dacryomycetineae Engler Engler and PrantI Nat. Pflanzenfam. in ITeil, Abt. 1**: iv (1900). Saprophytic Basidiomycetes producing small gelatinous fruits bearing holobasidia 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, Dacryomiin tra, Guepinia. Bodman ( 1938) observed the details of the cytological processes in the basidia. This insignificant order, sidered, is and the two great orders next like Tulasnella to be con- 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. 74 (1884). Suborders Exobasidiineae and Hymenomycetineae Engler in Engler and PrantI Nat. Pflanzenfam. I Teil, Abt.'l**: iv (1900). 1: 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 hymenium basidia is called the hymenium. In the lowest members of the group, on the mycelium, on the surface of the host or substratum; in higher examples, it is formed on the surface of more or less complicated 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 the is formed directly 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 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 least, part the conventionally accepted ones. Family 1. Pflanzenfam. sniall Exobasidiacea [Exobasidiaceae] Hennings I Teil, Abt. 1**: 103 (1900). The in Engler and Prantl Nat. basidia directly on the mycelium. A group, mostly parasitic on plants. Exohasidium. Family 2. Thelephoracea [Thelephoraceae] (Saccardo) Hennings (1900). Order Thelephorei Fries Hymen. Eur. 1 ( 1874) FamHy Thelephorei Winter ( 1884) Thelephoraceae Saccardo Sylloge 8: xiii (1889). Fruits of various form, gelatinous, fleshy . . or leathery, the hymenium covering the surface generally except where it faces upward. 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 Clavariacea [Clavariaceae] (Saccardo) Hennings (1900). Order (1874). Family Clavariei Winter (1884). Clavariaceae Saccardo (1889). Fruits fleshy, club-like or branched; stag-horn fungi. Clavaria, generally 3. Clavariei Fries edible. Family 4. Fries (1874). ium on Hydnacea [Hydnaceae] (Saccardo) Hennings (1900). Order Hydnei Family //yi/n^i Winter (1884). Hydnaceae Saccardo (1889). Hymen- 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 PolyporeiYries (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 Dacdalea, the pores are not cylinders but slits; this genus leads into Lenzites, in which the hymenium is borne on radiating plates, and which ing green when bruised. These conventionally stationed in is Agaricacea. Boletus has stout fleshy mushroom-shaped fruits are unattractive, yellow to brown, turn- fruits, but some species are eaten; others are supposed to be poisonous. Family 6. Agaricacea [Agaricaceae] Cohn in Hedwigia Agaricini Fries (1874). Family Agaricini Winter (1884). plates, radiating from a center, called gills. These are the Fungi whose fruits are called 11: 17 (1872). Order The hymenium on mushrooms or are mostly mushroom-shaped, sometimes shelf-like; the texture toadstools. is vertical The fruits 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 development 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 of five classes, white, pink to red, light Lower Organisms brown to rust color, dark brown or purple, or black), will find identification reasonably easy. Popular interest in agarics cerned, of course, with the edible and poisonous. is con- 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 Fruits of Pleurotus have of this character an excentric or is safely edible. lateral stalk, or none, bracket-like, fleshy, with white spores. All species are edible. being shelf- or The most familiar is 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 the oyster are known to be edible; others, as the fly agaric, A. muscaria, recognized by a red cap flecked with white, are extremely poisonous. 7. Podaxacea [Podaxaceae] Fischer in Engler and Prantl Nat. Pflanzenfam. 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- Family I Teil, 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 Sclcrodermatales 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 an unsatisfactory degree, and to to some [ 155 of which the suspicion of artificiality con- tinued to attach. Dodge, translating Gaumann (1928), took account of the course of development whose fruits are characteristically produced 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. of the fruits in rearranging those families 304. B. Fruits appearing Family are the on the surface of the ground. Lycoperdacea [Lycoperdaceae] Cohn in Hedwigia 6. common puffballs, 11: 17 Lycoperdon, Bovista, Calvatia, Lycogalopsis, ( 1872) . These The conglobular 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 tents of the in damp more or etc. less weather, exposing the inner peridium with its terminal pore. The appearance damp condition explains the common name, earth star, and the of the fruit in the 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 scentific 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. Sphaerobolacea [Sphaerobolaceae] Fischer in Engler and Prantl op. 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 Family cit. 9. 346. Sphaerobolus, a saprophyte 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 is not commonly observed. The growing stalk carries the the night or at dawn, and 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. X PHYLUM PROTOPLASTA Chapter 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 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 protoplasm, called pseudopodia, functional in motion and in predatory nutrition. Pseudopodia 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 or standard 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 1. Amoeboid 2. Producing rhizopodia; with 2. 1. Zoomastigoda. Class 3. Rhizopoda. Class 4. Heliozoa. Class 2. Mycetozoa. Glass 5. Sarkodina. shells, these usually calcareous 2. Class in the vegetative condition. Producing filopodia or axopodia; mostly with skeletons, these usually siliceous Producing lobopodia. flagellate reproductive 3. Producing cells; mostly macroscopic, subaerial 3. Not as above; without flagellate stages 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). The 158] Non-pigmented if Classification of flagellates Lower Organisms having acroneme or simple flagella; amoeboid stages, The standard is Bodo. Four orders are to be recog- they occur, having lobopodia. nized. 1. Flagella one or two 1. Flagella four to eight (in each neuromotor system, 2. if flagella; Rhizoflagellata. Order 2. PoLYMASXiGroA. more than one). homologous with parabasal body commonly abif present, Axostyles present, not homologous with flagella; parabasal body present, disap- pearing during mitosis 1. 1. these are Axostyles, sent 2. Order Flagella of indefinite large numbers Order 3. Trichomonadina. Order 4. Hypermastigina. 1. Rhizoflagellata [Rhizo-Flagellata] Kent Man. Inf. 1: 220 (1880). Orders Trypanosomata (the mere plural of a generic name) and FlagellatoPantostomata 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 Order : 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 Siisswasserfl. 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. 1. 2. Cells not notably slender 2. Cells notably slender Family Family 2. Family Family 4. 1. CERCOMONADroA. TRYPANOPLASMroA. Flagellum one. 2. Not 2. Conspicuously amoeboid regularly markedly amoeboid 3. Oicomonadacea. 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). Nonpigmented 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. Rhynchomonas, 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. which can lose their flagella and take on the appearance amoebas have repeatedly been discovered and variously named. So far as the pseudopodia are lobopodia and the flagella are unequal, these organisms belong Biflagellate organisms of ordinary flagella, many accounts fail to establish the equality or inequality of the with the result that the names used in them cannot be applied with confidence. This true of various organisms originally in this family; but is named under Pseudospora, Dimastiga- moeba, and Naegleria. The earliest generic name as described in Cercobodo Senn, 1910. definitely applicaple to organisms Kuhn (1915), and others have described mitosis 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 rhizoplast extends into the nucleus. The chromatin is reticulate, not massed in a karyosome, 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 Ringkorper. 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 membrane there appears a spindle with evident centrosomes at the poles. The centrosomes Belar (1914, 1916, 1920, 1921), in various come presently to the inner surface of the nuclear move membrane, while the blepharo- on the outside. Chromosomes duly assemble at the equator of the spindle and undergo division. Division of the nucleus is completed 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. plasts to adjacent positions — The 160] Classification of Lower Organisms 5. Fig. 30. Rhizoflagellata : a, Bodo sp. x 1,000. b, c, Cercomonas longicauda by Wenyon (1910) in material from a cholera patient; d, the same as e-h, Cryptobia spp.; e-g, cell and identified by Hovasse (1937) in swamp water, 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 & Thompson (1915). q. Division stage of Trypanosoma Brucii after Kiihn & Schuckmann (1911). r, Chaetoproteus [Mastigamoeba aspera) after Schulze (1875) x 100. as identified X 2,000 except as noted. Phylum Protoplasta [161 Bodo edax. Trypanoplasmida [Trypanoplasmidae] Hartmann and Jollos 1910. Family Cryptobiidae Poche in Arch. Prot. 30: 148 (1913). Family Trypanophidae Hollande in Grasse Traite Zool. 1, fasc. 1: 680 (1952). Organisms of essentially the AlexeiefF (1924) described fusions of pairs of cells of Family 2. 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 The numerous species may be and in the gut and blood of fishes. 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 blepharoplast 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 Protozoen 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 specialization, occurring in of the character of the family without particular contaminated water or soil, and as commensals in the intestine of animals. The bulk of the family consists of the slender-celled parasites celled trypanosomes in the broad sense of the word. From which may be 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 some and vertebrates; and some in vertebrates and in inverteand 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. plants; in insects brates other than insects, as ticks 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 1. Confined 2. 2. Phytomonas. to invertebrate animals. Trypanosoma Trypanosoma stage known stage unknown; Herpetomonas. crithidia 162 The ] stage 2. Classification of Lower Organisms known Crithidia. Trypanosoma and crithidia stages un- known 1. Leptomonas. Attacking vertebrate animals. 2. Trypanosoma Trypanosoma stage stage known unknown Trypanosoma. 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. 2. 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 divisions. 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 blepharoplast. Separated from the blepharoplast by a distance of one or two microns there is a conspicuous parabasal body (the kinetonucleus of Woodcock). Fine strands connecting the blepharoplast, parabasal body, and nucleus, have been observed. Most of 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 chromosomes. It divides before the chromosomes, the daughter centrosomes remaining connected by a fine fiber, the centrodesmose. An obscure spindle forms about the centrodesmose; thi' chromosomes undergo division within the spindle, and the daughter chromosome assemble about the centrosomes. Mitosis is completed by constriction the stainable material in the resting nucleus > of the nuclear membrane. 