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BERGEY'S MANUAL
OF
DETERMINATIVE
BACTERIOLOGY
BERGEY'S MANUAL
DETERMINATIVE
BACTERIOLOGY
BY
ROBERT S. BREED
Ixite Professor Emerilus, Cnrnell Universily, Geneva, New York
E. G. D. MURRAY
Research Professor, University of Western Ontario,
London, Ontario, Canada
NATHAN R. SMITH
Senior Bacteriologist, Retired, Plant Industry Station,
U. S. Department of Agriculture, Beltsville, Maryland
and
Ninety-four Contributors
Whose Names Appear on the Immediately Following Pages
SEVENTH EDITION
BALTIMORE
THE WILLIAMS & WILKINS COMPANY
1957
First Edition, August, 1923
Second Edition, December, 1925
Third Edition, January, 1930
Fourth Edition, March, 1934
Preprint of pages ix + 79 of Fifth Edition, October, 1938
Fifth Edition, April, 1939
Sixth Edition, January, 1948
Seventh Edition, October, 1957
COPYRIGHT ©, 1957
The Williams & Wilkins Company
Made in United States of America
Library of Congress
Catalog Card Number
57-11183
COMPOSED AND PRINTED AT THE
WAVERLY PRESS, INC.
Mt. Royal and Guilford Aves.
Baltimore 2, Md., U. S. A.
LIST OF CONTRIBUTORS
Allen, 0. N., Mr. & Mrs. Rhizobium 285
Andrewes, C. H. Virales 985
Barker, H. Albert Methanococcus 473
Butyrihacterium 577
Zymobacterium 577
Beger, H. Caulobacteraceae 212
Siderocapsaceae 217
Chlamydobacteriales 262
Bier, Otto Calymmatobacterium 418
Borman, E. K. Paracolobactrum 346
Branham, Sara E. Neisseria 480
Breed, R. S.f Introduction 1
Considerations influencing the classification 4
Methanomonadaceae 74
Thiobaderiaceae 78
Pholobacterium 193
Protaminobacter 200
Mycoplana 204
Caulobacteraceae 212
Siderocapsaceae 217
Vibrio 229
Methanobacterium 250
Cellvibrio 250
Cellfalcicula 252
Spirillum 253
Chlamydobacteriales 262
Chromobacterium 292
Alcaligenes 297
Achromobacter 300
Flavobacterium 309
Agarbacterium 322
Enter obacteriaceae 332
Escherichia 335
Aerobacter 341
Klebsiella 344
Serratia 359
Pasteurella 395
Bacteroidaceae 423
Sphaerophorus 441
Micrococcaceae 454
Micrococcus 455
Sarcina 467
Brevibacteriaceae 490
Cory neb acteriaceae 578
Caryophanales 830
Virales 985
Numerous contributions to various other taxa
Broom, J. C. Leptospira 907
t Deceased.
V
VI
Buchanan, R. E.
Burkholder, Walter H.
Campbell, L. Leon, Jr.
Clark, F. A.
Clise, Eleanore H.
Conn, H. J.
Couch, John N.
Davis, Gordon E.
Delwiche, Eugene A.
Doetsch, R. N.
Douglas, H. C.
Dumas, Julien
Elazari-Volcani, Benjamin
Eltinge, Ethel
Evans, James B.
Freundt, E. A.
Gordon, Ruth
Hanks, John H.
Hansen, Paul Arne
Haupt, Herbert
Haynes, William C.
Kitchens, A. Parkerf
Hofer, A. W.
Hoffman, Heiner
Holmes, Francis O.
Honigberg, B. M.
Hucker, George J.
Huddleson, I. F.
Janke, Alexander
LIST OF CONTRIBUTORS
How Bacteria are Named and Identified 15
Paraspirilluni 257
Beggiatoales 837
Beggiatoaceae 837
Leucotrichaceae 850
Pseudomonas 89
Xanthomonas 152
Agrobacterium 288
Erwinia 349
Corynebacterium 579
Beneckea 328
Cellulomonas 601
Pasteurella 395
Eubacterium 552
Catenabacterium 560
Ramibacterium 563
Cillobacterium 566
Agrobacterium 288
Alcaligenes 297
Actinomycetales 694
Actinoplanaceae 825
Spirochaetales 892
Propionibacteriaceae 569
Microbacterium 600
Hyphomicrobiales 276
Peptococcus 474
Shigella 384
Halubacterium 207
Chromobaclerium 292
Staphylococcus 464
Streptobacillus 451
Mycoplasmatales 914
Bacillus 613
Mycobacterium 695
Mycobacterium 695
Erysipelothrix 599
Actinobacillus 414
Pseudomonas 89
Dialister 440
Azotomonas 198
Azotobacteraceae 283
Fusobacterium 436
Sphaerophorus 441
Virales 985
Parasites of protozoa 927
Micrococcus 455
Gaffkya 466
Leuconostoc 531
Brucella 404
Thiobacteriaceae 78
Thiobacterium 79
t Deceased.
LIST OF CONTRIBUTORS
Kalz, Gertrude G.
Kelly, CD.
Kirby, Harold, Jr.
Kluyver, A. J.f
Langford, G. C.
Leathen, Wm. W.
Lessel, Erwin F., Jr.
Lochhead, A. G.
McClung, L. S.
McCoy, Elizabeth
Merchant, I. A.
Morse, E. V.
Murray, E. G. D.
Neitz, W, O.
Nellis, Lois
Niven, C. F.
0rskov, J.
Parker, CD.
Pederson, Carl S.
Pelczar, M. J.
Peshkoff, M. A.
Philip, Cornelius B.
Pittman, Margaret
Pringsheim, E. G.
Macromonas 80
Thiovulum 81
Thiospira 82
Salmonella 368
Noguchia 421
Bacteroides 424
Parasites of protozoa 927
Zymomonas 199
Erysipelothrix 599
Ferrohacillus 227
Photobacterium 193
Selenomonas 258
Myconostoc 260
Pasteurella 395
Euhacterium 552
Catenabacterium 560
Ramibacterium 563
Vilreoscillaceae 844
Arthrobacter 605
Clostridium 634
Clostridium 634
Pasteurella 395
Vibrio 229
Corynebacterium 579
Enter obacteriaceae 332
Moraxella 419
Bacteroidaceae 423
Neisseria 480
Lactobacillaceae 505
Diplococcus 507
Corynebacterium 579
Listeria 597
Anaplasma 981
Mycococcus 707
Lactobacillaceae 505
Streptococcus 508
Microcyclus 253
Thiobacillus 83
Lactobacillaceae 505
Pediococcus 529
Leuconostoc 531
Lactobacillus 542
Neisseria 480
Veillonella 485
Caryophanales 830
Microtatobiotes 931 and 933
Rickettsiales 934
Anaplasmataceae 980
Bordetella 402
Haemophilus 406
Vilreoscillaceae 844
t Deceased.
Vlll
LIST OF CONTRIBUTORS
Rake, Geoffrey W.
Reed, Guilford B.f
Robinson, Elliott S.
Robinson, George H.f
Schatz, Albert
Seeley, H. W.
Seeliger, H.
Sherman, James W.f
Skerman, V. B. D.
Smit, Jan
Smith, Louis DeSpain
Smith, N. R.
Snieszko, S. F.
Spray, R. S.
Stanier, R. Y.
Starkey, R. L.
Steinhaus, E. A.
Stuart, C. A.
Temple, Kenneth L.
Thj0tta, Th.t
Tobie, W. C.
van Niel, C. B.
Vaughn, Reese
Verona, Onorato
Waksman, S. A.
Wattie, Elsie (Mrs.
Lackey)
Weeks, Owen B.
Weinman, David
Wolff, J. W.
Yale, N. W.
ZoBell, Claude E.
Chlamydiaceae 957
Mycobacterium 695
Diplococcus 507
Spirochaetales 892
Hydrogenomonas facilis 76
Streptococcus 508
Corynebacterium 579
Lactobacillaceae 505
Streptococcus 508
Artificial Keys 987
Sarcina 467
Lactobacillaceae 505
Peptostreptococcus 533
Bacteroidaceae 423
Bacillus 613
Aeromonas 189
Clostridium 634
Myxobacterales 854
Nitrobacteraceae 68
Virales 985
Proteus 364
Thiobacillus 83
Alginornonas 202
Alginobacter 348
Chromobacterium 292
Rhodobacteriineae 35
Propionibacterium 569
Achromatiaceae 851
Acetobacter 183
Cellvibrio 250
Actinomycetales 694
Mycococcus 707
Actinomycetaceae 713
Streptomycetaceae 744
Zoogloea 206
Spirochaetales 892
Flavobacterium 309
Bartonellaceae 968
Leptospira 907
Escherichia 335
Aerobacter 341
Desulfovibrio 248
t Deceased.
PREFACE TO SEVENTH EDITION
The general format of the seventh edition of Bergey's Manual of Deter-
minative Bacteriology differs but Uttle from that of the sixth edition. How-
ever, examination will reveal many changes in the content as the result of a
thoroughgoing revision. Among these the following seem to be worthy of special
comment.
The most obvious change is that of the separation into two volumes of the
material comparable to that which appeared in the sixth edition. The present
volume is entitled the seventh edition of Bergey's Manual of Determinative
Bacteriology. This Manual contains an outlined classification of the bac-
teria and the descriptions of the taxa from Class to Species and Subspecies, to-
gether with the appropriate keys. Nearly all species regarded as having been
inadequately described or that could not be definitely placed have been excluded,
together with many of the less important synonyms of the accepted species.
These, together with the index to all the literature of both accepted and poorly
described organisms have been transferred to a volume to be known as the
Index Bergeyana. The latter volume will include all descriptions and citations
to species formerly found as appendices or indefinitely placed as species incertae
sedis. The host and habitat index will also be found in the Index Bergeyana.
The net result is that the Manual itself contains descriptions of many more
species with more adequate descriptions than have former editions; the transfer
of much material to the Index Bergeyana has meant a reduction in the number
of pages and a book of more convenient size and greater usefulness. The Index
Bergeyana should prove to be an invaluable tool for the research microbiologist,
containing, as it will, references to the whole field of systematic bacteriology and
an index to the names of described species, both valid and invalid.
Much material of historical value and interest in the sixth and earlier editions
of the Manual has been excluded, not because it is lacking in real value to the
student, but because repetition of its publication is now unnecessary.
This edition of the Manual represents the coordinated results of the work of
one hundred contributors, about thirty-five more than assisted in preparing the
sixth edition. The contributors to the Manual are to be regarded in all nomen-
clatural matters as strictly the authors of their sections. All new names of taxa
and the names of all new combinations are to be ascribed to these authors, and
not to the editors of the Manual. Contributors from countries other than the
United States are more numerous than in the earlier editions. In other words,
the Manual is rapidly assuming the character of an international publication.
To all of these contributors the sincere thanks of the Board of Editors, and par-
ticularly of the Editor-in-Chief, are due. The seventh edition is a tribute to the
patience, care and scientific acumen of these individuals. Special note should be
taken of the assistance rendered in the office of the Editor-in-Chief by Mr.
Erwin Lessel, Miss Maude Hogan, Mrs. Eleanore Heist CUse and Mrs. Margaret
Edson Breed.
X PREFACE TO SEVENTH EDITION
The keys to the several categories of taxa (orders, families, tribes, genera and
species) have been revised with a view to making them more reUable and useful.
There is included also an artificial key to the species prepared by Professor
V. B. D. Skerman, which key should prove helpful.
The Section on Nomenclature, including a synopsis of the Botanical Code of
Nomenclature, has been eliminated. At the time of preparation of the sixth
edition, the International Code of Bacteriological Nomenclature had not been
finally approved, and emphasis was properly laid upon the rules used in Botany.
This is no longer pertinent. The Bacteriological Code appeared in 1948 too late
for use in making appropriate revisions in the 6th Edition of the Manual. The
revised International Code of Nomenclature of the Bacteria and Viruses is about
to be published. This contains annotations that should prove of value to the
student, and should be regarded as a helpful aid in the understanding of the
nomenclature used in the 7th Edition of the Manual.
The naming and classification of the viruses, as published in the sixth edition
of the Manual, was regarded by some eminent virologists as perhaps inadvisable
because it was premature. They felt strongly that the problems of morphology,
physiology, pathogenesis and inter-relationships of the viruses were not as yet
sufficiently resolved to make satisfactory taxonomy and classification practicable.
After consultation with the International Subcommittee on Viruses it was de-
cided that the Virus Section should not be included in the seventh edition. This
deletion has been made with the full expectation that sufficient international
agreement will be reached to make possible adequate treatment in the eighth
edition. The Editorial Committee recognizes that a satisfactory system of
nomenclature and taxonomy for the viruses is imperative.
The Editors wish to repeat and emphasize a statement made in the Preface
of the first edition of the Manual:
"The assistance of all bacteriologists is earnestly solicited in the correction of possible
errors in the text."
Among the tasks of the several editions of the Manual has been the codification
of an increasingly satisfactory classification of the bacteria and the correction of
the nomenclature of the past. The present volume undoubtedly has many errors
that were not caught notwithstanding a most earnest effort. There are also many
unresolved questions. Inasmuch as this volume appears at almost the same time
as the Revised International Bacteriological Code, there are doubtless still some
inconsistencies.
E. G. D. Murray
N. R. Smith
R. S. Breed, Chairman
Editorial Committee
PREFACE TO SEVENTH EDITION XI
NOTE
The Board of Trustees of Bergey's Manual wish to record their profound
sorrow at the death on February 10, 1956, of Dr. Robert S. Breed, Chairman of
the Board of Trustees and Editor-in-Chief of the Manual. Most regrettable is
the fact that he did not live to see in printed form the results of his untiring
and devoted labor. At the time of his death, most of the manuscript for the
seventh edition had been placed in the hands of the publisher; the remainder was
in such shape that it could be promptly submitted. The Board of Trustees of
the Bergey Trust, including the Board of Editors, wish to pay tribute to the
devotion, energy and skill of Dr. Breed over a period of many years, as shown
in the organization of better bacterial nomenclature and classification. The
science of microbiology is his debtor.
Reconciliation of the nomenclature used in the seventh edition of the Manual
with the provisions of the revised International Code of Nomenclature of the
Bacteria and Viruses had not been entirely completed by Dr. Breed. As far as
possible, these discrepancies have been corrected before publication.
The death of Dr. Breed leaves the Editorial Committee without a chairman
and editor-in-chief. The Board of Trustees is unanimous in the belief that a suc-
cessor to Dr. Breed should be found outside its present membership. Dr. Breed
was also Chairman of the Board of Trustees. Following his death, reorganization
of the Board was undertaken by Dr. Conn, as Treasurer, and Dr. Buchanan was
designated as Chairman. The latter has agreed to serve until publication of the
seventh edition of Bergey's Manual and of the first edition of the companion
volume, planned by Dr. Breed and christened by him the Index Bergeyana.
It is recognized to be a matter of urgency and diflficulty to replace Dr. Breed
with an editor-in-chief for the eighth edition and to find a means of making this
onerous responsibility something more than the labor of love it always was to him.
E. G. D. Murray
N. R. Smith
H. J. Conn
R. E. Buchanan, Chairman
Board of Trustees
PREFACE OF FIRST EDITION
The elaborate system of classification of the bacteria into families, tribes
and genera by a Committee on Characterization and Classification of the
Society of American Bacteriologists (1917, 1920) has made it very desirable
to be able to place in the hands of students a more detailed key for the
identification of species than any that is available at present. The valuable
book on "Determinative Bacteriology" by Professor F. D. Chester, pub-
lished in 1901, is now of very little assistance to the student, and all previous
classifications are of still less value, especially as earlier systems of classifica-
tion were based entirely on morphologic characters.
It is hoped that this manual will serve to stimulate efforts to perfect the
classification of bacteria, especially by emphasizing the valuable features
as well as the weaker points in the new system which the Committee of the
Society of American Bacteriologists has promulgated. The Committee
does not regard the classification of species offered here as in any sense
final, but merely a progress report leading to more satisfactory classification
in the future.
The Committee desires to express its appreciation and thanks to those
members of the society who gave valuable aid in the compilation of material
and the classification of certain species. . . .
The assistance of all bacteriologists is earnestly solicited in the correction
of possible errors in the text ; in the collection of descriptions of all bacteria
that may have been omitted from the text; in supplying more detailed
descriptions of such organisms as are described incompletely; and in furnish-
ing complete descriptions of new organisms that may be discovered, or in
directing the attention of the Committee to publications of such newly
described bacteria.
David H. Bergey, Chairman
Francis C. Harrison
Robert S. Breed
Bernard W. Hammer
Frank M. Huntoon
Committee on Manual.
August, 1923.
CONTENTS
Introduction 1
Considerations Influencing the Classification Used in This Edition of the Manual. ... 4
How Bacteria are Named and Identified 15
Division I. Protophyta Sachs, 1874, cviend. Krassilnikov, 1949 29
Class I. Schizophyceae Cohn, 1879 30
Class II. Schizomycetes von Naegeli, 1857 33
Order I. Pseudomonadales Orla-Jensen, 1921 35
Suborder I. Rhodohacteriineae Breed, Murray and Kitchens, 1944 35
Family I. Thiorhodaceae Molisch, 1907 38
Genus I. Thiosarcina Winogradsky, 1888 39
Genus II. Thiopedia Winogradsky, 1888 40
Genus III. Thiocapsa Winogradsky, 1888 41
Genus IV. Thiodiclyon Winogradsky, 1888 41
Genus V. Thioihece Winogradsky, 1888 42
Genus VI. Thiocyslis Winogradsky, 1888 42
Genus VII. Lamprocystis Schroeter, 1886 43
Genus VIII. Amoebobacter Winogradsky, 1888 44
Genus IX. Thiopolycoccus Winogradsky, 1888 45
Genus X. Thiospirillum Winogradsky, 1888 46
Genus XI. Rhabdomonas Cohn, 1875 48
Genus XII. Rhodolhece Molisch, 1907 50
Genus XIII. Chromatium Perty, 1852 50
Family II. A thiorhodaceae Molisch, 1907 53
Genus I. Rhodopseudomonas Kluyver and van Niel, 1937, emend, van
Niel, 1944 53
Genus II. Rhodospirillum Molisch, 1907, emend, van Niel, 1944 58
Family III. Chlorobacteriaceae Lauterborn, 1913 61
Genus I. Chlorobium Nadson, 1912 62
Genus II. Pelodictyon Lauterborn, 1913 63
Genus III. Clathrochloris Geitler, 1925 64
Genus IV. Chlorobacterium Lauterborn, 1915 65
Genus V. Chlorochromatium Lauterborn, 1906 65
Genus VI. Cylindrogloea Perfiliev, 1914 66
Suborder II. Pseudomonadineae Breed, Murray and Smith, sub-ordo nov 67
Family I. Nitrobacteraceae Buchanan, 1917 68
Genus I. Nitrosomonas Winogradsky, 1890 68
Genus II. Nitrosococcus Winogradsky, 1892 69
Genus III. Nitrosospira Winogradsky, 1931 70
Genus IV. Nitrosocystis Winogradsky, 1931 70
Genus V. Nitrosogloea H. Winogradsky, 1935 71
Genus VI. Nitrobacter Winogradsky, 1892 72
Genus VII. Nitrocystis H. Winogradsky, 1935 73
Family II. Methanomonadaceae Breed, fam. nov 74
Genus I. Methanomonas Orla-Jensen, 1909 74
Genus II. Hydrogenomonas Orla-Jensen, 1909 75
Genus III. Carboxydomonas Orla-Jensen, 1909 77
Family III. Thiobacteriaceae Janke, 1924 78
Genus I. Thiobaclerium Janke, 1924 79
Genus II. Macromonas Utermohl and Koppe, 1923 80
Genus III. Thiovulum Hinze, 1913 81
xiii
'^'3664
CONTENTS
Genus IV. Thiospira Vislouch, 1914 82
Genus V. Thiobacillus Beijerinck, 1904 83
Family IV. Pseudomonadaceae Winslow et al., 1917 88
Genus I. Pseudomonas Migula, 1894 89
Genus II. Xanthomonas Dowson, 1939 152
Genus III. Acetobacier Beijerinck, 1898 183
Genus IV. Aeromonas Kluyver and van Niel, 1936 189
Genus V. Photohacterium Beijerinck, 1889, emend. Breed and Lessel,
1954 193
Genus VI. Azotomonas Stapp, 1940 198
Genus VII. Zymomonas Kluyver and van Niel, 1936 199
Genus VIII. Protaminobacter den Dooren de Jong, 1926 200
Genus IX. Alginomonas Thj0tta and K&ss, 1945 202
Genus X. Mycoplana Gray and Thornton, 1928 204
Genus XI. Zoogloea Cohn, 1854 206
Genus XII. Halobacterium Elazari-Volcani, 1940 207
Family V. Caulobacteraceae Henrici and Johnson, 1935, emend. Breed 212
Genus I. Caulobacter Henrici and Johnson, 1935 213
Genus II. Gallionella Ehrenberg, 1838 214
Genus III. Siderophacus Beger, 1944 216
Genus IV. Nevskia Famintzin, 1892 216
Family VI. Siderocapsaceae Pribram, 1929 217
Genus I. Siderocapsa Molisch, 1909 218
Genus II. Siderosphaera Beger, 1944 220
Genus III. Sideronema Beger, 1941 220
Genus IV. Ferribacterium Brussoff , 1916 221
Genus V. Sideromonas Cholodny, 1922 222
Genus VI. N aumanniella Dorff, 1934 223
Genus VII. Ochrobium Perfiliev, 1921 225
Genus VIII. Siderococcns Dorff, 1934 225
Genus IX. Siderobacter Naumann, 1922 226
Genus X. Ferrobacillus Leathen and Braley, 1954 227
Family VII. Spirillaceae Migula, 1894 228
Genus I. Vibrio Miiller, 1773 229
Genus II. Desulfovibrio Kluyver and van Niel, 1936 248
Genus III. Methanobacterium Kluyver and van Niel, 1936 250
Genus IV. Cellvibrio Winogradsky, 1929 250
Genus V. Cellfalcicula Winogradsky, 1929 252
Genus VI. Microcyclus 0rskov, 1928 253
Genus VII. Spirillum Ehrenberg, 1832 253
Genus VIII. Paraspirillum Dobell, 1912 257
Genus IX. Selenomonas von Prowazek, 1913 258
Genus X. Myconostoc Cohn, 1875 260
Order II. Chlamydobacteriales Buchanan, 1917 262
Family I. Chlamydobacteriaceae Migula, 1894 262
Genus I. Sphaerotilus Kiitzing, 1833 263
Genus II. Leptothrix Kiitzing, 1843 264
Genus III. Toxothrix Molisch, 1925 269
Family II. Peloplocaceae Beger, fam. nov 270
Genus I. Peloploca Lauterborn, 1913 270
Genus II. Pelonema Lauterborn, 1915 271
Family III. Crenotrichaceae Hansgirg, 1888 272
Genus I. Crenothrix Cohn, 1870 272
CONTENTS XV
Genus II. Phragmidiothrix Engler, 1883 273
Genus III. Clonothrix Roze, 1896 274
Order III. Hyphomicrobiales Douglas, ordo nov 276
Family I. Hyphomicrobiaceae Babudieri, 1950 276
Genus I. Hyphomicrobimn Stutzer and Hartleb, 1898 277
Genus II. Rhodomicrobium Duchow and Douglas, 1949 277
Family II. Pasteuriaceae Laurent, 1890, emend. Henrici and Johnson, 1935. . 278
Genus I. Pasteuria MetchnikofT, 1888 279
Genus II. Blasiocaulis Henrici and Johnson, 1935 279
Order IV. Enbacteriales Buchanan, 1917 281
Family I. Azotobacteraceae Bergey, Breed and Murray, 1938 283
Genus I. Azotobacter Beijerinck, 1901 283
Family II. Rhizobiaceae Conn, 1938 .' 285
Genus I. Rhizobium Frank, 1889 285
Genus II. Agrobacterium Conn, 1942 288
Genus III. Chromobacterium Bergonzini, 1881 292
Family III. Achromobacteraceae Breed, 1945 296
Genus I. Alcaligenes Castellani and Chalmers, 1919 297
Genus II. Achromobacter Bergey et al., 1923 300
Genus III. Flavobacteriurn Bergey et al., 1923 309
Genus IV. Agarbacterium Angst, 1929 322
Genus V. Beneckea Campbell, gen. nov 328
Family IV. Enterobacteriaceae Rahn, 1937 332
Tribe I. Escherichieae Bergey, Breed and Murray, 1938 334
Genus I. Escherichia Castellani and Chalmers, 1919 335
Genus II. Aerobacter Beijerinck, 1900 341
Genus III. Klebsiella Trevisan, 1885 344
Genus IV. Paracolobactrum Borman, Stuart and Wheeler, 1944 346
Genus V. Alginobacter Thj0tta and Kiss, 1944 348
Tribe II. Erwinieae Winslow et al., 1920 349
Genus VI. Erwinia Winslow et al., 1917 349
Tribe III. Serratieae Bergey, Breed and Murray, 1938 359
Genus VII. Serratia Bizio, 1823, emend. Breed and Breed, 1927 359
Tribe IV. Proteeae Castellani and Chalmers, 1919 364
Genus VIII. Proteus Hauser, 1885 364
Tribe V. Salmonelleae Bergey, Breed and Murray, 1938 368
Genus IX. Salmonella Lignieres, 1900 368
Genus X. Shigella Castellani and Chalmers, 1919 384
Family V. Brucellaceae, nom. nov 394
Genus I. Pasteurella Trevisan, 1887 395
Genus II. Bordetella Moreno-Lopez, 1952 402
Genus III. Brucella Meyer and Shaw, 1920 404
Genus IV. Haemophilus Winslow et al., 1917 406
Genus V. Actinobacillus Brumpt, 1910 414
Genus VI. Calymmatobacterium Aragao and Vianna, 1913 418
Genus VII. Moraxella Lwoff, 1939 419
Genus VIII. Noguchia Olitsky, Syverton and Tyler, 1934 421
Family VI. Bacteroidaceae Breed, Murray and Smith, fam. nov 423
Genus I. Bacieroides Castellani and Chalmers, 1919 424
Genus II. Fusobacterium Knorr, 1922 436
Genus III. Dialister Bergey et al., 1923 440
Genus IV. Sphaerophorus Pr6vot, 1938 441
Genus V. Streptobacillus Levaditi, Nicolau and Poincloux, 1925 451
XVI CONTENTS
Family VII. Micrococcaceae Pribram, 1929 454
Genus I. Micrococcus Cohn, 1872 455
Genus II. Staphylococcus Rosenbach, 1894 464
Genus III. Gaffkya Trevisan, 1885 466
Genus IV. Sarcina Goodsir, 1842 467
Subgenus I. Zyniosarcina Smit, 1930 468
Subgenus II. Methanosarcina Kluyver and van Niel, 1936 468
Subgenus III. Sarcinococcus Breed, 1948 468
Subgenus IV. Urosarcina Miquel, 1888 468
Genus V. Methanococcus Kku'ver and van Niel, 1936, emend. Barker,
1936 473
Genus VI. Peptococcus Kluyver and van Niel, 1936 474
Family VIII. Neisseriaceae Prevot, 1933 480
Genus I. Neisseria Trevisan, 1885 480
Genus II. Veillonella Prevot, 1933 485
Family IX. Brevibacteriaceae Breed, 1953 490
Genus I. Brevibacterium Breed, 1953 490
Genus II. Kiirthia Trevisan, 1885 503
Family X. Lactobacillaceae Winslow et al., 1917 505
Tribe I. Streptococceae Trevisan, 1889 506
Genus I. Diplococcus Weichselbaum, 1886 507
Genus II. Streptococcus Rosenbach, 1884 508
Genus III. Pediococcus Balcke, 1884, e77iend. Mees, 1934 529
Genus IV. Leuconostoc van Tieghem, 1878, eviend. Hucker and Pederson,
1930 531
Genus V. Peptostreptococcus Kluyver and van Niel, 1936 533
Tribe II. Laciobacilleae Winslow et al., 1920 541
Genus I. Lactobacillus Beijerinck, 1901 542
Subgenus I. Lactobacillus Beijerinck, 1901 543
Subgenus II. Saccharobacillus van Laer, 1892 543
Genus II. Eubacterium Prevot, 1938 552
Genus III. Catenabacterium Prevot, 1938 560
Genus IV. Ramibacterium Prevot, 1938 563
Genus V. Cillobacterium Prevot, 1938 566
Family XI. Propionibacteriaceae Delwiche, jam. nov 569
Genus I. Propionibacterium Orla-Jensen, 1909 569
Genus II. Butijribacterium Barker and Haas, 1944 577
Genus III. Zymohacterium Wachsman and Barker, 1954 577
Family XII. Corynebacteriaceae Lehmann and Neumann, 1907 578
Genus I. Corynebacterium Lehmann and Neumann, 1896 579
Genus II. Listeria Pirie, 1940 597
Genus III. Erysipelothrix Rosenbach, 1909 599
Genus IV. Microbacterium Orla-Jensen, 1919 600
Genus V. Cellulomonas Bergey et al., 1923, emend. Clark, 1952 601
Genus VI. Arthrobacter Fischer, 1895, emend. Conn and Dimmick, 1947. . 605
Family XIII. Bacillaceae Fischer, 1895 613
Genus I. Bacillus Cohn, 1872 613
Genus II. Clostridium Prazmowski, 1880 634
*Order V. Actinomycetales Buchanan, 1917 694
* Erratum: Due to a clerical error the orders Caryophanales and Actinomycetes of the
class Schizomycetales appear in the wrong sequence in numerous places throughout the
MANUAL: correctly, Caryophanales is Order V and should appear before Order VI, Acti-
nomycetales (see pages 12, 33, and 34 for reasons why Caryophanales precedes Actinomyce-
tales in the classification scheme).
CONTENTS XVll
Family I. Mycobacteriaceae Chester, 1901 695
Genus I. Mycobacterium Lehmann and Neumann, 1896 695
Genus II. Mycococcus Krassilnikov, 1938 707
Family II. Actinomyceiaceae Buchanan, 1918 713
Genus I. Nocardia Trevisan, 1889 713
Genus II. Actinomyces Harz, 1877 742
Family III. Streptonrycctaceae Waksman and Henrici, 1943 744
Genus I. Streptomyces Waksman and Henrici, 1943 744
Genus II. Micromunospora ^rskov, 1923 822
Genus III. Thermoactinomyces Tsiklinsky, 1899 824
Family IV. Aciinoplanaceae Couch, 1955 825
Genus I. Actinoplanes Couch, 1950 826
Genus II. Streptosporangiurn Couch, 1955 828
Order VI. Caryophanales Peshkoff, 1940 830
Family I. Caryophanaceae Peshkoff, 1940 830
Genus I. Caryophanon Peshkoff, 1940 831
Genus II. Lineola Pringsheim, 1950 832
Genus III. SimonsieUa Schmid, 1922 833
Family II. Oscillospiraceae Peshkoff, 1940 834
Genus I. Oscillospira Chatton and Perard, 1913 834
Family III. Arthromitaceae Peshkoff, 1940 835
Genus I. Arthromitus Leidy, 1849 835
Genus II. Coleomitus Duboscq and Grasse, 1930 836
Order VII. Beggiatoales Buchanan, ordo nov 837
Family I. Beggiatoaceae Migula, 1894 837
Genus I. Beggiatoa Trevisan, 1842 838
Genus II. Thiospirillopsis Uphof, 1927 840
Genus III. Thioploca Lauterborn, 1907 841
Genus IV. Thiothrix Winogradsky, 1888 842
Family II. Vitreoscillaceae Pringsheim, 1949 844
Genus I. Vitreoscilla Pringsheim, 1951 845
Genus II. Bactoscilla Pringsheim, 1951 848
Genus III. MicrosciUa Pringsheim, 1951 849
Family III. Leucotrichaceae Buchanan, farn. nov 850
Genus I. Leucothrix Oersted, 1844, emend. Harold and Stanier, 1955. . . . 850
Family IV. Achromatiaceae Massart, 1902 851
Genus I. Achromatiiim Schewiakoff, 1893 852
Order VIII. Myxobacterales Jahn, 1915 854
Family I. Cytophagaceae Stanier, 1940 858
Genus I. Cytophaga Winogradsky, 1929 858
Family II. Archangiaceae Jahn, 1924 863
Genus I. Archangium Jahn, 1924 863
Genus II. Stelangium Jahn, 1915 866
Family III. Sorangiaceae Jahn, 1924 866
Genus I. Sorangium Jahn, 1924 866
Family IV. Polyangiaceae Jahn, 1924 870
Genus I. Polyangium Link, 1809 870
Genus II. Synangium Jahn, 1924 877
Genus III. Podangium Jahn, 1924 877
Genus IV. Chondrojnyces Berkeley and Curtis, 1874 879
Family V. Myxococcaceae Jahn, 1924 882
Genus I. Myxococcus Thaxter, 1892 883
Genus II. Chondrococcus Jahn, 1924 886
XVlll CONTENTS
Genus III. Angiococcus Jahn, 1924 889
Genus IV. Sporocytophaga Stanier, 1940 890
Order IX. Spirochaeiales Buchanan, 1918 892
Family I. Spirochaetaceae Swellengrebel, 1907 892
Genus I. Spirochaeta Ehrenberg, 1833 893
Genus II. Saprospira Gross, 1911 894
Genus III. Cristispira Gross, 1910 895
Family II. Treponemataceae Robinson, 1948 896
Genus I. Borrelia Swellengrebel, 1907 897
Genus II. Treponema Schaudinn, 1905 904
Genus III. Leptospira Noguchi, 1917 907
Order X. Mycoplasmaiales Freundt, 1955 914
Family I. Mycoplasmaiaceae Freundt, 1955 914
Genus I. Mycoplasma Nowak, 1929 914
Addendum to Class II, Schizomycetes von Naegeli. Bacteria symbiotic or parasitic
in protozoa 926
Class III. Microtatobiofes Philip, 1956 931 and 933
Order I. iSz'c^^eWsmZes Buchanan and Buchanan, 1938, emend. Gieszczykiewicz, 1939 934
Family I. Rickettsiaceae Pinkerton, 1936 934
Tribe I. Rickettsieae Philip, trib. nov 935
Genus I. Rickettsia da Rocha-Lima, 1916 935
Subgenus A. Rickettsia Philip, 1943 937
Subgenus B. Zinssera Macchiavello, 1947 939
Subgenus C. Dermacentroxenus (Wolbach, 1919) Philip, 1943 941
Subgenus D. Rochalimaea Macchiavello, 1947 945
Genus II. Coxiella Philip, 1948 947
Tribe II. Ehrlichieae Philip, trib. nov 948
Genus III. Ehrlichia Moshkovskiy, 1945 949
Genus IV. Cowdria Moshkovskiy, 1945 950
Genus V. Neorickettsia Philip et al., 1953 951
Tribe III. Wolbachieae Philip, 1956 952
Genus VI. Wolbachia Hertig, 1936 953
Genus VII. Symhiotes Philip, 1956 956
Genus VIII. Rickettsiella Philip, 1956 957
Family II. Chlamydiaceae Rake, fam. nov 957
Genus I. Chlamydia Rake, 1956 958
Genus II. Colesiota Rake, 1948 959
Genus III. Ricolesia Rake, gen. nov 959
Genus IV. Colettsia Rake, nom. nov 961
Genus V. Miyagawanella Brumpt, 1938 961
Family III. Bartonellaceae Gieszczykiewicz, 1939 968
Genus I. Bartonella Strong, Tyzzer and Sellards, 1915 969
Genus II. Grahariiella Brumpt, 1911 971
Genus III. Haemobartonella Tyzzer and Weinman, 1939 972
Genus IV. Eperythrozoon Schilling, 1928 977
Family IV. Anaplasmataceae Yakimov, 1931 980
Genus I. Anaplasma Theiler, 1910 981
Order II. Virales Breed, Murray and Kitchens, 1944 985
Artificial Key 987
Index of Genera and Species 1093
INTRODUCTION
Suggestions for the Use of the Manual In Classifying
Unknown Organisms
No organism can be classified before its morphological, cultural, physiological
and pathogenic characters have been determined through a detailed study.
The characters used in the keys to orders, families and genera may ordinarily
be determined by the use of a dozen or more of the procedures described in the
]\Ianual of Microbiological Methods issued by the Committee on Bacteriological
Technic of the Society of American Bacteriologists. More complete examinations
are required in special cases to identify and to describe individual species ne-
cessitating resort to the original literature. When those prevailing are inad-
equate, new criteria are desirable. This desideratum extends to some higher taxa
to achieve more exact and distinctive definitions.
It is urged that beginning students be taught all of the techniques necessary
for the identification of species in the hope that the taxonomic work of the future
may be placed on a satisfactory basis.
After a complete study of the characters of the organism has been made,
either of two courses may be followed. (1) Use the Keys in the body of the text
as explained below. These follow what are believed to be the natural relationships
that exist between various groups of bacteria. (2) The beginning student may,
however, find the artificial key at the back of the Manual to be more helpful
than the natural keys in determining the identity of an unknown culture.
In all cases it should be kept in mind that many descriptions of species of
bacteria are not mentioned in this Manual. Failure to find agreement between
an unknown culture and any of the descriptions given in this Manual does not
prove that the unknown culture represents a species that has never been studied
and described.
If the student wishes to follow through the natural keys he should turn to
page 33 and ascertain first in which order the organism belongs. When the
order has been ascertained, turn to the page of the Manual on which the key to
that order is given. In this key ascertain the family or sub-family to which the
organism belongs.
When the family has been determined, again refer to the page of the Manual
on which the key to that family is given. In this key ascertain the tribe to which
the organism belongs.
When the tribe has been determined, again find the page of the Manual on
which the key to the tribe is given. In this key ascertain the genus to which the or-
ganism belongs.
When the genus has been determined, again refer to the page of the Manual
on which the key to that genus is given. In this key trace out the species under
investigation.
1
2 SUGGESTIONS FOR USE OF THE MANUAL
For example, if one wishes to identify a short, peritrichous. Gram-negative,
non-spore-forming, non-chromogenic rod that grows well on ordinary culture
media at 37 °C., fermenting glucose and lactose with the production of acid and
gas, not liquefying gelatin, with negative reaction for acetylmethylcarbinol,
citrate-negative, alginate-negative, pectinase-negative, producing indole and
reducing nitrates, consult the key to the orders on page 33.
In this key examine I: Cells rigid. Motile by means of polar flagella or non-motile.
This does not indicate our organism, which is peritrichous. We turn next to II:
Not as above. Under A: Cells rigid. Spherical or straight, rod-shaped cells. Occur
singly, in chains or in trichomes. Motile by means of peritrichous flagella or non-
motile. These characters agree with those of the organism in question.
We turn now to 1, in the same key: Cells spherical or rod-shaped. No trichomes,
though chains of cells may occur. This indicates that the organism in question
belongs in Order IV, Eubacteriales.
In the key to the families of Order Eubacteriales, p. 281, examine I: Cells
rod-shaped. Gram-negative. This indicates the organism in question, so we turn
next to A: Aerobic or facultatively anaerobic, which again indicates the organism
in question.
The next entry, 1: Large, ovoid to rod-shaped cells, sometimes yeast-like in
appearance. Free-living in soil. Fix free nitrogen, does not indicate the culture
under study so we turn to 2 : Not as above.
The heading, 2 : Heterotrophic rods which may not require organic nitrogen for
growth. Usually motile, with one to six flagella. Frequently form nodules or tubercles
on roots of plants or show violet chromogenesis. Colonies usually large and slimy,
especially on mannitol agar, does not indicate the organism in question. We then
turn to aa: Not as above.
Heading b: Straight rods which grow readily on ordinary peptone media. May
of may not ferment sugars anaerobically with the production of organic acids again
indicates the culture under study.
The heading c : Glucose usually attacked oxidatively or not at all does not indicate
the organism in question as it is an active fermenter of glucose and lactose,
cc: Ferment glucose anaerobically, frequently producing visible gas from glucose, and
sometimes lactose indicates that the culture under study belongs in the Family IV,
Enterobacteriaceae .
We now turn to the key to the tribes of family Enterobacteriaceae, page 334.
In this key we examine I: Lactose fermented anaerobically, usually within 48 hours.
This indicates the culture under study.
The entry A: Prodigiosin not produced also indicates the non-chromogenic
organism in question. Likewise, 1: Do not produce protopectinase. Not parasitic
on plants also applies to our culture, so we turn to the key to the genera of
Tribe I, Escherichieae, page 335.
We turn now to I: Alginic acid is not decomposed with the production of acid and
gas. This likewise indicates the organism is question. Under this. A: Lactose is
fermented within 48 hours also indicates the culture under study, as does 1:
SUGGESTIONS FOR USE OF THE MANUAL 3
Acetylmethylcarhinol not produced. Methyl red test positive. Salts of citric acid may
or may not he used as a sole source of carbon. This places the organism in Genus I
Escherichia.
We turn now to the key to the species of Genus I, Escherichia, page 335. On
tracing our organism in this key we find that its characters correspond with those
of Escherichia coli and turn to the description of this species for a final confirma-
tion of this identification.
It is self evident that where the characters of the original culture have not
been determined accurately or completely, the identity of the unknown can-
not be determined positively.
A second difficulty in the use of a key comes from inexperience in the use of
technical terms; that is, the student may not thoroughly understand the meaning
of the statement in the key and so cannot follow a route through the key with
certainty. For example in the keys used here, the student must know the dif-
ference between (1) chains of cells which are composed of dividing cells which
do not separate at once, and (2) trichomes which are composed of dividing cells
which remain more permanently together and are normally flattened against
each other on adjacent sides. The trichomes may show some differentiation into
holdfast cells and reproductive cells (conidia) . Both chains of cells and trichomes
are to be distinguished from the mycelial threads found in Actinomycetaceae : the
latter are unseptate and show true branching.
The student should be warned not to take descriptions in the Manual too
literally or too rigidly. Descriptions are usually drawn to represent average
findings. Especially among bacteria, characters such as sugar fermentations,
gelatin liquefaction, presence or absence of flagella and other things may vary
within a species. Sometimes these variations are due to slight, possibly un-
recognized variations in the techniques used in determining these characters.
Real knowledge of the characteristics of species may also be very incomplete.
This is true not only with respect to the physiological activities of these micro-
organisms but also to such detectable structural features as the number and
position of flagella. Dark-field movies of motile cells and photographs taken with
the electron microscope are revealing new and heretofore unsuspected facts
regarding structural features.
Source and habitat data are frequently helpful in aiding the student to rec-
ognize species of bacteria and may indicate that the pathogenicity of the culture
in question may need to be tried on some specific animal or plant. By habitat
is meant the kind of a place in which the organism normally grows; by source,
the particular material and place from which the culture was obtained. This
source may or may not indicate the natural habitat. The source of cultures is
invariably more limited in scope than the habitat, as bacteria normally occur
wherever their particular habitat may be found in a world-wide distribution.
CONSIDERATIONS INFLUENCING THE CLASSIFICA-
TION USED IN THIS EDITION OF THE MANUAL
Robert S. Breed
Cornell University, Geneva, New York
The development of the classification systems used in the various editions of
Bergey's Manual has caused those of us responsible for this work to give
considerable thought to the probable evolutionary development of the living
things that are included under the general terms bacteria and, more recently,
viruses.
For those who are not familiar with the principles of evolution, it might be
well to bear in mind that all living things, including bacteria and viruses, do but
represent the present form of a long line of ancestral forms. Customarily these
lines of development are thought of as being not lineal but like the twigs and
branches of a tree which trace their origin back to the trunk of the tree, living
species being regarded as the separate and distinct tips of the twigs.
Bacteria and viruses, endowed as they are with a simple morphology, are
naturally thought of as being primordial or primitive in nature. This concept
is fundamental in all systems of classification that have been developed for these
organisms. Nevertheless it should not be forgotten that the different species or
kinds of these morphologically simple living things now extant may have under-
gone many types of changes during the course of their evolutionary development.
However, because bacteria and viruses do not have hard parts that fossilize,
there is little that can be learned about their evolution directly from historical
geology (paleontology) .
It is difficult to picture the environment under which the undifferentiated,
unicellular organisms lived when they first appeared on the earth, but it is
certain that this environment was quite different from the environment in which
similar organisms live today. One important feature of the present-day envi-
ronment that would have been lacking in the earliest periods would be the
association of unicellular organisms with more highly developed types of living
plants and animals and with the resultant accumulation of organic materials
that must take place as the natural processes of life and death go forward.
Organisms which are saprophytic and, still less, those which are parasitic would
not have had conditions favorable for their existence in the earliest periods in
which life developed on this planet. This makes it necessary to assume that the
earliest living things must have existed on comparatively simple, largely inorganic
food materials. With this thought in mind, some students of the systematic
relationships of living things have thought of the chemoautotrophic bacteria
that still exist as being more like primordial living things than are other types of
bacteria.
CONSIDERATIONS INFLUENCING CLASSIFICATION 5
It is true that the chemoautotrophic organisms are able to Hve on simple
inorganic foods that were, in all probability, available to living things under
early conditions in the development of the earth. However, it does not necessarily
follow that chemoautotrophic forms are the only ones that could have existed
in the beginning. It seems even more reasonable to assume that early living
forms developed a pigment like chlorophyll that enabled primordial bacteria to
utilize the sun's energy in synthesizing organic matter. Such photosynthetic
pigments are found in purple or green bacteria. These photoautotrophic forms
could have existed on the simple foods available when life began as readily as
could chemoautotrophic forms.
In either case, it is necessary to assume that living protoplasm, with its com-
plex enzymatic systems, existed before primordial bacteria, which utilized
inorganic materials as food. In other words, complex proteins had to be in
existence before either chemoautotrophic or photoautotrophic bacteria of the
types now found on the earth could exist.
Even if it is granted that photoautotrophic living things were primordial, it
must also be granted that when the existence of such organisms is postulated
we are not starting with the beginning of life itself. So little is known about the
possibility of living proteins (protoplasm) developing out of inorganic com-
pounds that speculation regarding this development has brought but very little
information that is factual.
In the present edition of Bergey's Manual, the classification used has been
rearranged on the assumption that the photoautotrophic bacteria extant today
presumably are the living organisms that are most nearly like the primordial
types of bacteria.