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 The almost entirely new. The parabasal body undergoes constriction. The cell cuts in in such fashion as to divide the cell longitudinally. The blepharoplast 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. which is membrane Schaudinn described complicated processes by which a trypanosome generates 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 trypanosome. Still, the occurrence of syngamy among trypanosomes is inherently probable. differentiated 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 temporarily. In polluted soil or water, or commensal or pathogenic in animals. The oldest genus, Chaetoproteus Stein {Mastigamoeba F. E. Schulze, 1875; Dinamoeba Leidy ?) remains poorly known. This organism and Mastigella are described as fairly large; Craigia is much smaller. Rhizomastix is doubtfully distinct from Craigia. Early the family is names of much this family appear to refer to Rhizomastix as the type, but 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 multiplied; not of the definite characters of the following order. Free-living, chiefly in commensal or parasitic in animals. Polymastix is presumably the type group. It was listed with a query in Biitschli's original publication of family foul waters, or of the 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 neuromotor 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 Christianson (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. The 164] — Classification of Lower Organisms Fig. 31. PoLYMASTAcroA a, Polymastix Mclolonthae after Swezy (1916). b, c, d, Giardia cnterica after Kofoid Streblomastix Strix x 1,000 after Kidder ( 1929) : . & Swezy (1922). Trichomonadina: e, Hcxamastix Tcrmopsidis after Kirby i' Tricercomitus Termopsidis 2ihtv YJirhy (1930). (1930). g, Macrotrichomonas pulchra after Kirby (1938). h. Trichomonas tenax x 4,000 after Hinshaw (1926). Pentatrichomonas obliqua after Kirby (1943). Kirby 1929) x 2,000 except as noted. i, ( . j, Snydcrella Tabogae x 500 after 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 miscellany 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 1. With a 2. single Ochromonadalea. nucleus and neuromotor system. Cells not spirally twisted, at least not as wholes and not conspicuously 2. Entire conspicuously cells Family 1. TEXRAMiTroA. Family 2. Streblomastigida. Family 3. Dinenymphida. spirally twisted. 3. 3. With four free With four or flagella eight flagella proximal ends are grown cell 1. With one or whose fast to the membrane several nuclei and two or more neuromotor systems Family 4. Oxymonadida. 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 ) Family 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 Hominis (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. 1. . . ^Kofoid (1920) gave the history involved in this combination. Davaine, 1860, de- 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 respectively a Chilomastix and a Pentatrichomonas, under which genera they have various names. Kofoid named them respectively Chilomastix davainei and Trichomonas 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 conscribed the flagellates trary that an author to who designates a var. A designates the type in doing argument that the new combination here published the Cercomonas Hominis var. A of Davaine. on the basis of this is so. It is applied The 166] Family 2. Classification of Lower Organisms Streblomastigida [Streblomastigidae] Kofoid and Swezy fornia Publ. Zool. 20: 15 (1919). The only known species is in Univ. Cali- Strehlornastix Strix, a slender spirally twisted organism with four anterior flagella, free-swimming or attached 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 neuromotor 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 freeliving in fresh or foul waters and have been little studied. Hexamita occurs both freeliving 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), preoccupied 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 homologous 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 centrosome and blcpharoplast, the centroblcpharoplast of Kofoid and Swezy, 1919) lies Phylum Protoplasta outside the nuclear move membrane. This [167 structure divides apart along the nuclear membrane. and the daughter structures They remain connected, usually until mito- complete, by a stainable strand, the paradesmose. Definite chromosomes, usually few in number, and an intranuclear spindle, are formed. Mitosis is completed by constriction of the nuclear membrane. In what appears to be the typical course of cell sis is and blepharoplast divide when the centrosome does. Of other some may remain connected to one blepharoplast the other; some may disappear. The parts needed to complete a neuro- division, the rhizoplast parts of the neuromotor system, and some to motor system are regenerated in each daughter cell. 1. With a single nucleus and neuromotor system. 2. Lacking a cresta, costa, or undulating membrane 2. With a trailing flagellum whose base underlain by a cresta 2. With a trailing flagellum the cell grown Family 1. MoNOCERCOMONADroA. Family 2. DEVEScoviNroA. is fast to membrane, forming an undula- membrane underlain by a costa Family 3. Trichomonadida. 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 undulating 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, Metadevescovina. 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. 646 Family 3. Trichomonadida [Trichomonadidae] Wenyon Protozoology 1 (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 undulating membrane underlain by a costa. Entozoic in a wide variety of animals. Trichomonas, 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 beneficial 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. Pentatrichomonas 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 ting 1. With several nuclei . : (Kirby,^1943). icf. footnote, p. 165. The 168] Family Classification of Lower Organisms Calonymphida [Calonymphidae] Grass! in Atti Accad. Lincei ser. 5, Semestre: 730 (1911). Flagellates with radially symmetrical 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. 4. Rendiconti CI. Sci. 20, 1° 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. 4. Hypermastigina Grassi in Atti Accad. Lincei ser. 5, Rendiconti CI. Sci. 20, 1° Semestre: 727 (1911). Order 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 Spiratrichonymphina, Grasse Traite Zool. 1, fasc. 1: 837, 851, 862, 916 (1952). Flagellates, mostly large and of radial symmetry, with single nuclei and indefinitely numerous flagella. Entozoic in roaches and in termites excluding those of 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 family Termitidae. life of Termopsis is not prolonged by the presence of Streblomastix, and it is prolonged 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 division, 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 Lophomonas (Kudo, 1926), neuromotor system of a dividing cell is absorbed or discarded, with the exception of the centroblcpharoplasts, from which new systems generates the remaining parts of a complete system. In and Kofoidia (Light, 1927), the develop. In Trichonympha and Spirotrichonympha the details of nuclear division have 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. much Phylum Protoplasta [ 169 In species of Trichonympha, Leptospironympha, and Eucomonympha from the Cryptocercus, Cleveland (1947, 1948) observed the syngamy of undifferentiated 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- wood roach ternation of haploid and diploid stages. Diploid cells are produced when a centro- 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 recognized an essential feature of the evolution of the sexual cycle. His words suggest the blepharoplast fails to divide, idea that the sexual cycle may have impossibility; the sexual cycle is a originated within the present group. This is normal character of nucleate organisms, and an is 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. fully established in nucleate 2. Flagella distributed generally over the surface of the body or its 2. Flagella in spiral bands 2. Flagella in tufts. anterior part. . . . Family Family 1. TRiCHONYMPHroA. 2. HoLOMASTiGOTororoA. Family 3. LoPHOMONAoroA. 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. Holo3. Flagella in a single tuft 3. Flagella in two tufts 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 Zool. 29: 4. Hoplonymphida [Hoplonymphidae] Light in Univ. California Publ. The flagella assembled in two anterior tufts. Hoplonympha, 138 (1926). Barhulanympha, etc. Family 5. Staurojoeninda [Staurojoenindae] Grassi in Mem. Accad. Lincei CI. Sci. 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 ser. 5, 12: — The 170] 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). assimilis (1921). h, Hoplonympha Natator x 250 x 250 after Kirby (1926). j, after Light (1926). Tcratonympha i, Staurojoenina mirabilis after Koidzumi 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). Wettstein Handb. Stamm Myxophyta 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 2. 2. 1. fruits. Spores produced within the fruits Spores produced on the surfaces of the Order 1. Enteridiea. fruits Order Order 2. Exosporea 3. Phytomy.xii>a. Parasitic, not producing definite fruits Order 1. Enteridiea [Enteridieae] Rostafinski Vers. 3 (1873). Cohort Endosporeae and orders Anemeae, Heterodermeae, Reticularieae, Ainaurochaeteae, 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 Macbride op. cit. cd. 2 (1922). Order Liceales Ma. bride and Martin (1934). Suborder Eumycetozoina Hall Protozoology 230 (1953). 172 The ] Lower Organisms Classification of Predatory Mycetozoa producing macroscopic fruits, these producing internal uniThe type is Lycogala, the sole genus of the order as originally nucleate spores. 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 first published in Latin form, in the category of classes, by de Bary's student Rostafinski. Conventional botany continues to list Myxomycetes as a class of Fungi; conventional zoology makes the group an order of Rhizopoda or Sarcodina. apparently The spores germinate readily in water or appropriate solutions (Jahn, 1905; Gil- Smith, 1929). Their nuclei usually divide once or twice, during or just 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 disbert, 1929; after germination; thus each spore produces cerned definite centrosomes. When the one or two divisions associated with germination 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 connected to the base of the flagella by a conical body of clear protoplasm, the Geisselglocke of Jahn (Jahn, 1904; Howard, 1931). The flagellate cells are not spores, but gametes; they fuse with each other. Skupienski (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 Myxomyceten werde ich mit den Namen Plasmodium bezeichnen." The plasmodium nourishes itself in predatory fashion, on fungus spores, bacteria, and other digestible objects, and grows accordingly. Mitosis occurs simultaneously in all nuclei of the plasmodium, 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 : When an active plasmodium reaches a certain stage, its reand 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- the return of moisture. actions change; modia break up it moves out into the light 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. Harper (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 or of separate fruit many is called a sporangium. sporangia not completely separate, A is fruit of the form of a large mass, 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) Shasta (Cooke, 1949) consist entirely of familiar species. and from Mount Rostafinski (1873) arranged the genera in two cohorts, seven orders, and nineteen 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. tribes, the last first established by Lankester, mostly under names which had applied to tribes. Berlese (in Saccardo, 1888) provided a complete names in -aceae, valid under botanical rules; Poche provided a complete set in Definite families were Rostafinski set of -idae, valid list under zoological of families; here, 1. rules. Authorities somewhat Capillitium none have differed moderately as arbitrarily, fourteen are (order Cribrariales to the maintained. Mac- bride). 2. Producing separate sporangia, pseudocapillitium none. 3. Sporangia shattering irregularly or opening through a terminal oper- culum 3. like 2. Family 1 Family 2. . Liceacea. Sporangia opening through numerous pores, the walls becoming sieve- Fruits aethalioid, Cribrariacea. 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 ] 3. Classification of Lower Organisms Aethalia consisting of more or less separate sporangia. tubular, Sporangia opening through terminal pores 4. Sporangia indistinct, their walls becoming freely punctured and converted into a reticulate pseudocapillitium 3. Aethalia not consisting of distinguishable sporangia Capillitium present. 2. Fruits without considerable deposits of calcium carbonate. 3. Spores black or dark, capillitial 4. 1. smooth Macbride). hairs 4. Fruits Family 3, Tubiferida. Family 4. Retigulariacea. Family 5. Lycogalacttoa. (order Stemonitales capilHtium aethalioid, poorly defined, without a cen- Family tral axis 4. Fruits of separate 6. Amaurochaetacea. sporangia with a definite capillitium including a central axis (columella). Capillitium spreading hor- 5. from the colu- izontally 5. mella Capillitium 7. Stemonitea. Family 8. Enerthenemea. Family 9. Margaritida. Family 10. from the summit of chiefly the columella 3. Family spreading Spores pallid or yellow (order Trichiales Macbride). hairs smooth, unbranched or sparsely branched. 4. Capillitial 5. Capillitial threads attached zontal, at hori- both ends 5. Capillitial 4. threads run- ning at random, not attached at the ends Capillitium reticulate, sculptured, but not with spiral bands. 4. Capillitial or .Family 11. Arcyriagea. threads unbranched sparsely branched, sculp- tured with spiral bands 2. . Perichaenacea. Family 12. Trighiagea. Family 13. Physarea. Fruits containing considerable deposits of calcium carbonate (order Physarales Macbride). 3. Calcium and carbonate both in walls in capillitium Phylum Protoplasta 3. Calcium carbonate in walls [175 but not in capillitium Family 14. DrovMiACEA. 1. Liceacea [Liceaceae] (Rostafinski) Lankester in Enc. Brit. ed. 9, 19: (1885). Tribe Liceaceae Rostafinski Vers. 4 (1873). Order Liceaceae Lister Monog. Mycetozoa 149 (1894). Family Liceidae Doflein 1909. Family Orcadellidae 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- Family 841 litium or pseudocapillitium, the walls shattering irregularly or opening by means of a terminal operculum. Licea, Orcadella. Cribrariacea [Cribrariaceae] (Rostafinski) Lankester 1. c. Tribe Cribcit. 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- Family 2. rariaccac Rostafinski op. 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 Reticulariaceae Lister op. cit. 156. Family Dictydiaethaliidae Poche I.e. Aethalia of indistinct sporangia whose walls become porous and are converted into a reticulate pseudocapillitium. 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 Saccardo 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 Amaurochaete. Family 7. Stemonitea Lankester in Enc. Brit. ed. 9, 19: 841 (1885). Tribes Stemonitaceae 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 Brefeldiidae and Stemonitidae Poche op. cit. 202. Sporangia with dark spores and a capillitium of smooth threads spreading from a central axis, the columella. Stemonitis, 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 Enerthenemaceae 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 axis. been included, but the capillitium attached chiefly at the summit of the columella. Enerthenema, Clastoderma, Lamproderma, Echinostelium. — The 176] Classification of Lower Organisms Margaritida [Margaritidae] Doflein 1909. Order Margaritaceae Lister 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 Arcyriaceae 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- Family op. cit. 9. 202. Family Mycetozoa. a-f, Spore, germination, gametes, syngamy, and zygote of Fig. 33. 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. r-W, fruits of Myceq, Capillitium and spores of Lepidoderma Chailletii x 1,000. ( toza X 5; U, r, Sternonitis splcndens; Lepidoderma Chaillettii; v, s, Lycogala cpidcndrum; i, Lcocarpus notabile; w, Hemitrichia intorta. Physarum fragilis; Phylum Protoplasta [177 c; the genus Trichina does not belong to this family! Order Trichiaceae MacAm. 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. ter 1. bride N. Family 13. Spumariaceae Spumariaceae N. Am. Slime Physarea Lankester Rostafinski op. 1. c. cit. 9, 13. Tribes Cienkowskiaceae, Physaraceae, and Families Cienkowskiaceae , Physaraceae, and cit. 328, 329, 387. Order Physaraceae Macbride Molds 20 (1899). Family Physaridae Doflein 1909. Fruits sporangial Berlese in Saccardo op. 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 sessile little gray sporangia may be spherical or irregular, or stalked. Fuligo septica produces dirty yellow aethalia reaching several cen- timeters in diameter on vegetable trash; observed on spent tan bark, mon name of flowers of tan. Badhamia, Craterium, Leocarpus, it has the comChondrioderma, Spumaria, etc. Family 14. Didymiacea [Didymiaceae] (Rostafinski) Lankester 1. c. Tribe Didymiaceae Rostafinski op cit. 12. Order Didymiaceae Lister op. cit. 93. Family Didymidae Doflein 1909. Family Didymiidae Poche op. cit. 202. Family Collodermataceae 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, Each spore of the single superficial and ascends upon it before becoming walled. making the spores 4-nucleate; the chromosome number is of secreted material. layer generates a microscopic stalk Meiosis then takes place, cut from 16 to 8 (Gilbert, 1935). In germination, the contents of the spore are re- amoeboid protoplast, whose nuclei divide once; the cell then divides and these generate flagella (Rostafinski, 1873; Jahn, 1905; Gilbert, 1935). leased as a single into eight, Order 3. Phytomyxida Calkins Biol. Prot. 330 (1926). Class Phytomyxini Engler and Prantl Nat. Pflanzenfam. II Teil: Phytomyxinae op. cit. I Teil, Abt. 1: iii (1897). Order Phytomyxinae Campbell Univ. Textb. Class Plasmodiophorales Engler Syllab. ed. 3: 1 (1889); class Bot. 71 (1902). 1 (1903). Mycologia 34: 115 (1942). Suborder Plasmodia phorina Hall Protozoology 228 (1953). Order Plasmodiophorales Sparrow in Intracellular parasites chiefly of higher plants, attacking also algae, Oomycetes, beetles, being naked multicellular plasmodia producing walled resting cells, and — — The 1781 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-production 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 myxomycetes 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 uninucleate 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 become 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/Tnui wuium/^iiiMiivnirm- Fig. 34. Ceratiomyxa jruticulosa. a. Fruits x 5. b-q, reproductive processes 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. after Gilbert (1935); b, Phylum Protoplasta In the growing plasmodium, a nucleus which is [179 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 "cruciform" figure resembles some which have been seen in trypanosomes. The nuclear divisions which occur immediately before cleavage are of a different character: no endosome 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 Plasmodiophoraceae Engler Syllab. 1 (1892). Family Woroninaceae Minden 1911. Families : Phytomyxidae and Woroninidae Poche in Arch Prot. 30: 198 (1913). Plasmodiophora, 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 Mycetosporidium, 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). the pseudopodia of the character of rhizopodia, i.e., fine, branching and anastomosing; producing shells, these usually calcareous; commonly 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- Amoeboid organisms, freely inally applied in the categoiy of orders. The 180] Fig. 35. —Life Myers (1943); Classification of Lower Organisms i.e., Discorbis or Cymbalopora, from microspheric individuals, in 3 releasing young megalospheric cycle of "Tretomphalus," 1-3, individuals; 4-8 megalospheric individuals; 9-12, gametes and syngamy. Phylum [181 Protoplast a individual rhizopod originates as a minute amoeboid cell which secretes a 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, The shell having a cylindrical or irregular shell, enlarges this as of the group, the original shell, called the proloculus, it is grows. In the great majority of definite size and form and has a constricted orifice. When the protoplast reaches a certain stage, it expands, protrudes 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 transmitted 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, 1885, designated the smaller and larger proloculi reand megaspheres. Lister (1895), by study in culture of Elphidium crispiirn [Polystomella crispa Lamarck), showed that the two forms are alternate 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 jointly with Schlumberger, spectively microsperes minute biflagellate cells. Schaudinn (1902) confirmed much of what Lister had observed. in describing He was nuclear division (except just before the production of the mistaken 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 completed by its constriction. The spindle is blunt-ended; there is no evidence of centre- 182 The ] Classification of Lower Organisms The chromosomes are numerous, long, and slender; the mitotic figures resemble 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 producsomes. 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 had identified meiosis as occurring at the time of gametogenesis; it is the on the contrary, that it occurs in the last two nuclear divisions in the microspheric 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 coccf animals, fact, 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 coliths. 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 magnification 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 Economic micropaleontologists numbers of forms which are slightly, yet signifiexpedient not to name these, but to identify them by tens of thousands. find themselves dealing with great cantly, distinct. They find it 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 (foramina) but by cylindrical tubes. Dujardin ( 1835) found that his Rhizopodes are without 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 protoplasm. The effect of his discoveries was to [ 183 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 modern period of practical concern with the group, has been that of system, in the 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 followed 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 2. Not 3. Not 2. Miliolidea. Order 3. Foraminifera. Order 4. GLOBiGERiNroEA. Order 5. Nummulitinidea. specialized as in the following orders 3. Order as above. Shell hyaline, perforate, trochoid, having i.e., sively larger chambers the typically succes- helically ar- ranged so that all may be seen from one side and only the last whorl from the other 3. Order 1. Chambers fundamentally planispiral shell with specialized walls containing channels or producing chamberlets of the Monosomatia (Ehrenberg) Siebold Anat. 1: 11 in Siebold and Stannius Lehrb. vergl. (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 Lankester (1885). Family Astrorhizidaceae Lister. Families Rhizamminidae , Saccamminidae, and Hyperamminidae Cushman. Shell of agglutinated foreign material, usually 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. elongate, often branched, but not coiled. In Astrorhiza there The 184] Classification of - Fig. 36 bralina. lina. noted. j, — Shells of Rhizopoda. e, Peneroplis. Lagena. k, 1, a, ^ Lower Organisms / m\^^V^//'":^^^^^.^V^\\•\\ Ophthalimidium. h, c, Triloculina. d, Verte- Archaias x 25. g, Nodosaria. h, Dcntilina. i, FlabelNonion. m, n, Rotalia. o, Globigerina. x 50 except as f, Phylum Protoplasta Family Spirillinidea Reuss 1861. Family Spirillinina Lankester (1885). Family 3. Silicinidae Cushman. In Spirillina, the perforate planispirally coiled; the family stages if not throughout life. Patellina the spirally coiled Family 185 [ 4. Ammodiscida is hyaline one-chambered shell is distinguished by a shell of this form in the young a Jurassic fossil whose shell is silicified. In followed by others arranged in a heHx. [Ammodiscidae] Rhumbler 1895. Like the preceding Silicina first is chamber is 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, Miliolina, and Hauerinina Lankester (1885). Fisherinidae Cushman. The genus Cornuspira, known from the carboniferous, differs from Spirillina only in the texture. Evidently 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 chambers 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 Keramosphaeridae Cushman. Another group of specialized derivatives, the planispirally coiling chambers broadened and divided into many chamberlets with separate apertures, more or less globular. Borelis; Fasciolites Parkinson 1811 [Alveod'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. the entire body lina 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 19: 847 (1885). in Enc. Brit. ed. 9, The 186] 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- 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 Elphidium 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 lanous; not usually of trochoid form, ( 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 apparently 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 Trochamminina Lankester (1885). Family Plocapsilinidae Cushman. Cells planispiral or trochoid, becoming evolute or irregular; walls with imbedded grains of sand. Pennsylvanian 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 Loftusiidae 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 Vcrneulinidae Cushman 1927. Family Ataxophragmidae Galloway (1933). Having walls of agglutinated material and allied to the preceding families; chambers of the tending to form an elongate, screw-like spiral. Valvulina, Ataxophragmium, etc.; since