In support of this thought it should be kept in mind that the earliest living
forms must necessarily have been free-living forms, not saprophytes nor parasites.
This being the case, forms such as viruses that are very tiny in size and therefore
necessarily of a simple structure ought not to be regarded as primitive just
because of a comparatively simple structure. The viruses are adapted to life
within living protoplasm, and they represent an extreme degree of specialization
to a parasitic existence. They are known as organisms that invade the living cells
of higher plants and animals, including man. The latter are the living things
that were latest in development in geological time. Viruses could not have existed
before their host plants and animals were developed.
The term "viruses" ought not to be used for the hypothetical, very tiny,
free-living primordial organisms that must have existed before primordial
bacteria. Some investigators feel that such organisms may still exist in some as
yet unrecognized form.
It is not surprising that a great development has taken place in outline clas-
sifications since bacteriologists first tried to develop such classifications to express
the possible relationships of the organisms with which they have worked. While
O. F. Mueller (Animalcula infusoria et Marina. Hauniae. 1786) and C. G. Ehren-
berg (Die Infusionsthierchen als vollkommende Organismen. Leipzig, 1838) made
6 CONSIDERATIONS INFLUENCING CLASSIFICATION
simple beginnings along this line, their knowledge of bacteria as they are known
today was very limited indeed. Even in 1838, when Ehrenberg published his
description of the types of organisms found in infusions, microscopes had not
yet been developed to a place where even large bacteria could be studied with
any satisfaction.
By 1872, Ferdinand Cohn (Untersuchungen iiber Bacterien. I. Beitr. z. Biol,
d. Pflanzen., 1, Heft 2, 1872, 187-222), the botanist, began to understand that
a great variety of types of bacteria were in existence, and he was able to arrange
an outline classification on which later classifications of bacteria have been built.
However, his first outline classification of bacteria was scarcely pubUshed before
he felt that he should have expressed the relationships of the bacteria to the
simplest types of algae in a more intimate way. He therefore, in 1875 (Unter-
suchungen iiber Bacterien., II. ibid., 1, Heft 3, 1875, 141-207), drew up a second
classification in which he integrated the known groups of bacteria with known
groups of blue-green algae in a class, the Schizophyta. This arrangement assumed
that the bacteria had a much more intimate relationship to the blue-green algae
than the true fungi have to the green, red and brown algae.
It should be noted that early classifications of bacteria were based primarily
upon structural characters, particularly the shape of the cells. This was a
natural development, as morphological characters had been found to be useful
in drawing up natural classifications of higher plants and animals. It is also
(luite natural that workers who drew up these classifications should have regarded
the spherical organisms that they found as being primitive in nature. Little
was known at that time of the distribution of bacteria in nature. It was not
until later that it came to be realized that the bacteria that are spherical in
shape are normally found on the skin or in secretions of skin glands (milk and
other dairy products, etc.) of vertebrates. Few cocci exist as free-living forms in
water or soil. Likewise, when physiological studies were made, it was found that
the cocci require comparatively complex foods for their existence. Few modern
classifications retain the arrangement in which cocci are placed first as suggested
by Cohn in 1872.
Others have developed the early classifications* drawn up by Cohn, with many
individuals contributing to the development of a better and better understanding
of the evolutionary development of the bacteria. In the 1890's, two groups
of individuals undertook the publication of manuals describing the known species
of bacteria. These two groups exercised a great influence on the development of
systematic bacteriology.
Migula (Arb. Bact. Inst., Karlsruhe, 1, 1894, 235-238; in Engler and Prantl,
Naturlichen Pflanzenfamilien, Schizophyta, 1 Teil, la, 1895, 1-44) and his
students began their work at Karlsruhe, Germany, in the early part of the
1890's, publishing various papers and books, the last of which was Migula's
* For a more detailed discussion of outline classifications developed by bacteriologists,
3ee Manual, 3rd ed., 1930, 1-23; and Manual, 6th ed., 1948, 5-38.
CONSIDERATIONS INFLUENCING CLASSIFICATION Y
System der Bakterien (Bd. 1, 1897, 368 pp.; Bd. 2, 1900, 1068 pp., Jena). Only
one edition was published.
During the same period K. B. Lehmann and R. E. Neumann of Wiirzburg,
Germany, began the publication of their Bakteriologische Diagnostik, the first
edition of which was published, as were later editions, in two volumes (J. F.
Lehmann Verlag, Miinchen). The first edition was soon followed by a second and
later editions, the work being seriously interrupted by the first World War after
the publication of the 5th edition. Following the war they republished the 5th
edition with a supplement as the 6 edition and later carried through a complete
revision of this text which appeared as the 7th edition in 1927. No further editions
have been issued.
In the meantime, interest in taxonomic work had crystallized in the newly
organized (1899) Society of American Bacteriologists, led at first by F, D.
Chester. His Manual of Determinative Bacteriology, published in 1901 (The
MacMillan Co., New York), had great influence in guiding the thought of
American bacteriologists, but it never has been widely known outside of North
America.
As the Society developed, others took an active interest in this work, among
them R. E. Buchanan (Jour. Bact., 1, 1916, 591-596; 2, 1917, 155-164, 347-350,
603-617; 3, 1918, 27-61, 175-181, 301-306, 403-406, 461-474, 541-545), who
organized an outline classification of all bacteria as then known. This was pub-
lished just as another member of the Society, C.-E. A. Winslow, who had, with
his wife, completed a monographic study of the Coccaceae (Winslow, C. -E. A., and
Winslow, A. R. Systematic relationships of the Coccaceae, 300 pp., 1908, John
Wiley & Sons, New York), urged the Society to form a Committee to organize
a better classification for bacteria. The Society of American Bacteriologists'
Committee, of which Winslow was made Chairman, combined forces with Bu-
chanan and published first a preliminary (Jour. Bact., 2, 1917, 505-566) and then
a final report (Jour. Bact., 5, 1920, 191-229) on the classification of bacteria. The
report of this Committee was accepted with the thought that further revisions
of this outline classification were to be expected as knowledge developed.
Meanwhile in Europe, Orla-Jensen (Cent. f. Bakt., II Abt., 22, 1909, 305-346)
had made notable contributions to knowledge in this field. Still later A. J. Kluyver
and C. B. van Niel (Zent. f. Bakt., II Abt., 34, 1936, 369-403) and others con-
tinued the development of classifications of bacteria, but European workers have
been badly handicapped in their work because of the chaotic conditions that
have existed during two world wars fought largely in Europe.
Developments in the field of systematic bacteriology led to the publication by
D. H. Bergey of a manuscript on which he had been working for a long time, his
thought being that a new edition of Chester's Manual of Determinative Bac-
teriology was badly needed, as indeed it was. In order to aid Bergey in securing
publication of his manuscript, the Society of American Bacteriologists appointed
a Committee to assist him, Dr. F. C. Harrison, Chairman. The first edition of
8 CONSIDERATIONS INFLUENCING CLASSIFICATION
Bergey's Manual appeared in 1923 (The Williams & Wilkins Co., Baltimore).
Successive editions of this Manual were issued in 1925, 1930 and 1936. Before
his death in 1937, Bergey requested that an Editorial Board take over future
editions of Bergey's Manual.
At the same time, Bergey used the accumulated royalties that had previously
been placed in the custody of the Society of American Bacteriologists to organize
the so-called Bergey's Manual Trust. The publication of the 5th and 6th
editions of Bergey's Manual has been carried out by the Trustees of this
Trust, who, by the provisions of the Deed of Trust, must always be men trained
as bacteriologists. The Board of Trustees consisted at first of Dr. D. H. Bergey,
Professor R. S. Breed and Professor E. G. D. Murray. Dr. A. Parker Kitchens was
elected to this Board after Dr. Bergey's death, when Professor Breed was made
Chairman of the Board.
Because of the truly enormous development of our knowledge of bacteria,
viruses and related organisms, the Editorial Board asked students of special
groups to assist in the revisions of the groups in which they were interested.
Thus more than 40 specialists assisted in the preparation of the 5th edition,
and more than 60 individuals in the preparation of the 6th edition of the Manual.
Canadian bacteriologists as well as bacteriologists from the U. S. A. have
participated in the Manual work from the beginning. This participation by
Canadian workers has increased during the preparation of the manuscript for
the 7th edition, as has the participation from other countries. Fourteen countries
are represented among the more than 100 specialists who have contributed to
the 7th edition of the Manual.
After the death of Dr. A. Parker Hitchens, Dr. N. R. Smith was appointed
to the Board of Trustees and to the Editorial Board of the Manual, and shortly
thereafter the Board of Trustees was made a board of five members by the
election of Dr. R. E. Buchanan and Dr. Harold J. Conn to this Board.
In preparing manuscripts for the present edition of Bergey's Manual,
specialists have found many places where the relationships of described species
of bacteria have not been well presented in the literature. Consequently, they
have been stimulated to publish many papers reporting their findings. The
individual specialists have normally been persons who have actively worked
with cultures of the organisms that belong to the group for which they have
prepared the manuscript. Thus the development of the present edition of the
Manual has stimulated much research in the field of S3^stematic bacteriology
that would never have been accomplished under other conditions. It is hoped
that in the future the Bergey's Manual Trust can become a center for research
in the field of systematic bacteriology and virology. The work thus far accom-
plished has been carried out largely by volunteer workers. If adequate funds were
available for the support of such work its value could be greatly increased.
Out of studies by specialists of the accumulated knowledge of the systematic
relationships of the microorganisms considered in Bergey's Manual of Deter-
minative Bacteriology, the three of us chiefly responsible for organizing this
CONSIDERATIONS INFLUENCING CLASSIFICATION 9
work (Professor R. S. Breed, Professor E. G. D. Murray and Dr. N. R. Smith)
have developed an outline classification which expresses our ideas of the relation-
ships of the simplest types of living things. These are represented by such common
terms as true bacteria, filamentous bacteria, actinomycetes, slime bacteria,
spirochetes, rickettsias and related larger viruses and the filterable viruses. This
general classification also expresses our ideas of the relationships of these undif-
ferentiated types of living things to higher plants.
This outline may not express the views of other special students of this subject
adequately, as all such outlines represent compromises between differing view-
points. One such difference of viewpoint that has been discussed among the
three of us chiefly responsible for the outline given here has been the question
whether a third kingdom, the Protophyta as defined below, ought not to be
recognized in addition to the Plant and Animal Kingdoms. Prof. E. G. D. Murray
has been the one in our group who has felt most strongly that the bacteria and
related organisms are so different from plants and animals that they should be
grouped in a kingdom equal in rank with these kingdoms. It is quite probable
that support for this viewpoint would be stronger if early biologists had known
how different these important and widely diversified microorganisms are from
plants and animals. Even today it must be recognized that our knowledge of the
number of kinds of bacteria is growing rapidly as habitats not previously ade-
quately explored are studied. The human body is, as a matter of fact, practically
the only habitat that has been comprehensively studied as a source of bacteria.
Even in this case it is the bacteria that cause diseases that are best known.
Our knowledge of the still smaller types of parasitic and pathogenic organisms
such as the numerous kinds of organisms found in the Rickettsiales and Virales
is still more inadequate than our knowledge of the true bacteria. In fact our
present-day knowledge of the filterable viruses could perhaps best be compared
with Cohn's knowledge of the bacteria when he first drew up a system of classifi-
cation for bacteria in 1872.
Three groups are included in the outline presented here: (a) the blue-green
algae, (b) bacteria and related forms, and (c) the rickettsias and viruses.
These are placed in a single division of the plant kingdom for which the term
Protophyta has been used. This name was suggested by a botanist, Sachs (Lehr-
buch der Botanik, 4 Aufl., 926 pp., Wilhelm Engelmann, Leipzig). Recently
Sachs' concept of this group has been developed further by a Russian system-
atist, N. A. Krassilnikov (Guide to the Bacteria and Actinomycetes (Russian),
Izd. Akad. Nauk, Moskau, U.S.S.R., 1949, 830 pp.), and it is developed still
further in the present edition of the Manual.
Of the three names used for the different classes of Protophyta, Schizomycetes
was suggested by von Naegeli (Bericht iiber der Verhandlungen der bot. Section
der 33 Versammlung deutscher Naturforscher und Xrzter. Bot. Ztng., 15, 1857,
760) and Schizophyceae by Cohn (Jahresber. Schles. Ges. f . vaterl. Cultur f. 1879,
279-289), and these have been generally used. The development of our knowledge
of the rickettsias and viruses is so recent that no truly satisfactory class name
10 CONSIDERATIONS INFLUENCING CLASSIFICATION
has previously been suggested for this entire group. This has caused Dr. C. B.
Phihp, who has acted as editor of the section covering rickettsias and related
species in the present (7th) edition of the Manual, to suggest the name Microta-
tobiotes for Class III. The latter is a more appropriate name for the entire group
of organisms included in the orders Rickettsiales and Virales than any that has
previously been suggested. Dr. Philip has discussed the new developments in the
classification of the order Rickettsiales in a recent paper (Canadian Jour. Micro-
biol., 2, 1956, 261). Therefore the present discussion is limited to an explanation
of the reasons for increasing the number of orders recognized in Class II, Schizo-
mycetes von Naegeli, from five to ten.
The organisms placed in Class II, Schizomycetes von Naegeli, in the 6th edition
were arranged in five orders as follows:
Division I. Schizophyta Cohn, 1875. (Fission plants.)
Class I. Schizophyceae Cohn, 1879. (Fission algae. Blue-green algae.)
Class II. Schizomycetes von Naegeli, 1857. (Fission fungi, bacteria.)
Order I. Eubacteriales Buchanan, 1917 (The true bacteria.)
Order II. Aciinomycetales Buchanan, 1917. (The mycobacteria, actinomyces,
streptomyces and related forms.)
Order III. Chlamydohacteriales Buchanan, 1917. (The alga-like, filamentous
bacteria.)
Order IV. Myxobacteriales Jahn, 1911. (The slime bacteria.)
Order V. Spirochaetales Buchanan, 1918. (The spirochetes and related forms.)
Supplements: Groups whose relationships were regarded as uncertain.
Group I. Order Rickettsiales Buchanan and Buchanan, 1938, emend. Gieszczy-
kiewicz, 1939. (Rickettsias and related organisms.)
Group II. Order Virales Breed, Murray and Hitchens, 1944. (Filterable viruses.)
Group III. Family Borrelomycetaceae Turner, 1935. (Pleuro-pneumonia-like organ-
isms.)
This outline as given above is similar to the outline followed in earlier editions
of the Manual and is based upon the outline classification developed by Bu-
chanan (op. cit.) in 1916-18. It is expanded in the present edition of the Manual
as follows :
Division I. Protophyta Sachs, 1874, emend. Krassilnikov, 1949.* (Primitive plants.)
Class I. Schizophyceae Cohn, 1879. (Blue-green algae.)
Class II. Schizomycetes von Naegeli, 1857. (Bacteria and related forms.)
Order I. Pseudomonadales Orla- Jensen, 1921.
Order II. Chlamydohacteriales Buchanan, 1917.
Order III. Hyphomicrobiales Douglas, 1956.
Order IV. Eubacteriales Buchanan, 1917.
* Protophyta was previously used by Endlicher, S. (Genera Plantarum, Vindobonae,
1836, p. 1) in two different senses: (1) for Sectio I, Algae and Lichens of his Regio I Thal-
lophyta, (2) for Cohors II of his Sectio III, Acrobrya, to include horse tails, ferns, etc. This
use may be disregarded under Article 26 of the International Code of Botanical Nomen-
clature. This reads: "The rules of priority and typification do not apply to names of taxa
above the rank of orders.".
CONSIDERATIONS INFLUENCING CLASSIFICATION 11
Order V. Actinomycetales Buchanan, 1917.
Order VI. Caryophanales Peshkoff, 1940.
Order VII. Beggiatoales Buchanan, 1956.
Order VIII. Myxobacterales Jahn, 1911.
Order IX. Spirochaetales Buchanan, 1918.
Order X. Myco-phismatales Freundt, 1955.
Class III. Microtatobiotes Philip, 1955.
Order I. Rickettsiales Buchanan and Buchanan, 1938, emend. Gieszczy-
kiewicz, 1939.
Order II. Virales Breed, Murray and Hitchens, 1944.
Division II. Thallophyia Endlicher, 1836.
Division III. Bryophyta Haeckel, 1866.
Division IV. Pteridophyta Haeckel, 1866.
Division V. Spermatophyta Goebel, 1882.*
It has been felt desirable to subdivide the Order Eubacteriales, as defined in
the 6th edition of the Manual, into Order I, Pseudomonadales, which includes
all of the polar-flagellate types of true bacteria, and Order IV, Eubacteriales,
which includes the peritrichous types of true bacteria. As in the 6th edition, the
photosynthetic purple and green bacteria that are polar flagellate have been
included in the order with the colorless polar-flagellate bacteria. This arrangement
emphasizes a concept first introduced into the classification of bacteria by Migula
(Arb. Bact. Inst. Karlsruhe, 1, 1894, 235-238). This concept is analogous to the
concept used by protozoologists who recognize the orders Flagellata and Infusoria
in Protozoa.
Bacteriologists have recognized differences between polar flagellate and
peritrichous bacteria ever since Migula emphasized them, but there has always
been a residual protest against drawing a sharp line between the two groups of
bacteria. While there is good reason to draw a sharp line between the ordinary
polar flagellate types of bacteria and the peritrichous types, there are certain
groups such as legume nodule bacteria {Rhizobium) , the violet bacteria {Chromo-
bacterium), the agrobacteria (Agrobacterium) and certain motile forms placed in
the family Corynebacteriaceae that present a type of peritrichous flagellation that,
when studied superficially, is misleading. Some cultures of these organisms are
found to show only a single flagellum, while others closely related to these
monotrichous species show several flagella peritrichously arranged. On casual
examination these conditions appear to form a transition between the two types
of flagellation. However, this clearly is not the case. This apparently intermediate
type of flagellation seems to be a comparatively recent development in which
the flagella of certain peritrichously flagellated species have undergone a retro-
gressive specialization. In this the organisms have become primarily dependent
on one flagellum as their chief organ of locomotion. They therefore are included
in Order IV, Eubacteriales, with other peritrichous bacteria.
* Goebel, K., in his edition of Sach's Grundziige der Systematik und speciellen Pflanzen-
morphologie, p. 334, 1882, was apparently the first author to use this name, although he
used the incorrect spelling Spermaphyta.
12 CONSIDERATIONS INFLUENCING CLASSIFICATION
Eubacteriales is defined to include not only the bacteria that are peritrichously
flagellated but also such non-motile forms as seem by their physiology to be
closely related to these peritrichous species.
The placing of non-motile species of bacteria in systems of classification has
always caused difficulty. Some students think that lack of motility is a character
which should be used as a basis for separation of groups. However, evidence is
continually accumulating that indicates that separation of larger groups among
the bacteria solely by means of motility or lack of motility leads to a violent
disarrangement of natural groupings. Some non-motile bacteria present funda-
mental physiologies and other characters that show that they are much like
certain polar flagellate organisms. Such non-motile species are placed in the
classification used here in Order I, Pseudomonadales. However, where non-motile
species show fundamental physiologies and other characters more like those of
peritrichous species, then they have been placed in Order IV, Eubacteriales.
Organisms living in habitats where they are unable to use organs of locomotion
are usually found to be non-motile. This is very natural from the standpoint of
evolution.
Some bacteria develop into trichomes, which may be defined as chains (fila-
ments) of bacteria where the relationship between the cells in the chain have
become so intimate that the cells rarely live a separate, independent existence.
Sometimes the cells in the chain show a differentiation into hold-fast cells and/or
reproductive cells distinct from the usual vegetative cells. This differentiation
resembles that found among the simpler algae. Because the cells in these tri-
chomes sometimes develop flagella that are placed singly or in a tuft near or at
the pole of the cell, while others develop cells with peritrichous flagella, it has been
felt desirable to recognize two orders among these bacteria that occur in tri-
chomes: Order II, Chlamydobaderiales, for the polar flagellate types and Order VI,
Caryophanales, for the peritrichous types. Some non-motile species occur in these
orders also.
Little is known about the relationships of certain species of bacteria which
show a budding form of reproduction that is different from the simple cell division
(fission) that takes place in the four orders previously discussed. Only a few of
these species that reproduce by budding are well known, though some of them
occur abundantly in suitable habitats. Because the indications are that many
species of these organisms exist in nature. Prof. H. C. Douglas has set these apart
in a new order, Hijphomicrohiales, p. 276. Where flagellation has been observed
among these budding forms, it is of the polar type so that Order III has been
associated with Order I, Pseudomonadales, and Order II, Chlamydohacteriales,
in the arrangement of the 10 orders as given above.
Until recently everyone has thought of Order V, Actinomycetalcs, as including
species all of which were non-motile.* However Couch, in a series of papers,
the latest published in 1955 (Jour. Elisha Mitchell Sci. Soc, 71, 1955, 148-155),
* Also see footnote p. 713 for a discussion of motility in species of Nocardia by H. L.
Jensen.
CONSIDERATIONS INFLUENCING CLASSIFICATION 13
has shown that microorganisms that belong in this order sometimes exist in
water rather than as pathogens affecting animals or plants or in soil. These
water-inhabiting, saprophytic types of Actinomycetes have developed sporangia
in which motile or non-motile spores may develop. In a way they are analogous
to the so-called water molds. The structure of the vegetative cells and mycelia
of these water-inhabiting Actinomycetes is like that of the aerobic Actinomycetes.
Order VII, Beggiatoales, has been organized by Dr. R. E. Buchanan, page 837,
to include a group of bacteria, primarily ocurring in trichomes, that are motile
but which lack flagella. In spite of this lack they have the power to glide, roll or
oscillate as do certain species of blue-green algae. While none of these bacterial
types develop photosynthetic pigments, they are frequently and apparently
quite properly regarded as colorless, saprophytic forms of blue-green algae.
Certain species oxidize sulfur compounds with the liberation of free sulfur gran-
ules. Some specialists prefer to transfer this group to Class I, Schizophyceae, as
colorless species of blue-green algae rather than to include them with Class II,
Sch.izomycetes. As bacteriologists have been primarily responsible for developing
our knowledge of the species in this order, they are retained here in Class II,
Schizomycetes.
Our knowledge of Order VIII, Myxobacterales, the so-called slime bacteria,
was first developed by botanists rather than bacteriologists. These organisms
occur in leaf mold and on the dung of animals. Recently species causing diseases
of fish have been found. The cells of these species move with a flexuous motion
in a slime which normally grows up into fruiting bodies large enough to be visible
to the naked eye.
The organisms placed in Order IX, Spirochaetales, have always been set off
by themselves though certain species are knoA\Ti that are so much like other
species of bacteria placed in the genus Spirillum in Order I, Pseudomonadales,
that they may be regarded as transitional forms. Sometimes, without sufficient
justification, these spirally twisted organisms have been placed among the
Protozoa.
The tenth order of Class II, Schizomycetes, is the newly organized Order X,
Mycoplasmatales Freundt. Because a review of the nomenclature of the pleuro-
pneumonia-like organisms (Buchanan, Cowan and Wiken, Internat. Bull. Bact.
Nomen. and Taxon., d, 1955, 13-20) has shown that the first generic name applied
to these organisms that has a legitimate standing is M mycoplasma Nowak (Ann.
Inst. Past., 43, 1929, 1330-1352), this name has been adopted for use in the
classification of the pleuropneumonia-like organisms that has been prepared
by Freundt (Internat. Bull. Bact. Nomen. and Taxon., 6, 1955, 67-78). This
generic name has also been used by Edward (Internat. Bull. Bact. Nomen. and
Taxon., 5, 1955, 85-93). While other order names, such as Borrelomycetales
Turner (Jour. Path, and Bact., 41, 1935, 1-32), have been suggested, the generic
name Borrelomyces Turner on which the order name is founded has never come
into general use, and Borrelomyces is in fact an illegitimate homonym of Myco-
plasma Nowak. Acceptance of the order name Mycoplasmatales is in accordance
14 CONSIDERATIONS INFLUENCING CLASSIFICATION
with the principles of the Bacteriological Code of Nomenclature, and it should
tend to stabilize the nomenclature of this group.
In closing this discussion of the revised classification, it should again be em-
phasized that it has been developed as a result of a study of the ideas which
previous workers have expressed in preparing the outlines that they have sug-
gested. Previous workers have laid what appears, with our present knowledge,
to be a satisfactory foundation on which to build. The introduction of new ideas
has come about largely as a result of continuous study of the literature and
conferences with our colleagues, including the specialists who have contributed
and are contributing so much to the knowledge that has accumulated in past
years. A generous share of the credit for the things that constitute a real advance
in our knowledge should go to these specialists. Where the classification here
presented has defects, it is to be hoped that they will be discovered promptly
and eliminated.
HOW BACTERIA ARE NAMED AND IDENTIFIED
Prof. R. E. Buchanan
Dean of Graduate School {Emeritus) atid Director of Experiment Station (Emeritus),
Iowa State College, Ames, Iowa
A manual of determinative bacteriology, such as the present volume, has
several important functions. These should be recognized and understood by the
student if he is to use the volume with satisfaction.
First, the manual should list and describe all the kinds (species) of bacteria
and viruses known through adequate publication in bacteriology or virology.
Obviously, however, only those organisms that have usable published descrip-
tions can be included.
Second, the manual should arrange the descriptions of the kinds (species) in
smaller or larger groups (taxa, singular, taxon) on the basis of resemblances and
differences in an effort to show inter-relationships.
Third, the manual should indicate for each species its correct name, likewise
the correct name for each group (taxon) of related species.
That branch of biology which has for its purpose the orderly arrangement of
the descriptions of species and other taxa, together with the application of the
correct names, is termed taxonomy.
The manual, through its indices, should enable the student who knows the
correct name of an organism (or even a synonym of such name) to discover the
description of the organism and its characteristics, as well as something of its
relationships. If, on the other hand, he has an organism whose characteristics
and description he has determined or recorded, but whose name and relation-
ships he does not know, a satisfactory manual, through its keys, should enable
him to determine the correct name, its probable relationships and its position in
a classification.
Nomenclature. The necessity for applying names to species or kinds of bacteria
and to groups of inter-related organisms is self-evident. A name given by one
person should be understood by others, and as far as practicable all individuals
should use the same name for the same kind of organism. It is helpful, therefore,
if there can be agreement regarding the method of naming bacteria and agree-
ment as to the correct name for each kind or species. Nomenclature includes all
discussions as to methods of naming and of the correctness of particular names.
What kinds of names are used. Two kinds of names are commonly given
to the different species of plants and animals, (1) the common, provincial, ver-
nacular or casual names and (2), the international or scientific names. These
should be carefully differentiated, and their respective advantages and disad-
vantages noted.
It is inevitable, and on the whole probably desirable, that for each kind of
15
16 HOW BACTERIA ARE NAMED AND IDENTIFIED
familiar animal or plant in each language there will be coined a name. Usually
the name for the same organism will be different in each language. For example,
we have in English Oak, in German Eiche, in Latin Quercus, etc. For many less
common kinds, however, there may be no such vernacular names developed.
There have been, of course, many casual or vernacular names given to kinds of
bacteria. In English Ave speak of the tubercle bacillus, the typhoid germ, the
gonococcus, the Welch bacillus, the golden pus coccus, and many others. Simi-
larly, we find in German Typhusbazillus and in French bacille typhique, entero-
coque, etc. Not infrequently scientific names may be adopted into a modern
language and converted into vernacular names. For example, the English name
aster and the scientific generic name Aster are applied to the same group. This
is freciuently a convenience, and in general this practice is to be commended.
For example, many of the "scientific" generic names used in bacteriology are
also used as names in English and other languages. This adaptation is particu-
larly convenient when the organisms in the group under discussion are of im-
portance and are frequently referred to in the literature. Custom and nomen-
clatural rules suggest certain discretion and appropriateness in the use of these
casual or vernacular names. The following suggestions, based upon nomenclatural
precedent and custom, should prove useful to the student.
1. The name of a genus is a noun in the singular. It is not a collective noun
and should never be used with a plural verb. Do not use such an expression as
"The Salmonella are abundant."
2. However, custom since the beginning of binomial nomenclature has sanc-
tioned the use of the plural of generic names. One may say "The Sahnonellae
(Corynehacteria, Rhizohia, Sarcinae, Bacilli) are." These Latin plurals are used
with the meaning "The species of the genus Salmonella (etc.) are." They do not
connote the existence of more than one genus Salmonella.
3. Custom has also sanctioned the use of the generic name in the singular in
an expression such as "This Sarcina is yellow" with the meaning "This species
of Sarcina is j^ellow."
4. The Latin plural of a generic name should be employed whenever the name
is used as indicated in 2 above. "The Salmonellas or Sarcinas are ..." should be
avoided because of the use of the English plural endings.
5. An English (vernacular) name may be coined from any generic name. This
is done usually only for genera that are under length}^ discussion or of consider-
able economic significance.
6. An English, or vernacular, name of a genus may be used also in the plural,
as in "the corynehacteria are . . ." with the meaning "The species of coryne-
bacterium under consideration." When a Latin generic name is converted into
the English vernacular, either the English or the Latin plural may be used.
Perhaps the Latin plural is the better choice, but one finds "The salmonellas
are ... ." Note that when used as a vernacular (English) word the generic name
is never regarded as a proper noun and is not capitalized or italicized.
More than one form of a name may be derived in English (vernacular) from a
HOW BACTERIA ARE NAMED AND IDENTIFIED 17
generic name. A member of the genus Spironema may be termed a spironema
or a spironeme, a member of the genus Sireptomyces a streptomyces or a strepto-
mycete.
7. A genus includes usually se\eral to many species; it is the name of a group
of species. The expression "The genus Salmonella is . . ." or ''Salmonella is . . ."
should always be preferred to such ambiguous phrases as "The Salmonella
^roup is."
In contrast to common, vernacular or casual names, the scientific name for
each kind of organism is planned to be the same in all countries and in all lan-
guages. When a correct scientific name is used, no question should arise in any
language as to what organism is intended. The names thus applied are supposed
to conform to certain general rules.
International codes of nomenclature. In order that there be correct
scientific names, it is essential that there be international agreement as to the
rules governing their creation. Botanists and zoologists have met in numerous
international congresses in which delegates were accredited from the great
botanical and zoological societies, museums and educational institutions of the
world. Codes of nomenclature, designed to tell how names of taxa should be
published and to list the criteria of correctness, have been developed. These
codes or lists of rules and recommendations are quite similar in essentials for
botany and zoology, although they differ in some details.
The question arose in bacteriology: Are either or both of these codes satis-
factory or adaptable to the use of microbiologists? Three views have been ex-
pressed by various writers. Some few suggested that the naming of bacteria
cannot well conform to the approved international rules as their classification
involves considerations not familiar to botanists and zoologists generally. The
second group insisted that unicellular forms of life are neither plants nor animals,
but Protista, and that taxonomic rules, etc., should be distinct for this group
and coordinate with the corresponding rules for plants and for animals. The third
view, more commonly expressed, was that the bacteria are sufficiently closely
related to the plants and animals so that (in so far as they apply) the interna-
tional agreements of the botanists (or zoologists) should be used as a basis for
naming them.
International opinion on this topic was finally crystallized by resolutions
adopted by the First International Congress of the International Society for
Microbiology held in Paris in 1930. These resolutions, approved also by the ple-
nary session of the International Society for Microbiology, were in part as fol-
lows :
"It is clearly recognized that the living forms with which the microbiologists
concern themselves are in part plants, in part animals, and in part primitive. It
is further recognized that in so far as they may he applicable and appropriate the
nomenclatural codes agreed upon by International Congresses of Botany and
Zoology should be followed in the naming of micro-organisms. Bearing in mind,
however, the peculiarly independent course of development that bacteriology
18 HOW BACTERIA ARE NAMED AND IDENTIFIED
has taken in the past fifty years, and the elaboration of special descriptive cri-
teria which bacteriologists have of necessity developed, it is the opinion of the
International Society for Microbiology that the bacteria constitute a group for
which special arrangements are necessary. Therefore the International Society
for Microbiology has decided to consider the subject of bacterial nomenclature
as a part of its permanent program."
The International Society established a permanent Nomenclature Committee
to pass upon suggestions and to make recommendations. This committee is
composed of members (about 100 in all) from the participating nations. Two
permanent secretaries were named, one to represent primarily medical and
veterinary bacteriology, and one to represent other phases of bacteriology.*
It soon became apparent that the botanical and zoological codes of nomen-
clature included many items having no significance in bacteriology and virology
and that bacterial and viral nomenclature required special consideration.
In 1936, at the London International Microbiological Congress, it was decided
that an independent, but closely integrated. Code of Bacteriological Nomen-
clature be developed. In 1939, at the next International Congress, a Judicial
Commission of fourteen was appointed and directed to prepare a code for con-
sideration at the next Congress. The International Code of Bacteriological
Nomenclature prepared by the Commission was approved in 1947 by the Inter-
national Committee and by the plenary session of the Copenhagen Congress.
These rules were published in English in March, 1948, f and later in French,
Spanish, German and Japanese.
The Code was amended at Rio de Janeiro in 1950 and at Rome in 1953. The
present code should be accessible to all bacteriologists and virologists. It has
been edited and annotated by the Editorial Board of the Judicial Commission.}
It should be consulted by all who wish to determine the correctness of names used
in the literature and by those who describe new species or other taxa.
Some general principles of nomenclature. Every student of bacteriology
should be familiar with certain rules of nomenclature if he is to use names in-
telligently. If he wishes to correct names improperly used or if he desires to
name new species, some additional rules should be observed:
1. Each distinct kind of bacterium is called a species.
2. To each distinct species a name is given consisting usually of two Latin
words, as Bacillus suhtilis.
3. The first word is the name of the genus or group to which the organism
belongs. It is always written with a capital letter. It is a Latin or latinized Greek
* The permanent secretary for medical and veterinary bacteriology at the present time
is Dr. S. T. Cowan, National Collection of Type Cultures, Central Public Health Labora-
tory, Colindale Avenue, London, N.W. 9, England. The permanent secretary for general
bacteriology at the present time is Dr. T. Wiken, Laboratory for Microbiology, Technical
University, Delft, Holland.
t International Bacteriological Code of Nomenclature. Edited by R. E. Buchanan,
R. St. John-Brooks and R. S. Breed. Jour. Bact., 55, 1948, 287-306.
t In press, 1956.
HOW BACTERIA ARE NAMED AND IDENTIFIED
19
word, or a new word compounded from Latin or Greek stems, or it may be de-
rived from some other language; but whatever its origin, when used as a generic
name, it must he regarded as a Latin noun. If it is a word not found in classic
Latin, it is still to be treated as Latin. Some examples of generic names in bac-
teriology which are Latin or which are formed from Latin roots are: Bacillus
(masculine) a small rod; Cristispira (feminine) a crested spiral; Lactobacillus
(masculine) a milk small rod; Sarcina (feminine) a packet or bundle. Many
others are words from the Greek or are compounded from Greek roots, the words
transliterated into Latin letters and with endings modified in conformity with
Latin usage; some words of Greek origin are Micrococcus (masculine) a small
grain (sphere); Bacterium (neuter) a small rod; Clostridium (neuter) a small
spindle; Corijnebacterium (neuter) clubbed small rod; Actinomyces (masculine)
ray fungus. Other generic names have been given in honor of persons or places as
Beggiatoa (feminine), Borrelia (feminine), Eherthella (feminine), Pasteurella
(feminine), Erwinia (feminine), Zopfius (masculine).
4. The second word in the scientific name of a species is a specific epithet. It
is not capitalized (some authors capitalize species names derived from proper
nouns). The specific epithet may be:
(a) An adjective modifying the noun and indicating by its ending agreement
with the generic name in gender, as Bacterium album (white Bacterium), Bacillus
albus (white Bacillus), Sarcina alba (white Sarcina), Eberthella dispar (different
Eberthella), Bacterium variabile (variable Bacterium) , Brucella melitensis (maltese
Brucella), Bacillus teres (rounded Bacillus), Bacillus graveolens (sweet-smelling
Bacillus).
Typical adjectives
Masculine
Feminine
Neuter
white
albus
alba
album
black
niger
nigra
nigrum
delicate
tener
tenera
tenerum
sharp
acer
acris
acre
variable
variabilis
variabilis
variabile
different
dispar
dispar
dispar
Uke a berry
coccoides
coccoides
coccoides
gas-forming
aerogenes
aerogenes
aerogenes
(b) An adjective in the form of the present participle of a verb, as Clostridium
dissolvens (the dissolving Clostridium, in the sense of the Clostridium which is
able to dissolve). Bacillus adhaerens (the adhering Bacillus), Acetobacter ascen-
dens (the climbing Acetobacter), Bacillus esterifix^ans (the ester-producing Bacil-
lus). The endings for present participles used as adjectives are the same for all
genders. The past participle is used occasionally, as in Pseudomonas aptata (the
adapted Pseudomonas), Spirillum attenuatum (the attenuated Spirillum).
(c) A noun in the genitive (possessive) modifying the generic name. There is
no necessary agreement in gender or number. Examples, Clostridium welchii
20 HOW BACTERIA ARE NAMED AND IDENTIFIED
(Welch's Clostridium), Salmonella pullorum (the Salmonella of chicks), Strepto-
coccus lactis {the Streptococcus of milk), Brucella abortus (the Brucella of abortion),
Clostridium tetani (the Clostridium of tetanus), Diplococcus pneumoniae (the
Diplococcus of pneumonia), Salmonella anatum (the Salmonella of ducks).
(d) A noun in apposition, that is, an explanatory noun. This does not agree
necessarily with the generic name in gender. This method of naming is relatively
uncommon in bacteriology. Examples are Actinomyces scabies (the scurf or scab
Actinomyces) , Bacillus lacticola (the milk-dweller Bacillus), Rhizobium radicicola
(the root-dweller Rhizobium), Salmonella london (The London Salmonella).
5. The author of the name of a taxon is often cited by having his name follow
that of the species, as Bacillus subtilis Cohn. Sometimes the name of another
author is indicated also in parentheses, as Micrococcus luteus (Schroeter) Cohn.
This means that Schroeter first named the species, giving it the specific epithet
luteum, (placing it in the genus Bacteridium) . Cohn transferred it to the genus
Micrococcus. It should be noted that the name of a person following that of an
organism frequently is not that of the individual who first discovered or described
it, but of the person who first gave it the accepted name. For example, Clostrid-
ium welchii (Migula) Holland was first described by Dr. Wm. H. Welch, but
not named by him. It was named by Migula in honor of Dr. Welch and later
placed in the genus Clostridium by Holland.
6. Sometimes species of bacteria are subdivided into subspecies or varieties.
These are likewise given Latin designations, and the entire name written, as:
Streptococcus lactis subspecies (var.) maltigenes (the Streptococcus of milk produc-
ing malt flavor), or merely Streptococcus lactis maltigenes.
Some principles of taxonomy. The student of bacteriolog}^ should recognize
the meaning of certain terms used regularly in classifications.
(1) Species (plural species). A species of plant (or animal) is assumed above
to be one kind of plant. But how much difference must exist between two cul-
tures of bacteria before one is justified in regarding the organisms in them as
being of distinct kinds or species? No rule can be laid down. It depends largely
upon convenience and upon more or less arbitrary but considered decision. As
stated by Hitchcock (Descriptive Systematic Botany, New York, 1925, p. 8):
"The unit of classification is a coherent group of like individuals, called a species.
The term is difficult to define with precision because a species is not a definite
entity, but a taxonomic concept." Hucker and Pederson (New York Agric.
Exper. Sta. Tech. Bull. 167, 1930, p. 39) state: "The difficulty met with among
these lower forms in dividing them into well-defined groups has led many to
question whether these small groups of 'species' are natural groups and whether
such groups can be considered to be similar to 'species' among higher forms.
However this may be, it is necessary to arrange bacteria as well as possible into
groups or so-called 'species' for convenience in classification," and again (Hucker,
New York Agric. Exper. Sta. Tech. Bull. 100, 1924, p. 29), "characters appli-
cable to the differentiation of species must evidence a certain amount of con-
stancy when studied over a large series of tests. Furthermore, characters adapted
HOW BACTERIA ARE NAMED AND IDENTIFIED 21
to the differentiation of larger natural groups or genera should, in addition to
constancy, show some correlation with other constant characteristics. The pres-
ence of this relationship or correlation between characters for the division of
genera indicates that the groupings are being made along natural rather than
artificial lines."
Type culture. It is quite evident that when a new species of bacterium is de-
scribed, it must include the particular culture from which the species description
was made. This original culture is termed the type culture. One may develop a
definition as follows: A species of bacterium is the type culture or specimen
together with all other cultures or specimens regarded by an investigator as
sufficiently like the type (or sufficiently closely related to it) to be grouped with
it. It is self-evident that different investigators may not draw the same bound-
aries for a given species. There are some practical difficulties, but no better
definition has been evolved.
(2) Genus (plural genera). A genus is a group of related species. In some
cases a genus may include only a single species (is said to be monotypic); in
most cases several to many species are included in a genus. The ciuite pertinent
ciuestion should be asked: How close must be the resemblances (how close the
relationships) among the species of a group to entitle them to inclusion in the
same genus? In other words, how is it possible to delimit accurately the bound-
aries of a genus? This is a matter on which there is no agreement, and probably
can be none. Much of the confusion in modern bacteriological terminology is to
be attributed to this fact. Nevertheless, in the course of time experience tends to
delimit many genera with reasonable accuracy. As stated by Hitchcock (De-
scriptive Systematic Botany, New York, 1925, p. 9): "Convenience may play
a role in determining generic lines. Extremely large groups may be broken up
on the basis of differences of smaller degree not common to a group of closely
allied species, than if the group consisted of a few species. In general, the botanist,
in delimiting genera, keeps in mind two important rec^uirements, that of showing
natural aflftnities and that of aiding correct identification."
However, a genus may be defined helpfully in another way. One of the species
described as belonging to a genus is designated as the type species; a genus may
therefore be defined as including this type species together with such other
species as the investigator (or taxonomist) regards as sufficiently closely related.
It is apparent that some authors may draw the lines narrowly, others broadly.
Some early authors, for example, recognize only two genera of rod-shaped bac-
teria, one for those without endospores {Bacterium), and one for those producing
endospores {Bacillus). These genera thus defined are very large, each containing
hundreds, perhaps thousands, of species. Other students break up these large
genera into many smaller ones. There is not much point to the question as to
which is right and which is wrong. A better question is, which is the more con-
venient, better represents relationships, better facilitates diagnosis, and proves
most useful.
(3) Family. A family in taxonomy is a group of related genera one of which
22 HOW BACTERIA ARE NAMED AND IDENTIFIED
is designated as the type genus. In general the name of the family is formed from
the name of the type genus by affixing the suffix -aceac to the stem of the generic
name. The word is plural. Among bacterial families commonly recognized are
Bacillaceae, named from its type genus, Bacillus^ Pseudomonadaceae from Pseu-
domonas, Spirochaetaceae from Spirochaeia, Actinomycetaceae from Actinomyces
and Spirillaceae from Spirillum.
(4) Order. An order is a group of related families. It is usually named by
substituting the suffix -ales for -aceae in the name of the type family. Among
ordinal names that have been used in bacteriology are Actinomijcetales, Spiro-
chaetales, Myxohacterales.
(5) Class. A class is a group of related orders. In this treatise the bacteria
are treated as constituting the class Schizomycetes in Division I., Protophyta, of
the plant kingdom.
(6) Other categories. Other categories or ranks of names are used for higher
groups. Sometimes families are divided into sub-families, these into tribes, these
into subtribes, and these finally into genera.
How to identify an organism by name. One of the purposes of this Man-
ual OF Determinative Bacteriology, as noted previously, is to facilitate the
finding of the correct scientific name of a bacterium. It is well, however, to note
some of the reasons why this result, the identification of an unknown culture,
may not eventuate. Among these the following may be listed:
(1) The unknown organism awaiting identification b}^ the investigator may
possibly be one which has never been named; or, if named, perhaps was inade-
quately described. Of course it will not be listed in the Manual. Little effort
on the part of bacteriologists has been devoted to describing or naming bacteria
except as they have been found to have some economic significance or to possess
some striking or unusual characteristics. There are quite probably many times
as many species of unknown bacteria as have been described and named. Such
unknown species are all about us. It is not surprising, therefore, if one some-
times encounters undescribed species. When such unnamed species are found,
particularly if they are of economic importance or are related to such forms, it
is highly desirable that they should be adequately described and named, and
the results published and made accessible.
(2) The unknown organism may have been described and named in some
publication, but the description and name have been overlooked in the prepara-
tion of the Manual. Perhaps the description has been so inadequate or incom-
plete that it has not been possible to place it in a satisfactory classification. It
should be noted that the number of species that have been described is so great
that no one individual can know them all. Progress in classification comes about
largely as the result of the work of specialists in particular groups. Unfortunately
most groups of bacteria have not been adequately monographed. It is evidently
the function of a Manual such as this to draw largely upon the work of those
who have published monographs covering special groups of bacteria and to
supplement their achievements as far as possible by a necessarily less satisfactory
consideration of the unmonographed groups. It is clear that the fact that an
HOW BACTERIA ARE NAMED AND IDENTIFIED 23
organism cannot be identified from this text is no proof that it has not been
described and named.
(3) It is possible, of course, that an error has been made in the selection of
the correct name in this Manual. Bacteriological literature has, in recent years,
been engaged in the herculean task of rectifying the nomenclatural blunders of
the past. It is desirable, therefore, that users of the keys and descriptions of this
Manual should be familiar with the rules governing the correct choice of names,
and themselves propose suitable corrections where needed.
Some general rules governing nonienelature that should be known to
students of bacteriology. In summary, some of the more important rules and
recommendations of the Bacteriological Code may be briefly paraphrased. In
case of doubt, the Annotated Code itself should be consulted.
1. Every individual microorganism belongs to a species, every species to a
genus, every genus to a family, every family to an order, every order to a class.
Each one of these ranks is called a taxon (plural taxa) (Principle 7).
2. Each taxonomic group (taxon) with a given definition (circumscription),
position, and rank can bear only one correct name, the earliest name given to it
that is in accordance with the rules of nomenclature (Principle 9) .
3. The name of a species is made up of two words consisting of the name of
the genus followed by the specific epithet. The term "epithet" means a single
descriptive word or a single descriptive phrase. If the latter, the component
words are to be united or joined by a hyphen. Within the same genus, no two
species names may bear the same specific epithet (Rule 6).
4. Each taxon (species, genus, family, order) should have designated a nomen-
clatural type. The type of a bacterial species is preferably a designated culture
preferably maintained in a national type culture collection. When a new species
is described and named, a culture should be deposited by the author with such
type culture collection where it will be available as a standard and useful in
identification of other cultures believed to be related.
The nomenclatural type of a genus is a species of the genus selected in accord-
ance with the rules.
The nomenclatural type of a family is a genus contained within the family.
The family name is formed by adding the ending -aceae to the stem of the name
of the type genus. The nomenclatural type of the family Pseudomonadaceae is
the genus Pseudomonas (Rule 9).
5. Correct names. For the name of a taxon (species, genus, family, etc.) to
be correct it must meet certain requirements. The most important of these are
as follows :
a. The name must be the oldest that conforms to the rules.
b. The name must have been validly published. This means that the name
must have been distributed in printed matter (periodicals, books, other
publications) together with a description or clear reference to a previously
published description. The name must be accepted by the author. It is not
validly published if merely cited as a synonym. A name that has not been
validly published is without standing in nomenclature.
24 HOW BACTERIA ARE NAMED AND IDENTIFIED
c. A legitimate name is one that conforms to all the nomenelatural rules.
d. A correct name of a taxon is that legitimate name which for a given taxon
takes into consideration the boundaries or circumscription of the taxon.
For example, if one author recognizes two species in a genus, each species
will have a correct name determined by the application of the rules. An-
other author may unite the two into a single species, which will have a
correct name under the rules.
6. Citation of authors and names.
a. It is customary in formal use of the name of a species to cite the name of
the author, usually with the year of publication. This means exactly what
it says, one cites the author of the name of the taxon being used. This is
not necessarily the name of the author who first described the organism.
For example, one cites Bacillus suhtilis Cohn, 1872.
b. When a named species is transferred to another genus, the name of the
author who proposed the specific epithet is inserted in parentheses between
the new species name and the name of the author of the new combination.
For example, Neisser and Kuschbert in 1883 named an organism Bacillus
xerosis. Lehmann and Neumann in 1899 transferred this organism to their
newly created genus Corynebacterium, correctly retaining the original
specific epithet (as xerose to agree in gender with Corynebacterium), and
the new combination is cited as Corynebacterium xerose (Neisser and
Kuschbert, 1883) Lehmann and Neumann, 1899. Obviously citation of
author and date with the name of an organism is necessary only when the
organism is first mentioned in a publication.
7. Changes in names required by union or segregation of taxa.
a. When a genus is divided into two or more genera, the generic name must
be retained for one of them. The generic name must be retained for the
genus containing the type species.
b. When a species is divided into two or more species, the specific epithet
must be retained for one of them. The specific epithet of the species con-
taining the type must be retained for this species.
c. When a species is transferred from one genus to another, the specific
epithet is retained unless the resulting species name is a later homonym
or a tautonym or unless there is available an earlier validly published
specific epithet.
8. Rejection and replacement of names.
a. A name or epithet must not be rejected, changed or modified merely be-
cause it is badly chosen or disagreeable, or because another is preferable
or better known. Exceptions can be made only by international action
through the Judicial Commission of the International Committee.
b. A name must be rejected if it is illegitimate, that is, if it is contrary to a
rule. There are numerous defects which may make a name illegitimate,
for example, it may have been superfluous when proposed. Exceptions can
be made by international approval through action of the Judicial Com-
HOW BACTERIA ARE NAMED AND IDENTIFIED 25
9. Spelling and gender of names of taxa.
a. The original spelling of a name or epithet must be retained, except in the
case of a typographical error or of a clearly unintentional orthographic
error. It may be difficult to determine when a typographical or orthographic
error has occurred. In cases where there is doubt it is advisable to ask the
Judicial Commission to consider the matter and to render an OPINION
which ^\all be authoritative.
b. The gender of generic names is determined as follows:
(1) A Greek or Latin word adopted as a generic name retains the gender
of the Greek or Latin.
(2) Generic names which are modern compounds formed from two or more
Greek or Latin words take the gender of the last component. If the
ending is changed from that of the original Greek or Latin word, the
gender is determined by the rules of gender of the Greek or Latin
respectively.
(3) Arbitrarily formed generic names, i.e., those not formed from Latin
or Greek, take the gender assigned to them by their authors. Where
the original author did not indicate the gender, the next subsequent
author has the right of choice.
10. Provisions for exceptions to the rules or for their interpretation.
Whenever, in the opinion of any microbiologist, an interpretation of any rule
or recommendation of nomenclature is desirable because the correct application
of such rule or recommendation is doubtful, or the stability of nomenclature could
be increased by the conservation or by the rejection of some name which is a
source of confusion or error, it is recommended that he prepare a resume out-
lining the problem, citing pertinent references, and indicating reasons for and
against specific interpretations. This resume should be submitted to the Chair-
man of the Judicial Commission; if desired, through one of the Permanent
Secretaries. An OPINION will be formulated, which may not be issued until
it has been approved by at least eight members of the Commission.
Before the preparation of an OPINION, a preliminary statement is usually
published in the International Bulletin of Bacteriological Nomenclature and
Taxonomy, the official organ of the International Committee on Bacteriological
Nomenclature, Iowa State College Press, Ames, Iowa, U. S. A.
Those who are interested in the solution of special nomenclatural problems
have open to them as an avenue of communication and publication the columns
of the International Bulletin. The Board of Editors includes the chairman of the
Judicial Commission and the two permanent secretaries. Requests for assistance
in the solution of bacteriological nomenclatural problems may be sent to any
member of the Editorial Board at the following addresses:
Prof. R. E. Buchanan, Chairman of Judicial Commission and of the Editorial
Board. Room 316 Curtiss Hall, Iowa State College, Ames, Iowa, U. S. A.
Dr. S. T. Cowan, Permanent Secretary of the International Committee and
of the Judicial Commission. National Collection of Type Cultures, Central
Public Health Laboratory, Colindale Avenue, London, N.W. 9, England.
26 HOW BACTERIA ARE NAMED AND IDENTIFIED
Prof. Dr. Torsten Wiken, Permanent Secretary of the International Commit-
tee and of the Judicial Commission. Laboratory for Microbiology, Tech-
nical University, Delft, Holland.
Derivation, accentuation and pronunciation oj names of taxa and of specific epi-
thets. A serious attempt has been made in this Manual to give the derivation of
the words used as names of taxa (genera, families, etc.) and of the specific epi-
thets of the species names of the microorganisms described. Some guide to pro-
nunciation is given by designation of the principal accent. The rules clearly state
that all names of taxa are to be treated as Latin. But in modern times the pro-
nunciation of Latin words shows little uniformity. However, the principal accent
can be properly placed. Syllabication of the words may also be helpful.
L No Latin word consisting of two or more syllables is accented on the last
syllable.
2. A Latin word consisting of two or more syllables is accented either on the
next to the last syllable (the penult) or on the second to the last syllable (ante-
penult) .
How may one determine which of the two syllables is to be accented? The rule
is easily stated. If the next to the last syllable (penult) is long, it should be ac-
cented; if short, the preceding syllable (antepenult) is to be accented.
When is a syllable said to be long? There are several criteria; those most readily
recognized are as follows:
L If a syllable has a single long vowel, the syllable is long. A standard Latin
dictionary will indicate whether the vowel is long. In words derived from Greek
those syllables containing omega (w) or eta (77) are long, those with omicron (o) or
epsilon (e) are short. In a Greek lexicon the other vowels are usually marked to
indicate length.
2. If a syllable contains a diphthong, it is long.
3. If there is a double consonant or two consonants following a vowel, the
syllable is long. For example:
Ba.cil'lus. The accent is on the next to the last syllable (penult) because of the
double I.
Bac . te'ri . um. The accent is on the antepenult because the vowel in the penult
is short.
Ba.cil.la'ce.ae. The accent is on the antepenult because the vowel of the
penult is short.
Spi.ro. ne'ma. The accent is on the penult because the vowel of the penult
is long; it is the Greek eta (77).
Micros. pi'ra. The Greek epsilon iota (et) is a diphthong; when translated into
Latin, it becomes a long i, and the accent is on the penult.
Use of Greek and Latin in naming taxa. The Greek and Latin alphabets are not
identical. Greek words to be used as stems for the Latin names of taxa must be
transliterated into Latin (not translated) ; the Greek letters must be changed to
the Latin equivalents. The Latins developed well-recognized rules for doing this.
With most letters the shift is simple, in other cases, the changes are more compli-
HOW BACTERIA ARE NAMED AND IDENTIFIED 27
cated. Before a Latin name of a taxon is formed, the Greek word needs to be
spelled with Latin letters, and the whole word placed, when possible, in the corre-
sponding Latin declension with appropriate gender ending. How can the Greek
deri\'ations be indicated without confusion to the student who knows little or
nothing of Greek? In this Manual the following procedure has been adopted as
standard and as probably the most readily understood. The student must remem-
ber that the change is from Greek to Latin (not to English) orthography. The
system used here is not that usually found in giving derivations in medical or
general English dictionaries. Some illustrations may be helpful.
The Greek word for sulfur is delov. The first letter, theta (6), has no Latin equiv-
alent; the Latins used th. The second letter, epsilon (e), is the equivalent of short
e in Latin. The third letter, iota (t), is equivalent to i, the fourth, omicron (o), is
short 0, and the last, nu (v), is n. One may transliterate as theion. But the et of the
Greek, a diphthong, was transliterated by the Latins as a long i. The ending ov
of the Greek indicates that the noun is neuter. The corresponding neuter ending
in Latin is um. In final form we may write deiop = theion = thium. In the Manual
the statement given is simply Gr. neut. n. (Greek neuter noun) thium sulfur. Thi
is the stem from which a great number of new Latin names of taxa have been
constructed, as Thioploca, Thioderma, Thiocystis.
Some awkward transliterations are to be found in the literature. The Greek
diphthong at = ai was usually transliterated as ae by the Latins. The Greek
alfxa = haima = haema. Haemophilus is correctly spelled ; Hemophilus is not a
"simplified spelling" but an incorrectly spelled modern Latin word.
One finds many errors of transliteration in bacteriological nomenclature. If
corrected, the words should be regarded as alternative spellings (variants) of
the same word and not as two different words.
Sometimes there are incongruities in transliteration of Greek into Latin form
in a single word. For example, the specific epithet of the species Micrococcus lyso-
deikticus is an interesting mixture. The second component of the word is the Greek
8hktlk6(7. The first letter, delta (5), is d; the second, epsilon (e), is short e; the third
(and sixth), iota (t), is short i; the fourth (and seventh) is kappa (k), the Latin c;
the eighth, omicron (o), is short o; and the final, sigma (a-), is s. The Latins used
i for the diphthong et. There is no k in the Latin alphabet. The masculine ending
OS in Greek becomes us in Latin. Hence, beiKTiKocr = deicticos = dicticus. Correct
transliteration would have given lysodicticus instead of the current lysodeikticus.
However, in general, it is well to observe the rule that the original spelling of
the word be conserved, unless it can be regarded definitely as a slip of the pen.
A few generic names have been so commonly incorrectly accented as to consti-
tute accepted exceptions. Several examples may be cited.
Many generic names in bacteriology and protozoology have as the final com-
ponent -monas, as Pseudomonas and Xanthomonas. The Greek word is nopas.
The first vowel is short. Correct accentuation would give Pseu.do'mo.nas, Xan.-
tho'mo.nas, etc. with the accent on the antepenult. There is a tendency to regard
the as long and to place the accent on the penult, giving Pseu.do.mo'nas, the
pronunciation accepted by such dictionaries as Century and Borland.
28 HOW BACTERIA ARE NAMED AND IDENTIFIED
Again, many Modern Latin names of taxa have -myces as the last component.
The Greek is ixvK-q'i = myces in which the first vowel is definitely short. In these
generic names the accent would seem properly to be on the antepenult, as Ac-
ti.no'my.ces and Strep.to'my.ces. The commonly accepted accentuation is
Ac.ti.no. my' ces and Strep . to . my'ces.
There is also some confusion relative to syllabication and accentuation in
Modern Latin names of taxa ending in -oides. The derivation of the ending makes
it evident that the oi is not a diphthong, and the o and i should be differentiated
and separately pronounced. For example, the generic name Bacteroides should
be syllabicated and accented Bac.te.ro.i'des. There has been confusion with the
English diphthong oi, and pronunciation with one less syllable, Bac.te.roi'des, has
been recognized.
Abbreviations . The following areviations are used in the Manual in giving
derivations.
Gr. = Greek. The original Greek spelling is not given in the Manual. As noted
above, the word is transliterated into the Latin alphabet; the gender endings of
the Greek are changed usually to the Latin gender endings of the corresponding
Latin declension. This makes evident the stems* that may be used in construction
of the Modern Latin names. Gr. means latinized Greek.
L. = Latin. Usually this indicates that the word is one used in classic Latin
(or in some cases post-classic Latin) and found in an unabridged Latin dictionary.
M.L. = Modern Latin. A word used as the name of a taxon or as a specific
epithet, to be treated and used as a Latin word, of various derivations but not
classic Latin.
Med. L. = Medieval (sometimes pharmaceutical) Latin. Many words derived
from languages other than Latin were Latinized during the middle ages and uti-
lized in fields such as pharmacy, alchemy and biology. Some Modern Latin names
are derived from these.
fem.
= feminine gender.
n. =
noun.
mas.
= masculine gender.
part.
adj. = particip
neut.
= neuter gender.
V. =
verb.
part.
= participle.
nom.
= nominative.
adj. ■■
= adjective.
gen.
= genitive.
pi. = plural. Note that the names of all taxa higher than the genus are plural
and have plural endings, as Bacillaceae, Actinomycetales.
* The stem to be used in making compounds is not always complete in the nominative.
It is found by dropping the genitive ending. For example, the generic name Actinomyces
has as the genitive, Actinomycetis ; the stem used in compounds is Actinomycet-, hence the
famil}^ name eerived from Actinomyces is Actinomycetaceae, not Actinomycaceae. Note
should be taken of the fact that all Greek words that end in -ma are neuter and have as
genitive -malis. The stem combining form) always ends in mat. For example Treponema,
gen. Treponematis, has as its stem Treponemal- from which one may derive a family name
Treponemataceae (not Treponemaceae) .
DIVISION I. PROTOPHYTA* SACHS, 1874, EMEND.
KRASSILNIKOV, 1949.
(Sachs, Lehrbuch der Botanik, 4 Aufl., 1874, 249; Schizophyta Cohn, Beitr. z. Biol. d.
Pflanzen, /, Heft 3, 1875, 202; Krassilnikov, Guide to the Bacteria and Actinomycetes, Izd.
Akad. Nauk, U.S.S.R., Moskau, 1949, 41.)
Pro.to.phy'ta. Gr. combining form protos first (in time), primordial; Gr. noun phylum
plant; M.L. pi. noun Protophyta primordial plants.
Unicellular organisms and organisms which occur in trichomes. Generally these forms
are too small to be distinguishable to the naked eye. Ordinarily no differentiation of cells
is evident, although those forms that occur in trichomes may show some differentiation
into vegetative and specialized cells of various types (heterocysts, holdfast cells and re-
productive cells). Increase in number of individual cells is normally effected by simple cell
division (fission), rarely by budding; however among the most highly advanced forms,
spores of various types may be developed (endospores, conidia or gonidia). In the highly
specialized parasites such as the viruses, the processes of reproduction have become so
intimately associated with the living protoplasm of the host cells, and the virus particles
are so minute (less than 200 millimicrons in diameter) that the e.xact method of reproduction
has not yet been determined with certainty. For many years it was believed that these
organisms do not possess nuclei; however, in recent years simple types of nuclear bodies
have been demonstrated in many of these organisms, and a nucleus, or at least definite
nuclear material (chromatin), has been found to be present in all cases. Do not contain
chloroplastids, which are found in the cells of the green portions of higher plants. Ubiqui-
tous, occurring in the air, everywhere on the surface of the earth, in and on plants and ani-
mals and even far below the surface of the earth in mine waters.
Key to the classes of division Protophyta.
I. Organisms which possess the photosynthetic pigment phycocyanin in addition to
chlorophyll.
Class I. Schizophyceae, p. 30.
II. Organisms which usually do not contain photosynthetic pigments. None contain phyco-
c\'anin.
A. Reproduction by fission. Cells not normally filterable, though filterable stages are
known in some species.
Class II. Schizomycetes, p. 33.
B. Cells so minute that the exact form of reproduction is not clearly understood as yet.
All possess filterable stages.
Class III. Microtatobiotes, p. 931.
* The sections which characterize the Division Protophyta, the classes, the orders and in
some cases the families have been prepared by Prof. Robert S. Breed, Cornell University,
Geneva, New York.
29
CLASS I. SCHIZOPHYCEAE COHN, 1879.
(Myxophyceae Stizenberger, 1860; Phycochromophyceae Hahenhorst, 18G3 -jCyanophyceen
Sachs, Lehrbuch der Botanik, 4 Aufl., 1874, 249; Cohn, Jahresber. Schles. Ges. f. vaterl.
Cultur, f. 1879, 279-289.)
Schi.zo.phy'ce.ae. Gr. noun schizo cleft, fission; Gr. noun phycus seaweed, alga; M.L.
pi. noun Schizophyceae fission algae.
The organisms in this class are usualh' designated as the blue-green algae and are studied
in connection with other types of algae (green, brown and red) and the higher fungi in
courses in Cryptogamic Botany. However, the blue-green algae differ structurally from all
other types of Thallophyta. On the other hand they resemble the bacteria so that the blue-
green algae and the bacteria are commonly classed in the same Division of the Plant King-
dom.
In order to identify the species of blue-green algae, consult any of the following books:
O. Kirchner, Schizophyceae, in Engler and Prantl, Die Natiirlichen Pflanzenfamilien, I
Teil, Abt. la, 1900, 45-92; Gilbert H. Smith, Cryptogamic Botany, 2nd ed.. Vol. 1, Algae
and Fungi, New York, 1955, 526 pp.; Gilbert H. Smith, Freshwater Algae of the United
States, New York, 1950, 719 pp.
The Schizophyceae are not described further in the present Manual.
CLASS II.
SCHIZOMYCETES VON NAEGELI
By
ROBERT S. BREED
Late Professor Emeritus, Cornell University, Geneva, New York
E. G. D. MURRAY
Research Professor, University of Western Ontario,
London, Ontario, Canada
NATHAN R. SMITH
Senior Bacteriologist, Retired, Plant Industry Station, U. S. Department
of Agrictdture, Beltsville, Maryland
and
Specialists whose names appear on the following pages
in connection with the sections prepared by them
CLASS II. SCHIZOMYCETES VON NAEGELI, 1857.
(Von Naegeli, Bericht Verhandl. d. bot. Section d. 33 Versammling deutsch. Naturfonsch.
u. Arzt. Bot. Ztg., 1857, 760; Bacterien, Cohn, Beitr. z. Biol. d. Pflanzen, /, Heft 2, 1872,
127; Bacteriaceae Cohn, Arch. f. path. Anat., 55, 1872, 237; Schizomijcetaceae DeToni and
Trevisan, in Saccardo, Sylloge Fungorum, 8, 1889, 923; Bacteriales Clements (as an ordinal
name), The Genera of Fungi, Minneapolis, 1909, 8; Schizomycetacea Castellani and Chal-
mers, Manual of Tropical Medicine, 3rd ed., 1919, 924; Mychota Enderlein, Bakterien-
Cyclogenie, 1924, 236; Schizotnyceiae Stanier and van Niel, Jour. Bact., 4^, 1941, 458).
Schi.zo.m\-.ce'tes. Or. noun schiza cleft, fission; Gr. noun myces, mycetis fungus;
M.L. mas. pi. n. Schizomycetes the class of fission fungi.
Tj'pically unicellular plants. Cells usually small, sometimes ultramicroscopic. Fre-
quently motile. For many years it was thought that the cells of Schizomycetes and of the re-
lated Schizophyceae did not possess the nucleus invariably found in the cells of other plants.
However, using modern cytological techniques, investigators have now demonstrated a
true nucleus in bacterial cells. Individual cells maj- be spherical or straight, curved or spiral
rods. These cells may occur in regular or irregular masses, or even in cysts. Where they re-
main attached to each other after cell division, they may form chains or even definite
trichomes. The latter may show some differentiation into holdfast cells and into motile
or non-motile reproductive cells. Some grow as branching mycelial threads whose diameter
is not greater than that of ordinary bacterial cells, i.e., about one micron. Some species
produce pigments. The true purple and green bacteria possess pigments much like or related
to the true chlorophylls of higher plants. These pigments have photosynt^etic properties.
The phycocyanin found in the blue-green algae does not occur in the Schizomycetes . Multi-
plication is typically by cell division. Endospores are formed by some species included in
Eubacteriales. Sporocj'sts are found in Myxobacterales. Ultramicroscopic reproductive
bodies are found in Myco-plasmaiales . The bacteria are free-living, saprophytic, parasitic or
even pathogenic. The latter t3-pes cause diseases of either plants or animals. Ten orders
are recognized.
Key to the orders of class Schizomycetes.
I. Cells rigid. Spherical, rod-shaped (straight or curved) or spiral in form. Sometimes in
trichomes. Motile by means of polar flagella or non-motile.
A. Cells coccoid, straight or curved rods, or spiral in foi-m. Sometimes occur as chains
of cells. Cells may contain photosynthetic purple or green pigments. Not in tri-
chromes. Usually motile by means of polar fiagella. Occasionally non-motile.
Order I. Pseudomonadales, p. 35.
B. Not as above.
1. Cells in trichomes that are frequentl}' in a sheath. Occasionally motile (swarm
spores) or non-motile conidia are developed. The sheaths may contain a deposit
of ferric hydroxide, and the trichomes may be attached to a substrate.
Order II. Chlamydohacteriales , p. 262.
2. Cells reproduce by a process of budding rather than by ordinary cell division (fis-
sion). May be attached to a substrate by a stalk. One genus contains species with
photosynthetic pigments {Rhodomicrobium) .
Order III. Hyphomicrobiales , p. 276.
II. Not as above.
A. Cells rigid. Spherical or straight rod-shaped cells. Occur singly, in chains or in tri-
chomes. Motile by means of peritrichous flagella or non-motile. Not acid-fast.
1. Cells spherical or rod-shaped; no trichomes though chains of cells may occur.
Order IV. Eubacteriales, p. 281.
33
34 ORDER I. PSEUDOMONADALES
2. Cells in trichomes.
Order VI. Caryophanales, p. 830.
B. Not as above.
1. Cells rigid and maj- grow out into a branching mycelium-like .structure which
may even develop chains of aerial conidia giving colonies a superficial resem-
blance to mold colonies. In two genera spores develop within sporangia (sporan-
giospores), and in one of these genera the spores are motile. Where cells occur
singly or in simple branched forms, they are frequently acid-fast.
Order V. Actinomycetales, p. 694.
2. Not as above.
a. Cells rigid, usually large and may occur as coccoid cells or trichomes. Sulfur
granules may occur on the surface or within the cells. Move by a gliding, oscil-
lating or rolling, jerky motion like that of some blue-green algae. No flagella
present.
Order VII. Bcggiatoales, p. 837.
aa. Not as above.
b. Longer or shorter flexuous cells.
c. Cells flexuous, creeping on a substrate. Frequentlj^ pointed at both
ends. Fruiting bodies are usually developed from a thin spreading
colony (pseudoplasmodium). Slime bacteria.
Order VIII. Myxobacterales, p. 854.
cc. Cells in the form of longer or shorter spirals. Swim freely by flexion of
cells.
Order IX. Spirochaetales, p. 892.
bb. Non-motile, highly pleomorphic organisms of a very delicate character.
Possess filterable stages.
Order X. Mycoplasmatales, p. 914.
ORDER I. PSEUDOMONADALES ORLA -JENSEN, 1921.
(Jour. Bact., 6, 1921, 270.)
Pseu.do.mo.na.da'les. M.L. fern. pi. n. Pseudomonadaceae type family of the order; -ales
ending to denote an order; M.L. fem.pl.n. Pseudomonadales the Pseudomonadaceae order.
Straight, curved or spiral, rigid, rod -shaped bacteria. Rarely occur in pairs or chains.
The cells in a few species are ellipsoidal and are frequently spoken of as being coccoid or
even spherical in form. They are usually about 1.0 micron in diameter, but in a few species
the individual cell is larger than is normal for bacterial cells, reaching a size of 3.0 to 14.0
microns in diameter and as much as 100 microns in length. The cells are usually polar flagel-
late. When motile they sometimes bear a single flagellum, in other cases a tuft of flagella.
The flagella are normally found at one or both ends of the cell, but in one genus the curved
cells bear a tuft of flagella that is attached in the middle of the concave side (Selenomonas) .
Non-motile species whose characteristics indicate that thej' belong in this order with closely
related, motile species occasionally occur. Cells are Gram-negative so far as known. The
cells in one sub-order contain pigments that have the power of photosynthesis. The cells
in the second sub-order lack such pigments, as do all other groups of bacteria. The cells
in the first sub-order are photo-autotrophic, while chemo-autotrophic species occur in the
second sub-order. Energy is frequently secured by oxidative processes though there are also
many species that show a fermentative physiology. Cells quite frequently occur in zoogloeal
masses. No endospores are found, and reproduction is by means of fission. Many species
occur in coastal, swamp and pond waters and in soil. Some are parasitic and some are even
pathogenic, causing diseases of fishes and other cold-blooded vertebrates. There are a few
species (cholera, blue pus, etc.) that cause diseases of warm-blooded mammals, including
man.
Kerj to the sub -orders of order Pseudomonadales.
I. Cells contain red, purple, brown or green photosynthetic pigments. Sometimes also
enclose granules of free sulfur.
Sub-order I. Rhodobacteriineae , p. 35.
II. Cells do not contain photosynthetic pigments, although they may produce greenish,
brownish, rose or yellow, diffusible, water-soluble pigments or yellow or red non-water-
soluble pigments. Free sulfur granules may occur within or without the cells (Thio-
bacteriaceae). Ferric hydroxide may be deposited (Caulobacteriaceae) .
Sub -order II. Pseudomonadineae, p. 67.
Suborder I. Rhodobacteriineae Breed, Murray and Kitchens, 1944.*
(Family Rhodobacteriaceae Migula, Syst. d. Bakt., 2, 1900, 1042; Breed, Murray and
Hitchens, Bact. Rev., 8, 1944, 257.)
Rho.do.bac.te.ri.i'ne.ae. M.L. neut.n. Rhodobacteriuin a genus of bacteria; -ineue end-
ing to denote a suborder; M.L. fem.pl.n. Rhodobacteriineae the Rhodobacterium suborder.
* Rearranged and revised by Prof. C. B. van Niel, Hopkins Marine Station, Pacific
Grove, California, July, 1953.
35
V
36 ORDER I. PSEUDOMONADALES
Cells spherical, rod-, vibrio- or spiral-shaped. Diameter of individual cells from less than
1.0 to over 10 microns. Red, purple, brown or green bacteria which contain bacteriochloro-
phj-ll or other chloroph3dl-like green pigments, and which usually also possess one or more
carotenoid pigments. Capable of carrying out a photosynthetic metabolism which differs
from that of green plants in that it does not proceed with the evolution of oxygen, and de-
pends upon the presence of extraneous oxidizable compounds which are dehydrogenated
with the simultaneous reduction of carbon dioxide. As oxidizable substrates, a varietj' of
simple substances can be used, such as sulfide, or other reduced sulfur compounds, molecu-
lar hydrogen, alcohols, fatty acids, hydroxy- and keto-acids, etc. All can be grown in
strictly anaerobic cultures when illuminated. Those members which can grow in the pres-
ence of air can also be cultured in the dark under aerobic conditions. Color depends mark-
edly on environmental conditions; small individuals appear colorless unless observed in
masses. May contain sulfur globules. Described species have largely been found in fresh-
water habitats. Some species occur in marine habitats.
Key to the families of suborder Rhodobacteriineae.
I. Purple bacteria whose pigment system consists of bacteriochlorophyll and various
carotenoids capable of carrying out a photosynthetic metabolism.
A. Contain sulfur globules in the presence of hj'drogen sulfide. The sulfur purple bac-
teria.
Family I. TJiiorlwdaceae, p. 38.
B. Do not contain sulfur globules even in the presence of hydrogen sulfide. All require
organic growth factors. The non-sulfur purple and brown bacteria.
Family II. Athiorhodaceae, p. 53.
II. Green sulfur bacteria containing a pigment system which has the characteristics of a
chlorophyllous compound although it differs from the chlorophyll of green plants and
from the bacteriochlorophyll of the purple bacteria.
Family III. Chlorobacteriaceae , p. 61.
The organisms previously included in the order Thiohacteriales Buchanan do not consti-
tute a taxonomic entity; they represent rather a physiological-ecological community. In
this sense, however, a special treatment of this group as a unit has decided advantages from
a determinative point of view.
When first proposed as a systematic assemblage, the order Thiobacferia Migula (Sj'st. d.
Bakt., 3, 1900, 1039) was intended to include the morphologically conspicuous organisms
which, in their natural habitat, contain globules of sulfur as cell inclusions. Since Wino-
gradsky (Beitr. z. Morph. u. Physiol, d. Bact., I, Schwefelbacterien, 1888) had elucidated
the function of hydrogen sulfide and of sulfur in their metabolism, the characteristic inclu-
sions appeared linked with a hitherto unrecognized type of physiology, viz. the oxidation
of an inorganic substance instead of the decomposition of organic materials. From this
oxidation the sulfur bacteria derive their energy for maintenance and growth.
Two groups of sulfur bacteria could be distinguished, one consisting of colorless, the
other of red or purple organisms. The members of both groups presented an unusual mor-
phology apart from the sulfur droplets : in all cases the individual cells were considerably
larger than those of the common bacteria, while many species grew as distinctive colonial
aggregates. Migula separated these sulfur bacteria into two families, Beggiatoaceae and
Rhodobucteriaceae. Even at that time, however, some difficulties e.xisted as to just what
organisms should properly be considered as sulfur bacteria. Miyoshi (Jour. Coll. Sci., Imp.
Univ., Tokyo, 10, 1897, 143) had discovered a bacterium which forms strands, incrusted with
sulfur, in sulfur springs but which does not store sulfur globules in its cells. Although
physiologically this organism appeared to comply with Winogradsky's concept of a sulfur
bacterium, the absence of the typical cell inclusions made Miyoshi decide it could not be
ORDER I. PSEUDOMONADALES 37
considered as such. The problem was aggravated when Nathansohn, Beijerinck and Jacob-
sen published their studies on small, colorless, Psendomonas-like bacteria capable of oxidiz-
ing hydrogen sulfide, sulfur and thiosulfate, and evidently dependent upon this oxidation
process for their development. Morphologically these organisms have little in common with
the Beggiatoaceae; they were designated by Beijerinck as species of Thiobacillus and have
since been rightly considered as members of the order Psendomonndales (see p. 35). Never-
theless, these organisms are physiologically in no way different from the Beggiatoaceae, so
that if phj'siology only is considered, a good case could be made out for their incorporation
in the Thiohacterinles.
Furthermore, Molisch (Die Purpurbakterien, Jena, 1907, 95 pp.) described in some de-
tail a number of bacterial species which, in view of their characteristic pigment system,
appeared closely related to the Rhodobacieriaceae , but which develop only in organic media
and are, therefore, not sulfur bacteria in the sense of Winogradsk\' or Migula. In stressing
the importance of pigmentation, Molisch combined the red sulfur bacteria and the newly-
discovered purple bacteria into an order Rhodobacteria with the two families Thiorhodaceae
and Athiorhodaceae . It is this grouping that has been followed in the present edition of the
Manual.
Among the non-sulfur purple bacteria, or Athioihodaceae, is included an organism which,
on the basis of its morphology and manner of growth, does not conform to the criteria of
the order Pseudornonadalcs . This is Rhodomicrobium vannielii Duchow and Douglas (Jour.
Bact., 58, 1949, 409). Physiologically it is a typical non-sulfur purple bacterium in that it is
capable of development in strictly anaerobic media supplied with an appropriate oxidizable
substrate only when the cultures are illuminated and carries out a photosynthetic metabo-
lisii without oxygen evolution. Multiplication is not, however, by transverse fission but by
bud formation at the end of a thin filament growing out of a pole of the mother cell followed
by the formation of a cross wall in the connecting filament. This mode of development is
similar to that encountered in the non-photosynthetic bacterium Htjphomicrobium vulgare.
It should also be emphasized here that some of the sulfur purple bacteria (Thiopedia, for
example) and all of the green sulfur bacteria appear at present to be pernianentlj- immotile.
Only a very small number of t^'pical sulfur bacteria have been studied in pure cultures. As
a result the descriptions of genera and species rest mainly on observations made with
collections from natural sources or crude cultures. Most investigators have implicitly ac-
cepted differences in cell size or in colonial appearance as a sufficient justification for es-
tablishing independent species. Evidently this procedure presupposes a considerable degree
of constancy of such characteristics in the organisms in question. It is true that Wino-
gradskj^'s investigations have provided a reasonable basis for this belief, but later studies
with pure cultures of certain purple bacteria have established beyond a doubt that environ-
mental conditions, such as composition of the medium and temperature, may exert a pro-
found influence on the general morphology of these organisms. By this it is not intended to
infer that the previously proposed genera and species of sulfur bacteria should be aban-
doned, but it does follow that a cautious evaluation of the distinguishing features is neces-
sary. In the absence of carefully conducted investigations on morphological constancy and
variability of most of the previously recognized species of sulfur bacteria with pure cultures
grown under a variety of external conditions, the best approach appears to be a tentative
arrangement of these organisms based upon those characteristics which are readily ascer-
tainable. Experience with this group over the past twenty-five years has shown that, while
Winogradskj^'s fundamental work must remain the foundation of present taxonomic ef-
forts, it is advisable to simplifj' the much more elaborate classification developed by Bu-
chanan which was followed in previous editions of this Manual.
Certain genera of sulfur purple bacteria, created by Winogradsky, will very probably be
consolidated when detailed information concerning the morphology of the organisms is
available. Until such time it seems, however, best toj-etain most of them, even though the
38 ORDER I. PSEUDOMONADALES
distinguishing characteristics are not always very clear. For the benefit of those who are
familiar with previous methods of classification, it will be indicated where deviations have
been adopted.
The non-sulfur purple bacteria {Athiorhodacene Molisch; Rhodobacterioideae Buchanan)
have been subjected to a comparative morphological and physiological study comprising
more than 150 strains, among which all previously proposed genera and species are repre-
sented (van Niel, Bact. Rev., 8, 1944, 1-118). It has been found that the characteristics
upon which Molisch based the seven genera of this group are inadequate, and a new classi-
fication with only two distinguishable genera has been proposed. This system will be fol-
lowed here.
Nadson (Bull. Jard. Imper. Bot., St. Petersburg, 13, 1912, 64) described a new type of
small, green bacteria not containing sulfur globules in the presence of hydrogen sulfide but
excreting elemental sulfur. They arephotosynthetic and are capable of growing in anaerobic
culture when illuminated. The green pigment differs from the green plant chlorophylls and
from the bacteriochlorophyll of the purple bacteria but has the characteristics of a chloro-
phyllous compound. These are grouped in the family Chlorobacteriaceae.
FAMILY I. THIORHODACEAE MOLISCH, 1907.
(Die Purpurbakterien, Jena, 1907, 27.)
Thi.o.rho.da'ce.ae. Gr. noun thium sulfur; Gr. noun rhodum the rose; -aceae ending to
denote a family; M.L. fem.pl.n. Thiorhodaceae (probably intended to mean) the family of
sulfur red bacteria.
Unicellular organisms, often developing as cell aggregates or families of variable size
and shape. Single cells have the form of spheres, ovoids, short rods, vibrios, spirals, long
rods or, occasionally, chains. Thej^ occur in nature in environments containing sulfides and
require light for their development; infra-red irradiation of a wave-length extending to
about 900 millimicrons is effective. They produce a pigment system composed of green bac-
teriochlorophyll and yellow and red carotenoids. As a result they appear as bluish violet,
pale purple, brownish to deep red cell masses. Single cells, unless they are of considerable
size, usually appear to be unpigmented. These are anaerobic or microaerophilic organisms
with a photosynthetic metabolism in which carbon dioxide is reduced with the aid of spe-
cial hydrogen donors without the liberation of molecular oxygen. Where these organisms
are found in nature, hydrogen sulfide acts as a hydrogen donor, and sulfur, the first inter-
mediate oxidation product, accumulates as sulfur droplets in the cells. Probably all mem-
bers of the group can utilize a number of organic substances in place of hydrogen sulfide as
hydrogen donors for photosynthesis. Thus they are potentially mixotrophic.
Characterization of the genera in this group has, since Winogradsky's studies (Beitrage
zur Morphologie und Physiologie der Schwefelbacterien, Leipzig, 1888), been based upon
the mode of development of the cell aggregates. Pure-culture studies (Bavendamm, Die
farblosen und roten Bakterien, I. Schwefelbakterien, Pflanzenforschung, Heft 2, 1924, 74
pp.; van Niel, Arch. f. MikrobioL, 3, 1931, 1-112; Manten, Antonie van Leeuwenhoek, 8,
1942, 164 pp.) have shown, however, that not only the sequence of events in the formation
of the aggregates but also the appearance and form of the latter, even including the size
and shape of the component cells, are influenced to a considerable extent by environmental
conditions. This obviously casts doubt upon the usefulness of the previously used diag-
nostic criteria for genera and species. On the other hand, the scope of pure-culture studies
has not yet attained sufficient breadth to warrant the use of a different approach. As a
provisional measure, Winogradsky's genera are therefore maintained. Even the larger
taxonomic units must be regarded as being of tentative value only.
FAMILY I. THIOEHODACEAE 6\)
Key to the genera of family Thiorhodaceae.
I. Cells usually combined into aggregates.
A. Cells grouped as regular sarcina packets.
Genus I. TMosarcina, p. 39.
B. Cells not in sarcina packets.
1. Aggregates in the form of a flat sheet.
a. Cells in regular arrangement, with tetrads as the common structural unit.
Genus II. Thiopedia, p. 40.
aa. Cells in irregular aggregates.
Genus III. Thiocapsa, p. 41.
2. Aggregates in the form of three-dimensional masses.
a. Cells distinctly rod-shaped and arranged in a net-like structure.
Genus IV. Thiodictyon, p. 41.
aa. Cells not so arranged.
b. Cells in a common capsule, individuals rather scattered and loosely
grouped.
Genus V. Thiothece, p. 42.
bb. Cells in rather dense clumps.
c. Aggregates embedded in conspicuous common slime capsule.
d. Aggregates small, compact, often several of them enclosed to-
gether in a common capsule.
Genus VI. Thiocystis, p. 42.
dd. Aggregates large and solid, later break up into small clusters.
Genus VII. Lamprocystis , p. 43.
cc. Common capsule lacking or verj^ transient.
d. Aggregates as a whole exhibit amoeboid movements.
Genus VIII. Amoebobacter, p. 44.
dd. Aggregates devoid of amoeboid movements.
Genus IX. Thiopoly coccus, p. 45.
II. Cells usually occurring singly.
A. Cells clearly spiral-shaped.
Genus X. Thiospirillum, p. 46.
B. Cells not spiral-shaped.
1. Cells irregular, often swollen, distorted, or composed of long, crooked and bent
rods to filaments.
Genus XI. Rhabdomonas , p. 48.
2. Cells regular, spherical to short rods or bean-shaped.
a. Cells spherical, as a rule non-motile, and each one surrounded by a rather
wide capsule.
Genus XII. Rhodothece, p. 50.
aa. Cells ellipsoidal, ovoid, short rods or vibrios, actively motile.
Genus XIII. Chromatium, p. 50.
Genus I. Thiosarcina Winogradsky, 1888.
(Zur Morphologie und Phj'siologie der Bacterien, I. Schwefelbacterien, Leipzig, 1888,
104.)
Thi.o.sar.ci'na. Gr. noun thium sulfur; M.L. fem.n. Sarcina a genus of bacteria; M.L.
fem.n. Thiosarcina sulfur Sarcina.
Individual cells spherical, forming regular cubical packets of sarcina-shape, resulting
from consecutive division in three perpendicular planes. Packets commonly containing
8 to 64 cells. Infrequently motile. Non-spore-forming. Contain bacteriochlorophyll and
40
ORDER I. PSEUDOMONADALES
earotenoid pigments, hence, pigmented purplish to red. Capable of carrying out a photo-
sj'nthetic metabolism in the presence of hydrogen sulfide, cells then storing sulfur globules.
Anaerobic.
The type species is Thiosarcina rosea (Schroeter) Winogradsky.
1. Thiosarcina rosea (Schroeter, 1886)
Winogradsky, 1888. (Sarcina rosea Schroeter ,
Kryptog. -Flora von Schlesien, 3, 1, 1886, 154;
Winogradsky, Zur Morphologie und Physio-
logic der Schwefelbacterien, Leipzig, 1888,
104.)
ro'se.a. L. adj. roseus rosy, rose-colored.
Cells spherical, 2 to 3 microns in diameter,
occurring in packets containing 8 to 64 cells.
Infrequently motile. Color ranging from
purplish rose to nearly black.
Anaerobic.
Habitat : Occur less frequently than other
sulfur purple bacteria; probably widely dis-
tributed in mud and stagnant bodies of
water containing hydrogen sulfide and ex-
posed to light; sulfur springs.
Illustration: Issatchenko, Recherches sur
les microbes de I'ocean glacial arctique,
Petrograd, 1914, Plate II, fig. 5.
Genus II. Thiopedia Winogradsky , 1888.
(Zur Morphologie und Physiologie der Bacterien, I. Schwefelbacterien, Leipzig, 18S8, 85.)
Thi.o.pe'di.a. Gr.n. thium sulfur; Gr.n. pedium a plain, a flat area; M.L. fem.n. Thio-
pedia a sulfur plain.
Individual cells spherical to short rod-shaped, the latter shortly before cell division.
Arranged in flat sheets with typical tetrads as the structural units. These arise from divi-
sions of the cells in two perpendicular directions. Cell aggregates of various sizes, ranging
from single tetrads to large sheets composed of thousands of cells. Non-motile. Non-spore-
forming. Contain bacteriochlorophyll and earotenoid pigments. Capable of photosynthesis
in the presence of hydrogen sulfide, then storing sulfur globules. Anaerobic.
The type species is Thiopedia rosea Winogradsky.
1. Thiopedia rosea Winogradsky, 1888.
(Erythroconis littoralis Oersted, Naturhist.
Tidsskrift, 3, 1840-1841, 555; Winogradsky,
Zur Morphologie und Physiologie der
Schwefelbacterien, Leipzig, 1888, 85.)
ro'se.a. L. adj. roseus rosy, rose-colored.
Cells 1 to 2 microns, often appearing as
slightly elongated cocci regularly arranged
in platelets.
Color, pale red to nearly black, depending
upon the amount of sulfur stored. Red color
visible only with large cell masses, not in
individuals.
According to Winogradsky, the cells are
often embedded in a common slime capsule;
the e.xtensive studies of Utermohl (Archiv f .
Hydrobiol., Suppl. Vol. 5, 1925, 251-276)
make the regular occurrence of such cap-
sules extremely doubtful. On the other
hand, Utermohl emphasizes as quite charac-
teristic the common presence of a relatively
large pseudovacuole, or aerosome, in the
cells of this species encountered in plankton
samples. Winogradskj^ does not mention
this; nevei-theless, it appears to be a regular
and valuable distinguishing feature.
Anaerobic.
Habitat: Widely distributed in mud and
stagnant bodies of fresh, brackish and salt
water containing hydrogen sulfide and ex-
posed to light; sulfur springs. Common,
frequently giving rise to very extensive
mass developments.
Illustrations: Warming, Videnskab.
Meddel. naturhist. Forening, Kjobenhavn,
1876, Plate VIII, fig. 2; Winogradsky, op.
cit., 1888, 85, Plate III, fig. 18; Pringsheim,
Naturwissensch., £0, 1932, 481, the last one
a truly excellent photomicrograph.
FAMILY I. THIORHODACEAE 41
Genus III. Thiocapsa Winogradsky, 1888.
(Schwefelbacterien, Leipzig, 1888, 84.)
Thi.o.ca'psa. Gr.n. ihiiun sulfur; L.n. capsa a box; M.L. fem.n. Thiocapsa sulfur box.
Cells spherical, occurring in families of irregularly arranged individuals held together
in a common slime capsule. The aggregates are spread out flat on the substrate. Motility
not observed. As the colony grows, the capsule bursts, and the cells are spread apart. Gen-
eral morphology and development thus appear similar to that in the genus Aphanocapsa
among the blue-green algae. Contain bacteriochlorophyll and carotenoid pigments; capable
of photosynthesis in the presence of hydrogen sulfide. Under such conditions sulfur is stored
in the form of globules in the cells. This genus is so much like TMothece that it is doubtful
whether a distinction can be maintained.
The type species is Thiocapsa roseopersicina Winogradskj'.
Key to the species of genus Thiocapsa.
I. Individual cells about 3 microns in diameter.
1. Thiocapsa roseopersicina.
II. Individual cells about 1.5 microns in diameter.
2. Thiocapsa floridana.
1. Thiocapsa roseopersicina Winograd- Illustration: Winogradsky, loc. cit., Plate
sky, 1888. (Schwefelbacterien, Leipzig, 1888, IV, fig. 15.
• •/ T J- 2. Thiocapsa floridana Uphof, 1927.
ro.se.o.per.si.ci na. L. adj. roseus rosy; tt i u- i ^
Gr. noun persicus the peach, Persian apple,
Persian; M.L. adj. roseopersicinus rosy-
(Arch. f. Hydrobiol., 18, 1927, 84.)
flo.ri.da'na. M.L. adj . floridanus pertain-
ng to Florida.