early Mesozoic, abundant in the present. Family 8. Textularina Ehrcnberg (1839). Family Textularidac d'Orbigny (1839). shell Verneulina, 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 becoming curved or straight in the majorbecoming 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; Glandulina is shaped like a jug. Nodosaria is like a row of enlarging beads. Lagena is a onechambered form connected to Nodosaria by transitions, and evidently reduced, not ity; orifice in the earliest forms, ordinarily of radiating slits, primitive. Family 10. Polymorphinida [Polymorphinidae] d'Orbigny. Families Polymorphin- ina and Ramulinina Lankester ( 1885). Specialized irregular forms related to the preceding, 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, planispiral with successively larger chambers, a few of the highest trochoid; walls hyaline, N perforate; aperture generally a transverse slit. onion Montfort {Nonionina d'Orbigny) and Elphidium Montfort [Geophonus Montfort, Vorticialis Lamarck, Polystomella Lamarck, the apparent type of Foraminifera) are simply nautiloid; Hantkenina 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- and irregular forms, Piano pulvinulina, etc., and moderately specialized ones with conical or disk-shaped bodies of numerous chambers, Cymbalopora, Planorbuerate 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, sometimes 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, Asterigerina and Amphistegina, diverging from Rotalina in having each chamber divided two by an oblique wall. Doubtfully in the Cretaceous; Eocene to the present. Family 5. Chapmaniida [Chapmaniidae] Galloway (1933). The numerous chambers arranged in a low cone whose inside is filled with deposited solid material. Chapmania, Halkyardia, Dictyoconoides. Eocene and Oligocene. Family 6. Chilostomellida [Chilostomellidae] Brady (1881). Family Chilostomellaceae Lister. A few genera of reduced derivatives of Rotalina with few chambers. Allomorphina, Chilostomella, Sphaeroidina, etc. Jurassic to the present. into 188 The ] Classification of Lower Organisms Family 7. Orbulinida Schultze 1854. Family Globigerinida Carpenter 1862. few genera with the chambers mostly few, subglobular, clustered rather than arranged in a definite pattern. Orbulina. Globigerina, abundant, pelagic in all oceans, the shells abundant in the ooze on the bottom. Pennsylvanian, doubtful; Jurassic, rare; A 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, ina. Common, Mucron- 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 and comma-shaped orifices, evidently derived from the forego- high-spiralled shells ing family. Cassidulina, etc. Eocene to the present. Family 12 Uvigerinida [Uvigerinidae] Galloway and Wissler, 1927. Further varifrom 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. Pleurostomella, Nodosarella, Daucina, Ellipsoidina, etc. Cretaceous to the present, commonest in upper Cretaceous and Eocene. ants 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] chambers short and broad, numerous, in Moller 1878. Carboniferous fossils, a planispiral coil, forming bodies the which are usually fusiform or globular. Orobias, Fusidina, Triticina, Verbeekina, etc. Nummulitida [Nummulitidae] Reuss 1861. Family Camerinidae 1865. Family Nummulinidae Brady (1881). Family Nummulitina Lankester ( 1885). Family Nummulitaceae Lister. Mostly disk-shaped, planispiral, the walls not highly specialized. Camerina Bruguiere 1792 {Nxunrnulites Lamarck Family 2. Meek and Hayden 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 Carpenteri Brady. Phylum Protoplasta Class 4. HEUOZOA [ 189 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. Subclasses Proteomyxiae, Heliozoariae, Traite Zool. 1: 66, 156, 9, 19: 839 (1885). and Radiolariae Delage and Herouard 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 observations of these organisms has been scant and scattered; they remain poorly known. The Heliozoa as conventionally construed are also water. Ehrenberg observed cilia. some of them and took them mostly inhabitants of fresh for Infusoria with immobile There are only a few dozen species of Heliozoa sensu is no more than a reasonable order. strict o (Schaudinn, 1896) : the whole group The Radiolaria (this marine. Examples were name first also used at this point in its conventional sense) are 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 sublegions, twenty orders, 85 families, 739 genera, and more than four thousand species. The categories, Haeckel explained, are purely relative: Radiolaria would as well be . . The 190] Classification of . Lower Organisms a phylum, the legions classes, and so forth. This idea served him as license for confounding 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, a firm membrane surrounding i.e., without the inner part of the protoplast 1. Order 1. Radioflagellata. Order 2. Radiolaria. Order 3. Acantharia. Order 4. Monopylaria. Order 5. Ph.'SlEosphaeria. Cells with a central capsule. 2. Central capsule of spherical symmetry or with three planes of right angles, punctured 3. symmetry at by many pores. Pores of the central capsule evenly distributed; skeleton absent or present, without spicules which cross the central capsule or 3. meet in its center Pores of the central capsule clustered; skeleton including spicules which cross the central capsule or meet in its middle 2. Central capsule of radial symmetry, with one opening 2. Central capsule of isobilateral symmetry, with one main opening and two minor ones 225 ( 1880) 1. Radioflagellata Kent Man. Inf. 1 Subdivision or subclass Heliozoa (Haeckel), and orders Aphrothoraca (Hertwig), Chlamydophora (Archer), Desmothoraca (Hertwig and Lesser), and Order : 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 Zoosporidia; 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 without skeletons. The type, being the only genus assigned Actinomonas. 1. Pscudopodia unspecialized; amoeboid organisms with or without flagellate stages. to the order by Kent, is Phylum Protoplasta 2. Without 2. With [191 Family shells shells; Pseudosporea. 1. without known flagellate stages. 3. Shells chitinous, without siliceous Family scales 3. Shells bearing circular scales 1. 1. Pseudopodia slender, with apical knobs Pseudopodia of the character of typical axopodia, without apical knobs; the cells or their main bodies 2. Lagyntoa. siliceous Family Family 3. 4. Euglyphida. Vampyrellacea. Family 5. AcTiNOMONADroA. Family 6. AcTiNOPHRYroA. Family 7. Heterophryida. Family 8. Acanthocystida. usually regularly spherical. 2. Bearing flagella as well as axopodia in 2. Without the vegetative condition flagella in the vegetative condition. 3. Cells without a lifeless outer coat 3. Cells having a gelatinous outer coat 3. without siliceous spicules Cells having a gelatinous outer coat with siliceous spicules 3. Cells with a hard shell punctured by pores Family 9. Clathrulinida. 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 Family 1. . 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- in and Diplophysalis. In other genera, as Arachnula and Chlamydomyxa, no 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 spora. flagellate stages are 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). Monadineae 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. Vampyrclla, 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 t)v: %j^ h \ 37. Radioflagellata a-f, Diplophysalis stagnalis after Karling (1930); 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. Fig. a, b, : Phylum Protoplasta [ 193 Organisms bearing at the same time flagella and typical axopodia. Dimorpha, free-swimming, with two unequal flagella. Actinomonas, Pteridomonas, Ciliophrys, 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 Doflein. 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; Actonois 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 spaerium body of cytoplasm. Schaudinn and Belar (1923) observed conjugation. 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 spindle-like Pairs of gametes, 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 diatom shells imbedded in the envelope. Actinolophus is stalked. Sphaerastrum becomes colonial by incomplete division of the cells. Family 8. Acanthocystida [Acanthocystidae] Glaus 1874. Chalarothoraca Hertwig and Lesser in Arch. mikr. Anat. 10 Suppl. 193 ( 1874). Chalarothoraca seu Acanthocystidae 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 Pinaciophora, disks bearing a central spine which is in some species forked. The cell of Wagnerella (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 granule located outside the nucleus (in Wagnerella, in the head). Schaudinn (1896) reported 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 production of buds. The buds may lose their pseudopodia and develop paired flagella. It is . 194 The ] Classification of suspected that the flagellate cells may Lower Organisms be gametes. The central granule is said to 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 originate by extrusion 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 lying within globular shells, apparently of chitin, usually stalked, punctured by numerous 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 Jenaische 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 construed, 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 siliceous, 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 Thalassicolla; 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 capsule membrane and by capsule or meeting in its [ 195 the absence of skeletal spicules extending across the central middle. Except during reproduction, each central capsule contains a single nucleus, but the cells of many central capsules. many examples are coenocytic, containing several or Brandt (1885, 1905) observed reproduction particularly as it occurs by the producswimming cells by some of the coenocytic forms. The nucleus divides to produce very many, and the intracapsular cytoplasm divides to produce uninucleate tion of flagellate cells. In Collosphaera, all the nuclei are included in these cells. are of two The cells 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 appears 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 recogsizes, 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 numerous small spicules. Sphaerozoum, the cells globular, to 1 mm. in diameter; Raphidozoum, 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 Sphaeroidea, 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 Haeckel (1887). The skeleton with three unequal axes, or spiral. Cenolar- families, 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). — The 196] Classification of Lower Organisms f 0'^^' *lty -S^'^v"^ ^^ ^-'v-^if**^^^^ >o, Fig. 38. Radiolaria: b. Skeleton of Haliomma a, ::Q)t^^a ^/-T motile cells of Collosphaera Huxleyi after Brandt ( 1885). c, Skeleton of Actinomma Haeckel ( 1862) capillaris after Asteracanthion after Haeckel (1862). . d. Skeleton of //f/zorfz,?cu5 P/zaco^wcu^ after Haeckel (1887). Acantharia: e, Skeleton of Dorataspis costata after Haeckel Monopylaria: f, Central capsule of Tridictyopus elegans after R. Hertwig (1387). h. Skeleton f 1879) g, Skeleton of Lithocircus productus after R. Hertwig, op. cit. of Eucyrtidium carinatum after Haeckel (1862). Phaeosphaerl^: i. Typical central 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 pattern 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 elaborate, 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 acanthin. 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). Spicules numerous, radiating, not arranged according to Miiller's law. Litholophus, Astrolophus, 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). Family 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 radiating spicules forming a globular network, or two or more concentric networks. Dorataspis, 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. The 198] 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 Nassoidea, Plectoidea, Stephoidea, and Cyrtoidea, Haeckel Radiolarien Teil H (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 Halicryptina and Lithochytridina Ehrenberg 1847. Family Cyrtida Haeckel (1862). Order Cyrtoidea, with twelve families, Haeckel (1887). Skeleton a more or less basket-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 Order 5. more longitudinal grooves Phaeosphaeria Haeckel into as in Sitzber. Jenaische Gess. many lobes. Med. Naturw. 