^^^.. , ■ 1 r. r o • • 1- Cells spherical, about 1.5 microns in di-
Cells spherical, 2.5 to 3 microns in diam-
ameter. In groups of irregular colonies, each
eter, occurring in families of irregularly ,,,j.rounded by a common capsule, several
arranged individuals held together in a colonies being stuck together. Motility not
common slime capsule. Motility not ob- observed
served. Usually a distinct rose-red. Stored Source: Palm Springs, Florida and Lake
sulfur droplets may attain a considerable Sakskoje, near Eupatoria, Crimea,
size. Habitat: Mud and stagnant water con-
Habitat: Mud and stagnant bodies of taining hydrogen sulfide and exposed to
water containing hydrogen sulfide and ex- light; sulfur springs. Probably ubiquitous,
posed to light; sulfur springs. Illustration: Uphof, ibid., 83, fig. VI.
Genus IV. Thiodictyon Winogradsky, 1888.
(Winogradsky, Schwefelbacterien, Leipzig, 1888, 80; Rhododictyon Orla-Jensen, Cent. f.
Bakt., II Abt., 22, 1909, 334.)
Thi.o.dic'ty.on. Gr. noun thium sulfur; Gr. noun dictyum or dictyoii net; M.L. neut.n.
Thiodictyon sulfur net.
Cells rod-shaped, frequently with pointed ends, somewhat resembling spindles. Form
aggregates in which the cells become arranged end to end in a net-like structure, somewhat
reminiscent of the shape of the green alga Hydrodictyon. The shape is not constant; cells
may also form more compact masses. Sometimes groups of cells separate from the main
aggregate by active movements. Common gelatinous capsule not observed. Contain bac-
teriochlorophyll and carotenoid pigments; cells usually very faintly colored. Capable of
photosynthesis in the presence of hydrogen sulfide, the cells then storing sulfur as small
globules.
42 ORDER I. PSEUDOMONADALES
The type species is Thiodictyon elegans Winogradsky.
1. Thiodictyon elegans Winogradsky, Issatchenko (Etudes niicrobiologiques des
1888. (Schwefelbacterien, Leipzig, 1888, 80.) Lacs de Boue, Leningrad, 1927, 113-114)
e'le.gans. L. adj. elegans choice, elegant. recognizes a forma vmius and a forma
Rods 1.5 to 1.7 by 2.5 to 5 microns; or ^agna, differentiated mainly by the size of
longer just prior to cell division. Usually
contain a large pseudovacuole (aerosome),
leaving a rather thin protoplasmic sheath
along the cell wall
the individual rods.
Habitat: Mud and stagnant water con-
taining hj^drogen sulfide and exposed to
Sulfur droplets generally quite small; light; sulfur springs,
deposited exclusively in the thin proto- Illustrations: Winogradsky, ioc. ci<., Plate
plasmic layer. Ill, fig. 13-17.
Genus V. Thiothece Winogradsky, 1888.
(Schwefelbacterien, Leipzig, 1888, 82.)
Thi.o.the'ce Gr. noun thium sulfur; Gr. noun thece a box, chest; M.L. fem.n. Thiothece
sulfur box.
Sulfur purple bacteria which, in their growth characteristics, resemble the blue-green
alga Aphanothece. Cells spherical to relatively long cylindrical-ellipsoidal, embedded in a
gelatinous capsule of considerable dimensions. Following cell division the daughter cells
continue to secrete mucus which causes the individual bacteria to remain clearly separated
by an appreciable distance; the common capsule thus appears only loosely filled. The cells
may become actively motile and separate themselves from the colony. Such swarmers
closely resemble the cells of certain species of Chromatium. Contain bacteriochlorphyll
and carotenoid pigments. Capable of photosynthesis in the presence of hydrogen sulfide,
producing elemental sulfur as an intermediate oxidation product which is stored as sulfur
globules inside the cells.
The type species is Thiothece gelatinosa Winogradsky.
1. Thiothece gelatinosa Winogradsky, in outermost layers of protoplasm and
1888. (Schwefelbacterien, Leipzig, 1888. 82.) generally small.
ge. la. ti. no'sa. L. part. adj. gre/aiMS frozen. Habitat: Mud and stagnant water con-
stiffened; M.L. noun ffeZafznum gelatin, that taining hydrogen sulfide and exposed to
which stiffens; M.L. adj. gelatinosus gelati- jj^^^. ^^^^^^ springs.
Illustrations: Winogradsky, lac. cit., PI.
nous.
Cells 4 to 6 by 4 to 7 microns, spherical
to cylindrical. Color of individual cells,
faint, often grayish violet or even dirty Univ. Tokyo, Japan, 10, 1897, 170, PI. XIV
III, fig. 9-12; Miyoshi, Jour. Coll. Sci., Imp.
Univ. r
yellowish. Sulfur globules usually deposited fig. 25.
Genus VI. Thiocystis Winogradsky, 1888.
(Schwefelbacterien, Leipzig, 1888, 60.)
Thi.o.cys'tis. Gr. noun thium sulfur; Gr. noun cystis the bladder, a bag; M.L. fem.n.
Thiocystis sulfur bag.
Sulfur purple bacteria which form compact colonies, many of which may be loosely em-
bedded in a common gelatinous capsule. Individual cells spherical to ovoid, often diplo-
coccus-shaped. Colonies may emerge as more or less large units from out of the common
capsule and break up afterwards, sometimes into single swarmers; or the aggregates may
split up inside the original capsule and release small motile units or single swarmers. In
pure cultures frequently develop as single cells and diplococci. Produce bacteriochlorophyll
FAMILY I. THIORHODACEAE 43
and carotenoid pigments, coloring the cell masses purplish to red. Capable of photosyn-
thesis, in the presence of hydrogen sulfide, whereby elemental sulfur is formed as an inter-
mediate oxidation product which is deposited as droplets inside the cells.
The type species is Thiocystis violacea Winogradsky.
Key to the species of genus Thiocystis.
I. Individual cells more than 2 microns in width.
1. Thiocystis violacea.
II. Individual cells about 1 micron or less in width.
2. Thiocystis rufa.
1. Thiocystis violacea Winogradsky, taining hydrogen sulfide and exposed to
1888. (Schwefelbacterien, Leipzig, 1888, 65.) light; sulfur springs.
vi.o.la'ce.a. L. adj. violaceus violet- Illustrations: Zopf, Zur Morphologie der
colored. Spaltpflanzen, Leipzig, 1882, PI. V, fig. 12;
Cells about 2.5 to 5.5 microns in diam- Winogradsky, op. cit., 1888, 65, PI. II. Fig.
eter, spherical to ovoid. Swarmers actively 1-7.
motile by means of polar flagella.
^ , . f, 1, . . , , 2. Ihiocystis rufa vVmogradsky, 1888.
Colonies: Small, inside a common capsule, f lu + • t • • looo /=\
containing not over 30 cells. Several such
colonies form loosely arranged aggregates,
most characteristically composed of about _, , , n i , ^i. • ., ,
,- ^ _„ , . . • 1 IT., Color red, usually darker than in the type
10 to 20 colonies in a single capsule. The re- . ' ,/ n «• j • l
(Schwefelbacterien, Leipzig, 1888, 65.)
ru'fa. L. adj. rujus red, reddish.
Cells less than 1 micron in diameter.
suit is a nearly spherical zoogloea. In small
colonies, the cells appear as rather distinct ""oT bl*'"k
species. When the cells are stuffed with
sulfur globules, the aggregates appear al-
tetrads; in larger colonies, the cells become
The common gelatinous capsule usually
somewhat compressed and the tetrad-like .obtains a far greater number of closely
arrangement may be lost. p^^j^^^ individual colonies than is the case
In pure cultures, the species often fails to Jq Thiocystis violacea.
produce the characteristic capsules; the or- Habitat: Mud and stagnant water con-
ganisms then occur as actively motile single taining hydrogen sulfide and exposed to
cells or diplococci, with little or no slime light; sulfur springs,
formation. No pseudocapsules are formed. Illustration: Winogradsky, loc. cit., PI.
Habitat: Mud and stagnant water con- II, fig. 8.
Genus VII. Lamprocystis Schroeter, 1886.
(In part, Clathrocystis Cohn, Beitr. Biol. Pfl., /, Heft 3, 1875, 156; in part, Cohnia Winter,
in Rabenhorst, Kryptogamen-Flora, 2 Aufl., 1884, 48; not Cohnia Kunth, Enumeratio plan-
tarum, 5, 1850, 35; Schroeter, Die Pilze Schlesiens, in Cohn, Kryptogamen-Flora von
Schle-sien, 3, 1, 1886, 151.)
Lam.pro.cys'tis. Gr. adj. lamprus bright, brilliant; Or. noun cystis the bladder, a bag;
M.L. fem.n. Lamprocystis a brilliant bag.
Sulfur purple bacteria which form more or less large aggregates of cells enclosed in a
common gelatinous capsule. Individual cells spherical to ovoid. Small aggregates closely
resemble those of Thiocystis, even to the extent of the tetrad-like arrangement of cells in
the small colonies. Behavior of the large aggregates during development appears to be
different; the small individual cell groups or colonies do not emerge from the slime capsule
until the initially relatively compact cell mass becomes broken up into smaller clusters,
these eventually forming a somewhat net-like structure. This behavior has been ascribed
to a change in the mode of cell division which at first appears to take place in three per-
pendicular planes and later presumably changes to a division in only two directions. Cells
44 ORDER I. PSEUDOMONADALES
when free are motile by means of polar fiagella. In pure culture also this type rarely, if ever,
produces large aggregates with the development here mentioned as characteristic for the
genus (Bavendamm, Die farblosen und roten Schwefelbakterien, Pflanzenforschung, Heft
2, 1924, 76). This, along with the other similarities, makes it doubtful whether future studies
will result in the retention of the genera Lamprocijsiis and Thiocystis side by side. Produce
bacteriochlorophyll and carotenoid pigments, coloring the cell masses purplish pink to red.
Capable of photosynthesis in the presence of hj'drogen sulfide, storing elemental sulfur as
globules inside the cells.
The type species is Lamprocystis roseopersidna (Kiitzing) Schroeter.
1. Lamprocystis roseopersicina (Kiitz- Winogradsky reports that the cells fre-
ing, 1849) Schroeter, 1886. (Protococcus quently contain pseudovacuoles.
roseopersicinus Kiitzing, Species Algarum, Habitat: Mud and stagnant water con-
Leipzig, 1849, 196; Schroeter, in Cohn, taining hydrogen sulfide and exposed to
Kryptogamen-Flora von Schlesien, 3, 1, light; sulfur springs.
1886, 151.) Illustrations: Warming, Videnskab.
ro.se.o.per.si.ci'na. L. adj. roseus rosy; Meddel. naturhistor. Foren., Kjobenhavn,
Gr. noun persiCMS the peach (Persian apple); 1876, PI. VIII, fig. 3 g; Zopf, Z. Morphol.
M.L. adj. roseopersicinus rosy peach (-col- d. Spaltpflanzen, Leipzig, 1882, PI. V, fig. 8,
ored). 13; Winogradsky, Schwefelbacterien, Leip-
Cells spherical to ovoid, 2 to 2.5 microns zig, 1888, PI. II, fig. 9-15; Bavendamm, Die
in diameter, up to 5 microns long before cell farblosen und roten Schwefelbakterien,
division. Motile by means of polar flagella. Jena, 1924, PI. II, fig. 3.
Genus VIII. Amoebobacter Winogradsky , 18SS.
(Schwefelbacterien, Leipzig, 1888, 71.)
A.moe.bo.bac'ter. M.L. noun Amoeba a protozoan genus; Gr. noun amoebe change, trans-
formation; M.L. noun bacter a rod; M.L. mas.n. Amoebobacter changeable rod.
Sulfur purple bacteria, usually occurring in aggregates composed of many individuals
without a characteristic common capsule. Slime formation can, nevertheless, be observed
with very small colonies. With growth of the individual cells, the capsule bursts and the
cell mass slowly moves out while the bacteria remain united. The colonies change their
shape during growth and in response to environmental influences; the individual cells ap-
pear motile and cause the movements of the entire colon3^ Winogradsky ascribes the co-
herence of the cell masses to the existence of interconnecting protoplasmic filaments be-
tween cells, but these have never been observed, and their occurrence is extremely doubtful.
It is much more probable that the bacteria are held together by mucus, though not so much
of the latter is produced as to form a clearly discernible capsule. Produce bacteriochloro-
phyll and carotenoid pigments. Capable of photosynthesis in the presence of hydrogen
sulfide, then storing sulfur as droplets inside the cells.
The type species is Amoebobacter roseus Winogradsky.
The characterizations of the genera Amoebobacter, Lamprocystis, Thiocystis, Thiocapsa
and Thiothece are based upon the arrangement of individual bacteria in a common capsule.
However, from Winogradsky's descriptions of Amoebobacter and from pure-culture studies
with Thiocystis and Lamprocystis, the capsules have been shown to vary considerably, de-
pending upon developmental stages and environmental conditions. Therefore it is quite
possible that future investigations will show the desirability of restricting the number of
genera.
Key to the species of genus Amoebobacter.
I. Cells spherical to ovoid, al)Out 2.5 to 3.5 microns in diameter and up to 6 microns in
length prior to cell division.
1. Amoebobacter roseus.
FAMILY I. THIORHODACEAE
45
II. Cells distinctly rod-shaped, about 1.5 to 2 microns in width by 2 to 4 microns in length.
2. Amoebobacter bacillosus.
III. Cells spherical, quite small, about 0.5 to 1 micron in diameter.
3. Amoebobacter granula.
1. Anioebobacler roseus Winogradsky,
1888. (Schwefelbacterien, Leipzig, 1888, 77.)
ro'se.us. L. adj. roseus rosy.
Cells spherical to ovoid, 2.5 to 3.5 microns
in width and up to 6 microns in length.
Motile. Often contain pseudovacuoles. Cell
aggregates often form transitory hollow
spheres or sacks with the bacteria occupy-
ing the peripherj^ as a shallow layer. These
are reminiscent of stages in the development
of Lamprocystis.
Habitat: Mud and stagnant water con-
taining hj^drogen sulfide and exposed to
light; sulfur springs.
Illustrations: Winogradsky, loc. cit., PI.
Ill, fig. 1-6.
2. Amoebobacter bacillosus Winograd-
sky, 1888. (Winogradsky, Schwefelbac-
terien, Leipzig, 1888, 78; Thioderma roseum
Miyoshi, Jour. Coll. Sci., Imp. Univ.
Tokyo, Japan, 10, 1897, 158.)
ba.cil.lo'sus. L. dim. noun bacillus a small
staff or rod; M.L. adj. bacillosus full of
(made up of) small rods.
Cells rod-shaped, about 1.5 to 2 microns
by 2 to 4 microns. Cells contain pseudo-
vacuoles (aerosomes). Sulfur globules de-
posited exclusively in peripheral proto-
plasmic layer, usuallj' quite small.
Mij'oshi's incomplete description of Thio-
derma roseum (loc. cit.), type species of
genus Thioderma, is sufficient to make prac-
tically certain that it is identical with
Amoebobacter bacillosus. The description of
Thiodictyon elegans Winogradsky {op. cit.,
1888, 80) suggests that it cannot be distin-
guished from this species.
Habitat: Mud and stagnant water con-
taining hydrogen sulfide and exposed to
light; sulfur springs.
Illustrations: Zopf, Z. Morphol. d.
Spaltpfl., Leipzig, 1882, PI. V, fig. 26-27;
Winogradsky, op. cit., 1888, PI. Ill, fig. 7.
3. Amoebobacter granula Winograd-
sky, 1888. (Schwefelbacterien, Leipzig,
1888, 78.)
gra'nu.la. L. dim. noun granulum a small
grain; M.L. fem.n. granula a small grain.
Cells spherical, small, about 0.5 to 1.0
micron in diameter. Faint pigmentation;
the sulfur inclusions give the cell masses a
black appearance. Aggregates are apt to
consist of closely-knit masses which are
difficult to separate.
When sulfur is stored, a single droplet
usually fills most of the cell. Because of the
high refractive index of this globule, it be-
comes difficult if not impossible to make
accurate observations of the cell shape.
Habitat: Mud and stagnant water con-
taining hj-drogen sulfide and exposed to
light; sulfur springs.
Illustration: Winogradsky, loc. cit., PI.
Ill, fig. 8.
Genus IX. Thiopolycoccus Winogradsky, 1888.
(Schwefelbacterien, Leipzig, 1888, 79.)
Thi.o.po.ly.coc'cus. Or. noun thium sulfur; Gr. adj. poly many; Gr. noun coccus a berry;
M.L. noun coccus; M.L. mas.n. Thiopolycoccus with man}'- sulfur cocci.
Sulfur purple bacteria which form dense aggregates of rather solid construction and ir-
regular shape. The colonies appear, in contrast with those of Amoebobacter, non-motile and
do not tend to form hollow zoogloeal structures by which they are differentiated from
Lamprocystis. Cell masses held together by mucus which does not, however, appear as a
regular capsule. Large clumps may fissure with the formation of irregular shreds and lobes
which continue to break up into smaller groups of cells. Individual bacteria spherical, mo-
tility not observed. Contain bacteriochlorophyll and carotenoid pigments, so that the
aggregates, in accord with the dense packing with individual cells, appear distinctly red.
46 ORDER I. PSEUDOMONADALES
Capable of photosynthesis in the presence of hydrogen sulfide, when the cells store elemen-
tal sulfur as droplets inside the cells.
The type species is Thiopoly coccus ruber Winogi'adsky.
1. Thiopolycoccus ruber Winogradsky, taining hydrogen sulfide and exposed to
1888. (Schwefelbacterien, Leipzig, 1888, 79.) light; sulfur springs.
ru'ber. L. adj. rwber red. Illustrations: Winogradsky, loc. cil., PI.
Cells spherical, about 1.2 microns in di- IV, fig. 16-18; Issatchenko, Recherches sur
ameter. No motility observed. les microbes de I'ocean glacial arctique.
Habitat: Mud and stagnant water con- Petrograd, 1914, PI. II, fig. 7.
Genus X. Thiospirillum Winogradsky, 1888.
{Ophidomonas Ehrenberg, Die Infusionstierchen, Leipzig, 1838, 43; Winogradsky, Schwe-
felbacterien, Leipzig, 1888, 104.)
Thi.o.spi.ril'lum. Or. noun thium sulfur; M.L. dim.neut.n. Spirillum a bacterial genus;
Gr. noun spira a spiral; M.L. neut.n. Thiospirillum suUur Spirillum.
Sulfur purple bacteria, occurring singly as spirally wound cells, motile by means of polar
flagella. Contain bacteriochlorophyll and carotenoid pigments, coloring the cells brownish
to purplish red. Capable of photosynthesis in the presence of hydrogen sulfide, during which
they produce and store, as an intermediate oxidation product, elemental sulfur in the form
of droplets inside the cells.
The differentiation of species in this group has been based exclusively on observations
with material from natural collections and from laboratory mass cultures. The criteria
used are the size and shape of the spirals and the color of the organisms. Not a single repre-
sentative has so far been obtained and studied in pure culture, so that no information is
available concerning the constancy or variability of these characteristics. It is likely, how-
ever, that such properties may be greatly influenced by environmental factors. Hence, the
following key and descriptions of species are apt to be modified when more extensive studies
have been made. The published descriptions of some species make it seem probable that
they should not even be incorporated in Thiospirillum.
The type species is Thiospirillum jenense (Ehrenberg) Winogradsky.
Key to the species of genus Thiospirillum.
I. Width of cells 2.5 microns or more.
A. Color of cells, especially in masses, yellowish brown to orange-brown.
1. Thiospirillum jenense.
B. Color of cells deep red or violet.
1. Cells long, typical spirals; clearly red.
2. Thiospirillum sanguineum.
2. Cells short, slightly curved, vibrio-shaped; color purple to violet-red.
3. Thiospirillum viokiceum.
II. Width of cells less than 2.5 microns.
A. Width of cells 1.5 to 2.5 microns.
4. Thiospirillum rosenbergii.
B. Width of cells about 1 micron.
5. Thiospirillum rufum.
1. Thiospirillum jenense (Ehrenberg, bacterien, Leipzig, 1888, 104; Thiospirillum
1838) Winogradsky, 1888. (Ophidomonas cmssi<A« Hama, Jour. Sci. Hiroshima Univ.,
jenensis Ehrenberg, Die Infusionstierchen, Ser. B, Div. 2, Bot., 1, 1933, 157.)
Leipzig, 1838, 44; Winogradsky, Schwefel- je.nen'se. M.L. adj. jenensis pertaining to
FAMILY I. THIORHODACEAE
47
Jena; named for the city of Jena, Germany,
where Ehrenberg discovered this organism.
Cells 2.5 to 4.0 microns thick, cylindrical,
sometimes pointed at the ends; coiled as
spirals, generally 30 to 40 microns in length,
but may be as long as 100 microns. Complete
turns may measure from 15 to 40 microns
with a wave depth of 3 to 7 microns. Polar
flagellate. Tufted at both ends. Olive-
brown, sepia-brown and reddish brown.
This coloring appears to be the only rec-
ognizable difference from Thiospirilhan san-
(juineiim. Thiospirillum crassum Hama (loc.
cit.), reported to be 3.7 to 4 by 12 to 40
microns and yellowish brown in color, thus
becomes indistinguishable from Thiospiril-
lumjenense; the 80-microns-long Thiospiril-
lum jenense forma maxima Szafer (Bull.
Acad. Sci. Cracovie, S6r. B, 1910, 162) does
not, at present, justify recognition as a
special taxonomic entity.
It is even doubtful whether the observed
color difference between Thiospirillum
jenense and Thiospirillum sanguineum con-
stitutes a valid criterion for their mainte-
nance as two distinct species (Buder, Jahrb.
wiss. Bot., 56, 1915, 534; Bavendamm, Die
farblosen und roten Schwefelbakterien,
Pflanzenforschung, Heft 2, 1924, 131).
Habitat: Mud and stagnant water con-
taining hydrogen sulfide and exposed to
light; more rarelj' in sulfur springs.
Illustrations: Zettnow, Ztschr. f. Hyg.,
24, 1897, PI. II, fig. 49-52; Buder, op. cit.,
1915, 534, fig. 1; Szafer, op. cit., 1910, PI. IV,
fig. 4; Hama, op. cit., 1933, PI. 18, fig. 1, 8a;
PI. 19, fig. 1.
2. Thiospirillum sanguineum (Ehren-
berg, 1840) Winogradsky, 1888. (Ophido-
monas sanguinea Ehrenberg, Verhandl.
Akad. Wiss. Berlin, 1840, 201 ; Spirillum san-
guineum Cohn, Beitr. Biol. Pfl., 1, Heft 3,
1875, 169; Winogradsky, Schwefelbacterien,
Leipzig, 1888, 104.)
san.gui'ne.um. L. adj. sanguineus blood-
colored, blood-red.
Cells cylindrical, sometimes attenuated
at ends, spirally coiled; 2.5 to 4.0 microns in
width, commonly about 40 microns long
with a range of from 10 to 100 microns.
Size and shape of coils variable, complete
turns measuring from 15 to 40 microns in
length and from }4 to }{o of the length in
width. Polar flagellate, usually tufted at
both ends. Individual cells rose-red with a
grayish hue, groups of cells deep red. Sulfur
droplets numerous under appropriate con-
ditions.
Habitat: Mud and stagnant water con-
taining hydrogen sulfide and exposed to
light; rarely in sulfur springs.
Illustrations: Cohn, op. cit., 1875, PI. VI,
fig. 15; Warming, Vidensk. Meddel. natur-
hist. Foren., Kjobenhavn, 1876, PI. VII, fig.
8; Buder, Jahrb. wiss. Bot., 56, 1915, 534,
fig. 2.
3. Thiospirillum violaceum (Warming,
1876) Winogradsky, 1888. {Spirillum vio-
laceum Warming, Vidensk. Meddel. natur-
hist. Foren., Kjobenhavn, 1876, 395; Wino-
gradsky, Schwefelbacterien, Leipzig, 1888,
104.)
vi.o.la'ce.um. L. adj. violaceus violet-
colored.
Cells short and fat, 3 to 4 by 8 to 10
microns, ends smoothly rounded. Slightly
curved, bean- or vibrio-shaped. Onl}^ rarely
are they twisted suggesting a spirillum.
Polar flagellate.
The shape of cell seems to fit the genus
Chromatium rather than Thiospirillum, and
Warming (op. cit., 1876, 395) emphasizes the
resemblance to Chromatium okenii.
Color: Bluish violet; this color may be re-
lated to a scarcity of sulfur droplets in the
cells.
Habitat: Mud and stagnant water.
Illustration: Warming, op. cit., 1876, PI.
VII, fig. 3.
4. Thiospirillum rosenbergii (Warm-
ing, 1875) Winogradsky, 1888. (Spirillum
rosenbergii Warming, Vidensk. Meddel. na-
turhist. Foren., Kjobenhavn, 7, 1875, 346;
Winogradsky, Schwefelbacterien, Leipzig,
1888, 104.)
ro.sen.ber'gi.i. M.L. gen. noun rosen&er^n
of Rosenberg; named for Rosenberg, a
Danish algologist.
Cells 1.5 to 2.5 by 4 to 12 microns; coiled,
with turns of about 6 to 7.5 microns in
length and variable width up to 3 or 4 mi-
crons. Color very dark, due to numerous
48 ORDER I. PSEUDOMONADALES
sulfur globules. Color of protoplasm not losen und roten Schwefelbakterien Jena,
recorded. 1924, 132) or Huber-Pestalozzi (Die Bin-
Habitat: Mud and stagnant water con- nengewasser, ^6, Heft 1, Das Phytoplankton
taining hj'-drogen sulfide and exposed to des Siisswassers, Stuttgart, 1938, 304) that
light. Probably widely distributed, but less the cells ever contain sulfur globules. Only
frequently recorded as the organism is not the red color is emphasized. Consequently,
so spectacular as the large Thiospirillum it is quite possible that this organism be-
jenense and Thiospirillum sanguineum. longs in the genus Rhodospirillum.
Illustration: Warming, op. cit., 1876, PI. Cells 1.0 by 8 to 18 microns; coiled to
X, fig. 12. occupy 1}4 to 4 turns, the latter commonly
4 microns wide by 4 microns long. These
5. Thiospirillum rufum (Perty, 1852) dimensions agree with those of Rhodospiril-
Migula, 1900. {Spirillum rufum Perty, Bern, lum ruhrum (Esmarch) Molisch, and it
1852, 179; Migula, S3^st. d. Bakt. 2, 1900, seems probable that the two organisms are
1050.) identical.
ru'fum. L. adj. rujus red, reddish. Habitat: Found in red slime spots on the
General characteristics presumably those side of a well. Mud and stagnant bodies of
of the genus, although it does not appear water.
either from Perty's description or from that Illustration: Migula, Syst. d. Bakt., 1,
of Migula {loc. cit.), Bavendamm (Die farb- 1897, PI. Ill, fig. 7.
Genus XI. Rhabdomonas Cohn, 1875.
(Cohn, Beitr. Biol. Pfl., 1, Heft 3, 1875, 167; Rhahdochromalium Winogradsky, Schwefel-
bacterien, Leipzig, 1888, 100.)
Rhab.do.mo'nas. Or. noun rhabdus a rod; Or. noun monas a unit, monad; M.L. noun
Manas a protozoan genus; M.L. fem.n. Rhabdomonas the rod monad.
Sulfur purple bacteria, as a rule occurring singly in the form of rather irregular, long
rods to filaments, exhibiting more or less pronounced swellings, or club and spindle shapes.
Filamentous structures, sometimes with constrictions, giving the filament the appearance
of a string of beads. These may be surrounded by a relatively inconspicuous slime capsule
which can be rendered visible by India ink. The less distorted cell t3^pes are frequently
motile b}^ means of polar flagella. Produce bacteriochlorophyll and carotenoid pigments,
coloring the cells pinkish to purplish red. Capable of photosynthesis in the presence of
hydrogen sulfide and then storing sulfur globules as an intermediate oxidation product in-
side the cells.
The status of this genus is doubtful. Winogradsky (loc. cit.) recognized the similarity of
its members to species of Chromatium and the occurrence of many intermediate forms which
make a sharp distinction between the two genera impossible. He preferred the designation
of Rhabdochromatium as a sub-genus. Warming (Videnskab. Meddel. naturhist. Foren.,
Kjobenhavn, 1876, 320 ff.), Nadson (Bull. Jard. Imp(5r. Bot. St. P^tersb., 3, 1903, 116), van
Niel (Arch. f. Mikrobiol., 8, 1931, 61) and Ellis (Sulphur Bacteria, London and New York,
1932, 151) considered the species of Rhabdochroinatium as abnormal growth forms (involu-
tion forms) of corresponding species of Chromatium, while Lauterborn (Verhandl. natur-
histor.-medizin. Vereins, Heidelberg, N.F., 13, 1915, 424), Buder (Jahrb. wiss. Bot., 68,
1919, 534) and Bavendamm (Die farblosen und roten Schwefelbakterien, Pflanzenforschung,
Heft 2, 1924, 129) favor generic rank.
The type species is Rhahdoinonas rosea Cohn.
Key to the species of genus Khabdomonas.
I. Cells not containing calcium carbonate inclusions in addition to sulfur globules.
A. Cells more than 3 microns in width.
1. Rhabdomonas rosea.
FAMILY I. THIORHODACEAE
49
B. Cells less than 3 microns in width.
2. Rhabdomonas gracilis.
II. Cells containing calcium carbonate inclusions in addition to sulfur globules.
3. Rhabdomonas linsbaueri.
1. Rhabdomonas rosea Cohn, 1875.
(Cohn, Beitr. Biol. Pfl., 1, Heft 3, 1875, 167;
Beggiatoa roseo-persicina Zopf, Z. Morphol.
d. Spaltpfianzen, Leipzig, 1882, 30; Rhabdo-
chromatium roseiim Winogradsky, Schwefel-
bacterien, Leipzig, 1888, 100; Rhabdochro-
matium fusiforyne Winogradsky , ibid., 102.)
ro'se.a. L. adj. roseus rosy, rose-red.
Cells uneven in width and length, often
swollen to spindle-shaped, sometimes tend-
ing towards filamentous growth. The great-
est width of a spindle-shaped or fusiform
cell may be close to 10 microns; in the more
filamentous structures it is usually around
5 microns. The length varies between 10 and
30 microns for single cells; filamentous
forms, frequently showing bulges and con-
strictions suggestive of compound struc-
tures in which cell division has been incom-
plete, may attain considerably greater
lengths, up to 100 microns. The ends of
spindle-shaped cells often taper to very fine
points or attenuated fibers; also, filaments
are generally thinner toward the extrem-
ities. Single individuals and short filaments
are motile by means of polar flagella, long
filaments rarely motile. The ends of a fila-
ment may become pinched off and swim
away.
Color rose-red; cells are usually filled with
sulfur globules.
There is no good reason for maintaining
Rhabdomonas fusiformis (Rhabdochromatium
fusiforme Winogradsky) as a separate
species; the variations in size and shape
bring this form well within the range of
Rhabdomonas rosea. Present indications
strongly suggest that the latter species
should be regarded as a peculiar develop-
mental form of Chromatium okenii.
Habitat: Mud and stagnant water con-
taining hydrogen sulfide and exposed to
light; sulfur springs.
Illustrations: Cohn, op. cit., 1875, PI. VI,
fig. 14; Warming, Vidensk. Meddel. natur-
histor. Foren., Kjobenhavn, 1876, PI. VII,
fig. Ic-e; Zopf, op. cit., 1882, PI. V, fig. 2b;
Winogradsky, op. cit., 1888, PI. IV, fig. 9-11,
13-14.
2. Rhabdomonas gracilis (Warming,
1876) Bergey et al., 1923. {Monas gracilis
Warming, Vidensk. Meddel. naturhist.
Foren., Kjobenhavn, 1876, 331; Rhabdochro-
matium minus Winogradsky, Schwefelbac-
terien, Leipzig, 1888, 102; Rhodocapsa sus-
pensa Molisch, Die Purpurbakterien, Jena,
1907, 17; Bergey et al.. Manual, 1st ed.,
1923, 402.)
gra'ci.lis. L. adj. gracilis thin, slender.
Cells much smaller than those of Rhabdo-
monas rosea and with less tendency to form
fusiform cells. Usually filamentous, more or
less cylindrical, often with constrictions,
but found up to 60 microns in length.
Shorter filaments motile. Polar flagellate.
Slime formation may occur under special
conditions. Rose-red. Sulfur globules.
Probably an abnormal growth form of Chro-
matium virosum.
Habitat: Mud and stagnant water con-
taining hydrogen sulfide and exposed to
light; sulfur springs.
Illustrations: Warming, op. cit., 1876, PI.
VII, fig. 5; Winogradsky, op. cit., 1888, PI.
IV, fig. 12; Molisch, op. cit., 1907, PI. II, fig.
11-12.
3. Rhabdomonas linsbaueri (Gickl-
horn, 1921) van Niel, 1948. (Rhabdochroma-
tium linsbaueri Gicklhorn, Ber. d. deut. bot.
Ges., 39, 1921, 312; van Niel, in Manual, 6th
ed., 1948, 855.)
lins.bau'er.i. M.L. gen. noun linsbaueri of
Linsbauer; named for K. Linsbauer, an
Austrian botanist.
Cells resemble those of Rhabdomonas rosea,
irregular, rod-shaped, 3 to 5 microns wide,
up to 30 microns in length.
The characteristic feature of the species,
and the chief means of differentiation, is the
occurrence of calcium carbonate inclusions
in addition to the sulfur globules in the cells.
Whether this is strictly an environmentally
conditioned characteristic, due to the
photosynthetic development of the bacteria
50 ORDER I. PSEUDOMONADALES
in a medium rich in calcium ions, so that identity of this species with Rhabdomonas
calcium carbonate is precipitated as the rosea would become evident,
alkalinity increases, has not yet been estab- Source: From a pond near Graz, Austria,
lished but seems possible. In that case the Habitat: Fresh water.
Genus XII. Rhodothece Molisch, 1907.
(Die Purpurbakterien, Jena, 1907, 19.)
Rho.do.the'ce. Gr. noun rhodum the rose; Or. noun thece box (capsule); Rhodothece the
rose capsule.
Sulfur purple bacteria, occurring singly, not aggregated in families. Cells spherical, each
surrounded by a rather wide capsule which is, however, rarely visible without special stain-
ing. Motility not observed. Contain bacteriochlorophyll and carotenoid pigments, coloring
the cells reddish. Capable of photosynthesis in the presence of hydrogen sulfide; the cells
then store sulfur globules, arising as an intermediate oxidation product of the sulfide.
In view of the experiences of Bavendamm and others that a number of representatives of
the sulfur purple bacteria, characterized by typical colonial aggregates when found in
nature, may develop as single cells in pure culture, it is quite conceivable that the genus
Rhodothece is identical with some other genus, e.g., Thiopedia or Lamprocystis, and that
these genera represent different growth forms induced by environmental conditions.
The type species is Rhodothece pendens Molisch.
1. Rhodothece pendens Molisch, 1907. nomena due to the pseudovacuoles and to
(Die Purpurbakterien, Jena, 1907, 19.) the sulfur globules distort the cell shape
pen'dens. L. part. adj. pendens hanging. under ordinary illumination so that bacteria
Cells spherical, frequentlj^ occurring as appear as polygons rather than round cells,
diplococci, occasionally as very short chains Usually 2 aerosomes and 2 sulfur globules
or clumps of 3 to 5 individuals. 1.8 to 2.5 per cell. Color not observable in individual
microns in diameter. Produce rather abun- bacteria. Cell groups are rose-red. Motility
dant slime. Cells embedded in individual not observed.
capsules which are rarely visible without Habitat: Mud and stagnant water con-
staining (India ink). Characteristic is the taining hydrogen sulfide and exposed to
regular occurrence of pseudovacuoles (aero- light. Not reported from sulfur springs,
somes) which are supposed to keep the cells Illustrations: Molisch, Die Purpurbak-
suspended in liquid media. Refractive phe- terien, Jena, 1907, PI. II, fig. 13-14.
Genus XIII. Chromatium Perty, 1852.
(Zur Kenntniss kleinster Lebensformen, Bern, 1852, 174.)
Chro.ma'ti.um. Gr. noun chromatium color, paint.
Cells occur singly, more or less ovoid, bean- or vibrio-shaped or short rods. The last-
mentioned are often thick-cj'lindrical with rounded ends. Motile by means of polar flagella.
Contain bacteriochlorophyll and carotenoid pigments, coloring the cells various shades of
red. Capable of photosynthesis in the presence of hydrogen sulfide and storing elemental
sulfur as an incomplete oxidation product in the form of globules inside the cells.
At present the genus contains twelve described species. Differentiation of these species
has, in the past, been based almost entirely upon size and shape of individual cells, often
with complete disregard for the variability of these criteria. The unsatisfactory and arbi-
trary nature of such a classification has occasionallj^ been pointed out, and with much
justification. Winogradsky (Schwefelbacterien, Leipzig, 1888, 98) mentions the many transi-
tional stagfes that can be observed between Chromatium okenii and Chromatium weissei;
Strzeszewski (Bullet. Acad. Sci., Cracovie, Ser. B, 1913, 321) holds that it is impossible to
distinguish, on the basis of sizes or otherwise, between Chromatium weissei and Chromatium
minus. Such contentions, derived from observations on material from natural collections or
FAMILY I. THIORHODACEAE
51
crude cultures, have been greatly strengthened by studies with pure cultures of species of
Chromatium. Thus van Niel (Arch. f. Mikrobiol., 3, 1931, 59) reported variations in width
from 1 to 4 microns, and in length from 2 to 10 microns or even up to 50 microns; Manten
(Antonie van Leeuwenhoek, 8, 1942, 164 S.) found size differences of 1 to 14 microns with a
pure culture of an organism that he identified as Chromatium okenii. Often the differences
in size of a pure culture can be related to special environmental conditions. On account of
such results a designation of species on the basis of size relations alone is manifest)}^ un-
satisfactory. Moreover, the available data do not suggest that differences in shape, color or
arrangement of sulfur globules can be used more effectively. Lack of adequate experimental
results with a sufficiently large number and variety of pure cultures prevents a more rational
classification at present.
The previously proposed species have been listed below with their respective character-
istics and arranged as far as possible in the order of decreasing width.
Two Chromatium species have been described as containing inclusions of calcium carbon-
ate in addition to sulfur globules. As in the case of Rhabdomonas linsbaueri, it is not known
w'hether this feature may be a direct consequence of the calcium ion content and pH of the
environment and thus fail to have taxonomic significance.
The type species is Chromatium okenii Perty.
1. Chromatium gobii Issatchenko, 1914.
(Recherches sur les microbes de I'oc^an
glacial arctique, Petrograd, 1914, 253.)
go'bi.i. M.L. gen.noun gobii of Gobi;
named for Prof. X. Gobi.
Cells 10 microns by 20 to 25 microns.
Source: From sea water of the Arctic
Ocean.
Habitat: Presumably ubiquitous in the
colder portions of the Ocean at least.
Illustration: Issatchenko, loc. cit., PI. II,
fig. 12.
2. Chromatium warmingii (Cohn, 1875)
Migula, 1900. (Monas warmingii Cohn,
Beitr. Biol. Pfl., i. Heft 3, 1875, 167; Migula,
Syst. d. Bakt., 2, 1900, 1048.)
war.min'gi.i. Named for Eugene Warm-
ing, a Danish botanist; M.L. gen.noun
warmingii of Warming.
Cells 8 by 15 to 20 microns, also smaller
(Cohn).
Illustration: Cohn, op. cit., 1875. PI. VI,
fig. 11.
3. Chromatium linsbaueri Gicklhorn,
1921. (Ber. d. deut. botan. Ges., 39, 1921,
312.)
lins.bau'er.i. Named for K. Linsbauer, an
Austrian botanist; M.L. gen.noun linsbaueri
of Linsbauer.
Cells 6 by up to 15 microns (Gickl-
horn) ; 6 to 8 microns in width (Ellis, Sul-
phur Bacteria, London and New York, 1932,
147) . Special characteristic is the occurrence
of calcium carbonate inclusions. Otherwise
resembles Chromatium okenii.
Source: From a pool in the Stiftingtal,
near Graz, Austria.
Habitat: Fresh water.
Illustrations: Gicklhorn, op. cit., 1921,
314, fig. 1; Ellis, op. cit., 1932, 148, fig. 31.
4. Chromatium okenii (Ehrenberg,
1838) Perty, 1852. {Monas okenii Ehrenberg,
Infusionsthierchen, Leipzig, 1838; Perty,
Zur Kenntniss kleinster Lebensformen,
Bern, 1852, 174.) This is the type species of
genus Chromatium.
o.ke'ni.i. Named for L. Oken, a German
naturalist; M.L. gen.noun okenii of Oken.
Cells 5.6 to 6.3 by 7.5 to 15 microns
(Cohn) ; minimum width 4.5 microns (Issat-
chenko, Borodin Jubilee Vol., 1929?, 8);
with many transitions to Chromatium weis-
sei (Winogradsky, Schwefelbacterien, Leip-
zig, 1888, 92). Also: 3.5 by 8 to 12 microns
and varj'ing in size from 1 to 15 microns
(Manten, Antonie van Leeuwenhoek, 8, 1942,
164).
Illustrations: Cohn, Beitr. Biol. Pfl., 1,
Heft 3, 1875, PI. VI, fig. 12; Winogradsky,
op. cit., 1888, PI. IV, fig. 3-4; Issatchenko,
Recherches sur les microbes de I'ocean
glacial arctique, Petrograd, 1914, PI. II,
fig. 9.
5. Chromatium weissei Perty, 1852.
oz
ORDER I. PSEUDOMONADALES
(Zur Kenntniss kleinster Lebensformen,
Bern, 1852, 174.)
weis'se.i. Named for J. F. Weisse, a zoolo-
gist; M.L. gen. noun weissei of Weisse.
Cells 4.2 by 5.7 to 11.5 microns (Perty);
also 3 to 4 by 7 to 9 microns (Issatchenko,
Borodin Jubilee Volume, 1929?, 8); transi-
tions to Chromatium okenii (Winogradsky,
Schwefelbacterien, Leipzig, 1888, 92);
transitions to Chromatium minus (Strzes-
zewski, Bull. Acad. Sci., Cracovie, Ser. B,
1913, 321).
Illustrations: Winogradsky, op. cit., 1888,
PI. IV, fig. 1-2, Miyoshi, Jour. Coll. Sci.,
Imp. Univ. Tokyo, Japan, 10, 1897, PI.
XIV, fig. 15.
6. Chromatium cuculliferum Gickl-
horn, 1920. (Cent, f . Bakt., II Abt., 50, 1920,
419.)
cu.cul.li'fe.rum. L. noun cucullus hood,
cap; L. V. few to bear; M.L. adj. cucullifer
cap-bearing.
Cells 4 by 6 to 8 microns (Gicklhorn) ;
according to Bavendamm (Schwefelbak-
terien, Jena, 1924, 127), identical with
Chromatium warmingii forma minus. Gickl-
horn claims this organism to be colorless,
which appears very doubtful.
Source: From the pond in the Annen
Castle Park, Graz, Austria.
Habitat: Fresh-water ponds.
Illustration: Gicklhorn, op. cit., 1920, 419,
fig. 2.
7. Chromatium minus Winogradsky,
1888. (Schwefelbacterien, Leipzig, 1888, 99.)
mi'nus. L. comp.adj. minor (neut. minus)
less, smaller.
Cells 3 by 3.5 to 7 microns (Winograd-
sky) ; also 1.7 to 3 microns in width and up to
8.5 microns in length (Issatchenko, Borodin
Jubilee Volume, 1929?, 9); all transitions to
Chromatium weissei from which it cannot be
distinguished (Strzeszewski, Bull. Acad.
Sci., Cracovie, Ser. B, 1913, 321).
Illustrations: Winogradsky, op. cit., 1888,
PI. IV, fig. 5; Miyoshi, Jour. Coll. Sci., Imp.
Univ., Tokyo, Japan, 10, 1897, PI. XIV, fig.
16; Issatchenko, Recherches sur les microbes
de I'ocean glacial arctique, Petrograd, 1914,
PI. II, fig. 10-11.
8. Chromatium vinosum (Ehrenberg,
1838). Winogradsky, 1888. {Monas vinosa
Ehrenberg, Die Infusionstierchen, Leipzig,
1838, 11; Winogradsky, Schwefelbacterien,
Leipzig, 1888, 99.)
vi.no'sum. L. adj. vinosus full of wine.
Cells 2 by 2.5 to 5 microns; also 1.4 to 3 by
1.5 to 5 microns (Jimbo, Botan. Magaz.
Tokyo, 51, 1937, 872); 1.7 to 2 by 2 to 9
microns (Issatchenko, Borodin Jubilee
Volume, 1929?, 9) ; or 1 to 1.3 microns by 2.5
to 3 microns (Schrammeck, Beitr. Biol. d.
Pflanzen, 22, 1935, 317). Jimbo considers
Thioderma roseum Miyoshi to be identical
with Chromatium vinosum.
Illustrations: Winogradsky, op. cit., 1888,
PI. IV, 6-7; Miyoshi, Jour. Coll. Sci., Imp.
Univ. Tokyo, Japan, 10, 1897, PI. XIV, fig.
17; Nadson, Bull. Jard. Imp. Botan., St.
Petersbourg, 12, 1912, PI. Ill, fig. 1-2.
9. Chromatium violaceum Perty, 1852.
(Zur Kenntniss kleinster Lebensformen,
Bern, 1852, 174.)
vi.o.la'ce.um. L. adj. violaceus violet-
colored.
Cells about 2 by 2 to 3 microns. Accord-
ing to Cohn (Beitr. Biol. Pfl., 1, Heft 3,
1875, 166), probably identical with Chro-
matium vinosum. Apparently includes var-
ious sizes.
10. Chromatium molischii (Bersa,
1926) van Niel, 1948. {Pseudomonas molischii
Bersa, Planta, 2, 1926, 375; van Niel, in
Manual, 6th ed., 1948, 858.)
mo.li'schi.i. Named for H. Molisch, an
Austrian botanist; M.L. gen. noun molischii
of Molisch.
Cells about 2 by 2.5 to 8 microns. Sup-
posedly contain calcium carbonate as in-
clusions.
Illustration: Bersa, op. cit., 1926, 376, fig.
3.
11. Chromatium gracile Strzeszewski,
1913. (Bull. Acad. Sci., Cracovie, Ser. B,
1913, 321.)
gra'ci.le. L. adj. gracilis thin, slender.
Cells 1 to 1.3 by 2 to 6 microns; also to 1.5
microns in width (Issatchenko, Etudes mi-
crobiologiques des Lacs de Boue, Leningrad,
1927, 114).
Illustration: Strzeszewski, op. cit., 1913,
FAMILY I. THIORHODACEAE 53
PI. XXXIX, fig. 1-2; Tokuda, Botau. from 0.5 to 0.7 by 0.6 to 1 micron (Issat-
Magaz., Tokyo, 50, 1936, 339, fig. 1-23. chenko, Recherches sur les microbes de
I'ocean glacial arctique, Petrograd, 1914,
12. Chroniatium miniitissiniuin Wino- 253), and 1 to 3 by 2 to 5 microns (Issat-
gradskjr, 1888. (Schwefelbacterien, Leipzig, chenko, Borodin Jubilee Volume, 1929?, 9).
1888, 100.) Illustrations : Winogradsky, op. cit., 1888,
mi.nu.tis'si.mum. L. sup. adj. minutissi- PI. IV, fig. 8; Miyoshi, Jour. Coll. Sci., Imp.
mtfs very small. Univ., Tokyo, Japan, 10, 1897, PI. XIV,
Cells about 1 to 1.2 by 2 microns. Also fig. 18.
FAMILY II. ATHIORHODACEAE MOLISCH, 1907.
(Die Purpurbakterien, Jena, 1907, 28.)
A.thi.o.rho.da'ce.ae. Or. pref. a without; Gr. noun ihium sulfur; Gr. noun rhodum the
rose; -aceae ending to denote a family; M.L. fem.pl.n. Athiorhodaceae (probably intended
to mean) the family of the non-sulfur red bacteria.
Unicellular bacteria, of relatively small size, occurring as spheres, short rods, vibrios,
long rods and spirals. Motility is due to the presence of polar flagella. Gram-negative. They
produce a pigment sj^stem composed of bacteriochlorophyll and one or more carotenoids,
coloring the cells yellowish brown, olive-brown, dark brown or various shades of red. Color
usually not observable with single cells but only with cell masses. Generally microaero-
philic, although many representatives may grow at full atmospheric oxygen tension. Capa-
ble of development under strictly anaerobic conditions, but only in illuminated cultures by
virtue of a photosynthetic metabolism. The latter is dependent upon the presence of ex-
traneous hydrogen donors, such as alcohols, fatty acids, hydroxy- and keto-acids, and does
not proceed with the evolution of molecular oxygen. Those members which can grow in the
presence of air can also be cultivated in darkness, but only under aerobic conditions.
The growth requirements of some of the species in this family have been reported by
Hutner (Arch. Biochem., 3, 1944, 439; Jour. Bact., 52, 1946, 217; Jour. Gen. Microbiol., 4,
1950, 286) ; his findings are incorporated in the descriptions which follow.
Key to the genera of family Athiorhodaceae.
I. Cells rod-shaped or spherical, not spiral-shaped.
Genus I. Rhodopseudomonas , p. 53.
II. Cells spiral-shaped.
Genus II. Rhodospirillum, p. 58.
Genus I. Rhodopseudomonas Kluyver and van Niel, 19S7, emend, van Niel, 1944-
(Includes Rhodobacillus Molisch, Die Purpurbakterien, Jena, 1907, 14; Rhodobacterium
Molisch, ibid., 16; Rhodococcus Molisch, ibid., 20; Rhodovibrio Molisch, ibid., 21; Rhodocystis
Molisch, ibid., 22; Rhodonostoc Molisch, ibid., 23; Rhodosphaera Buchanan, Jour. Bact., S,
1918, 472; Rhodorhagus (sic) Bergey et al.. Manual, 2nd ed., 1925, 414; Rhodomonas Kluyver
and van Niel, Zent. f. Bakt., II Abt., 94, 1936, 397; not Rhodomonas Orla-Jensen, Cent. f.
Bakt., II Abt., 22, 1909, 331 ; Kluyver and van Niel, in Czurda and Maresch, Arch, f . Mikro-
biol., 8, 1937, 119; Rhodorrhagus Bergey et al.. Manual, 5th ed., 1939, 905; van Niel, Bact.
Rev., 8, 1944, 86.)
Rho.do.pseu.do.mo'nas. Gr. noun rhodum the rose; Gr. adj. pseudes false; Gr. noun
monas monad, unit; M.L. fem.n. Pseudotnonas a bacterial genus; M.L. fem.n. Rhodopseudo-
monas the rose Pseudomonas.
Spherical and rod-shaped bacteria, motile by means of polar flagella. Gram -negative.
54 ORDER I. PSEUDOMONADALES
Contain bacteriochlorophjdl which enables them to carry out a photosynthetic metabolism.
The latter is dependent upon the presence of extraneous oxidizable substances and proceeds
without the evolution of molecular oxygen. Though some members can oxidize inorganic
substrates, none appears to be strictly autotrophic due to the need for special organic growth
factors. Produce accessory pigments causing the cultures, especially when kept in light, to
appear in various shades of brownish yellow to deep red.
The type species is Rhodopsendornonas palnstris (Molisch) van Niel.
Keys to the species of genus Rhodopseudomonas.
I. Based upon morphological characters.
A. Cells clearly rod-shaped in all media.
1. Cells short, somewhat curved, to long branched rods, size of j'oung and short
cells 0.6 to 0.8 by 1.2 to 2 microns; in older cultures up to 10 microns long; do
not form slime; liquid cultures, when young, or after shaking, evenly turbid.
Color red to dark brown-red.
1. Rhodopseudomonas palustris.
2. Cells slender rods, 0.5 by 1.2 microns, usually clumped together in extensive
slime masses. Cultures pale brown to peach-colored.
2. Rhodopseudomonas gelatinosa.
B. Cells more or less spherical in media at pH below 7.
1. In media at pH about 7 clearly rod-shaped, 1 by 1 to 2.5 microns. Chains of cells
frequent and in characteristic zigzag arrangement.
3. Rhodopseudomonas capsulaia.
2. In media at pH above 7 cells still predominantly spherical, 0.7 to 4 microns in
diameter. Mostly single, little tendency to chain formation.
4. Rhodopsendomonas spheroides.
II. Based chiefly on physiological properties.
A. Gelatin liquefied.
2. Rhodopseudomonas gelatinosa.
B. Gelatin not liquefied.
1. Does not produce mucus in media at pH above 8. Color the same under aerobic
and anaerobic conditions of growth.
1. Rhodopseudomonas palustris.
2. Produce mucus in media at pH above 8. Color brow^n in anaerobic, red in aerobic
culture.
a. Develops readily in media with 0.2 per cent propionate as the chief oxidation
substrate. Mucus production marked at pH above 8, but very limited be-
tween 7 and 8.
3. Rhodopseudomonas capsulata.
aa. Does not develop in media with 0.2 per cent propionate as the main oxida-
tion substrate. Slime formation extensive at pH above 7.
4. Rhodopseudomonas spheroides.
III. Based principally upon biochemical characters.
A. Thiosulfate used as main oxidation substrate.
1. Rhodopseudomonas palustris.
B. Thiosulfate not used.
1. Propionate (0.2 per cent) used.
3. Rhodopseudomonas capsulata.
2. Propionate not used.
a. Mannitol and sorbitol (0.2 per cent) used.
4. Rhodopseudomonas spheroides.
aa. Mannitol and sorbitol not used.
2. Rhodopseudomonas gelatinosa.
FAMILY II. ATHIORHODACEAE
55
1. Rhodopseudonionas palustris (Mo-
lisch, 1907) van Niel, 194-4. [Rhodobacillus
palustris ]Molisch, Rhodobacterium capsula-
tum Molisch and Rhodovibrio parvus Mo-
lisch, Die Purpurbakterien, Jena, 1907, 14,
18 and 21 ; van Niel, Bact. Rev., 8, 1944, 89.)
pa.lu.s'tris. L. adj. paluster marshy,
swampy.
Cells usually distinct!}^ rod-shaped,
though in young cultures very short, lightly
curved rods may often predominate. Size
variable, even for the same strain, and
strongly influenced by age of culture and
composition of medium. Rather consistently
short cells in young cultures in yeast ex-
tract, especially when incubated anaerobi-
cally in the light or in anaerobic cultures
with substrates, such as malonate, which
permit only a slow and scant develop-
ment. Dimensions in such cultures 0.6 to 0.8
by 1.2 to 2 microns. More often, especially
in older cultures, cells are much longer, up
to 10 microns. Highly characteristic is the
pronounced tendency to the formation of
irregularly shaped, bent and crooked long
rods, occasionally swollen at one or both
e.xtremities, and frequently suggesting
branching. Such cells usually form clusters
reminiscent of Corynebacterium and Myco-
bacterium cultures.
Cells in young cultures actively motile by
means of polar flagella; irregular and long
cells as a rule non -motile. Gram-negative.
Growth in liquid media never mucoid;
sediment in older cultures homogeneous and
smooth, readily redispersible.
Color varies considerably, depending upon
the medium, and especially in anaero-
bic illuminated cultures. Where develop-
ment is slight (as in malonate, thiosulfate
and, usually, glycerol media), the color is a
light pink; in media containing fatty acids,
more nearly dark reddish brown. Color due
to bacteriochlorophyll and a number of dif-
ferent carotenoid pigments; most strains
produce, in addition, a water-soluble, non-
carotenoid, bluish red pigment which dif-
fuses into the culture medium.
In yeast extract cultures growth is pos-
sible over the range pH 6 to 8.5. With cer-
tain substrates, especially fatty acids, the
combined effect of low pH and a substrate
concentration of 0.1 to 0.2 per cent may pre-
vent growth. No characteristic odors save
that old cultures may develop a distinct
ionone-like fragrance. Gelatin is not lique-
fied; leucine is generally utilized as a sub-
strate.
Most strains are able to grow on the sur-
face of agar plates or slants ; a few, especially
when first isolated, appear more sensitive to
oxygen and develop only in stabs in which
the upper region may remain free of growth.
Generally such strains can be adapted to
grow at full atmospheric oxygen tension.
Most fatty acids and hydroxy acids are
adequate oxidation substrates. All cultures
can grow at the expense of thiosulfate and
produce rapid and profuse growth in glu-
tarate and ethanol media. No development
in media containing, as the chief oxidation
substrate, 0.2 per cent sorbitol, glucose or
mannose, even though these substances are
not inhibitory. Molecular hydrogen can be
oxidized.
All cultures can develop anaerobically in
illuminated cultures by photosj^nthesis.
p-amino-benzoic acid is required for
growth (Hutner).
Optimum temperature generally rather
high, good development being possible up
to 37° C. However, certain strains exhibit a
lower optimum temperature.
Distinctive characters : Morphological
resemblance to species of Mycobacterium in
old cultures, ability to grow with thiosulfate
as the chief oxidizable substrate, and failure
to develop in media which contain carbohy-
drates or sugar alcohols in a concentration
of 0.2 per cent as the main oxidizable com-
pounds.
Habitat: Regularly found in mud and
stagnant bodies of water.
Illustrations: Molisch, op. cit., 1907,
Plate I, fig. 1, 2; Plate II, fig. 10; van Niel,
op. cit., 1944, fig. 1-3, p. 18, and fig. 18-26,
p. 90.
2. Rhodopseudomonas gelatinosa (Mo-
lisch, 1907) van Niel, 1944. (Rhodocystis
gelatinosa Molisch, Die Purpurbakterien,
Jena, 1907, 22; van Niel, Bact. Rev., 8, 1944,
98.)
ge.la.ti.no'sa. L. part. adj. gclatus frozen,
stiffened; M.L. gelatinum gelatin, that which
56
ORDER I. PSEUDOMONADALES
stiffens; M.L. adj. gelatinosus full of gelatin,
gelatinous.
Cells in young cultures short and small
rods, approximately 0.5 by 1 to 2 microns.
In old cultures much longer, up to 15 mi-
crons, and then irregularly curved rods,
often swollen and gnarled in places up to 1
micron in width. In this stage the cells bear
some resemblance to those found in old cul-
tures of Rhodopseudomonas palustris, but
the characteristic Mycobacterium-Yike clus-
ters of the latter are absent. Single cells in-
frequent due to a copious mucus produc-
tion in all media which causes the cells to
clump together. While young cells are ac-
tively motile by means of polar flagella,
motility is often difficult to ascertain as a
result of the pronounced tendency to con-
glomerate; the individuals in the clumps
appear to be non-motile. Gram-negative.
Gelatin is liquefied; of the single amino acids
alanine, asparagine, aspartic and glutamic
acids appear generally satisfactory sub-
strates.
Color quite distinctive in most anaerobic
cultures as a pale, delicate, pinkish shade,
rather peach-colored. Only in the presence
of rather high concentrations of yeast ex-
tract (when a much heavier growth is ob-
tained than with low concentrations supple-
mented with 0.2 per cent of various single
oxidation substrates) do the slimy cell
masses appear a dirty, faded brown. Color
is due to bacteriochlorophyll and carotenoid
pigments. Occasionally a water-soluble,
non-carotenoid, bluish red pigment is
produced which diffuses into the culture
medium.
In j^east extract, growth occurs over a pH
range extending from at least 6.0 to 8.5.
Cultures produce a characteristic acrid
odor.
More sensitive to fatty acids than are
other species of Rhodopseudomonas; with
0.2 per cent propionate no growth occurs.
The best single oxidizable substrates appear
to be ethanol, glucose, fructose and man-
nose, as well as a variety of amino acids.
Citrate also permits good growth; not, on
the other hand, glycerol, mannitol, sorbitol
or tartrate in the usual concentration of
0.2 per cent.
Thiosulfate is not oxidized; behavior
towards molecular hydrogen unknown.
More pronouncedly microaerophilic than
the other Rhodopseudomonas species; most
cultures cannot develop on aerobically in-
cubated slants or agar plates.
Capable of strictly anaerobic development
in illuminated cultures by virtue of a photo-
synthetic metabolism.
Thiamin plus biotin is required for growth
(Hutner).
Temperature relations so far unknown.
Distinctive characters: The small size of
the individual cells and the pronounced
clumping which causes the cultures to be
exceptionally stringy; the unusual color of
the cell masses; the ability to liquefy gela-
tin, to utilize citrate and a number of amino
acids. Correlated with these is the failure to
grow in media with 0.2 per cent propionate,
tartrate and glycerol.
Habitat: Regularly' present in stagnant
bodies of water and in mud.
Illustrations : Molisch, op. cit., 1907, Plate
I, fig. 8; van Niel, op. cit., 1944, fig. 55-60, p.
99; fig. 61-66, p. 100.
3. Rhodopseudomonas capsulata (Mo-
lisch, 1907) van Niel, 1944. {Rhodonostoc
capsulatum Molisch, Die Purpurbakterien,
Jena, 1907, 23; Rhodopseudomonas capsulatus
(sic) van Niel, Pact. Rev., S, 1944, 92.)
cap.su.la'ta. L. dim. noun capsula a small
chest, capsule; M.L. adj. capsulatus capsu-
lated.
Depending upon the pH of the medium,
cells nearly spherical, or as distinct rods,
often devoid of motility. Motility due
to polar flagella. The spherical cells are
found in media with a pH below 7; thej' are
usually arranged in chains resembling strep-
tococci. Rod-shaped cells are characteristic
for media with pH above 7; the higher the
pH, the longer the rods. Individual cells
slightly less than 1 micron wide, although
attenuated rods (about 0.5 micron in width)
are frequent at pH above 8, and slightly
swollen cells (to 1.2 microns) are found in
media containing sugars. Length varies from
1 to 6 microns; most common dimensions in
approximately neutral media, 2 to 2.5 mi-
crons. At pH above 8, abnormal growth in
FAMILY II. ATHIORHODACEAE
57
the form of irregular filaments. Outstand-
ingly characteristic is the zigzag arrange-
ment of the cells in chains. Gram-negative.
Cultures in media of pH 8 or above are
distinctly mucoid.
Color: Anaerobic cultures develop with a
brown color, the shade ranging from a light
3'ellowish l^rown to a deep mahogany-
brown. When grown in the presence of oxy-
gen, the cultures are dark red. Even the pig-
mentation of the brown-colored organisms
from an anaerobic culture can be changed
into a distinct red by shaking a suspension
with air for some hours; light enhances the
rate of this color change. Color due to bac-
teriochlorophyll and carotenoid pigments.
No diffusible water-soluble pigment is pro-
duced.
Growth possible over a pH range from at
least 6 to 8.5, morphology becoming abnor-
mal in the alkaline media.
Most cultures are odorless, although occa-
sionally a faint peach-like odor can be de-
tected.
Growth is not inhibited by the presence of
oxygen, although the pigmentation is
thereby affected.
Fatty acids and most substituted acids
are satisfactory substrates. Rapid and
abundant growth with propionate at a
concentration of 0.2 per cent. At this same
concentration glutaric acid leads, at best, to
very meager cultures, while tartrate, citrate
and gluconate fail to induce growth, as do
also ethanol, glycerol, mannitol and sorbi-
tol. In media with 0.2 per cent glucose or
fructose good growth is obtained. No growth
with mannose. Thiosulfate is not, but
molecular hydrogen can be, oxidized by this
species.
Gelatin is not liquefied; of the amino
acids, alanine and glutamic acid are satis-
factory substrates while leucine is not uti-
lized.
Distinctive characters: Cell shape and
arrangement in chains; brown color of
anaerobic, red pigmentation of aerobic cul-
tures; ability to grow in media with 0.2 per
cent propionate, glucose, fructose, alanine
and glutamic acid; failure to develop with
leucine, as well as with ethanol, glycerol,
mannitol and sorbitol in the above-men-
tioned concentration.
All cultures can develop anaerobically in
illuminated cultures by a photosynthetic
metabolism.
Thiamin is required for growth; u few
strains require biotin and nicotinic acid in
addition (Hutrier).
Optimum temperature distinctly lower
than for Rhodopseudomonas palustris, and,
as a rule, around 25° C.
Habitat: Regularly found in stagnant
bodies of water and in mud.
Illustrations: Molisch, op. cit., 1907,
Plate II, fig. 9; van Niel, op. cit., 1944, fig.
4-6, p. 19; fig. 27-32, p. 92; and fig. 33-38, p.
93.
4. RhodopseudonionaH spheroides van
Niel, 1944. {Rhodococcus capsulatus Molisch,
Die Purpurbakterien, Jena, 1907, 20; Rhodo-
coccus minor Molisch, ibid., 21; van Niel,
Bact. Rev., 8, 1944, 95.)
sphe.ro. i'des or sphe.roi'des. Gr. adj.
sphaeroides globular.
Cells generally single, nearly spherical,
diameter without slime capsule variable, de-
pending upon medium, ranging from 0.7 to
4 microns. In young cultures actively motile
by means of polar flagella; motility soon
ceases in media which are or become alka-
line. Copious slime production in media at
pH above 7. In strongly alkaline cultures
abnormal cell-shapes occur in the form of
irregular, swollen and distorted rods, often
having the appearance of spore-bearing
cells, simulated by the production of fat
bodies. In sugar-containing media egg-
shaped cells, measuring as a rule 2.0 to 2.5
by 2.5 to 3.5 microns, are frequently found.
Gram-negative.
Color: Anaerobic cultures develop with
brown color, ranging in shade from a light,
dirty greenish brown to a dark brown. Cul-
tures grown in the presence of oxygen are
distinctly red. As in the case of Rhodopseudo-
monas capsulata, the brown color of an
anaerobic culture can be changed to red by
shaking with air, light stimulating the color
change. Color due to bacteriochlorophyll
and carotenoid pigments. The large major-
ity of cultures of this species produce, in
addition, a water-soluble, non-carotenoid,
bluish red pigment which diffuses into the
culture medium.
58
ORDER I. PSEUDOMONADALES
Gelatin is not liquefied, and growth with
single amino acids appears somewhat er-
ratic. No definite correlations have been
observed.
Development is possible over a wide pH
range, extending from at least 6.0 to 8.5.
All cultures exhibit an unpleasant putrid
odor.
Requires for optimal development higher
concentrations of yeast extract as a supply
of growth factors than either Rhodopseudo-
7nonas palustris or Rhodopseudomonas capsu-
lata and is more sensitive to low fatty acid
concentrations. With 0.2 per cent propionate
in a neutral medium, no growth occurs;
caproic and pelargonic acids are toxic in
concentrations below 0.1 per cent. On the
other hand, tartrate and gluconate can serve
as oxidation substrates, as can also ethanol,
glycerol, mannitol, sorbitol, glucose, fruc-
tose and mannose in 0.2 per cent concentra-
tions.
In sugar-containing media, acid is pro-
duced; the pH ma}^ drop to below 4.0 before
development ceases. Acid production from
glucose occurs both in presence and absence
of air, and in illuminated as well as in non-
illuminated cultures. In cultures exposed to
light, the acid usually disappears later on.
Thiosulfate is not oxidized; hydrogen
oxidation has not been observed.
Oxygen does not prevent growth; colonies
develop on the surface of agar plates ex-
posed to air, with a red pigmentation.
Capable of strictly anaerobic development
in illuminated cultures by photosynthesis.
Thiamin, biotin and nicotinic acid are
required for growth (Hutner).
Optimum temperature, below 30° C.
Distinctive characters: Spherical cell-
shape in most media; brown color of anaero-
bic and red pigmentation of aerobic cul-
tures; growth with 0.2 per cent tartrate,
gluconate, ethanol, glycerol, mannitol,
sorbitol, glucose, fructose and mannose;
failure to grow with 0.2 per cent propio-
nate.
Habitat: Regularly found in stagnant
bodies of water and in mud.
Illustrations : INIolisch, op. cit., 1907, Plate
II, fig. 15; van Niel, op. cit., 1944, fig. 7-8, p.
19; fig. 39-45, p. 96; fig. 46-54, p. 97.
Genvs II. Rhodospirillum Molisch, 1907, emend, van Niel, 1944-
(Molisch, Die Purpurbakterien, Jena, 1907, 24; van Niel, Bact. Rev., 8, 1944, 86; the genus
now includes the genus Phaeospirilhtm Kluyver and van Niel, Zent. f. Bakt., II Abt., 94,
1936, 396.)
Rho.do.spi.ril'lum. Gr. noun rhodum the rose; Gr. noun spira a coil, a spiral; M.L.
dim.neut.n. Spirillum a bacterial genus; M.L. neut.n. Rhodospirillum the rose Spirillum.
Spiral-shaped bacteria, motile by means of polar flagella. Gram-negative. Contain bac-
teriochlorophyll and are potentially photosynthetic in the presence of extraneous oxi-
dizable substances. Molecular oxygen is not produced. Unable to grow in strictly mineral
media, even when possessed of the ability to utilize hydrogen as oxidizable substrate, due
to the need for organic nutrilites. Produce accessory pigments causing the cultures, espe-
cially when grown in the light, to appear in various shades of red to brown.
The type species is Rhodospirillum rubrum (Esmarch) Molisch.
Key to the species of genus Rhodospirillum.
I. Cultures deep red without brownish tinge; characteristic absorption band around 550
millimicrons.
1. Rhodospirillum ruhrum.
II. Cultures reddish brown to orange; characteristic absorption maximum around 520, not
550, millimicrons.
A. Cells 0.5 or less micron in width.
2. Rhodospirillum Julvum.
B. Cells more than 0.5 micron wide.
1. Size of cells 0.7 to 0.9 by 5 to 10 microns.
3. Rhodospirillum molischianum.
2. Size 1.2 to 1.5 by 14 to 30 microns.
4. Rhodospirillum photometricum.
FAMILY II. ATHIORHODACEAE
59
1. Rhotlospirillum rubruni (von Es-
march, 1887) Molisch, 1907. {Spirillum rub-
rum von Esmarch, Cent. f. Bakt., 1, 1887,
225; Rhodospirillum giganteum Molisch, Die
Purpurbakterien, Jena, 1907, 24; Molisch,
ibid., 25.)
rub'rum. L. adj. ruber red.
Cells characteristically spiral-shaped, but
size of elements variable within wide
limits, depending upon environmental con-
ditions during growth. Width of cells from
0.5 to 1.5 microns; length from 2 to 50 mi-
crons, and over ; even in a single culture such
differences may be found. Also the shape and
size of the spiral coil varies much; it usually
ranges between 1 to 4 microns in width, and
from 1.5 to 7 microns in length. In alanine
media the majority of the cells occur in the
form of half -circles to complete rings;
malate media tend to produce much flat-
tened spirals. In old cultures involution
forms appear, straightened spirals and ir-
regularly swollen cells, the latter common in
media with higher fatty acids. Such cells
stain irregularly, contain fatty inclusions,
and are occasionally branched. Young cul-
tures show active motility, due to polar
fiagella. Gram-negative.
Mucus is not produced. In calcium-defi-
cient media the growth is flocculent, as if
agglutinated. With an adequate calcium
supply the growth in liquid media is ho-
mogeneous, suspended and consists of single
cells.
Gelatin is not liquefied; the amino acids
alanine, asparagine, aspartic and glutamic
acids are satisfactory oxidizable compounds.
Color: Ordinarily deep and dark red,
without any brownish tinge. In ethanol
media lighter, and a characteristic pink.
Pigment production markedly influenced by
oxygen and light. Slants incubated in dark-
ness present a pale grayish surface growth
with a faint reddish hue, while often show-
ing deep-red cell masses in the region
between glass wall and agar surface where
development proceeds at low oxygen ten-
sion. The color is due to bacteriochlorophyll
and carotenoid pigments. Among the latter,
spirilloxanthin is quantitatively predomi-
nant and is responsible for the characteristic
absorption band at 550 millimicrons. Water-
soluble, diffusible pigments are not pro-
duced.
Development possible over a pH range of
at least 6 to 8.5, although, as in other cases,
the combination of an acid reaction and the
presence of fatty acids may prevent growth.
Cultures produce a distinctive odor,
reminiscent of slightly putrid yeast.
In general, grow well with fatty acids as
the chief oxidizable substrate; however, are
prevented from growing by 0.2 per cent
propionate in a neutral medium. Most sub-
stituted acids are equally satisfactory, with
the exception of tartrate, gluconate and
citrate. In a concentration of 0.2 per cent,
ethanol is a suitable substrate, whereas
the carbohydrates and their corresponding
polyalcohols are not utilized.
Thiosulfate is not oxidized; molecular hy-
drogen can be used by some strains.
Rather microaerophilic; many strains,
upon initial isolation, incapable of growth
at atmospheric oxygen tension. Subse-
quent adaptation can be induced, but even
such adapted cultures exhibit negative
chemota.xis to air.
Capable of strictly anaerobic develop-
ment in illuminated cultures on the basis
of a photosynthetic metabolism.
Biotin is required for growth (Hutner).
Optimum temperature generally between
30° and 37° C.
Distinctive characters: The most im-
portant characteristics of the species are
the spiral shape, combined with the ability
to produce a red pigment with a definite
absorption maximum at 550 millimicrons
in the intact cells. Diagnostically useful are
the good growth in media with 0.2 per cent
ethanol, alanine, asparagine, aspartate or
glutamate and the inadequacy of similar
concentrations of carbohydrates and thio-
sulfate as substrates.
Habitat: Regularly present in stagnant
bodies of water and in mud.
Illustrations: Molisch, ibid., Plate I, fig.
5-7; van Niel, Bact. Rev., 8, 1944, fig. 9-10,
p. 19; fig. 11-16, p. 24; fig. 67-75, p. 103; fig.
76-84, p. 104; fig. 85-90, p. 106; fig. 91-96, p.
107.
2. Rhodospirillum fulvum van Niel,
1944. (Bact. Rev., 8, 1944, 108.)
ful'vum. L. adj. fulvus deep or reddish
j^ellow, tawn3^
Characteristic for the species is the very
60
ORDER I. PSEUDOMONADALES
small size of the individual cells. These are
not over 0.5 micron wide and generally not
longer than 2.5 microns. The most common
shape consists of a complete turn of about
1 by 1.5 microns. In media with fatty acids
as a substrate the spirals appear somewhat
steeper than in fumarate, succinate or
malate cultures. Swollen individuals re-
sembling vibrios are encountered in cultures
which do not appear quite healthy. Forma-
tion of mucus or clumping has not been ob-
served.
Gelatin is not liquefied; aspartate has been
the only amino acid capable of inducing
growth. Thiosulfate is not oxidized.
Color: Quite distinct from that of Rhodo-
spirillum rubrimi; colonies and stab cul-
tures are a reddish brown while liquid cul-
tures often appear brownish orange. The
color is due to bacteriochlorophyll and
carotenoid pigments; among the latter
spirilloxanthin, as evidenced by the absence
of an absorption maximum at 550 milli-
microns, is not represented as a major con-
stituent. Does not produce water-soluble,
diffusible pigments.
Capable of strictly anaerobic develop-
ment in illuminated cultures, due to photo-
synthetic metabolism.
Fatty acids and the four-carbon dicar-
boxylic acids are imiformly good substrates;
glutarate is not used. Ethanol and glucose,
in a concentration of 0.2 per cent, have
yielded satisfactory cultures; other carbo-
hydrates, as well as the corresponding poly-
alcohols, have given negative results.
Little information available concerning
pH and temperature relations. Behaves
generally as a strict anaerobe; adaptation
to microaerophilic conditions has not been
achieved. Negative aerotaxis very pro-
nounced.
Distinctive characters : The small size and
the color of the cultures serve as adequate
criteria for its differentiation from Rhodo-
spirilhim ruhrum. The strictly anaerobic
nature and the failure to grow with glu-
tarate and various amino acids except as-
partate can probably be used as supple-
mentary specific properties.
Habitat: Bodies of stagnant water and
mud .
Illustrations: Van Niel, ibid., fig. 97-102,
p. 109; Giesberger, Jour. Microb. and Serol.,
13, 1947, fig. 1-2, p. 141.
3. Rhodospirilluin niolischianuni
Giesberger, 1947. (Jour. Microbiol, and
Serol., 13, 1947, 137.)
mo.li.schi.an'um. M.L. adj. molischianus
pertaining to Molisch.
Cells characteristically spiral -shaped,
moderately large, 0.7 to 0.9 by 5 to 10 mi-
crons. Mostly with one or two complete
turns which vary in width from 1.3 to 2
microns and in length from 4 to 6 microns;
this depends upon environmental condi-
tions.
Color: Distinctly reddish brown due to
the presence of bacteriochlorophyll and
carotenoids, the latter responsible for the
absorption maxima at 520 and 485 milli-
microns. Spirilloxanthin absent.
Capable of development under stricth^
anaerobic conditions in media containing
ethanol or fatty- or hydroxy-acids as oxidiz-
able substrates. Citrate can also be utilized
in this manner, but not glycerol, glucose,
hydrogen sulfide or thiosulfates. Tends to be
strictl}^ anaerobic, and hence capable of
development onlj- in illuminated cultures.
Does not liquefy gelatin.
Distinctive characters : The absence of an
absorption band at 550 millimicrons and the
ability to utilize citrate serve to distinguish
this type from Rhodospirillum ruhrum,
which it closely resembles in size and shape.
The individual cell size differentiates this
species from R. fulvum and R. photometri-
cum.
Habitat: Widely distributed. Regularly
present in stagnant water and mud; can be
found in abundance in anaerobic cultures of
hay extract inoculated with such materials
and incubated in light.
Illustrations: Giesberger, ibid., fig. 3-5,
p. 141.
4. Rhodospirillum photonietricum
Molisch, 1907. (Die Purpurbakterien, Jena,
1907, 24.)
pho.to.me'tri.cum. Gr. noun phos light;
Gr. adj. metricus measured; M.L. adj.
photometriciis light-measured.
Cells large, stout, spiral -shaped. Mostly
with one or two complete turns whose wave
FAMILY III. CHLOROBACTERIACEAE 61
length varies in width from 4 to 6 microns glucose, hydrogen sulfide or thiosulfates.
and in length from 7 to 10 microns; this is Strongly microaerophilic; tends to be
dependent upon environmental conditions. strictly anaerobic, and hence capable of
Actively motile by means of a single polar development only in illuminated cultures,
flagellum. Does not liquefy gelatin.
Color: Distinctly reddish brown due to Distinctive characters: Readily distin-
the presence of bacteriochlorophyll and guishable from Rhodospirillum rubrum by
carotenoids, the latter responsible for the the absence of an absorption band at 550
absorption maxima at 520 and 485 milli- millimicrons and from R. fulvum and R.
microns. Spirilloxanthin not formed. molischianum by the greater size of its cells.
Capable of development under strictly Habitat: Stagnant water and mud; widely
anaerobic conditions in media containing distributed.
ethanol, fatty acids or hydroxy acids as Illustrations: Molisch, ibid., Plate 1, fig.
oxidizable substrates. Citrate can also be 5-6; Giesberger, Jour. Microbiol, and Serol.,
utilized in this manner, but not glycerol, 13, 1947, fig. 6-9, p. 141.
FAMILY III. CHLOROBACTERIACEAE LAUTERBORN, 1913.
{Chlorobakteriaceae (sic) Lauterborn, Alg. Bot. Ztschr., 19, 1913, 99.)
Chlo.ro. bac.te.ri.a'ce.ae. M.L. neut.n. Chlorobacterium type genus of the family; -aceae
ending to denote a family; M.L. fern. pi. n. Chlorobacteriaceae the Chlorobacterium family.
Green bacteria, usually of small size, occurring singly or in cell masses of various shapes
and sizes, developing in environments containing rather high concentrations of hydrogen
sulfide and exposed to light. As a rule not containing sulfur globules but frequently de-
positing elemental sulfur outside the cells. Contain green pigments of a chlorophyllous
nature, though not identical with the common green plant chlorophylls nor with bacterio-
chlorophjdl. Capable of photosynthesis in the presence of hydrogen sulfide; do not liberate
oxygen.
A number of genera have been proposed; some are characterized by special colonial growth
forms while others are characterized on the basis of a supposed symbiotic habitus where
the green bacteria grow in more or less characteristic aggregates together with other micro-
organisms. In view of the variations in size and shape exhibited by the only member of this
group which has so far been obtained and studied in pure culture (van Niel, Arch, f . Mikro-
hio\.,3, 1931, 65ff.), the validity of many of these genera is doubtful. The following keys and
descriptions, therefore, bear a strictly provisional character. Here, as in the case of the
sulfur purple bacteria, significant advances can only be expected from pure-culture studies
under controlled environmental conditions.
Key to the genera of family Chlorobacteriaceae.
I. Free-living bacteria not intimately associated with other microbes.
A. Bacteria not united into well defined colonies.
Genus I. Chlorobium, p. 62.
B. Bacteria united into characteristic aggregates.
1. Bacteria without intracellular sulfur globules.
Genus II. Pelodictyon, p. 63.
2. Bacteria with intracellular sulfur globules.
Genus III. Clathrochloris, p. 64.
II. Green bacteria found as symbiotic aggregates with other organisms.
A. Aggregates composed of green bacteria and protozoa.
Genus IV. Chlorobacterium, p. 65.
62 ORDER I. PSEUDOMONADALES
B. Aggregates composed of two different types of bacteria.
1. Aggregates small, barrel -shaped, actively motile and consisting of a central, polar
flagellate, rod-shaped bacterium with a covering of sulfur green bacteria.
Genus V. Chlorochromatium, p. 65.
2. Aggregates large, cylindrical, non-motile and composed of a central filamentous
bacterium with a more or less extensive covering of sulfur green bacteria.
Genus VI. Cylindrogloea, p. 66.
Genus I. Chlorobium Nadson, 1912.
(Bull. Jard. Imper. Botan., St. P^tersb., 12, 1912, 64 (Russian), 83 (German).)
Chlo.ro 'bi.um. Gr. adj. chlorus greenish yellow, green; Gr. noun hios life; M.L. neut.n.
Chlorobium green life.
Sulfur green bacteria, occurring singly or in chains, individual cells of various sizes and
shapes, from spherical to relatively long rod -shaped, the latter sometimes coiled into tight
spirals; often united in chains and generally embedded in a slime capsule. Non-motile.
Gram-negative. Contain a chlorophjdlous pigment different from the common green plant
chlorophylls and from bacteriochlorophyll. Capable of photosynthesis in the presence of
hydrogen sulfide, during which thej^ produce elemental sulfur which is excreted outside the
cells. Do not form spores.
The tj^pe species is Chlorobium limicola Nadson.
Key to the species of genus Chlorobium.
I. Does not utilize thiosulfates as oxidizable substrate.
1. Chlorobium limicola.
II. Utilizes thiosulfates as oxidizable substrate.
2. Chlorobium thiosulfatophihan.
1. Chlorobium limicola Nadson, 1912. appearance of the cultures. In healthy cul-
(Bull. Jard. Imper. Botan., St. Petersb., 12, tures the bacteria rem.ain evenly dispersed
1912, 64 (Russian), 83 (German).) and settle very slowly. Non-motile.
li.mi'co.la. L. noun limus mud; L. v. colo Color: Intensely green in healthy cul-
to dwell; M.L. fem.n. limicola the mud tures; poor pigmentation and then yellow-
dweller, ish green in media deficient in iron.
Cells occur in various sizes and shapes Strictly anaerobic, obligatory photosyn-
which are markedly dependent upon en- thetic bacteria whose occurrence in nature
vironmental conditions. In young and is dependent upon the presence of hydrogen
healthy cultures, predominantly small, sulfide. They utilize this substance, as well
ovoid to short rods, 0.7 by 0.9 to 1.5 microns, as elemental sulfur and molecular hydrogen,
frequently united in chains resemblingstrep- as oxidizable substrates; produce sulfur
tococci. Greatly elongated and irregularly from sulfides but do not store sulfur glob-
bent and curved rods also occur as involu- ules inside the cells. Oxidation of sulfide
tion forms; these rods may likewise remain may yield sulfur as an end product, but
united in chains. Club-shaped and spirally under optimal conditions the sulfur is
wound to tightly coiled involution forms further oxidized to sulfate. Unable to use
have been described, but the conditions for thiosulfate and tetrathionate as oxidizable
their formation are not understood, and in substrates. Development in organic media
recent pure-culture studies these have never free of sulfide has not been obtained,
been encountered (Larsen, Jour. Bact., 64, Source: Isolated from mud and stagnant
1952, 187). Regularly produce mucus; in water, containing hydrogen sulfide, from
media of inadequate composition this may the St. Petersburg Botanical Garden. Also
lead to the formation of cell conglomerates found by Bicknell (Lloydia, 12, 1949, 183) in
of different sizes and shapes and a stringy Sodon Lake, Bloomfield Hills, Michigan.
FAMILY III. CHLOROBACTERIACEAE
63
Habitat: Widely distributed in mud and
stagnant water. Mass development under
conditions of relatively high sulfide con-
centrations and low pH in environments
exposed to light.
Illustrations: Nadson, up. cit., 1912, PI.
Ill, fig. 3-12; van Niel, Arch. f. Mikrobiol.,
3, 1931, fig. 8, p. 66.
2. Chlorobiuin ihiosulfatophilum Lar-
sen, 1952. (Jour. Bact., 64, 1952, 187.)
thi.o.sul.fa.to'phi.lum. M.L. noun thio-
sulfatum thiosulfate; Gr.adj. phibis loving;
M.L. adj. thiosulfatophilus thiosulfate-
loving.
Cells indistinguishable from those of
Chlorobium limicola.
Color: As in Chlorobium limicola.
Strictly anaerobic, obligatory photosyn-
thetic bacteria. Utilize sulfides, sulfur,
thiosulfate, tetrathionate and molecular
hydrogen as o.xidizable substrates; produce
sulfate from inorganic sulfur compounds.
Unable to grow in organic media free of oxi-
dizable, inorganic sulfur compounds.
Distinctive characters: Differs from
Chlorobium limicola in its ability to oxidize
thiosulfate and tetrathionate.
Source: Isolated from marine and fresh-
water mud.
Habitat: Same as for Chlorobium limicola.
Genus II. Pelodictyon Lauterborn, 1913.
(Allgem. botan. Ztschr., 19, 1913, 98; Verhandl. naturhistor.-medizin. Vereins, Heidel-
berg, N.F. 13, 1915, 431.)
Pe.lo.dic'ty.on. Gr. adj. pelos dark-colored; Gr. noun dictyon net; M.L. neut.n. Pelodic-
tyon a dark-colored net.
Sulfur green bacteria, individual cells ovoid to distinctly rod-shaped, producing rather
extensive mucoid capsules and generally united into large colonies of characteristic shapes.
Non-motile. Contain chlorophyllous pigments different from the common green plant
chlorophylls and from bacteriochlorophyll. Capable of photosynthesis in the presence of
hydrogen sulfide, but do not store sulfur globules inside the cells.
The type species is Pelodictyon clathratiforme (Szafer) Lauterborn.
Key to the species of genus Pelodictyon.
I. Cells united in colonies in a net-like fashion.
1. Pelodictyon clathratiforme.
II. Cells arranged in tightly packed colonies without net-like structure.
A. Colonies composed of irregularly arranged cell-masses, extending in three dimen-
sions.
2. Pelodictyon aggregatum.
B. Colonies consisting of parallel strands and extending in two dimen.sions.
3. Pelodictyon parallelum.
1. Pelodictyon clathratiforme (Szafer,
1910) Lauterborn, 1913. (Aphanothece clath-
ratiforme Szafer, Bull. Acad. Sci., Cracovie,
Ser. B, 3, 1910, 162; Lauterborn, Allgem.
botan. Ztschr., 19, 1913, 98; also see Ver-
handl. naturhist.-medizin. Vereins, Heidel-
berg, N.F. 13, 1915,430.)
clath.ra.ti.for'me. L. part. adj. clathratus
latticed; L. noun forma shape, form; M.L.
adj. clathratiformis lattice-like.
Cells generally rod-shaped, ranging from
slightly elongated ovoids to distinct rods,
often vacuolated, about 0.5 to 1.5 by 2 to 4
microns, producing rather wide slime cap-
sules. Characteristically united into three-
dimensional colonies which present a
net-like appearance with mazes of about 10
to 50 microns. Non-motile.
Color: Yellowish green.
Abnormal cell forms (involution forms)
not uncommon, consisting of elongated and
curved, forked, or club-shaped and swollen
rods, occasionally suggesting rudimentary
branching at the extremities. Such cells
64
ORDER I. PSEUDOMONADALES
may be found as elements in chains for the
greater part composed of normal individ-
uals.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hy-
drogen sulfide and exposed to light; sulfur
springs.
Illustrations: Szafer, op. cit., 1910, PI.
VI, fig. 5; Perfiliev, Jour. Microbiol. (Rus-
sian), 1, 1914, PI. II, fig. 1, 5-12; Lauter-
born, op. cit., 1915, PI. Ill, fig. 33.
2. Pelodictyon aggregatuin Perfiliev,
1914. (Aphanothece luteola Schmidle, Bei-
hefte Botan. Cent., 10, 1901, 179; Perfiliev,
Jour. Microbiol. (Russian), 1, 1914, 197.)
ag.gre.ga'tum. L. part. adj. aggregatus
added to a flock, aggregated, clumped.
Cells usually rod-shaped, about 1 to 1.5
by 2 to 4 microns, often vacuolated, produc-
ing slime capsules and united into irregu-
larly shaped, three-dimensional colonies in
which the cells are more or less tightly
packed without orderly arrangement. Colo-
nies may attain a size of up to 1 mm; fre-
quently they are not fully compact but
contain less dense areas or appear perfo-
rated, thus forming transition stages to
Pelodictyon claihratiforme. Non-motile.
Color: Yellowish green.
Abnormal cell forms (involution forms)
usually in the shape of elongated and
curved, forked or club-shaped and swollen
rods, occasionally suggesting branching at
extremities.
Source: Isolated from sulfureted water in
Europe ; also reported by Button and Juday
(Ecology, 25, 1944, 277) from Scaffold Lake,
Wisconsin.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hydro-
gen sulfide and exposed to light; sulfur
springs.
Illustrations: Perfiliev, ibid., PI. II, fig.
2; Lauterborn, Verhandl. naturhistor.-
medizin. Vereins, Heidelberg, N.F. 13, 1915,
PI. Ill, fig. 29-31.
3. Pelodictyon parallelum (Szafer,
1910) Perfiliev, 1914. (Aphanothece parallela
Szafer, Bull. Acad. Sci., Cracovie, Ser. B, 3,
1910, 163; Perfiliev, Jour. Microbiol. (Rus-
sian), 1, 1914, 198.)
pa.ral.le'lum. Gr. adj. parallelus parallel.
Cells rather small, spherical to ovoid, or
even rod-shaped; about 0.5 to 1 by 1 to 3
microns, occurring in chains and forming
flat, plate-like, two-dimensional aggre-
gates in which the chains are arranged as
parallel strands. Non-motile.
Color: Yellowish green.
Abnormal cell forms not specifically men-
tioned, but likely to occur and to resemble
those of other species.
This species may well be a special growth-
form of Chlorobium limicola.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hydro-
gen sulfide and exposed to light; sulfur
springs.
Illustrations: Szafer, op. cit., 1910, PI.
VI, fig. 7; Perfiliev, op. cit., 1914, PI. II, fig.
2.
Genus HI. Clathrochloris Geitler, 1925.
(Geitler, in Pascher, Die Siisswasserflora Deutschlands, Osterreichs und der Schweiz,
Jena, 12, 1925, 457.)
Clath.ro. chlo'ris. L. pi. noun clathri lattice; Gr. adj. chlorus green; M.L. fem.n. Clathro-
chloris green lattice.
Sulfur green bacteria of small size, generally spherical and arranged in chains which are
united into loose, trellis-shaped aggregates, somewhat similar to those of Pelodictyon cla-
thratiforme and Pelodictyon aggregatum. Cells usually contain sulfur globules. Color is yel-
loivish green. Non-motile.
The type species is Clathrochloris sulphurica (Szafer) Geitler.