1879: 156(1879). Phaeodariae, with orders Phaeoconchia Haeckel op. Phaeocystia, cit. Phaeogromia, Phaeosphaeria. and 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 membrane; 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 cytoplasm 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 bowlshaped, and finally globular, after which nuclear membranes form about them. While the nucleus divides, the central capsule membrane becomes constricted by longitudinal 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 cytoplasm. 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 reports, in part illustrated with photographs, of similar processes in family Thalassicollida (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 symmetrical, or distinctly dorsiventral. — The 200] 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 Chal- pitted. lengeron. Family Medusettida Haeckel (1887). Shell smooth or with small Medu- spines. 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 carolinensis by Wilson (1900); c, x 25, after the original figures of Pelomyxa 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 Sarkodina 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 presence of shells. is obviously and superficially divisible into two by the absence or 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 pseudoplasmodial communities. 2. Free-living 2. 1. Entozoic 3. Amoebaea. MAYORELLroA. TnECAMOEBroA. 4. Hyalodiscida. 5. ENDAMOEBroA. Family 6. Labyrinthulida. Family 7. Guttulinacea. Family Family Family Family Family 1. 2. Protoplasts assembling and acting in unison in pseudoplasmodial communities. 2. Parasitic in plants 2. Predatory on bacteria, in air on moist surfaces; mostly producing complicated fructifications Amoebaea Ehrenberg Infusionsthierchen 125 (1838). Family Amoebidae Bronn 1859. Family Chaidae Poche in Arch. Prot. 30: 171 (1913). Family Family 1. Chaosidae Chatton in Grasse Traite Zool. 1, fasc. 2: 58 (1953). The ordinary freeamoebas. SchaeflFer (1926) limited the family to forms which produce numerous indefinite granular pseudopodia. There has been much confusion as to the identity 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 living 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 divergens Bory Diet. Class. Hist. Nat. 1: 261. 1822; Amoeba Proteui 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 Proteus; Amiba Leidy). It 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 peculiar twisting of the spindle of Chaos Protheus. Schaeffer included in the present family three further genera, Trichamoeba Fromentel, 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 different 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( , 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 nonflagellate 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 fia. (1934). Similar to the foregoing but without the tough pellicle. Commonly domeshaped, 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 desmose is is apparently no centrosome; at least, no intra- seen (Kirby, 1927). Entamoeba Casagrandi and Barbagello, named at nearly the same time as the foregoing and regrettably similarly, is widely distributed in invertebrate and vertebrate hosts. E. dysenteriae (Councilman and Lafleur) Craig {Endamoeba histolytica Schaudinn) 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. Mitosis begins with division of the centrosome into two, which remain connected, as they draw apart, by a stainable strand, the intradesmose (the term The karyosome is of Kofoid and Swczy, breaks up into chromosomes, six in the species mentioned. Spindle fibers connecting these to the centrosomes have been seen; Child (1926) 1921). Phylum Protoplasta [ 203 found that the two halves of the spindle swing apart as the centrosomes move apart compass being extended. There is no doubt that Endamoeba and like the legs of a 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 different 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 Guttulineen 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 activity 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- and Coenenia produce slightly more elaborate 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, lar cluster of spores; Polys phondylium fruits of the same general nature. In Dictyostelium, The 204] — Classification of Lower Organisms Labyrinthula as a parasite in cells of Ectocarpus Mitchelliae x 1,000 (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, Sappinia 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 Fig. 40. after a, Karling . : o, p. Fruits of 1, m, again heaps itself up and becomes a fruit, n. Dictyostelium mucoroides; q, of Polysphondylium violaccum; x 10, after Bonner, light op. and creeps; 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 afterward creeps some distance in the same direction. When this has happened, the foremost 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. Lampramoebae Haeckel Gen. Morph. 2: xxiv (1866). Testacea Schultze 1854, non L. (1758). Thecamoebae Haeckel. Conchulina Cash and Hopkinson British Freshw. Rhizop. 1: 37 (1905). Suborder Testaceolobosa de Saedeleer in Mem. Mus. Roy. Hist. Nat. Belgique Order 2. Order Order Order 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 associated with these, have here been placed among Rhizopoda or Heliozoa, by de Saedeleer (1934) and Grasse (1953). Deflandre (in Grasse, op. as suggested cit.) distin- guishes several families beside the following: 1. Shell without secreted scales of silica. Family 1. Arcellina. Family 2. Difflugiida. 1. Shell with secreted scales of silica Family 3. NEBELroA. Family 1. Arcellina Ehrenberg Infusionsthierchen 129 (1838). Family Arcellidae 2. Shell of uniform secreted material 2. Shell with imbedded grains of sand 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 groups: (a) the gregarines, first by Leuckart included the following described by Dufour (1826) as name Gregarina Dufour (1828) beetles: the generic crowds; (b, c) coccidians and psorosperms, worms parasitic in refers to their occurrence in different sorts of parasites discovered in 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^osema 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- fishes erous. The number of species duly some two or three thousand; this is registered by name and description surely a small fraction of the is apparently number which exist. 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 subordinate groups, the first including the gregarines and coccidians, the second including the organisms which were formerly called psorosperms (Myxosporidia or Neosporidia). 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 transmission of disease by biting arthropods was first 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 the 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 possible exception of is natural. The classes are distinguished as follows: Phylum 1. 1. Fungilli [ Producing resting cells protected by cell walls and not containing polar capsules; or not producing resting cells Producing resting cells whose walls consist (at least usually) of modified cells, and which contain "polar capsules" enclosing coiled threads Class 1. SPOROZOA Class Sporozoa Leuckart Parasiten der 207 Class 1. Sporozoa. Class 2. Neosporidia. Leuckart 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, Coccidiomorpha, 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 parasitic 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 producing 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 apparently 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 centipede 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: The 208] Classification of Sporozoite, the original infective Lower Organisms 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; 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. b-d, developing trophozoites; e, schizogony; f, Phylum Fungilli [ 209 monocystid gregarines, Muslow (1911) and Calkins and Bowling (1926) described 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 immediately after karyogamy. They conclude that all nuclei except those of zygotes are haploid, 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 a reduction of the chromosome gametes. The Sporozoa are classified primarily by whether or not the trophozoites are intraby 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. cellular; 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 3. 3. not attached in pairs. Producing walled spores Not producing walled spores. 4. Intracellular in erythrocytes Producing macroscopic bodies in muscle The gametocytes pairing before gametogenesis; sperms few; with or without walled spores Order 1. Oligosporea. Order 3. GYMNOSPORiDnoA. Order 4. Dolichocystida. Order 2. Polysporea. Order Order 5. 8. Schizogregarinida. Haplosporidhdea. Order 6. Monogystidea. Order 7. Polycystidea. 4. 2. 1. Gametes tiated, slightly differentiated or undifferen- produced by the gametocytes in more or less equal, usually large, numbers. 2. The organisms multiplying also asex- ually. 2. 3. Spores producing several sporozoites. 3. Each spore producing one The organisms 3. sporozoite. Cells elongate parts . . . . . not multiplying asexually. Cells not elongate and divided into two parts 3. . and divided into two 1. OUgosporea Lankester in Enc. Brit. ed. 9, 19: 855 (1885). Tribe Monosporees and groups Disporees and Tetrasporees Schneider Zool. Exp. Gen. 9: 387 (1881). Order in Arch. The 210] 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 Eimerioidae, 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 vertebrate 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 centipedes, has spores whose walls split lengthwise into two valves. The zygote of Isospora, 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 binuclcata, 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 family 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 [211 Fungilli 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 reproduction 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 Protozoology 296 (1953). It is characteristic of this order that pairs of reproductive cells, essentially merozoites, which are to become gametocytes, become attached to each other. The macrogametocyte 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 Adelea and Adelina chiefly in centipedes; Klossia and Orcheobius in sporozoites. 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. Tropenkrankheiten 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 Protozoology 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 arthropods. The gametocytes do not become associated in pairs; the male gametocytes produce 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. The 212] Classification of The name Haemosporidia, commonly Lower Organisms 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 nature as the Aggregatida, Adeleida, and Haemogregarinida. — a-m. Life cycle of Plasmodium compiled from various sources: a, infecan erythrocyte by a sporozoite; b-e, trophozoites, plasmotomy, and mero- Fig. 42. tion of spermatocyte; g, oocyte; h, production of sperms; i, fertilization; j, k, production of sporozoites in cells of the gut epithelium of the mosquito; 1, sporozoites; n-q. Stages of division sporozoites entering the salivary gland of the mosquito, zoites; f, m of cells of Babesia bigemina in erythrocytes of cattle The Gymnosporidiida Family 1. are organized, somewhat x 2,000 after Dennis (1930). arbitrarily, as three families. Halteridiida [Halteridiidae]Hartmann and Jollos 1910. Family Leu- Hartmann and Jollos. Family Hacmoproteidae Doflcin. Hacmoproteus Kruse {Haltcridium Labbc) occurs in reptiles and birds. Vegetative growth and reproduction 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 cocytozoidae Phylum (Argao, 1908), the alternate host is a fly. Fungilli In the gut of the [ fly, 213 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 comparatively 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 erythrocytes then break up and release the merozoites. The chill and fever of malaria are associated 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 requires 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- The body cavity of the mosquito, migrate to 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 Nomenclature 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 ites. sporozoites are released into the the salivary gland, 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 Marchiafava 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. vivax, the agent of tertian malaria, and H. malariae, that of quartan malaria; it is believed 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; The 214] 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 expedient 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, ton) is by which ticks are infected. Babesia Stercovici [Piro plasma Pat- 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 division, 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 Order 4. this family. Dolichocystida Delage and Herouard Traite Zool. 1 : 289 (1896). Sarcosporidia Balbiani 1882. Class Sarcosporidia Biitschli in Bronn Kl. u. Ord. Thierreichs (1882). Subclass Sarcocystidca Lankester in Enc. Brit. ed. 9, 1, Abt. 1 : Inhalt 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). characters are those of the single family and genus: The 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. Miescheriana. The visible body is a mass The mass originates as host. of cells, the whole walled by modified muscle of the 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 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 the host, they 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. (1929). Orders Archegregarina and Neogregarina Grasse Traite Zool. 665 (1953). 