1. Clathrochloris sulphurica (Szafer,
1910) Geitler, 1925. {Aphanothece sulphurica
Szafer, Bull. Acad. Sci., Cracovie, Ser. B, 3,
1910, 162; Geitler, Die Siisswasserflora
Deutschlands, Osterreichs und der Schweiz,
Jena, 12, 1925, 457.)
FAMILY III. CHLOROBACTERIACEAE
65
sul.phur'i.ca. L. noun sulfw' (sometimes
sulphur) sulfur; IM.L. adj. sulphuricus sul-
furic.
Cells spherical, about 0.5 to 0.7 micron
in diameter, usually containing sulfur glob-
ules. Non-motile.
Color: Yellowish green.
The reported occurrence of sulfur globules
in the cells of this very small species is sur-
prising; it is the only one among the sulfur
green bacteria in which these inclusions
have been encountered. The published de-
scriptions are even more fragmentary than
those of other members of the group.
Source : Reported only from sulfur springs
in Lubien Wielki, near Lwow, Poland.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hj^-
drogen sulfide and exposed to light; sulfur
springs.
Illustration: Szafer, op. cit., 1910, PI. VI,
fig. 6.
Genus IV. Chlorohaclerium Lauterborn, 1915.
(Luuterborn, Verhandl, naturhist.-medizin. Vereins, Heidelberg, N.F., 13, 1915, 429; not
Chlorobacteriwn* Guillebeau, Landw. Jahrb. d. Schweiz, 4, 1890, 32.)
Chlo.ro. bac.te'ri.um. Gr. adj. chlonis green; L. noun bacterium a small rod; M.L. neut.n.
Chlorobacterium a green rodlet.
Sulfur green bacteria(?) which grow sj'mbiotically as an outside covering on cells of pro-
tozoa, such as amoeba and flagellates. Cells rod-shaped, often slightly curved, greenish.
Non-motile.
The type species is Chlorobacterium symhioticum Lauterborn.
1. Chlorobacterium symbioticum Lau-
terborn, 1915. (Verhandl. naturhist.-medi-
zin. Vereins, Heidelberg, X.F., IS, 1915,
429.)
sym.bi.o'ti.cum. Gr. adj. stjnibioticus of
companionship, sj'mbiotic.
Cells rod-shaped, about 0.5 by 2 to 5
microns, often slightly curved. Non-motile.
Occur as a peripheral covering of certain
protozoa with which they may form a sym-
biotic unit.
It is not certain that this is a sulfur green
bacterium: the descriptions of localities
Avhere it was found fail to mention the pres-
ence of hydrogen sulfide in the environ-
ment; this should be a prerequisite for a
member of this group.
Source : Reported from a number of pools
in Germany.
Habitat: Stagnant water.
Illustrations: Lauterborn, loc. cit., PL III,
fig. 34-36; Pascher, Die Siisswasserflora
Deutschlands, Osterreichs und der Schweiz,
Jena, 12, 1925, fig. 149.
Genus V . Chlorochromatiuni Lauterborn, 1906.
(Allgem. botan. Ztschr., 19, 1906, 196.)
Chlo.ro. chro.ma'ti.um. Gr. adj. chlorus green; Gr. noun chromatium color, paint; M.L.
neut.n. Chromatium a bacterial genus; M.L. neut.n. Chlorochromatium a green Chromatium.
Sulfur green bacteria, ovoid to rod -shaped with rounded ends. Occur as barrel -shaped
aggregates consisting of a rather large, colorless, polar flagellate bacterium as the center
which is surrounded by green bacteria, arranged in 4 to 6 rows, ordinaril}' from 2 to 4 cells
high. The entire conglomerate behaves like a unit, is motile, and multiplies by the more or
less simultaneous fission of its components.
The green constituents contain a chlorophyllous pigment which is not identical with the
common green plant chlorophylls or with bacteriochlorophjdl. Capable of photosj-nthesis
in the presence of hydrogen sulfide but do not store sulfur globules in the cells.
The type species is Chlorochromatium aggregatum Lauterborn.
* It has been proposed that Chlorobacterium Guillebeau be rejected as a generic name in
bacteriology and placed in the list of nomina generica rejicienda (Internat. Bull. Bact.
Nomen. and Tax., 1, 1951, 43 and 2, 1952, 110).
GG
ORDER I. PSEUDOMONADALES
1. Chlorochromatium aggregatum
Lauterborn, 1906. (AUgem. botan. Ztschr.,
19, 1906, 196.)
ag.gre.ga'tum. L. part. adj. aggregatus
flocked together, clumped.
Cells of the green component 0.5 to 1.0 by
1.0 to 2.5 microns, mostly from 8 to 16 in-
dividuals surrounding the central bac-
terium. Size of the total barrel-shaped unit
variable, generally 2.5 to 5 by 7 to 12 mi-
crons. Occasionally a group of the complex
colonies may remain attached in a chain.
Anaerobic.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hydro-
gen sulfide and exposed to light.
There is at present no good reason for
distinguishing 2 varieties (forma typica and
forma minor) or even species, on the basis
of size differences of the colony, as Geitler
proposed (Die Siisswasserflora Deutsch-
lands, Osterreichs und der Schweiz, Jena,
12, 1925, 460). The reported and personally
observed sizes of such units show that the
extreme limits are linked by a complete
series of transitions.
Illustrations: Buder, Ber. deut. botan.
Ges., 31, 1914, Generalversam., PI. XXIV,
fig. 1-5; Perfiliev, Jour. Microbiol. (Rus-
sian), 1, 1914, fig. 1-5, p. 213.
Genus VI. Cylindrogloea Perfiliev, 1914-
(Jour. Microbiol. (Russian), 1, 1914, 223.)
Cy.lin.dro.gloe'a. Gr. noun cylindrus cylinder; Gr. novm gloea gum; M.L. fem.n. Cylin-
drogloea cylindrical gum.
Sulfur green bacteria consisting of small ovoid to rod-shaped cells growing in association
with a filamentous, colorless, central bacterium, thus forming colonies of a cylindrical
shape. Non-motile. The green component contains a chlorophyllous pigment different from
the common chlorophylls of green plants and from bacteriochlorophyll. Capable of photo-
synthesis in the presence of hydrogen sulfide without depositing sulfur globules in the cells.
The type species is Cylindrogloea bacterifera Perfiliev.
1. Cylindrogloea bacterifera Perfiliev,
1914. (Jour. Microbiol. (Russian), 1, 1914,
223.)
bac.te.ri'fe.ra. Gr. neut.n. bactrum a rod;
M.L. mas.n. bacter rod (a combining form);
L. verbal suf. fer bearing; M.L. adj. bac-
terifera rod-bearing.
Individual green components ovoid to
rod-shaped, about 0.5 to 1 by 2 to 4 microns,
very similar to those of the complex Chloro-
bacterium symbioiicum and Chlorochromatium
aggregatum with which they may well be
identical. The central filamentous bac-
terium is embedded in a slime capsule of
considerable dimensions. This, in turn, is
surrounded by a layer of green bacteria,
usually one cell thick. The green organisms
may form a very dense outer covering, or
they may be more sparsely distributed over
the slime capsule. The entire unit is again
surrounded by a sizeable slime zone. Aggre-
gates measure about 7 to 8 microns in width
and up to 50 microns in length; they are non-
motile. Both components appear to be
non -spore-forming.
Habitat: Mud and stagnant water con-
taining rather high concentrations of hj^dro-
gen sulfide and exposed to light.
Illustration: Perfiliev, ibid., fig. 6-11, p.
213.
Perfiliev rightly emphasizes, as Buder had
done for Chloronium mirabile, the provi-
sional nature of thus using a generic desig-
nation for an apparently stable complex
composed of two different organisms. It re-
mains possible that the last three genera of
symbiotic entities represent fortuitous com-
binations whose occurrence is conditioned
by environmental factors. If so, the generic
terminology would be devoid of any taxo-
nomic significance, and the green bacteria
should be relegated to more appropriate
genera. Indications suggestive of this state
of affairs can be found in the literature ; for
example in Utermohl's observation (Archiv
FAMILY I. NITROBACTERACEAE 67
f. Hydrobiol., Suppl. 5, 1925, 279) that the grate, whereupon the green constituents
complex Chlorochromatium aggregalum may, appear as small Pelodictyon aggregatum
especially in the presence of oxygen, disinte- {Schmidlea luteola) colonies.
Suborder II. Pseiidomonadineae Breed, Murray and Smith, Sub-Ordo Nov.
Pseu.do.mo.na.di'ne.ae. M.L. fem.n. Pseudomonas, -adds a genus of bacteria; -ineae end-
ing to denote a suborder; ALL. fern. pi. n. Pseiidovionadineae the Pseudomonas suborder.
Cells normally about 1 micron in diameter, but among the colorless sulfur bacteria they
may be as much as 14 microns in diameter. The cells do not contain photosynthetic pig-
ments, but diffusible, water-soluble pigments of a type not found elsewhere among bacteria
occur in many species. Also, non-water-soluble yellow or red pigments occur in some gen-
era. The motile species are invariably polar flagellate. Some groups are strictly autotrophic,
oxidizing simple inorganic compounds; others are also oxidative but are facultatively
heterotrophic or heterotrophic in their physiology. A few genera include species that fer-
ment simple sugars, even producing H2 and CO2 as do the common acid- and gas-producing
coliform bacteria. The organisms in one genus {Zijmomonas) even carry out an alcoholic
fermentation similar to that of j^easts. The majority of the species grow well and fairly
rapidly on the surfaces of ordinary culture media. Some species that attack agar or cellulose
or that show other unusual types of physiology are more fastidious in their requirements.
Only a few species are strictly anaerobic as in Vibrio and Desulfovibrio . The species in this
suborder are largely found in salt- or fresh-water or in soil. Some are parasitic and a few
are pathogenic to vertebrates including man.
Key to (he families of siihordcr Pseiidomonadineae.
I. Coccoid to rod-shaped cells. Occasionally individual rods may be curved although the
majority of the cells are straight.
A. Cells not attached to a substrate.
1. Cells have the power to oxidize simple compounds such as ammonia, nitrites,
methane, hydrogen, carbon monoxide, sulfur or sulfur compounds. Chemo-auto-
trophic or facultatively chemo-autotrophic.
a. Do not secure their energy from the oxidation of sulfur compounds,
b. Oxidize ammonia to nitrites or nitrites to nitrates. Autotrophic.
Family I. Nitrobacteraceae, p. 68.
bb. Oxidize methane, hydrogen or carbon monoxide. Autotrophic.
Family II. Methanomonadaceae , p. 74.
aa. Oxidize sulfur compounds, frequently with a deposit of free sulfur granules
or crystals within or without the cells.
Family III. Thiobacteriaceae, p. 78.
2. Cells frequently oxidative, although they are sometimes fermentative in their
physiology. Usually heterotrophic. Rarely facultatively autotrophic
Family IV. Pseudomonadaceae, p. 88.
B. Cells in free-floating films or attached to a substrate.
a. Cells attached to the substrate by means of a stalk, usually with a holdfast.
Family V. Caulobacteraceae , p. 212.
aa. Cells in free-floating films or attached to the substrate by means of capsular
material.
Family VI. Siderocapsaceae, p. 217.
II. Curved, vibrio-like to spiral-shaped cells.
Family VII. Spirillaceae, p. 228.
68 ORDER I. PSEUDOMONADALES
FAMILY I. NITROBACTERACEAE BUCHANAN, 1917.*
(Jour. Bact., 2, 1917, 349 and Jour. Bact., S, 1918, 179.)
Ni.tro.bac.te.ra'ce.ae. M.L. A^ z'^rofeacier name of type genus of the family; -aceae ending
to denote a family; M.L. plural noun Nitrohacteraceae the Nitrobacter family.
Cells without endospores; rod-shaped, ellipsoidal or even spherical (Nitrosococcus) or
spirillar in shape (Nitrosospira) . Flagella polar, occasionally absent. Gram-negative. Or-
ganisms which derive energy from the oxidation of ammonia to nitrite or from the oxida-
tion of nitrite to nitrate; these bacteria depend on this oxidation for growth and fail to grow
on media containing organic matter in the absence of the specific inorganic materials used
as sources of energy. Many organic compounds commonly used in standard culture media
are toxic to these bacteria. Not parasitic. Commonly found in soil and fresh water.
The type genus for the family is Nitrobacter Winogradsky.
As it appears to be a more logical arrangement, the genera that include the species that
oxidize ammonia are discussed first although the type genus does not belong in this group.
Key to the genera of family Nitrohacteraceae.
I. Ammonia oxidized to nitrite.
A. Zoogloeae not formed. Cells occur separately, free or in dense aggregates.
1. Cells not spiral -shaped.
a. Cells ellipsoidal.
Genus I. Nitrosomonas, p. 68.
aa. Cells spherical.
Genus II. Nitrosococcus, p. 69.
2. Cells spiral.
Genus III. Nitrosospira, p. 70.
B. Zoogloeae formed.
1. Zoogloea surrounded by a common membrane forming a cyst.
Genus IV. Nitrosocystis, p. 70.
2. No common membrane surrounds the cells. The massed cells are embedded in
slime.
Genus V. Nitrosogloea, p. 71.
II. Nitrite oxidized to nitrate.
A. Zoogloeae not formed.
Genus VI. Nitrobacter, p. 72.
B. Zoogloeae formed.
Genus VII. Nitrocystis, p. 73.
Genus I. Nitrosomonas Winogradsky , 1890.
{Nitromonas Winogradsky, Ann. Inst. Past., 4, 1890, 257; not Nitromonas Orla-Jensen,
Cent. f. Bakt., II Abt., 22, 1909, 334; Arch. Sci. biol., St. Petersburg, 1, 1892, 127; emend.
S. and H. Winogradsky, Ann. Inst. Past., 50, 1933, 393.)
Ni.tro.so.mo'nas. M.L. nitrosus nitrous; Gr. monas, monadis a unit, monad; M.L. fem.n.
Nitrosomonas nitrous monad, i.e., the monad producing nitrite.
Cells ellipsoidal, non-rhotile or with a single polar flagellum, occurring singly, in pairs,
short chains or irregular masses which are not enclosed in a common membrane. O.xidize
ammonia to nitrite more rapidly than the other genera of this family.
The type species is Nitrosomonas europaea Winogradsky.
* Completely revised by Dr. R. L. Starkey, New Jersey Agricultural Experiment Station,
New Brunswick, N. J., March, 1943; minor revisions, November, 1953.
FAMILY I. NITROBACTERACEAE
1. Nitrosomonas europaea Winograd-
sky, 1892. (Arch. Sci. biol., St. Petersburg,
1, 1892, 127.)
eu.ro. pae'a. Gr. adj. europaeus of Europe,
European.
Rods, 0.9 to 1.0 by 1.1 to 1.8 microns, oc-
curring singly, rarely in chains of three to
four. Possess a single polar flagellum 3 to 4
times the length of the rods, or rarely one
at either end.
Grow readily in aqueous media without or-
ganic matter and containing ammonium
sulfate, potassium phosphate and mag-
nesium carbonate. The cells accumulate in
soft masses around the particles of mag-
nesium carbonate at the bottom of the flask.
The liquid is occasionally turbid through
development of motile swarm cells or
monads.
Small, compact, sharply defined colonies
brownish in color on silica gel.
Aerobic.
Strictly autotrophic.
Source: Soils of Zurich, Switzerland; of
Gennevilliers, France; and Kazan, Russia.
Habitat: Presumably widely distributed
in soil.
2. Nitrosomonas monocella Nelson,
1931. (Zent. f. Bakt., II Abt., 83, 1931, 287.)
mon.o.cel'la. Gr. monus single; L. cella
oom, cell; M.L. adj. monocellus one-celled.
Ovoid rods, 0.6 to 0.9 micron, often oc-
curring in pairs. Young cells nearly spheri-
cal. Motile by means of a single polar
flagellum 3 to 5 times as long as the rod.
Gram-positive (Nelson). Found negative by
H. J. Conn (personal communication).
No growth in nutrient broth, nutrient
agar, nutrient or plain gelatin, plain or lit-
mus milk, glucose or plain yeast water, or on
potato.
Silica gel or agar plates of inorganic
medium: No typical colonies, but yellowish
brown masses of growth around particles of
CaCOs in the medium.
Inorganic liquid medium containing
ammonium salts: Uniform development
throughout the liquid as well as in the
carbonate sediment.
Even low concentrations of organic mat-
ter retard or completely inhibit the initia-
tion of growth. Plant extracts are toxic.
Free CO 2 and O2 necessary for growth.
Optimum pH, 8.0 to 9.0. Poor growth
below pH 7.0. Some growth above pH 9.0.
Optimum temperature for growth and oxi-
dation, 28° C.
Aerobic .
Strictly autotrophic.
Source: Isolated from field soil.
Habitat: Presumably widely distributed
in soil.
Genus II. Nitrosococcus Winogradshj, 1892.
(Arch. Sci. biol., St. Petersburg, 1, 1892, 127.)
Ni.tro.so.coc'cus. M.L. nitrosus nitrous; Gr. coccus grain, berry; M.L. mas.n. Nitroso-
coccus nitrous sphere.
Cells large spheres, non-motile, not producing zoogloeae. Oxidize ammonia to nitrite.
The type species is Nitrosococcus nitrosus (Migula) Buchanan.
1. Nitrosococcus nitrosus (Migula,
1900) Buchanan, 1925. {Nitrosococcus Wino-
gradsky, Ann. Inst. Past., 5, 1891, 577;
Arch. Sci. biol., St. Petersburg, 1, 1892, 127;
Micrococcus nitrosus Migula. Sj^st. d. Bakt.,
2, 1900, 194; Buchanan, Gen. Syst. Bact.,
1925, 402.)
ni.tro'sus. M.L. adj. nitrosus nitrous.
Large spheres, 1.5 to 1.7 microns in size,
with thick cell membranes. Motility could
not be demonstrated. Stain readily with
aniline dyes. Zoogloea formation not ob-
served. Gram-positive (Omelianski, Cent. f.
Bakt., II Abt., 19, 1907, 263).
Liquid medium: Turbid.
Silica gel: Both dark and light colonies.
Surface colonies look like small drops of a
turbid yellowish liquid.
Aerobic.
Optimum temperature, lietween 20° and
25° C.
Source: Isolated from .soil from (^uito.
70 ORDER I. PSEUDOMONADALES
Ecuador; Campinas, Brazil; and Melbourne, Habitat: Presumably widely distributed
Australia. in soil.
Genus III. Nitrosospira Winogradsky, 1931.
(S. Winogradsky, Compt. rend. Acad. Sci., Paris, 192, 1931, 1004; S. Winogradsky
and H. Winogradsky, Ann. Inst. Past., 50, 1933, 394 and 406.)
Ni.tro.so.spi'ra. M.L. nifrosus nitrous; Gr. spira a coil, spiral; M.L. fem.n. Nitrosospira
nitrous spiral.
Cells spiral-shaped. Oxidize ammonia to nitrite very slowly.
The type species is Nitrosospira briensis S. Winogradsky and H. Winogradsky.
1. Nitrosospira briensis S. Winogradsky Habitat: Presumably widely distributed
and H. Winogradsky, 1933. (Ann. Inst. in soil.
Pasteur, 50, 1933, 407.)
bri.en'sis. French Brie, place name; M.L. 2. Nitrosospira antarctica S. Wino-
adj. briensis of Brie. gradsky and H. Winogradsky, 1933. (Ann.
Spirals wound tightly to form very small Inst. Past., 50, 1933, 407.)
cylinders as long as 15 to 20 microns. Short ant. arc 'tic. a. Gr. anti opposite; Gr. arcius
spirals have the appearance of short rods a bear; Gr. adj. antarcticus opposite the
and ellipsoidal cells. Small pseudo-cocci ob- bear, antarctic,
served in old cultures. Cells and colonies similar to those of N .
Colonies on silica gel: Small, occasionally briensis except that the cells are generally
containing cyst-like aggregates of cells. The wound together to form more compact
cysts are more poorly developed than in spirals.
Nitrosocystis. Aerobic.
Aerobic. Optimum pH, between 7.0 and 7.2.
Optimum pH, between 7.0 and 7.2. Source: Soil from the Antarctic.
Source: Uncultivated pasture soil of Brie, Habitat: Presumably widely distributed
France. in soil.
Genus IV. Nitrosocystis Winogradsky, 1931.
(S. Winogradsky, Compt. rend. Acad. Sci., Paris, 192, 1931, 1003; also see S. Winogradsky
and H. Winogradsky, Ann. Inst. Past., 50, 1933, 394 and 399.)
Ni.tro.so.cyst'is. M.L. adj. nitrosus nitrous; Gr. noun cystis bladder, cyst; M.L. fem.n.
Nitrosocystis nitrous cyst.
Cells ellipsoidal or elongated, uniting in compact, rounded aggregates surrounded by a
common membrane to form cysts. The cysts disintegrate to free the cells, particularly when
transferred to fresh media. Within the cyst the cells are embedded in slime. Ammonia is
oxidized to nitrite at a rate intermediate between that of Nitrosoijionas and that of Nitroso-
spira.
Winogradsky and Winogradsky (ibid., 393) differentiated between Nitrosomonas and
Nitrosocystis in that the former produced soft (or clear) colonies and the latter produced
hard (or dark) colonies on silica gel. However, Kingma Boltjes (Arch. Mikrobiol., 6,
1935, 79) was able to obtain both hard and soft colonies in cultures of Nitrosomonas derived
from single-cell isolates. Meiklejohn (Nature, 168, 1951, 561 ; also see Jour. Soil Sci., 4, 1953,
62), furthermore, states that the appearance of hard or soft colonies is dependent upon the
density of the silica gel and upon whether the colonies are in the gel or on the surface;
consequently she regards Nitrosocystis as probably identical with Nitrosomonas. Some ob-
servers (Imsenecki, Nature, 157, 1946, 877; and Grace, Nature, 168, 1951, 117; also see
Riassunti d. Comunicazione, VI Cong. Internaz. d. Microbiol., Roma, 1, 1953, 53) have
FAMILY I. NITROBACTERACEAE
71
suggested that the organisms described in this genus are myxobacters or that the cultures
were contaminated with myxobacters; however, this does not seem probable.
The type species is Nitrosocystis javanensis (Winogradsky) Starkey.
1. Nitrosocystis javanensis (Wino-
gradsky, 1892) Starkey, 1948. {Nitrosomonas
javanensis Winogradsky, Arch. Sci. biol.,
St. Petersburg, 1, 1892, 127; Starkey, in
Manual, 6th ed., 1948, 72.)
jav.a.nen'sis. Java, a place name; M.L.
adj. javanensis of Java, Javanese.
Small ellipsoidal cells having a diameter of
0.5 to 0.6 micron. Possess a polar flagellum
20 times as long as the rods.
In liquid medium produces very compact
zoogloeal masses of cells and motile swarm-
ers. The large zoogloeae are themselves
composed of smaller compact aggregates of
cells.
On silica gel the colonies are circular to
elliptical becoming clear or light brown.
Aerobic.
Strictly autotrophic.
Source: Soil of Buitenzorg, Java; Tokyo,
Japan; and La Reghaia, Tunisia.
Habitat: Presumably wddely distributed
in soil.
2. Nitrosocystis coccoides Starkey,
1948. {Nitrosocystis a, S. Winogradsky and
H. Winogradsky, Ann. Inst. Past., 50, 1933,
401; Starkey, in Manual, 6th ed., 1948, 72.)
coc.co.i'des. Gr. coccus grain, berry; Gr.
idus form, shape; M.L. adj. coccoides coccus-
shaped.
Ellipsoidal cells about 1.5 microns in di-
ameter. Occur as compact aggregates of
cells imbedded in mucus and surrounded by
a thickened capsule to form cyst-like bod-
ies. Cells rarely solitary but more often in
pairs and in small groups of four or more.
Probably motile. The mucus which sur-
rounds the cells is not readily stained
whereas the outside coating stains more
easily.
Colonies on silica gel : As colonies develop,
the coating of CaCOs on the gel becomes
yellowish and dissolves, and the colony ap-
pears as a bulbous, angular, brown body
which may become 0.5 mm in diameter.
The cells are held firmly together in these
irregularly shaped bulbous aggregates.
Aerobic.
Source: Poor soils of Brie and elsewhere
in France.
Habitat: Presumably widely distributed
in forest and manured soils.
Genus V. Nitrosogloea H. Winogradsky, 1935.
(Compt. rend. Acad. Sci., Paris, £00, 1935, 1887; Ann. Inst. Pasteur, 58, 1937, 335.)
Ni.tro.so.gloe'a. M.L. nitrosus nitrous; Gr. gloea glue, jelly; M.L. fem.n. Nitrosogloea
nitrous jelly.
Cells ellipsoidal or rod-shaped. Embedded in slime to form zoogloeae. No common mem-
brane surrounds the cell aggregates. Oxidize ammonia to nitrite.
It has been suggested that these organisms were contaminated with myxobacters. See
note under Nitrosocystis for references.
The type species is Nitrosogloea merismoides H. Winogradsky.
1. Nitrosogloea merismoides H. Wino-
gradsky, 1935. {Nitrosocystis "I", H. Wino-
gradsky, Trans. Third Intern. Cong. Soil
Sci., Oxford, 1, 1935, 139; H. Winogradsky,
Compt. rend. Acad. Sci., Paris, 300, 1935,
1887; also see Ann. Inst. Past., 58, 1937,
333.)
mer.is.mo.i'des. Gr. merismus a divi-
sion; Gr. idus form, shape; M.L. adj. meris-
moides resembling a division.
Ellipsoidal cells 0.5 by 1.5 microns. Oval
cells or short rods forming tetards or
chains, each group with its own sheath. The
groups vary in shape producing branched
chains, irregular or compact aggregates.
Colonies on silica gel: Cells encased in a
pale yellow mucilage giving the colony a dull
appearance. Colony surface studded with
little humps.
Aerobic.
ORDER I. PSEUDOMONADALES
Source: Activated sludge.
Habitat: Unknown.
Source: Activated sludge.
Habitat: Unknown.
2. Nitrosogloea schizobacteroides H.
Winogradsky, 1935. (Nitrosocystis "11", H.
Winogradsky, Trans. Third Intern. Cong.
Soil Sci., Oxford, 1, 1935, 139; H. Winograd-
sky, Compt. rend. Acad. Sci., Paris, 200,
1935, 1887; Ann. Inst. Past., 58, 1937, 333.)
schiz.o.bac.te.ro.i'des. Gr. schiza cleft,
fission; Gr. noun bactrmn a rod; Gr. noun
idus form, shape; M.L. adj. schizobacteroides
shaped like a fission rod.
Elongated rods or short filaments 3 to 4
microns long.
Colonies on silica gel : Flat groups of cells
are produced which are united in a common
sheath. The aggregates form a pseudo-tissue
of interwoven filaments suggestive of a
fungus pad. The pad can be removed as a
unit from the medium.
Aerobic.
3. Nitrosogloea membranacea H.
Winogradsky, 1935. (Nitrosocystis "III", H.
Winogradsky, Trans. Third Intern. Cong.
Soil Sci., Oxford, 1, 1935, 139; Compt. rend.
Acad. Sci., Paris, 200, 1935, 1887; Ann. Inst.
Past., 58, 1937, 333.)
mem.bran.a'ce.a. L. adj. membranaceus
membranaceous .
Ellipsoidal cells commonly in pairs and
also solitary.
Colonies on silica gel: Appear as dull mu-
coid material with a pale straw color. The
cells are held firmly together so that the
entire colony is easily picked up with the
transfer needle. No structural units within
the colony.
Aerobic.
Source: Activated sludge.
Habitat: Unknown.
Genus VI. Nitrobacter Winogradsky, 1892.
(Arch. Sci. biol., St. Petersburg, 1, 1892, 127.)
Ni.tro.bac'ter. Gr. noun nitrum nitre, M.L. nitrate; M.L. noun bacter the masculine
form of the Gr. neut.n. bactrum a rod; M.L. mas.n. Nitrobacter nitrate rod.
Cells rod-shaped. Oxidize nitrites to nitrates.
The type species is Nitrobacter winogradshji Winslow et al.
1. Nitrobacter winogradskyi Winslow
et al., 1917. (Nitrobacter Winogradsky,
Arch. Sci. biol., St. Petersburg, 1, 1892, 127;
Bacterium nitrobacter Lehmann and Neu-
mann, Bakt. Diag., 2 Aufl., 2, 1899, 187;
Winslow et al.. Jour. Bact., 2, 1917, 552.)
wi.no.grad'sky.i. Named for S. Wino-
gradsky, the microbiologist who first iso-
lated these bacteria; M.L. mas.gen.n.
winogradskyi of Winogradsky.
Description taken from Gibbs (Soil Sci.,
8, 1919, 448).
Short, non-motile rods with gelatinous
membrane, 0.6 to 0.8 by 1.0 to 1.2 microns.
Do not stain readily. Gram-negative
(Omelianski, Cent. f. Bakt., II Abt., 19,
1907, 263).
Can be cultivated on media free of organic
matter. Sensitive to certain organic com-
pounds.
Washed agar colonies: In 7 to 10 days
very small, light brown, circular to irregular
colonies, becoming darker.
Silica gel: Colonies smaller but more
dense than those on washed agar.
Washed agar slant: In 7 to 10 days scant,
grayish streak.
Inorganic solution medium: After 10 daj's
flocculent sediment. Sensitive to ammonium
salts under alkaline conditions.
Nitrite is oxidized to nitrate.
Aerobic.
Strictly autotrophic.
Optimum temperature, between 25° and
28° C.
Source: Soil.
Habitat: Presumably widely distributed
in soil.
2. Nitrobacter agilis Nelson, 1931.
(Zent. f. Bakt., II Abt., 83, 1931, 287.)
FAMILY I. NITROBACTERACEAE
73
ag'il.is. L. adj. agilis agile, quick.
Rods, 0.5 by 0.8 to 0.9 micron, occurring
singly, sometimes in pairs or larger aggre-
gates. Rapidly motile with a long, thin,
polar flagellum often 7 to 10 times as long as
the rod. (Non-motile culture obtained by
Kingma Boltjes, Arch. f. Mikrobiol., 6,
1935, 79.) Gram-negative.
No growth in nutrient broth, nutrient
agar, nutrient or plain gelatin, litmus or
plain milk, glucose or plain yeast water, or
on potato.
Nitrite agar: After two weeks, produces
semi -spherical, minute, nearly transparent
colonies. Oxidation usually complete in 10
to 14 days.
Inorganic liquid medium containing ni-
trite: Produces uniformly dispersed growth.
Optimum pH, between 7.6 and 8.6. Limits
of growth, 6.6 to 10.0.
Temperature relations: Optimum for
growth, between 25° and 30° C. Optimum
for oxidation, 28° C. No oxidation at 37° C.
Thermal death point, 60° C. for five min-
utes.
Strictly autotrophic.
Aerobic.
Source: Isolated from greenhouse soils
and from sewage effluents in Madison,
Wisconsin.
Habitat: Presumably widely distributed
in soil.
Genus VII. Nitrocystis H. Winogradsky , 1935.
(Trans. Third Intern. Cong. Soil Sci., Oxford, 1, 1935, 139.)
Ni.tro.cyst'is. Gr. noun nitrum nitre, M.L. nitrate; Gr. noun cyslis bladder, cj'st; M.L.
fem.n. Nitrocystis nitrate cyst.
Cells ellipsoidal or rod-shaped. Embedded in slime and united into compact zoogloeal
aggregates. Oxidize nitrites to nitrates.
It has been suggested that these organisms were really myxobacters. See note under
Nitrosocystis for references.
The type species is Nitrocystis sarcinoides H. Winogradsky.
1. Nitrocystis sarcinoides H. Wino-
gradsky, 1937. {Nitrocystis B. A., H. Wino-
gradsky, Compt. rend. Acad. Sci., Paris,
200, 1935, 1888; also see Ann. Inst. Past.,
58, 1937, 336.)
sar.cin.o.i'des. L. sarcina a packet; Gr.
idus form, shape; M.L. adj. sarcinoides re-
sembling Sarcina, a genus of bacteria.
Small rods 0.5 by 1.0 micron. Cells ellip-
soidal or wedge-shaped and grouped in sar-
cina-like packets.
Colonies on silica gel : On the surface of
gel coated with kaolin the colonies appear
as small, raised, amber warts. The colonies
grow up to 5 mm in diameter. The colonies
are viscous and sticky when young, and
they become brown with age, shrink, and
look like scales and become hard like grains
of sand. Each colony is enveloped in several
layers of a thick slime which holds the cells
together so that the entire colony can be
removed with a transfer needle.
Aerobic.
Source: Activated sludge.
Babitat: Unknown.
2. Nitrocystis micropunctata (H.
Winogradsky, 1935) H. Winogradsky, 1937.
(Nitrocystis "III", H. Winogradsky, Trans.
Third Intern. Cong. Soil Sci., Oxford, 1,
1935, 139; Nitrogloea micropiinctata H.
Winogradsky, Compt. rend. Acad. Sci.,
Paris, 200, 1935, 1888; H. Winogradsky,
Ann. Inst. Past., 58, 1937, 326.)
mi.cro.punc.ta'ta. Gr. micrus small; L.
punctatus spotted; M.L. adj. micropunctatus
full of small spots.
Cells are ellipsoidal rods, about 0.5 mi-
cron in diameter, which stain poorly except
at the ends. Encased in a viscous slime.
Colonies on silica gel: Like those of A^.
sarcinoides except that they are clearer and
have a more plastic consistency. The cells
are not held together by the slime in the
colony as with N. sarcinoides. The capsule
is more readily differentiated in old colonies.
Aerobic.
Source: Activated sludge.
Habitat: Unknown.
74 ORDER I. PSEUDOMONADALES
FAMILY II. METHANOMONADACEAE BREED Fam. Nov.*
(Oxydobacteriaceae Orla-Jensen, pro parte, Cent. f. Bakt., II Abt., 22, 1909, 329;
Protohacterieae Rahn, Cent. f. Bakt., II Abt., 96, 1937, 273.)
Me. tha.no. mo. na.da'ce.ae. M.L. noun Methanonwnas, -adis a genus of bacteria; -aceae
ending to denote a familj^; M.L. fern. pi. n. Methanomonadaceae the Methanonwnas family.
Rod -shaped organisms deriving their life energy from the oxidation of simple compounds
of hydrogen or carbon. Polar flagellate when motile. Gram-negative. Found in soil and
water.
It is clear that the species placed in the genera in this family belong with other polar
flagellate bacteria (the group of pseudomonads in the broad sense). Their method of de-
riving energy from oxidative processes is in accord with that of many other polar-fiagellate
bacteria. As a matter of convenience and as a means of emphasizing the fact that the species
included here secure their energy from the oxidation of simple hydrogen and carbon com-
pounds, the genera that have been proposed to include these species are grouped into a
family separate from those of the species that secure their energy from the oxidation of
simple nitrogen or sulfur compounds on the one hand, and those that normally secure their
energy from the oxidation of glucose or other organic compounds on the other hand. Further
studies of the differences in physiology found among the polar flagellate bacteria are badly
needed.
Key to the genera of the family Methanomonadaceae.
I. Organisms deriving their life energy from the oxidation of simple compounds of hy-
drogen.
A. Cells capable of securing growth energy by the oxidation of methane.
Genus I. Methanomonas, p. 74.
B. Cells capable of securing growth energy by the oxidation of hydrogen.
Genus II. Hydrogenomonas , p. 75.
II. Organisms deriving their life energy from the oxidation of carbon monoxide.
Genus III. Carhoxydomonas , p. 77.
Genus I. Methanomonas Orla-Jensen, 1909.
(Cent. f. Bakt., II Abt., 22, 1909, 311.)
Me.tha.no.mo'nas. Gr. methy wine; Gr. methe strong drink; M.L. methanum methane;
Gr. monas a unit, monad; M.L. fem.n. Methanomonas methane monad.
Cells monotrichous, capable of obtaining energj' from oxidation of methane to CO2
and water.
The type species is Methanomonas methanica (Sohngen) Orla-Jensen.
1. Methanomonas methanica (Sohn- motile in young cultures by means of a
gen, 1906) Orla-Jensen, 1909. {Bacillus single flagellum. In older cultures nearly
me/AamciiS Sohngen, Cent. f. Bakt., II Abt., spherical. Can be cultivated in an atmos-
15, 1906, 513; Orla-Jensen, Cent. f. Bakt., phere composed of one part CH4 and two
II Abt., ^^, 1909, 311.) parts air on washed agar containing the
me.tha'ni.ca. M.L. noun methanum necessary inorganic salts. The growth is
methane; M.L. adj. melhanicus relating to membranous,
methane. At the end of two weeks, the organisms
Short rods, 0.5 to 0.8 by 2.0 to 3.0 microns, changed an atmosphere containing 225 ml
* Revised })y Prof. Robert S. Breed, Cornell University, Geneva, New York, January,
1954.
FAMILY II. METHANOMONADACEAE 75
CH4 and 321 ml O2 to the following: In addition, 21 ml CO2 were dissolved in
CH4 ml the liquid.
CO2 78 ml Habitat : Presumably widely distributed
O2 172 ml in soil.
Genus II. Hydrogenomonas Orla-Jensen, 1909.
(Cent. f. Bakt., II Abt., 22, 1909, 311.)
Hj-.dro.ge.no.mo'nas. Gr. hydro water; Gr. genus race, offspring; whence, M.L. hy-
drogenum hydrogen, that which produces water; Gr. monas a unit, monad; M.L. fem. noun
Hydrogenomonas hydrogen monad.
Short rods that are polar flagellate when motile. Cells capable of deriving energy from the
oxidation of hydrogen. They may grow well on organic media without hydrogen although
this has not been shown to be true in all cases. Gram-negative. Found in soil and water.
This group of bacteria is characterized by the ability to grow in substrates containing no
organic matter and to use elemental hydrogen as the source of energj^ for growth. Under
these conditions CO2 is used as the source of carbon. Bacteria with similar phj'siological
characteristics but differing in morpholog}' are placed in the genera Bacillus and Clos-
tridium. Although other bacteria and even certain algae have enzyme systems which can
activate hydrogen and reduce CO2 in the process, there is no evidence that these organisms
are able to grow in inorganic media with hydrogen as the exclusive source of energj^ (see
Stephenson and Strickland, Biochem. Jour., 25, 1931, 205-215; Woods, Biochem. Jour., 30,
1936, 515; Lee and Umbreit, Zent. f. Bakt., II Abt., 101, 1940, 354; Gaffron, Amer. Jour.
Bot., 27, 1940, 273).
The tj'pe species is Hydrogenomonas pantotropha (Kaserer) Orla-Jensen.
Key to the species of genus Hydrogenomonas.
I. Not sensitive to high concentrations of O2 .
A. When growing autotrophically, no pellicle on liquid media.
1. Hydrogenomonas pantotropha.
B. When growing autotrophically, pellicle formed on liquid media.
2. Hydrogenomonas facilis.
II. Sensitive to high concentrations of O2 .
A. When growing autotrophically, no pellicle on liquid media.
3. Hydrogenomonas flava .
B. When growing autotrophically, pellicle formed on liquid media.
4. Hydrogenomonas vitrea.
1. Hydrogenomonas pantotropha (Ka- Inorganic solution: When cultivated
serer, 1906) Orla-Jensen, 1909. (Bacillvs under an atmosphere of O2 , CO2 and Ho ,
paniotrophus Kaserer, Cent. f. Bakt., II the liquid becomes turbid without pellicle
Abt., 16, 1906, 688; Orla-Jensen, Cent. f. formation.
Bakt., II Abt., .?^, 1909, 311.) Inorganic solid media: When cultivated
pan.to'troph.a. Gr. prefix panto all; Gr. ""^er an atmosphere of O2 , CO2 and H2 ,
, 7 r 1 TiTT J- I i I the colonies are yellow and slimy, and the
trophus feeder; M.L. adj. pantotrophus om- , , ^ , , ,- ,
agar plates have an odor resembling hot,
mvorous. x
soapy water.
Rods, 0.4 to 0.5 by 1.2 to 1.5 microns, with G^\^t\,, colonies: Yellow, smooth, rarely
rounded ends. Occur singly, in pairs and in concentrically ringed or greenish,
chains. Encapsulated. Actively motile by Gelatin stab: Growth only at surface. As
means of a single, long, polar flagellum. a rule no liquefaction.
Gram stain not recorded. Bipolar staining Agar colonies: Same as on gelatin, green-
in old cultures. ish, often slimy.
76
ORDER I. PSEUDOMONADALES
Broth: Turbid, somewhat slimy; pellicle
occasionally produced.
Milk: No coagulation. A yellow pellicle
forms. Medium becomes slimy and assumes
a dirty flesh color.
Potato: Moist, yellow, glistening.
Indole not produced.
Hydrogen sulfide not produced.
Nitrites not produced from nitrates.
Carbohydrates not utilized.
Aerobic.
Optimum temperature, between 28° and
30° C.
Facultatively autotrophic.
Distinctive characters: Develops auto-
trophically in inorganic medium under an
atmosphere of H2 , O2 and CO2 . Oxidizes
hydrogen to water and uses CO2 as the
source of carbon for growth.
Source: Isolated from soil near Vienna.
Habitat: Probably widely distributed in
soil.
2. Hydrogenomonas facilis Schatz and
Bovell, 1952.* (An undescribed Hydrogeno-
monas, Schatz, Proc. Soc. Amer. Bact.,
Baltimore Meeting, 1950, 124; Schatz and
Bovell, Jour. Bact., 63, 1952, 87.)
fa'ci.lis. L. Sid\.Jacilis ready, quick.
Rods 0.3 bj^ 2.0 microns in autotrophic
and 0.4 by 2.5 microns in heterotrophic cul-
tures. Occur singly, in pairs and in short
chains. Motile by means of one or two polar
flagella. Gram-negative.
Gelatin stab: Rapidly liquefied.
Agar colonies: Round, raised, glistening,
translucent, non-fluorescent and non-mu-
coid. No distinctive odor developed.
Autotrophic media: Cultures readily
maintained in media of this type.
Autotrophic gas uptake : The same overall
reaction is effected as that carried out by
certain anaerobically adapted green algae
(6H2 + 2O2 + CO2 -^ CH2O + 5H2O). In an
atmosphere of CO 2 and H2 , no CO2 fixation
accompanies the quantitative reduction of
nitrate to nitrite by molecular H2 ; nor is
there any change in concenti-ation of bicar-
bonate or in total gas pressure (Warburg
apparatus) when acetone, pyruvate ora-ke-
toglutarate are added (Schatz, Jour. Gen.
Microbiol., 6, 1952, 329).
Broth: Turbid with pellicle.
Milk: Slowly digested with alkalinization.
Potato: Abundant, spreading, non-pig-
mented growth.
Indole not produced.
Hydrogen sulfide not produced.
Acetylmethylcarbinol not produced.
Nitrites produced from nitrates.
Aerobic, obligate.
Non-hemolytic.
Optimum temperature, 28° C.
Source: Isolated from soil.
Habitat: Presumably widely distributed
in soil.
3. Hydrogenomonas flava Niklewski,
1910. (Jahrb. f. wissensch. Botanik, 48,
1910, 113; emend. Kluyver and Manten, An-
tonie V. Leeuwenhoek, 8, 1942, 71.)
fla'va. L. flavus yellow.
Rods 1.5 microns in length. Motile by
means of polar flagella. Gram-negative.
Agar colonies on inorganic medium in
presence of H2 ,02andC02 : Small, smooth,
yellow, shining, adhering to medium. De-
velop well below surface of medium, but
growth is paler.
Gelatin not liquefied.
Inorganic liquid medium in presence of
H2 , O2 and CO2 : No pellicle formation.
Good development when there is from 2 to 8
per cent oxygen in the gas. At higher O2
concentrations good growth occurs only in
association with H. vitrea or other bacteria.
Oxidizes hydrogen to water.
Microaerophilic, growing in an atmos-
phere of low oxygen tension, not exceeding
8 per cent.
Facultatively autotrophic.
Distinctive characters: Found singly on
slides whereas the rod-shaped cells of Hydro-
genomonas vitrea tend to cling together in
masses. Colonies on agar opaque, not trans-
parent.
Source: Isolated from mud, garden soil,
pasture land, vegetable mold and peat.
Habitat: Presumably widely distributed
in soil.
* Prepared by Prof. Albert Schatz, National Agricultural College, Farm School P.O.,
Bucks Co., Pennsylvania, December, 1953.
FAMILY II. METHANOMONADACEAE
77
4. Hydrogenomonas vitrea Niklewski,
1910. (Jahrb. f. wissensch. Botanik, 48, 1910,
113.)
vit're.a. L. vitreus of glass.
Rods 2.0 microns in length, cells adhering
to each other as by slime. Motility not ob-
served.
Agar colonies on inorganic medium in
presence of H2 , O2 and CO2 : Delicate,
transparent, with slight fluorescence and
yellow center. Surface folded. Do not de-
velop readily beneath the surface of me-
dium.
Agar streak on inorganic substrate : Same
as agar colonies e.xcept that growth is
spreading.
Inorganic liquid medium in presence of
H2 , O2 and CO2 : Pellicle, adherent to wall
of tube. Good development when there is
from 2 to 8 per cent oxygen in the gas. At
higher O2 concentrations good growth occurs
only in association with H. flava or other
bacteria.
Oxidizes hydrogen to water.
Microaerophilic, growing in an atmos-
phere of low oxygen tension, not exceeding
8 per cent.
Facultatively autotrophic.
Distinctive characters: Grows in sub-
strates containing no organic matter and
produces a pellicle.
Source: Isolated from mud, garden soil,
pasture land, vegetable mold and peat.
Habitat: Presumably widely distributed
in soil.
Gemis III. Carboxydonionas Orla-Jensen, 1909.
(Cent. f. Bakt., II Abt., 22, 1909, 311.)
Car.box.y.do.mo'nas. L. noun carbo charcoal, carbon; Gr. adj. oxys sharp; Gr. noun
monas a unit, monad; M.L. fem.n. Carboxydomonas the carbon-oxidizing monad.
Autotrophic, rod-shaped cells capable of securing growth energy by the oxidation of CO,
forming CO2 .
The tj^pe species is Carboxydomonas oligocarbophila (Beijerinck and van Delden) Orla-
Jensen.
1. Carboxydonionas oligocarbophila
(Beijerinck and van Delden, 1903) Orla-
Jensen, 1909. (Bacillus oligocarbophilns
Beijerinck and van Delden, Cent. f. Bakt.,
II Abt., 10, 1903, 33; Orla-Jensen, Cent. f.
Bakt., II Abt., 22, 1909, 311.)*
o.li.go.car.bo'phi.la. Gr. adj. nligus
little, scanty; L. noun carbo charcoal, car-
bon; Gr. adj. ■philus loving; M.L. adj. oligo-
carbophilus loving little carbon.
Rods very small, 0.5 by 1.0 micron, color-
less, united into irregular masses by a
slimy substance. Non-motile. There is little
cytoplasm within the slimy, cellulose-like
wall of the cells.
Growth occurs in culture fluids free from
organic matter and on washed agar con-
taining the necessarj^ inorganic salts.
Media containing carbonaceous materials :
No growth.
Liquid media: A thick, slimy film is pro-
duced.
CO is utilized as food and as such is oxi-
dized to CO 2 • In symbiosis with other bac-
teria, hj'drogen in water is o.xidized by the
catalytic reduction of CO2 to CO. The CO is
then metabolized, again forming CO2 (Ka-
serer. Cent. f. Bakt., II Abt., 16, 1906, 681).
Growth best in the dark.
Optimum temperature, 25° C.
* Kistner (Proc. Kon. Nederl. Akad. van Wetenschappen, Amsterdam, Series C, 56,
1953, 443) , in a paper received after the section covering Carboxydomonas was prepared , ques-
tions the data gathered by Beijerinck, Kaserer, Lantzsch and others. He concluded that
their reports were based on doubtful and imperfect ol)servations. Using a carefully con-
trolled technique, he was able to isolate an organism which oxidized CO to CO2 and which
had the characters of a pseudomonad (polar flagellate, straight rod). On further testing,
however, because it also oxidized H2 , he concluded that it belonged in the genus Hydro-
genomonas. Further studies on the species are promised.
78 ORDER I. PSEUDOMONADALES
Discussion : In spite of the fact that reaction would be helpful in clarifying this
several able bacteriologists have studied situation. If the species is to be accepted as
this species and the actinomycete that has a non-motile but related to polar-flagellate
similar physiology, several important points bacteria, it must be Gram-negative. If an
actinomycete, it would be Gram -positive.
Lantzsch reports the organism he studied
are left in doubt; the most important of
these is whether Beijerinck was right ii
thinking the actinomycete something dis , , . , ,. , ^ , , ^
^. . r \- T> 11 T h i,-i „>. (which was an actinomj^cete) to be Gram-
tinct from his Bacillus ohgocarbophilus, or ^ j /
whether Lantzsch (Cent. f. Bakt., II Abt., Positive.
57, 1922, 309) was right in thinking of them as Source: Isolated from garden soil.
but stages in the growth cycle of a single Habitat: Presumably widely distributed
species. Definite data in regard to the Gram in soil.
FAMILY III. THIOBACTERIACEAE JANKE, 1924.*
(Allgem. Tech. Mikrobiol., Dresden and Leipzig, I Teil, 1924, 68.)
Thi.o.bac.te.ri.a'ce.ae. M.L. neut.n. Thiobacterium type genus of the family; -aceae
ending to denote a family; M.L. fem.pl.n. Thiobacteriaceae the Thiobacterium family.
Coccoid, straight or curved rod-shaped bacteria. Oxidize sulfur compounds, usually de-
positing free sulfur granules within or without the cells. Never filamentous. Colorless sulfur
bacteria that are sometimes embedded in gelatinous pellicles or in gelatinous bladder-like
colonies. Polar flagellate when motile. Presumably Gram-negative. Found in places where
hydrogen sulfide occurs or may oxidize free sulfur, thiosulfates or related compounds.
While all of the species placed in this family have been described as colorless sulfur bac-
teria, they are still inadequately known and may not all deserve to be designated as sulfur
bacteria. It is hoped that placing them together in one family will cause comparative studies
to be made.
The type genus is Thiobacterium Janke.
Key to the genera of family Thiobacteriaceae.
I. Free sulfur granules deposited within or without the cells. Usually found in sulfurous
waters or soil.
A. Cells coccoid or straight rods.
1. Non-motile so far as known. ^
Genus I. Thiobacterium, p. 79.
2. Motile by means of polar flagella so far as known.
a. Cells rod-shaped, very large.
Genus II. Macromonas, p. 80.
aa. Cells round to ovoid, large.
Genus III. Thiovuhim., p. 81.
B. Cells large, curved rods, somewhat pointed.
Genus IV. Thiospira, p. 82.
II. Oxidize free sulfur, thiosulfates and related sulfur compounds to sulfates. Autotrophic
or facultatively autotrophic.
Genus V. Thiohacillus, p. 83.
* Revision of Thiobacteriaceae Janke prepared by Prof. Dr. Alexander Janke, Technische
Hochschule, Vienna, Austria, December, 1954, with the assistance of Prof. Robert S. Breed,
Cornell University, Geneva, New York.
FAMILY III. THIOBACTERIACEAE
79
Genus I. Thiobacterium Janke, 1924*
(Janke, Allgem. Tech. Mikrobiol., I Teil, 1924, 68; not Thiobacterium Issatchenko and
Salimowskaja, Zur Morphologie u. Physiol, der Thionsaurebakterien (Russian),
Izyiestia Gosud. Gidrobiol. Inst. (Memoirs State Hydrobiol. Inst.
Leningrad), No. 21, 1928, 61.)
Thi.o.bac.te'ri.um. Gr. noun thi iwi suUur; Gr. dim. noun bacterium a small rod; M.L.
neut.n. Thiobacterium small sulfur rod.
Rod-shaped, sulfur bacteria found in fresh or salt water or soil. Cells 1.0 micron or less
in diameter. Motility not observed. Sulfur granules sometimes found inside, sometimes
outside the cells. These cells may or may not be embedded in pellicles or in spherical,
bladder-like colonies.
The type species is Thiobacterium bovista Janke.
Key to the species in genus Thiobacterium.
I. Sulfur grains are found within the cells. Forms colonies in bladder-like masses which
resemble puff balls.
1. Thiobacterium bovista.
II. Sulfur grains are found outside of the cells.
A. Produces colonies on the surface of water containing proper nutrients. Sulfur crys-
tals are found among the cells.
2. Thiobacterium cristalliferum.
B. Produces a surface film in the form of a network on water. Sulfur globules are found
among the cells.
3. Thiobacterium retiformans.
1. Thiobacterium bovista (Molisch,
1912) Janke, 1924. {Bacterium bovista Mo-
lisch, Cent. f. Bakt., II Abt., 33, 1912, 59;
Janke, Allgem. Tech. Mikrobiol., I Teil,
1924, 68.)
bo.vis'ta. M.L. noun Bovista a genus of
puff balls; from German bovist puff ball;
M.L. fem.n. bovista puff ball.
Rod-shaped bacteria embedded in the
wall of bladder-like gelatinous colonies, the
interiors of which are filled with a clear
liquid. The cells are 0.6 to 1.5 by 2.0 to 5.0
microns, occurring by the thousands in
each colony. Each cell contains from one to
four sulfur granules. No motility observed.
The cells stain well with gentian violet while
the gelatinous matrix stains poorly, if at all.
The spherical colonies increase in number
by a kind of budding process that produces
smaller colonies. The colonies are white by
reflected light, black or bluish black by
transmitted light. Groups of these colonies
have the appearance of groups of puff balls
of variable sizes. They occur near the surface
of the water.
These organisms have not been cultivated
in pure culture.
Source: Found commonly in sulfurous
sea-water in the harbor at Trieste.
Habitat: Presumably widely distributed
in coastal waters containing hydrogen sul-
fide.
2. Thiobacterium cristalliferum (Gickl-
horn, 1920) Janke, 1924. (Bacterium cris'
talliferum Gicklhorn, Cent. f. Bakt., II Abt.,
50, 1920, 420; Janke, Allgem. Tech. Mikro-
biol., I Teil, 1924, 68.)
cris. tal.li'fe. rum. Gr. noun crystallus a
crystal; L. v.fero to bear; M.L. adj. cristal-
liferus crystal-bearing.
Straight to curved, rod-shaped bacteria.
0.3 to 0.5 by 1.0 to 2.4 microns. Deposit sul-
fur crystals outside of the cells. Non-motile.
Stain readily in gentian violet.
Colonies developed on the surface of water
* Prepared by Prof. Dr. Alexander Janke, Technische Hochschule, Vienna, Austria,
December, 1954.
80
ORDER I. PSEUDOMONADALES
containing potassium sulfide (K2S) which
was inoculated with a handful of garden soil
from Graz, Austria. At the end of three
weeks, numerous, snow-white colonies de-
veloped on the surface of the water.
Colonies which at first are of microscopic
size maj^ become 0.8 to 1.5 mm in diameter.
Sulfur crystals appear by transmitted light
as a black mass in the center of the smaller
colonies, but these crystals extend to the
margin in older colonies.
Habitat: Garden soil.
3. Thiobacteriuni retifornians (Gickl-
horn, 1920) Janke, 1924. (Bacterium retifor-
nians Gicklhorn, Cent. f. Bakt., II Abt., 50,
1920, 421; Janke, Allgem. Tech. Mikrobiol.,
I Teil., 1924, 68.)
re. ti. for 'mans. L. noun rete a net; L. v
formo to form; M.L. part. adj. retifonnans
net-forming.
Rod-shaped bacteria, 0.5 to 1.0 by 2.0 to
4.5 microns. Globular sulfur granules found
among the cells. Non-motile. Forms pelli-
cles and zoogloeal masses.
Developed in water containing potassium
sulfide (K2S) which was inoculated with the
decaying roots of nettle plants. This species
developed a delicate pellicle in the form of a
network on the surface of the water. It
also formed zoogloeal masses attached to
the wall of the culture flask.
Source: Soil containing decaying roots,
Graz, Austria.
Habitat: Presumably widely distributed.
Gemis II. Macromonas Utermohl and Koppe, 1923.*
(Utermohl and Koppe, Verhandl. Intern. Ver. f. Theoret. u. angew. Limnologie, 1923, 86;
Thiovibrio Janke, Allgem. Tech. Mikrobiol., I Teil, 1924, 68.)
Mac.ro.mo'nas. Gr. adj. macrus large; Gr. noun monas a unit, monad; M.L. fem.n. Macro-
monas a large monad.
Colorless, cylindrical to bean -shaped bacteria, actively motile by means of a single polar
flagellum. Cells large, 3.0 to 14.0 microns in diameter. Multiplication by constriction (fis-
sion). Chiefly characterized by the occurrence of calcium carbonate inclusions in the form
of large spherules. In their natural habitat they may also contain small sulfur globules.
Two species have been distinguished, primarily on the basis of cell size. Whether this is
sufficiently constant to serve as a specific character has not been definitely established.
From studies on the organisms in their natural habitat, which are still limited in scope and
extent, it appears at present that the two species should be maintained, at least provi-
sionally. It is possible, however, that further observations, especially with cultures under
different environmental conditions, will show the occurrence of intermediate types and of a
greater range of variation in size of pure cultures than has previously been reported.
The type species is Macromonas mobilis (Lauterborn) Utermohl and Koppe.
Key to the species of genus Macromonas.
I. Cells measure 12 microns or more in length and 8 microns or more in width.
1. Macromonas mobilis.
II. Cells measure less than 12 microns in length and 5 microns or less in width.
2. Macromonas bipunctata.
1. Macromonas mobilis (Lauterborn,
1915) Utermohl and Koppe, 1923. {Achro-
matiitm mobile Lauterborn, Verhandl.
Naturhist.-medizin. Vereins, Heidelberg,
N. F., IS, 1915, 413; Utermohl and Koppe,
Verhandl. Intern. Ver. f. theoret. u. angew.
Limnologie, 1923, 86 and Utermohl and
Koppe, Arch. f. Hydrobiol., Suppl. Bd. 5,
1925, 234.)
mo'bi.lis. L. adj. mobilis movable, motile.
* Prepared by Prof. Dr. Alexander Janke, Technische Hochschule, Vienna, Austi
December, 1954.
FAMILY III. THIOBACTERIACEAE
81
Colorless sulfur bacteria always occurring
singly; slightly curved, elongated ellipsoids
or cylinders with broad, hemispherical ends.
Width varies from 8 to 14 microns, length
from 12 to 30 microns; most common size, 9
by 20 microns. Multiplication by constric-
tion in the middle.
Cells actively motile by means of a single
polar flagellum distinctly visible without
special staining. It is 20 to 40 microns long,
and, with respect to the direction of mo-
tion, always posteriorly placed. Rate of
movement somewhat sluggish, about 800
microns per minute, probably on account
of the high specific gravity of the cells.
Normally contain small sulfur droplets
and, in addition, large, roughly spherical
inclusions of calcium carbonate. Two to
four such crystal masses almost fill a single
cell. Under unfavorable conditions the
calcium carbonate crystals may disappear
before the sulfur globules.
Microaerophilic; apparently require hy-
drogen sulfide.
Habitat: Found in fresh-water environ-
ments containing sulfide and calcium ions,
as in shallow basins and streams in the upper
layers of the mud.
2. Macromonas bipunctata (Gicklhorn,
1920) Utermohl and Koppe, 1925. (Pseudo-
nonas bipunctata Gicklhorn, Cent. f. Bakt.,
[I Abt., 50, 1920, 425; Utermohl and Koppe,
A.rch. f. Hydrobiol., Suppl. Bd. 5, 1925, 235.)
bi.punc.ta'ta. L. bis twice; L. part. adj.
punctatus punctate, dotted; M.L. adj. bi-
punctatiis twice punctate.
Cells colorless, occurring singly; cylindri-
cal with hemispherical ends, after cell divi-
sion often temporarily pear-shaped. 3 to 5
by 8 to 12 microns. Multiplication by con-
striction in the middle.
Actively motile by means of a single polar
flagellum, about 10 to 15 microns long, al-
ways posteriorlj^ placed with respect to the
direction of movement. Flagellum delicate,
not visible without staining. Rate of move-
ment sluggish, about 600 microns per min-
ute. Probably this slow motion is due to the
high specific gravity of the cells.
Normally contain calcium carbonate
crystals as inclusions. These are in the form
of large spherules, one or two of which
nearly fill the individual cells. Sulfur glo-
bules have not been demonstrated with
certainty as yet.
Microaerophilic, but it is uncertain
whether hj^drogen sulfide is required.
A second species that is like Macromonas
bipunctata, except that the cells are smaller
in size, has been named by Gicklhorn (op.
cit., 50, 1920, 425). Pure-culture studies may
show the two species to be identical as dif-
ference in size of cells has not been found to
be significant elsewhere among sulfur bac-
tei'ia.
Source: From stems, leaves, etc. of fresh-
water plants in ponds near Graz, Austria.
Habitat: Found in fresh-water environ-
ments containing calcium ions; but it has
been found in sulfide-containing as well as
in sulfide-free water. Also found in shallow
basins and streams in upper layers of the
mud.
Genus III. Thiovulum Hinze, 1913.*
(Ber. d. deutsch. bot. Ges., 31, 1913, 195.)
Thi.o'vu.lum. Gr. noun thium sulfur; L. noun ovum egg; M.L. neut.dim.n. Thiovulum
small sulfur egg.
Unicellular organisms, round to ovoid, 5.0 to 20.0 microns in diameter. Cytoplasm often
concentrated at one end of the cell, the remaining space being occupied by a large vacuole.
Multiplication by constriction which, in late stages, merges into fission. Actively motile;
movements accompanied by rapid rotation. Flagellation not definitely demonstrated, but
type of locomotion suggests polar flagellation. Normally contain sulfur globules in the cj^to-
plasm; hence, these are frequently concentrated at one end of the cell.
It is difficult to establish distinct species. Those that have been described differ only in
* Prepared by Prof. Dr. Alexander Janke, Technische Hochschule, Vienna, Austria, De-
cember, 1954.
82
ORDER I. PSEUDOMONADALES
size, and the differences appear to be far from constant. The ovoid cells of Thiovulum majiis
are noted as being 11 to 18 microns long and 9 to 17 microns wide, while Thiovulum minus
comprises the smaller forms from 9.6 to 11.0 microns long by 7.2 to 9.0 microns wide. In
view of the regular occurrence of all intermediate sizes, it seems best to recognize only a
single species at present.
The type species is Thiovulum majus Hinze.
1. Thiovulum majus Hinze, 1913.
(Hinze, Ber. d. deutsch. bot. Ges., 31, 1913,
195; including Thiovulum minus Hinze,
loc. cit.; Thiovulum muUeri Lauterborn,
Verhandl. Naturhist.-medizin. Vereins,
Heidelberg, N. F., 13, 1915, 414.)
ma 'jus. L. comp.adj. major larger.
Unicellular organisms, spherical to ovoid.
Cytoplasm often concentrated at one end
of the cell, the remainder being occupied by
a vacuole. Multiplication by constriction
which, in late stages, merges into fission.
Size of cells, 5 to 20 microns in diameter.
The most characteristic feature is its mo-
tility; it is the only one of the spherical to
ovoid, colorless sulfur bacteria capable of
rapid movement. Flagellation has not been
definitely demonstrated, but the type of
locomotion suggests the presence of polar
flagella.
Normally contains sulfur droplets in cyto-
plasm, frequently concentrated at one end
of cell.
Microaerophilic ; apparently' requires hj^-
drogen sulfide.
Habitat: Found in sulfide-containing
water, usually accumulating near the sur-
face. Often found in cultures of decaying
algae and in both fresh-water and marine
environments.
Genus IV. Thiospira Vislouch, 1914*
(Vislouch, Jour, de Microbiologic, 1, 1914, 50; Sulfospirillum Kluyver and van Niel, Zent.
f. Bakt., II Abt., 94, 1936, 396; Thiospirillum Janke, Allgem. Tech. Mikrobiol., I Teil,
1924, 68; not Thiospirillum Winogradsky, Schwefelbakterien, Leipzig, 1888, 104.)
Thi.o.spi'ra. Gr. noun thium sulfur; Gr. noun spira a coil; M.L. fem.n. Thiospira sulfur
coil or spiral.
Colorless, motile, slightly bent, large rods, somewhat pointed at the ends, with granules
of sulfur within the cells and a small number of flagella at the ends.
The type species is Thiospira winogradskyi (Omelianski) Vislouch.
Key to the species of genus Thiospira.
I. Large spirilla containing numerous sulfur granules.
1. Thiospira winogradskyi.
II. Clear center of spirilla cells contains two, occasionally one or three, sulfur granules.
2. Thiospira bipunctata.
1. Thiospira winogradskyi (Omelian-
ski, 1905) Vislouch, 1914. (Thiospirillum
winogradskyi Omelianski, Cent. f. Bakt., II
Abt., 14, 1905, 769; Vislouch, Jour, de Mi-
crobiologic (Russian), 1, 1914, 50.)
wi.no.grad'sky.i. M.L. gen. noun wino-
gradskyi of Winogradsky; named for S. N.
Winogradsky, a Russian bacteriologist.
Large sulfur spirilla, somewhat pointed at
the ends, 2.0 to 2.5 by 50 microns. Numerous
granules of sulfur. Very motile, with one to
two polar flagella.
The large, very active sulfur spirillum
found by Gicklhorn (Cent. f. Bakt., II Abt.,
50, 1920, 418) may have belonged to this
species.
Habitat: Curative mud.
2. Thiospira bipunctata (Molisch, 1912)
Vislouch. 1914. (Svirillum bivunctatum
. Thiospira bipunctata (Molisch, 1912)
louch, 1914. {Spirillum bipunctatum
* Prepared by Prof . Dr. Alexander Janke, Technische Hochschule, Vienna, Austria, De-
cember, 1954.
FAMILY III. THIOBACTERIACEAE 83
Molisch, Cent. f. Bakt., II Abt., 33, 1912, of the cell). Both ends are more or less filled
55; Vislouch, Jour, de Microbiologie (Riis- ^vith large volutin (metachromatic) gran-
sian),/, 1914,50.)
bi.punc.ta'ta. L. bis twice; L. noun
punctum a point, spot; M.L. adj. bipunc-
tatus two-spotted
ules. Several minute granules of sulfur are
present in the clear center and sometimes at
the ends. Old cells possess one flagellum at
Small, slightly bent sulfur spirilla, mark- each end; young cells have a flagellum at one
edly pointed at the ends; 6.6 to 14 microns end.
long, 1.7 to 2.4 microns wide (in the center Habitat: Sea and salt waters.
Genus V. Thiobacillus Beijerinck, 1904*
(Beijerinck, Cent. f. Bakt., II Abt., 11, 1904, 593; not Thiobacillus Ellis, Sulphur Bacteria,
London, 1932, 130; Sulfomonas Orla-Jensen, Cent. f. Bakt., II Abt., 22, 1909, 314.)
Thi.o.ba.cil'lus. Gr. noun^ium sulfur; L. noun bacillus a small rod; M.L. mas.n. Thio-
bacillus a sulfur rodlet.
Small, Gram-negative, rod-shaped cells. Non-motile or motile, usually by means of a
single polar flagellum. Energy derived from the oxidation of incompletely oxidized sulfur
compounds, principally from elemental sulfur and thiosulfate but in some cases also from
sulfide, sulfite and polythionates. The principal product of oxidation is sulfate, but sulfur
is sometimes formed. Grow under acid or alkaline conditions and derive carbon from carbon
dioxide or from bicarbonates in solution; some are obligate and some facultatively auto-
trophic. Some species are anaerobic in the presence of nitrate. Found in soil, mine waste-
waters, sewage, effluents and related sources.
The type species of this genus is strictly autotrophic as are the majority of the species in
the genus. It has been suggested that Thiobacillus should be restricted to these autotrophic
species and that the facultatively autotrophic species be placed in the genus Psexidomonas.
Some heterotrophic species now placed in Pseudomonas are known to have the ability to oxi-
dize thiosulfates (Starkey, Soil Sci., 89, 1935, 325).
The type species is Thiobacillus thioparus Beijerinck.
Key to the species of genus Thiobacillus.
I. Thiosulfate oxidized with increa.sed acidity.
A. Tetrathionate not formed as an intermediate product.
1. Strictly autotrophic.
a. Does not oxidize ferrous salts.
1. Thiobacilhis thioparus.
aa. Oxidizes ferrous salts.
2. Thiobacillus ferrooxidans .
2. Facultatively autotrophic.
a. Aerobic.
b. Does not oxidize free sulfur.
3. Thiobacillus novellus.
bb. Oxidizes free sulfur to sulfate.
4. Thiobacillus coproliticus .
aa. Facultatively anaerobic in presence of nitrate.
5. Thiobacillus denitrificans .
B. Tetrathionate formed as intermediate product.
1. Final pH, 3.0.
6. Thiobacillus neopoUtanus.
* Revised by Dr. C. D. Parker, South Melbourne, Australia, with the assistance of Dr.
Kenneth L. Temple, Morgantown, West Virginia, June, 1954.
84
ORDER I. PSEUDOMONADALES
2. Final pH, 1.0 or less,
a. Nitrate utilized.
7. Thiohacilhis concretivonis.
aa. Nitrate not utilized.
8. Thiobacillus thiooxidans .
II. Thiosulfate oxidized with increased alkalinity.
9. Thiobacillus trautweinii.
1. Thiobacillus thioparus Beijerinck,
1904. (Arch. d. Sci. Exact, et Nat. Haar-
lem, Ser. 2, 9, 1904, 153; also see Cent. f.
Bakt., II Abt., 11, 1904, 593.)
thi.o'par.us. Gr. noun thium sulfur; L.v.
paro to produce; M.L. adj. thioparus sulfur-
producing.
Thin, short rods, 0.5 by 1.0 to 3.0 microns,
averaging 0.5 by 1.7 microns. Motile.
Starkey (Soil Sci., 39, 1935, 209) reports the
isolation of cultures (C) that he regards as
practically identical with this species
though they were non-motile and of coccoid
form. Gram-negative.
Thiosulfate liquid medium: Pellicle con-
sists of cells and free sulfur. Medium be-
comes turbid. pH drops to 4.5.
Thiosulfate agar colonies: Small (1 to 2
mm in diameter) circular, whitish yellow due
to precipitated sulfur. Turn brown in old
cultures.
No growth on organic media.
Optimum reaction, close to neutrality.
Growth occurs between pH 7.8 and 4.5.
Strictly autotrophic. Derives its energy
by the oxidation of thiosulfate to sulfate
and sulfur without the intermediate forma-
tion of tetrathionate. Also oxidizes ele-
mental sulfur. Does not oxidize hydrogen
sulfide or sulfides.
Utilizes nitrate and ammonium salts as
sources of nitrogen.
Aerobic.
Source: Canal water, mud and soil.
Habitat: Presumably widely distributed.
2. Thiobacillus ferrooxidans Temple
and Colmer, 1951. (Iron oxidizing bacter-
ium, Colmer, Temple and Hinkle, Jour.
Bact., 59, 1950, 317; Temple and Colmer,
Jour. Bact.,6^, 1951,605.)
fer.ro.o'xi.dans. L. noun Jerrum iron;
Gr. adj. oxys sharp, acid; M.L. v. oxido to
oxidize or make acid; M.L. part. adj. Jer-
rooxidans iron-oxidizing.
Description prepared by Dr. Kenneth L.
Temple, Morgantown, West Virginia.
Short rods, 0.5 by 1.0 micron, with rounded
ends. Occur singly or in pairs, rarely in
chains. Motile, presumably polar flagel-
late. Gram -negative.
Thiosulfate agar colonies: Very thin and
small with irregular margins, becoming
whitish in center upon aging.
Thiosulfate liquid medium: Uniform tur-
bidity; delicate pellicle in two or three
weeks.
Ferrous agar: Colonial appearance varies
with ferrous-iron content of agar: on low to
moderate iron concentration, an amber zone
reveals the presence of microscopic colonies
which become lobed and coated with hy-
drated ferric oxide; on high ferrous iron
concentration, growth is abundant becom-
ing heavily encrusted with hydrated ferric
oxide.
Ferrous liquid medium: Clear, rapidly
turning amber to reddish brown due to pro-
duction of ferric iron; ferric hydrate pre-
cipitated. Pellicle composed of cells and
ferric hydrate.
Nitrogen sources: Utilizes ammonia; ni-
trate to a lesser extent.
Aerobic.
Optimum pH, between 2.5 and 5.8. No
growth above pH 6.0. There is some step-
wise adaptation to a lower pH than 2.5.
Strictly autotrophic, deriving its energy
from the oxidation of thiosulfates or inor-
ganic ferrous iron. Sulfur not appreciably
utilized.
Distinctive characters: The pH range ap-
proaches that of Thiobacillus thiooxidans but
does not extend below pH 2.0, and elemental
sulfur is not appreciably used. Thiosulfate
is oxidized rapidly but both liquid and agar
FAMILY III. THIOBACTERIACEAE
85
cultures differ in appearance from Thio-
bacillus thioparus. Ferrous iron serves as a
sufficient energy source with the concomi-
tant formation of enormous quantities of
ferric ions in acid media where ferric iron is
not otherwise produced in quantity. The
lobed, iron-encrusted colony formed on
ferrous agar is unique. Cultures maintained
on ferrous media lose the ability to oxidize
thiosulfate, but colonies maintained on
thiosulfate media retain their iron-oxidiz-
ing capacity.
Source: Isolated from bituminous coal
mine drainage waters which were strongly
acid and high in ferrous iron. Found in
West Virginia and Pennsylvania.
Habitat : Acid waters of high iron content
including drainage from several types of
mines and soils containing pyrite or marca-
site.
3. Thiobacillus novellus Starkey,
1934. (Jour. Bact., 28, 1934, 365; Jour. Gen.
Physiol., 18, 1935, 325; Soil Sci., 39, 1935,
207, 210.)
no.vel'lus. L. dim. adj. novellus new.
Short rods or ellipsoidal cells 0.4 to 0.8
by 0.6 to 1.8 microns. Non-motile. Gram-
negative.
Gelatin stab: Mucoid growth at point of
inoculation. Sub-surface growth meager,
slow liquefaction.
Agar plate : Growth slow, colorless, moist,
raised, circular, 1 mm in diameter. Deep
colonies tiny, lens-shaped.
Thiosulfate agar plate: Growth slow, be-
coming white from precipitated sulfur.
Surface colonies small, circular, moist.
Crystals of CaS04 appear throughout the
agar.
Agar slant: Growth fairly abundant, soft,
somewhat ropy, raised, shining, moderately
spreading; whitish in reflected light, brown-
ish opalescence in transmitted light.
Thiosulfate agar slant: Growth very thin,
practically colorless. No sub-surface
growth. Sulfur usually precipitated as
white, frosty film on the surface.
Agar stab : White to cream-colored growth
confined close to point of inoculation; pene-
trates to bottom of tube.
Thiosulfate agar stab: No appreciable
surface growth.
Broth: Slightly turbid. Gelatinous pel-
licle. Forms long, streamer-like network
extending from surface to the bottom. Some
sediment.
Thiosulfate broth: Uniform turbidity. No
pellicle. Whitish sediment with thin, incom-
plete membrane on the bottom of the flask.
Reaction acid in a few days, changes pH 7.8
to 5.8 with decomposition of a small quantity
of thiosulfate.
Sulfur solution medium of slightly alka-
line reaction: No growth.
Potato slant: Growth limited, cream-
colored, moist, shining, slightly brown.
Litmus milk: Slow development of slight
alkalinity.
Facultatively autotrophic.
Optimum reaction: Close to neutrality
(limiting reactions, pH 5.0 to 9.0).
Aerobic.
Distinctive characters: Oxidizes thiosul-
fate to sulfate and sulfuric acid. Does not
oxidize free sulfur.
Source: Isolated from soils.
Habitat: Soils.
4. Thiobacillus coproliticus Lipman
and McLees, 1940. (Soil Sci., 50, 1940, 432.)
co.pro.Iit'i.cus. Gr. noun coprus dung;
Gr. noun lithus a stone; whence coprolite, a
fossil dung; M.L. adj . coproliticus of a copro-
lite.
Long, thin rods 0.1 to 0.2 by 6 to 8 (may
measure 3 to 40) microns. Straight, S-shaped
and curved cells. Motile by means of a sin-
gle polar flagellum.
Peptone soil extract agar: Slight growth.
Nutrient broth: Little or no growth.
Thiosulfate agar: Slow development. Pro-
duces small, watery colonies raised above
the agar surface. Colonies have been noted
which were white from precipitated sulfur.
Thiosulfate broth: Thiosulfate is oxi-
dized. Little or no turbidity. No pellicle.
No sediment. Change in reaction from pH
7.6 to 6.1.
Sulfur broth: Sulfur is oxidized. No tur-
bidity.
Facultatively autotrophic.
Aerobic.
86
ORDER I. PSEUDOMONADALES
Distinctive characters: Develops in inor-
ganic media and oxidizes thiosulfate and
sulfur to sulfate. Media with slightly alka-
line reactions most favorable for growth.
Source: Isolated from coprolite rock ma-
terial from Triassic period (Arizona).
Habitat: Unknown.
5. Thiobacillus denitrificans Beijer-
inck, 1904. (Beijerinck, Cent. f. Bakt., II
Abt., 11, 1904, 597; Sulfomonas denitrificans
Orla-Jensen, Cent. f. Bakt., II Abt., 22,
1909, 314.)
de.ni.tri'fi.cans. L. pref. de off, removed;
M.L. noun niter saltpeter, nitrate; M.L. v.
nitrifico to make nitrate, to nitrify; M.L.
denitrifico to denitrify; M.L. part. adj. de-
nitrificans denitrifying.
Short rods, 0.5 by 1 to 3 microns. Motile
by means of a single polar flagellum (Tjulpa-
nova-Mossevitch, Arch. d. Sci. Biol.,
U.S.S.R.,SO, 1930, 203).
Inorganic broth: Growth with production
of gas, predominantly nitrogen.
Thiosulfate agar colonies: Thin; clear or
weakly opalescent.
Optimum reaction: Neutral or slightly
alkaline.
Autotrophic, utilizing carbon from COo ,
carbonates and bicarbonates. Considered to
be strictly autotrophic by Lieske (Ber. d.
deutsch. botan. Gesell., 30, 1912, 12) and
facultatively by Tjulpanova-Mossevitch (op.
cit., 30, 1930, 203). Beijerinck stated (Kon.
Akad. V. Wetenschappen Amsterdam, 42,
1920, 899) that whereas the organism de-
veloped initially in an inorganic medium, it
lost the autotrophic habit bj^ cultivation in
an organic medium.
Facultatively anaerobic. Can live in the
absence of free O2 in the presence of nitrate.
Distinctive characters: Oxidizes thiosul-
fate to sulfate under anaerobic conditions
using nitrate as the hydrogen acceptor which
is reduced to N2 . Also oxidizes sulfide, ele-
mental sulfur and dithionate.
Habitat: Canal and river water, salt
water, soil, peat, composts and mud.
6. Thiobacillus neapolitanus Parker,
nom. nov. (Neue Gruppe von Schwefelbak-
terium, Nathansohn, Mitt. Zool. Sta.,
Neapel, 15, 1902, 655; Thiobacillus X,
Parker, Jour. Gen. Microbiol., 8, 1953, 344.)
ne.a.po.li.ta'nus. L. adj. neapolitanus
pertaining to Naples.
Short rods, 0.5 by 1.0 to 1.5 microns. Non-
motile. Gram-negative.
Thiosulfate agar colonies: Small (1 to 2
mm in diam.) circular, convex; whitish yel-
low due to precipitated sulfur.
Thiosulfate liquid medium; Uniform tur-
bidity with pellicle which contains free
sulfur. pH drops to 3.0.
Sulfur liquid medium: Very slight, uni-
form turbidity.
Optimum pH, about 6.0; growth occurs
between pH 8.5 to 3.0.
Temperature relations: Optimum, 28° C;
slow growth at 10° and 37° C; death occurs
at 55° C.
Strictly autotrophic. Derives energy by
the oxidation of thiosulfate, tetrathionate,
elemental sulfur and hydrogen sulfide.
Thiosulfate is oxidized to tetrathionate,
sulfate and sulfuric acid; tetrathionate is
oxidized to sulfate and sulfuric acid while
free sulfuric acid only is formed from ele-
mental sulfur and hydrogen sulfide. Utilizes
atmospheric CO 2 as a source of carbon.
• Nitrogen sources: Ammonium; nitrate-
and nitrite-nitrogen.
Aerobic.
Comments: It has not been clear until re-
cently (Parker, Jour. Gen. Microbiol., 8,
1953, 344) that this organism is a species
separate from Thiobacillus thioparus Beijer-
inck. The isolation and detailed study of an
organism from concrete identical in most
respects with Nathansohn 's description of
his isolate but different from Beijerinck's
and Starkey's description of Thiobacillus
thioparus makes it clear that two separate
species are involved. Thiobacillus neapoli-
tanus produces tetrathionate and sulfate
from thiosulfate and oxidizes HoS and tetra-
thionate whereas Thiobacillus thioparus
produces sulfur and sulfate from thiosulfate
and does not oxidize H2S or tetrathionate.
Source: Originally isolated by Nathan-
sohn from sea water at Naples, Italy. Iso-
lated by Parker from early stages of the cor-
rosion of concrete sewers and other concrete
structures.
Habitat: Presumably widely distributed
in soil and water, including sea water.
7. Thiobacillus concretivorus Parker,
FAMILY III. THIOBACTERIACEAE
87
1945. (Austral. Jour. Exper. Biol, and Med.
Sci., S3, 1945, 81; also see Jour. Gen. Micro-
biol., 8, 1953, 344.)
con.cre.ti'vo.rus. L. noun concretum firm
or solid matter; L. v. voro to devour or de-
stroy; M.L. part. adj. co/ic?'e<iwrM.s concrete-
destroying.
Short, straight rods 0.5 by 1.5 to 2.0
microns with square ends. Stain irregularly,
showing deeply stained granules in poorly
stained slender rods. Motile, presumably
polar flagellate. Motility lost in older cul-
tures. Single polar flagellum, two to three
times the length of the organism (unpub-
lished data). Gram-negative.
Thiosulfate agar colonies: Minute, water-
clear, whitish yellow on prolonged incuba-
tion. No confluent growth.
Thiosulfate liquid medium: Uniform tur-
bidit\-, slight deposit of sulfur. No pellicle.
Sulfur liquid medium: Uniform turbidity;
floating sulfur granules fall to the bottom.
Strictly aerobic.
Temperature relations: Optimum, 28° C.;
slow growth at 10° and 37° C.; death occurs
at 55° C.
Strictly autotrophic, utilizing atmos-
pheric CO2 as the source of carbon; growth
inhibited by higher concentrations of glu-
cose, glycerol and lactate. Derives energy
from its oxidation of elemental .sulfur, thio-
sulfate and hydrogen sulfide, o.xidizing
them ultimately to sulfate and sulfuric acid.
Thiosulfate is oxidized with the intermedi-
ate production of tetrathionate.
Nitrogen sources: Utilizes ammonium-
and nitrate-nitrogen equally well. Nitrate
is not toxic whereas nitrite is.
Optimum reaction: Growth occurs be-
tween pH 6.0 and acid concentrations up to
10 per cent. Sulfuric acid optimum is be-
tween pH 2.0 and 4.0.
Distinctive characters: Responsible for
the rapid corrosion of concrete sewers and
other concrete structures where the sewer
air contains hydrogen sulfide.
Habitat: Corroding concrete in sewers;
also found in sewage and presumably in soil
and fresh water.
8. Thiobacillus thiooxidans Waksman
and Joffe, 1922. (Jour. Bact., 7, 1922, 239.)
thi.o.ox'i.dans. Gr. noun thiuvi sulfur;
Gr. adj. oxys sharp, acid; M.L. v. oxido to
make acid, to o.xidize; M.L. part. adj. thio-
oxidans oxidizing sulfur.
Short rods: 0.5 by 1.0 micron with rounded
ends. Occur singly, in pairs or in chains.
Motile by means of a single polar flagellum.
Gram-negative (Starkey, Soil Sci., 39, 1935,
210).
Thiosulfate agar: Scant growth. Nearly
transparent colonies.
Sulfur broth: Uniform turbidity. No sedi-
ment or surface growth. Medium becomes
very acid (below pH 1.0).
Thiosulfate broth: Uniform turbidity.
Medium becomes acid, and sulfur is precipi-
tated.
Nitrogen sources: Utilizes ammonia-
nitrogen but not nitrate-nitrogen, which is
toxic. Asparagin, urea and peptone not
utilized.
Temperature relations: Optimum, be-
tween 28° and 30° C. Slow growth at 18°
and 37° C. Death occurs between 55° and
60° C.
Optimum reaction, between pH 2.0 and
3.5. (Limiting reactions, pH 6.0 to less than
pH 0.5.)
Strictly autotrophic, deriving its energy
from the oxidation of elemental sulfur and
thiosulfate, o.xidizing these to sulfuric acid.
It utilizes the CO 2 of the atmosphere as a
source of carbon.
Strictly aerobic.
Distinctive characters: This species pro-
duces more acid, from oxidation of sulfur,
and continues to live in a more acid medium,
than any other living organism yet reported,
the hydrogen-ion concentration of the me-
dium increasing to a pH 0.6 and less.
Source: Isolated from composts of soil,
sulfur and rock phosphate, and soils con-
taining incompletel}' oxidized sulfur com-
pounds.
Habitat: Soil.
9. Thiobacillus trautweinii Bergey et
al., 1925. (Thionsaurebakterium, Trautwein,
Cent. f. Bakt., II Abt., 53, 1921, 513; also
see ibid., 61, 1924, 1; Bergey et al.. Manual,
2nd ed., 1925, 39; Bacterium thiogenes
Lehmann, in Lehmann and Neumann,
Bakt. Diag., 7 Aufl., 2, 1927, 516.)
traut.wein'i.i. M.L. gen. noun trautweinii
of Trautwein; named for K. Trautwein, who
first isolated and studied this species.
ORDER I. PSEUDOMONADALES
Short rods, 0.5 by 1.0 to 2.0 microns. Mo-
tile by means of six to eight long flagella.
Gram-negative.
Gelatin stab: Slow liquefaction. No
chromogenesis.
Thiosulfate agar: Colonies small, white,
1 mm in diameter.
Thiosulfate liquid medium: Verj' little
visible turbiditjs no sulfur precipitated.
Produces sulfate and tetrathionate with in-
crease in pH. Rate of thiosulfate oxidation
increased by presence of organic com-
pounds.
No acid or gas from sugars.
Nitrites and gas produced from nitrate-
peptone broth; no ammonia produced. May
live anaerobically in the presence of ni-
trates.
Indole not produced.
Hydrogen sulfide not produced.
Starch is hydrolyzed.
Lipolytic.
Catalase-positive.
Non-hemolytic.
Temperature relations: Optimum, 27° C.
Minimum, 6.9° C. Maximum, l^etween 36.5°
and 40° C. Death occurs in 2 to 5 minutes at
55° C.
Optimum pH, between 7.9 and 8.5. pH
limits, 6.0 and 10.0.
Comments: Trautwein (Cent. f. Bakt., II
Abt., 61, 1924, 5) regards his bacterium as
being closely related to the fluorescent
group and to the denitrifying bacteria of
Burri and Stutzer. Starkey (Jour. Gen.
Physiol., 18, 1935, 346) reports this species
to be heterotrophic. However, Lehmann (in
Lehmann and Neumann, Bakt. Diag., 7
Aufl., ^, 1927, 516), under whom Trautwein
did his work, reports that this species is a
facultative autotroph as does Parker also
(Jour. Gen. Microbiol., 3, 1953, 344). As
facultatively autotrophic species are in-
cluded in Thiobacillus as defined, this
species has again been included in Thio-
bacillus (see Manual, 2nd ed., 1925, 39).
Starkey 's culture B and Parker's M cul-
ures appear to be identical with Thioba-
cillus trautweinii.
Source: Isolated from soil and water
(Trautwein) and from purified sewage from
Wiirzburg (Lehmann).
Habitat: Widely distributed in polluted
waters and soil.
FAMILY IV. PSEUDOMONADACEAE WINSLOW ET AL., 1917.
(Winslow, Broadhurst, Buchanan, Krumwiede, Rogers and Smith, Jour. Bact., 2, 1917, 555.)
Pseu.do.mo.na.da'ce.ae. M.L. fem.n. Pseudomonas tj'pe genus of the family; -aceae
ending to denote a family; M.L. fern. pi. n. Pseudomonadaceae the Pseudomonas family.
Cells elongate, straight rods, occasionally coccoid. Motile by means of polar flagella
which are either single or in small or large tufts. A few species are non-motile. Gram-nega-
tive. May possess either water-soluble pigments that diffuse through the medium or non-
water-soluble pigments. Usually grow well and fairly rapidly on the surface of culture
media. Aerobic. Frequently oxidative in their physiology but may be fermentative. Usually
found in soil or water, including sea water or even heavy brines. Many plant and a few
animal pathogens.
Key to the genera of family Pseudomonadaceae.
I. Attack glucose and other sugars either oxidatively or fermentatively.
A. Genera in which the species are either known or are thought to attack glucose oxi-
datively.
1. Bacteria which do not produce readily detectable acetic acid though they may
oxidize ethanol. May produce a water-soluble pigment which diffuses through
the medium,
a. Cultures may or maj' not produce a water-soluble pigment which is bluish,
greenish or brownish in color. Rose, lilac- and yellow-colored, diffusible pig-
ments occasionally occur.
FAMILY IV. PSEUDOMONADACEAE »«
Genus I. Pseudomonas, p. 89.
aa. Cultures develop a yellow, non-water-soluble pigment. Cells normally mono-
trichous. Mostlj^ plant pathogens which cause a necrosis.
Genus II. Xanthomonas, p. 152.
2. Bacteria which produce readily detectable amounts of acetic acid bj- the oxida-
tion of ethanol. The vinegar bacteria.
Genus III. Acetobacter, p. 183.
B. Genera in which the species ferment glucose, usually with the production of H2 and
CO2.
1. Cells carry out a fermentation like that of the coliform bacteria. Usually produce
acid and gas from glucose.
a. Cells not known to fix free atmospheric nitrogen.
b. Water organisms. Common species cause diseases of fishes. Also found in
leeches. Not luminescent.
Genus IV. Aeromo?ias, p. 189.
bb. Luminescent bacteria commonly found on dead fishes and Crustacea on
salt-water beaches.
Genus V. Photobacterium , p. 193.
aa. Cells fix free atmospheric nitrogen.
Genus VI. Azotomonas, p. 198.
2. Cells carry out an alcoholic fermentation similar to that of yeasts.
Genus VII. Zymomonas, p. 199.
II. Do not attack carbohydrates or, if so, produce only slight amounts of acid from glu-
cose and similar sugars. Includes certain species which require at least 12 per cent salt
for growth.
A. Do not require salt in excess of 12 per cent for growth.
1. Cells not embedded in a gelatinous matrix.
a. Cells rod-shaped.
b. Soil and water bacteria that are known to dissimilate alkylamines.
Genus VIII. Protaminobacter , p. 200.
bb. Soil and water bacteria that are known to dissimilate alginic acid.
Genus IX. Alginovwnas , p. 202.
aa. Soil bacteria that are known to utilize phenol and similar aromatic com-
pounds. Cells may be branched.
Genus X. Mycoplana, p. 204.
2. Cells embedded in a gelatinous matrix; this matrix may be of a branching form.
Genus XI. Zoogloea, p. 206.
B. Requires at least 12 per cent salt before growth will take place.
Genus XII. Halobacterium, p. 207.
Genus I. Pseudomonas Migula, 1894*
{Chlorobacterium Guillebeauf, Landw. Jahrb. d. Schweiz, 4, 1890, 32; Migula,
Arb. bakt. Inst. Karlsruhe, 1, 1894, 237.)
Pseu.do'mo.nas or Pseu.do.mo'nasJ. Gr. pseudes false; Gr. monas a unit, monad; M.L.
fem.n. Pseudomonas false monad.