1, 5, 3 fasc. 2: : 872 622, 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. Schizocystis, 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 Caulleryellidae Keilin. Merogregarina, Caulleryella, Tipulocystis. Spirocystida [Spirocystidae] Calkins Biol. Prot. 435 (1925). Family 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 4. Spirocystidees Leger and Family Dischizac Weiser in Jour. Protozool 2: 10 (1955). In Ophryocystis Schneider (Leger, 1907), the vegetative individuals, attached to the walls of the Malpighian grow and become multinucleate and send out branches whose ends develop into additional individuals. Eventually, different individuals become attached 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. tubules of beetles, 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. : ( The 216] The genus which are common is best Classification of known The in earthworms. is Lower Organisms Monocystis Stein, including several species which grow within epithelial cells of the seminal fun- cells 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 nels; they Several observers, as Brasil (1905) and Mulsow (1911); also, as 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 paired cells divide. to related genera, observed by Wolters (1891). The gametes from the respective paired cells are presumably 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). Monocystidae Biitschli 1882). Monocystis, etc. Family 2. Rhynchocystida [Rhynchocystidae] Bhatia in Parasitology 22: 158 ( (1930). Rhynchocystis. Family 3. Stomatophorida [Stomatophoridae] Bhatia op. Choanocystis, Family Family Family Family Family 4. 5. 6. 7. 8. cit. 159. Stomatophora, etc. Zygocystida [Zygocystidae] Bhatia op. cit. 160. Zygocystis, Pleurocystis. Akinetocystida [Akinetocystidae] Bhatia op. cit. 160. Akinetocystis. Syncystida [Syncystidae] Bhatia op. cit. 161. Syncystis. Diplocystida [Diplocystidae] Bhatia op. cit. 161. Diplocystis, Lankcsteria. 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 n..s. 11. Ganymedida [Ganymcdidae] J. S. Iluxlcy in Quart. Jour. Micr. Sci. 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 Subclass Gregarinida Biitschli op. cit. [217 Fungilli 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, The 428 (1926). and divided by more more than two) parts; not typical gregarines, the vegetative cells elongate definite constrictions into two (or, occasionally, ducing asexually. Typical gregarines occur chiefly in or less repro- The vegetative cell consists of an anattachment and a posterior portion (deutomerite), containing the nucleus, lying in the gut cavity of the host. Both parts have insects. terior portion (protomerite) serving for a thick outer layer, less commonly differentiated upon the protomerite elaborate knob, the epimerite. Longitudinal present. The cells fibrils, presumably into a more or contractile, are writhe actively. commonly found in pairs, one member attached to the epithelium 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 The individuals are 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 sporozoites 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 extruded through them in uniseriate rows. In connection with this behavior, the spores cell is left have flat ends like barrels. Family 1. Stenophorida [Stenophoridae] Crawley 1903. Protomerite a mere knob. Stenophora. Gregarinida [Gregarinidae] Greene 1859. Family Gregarinarien Stein 187 (1848). Gregarines which are without epimerites and are not notably elongate. There are about a dozen genera. Cysts without sporoducts: 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 orthopteran. 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. Family 2. in Arch. Anat. Phys. 1948: , cuneata, which Stein observed in cockroaches. The 218] Classification of Lower Organisms Family 3. Didymophyida [Didymophyidae] Wasilewski 1896. Family Didymophyiden Stein (1848). Like the foregoing, but the cells extremely elongate. Didymophyes. Family The Acanthosporida [Acanthosporidae] Labbe 4. in Thierreich 5: 27 (1899). spores with polar or equatorial bristles. Acanthospora. Stylocephalida [Stylocephalidae] Ellis 1912. Family Stylorhynchidae 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, Hoplorhynchus, Pileoccphalus, etc. Family 7. Menosporida [Menosporidae] Labbe op. cit. 29. Epimerite with a long Family Labbe op. 5. cit. branched and bearing appendages. Menospora. Family 8. Dactylophorida [Dactylophoridae] Wasilewski 1896. Epimerite distally broadened, clinging to the host epithelium by means of numerous filiform prostalk, distally Dactylophorus, Nina [Pterocephalus), etc. 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. cesses. Family Order 8. Haplosporidiidea Poche in Arch. Prot. 30: 178 (1913). Order Aplosporidies Caullery and Mesnil 1899. Order aplosporidies Caullery and Mesnil in Arch. Zool. Exp. Gen. H 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 multiplying 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 understood, operculum and releasing the contents. we may suppose If the life cycle is correctly 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 Caullery 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 chiefly [219 Fungilli 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. Myxosporidia Biitschli Subclass Myxosporidia NEOSPOR!D!A (Schaudinn) in Zool. Jahresber. 1880: Biitschli in Bronn Kl. Calkins 162 (1881). Ord. Thierreichs u. 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, 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 by a layer of modified external budding. cells; An entire small plasmodium may become converted into one may be cut out internally and produced continually. The spores, or the spores or two spores, 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 converted into the structures called polar capsules. These resemble the nematocysts of they contain a coiled hollow thread capable of swift extroversion. coelenterates Extroversion occurs during germination. Its significance is unknown. The presence at least in the : of polar capsules marks the class as a natural group. Three orders are recognized: 1. Spores covered by two valves formed from accessory cells 1. accessory cells 1. Order 1. Phaenocystes. Order 2. Actinomyxida, Order 3. Cryptocystes. Spores covered by three valves formed from Spores very minute, with a continuous brane mem- Order 1. Phaenocytes Gurley in Bull. U. S. Fish Comm. 11 Order N^maiocj^^ffrfa Delage and Herouard Traite Zool. Order Phaenocystida Labbe in Thierreich 5 85 (1899). : : 410 (1893). 1: 291 (1896). The 220] 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 animals, 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 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. the order. Phylum The spore-forming [221 Fungilli structure (sporoblast) is a protoplast with several nuclei; it is whole small plasmodium, or half of one, or a protoplast cut out endogenously 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 reduction 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 multinucleate. The two haploid nuclei of the spore, which unite after germination, are derived from a single diploid nucleus. Authors who have described fusions of protoplasts, 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 either a 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 suborder. Myxoceros nom. nov. [Ccratomyxa Thelohan 1892, non Ceratiomyxa Schroter ever names are homonymous without being 1889; if Some thirty-five species; the type is absolutely identical, these are.) M. sphaerulosa (Thelohan) comb, nov.; studied mitosis in M. Blennius (Noble) comb. nov. Leptotheca, Myxoproteus, Noble 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 capSphaerospora, Sinuolinea. C. Valves saucer- or boat-shaped, spores disk-shaped or fusiform sules. (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 into The ] Classification of Lower Organisms an individual whose one or two nuclei remain undivided until the commenceof the ill-understood process by which the complicated spores, with three valves ment 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 Euactinomyxidae 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 considerable horns, the whole horse-shoe shaped. Triactinomyxon and Hexactinomyxon, all three valves drawn out into long horns, the whole caltrop- or anchor-shaped. Order Cryptocystes Gurley in Bull. U. 3. S. Fish Comm. 11 : 409 ( Microsporidies Balbiani 1882. Order Microsporidiida Labbe in Thierreich 5: 104 (1899). The parasites of this order attack chiefly arthropods and fishes. asexually and produce serious epizootics. The spores are very minute, 1893). They multiply and the details 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 of the processes by , . the oligochaet Family 3. exceedingly worm Slavina. Mrazekiida [Mrazekiidae] Leger and Hesse 1922. Spores elongate, minute, resembling bacteria. Mrazekia, Octospora, Spironema, Toxonema. Family 4. Telomyxida [Telomyxidae] Leger and Hesse 1910. Telornyxa glugeibody of the larva of Ephemera vulgata, producing ellipsoid spores formis, in the fat with a polar capsule at each end. Chapter XII PHYLUM CILIOPHORA Phylum CIUOPHORA 8. Class Infusoires (Doflein) nomen phylare novum Lamarck Phil. Zool. 1: 127 (1809). Lamarck Anim. sans Verteb. 1: 392 (1815). Class Infusoria Isis 1818: 1008 (1818). Class Polygastrica Ehrenberg Infusionsthierchen p.* (1838). Class Protozoa Goldfuss in 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 Vorticella, the only genus cell. 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 scientific name it has status from its Protozoa, applied to a class in original publication by Goldfuss, of Infusoria. In treating the group as a name, and takes up The as such the Lamarck (1815). The name is a later synonym phylum, one finds it necessary to apply a new application to a class by its name which essential point in the definition of the Doflein applied to phylum is the as a it word cilia. subphylum. Cilia are cell- 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- organs of the same nature as flagella, differing in 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- term ciHum is to be restricted to two things, (a) the swimming organelles and (b) moving fibrils protruding abundantly from certain epitheHal 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 tion of the of the Ciliophora, 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 individual 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 upon part or all of the surface, but bear a variety of strucMembranelles are triangular appendages conciHa; undulating membranes represent long rows; cirri represent are without separate cilia tures which consist of coalescent cilia. sisting of brief tufts. rows of The organisms of class Tentaculifera bear cilia only in the juvenile condition. At maturity they bear extensible tubular structures called which they capture free-swimming ciUates and absorb their The and tentacles, basal granules of the cilia are linked together by a system of fibrils make up by means of contents. fibrils; the cilia the neuromotor apparatus. This term was coined by Sharp, in his study of Diplodinium (1914). The neuromotor fibrils form a highly elaborate network, not connected with the nucleus, as in flagellates, but to a central structure, apparently regulative, called the motorium. The motorium of Diplodinium is a massive 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 struchomologous with those of animals, a cytostome and cytopharynx. The tures are not cytopharynx is a more or less funnel-shaped impression in the cell. It is bounded 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 cytopharynx 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 undergoes 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 laterally 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 convacuole occurs by dissolution of the bounding membranes of gelled protoplasm where the two are in contact, followed by contraction of the membrane of the canal. The proper contractile vacuole discharges by essentially the same mechanism. 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 memtractile brane breaks, and the membrane of the vacuole contracts. Phylum Ciliophora Earlier biologists supposed 225 the contractile vacuole is an excretory mechanism. purely hydrostatic, to rid the cell of the water which constantly entering by osmosis. Marine Ciliophora have no contractile vacuoles. More is [ In probably, function its tliat is many members of the family Opalinoea each cell has many similar nuclei. These from time to time, by mitosis. Cell division takes place independently of nuclear division, by transverse constriction, when a certain size has been reached. divide, In the generaHty of Ciliophora, each cell has one or more nuclei of each of two 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 micronucleus 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: types, macronuclei, Stentor coeruleus 28 Didinium nasutum 16 Chilodon uncinatus 4 Carche 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 different ends of the nuclear cavity. The nucleus becomes greatly elongate and its membrane 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 cells, apparently undifferentiated, join in pairs but maintain their individ- which the uality. The uniting cells Paramaecium, by become attached their ventral or to each other in definite positions: in 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 conjugants, a fixed number of times, in Paramaecium three, in Euplotes four. These divisions 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 Paramaecium, the spindles of the mitotic final nuclear divisions extend through this connection, 226 The ] when Classification of Lower Organisms complete each protoplast contains two haploid nuclei of difsame result is attained, apparently, by the migration of one nucleus of each pair. Karyogamy takes place in each conjugant. The cytoplasmic 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 Tentaculifera has not been much studied, but here also the nuclear changes are as in the generahty of ciliates (Noble, 1932). so that mitosis is ferent origin. In other ciliates the The possibility of conjugation is limited by the occurrence of mating types. Certain 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 early observations current knowledge, then: Paramaecium caudatum includes four mating types divided into two groups; types 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 conjuI gate with any other Euplotes Patella includes with any other. The six heredity of mating types dclian heredity. In mating types is one group; each can conjugate all in not understood. It Paramaecium Bursaria group I, is not a matter of simple Men- 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 macronuclei 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 term cndomixis. It is not possible that this process is the genetic equivakaryogamy. It is, presumably, the physiological equivalent of conjugation in of nuclei the lent of 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 conjugation, may simultaneously undergo autogamy instead of exchanging nuclei. In the normal conjugation of ciliates, the gamete nuclei produced in each cell, being duced sister products of mitosis, are genetically identical; and the zygote nuclei pro- 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 increased by exposure to high temperature that they would continue to divide abnormally 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 reproduced themselves through many generations, and have proved capable eventually of giving rise to normal individuals. Jollos (1913) designated as Dauermodifikationen, They are actually is, enduring changes, modifications of the type described. 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 that 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 artificially 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 produced after division or excystment. The less elaborate ciliates, as Opalina and 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 Paramaecimn, are usually 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 significant 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 considerable 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 Class Ciliata Haeckel Gen. 1. INFUSORIA Lamarck 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 about the cytostome With a spiral band of membranelles or cilia about the cytostome. or 2. cilia 3. The 4. Not The Holotricha. Order 3. Heterotricha. Order Order 4. Hypotricha. Stomatoda. of the character of the fol- Flattened, cirri and most cilia confined to the ventral surface 3. 2. spiral sinistrorse. lowing order 4. Order spiral dextrorse Order 5. 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 Opalinidae 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. 169 ( 1867) 2. Holotricha Stein Org. Inf. 2 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. Order : Hist. 1: 137 (1909). Order Holotricha with suborders Anoplophryinea, Gymnostomata, and Hymenostomata Poche in Arch. Prot. 30: 250-255 (1913). Order Holotrichida Calkins Biol. Prot. 376 (1926). Infusoria with differentiated macronuclei and micronuclei, with simple cilia distributed 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 families, 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. 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 a. 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 confined 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 Loxodes, mouth at the base of this. Trachelius, Dileptus, Lionotus, etc. Family Chlamydodontida [Chlamydodontidae] Glaus 1874. Family Chlamydodonta Stein, the mere plural of a generic name. Family Chilodontida Biitschli. Family 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 AnoplophryINEA Poche; suborder Astomina Hall. Family Anoplophryida [Anoplophryidae] and seven other Cepede, 1910. families, all named by 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. Order : : 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 peculiar as to appear to have evolved only once: in short, the order appears natural. There are numerous subordinate groups. Several of these, of character, have been segregated as additional orders; many it is species or of exceptional by an arbitrary decision that they are here treated as suborders. 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, parasitic 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 a. area anterior, more or less transverse. Stcntor, sessile and obconic, familiar. Follicu- Phylum alio phor [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 monographed 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, Spirodinium, 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 polysaccharide carbohydrate. Animals are infected by eating food contaminated with the saliva of others. The 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 carrying 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 Grobciliates — The 232 Classification of Lower Organisms Infusoria, order Hypotricha: a, Aspidisca x 800. b, Stylonychia 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 conjugation 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 Fig. 44. X 400. C, undergo dissolution. Phylum Ciliophora ben. Spirochona marine, best [ 233 and a few other genera, attached to aquatic animals, fresh-water or the crustacean Gammarus. known from 4. Hypotricha Stein Org. Inf. 2 168 ( 1867) Section Hypotricha Biitschli in Bronn Kl. u. Ord. Thierreichs 1: 1674 (1889). Order : Order Hypotrichaceae Hartog in Cambridge Nat. Hist. Order Hypotrichida Calkins Biol. Prot. 389 (1926). 1: 137 (1909). 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 ceding order. is evidently natural, and evidently a specialized offshoot from the pre- It might reasonably be treated as a subordinate group of the preceding 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 Oxytrichidae Kent. Family Pleurotrichidae Biitschli. Cirri present; cilia in one or two marginal rows, few or absent on the ventral surface. Oxtricha, Stylonychia, Pleurotricha, order; Biitschli, 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 individual with a larger attached one. Vorticella names Stomatoda and is the apparent type of the old ordinal 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 microscopic 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. Ciliata, — The 234] Fig. 45 Lower Organisms Tokophyra Lemnarum after A. Noble (1932) a, representative individswimming bud; d, conjugation; e, feeding on a cell of Euplotes; tentacles, feeding, expanded, and contracted, g x 2,000, all others x 400. ual; b, budding; c, t, cyst; g, Classification of : Phylum Ciliophora 235 [ Family Urceolarina Perty (1852). Family Trichodinidae Glaus. Family Urceolaridae 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 micromacronucleus may extend to all Asexual reproduction is by budding, often endogenous. Conjugation occurs either between attached individuals or between an attached individual and a swimming 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 remainder may be construed as a single order: nuclei; in branching or colonial individuals, a single parts. Order Suctoria Kent (1880). Lankester chose which Kent published at the same time. a. this as between two ordinal names 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 Lachmann (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. 4. Dendrosomida [Dendrosomidae] Kent Man. Inf. 2: 215 (1882). Family Dendrosomina BiitschU (1889). Family Dendrosomatidae Poche (1913). Dendro- Family soma, etc. 236 The ] Family 5. Classification of Lower Organisms Ophryodendrida [Ophryodendridae] Kent Ophryodendron, etc. Dendrocometida [Dendrocometidae] Kent I.e. Family Ophryodendrina I.e. Family Dendrocometina Biitschli (1889). Family 6. Biitschli (1889). c. With Family 7. Dendrocometes, Stylocometes. differentiated tentacles for piercing Ephelotida [Ephelotidae] Kent. and sucking. Family Ephelotina Sand 1899. I.e. 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 LIST N omenclatural Novelties [ 237 OF NOMENCLATURAL NOVELTIES Page P'amily Kurthiacea fam. nov Family Pasteurellacea nom. nov Family Chromatiacea nomen familiare novum Family Rhodobacillacea nom. nov Family Chlorobiacea nom. nov Order Sphaerotilalea nom nov Lagenocystis, nom. nov., and L. radicicola, comb, nov Family Dinamoebidina nom nov Phylum Opisthokonta phylum novum Chilomastix hominis comb, nov Pentatrichomonas obliqua comb, nov Goussia Schubergi comb, nov Family Myxoceratida and Myxoceros, nomina nova; M. sphaerulosa and M. 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Arch. INDEX OF NAMES OF ORGANISMS AND GROUPS Absidia, 123, 124 Acantharia, 189, 190, 195, 196, 197 Actiniscea, 61, 62 Actinisceae, 62 Acanthochiasma, 197 Actinocephalida, 217, 218 Actinocephalidae, 218 Actinocephalus, 218 Actinolophus, 193 Actinomma, 195 Actinomma Asteracanthion, 196 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 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 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 Adeleoidae, 21 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, Agarics, 152 Aggregata, 209 Ae;gregata Eberthi, 210 Aggregatida, 210, 212 Aggregatidae, 210 Aglaozonia, 88 Agrobacterium, 23 Agrobactcrium tumefaciens, 23 Agrostis, 148 Agyriales, 137 ' 152, 153 The 272] Classification of Amoebina, 201 Amoebodiniaceae, 101 Amoebogeniae, 219 Amoebosporidia, 215 Amoebosporidies, 215 Amoebosporidiidae, 215 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 muscaria, 152 Amaurochaetacea, 174, 175 Amaurochaetaceae, 175 Amaurochaete, 175 Amaurochacteae, 171 Amaurochaetidae, 175 Amaurosporales, 171 diffluens, 37, 157, 201 divergcns, 202 Ammodiscida, 185 Ammodiscidae, 185 Ammodiscus, 185 Ammodochidae, 62 Amoeba, 71, 118, 124, Amoeba Proteus, 202 Amorbaca, 201 Amocbida, 10,201 Amocbidac, 10, 201 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 Anabaena circinnalis, 13 inaequalis, 32 Ancylistales, 81 Ancylistes, 125 Ancylistinaeae, 81 Anemeae, 171 Angiococcus, 28 Angiogastres, 152 Angiospermeae, 82, 91 Animacule, 18 Animal kingdom, Animalia. Animals, 10,' 157, 189, 201 1, 2, 68, 95, 111, 113, 159, 163, 167, 206, 214, 220, 223, 231, 233, 235 Anisochytridiales, 69 Anisochytrids, 57 4, 6, Anisolpidiaceae, 69 Amanita, 152 Amiba Amiba Lower Organisms 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 [273 Index 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 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 1 The 274] Classification of Lower Organisms Bacteriaceae, 21 Bacteriophyta, 17 Bacteroides, 22 Blastodinida, 100, 102 Blastodinidae, 102 Blastodinides, 102 Badhamia, 177 Balantidium, 230 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 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 Blastocladiclla, 112, 113 Blastocladiclla cystogena, 115 Blastocladiineae, 111 Blastodcrma, 130 Bursaria, 230 Bursariidae, 230 Bursarina, 230 1 78 Calcaroae, 1 71 Calcarina, 187 Calcarinidae, 187 Calciconus, 60 Galciconus vitrcus, 56 Cabbage, Calrisolcnia. 60 Calcisolonidae, 60 Callocolax, 50 Callophyllis, 50 [275 Index 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 1 The 276 Classification of 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 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 Lower Organisms 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 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 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 [277 278] The Classification of 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 Lower Organisms 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 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, Cytophagacea, 28 Cytophagaceae, 28 Cytosporidia, 207 Cyttariacea, 135 26, 28 [279 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, Delesseriea, 51 49 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 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 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 Dictydiaethaliidae, 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 varians, 101, 104 Dinamocbidium Lower Organisms 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 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 [281 Elphidium, 186, 187 Elphidium crispum, 181 Elvella, 135 Empusa, 125 Enchelia, 229 Enchelina, 229 Enchelis, 229 Enchelyidae, 229 Endamoeba, 202, 203 Endamoeba Endamoeba 202 202 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 disparita, histolytica, Endosporinei, 171 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 Endothia parasitica, 139 Endothyra, 186 Endothyridae, 186 Endothyrina, 186 Enerthenema, 175 Enerthenemaceae, 175 Enerthenemea, 174, 175 Entamoeba, 202, 203 Entamoeba coli, 202 Entamoeba Entamoeba dystenteriae, 202 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 The 282] Classification of Eunotiaceae, 75 Eunotiea, 75 Eunotieae, 75 Euphorbiaceae, 161 Euplotes, 227, 232, 233, 234 Euplotes Patella, 225, 226, 232 Eupodiscales, 73 Eurotium, 131 Epichrysis, 56, 62 Epiclintes, 233 Epidinium, 231 Epipyxis, 60 Epipyxis utriculus, 54 233 Eremascus, 130 Epistylis, Eremascus albidus, 127 Eremospermeae, 77 Eurychasma, 81 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, Lower Organisms 22 Ethmosphaerida, 195 Euactinomyxidae, 222 Euasci, 130 Eubacteria, 18, 25 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 Eubacteriales, 18 Eubasidii, 145 Euchrysomonadina, 61 Euchrysomonadinae, 61 Eucomonympha, 169 Fasciolites, 185 Fauchea, 51 Faucheocolax, 51 Eucyrtidina, 198 Felis Catus, 6 Eucyrtidium, 198 Eucyrtidium carinatum, 196 Eudesme, 88 Ferns, 125, 148 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 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 Glenodinium, 94, 103 Globigerina, 184, 188 Globigerinida, 188 Frondicularia, 187 Fucaceae, 91 Fucales, 91 Fucea, 91 Fuceae, 91 Fucineae, 91 Fucacees, 82 Fucoidea, 83, 86, 91 Globigerinidea, 183, 187 Globorotalia, 187 Globorotaliidae, 187 Gloeocapsa, 33 Gloeochaete, 33 Gloeochrysis, 62 Gloeodiniaceae, 100 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 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 1 1 The 284] Classification of 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 Haemamoeba malariac, 213 vivax, 213 Hacmogregarina, 2 1 Haemogregarinida, 211, 212 Haemogregarinidac, 211 Haemogrcgarinina, 2 1 Haemoproteidae, 212 Haemoproteus, 213 Haemoproteus Columbae, 212, 213 Hacmosporidae, 211 Hacmosporidia, 207, 211, 212 Hacmosporidiida, 211 Haliarchnion, 49 Halicryptina, 198 Haliomma, 195 Lower Organisms 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 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 [285 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, 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 1 11 The 286] Classification of 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 Kelps, 82, 83, 89, 90 Keramosphaera, 185 Keramosphacridac, 185 Keramosphaerina, 185 Kerona, 233 Klebsiella (bacterium), 7, 22 1 Lower Organisms 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 Latrostium, 69 Laurencia, 52 Leangium, 177 Lithocampe, 198 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 [287 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, 213,230 Margarita, 176 Margaritaceae, 176 Margaritida, 174, 176 6, 159, 165, 210, 7 7 5 288] 1 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 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 Melanophycea, 11, 55, 82 Melanophyceae, 82 Melanospermeae, 82 Melitangium, 28 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 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 Merolpidiaceae, 1 1 Meroselenidium, 215 Mesocaena, 63 Mesogloia, 88 Mesogloiacca, 88 Metachaos, 202 Metadevescovina, 167 Metaphyta, 6 Metasporeae, 1 1 Metazoa, 6 Metchnikovclla, 219 Metchnikovellida, 219 Mctchnikovellidae, 219 Methanomonas, 24 Micrococcacea, 19, 20 Micrococcaceae, 20 Micrococcus, 20 Milioles, 179 Miliolida, 185 Miliolidae, 185 Miliolidea, 183, 185 Miliolina, 185 Mindeniella, 79 Monadineae Tetraplasteae, 191 Monadineae Zoosporeae, 191 Monads, collared, 38 Monas, 38, 54, 59, 60, 158 Monas Monas amyli, 189 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 Monoblcpharideae, 110 Monoblcpharidineae, 110, 111 Monoblepharis, 111, 112 Monocercomonadida, 167 Index 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 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 [289 290] The Classification of 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 Lower Organisms 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 [291 Index Octomyxa, 179 Orbitolinidae, 186 Octospora, 222 Orbitolites, 185 Oicomonadacea, 159, 161 Oicomonadaceae, 161 Oicomonadidae, 161 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 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, 111, 118, 119, 121, 125, 127, 177, 178, 179 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 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 Oxyphysis, 103 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 Paramaecium Paramaecium Paramaecium Aurelia, 226, 227 Bursaria, 226 caudatum, 226 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 The 292] Classification of 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 Pelomyxa carolinensis, 200, 201 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 Lower Organisms 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 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 Phaeocystis globosa, 54 Phaeodaria, 199 Phacodariae, 198 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 Phaeothamnion, 61 Phaeothamnionacea, 58, 60 Phaeothamnionaccae, 60 Phaeozoosporea, 85, 86, 87 Phaeozoosporeac, 86 Phagomyxa, 179 Phalanastrriaccae, 67 Phalanastcriidae, 67 Phalanasterium, 67 Phalanasterium digitatum, 68 Pcritromidae, 233 Peritromina, 233 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 Phallaccae, 155 Phallales, 152 Phallineac, 152 Phalloidca, 155 Phalloidei, 155 Index [293 Phallus, 155 Pines, 148 Phlebotomus, 21 Phleospora, 139 Pinnularia, 72, 75 Pipetta, 195 Phlyctidiacea, 115, 117 Phlyctidiaceae, 117 Piptocephalidacea, 123, 124 Piptocephalidaceae, 124 Piptocephalis, 123, 124 Piroplasma, 214 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 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 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 Lower Organisms 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 [295 Index 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 86 Pyrenomycetales, 138 Pyrenomycetes, 137 Pyrenomycetineae, 137 Pyrgo, 185 Pyrocystis, 100 Pylaiella, 137 Pyronema confluens var. igneum, 127 Pyronemacea, 135 Pyrrhophycophyta, 94 Pyrrhophyta, 39, 94, 182 Pyrsonympha, 166 Pyrsonymphina, 163 Pythiacea, 80 Pyronema, 127, 134, Pythiaceae, 80 135,_ 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 The 296] Classification Rodents, 211 Roesia, 69 157, 158, 160, 178, 192 Rhizomastigaceae, 163 Rhizomastigida, 158 Rhizomastigina, 158, 163 Rhizomastix, 163 Rhizopoda, 6, 63, 157, 172, 179, 184, 200, Rhizoflagellata, 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 Lower Organisms Rickettsiaceae, 20 Rivularia, 36 Rivulariacea, 34, 36 Rivulariaceae, 36 Roach, 166, 168,170 Rhizochrysidae, 63 Rhizochrysidina, 61 Rhizochrysidinae, 61 Rhizochrysidineae, 55 Rhizochrysis, 61, 63 Rhizochrysis Scherffeli, 56 Rhizocryptineae, 95 Rhizoctonia, 142 Rhizodiniales, 99, 101 Rhizodininae, 95, 99 205 Rhizopoda of Rosaceae, 148 Rotalia, 184, 187 Rotaliaceae, 187 Rotalida, 187 Rotalidae, 187 Rotalidea, 187 Rotalina, 187 Rotifers, 113, 118, Rozella, 118 219 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, Sarcocystis Muris, 214 214 Index Sarcodina, 6, 172, 200 Sarcosporidia, 207, 214 Sargassaceae, 91 Sargassea, 92 Sargasseae, 92 [297 Serratia, 22 Siderocapsa, 27 Sideromonas, 27 Siedleckia, 215 Silicina, 185 Sargassum, 93 Sargassum Horneri, 93 Silicoflagellata, 55, 56, 57, 61, 62, 64, 67, 157,200 Schaudinella, 216 Schaudinellida, 216 SchaudinelHdae, 216 Schinzia Leguminosarum, 23 Schizocystida, 215 Schizocystidae, 215 Schizocystinea, 215 Schizocystis, 215 Silicoflagellatae, 55, Silicoflagellidae, 62 Silicoflagellina, 61 Saricodina, 63, 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 69 62 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 The 298] Classification of 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 Lower Organisms 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 Sphaerozoum, 189, 195 Squamarieae, 50 Spirillacea, 19, 23 Spirillaceae, 23 Squids, 210 Staphylococcus, 20 Staurocyclia, 195 Staurojoenina, 169 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 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, 225 Stentoridae, 230 Stcntorina, 230 Stephanida, 198 Stephanonympha, 168 Stephida, 198 Stephoidea, 198 Stereotestales, 62 Stereum, 151 Stictaceac, 134 Stictea, 134 Sticteac, 134 Stictidaceac, 134 Stictideae, 133 Stigonema, 36 Stigoncmataceae, 36 Stiibaceae, 142 Index Stilbeae, 142 Stilbellacea, 142 Stilbellaceae, 142 Stilbosporei, 141 Stilbum, 142 Stilophora, 88 Stilotricha, 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, Syncryptaceae, 59 Syncryptida, 58, 59 Syncryptidae, 59 Syncystida, 216 Syncystidae, 216 Syncystis, 233 Stipitochloridae, 66 Stipitococcacea, 65, 66 Stipitococcaceae, 66 Stipitococcus, 66 Stokesiella, [299 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, 59 216 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 The 300] Classification of Lower Organisms Toadstools, 151 Textularidae, 186 Textularidea, 185 Textularlna, 186 Textulinida, 186 Tokophrya, 235 Tokophrya Lemnarum, 234 Thalamophora, 179 Thalassicolla, 189, 194, 195 Thalassicollen, 194, 195 Thalassicollida, 195, 199 Thallochrysidacea, 62, 63 Tolypothrix, 35, 36 Torula, 130 Torulopsis, 130 Toxonema, 222 Thallochrysidaceae, 63 Tracheliidae, 230 Trachelina, 230 Trachelius, 230 Thallochrysis, 63 Trachelomonas, 94, 106, 107 Thamnidium, 124 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 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, Thiorhodaceae, 31 Treponema Treponema Treponema Treponema 35 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 macrodentium, 29 microdentium, 29 pallidum, 28, 29 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 [301 Index 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, 145 Tulasnellales, 149 Tulostoma, 155 Tulostomataceae, 155 Tulostomea, 155 Tulostomei, 155 Tunicates, 216 Turillina, 188 Turkeys, 210 Turtles, 211 Tuscarilla, 200 Tuscarora, 200 Tuscarorida, 200 24 24 Uncinula, 132, 133 Uniflagellatae, 110 Urceolaria, 235 Urceolaridae, 235 Urceolarina, 235 Urceolus, 109 Uredinacea, 148 Uredinaceae, 148 Uredinales, 145, 147 Uredineae, 147 Ulvina aceti, Umbina aceti, 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 302] The Classification of 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 Lower Organisms 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