Cells monotrichous, lophotrichous or non-motile. Gram-negative. Frequently develop
* Completely revised by Dr. Wm. C. Haynes, Northern Utilization Research Branch,
U.S.D.A., Peoria, Illinois (Species Nos. 1-58) and by Prof. Walter H. Burkholder, Cornell
University, Ithaca, New York (Species Nos. 59-149), September, 1953.
t See Footnote, p. 65. Also see Internat. Bull. Bact. Nomen. and Tax., 2, 1952, 121, foi
a proposal to conserve Pseudomonas Migula.
t The former accords with the Latin rules of accentuation; the latter is commonly used.
90 ORDER I. PSEUDOMONADALES
fluorescent, diffusible pigments of a greenish, bluish, violet, lilac, rose, yellow or other
color. Sometimes the pigments are bright red or yellow and non-diffusible; there are many
species that fail to develop any pigmentation. The majority of species oxidize glucose to
gluconic acid, 2-ketogluconic acid or other intermediates. Usually inactive in the o.xidation
of lactose. Nitrates are frequently reduced either to nitrites, ammonia or to free nitrogen.
Some species split fat and/or attack hydrocarbons. Many species are found in soil and
water, including sea water or even heavy brines. Many are plant pathogens; verj^ few are
animal pathogens.
The borderline between the straight rods found in Pseudomonas and the curved rods
found in Vibrio is not sharp : occasionally curved rods ma.y occur in species that normally
are composed of straight rods, this variation sometimes being dependent upon the medium
used. Recently, however, Shewan, Hodgkiss and Liston (Nature, 173, 1954, 208) have de-
scribed a method employing antibiotics and a vibriostatic agent whereby a sharper differen-
tiation between pseudomonads and vibrios may possibly be effected. Future studies of this
nature may show that some of the species in the genus Pseudomonas should be transferred
to the genus Vibrio, and vice versa.
The type species is Pseudomonas aeruginosa (Schroeter) Migula.
Key to the species of genus Pseudomonas.
I. Soil and water forms. A few species are pathogenic to warm- and cold-blooded verte-
brates.
A. Soil and fresh-water forms (a few are pathogenic).
1. Produce diffusible pigments, usually of a yellow, green or blue color; may be
fluorescent. (Soluble pigments are not formed in all media. Furthermore, the
ability to produce such pigments may be lost. Therefore, failure to observe sol-
uble-pigment formation does not preclude identity with species listed in this
category.)
a. Grow in gelatin.
b. Gelatin liquefied.
c. Polar flagellate.
d. Grows readily at 42°C. on ordinary media,
e. Milk becomes alkaline.
1. Pseudomonas aeruginosa.
ee. Milk acidified.
2. Pseudomonas pseudomallei.
dd. Grow poorly or not at all at 42° C.
e. Grow readily at 37° C.
f. Not known to attack cellulose.
g. Milk becomes alkaline, indole not produced.
3. Pseudomonas reptilivora.
gg. Milk acidified, indole produced.
4. Pseudomonas caviae.
ggg. Action on milk and indole production unre-
corded.
5. Pseudomonas horeopolis.
ff. Attack cellulose.
g. Milk becomes alkaline, coagulated and pep-
tonized, and litmus is reduced.
6. Pseudomonas effusa.
gg. No growth in milk.
7. Pseudomonas ephemerocyanea.
FAMILY IV. PSEUDOMOXADACEAE 91
ee. Grow poorly or not at all at 37°C.
f. Reaction in milk becomes acid or alkaline,
g. Acid reaction produced in milk.
8. Pseudotnonas fairmontensis .
gg. Alkaline reaction produced in milk.
h. Produces crj'stals of chlororaphine.
9. Pseudomonas chlororaphis.
hh. Chlororaphine not produced.
i. Indole produced.
10. Pseudomonas myxogenes.
11. Pseudomonas schuylkilliensis.
ii. Indole not produced.
j. Produces an intense, diffusible, yel-
low to orange pigment in cream or in
cream layer of milk.
12. Pseudomonas sijnxantha.
jj . Fail to produce diffusible, yellow pig-
ment in cream or in cream laj-er of
milk,
k. Nitrites produced from nitrates.
13. Pseudomonas fluorescens.
kk. Nitrites not produced from ni-
trates.
14. Pseudomonas pavonacea.
15. Pseudomonas geniculata.
16. Pseudomonas sepiica.
ii. Reaction in milk unchanged. Becomes blue in asso-
ciation with lactic-acid bacteria.
17. Pseudomonas syncyanea.
cc. Non-motile.
d. Produces iodinin.
18. Pseudomonas iodinum.
dd. Iodinin not produced.
e. Grows poorly or not at all at 37° C.
13. Pseudotnonas fluorescens (non-
motile variety),
ee. Grows readilj' at 37° C.
19. Pseudomonas smaraydina.
bb. Gelatin not liquefied.
c. Polar flagellate.
d. Grow readily at 37° C.
e. Reaction in milk unchanged.
20. Pseudomonas puiida.
ee. Alkaline reaction in milk.
f. Litmus reduced.
21. Pseudomonas striata,
ii. Litmus not reduced.
22. Pseudomonas oralis.
dd. Grow poorly or not at all at 37° C.
e. Reaction in milk acid.
f. Musty odor produced in culture media.
23. Pseudomonas taetrolens.
92 ORDER I. PSEUDOMONADALES
ff. Musty odor not produced in culture media.
24. Pseudomonas incognita.
25. Pseudomonas rugosa.
ee. Reaction in milk alkaline.
26. Pseudomonas mildenhergii.
27. Pseudomonas convexa.
cc. Non-motile.
28. Pseudoinonas eisenbergii.
aa. No growth in gelatin.
29. Pseudomonas erythra.
2. Soluble pigments not produced or not reported.
a. Gelatin liquefied.
b. Polar flagellate.
c. Grow readil}" at 42° C.
d. Alkaline reaction in milk.
1. Pseudomonas aeruginosa (achro-
mogenic variety' ).
dd. Acid reaction in milk.
2. Pseudomonas pseudomallei
(achromogenic variety).
cc. Grow readily at 25° C. but poorly or not at all at 37° C.
d. Acid reaction in milk.
e. May-apple odor produced in milk. Nitrites not produced
from nitrates.
30. Pseudomonas fragi.
ee. Musty odor produced from all media. Nitrites and am-
monia produced from nitrates.
31. Pseudomonas perolens.
dd. Alkaline reaction in milk.
32. Pseudomonas mephitica.
33. Pseudomonas putrefaciens .
34. Pseudomonas cohaerens.
aa. Gelatin not liquefied.
b. Polar flagellate.
c. Grow readily at 37° C.
d. Action on cellulose not known or not reported,
e. Acid produced in milk.
35. Pseudomonas avMgua.
ee. Milk imchanged.
36. Pseudomonas oleovorans.
eee. Action on milk unknown or unreported.
f. Utilize hydrocarbons.
37. Pseudomonas arvilla.
38. Pseudomonas dacunhae.
39. Pseudomonas desmolytica.
40. Pseudomonas rathonis.
41. Pseudomonas salopia.
ff. Ability to utilize hydrocarbons unreported.
g. Nitrites not produced from nitrates.
42. Pseudomonas cruciviae.
gg. Nitrates, nitrites, nitramids and NoO reduced
to elemental nitrogen.
43. Pseudomonas stutzeri.
FAMILY IV. PSEUDOMONADACEAE 93
dd. Attack cellulose
e. Acid produced in milk.
44. Pseudomonas tralncida.
ee. Milk unchanged.
45. Pseudomonas lasia.
cc. Grow readily at 25° C. but poorly or not at all at 37°C.
d. Action on cellulose unknown or unreported.
e. Alkaline reaction in milk. Attacks riboflavin converting
it to lumichrome.
46. Pseudomonas riboflavina.
ee. Reaction in milk unknown or unreported.
f. Nitrates reduced to elemental nitrogen.
47. Pseudomonas denitrificans .
ff . Nitrates reduced to nitrites. Indole decomposed with
formation of blue crj'stals of indigotin.
48. Pseudomonas indoloxidans.
dd. Attacks cellulose.
49. Pseudomonas mira.
B. Sea-water and brine forms (a few are pathogenic).
1. Produces pigments which are soluble in culture media. Gelatin liquefied.
50. Pseudomonas nigrifaciens.
2. Pigments soluble in culture media not produced.
a. Gelatin liquefied.
b. Polar flagellate.
c. No growth in milk.
d. "Indole produced; nitrites produced from nitrates.
51. Pseudomonas ichthyodermis.
dd. Indole not produced; nitrites not produced from nitrates.
e. Produces hydrogen sulfide and ammonia from tr5^ptone:
no acid from glucose.
52. Pseudomonas marinoglutinosa .
ee. Hydrogen sulfide not produced; glucose acidified.
53. Pseudomonas memhranoformis .
cc. Action in milk unknown or unreported.
d. Digests agar.
54. Pseudomonas gelatica.
dd. Do not digest agar.
e. Deposit CaCOs in sea-water gelatin and in agar media in
old cultures. Do not grow in 12 to 30 per cent salt solu-
tions.
55. Pseudomonas calcis.
56. Pseudomonas calciprecipitans.
ee. Does not deposit CaCOs in sea-water gelatin or in agar
media. Grows well in 12 to 30 per cent salt solutions.
56a. Pseudomonas halestorga.
bb. Non-motile. Cellulose attacked. Insoluble yellow pigment produced.
57. Pseudomonas iridescens (gela-
tin-liquefying variety),
aa. Gelatin not liquefied,
b. Polar flagellate.
c. Does not attack cellulose. Produces an insoluble purjjle pigment
in vegetable extracts.
58. Pseudomonas beijerinckii .
94 ORDER I. PSEUDOMONADALES
bb. Non-motile.
c. Cellulose attacked. Produces an insoluble yellow pigment.
57. Pseudomonas iridescens (non-
gelatin-liquefying variety) .
II. Plant pathogens, causing leaf spot, leaf stripe and similar diseases. (Also see Host Plant
Key, p. 96.)
A. Green fluorescent pigment produced.
1. Gelatin liquefied.
a. Acid from sucrose.
b. Growth in 5 per cent salt.
59. Pseudomonas aceris.
60. Pseudomonas angulata.
61. Pseudomonas aptata.
62. Pseudomonas primulae.
63. Pseudomonas viridilivida.
bb. No growth in 5 per cent salt.
c. Beef peptone agar turns deep brown.
64. Pseudomonas delphinii.
cc. Beef peptone agar imcolored.
d. Colonies yellow.
65. Pseudomonas cepacia.
dd. Colonies white to cream.
66. Pseudomonas apii.
67. Pseudomonas asplenii.
68. Pseudomonas berberidis.
69. Pseudomonas coronafaciens .
70. Pseudomonas lachrymans.
71. Pseudomonas maculicola.
72. Pseudomonas mangiferaeindicae.
73. Pseudomonas marginata.
74. Pseudomonas medicaginis.
75. Pseudomonas phaseolicola .
76. Pseudomonas pisi.
77. Pseudomonas syringae.
78. Pseudomonas tomato.
bbb. Growth in salt solution not recorded.
79. Pseudomonas atrofaciens.
80. Pseudomonas cumini.
81. Pseudomonas desaiana.
82. Pseudomonas erodii.
83. Pseudomonas lapsa.
84. Pseudomonas martyniae.
85. Pseudomonas matthiolae.
86. Pseudomonas 7uorspru?iorum.
87. Pseudomonas papulans.
88. Pseudomonas pseudozoogloeae.
89. Pseudomonas rimaefaciens .
90. Pseudomonas striafaciens.
91. Pseudomonas tabaci.
aa. No acid from sucrose,
b. Lipolytic.
92. Pseudomonas polycolor
FAMILY IV. PSEUDOMONADACEAE
95
bb. Not lipolytic,
bbb. Lipolytic action not reported.
93. Pseudomonas viridiflava.
aaa. Acid from sucrose not i-eported.
2. Gelatin not liquefied,
a. Acid from sucrose.
I. No acid from sucrc
b. Non-motile.
bb. Motile.
aaa. Acid from sucrose not reported.
94. Pseudomonas
95. Pseudomonas
96. Pseudomonas
97. Pseudomonas
98. Pseudomonas
99. Pseudomonas
100. Pseudomonas
101. Pseudomonas
102. Pseudomonas
103. Pseudomonas
104. Pseudomonas
105. Pseudomonas
106. Pseudomonas
107. Pseudomonas
108. Pseudomonas
ananas,
bowlesiae .
ligustri.
marginalis.
sesami.
setariae.
tolaasii.
washingtoniae .
harkeri.
be tie.
gladioli.
niellea.
panacis.
ribicola .
xanthochlora.
109. Pseudomonas aleuritidis.
110. Pseudomonas glycinea.
111. Pseudomonas savastanoi.
112. Pseudomonas tonelliana.
113. Pseudo?nonas cissicola.
114. Pseudomonas calendulae.
115. Pseudomonas dehor ii.
116. Pseudomonas nectarophila.
117. Pseudomonas viburni.
B. Green fluorescent pigment
1. Gelatin liquefied.
a. Acid from sucrose,
b. Beef -peptone
118. Pseudomonas mori.
119. Pseudomonas stizolobii.
120. Pseudomonas viciae.
not produced or not reported.
agar turns dark brown.
121. Pseudomonas alliicola.
122. Pseudomonas gardeniae.
bb. Beef-peptone agar uncolored or only slightly so.
c. Colonies tan to brown.
123. Pseudomonas caryophylli .
124. Pseudomonas solanacearum.
cc. Colonies white or colorless.
125. Pseudomonas casianeae.
126. Pseudomonas passiflorae.
127. Pseudomonas seminum.
128. Pseudomonas vitiswoodrowii.
96
ORDER I. PSEUDOMOXADALES
aa. No acid from sucrose,
aaa. Acid from sucrose not reported.
2. Gelatin not liquefied,
a. Acid from sucrose.
aa. No acid from sucrose.
aaa. Acid from sucrose not reported.
3. Gelatin liquefaction not reported.
129. Pseudomonas fabae.
130. Pseudouionas
131. Pseudomonas
132. Pseudomonas
133. Pseudomonas
134. Pseudomonas
135. Pseudomonas
136. Pseudomonas
astragali.
colurnae.
iridicola.
levistici.
viauhlancii.
polygoni.
radiciperda.
137. Pseudomonas cattleyae.
138. Pseudomonas dysoxyli.
139. Pseudomonas helianthi.
140. Pseudomonas melophthora.
141. Pseudomonas alhoprecipitans .
142. Pseudomonas andropogonis.
143. Pseudomonas lignicola.
144. Pseudomonas petasitis.
145. Pseudomonas woodsii.
146. Pseudomonas eriobotryae.
147. Pseudomonas panicimiliacei .
148. Pseudomonas salicivp.rda
149. Pseudomonas wieringae.
HOST PLANT KEY
Where the host plant is known, the following key will be foimd useful.
I. Cause of necrotic spots on mushrooms.
100. Pseudomonas tolaasii.
II. Cause of spots on ferns, Asplenium nidus.
67. Pseudomonas asplenii.
III. Cause of leaf blights and streaks on monocotyledonous plants.
A. Attack members of the family Amaryllidaceae.
121. Pseudomonas alliicola.
65. Pseudomonas cepacia.
B. Attacks members of the family Bromeliaceae.
94. Pseudomonas ananas.
C. Attack members of the family Gramineae.
141. Pseudomonas alboprecipitans.
142. Pseudomonas andropogonis.
79. Pseudomonas atrofaciens.
69. Pseudomonas coronafaciens .
81. Pseudomonas desaiana.
83. Pseudomonas lapsa.
147. Pseudomonas panicimiliacei.
99. Pseudomonas setariae.
90. Pseudomonas striafaciens .
FAMILY IV. PSEUDOMONADACEAE 97
D. Attack members of the family Iridaceae.
104. Pseudoinonas gladioli.
132. Pseudomonas iridicola.
73. Pseudomonas marginata.
E. Attacks members of the family Musaceae.
134. Pseudomonas matihlancii.
F. Attacks members of the family Orchidaceae.
137. Pseudomonas cattleyae.
G. Attacks members of the family Palmaceae.
101. Pseudomonas washingtoniae .
IV. Cause of leaf, stem and fruit spots on dicotyledonous plants.
A. Attacks members of the famih' Aceraceae.
59. Pseudomonas aceris.
B. Attacks members of the family Anacardiaceae.
72. Pseudomonas mangiferaeindicae.
C. Attacks members of the family Apocynaceae.
112. Pseudomonas tonelliana.
D. Attacks members of the family Araliaceae.
106. Pseudomonas panacis.
E. Attacks members of the family Berberidaceae .
68. Pseudomonas berberidis.
F. Attacks members of the family Betulaceae .
131. Pseudomonas colurnae.
G. Attacks members of the family Caprifoliaceae.
117. Pseudomonas vihurni.
H. Attack members of the family Caryophyllaceae.
123. Pseudomonas caryophylli.
145. Pseudomonas woodsii.
I. Attacks members of the family Chenopodiaceae.
149. Pseudomonas wieringae.
J. Attack members of the family Compositae.
114. Pseudoynonas calendulae.
115. Pseudomonas cichorii.
139. Pseudomonas helianthi.
97. Pseudomonas marginalis.
144. Pseudomonas petasitis.
63. Pseudomonas viridilivida.
K. Attack members of the family Cruciferae.
71. Pseudomonas maculicola.
85. Pseudomonas matthiolae.
L. Attacks members of the family Cucurbitaceae.
70. Pseudomonas lachrymans.
M. Attacks members of the family Euphorbiaceae.
109. Pseudomonas aleuritidis.
N. Attacks members of the family Fagaceae.
125. Pseudoinonas castaneae.
O. Attacks members of the family Geraniaceae.
82. Pseudomonas erodii.
P. Attack members of the family Legmninosae.
130. Pseudomonas astragali.
129. Pseudomonas fabae.
110. Pseudomonas glycinea.
98 ORDER I. PSEUDOMOXADALES
74. Pseudomonas medicaginis.
75. Pseudomonas phaseolicola.
76. Pseudomonas pisi.
136. Pseudomonas radiciperda.
127. Pseudomonas seminum.
119. Pseudomonas stizolohii.
110. Pseudomonas viciae.
93. Pseudomonas viridiflava.
Q. Attacks members of the family Martyniaceae.
84. Pseudomonas martyniae.
R. Attacks members of the family Meliaceae.
138. Pseudomonas dysoxyli.
S. Attacks members of the family Moraceae.
118. Pseudomonas mori.
T. Attack members of the family Oleaceae.
96. Pseudomonas ligustri.
111. Pseudomonas savastanoi.
U. Attacks members of the famih' Passifloraceae.
126. Pseudomonas passiflorae.
V. Attacks members of the family Pedaliaceae.
98. Pseudomonas sesami.
W. Attacks members of the iamily Piperaceae.
103. Pseudomonas belle.
X. Attacks members of the family Polygonaceae.
135. Pseudomonas polygoni.
Y. Attacks members of the family Primulaceae.
62. Pseudomonas primulae.
Z. Attacks members of the famih^ Ranunculaceae.
64. Pseudomonas delphinii.
AA. Attack members of the family Rosaceae.
102. Pseudomonas harkeri.
146. Pseudomonas eriobotryae.
140. Pseudomonas melophthora.
86. Pseudomonas morsprunorum.
116. Pseudomonas nectarophila.
87. Pseudomonas papulans.
BB. Attacks members of the family Rubiaceae.
122. Pseudomonas gardeniae.
CC. Attacks members of the famil}^ Salicaceae.
148. Pseudomonas saliciperda.
DD. Attacks members of the family Saxifragaceae.
107. Pseudomonas ribicola.
EE. Attack members of the family Solanaceae.
60. Pseudomonas angulata.
105. Pseudomonas mellea.
92. Pseudanonas polycolor.
88. Pseudomonas pseudozoogloeae.
91. Pseudomonas tabaci.
78. Pseudomonas tomato.
FF. Attacks members of the family Ulmaceae.
143. Pseudomonas lignicola.
GG. Attack members of the family Umbelliferae.
66. Pseudomonas apii.
FAMILY IV. PSEUDOMONADACEAE
99
95. Pseudomonas bowlesiae.
80. Pseudomonas cumini.
133. Pseudomonas levistici.
HH. Attacks members of the family Vitaceae.
113. Pseudomonas cissicola.
II. Attack members of numerous families.
61. Pseudomonas aplata.
124. Pseudomonas solanacearum.
11 . Pseudomonas syringae.
108. Pseudomonas xanthochlora.
1. Pseudomonas aeruginosa (Schroe-
ter, 1872) Migula, 1900. {Bacterium aerugi-
nosum Schroeter, in Cohn, Beitrage z.
Biologic, 1, Heft 2, 1872, 126; Bacillus pyo-
cyaneus Gessard, Compt. rend. Acad. Sci.,
Paris, 94, 1882, 536; Pseudomonas pyocyanea
Migula, in Engler and PrantI, Die natiirl.
Pflanzenfam., 1, la, 1895, 29; Migula, Syst.
d. Bakt., 2, 1900, 884.)
ae.ru. gi.no'sa. L. adj. aeruginosus full of
copper rust or verdigris, hence green.
Common name: Blue pus organism.
Rods, 0.5 to 0.6 b}' 1.5 microns, occurring
singly, in pairs and short chains. Motile,
possessing one to three polar flagella.
(Monotrichous according to Reid, Naghski,
Farrell and Haley, Penn. Agr. E.xp. Sta.,
Bull. 422, 1942, 6.) Gram-negative.
Gelatin colonies: Yellowish or greenish
yellow, fringed, irregular, skein-like, granu-
lar, rapidly liquefying.
Gelatin stab: Rapid liquefaction. The
fluid assumes a yellowish green or bluish
green color.
Agar colonies: Large, spreading, grayish
with dark center and tran.slucent edge, ir-
regular. Medium greenish.
Agar slant: Abundant, thin, white, glis-
tening, the medium turning green to dark
brown or black, fluorescent.
Broth: Marked turbidity with thick pel-
licle and heavy sediment. Medium yellowish
green to blue, with fluorescence, later
brownish. Often produces pyocyanine, fluo-
rescein and pyrorubrin (Meader, Robinson
and Leonard, Am. Jour. Hyg., 5, 1925, 682).
Litmus milk: A soft coagulum is formed,
with rapid peptonization and reduction of
litmus. Reaction alkaline.
Potato: Luxuriant, dirty brown, the me-
dium becoming dark green.
Indole usually not produced (Sandiford,
Jour. Path, and Pact., U, 1937, 567).
Nitrates reduced to nitrites and nitrogen.
Glucose, fructose, galactose, arabinose,
maltose, lactose, sucrose, dextrin, inulin,
glycerol, mannitol and dulcitol are not fer-
mented. Glucose oxidized to gluconic acid,
2-ketogluconic acid and other intermediates
(Lockwood, Tabenkin and Ward, Jour.
Bact., 42, 1941, 51; Hajmes, Jour. Gen.
Microbiol., 5, 1951,939).
Blood serum: Liquefied. Yellow liquid,
greenish on surface.
Blood hemolyzed.
Cultures have marked odor of trimethyl-
amine.
Aerobic, facultative.
Optimum temperature, 37° C. Good
growth at 42° C.
Pathogenic for rabbits, guinea pigs, rats
and mice.
Distinctive characters: Some strains pro-
duce p.yocyanine, a phenazine derivative
which is extractable from alkaline media
with chloroform as a deep blue pigment.
Upon addition of acid, the color is trans-
formed to red and becomes insoluble in
chloroform. The ability to grow well at
42° C, to oxidize gluconic acid to 2-keto-
gluconic acid and to produce a slime in
potassium gluconate media permits identifi-
cation even though pyocyanine is not
formed (Haynes, loc. cit.).
Source: Pus from wounds. Regarded as
identical with one of the plant pathogens
(Pseudomonas polycolor) by Elrod and Braun
(Jour. Bact., U, 1942, 633).
Habitat: Cause of various human and
animal lesions. Found in polluted water and
100
ORDER I. PSEUDOMONADALES
2. Pseudoiiionas pseudomallei (Whit-
more, 1913) Haynes, comb. nov. {Bacilhis
pseudomallei Whitmore, Jour. Hyg., 13,
1913, 1; Bacillus whitmori Stanton and
Fletcher, Trans. 4th Cong. Far East Assn.
Trop. Med., ^, 1921, 196; also see Jour. Hyg.,
23, 1925, 347; Malleomyces pseudomallei
Breed, in Manual, 5th ed., 1939, 300;
Loefflerella pseudomallei Brindle and Cowan,
Jour. Path, and Bact., 63, 1951, 574.)
pseu.do.mal'le.i. Or. adj. pseudes false;
L. noun malleus the disease glanders; M.L.
noun pseudomalleus false glanders; M.L.
gen. noun pseudoviallei of false glanders.
Short rods, with rounded ends, occurring
singly and in short chains. Motile. Possess
1 to 4 polar flagella (Brindle and Cowan,
ibid., 571) ; this was confirmed by de Lajudie,
Fournier and Chambon (Ann. Inst. Past.,
85, 1953, 112). Show bipolar staining. Gram-
negative.
Gelatin stab: Moderate, crateriform
liquefaction.
Agar colonies: Circular, slightly raised,
thick, opaque, cream-colored with irregular
margin.
Glycerol agar slant: Wrinkled, thick,
rugose, cream-colored growth.
Broth: Turbid with pellicle.
Litmus milk: Curdling with slowly de-
veloped acidity; pink sediment; may be
digested.
Potato: Vigorous, cream-colored growth.
Indole not produced.
Acid from glucose, maltose, lactose,
sucrose and mannitol.
Grows in simple, chemically defined media
containing single amino acids or the am-
monium salt of certain organic acids as the
sole carbon, nitrogen and energy source in a
mineral salt base (Levine, Dowling, Even-
son and Lien, Jour. Bact., 67, 1954, 350).
Blood serum slowly liquefied.
Aerobic, facultative.
Optimum temperature, 37° C; but will
grow readily at 42° C. (Cowan, personal
communication, March, 1955).
Distinctive character: Brygoo and Rich-
ard (Ann. Inst. Past., 83, 1952, 822) report
that a large number of strains, isolated in
Saigon, produce a yellow pigment which is
extractable in 2 per cent boiling HCl ; a few
of these strains become non-pigmented
when cultured on glycerol agar media. While
this pigment has sometimes been described
as water-soluble, Brindle and Cowan (op.
cit., 1951, 574) suggest that this species may
be more closely related to the species placed
in Xanthomonas than to those placed in
Pseudomonas. The xanthomonads develop
yellow, non-diffusible, carotenoid pigments.
Source: Isolated from lesions and blood in
rats, guinea pigs, rabbits and man; also iso-
lated once from a transient nasal discharge
in a horse, once from a splenic abscess in a
cow and once from a fatal case of an infected
sheep. Virulent and avirulent strains can
also be readily isolated from water at
Saigon, Indochina, if appropriate media are
used (Fournier and Chambon, personal
communication, 1955).
Habitat: Glanders-like infections (melioi-
dosis) in rats, guinea pigs, rabbits and man
in India, Federated Malay States and Indo-
China.
3. Pseudomonas reptilivora Caldwell
and Ryerson, 1940. (Pseudomonas reptilivor-
ous (sic) Caldwell and Ryerson, Jour. Bact.,
39, 1940, 335.)
rep.ti.li'vo.ra. L. n. reptile a reptile;
L. V. voro to devour; M.L. adj. reptilivorus
reptile-destroying.
Rods, 0.5 by 1.5 to 2.0 microns, occurring
singly, in pairs and in short chains. Actively
motile with two to six polar flagella. Gram-
negative.
Gelatin colonies: After 24 hours, small,
circular, smooth, entire. Liquefaction.
Medium becomes yellowish green fluores-
cent.
Gelatin stab: Infundibuliform liquefac-
tion becoming stratiform. Putrid odor
present.
Serum slant: Liquefied.
Agar cultures: Circular, smooth, glisten-
ing, slightly raised, butyrous, translucent, 2
mm in diameter.
Agar slant: Growth abundant, smooth,
filiform, glistening, butyrous and translu-
cent.
Broth: Turbid with pellicle and sediment.
Putrid odor.
Litmus milk: Alkaline, peptonization,
complete reduction. Disagreeable odor.
Potato: Growth moderate, spreading.
FAMILY IV. PSEUDOMONADACEAE
101
i^listening, yellowish gray to creamy. Dis-
agreeable odor. Medium becomes brownish
gray.
Indole not produced.
Nitrites not produced from nitrates.
Acetylmethjdcarbinol not produced.
Hydrogen sulfide not produced.
Slightly acid, becoming alkaline in glu-
cose. No acid from arabinose, .xylose, lactose,
sucrose, maltose, trehalose, raffinose, man-
nitol, dulcitol, inositol or salicin.
Starch not hydrolyzed.
Pathogenic for guinea pigs and rabbits,
horned lizards, Gila monsters and chuck-
wallas. Marked hemolysis of rabbit cells
and slight hemolysis of Gila monster cells
suspended in agar.
Temperature relations: Optimum, 20° to
25° C. Maximum, 37° C. A retest of several
strains of this organism by Haynes shows
that it grows well at 37° C. and is closelj^
related to, though not identical with,
Pseudomonas aervginosa Migula.
Distinctive characters: Yellowish green
fluorescence present only in meat infusion
media. The pigment is water-soluble, but
insoluble in chloroform. Pathogenic for
guinea pigs, rabbits, horned lizards and
chuckwallas.
Source: Isolated from a bacterial disease
of horned lizards and Gila monsters.
Habitat: Pathogenic for lizards.
4. Pseudomonas caviae Scherago, 1936.
(Jour. Bact., 31, 1936, 83; also see Jour. Inf.
Dis., 60, 1937, 245.)
ca'vi.ae. M.L. fem.noun Cavia generic
name of the guinea pig; from So. American
Indian, "cabiai", a guinea pig; caviae of
Cavia.
Rods, 0.6 to 1.0 by 1.5 to 3.0 microns, oc-
curring singly and in pairs; rounded ends.
Motile by means of 1 to 3 polar flagella.
Encapsulated. Gram-negative.
Gelatin stab: Infundibuliform liquefac-
tion.
Agar colonies: Circular, convex, smooth,
iridescent and translucent, finely granular,
entire.
Agar slant: Growth abundant, grayish
white, butyrous, filiform, glistening, trans-
lucent, markedly iridescent. Medium even-
tually tinged greenish j'ellow, becoming
brownish yellow.
Broth: Cloudy, pellicle, abundant light
yellow granular sediment, becoming brown.
IVIedium becomes j^ellow.
Litmus milk: Acidified, coagulated, pep-
tonized, litmus partially reduced.
Potato: Growth scant, filiform, glisten-
ing, light yellow to light orange, becoming
light brown.
Hydrogen sulfide not produced.
Indole produced.
Nitrites produced from nitrates.
Blood serum not liquefied.
Blood not hemolyzed.
Sodium formate decomposed.
Catalase-negative.
Methyl red positive; acetylmethylcar-
binol not produced.
Citrate broth: No growth.
Methylene blue reduced.
Acid but no gas from glucose, fructose,
galactose, maltose, cellobiose, mannitol,
lactose, arabinose, sucrose, trehalose, sor-
bitol, mannose, dextrin, salicin, glycerol,
aesculin, am3^gdalin and starch. No acid
from xylose, dulcitol, rhamnose, inulin,
adonitol, raffinose, erythritol or inositol.
Aerobic, facultative.
Optimum temperature, 37° C. Grows at
25° C.
Source: Isolated from guinea pigs dead
from epizootic septicemia.
Habitat : From infected guinea pigs so far
as known.
5. Pseudomonas boreopolis Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 92.)
bo.re.o'po.lis. Gr. boreas north; Gr. polis
a city; M.L. fern. gen. n. boreopolis of North
City.
Rods, 0.5 to 1.0 by 2.0 to 3.0 microns, oc-
curring singly and in pairs. Motile with one
to five polar flagella. Gram-negative.
Gelatin colonies: Liquefied.
Gelatin stab: Liquefied. Medium red-
dened by some strains.
Agar colonies: Circular or amoeboid,
white to buff, flat to convex, smooth,
glistening, translucent border.
Agar slant: Filiform, whitish, raised,
smooth, glistening, fluorescent.
102
ORDER I. PSEUDOMONADALES
Broth: Turbid.
Nitrates reduced to nitrites by some
strains.
Starch not hydrolyzed.
Acid produced from glucose by most
strains.
Attacks naphthalene.
Aerobic, facultative.
Grows at 35° to 37° C.
Source: Isolated from soil.
Habitat: Soil.
6. Pseudomonas efifusa Kellerman et al.,
1913. (Kellerman, McBeth, Scales and
Smith, Cent. f. Bakt., II Abt., 39, 1913, 515;
also see Soil Science, 1, 1916, 472.)
ef.fu'sa. L. adj. effusus spread out.
Rods 0.4 by 1.7 microns. Motile by means
of one to three polar flagella. Gram-nega-
tive.
Gelatin stab: Liquefied. A non-liquefying
variety is also found.
Agar slant: Luxuriant, glistening, moist,
creamy, spreading growth. Medium be-
comes greenish fluorescent.
Peptone starch agar slant : Abundant, flat,
moist, rich creamy growth. Medium shows
greenish fluorescence.
Broth: Turbid; viscid sediment. Medium
becomes greenish fluorescent.
Litmus milk: Alkaline. Coagulation and
digestion. Litmus reduced. A variety that
acts more slowly on litmus milk is also
found.
Potato: Abundant, creamy, glistening,
brownish flesh-colored growth.
Indole not produced.
Nitrites produced from nitrates.
Ammonia is produced.
No acid from glucose, starch, lactose,
sucrose, maltose, glj'cerol or mannitol.
Starch hydrolysis weak.
Cellulose is attacked.
Aerobic, facultative.
Temperature relations: Optimum, 37° C.
Survives 60° C, but not 70° C, for 15 min-
utes.
Source: Isolated from soils in Utah.
Habitat: Soil.
7. Pseudomonas ephemerocyanea Ful-
ler and Norman, 1943. (Jour. Bact., 46,
1943, 274.)
e.phe.me.ro.cy.a'ne.a. Gr. adj. epheme-
rus short-lived; Gr. adj. ajaneus blue; M.L.
adj. ephemerocyaneus ephemeral blue.
Rods, 0.3 to 0.4 by 2.2 to 2.8 microns,
straight to slightly bent with rounded ends,
arranged singlj-. Motile by means of 1 to 3
polar flagella. Gram-negative.
Gelatin stab: Liquefied.
Starch agar colonies: Pinpoint colonies
in three days, 1 to 2 mm in 5 days. White
becoming tan, raised, glistening, smooth,
entire.
Water-insoluble de.xtrin colonies: Pin-
point colonies show an enzymic zone, white,
convex, entire.
Starch agar slant: Heavy gelatinous,
light brown becoming deeper brown.
Litmus milk: No visible growth.
Indole not produced.
Nitrites produced from nitrates.
Starch hydrolyzed.
Attacks glucose, lactose, maltose, galac-
tose, arabinose and xylose. Utilizes cellu-
lose, cellulosan, water-soluble and water-
insoluble cellulose dextrins and pectin. Slow
utilization of gum arable and calcium glu-
conate.
In mineral nutrient media, filter paper
strips are disintegrated at the air-liquid
interface with the formation of a transitory
violet or blue color which becomes light
brown.
Peptone, yeast extract, nitrate and am- ■
monia serve as nitrogen sources.
Aerobic.
Optimum temperature, 22° to 35° C.
Distinctive characters: In media con-
taining cellulose a transitory intense blue
or violet color develops. In aerated cultures
the entire medium becomes blue. The pig-
ment appears to be water-soluble. After a
few hours the color becomes light brown.
Source: Isolated from soil.
Habitat: Soil.
8. Pseudomonas fairmontensis
(Wright, 1895) Chester, 1901. (Bacillus Fair-
montensis (sic) Wright, Memoirs Nat. Acad.
Sci., 7, 1895, 458; Chester, Man. Determ.
Bact., 1901, 311.)
fair.mon.ten'sis. Fairmount Park (Phila-
delphia) place name; M.L. adj. fairmonten-
sis pertaining to Fairmount.
Medium-sized rods, occurring singly, in
FAMILY IV. PSEUDOMONADACEAE
103
pairs and in chains. Motile, possessing a
single polar flagellum. Gram-negative.
Gelatin colonies: Circular, white, trans-
lucent. Dark centers with a greenish shim-
mer, thinner edges and faint radial lines.
Gelatin stab: Crateriform liquefaction.
Agar slant: Grayish white, glistening.
Agar becomes green.
Broth: Turbid; delicate pellicle; white
sediment. Becomes green.
Litmus milk: Acid, coagulated; litmus
reduced.
Potato: Raised, granular, spreading, vis-
cid. Becomes brownish.
Indole produced.
Action on nitrates unknown.
Aerobic, facultative.
Optimum temperature, 20° to 25° C. Fails
to grow at 35° C.
Source: From water from the Schuylkill
River.
Habitat: Water.
9. Pseudomonas chlororaphis (Guig
nard and Sauvageau, 1894) Bergey et al.
1930. (Bacillus chlororaphis Guignard and
Sauvageau, Compt. rend. Soc. Biol., Paris
1, 10 ser., 1894, 841; Bergey et al.. Manual
3rd ed., 1930, 166.)
chlo.ro'ra.phis. Gr. chlorus green; Gr
noun rhaphis a needle; M.L. fem.n. chloro
r aphis a green needle.
Description taken from Lasseur (Ann. de
la Sci. Agron., Ser. 4,2« Annee, 2, 1913, 165)
While Guignard and Sauvageau {op. cit.
1894, 841) found .spores in this species
Gessard, on reisolation, could find no spores
(Ann. de la Sci. Agron., Ser. 3, 6^ Ann^e, 2
1911, 374). The identification of the reiso
lated culture was confirmed by Guignard
The original description is brief and inade-
quate and is probably based on a contami-
nated culture.
Rods, 0.8 b}' 1.5 microns, with rounded
ends, occurring singly and in pairs. Motile
with one to si.\ polar flagella. Gram-nega-
tive. After continued cultivation some cells
decolorize slowly.
Gelatin colonies: Circular, viscid, trans-
parent, glistening, lobate margin with
fluorescent corona. Dissociates readilj^
(Lasseur and Dupai.x-Lasseur, Trav. Lab.
Microbiol. Fac. Pharm. Nancy, Fasc. 9,
1936, 35).
Gelatin stab : Rapid liquefaction. Fluores-
cent. Chlororaphine crystals may form.
Broth: Turbid, greenish, fluorescent.
Crystals of green chlororaphine may form.
Broth becomes viscous.
Litmus milk: Alkaline; coagulated. Be-
comes viscous. Chlororaphine crystals may
form in the central part of the culture. Odor
of coumarin.
Potato: Citron-yellow layer. Crystals of
chlororaphine are formed.
Nitrites produced from nitrates.
Indole not produced.
Pigment formation: Asparagine, potas-
sium phosphate, glycerol, sulfate of mag-
nesium and sulfate of iron are indispensable
to the formation of crystals of chlorora-
phine. Green crystals develop slowly and
poorly in peptone solutions, best in syn-
thetic media.
Aerobic, facultative.
Optimum temperature, between 25° and
30° C. Cultures killed in ten minutes at
63° C.
Pathogenic for mice, guinea pigs, frogs,
fresh-water fishes and crayfishes. An e.xo-
toxin is formed.
Distinctive character: Produces a beauti-
ful emerald-green pigment which crystal-
lizes in cultures as fine needles in bundles
or as needles radiating from a center. The
crystals form slowly and are not always
present. Other species of pseudomonads,
e.g. Pseudomonas iodinum, form crystals.
As this power is readily lost, it raises the
question whether other species of green,
fluorescent pseudomonads may not form
crystals under proper conditions.
Source: Isolated from dead larvae of the
cockchafer. Later reisolated bj^ various
French bacteriologists from contami-
nated water supplies.
Habitat: Decomposing organic matter
and fresh water so far as known.
10. Pseudomonas myxogenes Fuhr-
mann, 1907. (Cent. f. Bakt., II Abt., 17,
1907, 356.)
myx.o'ge.nes. Gr. myxa slime; Gr. gennao
to produce, beget; M.L. adj. myxogenes
slime-producing.
104
ORDER I. PSEUDOMONADALES
Rods, 0.4 to 0.5 by 1.0 to 1.5 microns, oc-
curring singly and in pairs. Motile, possess-
ing a bundle of five to seven polar flagella.
Gram-negative.
Gelatin colonies: Smooth, soft, flat,
spreading, brownish yellow, entire. Medium
becomes yellowish green fluorescent.
Gelatin stab: Growth along stab. Lique-
faction with yellowish white sediment.
Agar colonies: Circular, raised, smooth,
amorphous, entire.
Agar slant: Lemon-yellow, moist, mu-
coid, gistening, becoming light green-
fluorescent.
Broth: Turbid, with slimy white sedi-
ment. No pellicle.
Litmus milk: Flocculent precipitation.
Slow peptonization with yellow serum.
Alkaline.
Potato: Dirty yellow, moist, glistening,
entire.
Indole produced.
Nitrates reduced to nitrites and am-
monia. No gas formed.
Acid from glucose. No acid from lactose
or sucrose.
Aerobic, facultative.
Optimum temperature, 22° C. Scant
growth at 35° C.
Distinctive character: Grows in broth
containing up to 6 per cent by volume of
alcohol.
Source: Isolated from beer.
Habitat: Found in materials undergoing
alcoholic fermentation, but probably also
occurs elsewhere.
11. Pseudomona.s schuylkilliensis
Chester, 1901. (Bacillus fluorescens schuyl-
killiensis Wright, Memoirs Natl. Acad.
Sci., 7, 1895, 448; Chester, Man. Determ.
Bact., 1901, 320.)
schuyl.kil.li.en'sis. Schujdkill, name of a
river; M.L. adj. schuylkilliensis of the
Schuylkill.
Short rods, with rounded ends, occurring
singly, in pairs and in chains. Motile, posses-
sing a polar flagellum. Gram-negative.
Gelatin colonies: Grayish white, translu-
cent. Medium becomes bluish green fluores-
cent.
Gelatin stab: Slow crateriform liquefac-
tion, with blue-green fluorescence.
Agar slant: Grayish, translucent growth.
Medium shows greenish fluorescence.
Broth: Turbid, with delicate pellicle and
blue-green fluorescence. Stringy sediment.
Litmus milk: Alkaline. Coagulated, with
slow reduction of litmus; peptonized.
Potato: Brownish, spreading, viscid,
thick.
Indole jjroduced (trace).
Aerobic, facultative.
Does not grow at 35° to 36° C.
Source: Isolated from Schuylkill River
water.
Habitat: Water.
12. Pseudomonas synxantha (Ehren-
berg, 1840) Holland, 1920. (Vibrio synxan-
thus Ehrenberg, Verhandl. d. Berl. Akad.,
1840, 202; Holland, Jour. Bact., 5, 1920, 220.)
syn.xan'tha. Gr. pref. syn- along with,
together; Gr. adj. xanthus yellow; M.L.
adj . synxanihus with yellow.
Description from Hammer (Res. Bull. 20,
Iowa Agr. Exp. Sta., 1915); also see Zim-
mermann (Bakt. unserer Trink- und Nutz-
wasser, Chemnitz, 2, 1890, 44).
Rods, 0.5 to 0.6 by 1.3 to 2.2 microns, oc-
curring singly and in pairs. Motile with
polar flagella (Hammer, personal communi-
cation, 1944). Gram-negative.
Gelatin stab: Liquefied; a greenish tinge,
a heavy, flocculent sediment and a partial
membrane and ring appear in two weeks.
Agar colonies: After 72 hours, large,
spreading, transparent; bluish cast by re-
flected light. Colonies may show flesh
color (Zimmermann).
Agar slant: Growth raised, shiny, white,
becoming brown and heavy.
Agar stab : Growth heaviest near the sur-
face, becoming light brown, heavy, spread-
ing.
Broth: Turbid, becoming alkaline and
green; pellicle and brittle membrane form
in older cultures. With the addition of glu-
cose or galactose, black granules form on
the membranes of older cultures.
Uschinsky's and Dunham's solutions:
Turbid, occasionally becoming green.
Litmus milk: Coagulated; casein digested
in older cultures. Litmus reduced.
Potato: Growth spreading, brown with
greenish edges.
FAMILY IV. PSEUDOMONADACEAE
105
Indole not produced.
Acid but no gas from glucose, fructose,
galactose and glycerol. No acid or gas from
salicin or raffinose.
Aerobic.
Grows well at 20° C.
Distinctive character: Produces an in-
tense, diffusible, yellow to orange color in
cream or in the cream layer of milk.
Source: Isolated from bitter milk.
Habitat: Milk and cream so far as is
known.
13. Pseudoinonas fluorescens INIigula,
1895. (Bacilhis fluorescens liquefadens
Fliigge, Die Mikroorganismen, 1886, 289;
Migula, in Engler and Prantl, Die natiirl.
Pflanzenfamilien, 1, la, 1895, 29.)
flu.o.res'cens. L. fluor a flux; M.L.
fluoresco to fluoresce; fluor-spar, a fluxing
mineral which is fluorescent; M.L. part.
ad] . fluorescens fluorescing.
Rods, 0.3 to 0.5 by 1.0 to 1.8 microns, oc-
curring singly and in pairs. Motile, posses-
sing a polar flagellum; occasionally non-
motile. Gram-negative.
Gelatin colonies: Circular, with greenish
center, lobular, liquefying quickly; occa-
sionally viscid.
Gelatin stab: Infundibuliform liquefac-
tion, with whitish to reddish gray sediment.
Agar slant: Abundant, reddish layer, be-
coming reddish gray. The medium shows
greenish to olive-brown coloration.
Broth: Turbid, flocculent, with yellowish
green pellicle and graj'ish sediment.
Litmus milk: No coagulation; becoming
alkaline.
Potato: Thick, grayish yellow, spreading,
becoming light sepia-brown in color; occa-
sionally viscid.
Indole not produced.
Nitrates reduced to nitrites and am-
monia.
Acid from glucose.
Blood serum liquefied.
Aerobic.
Optimum temperature, between 20° and
25° C.
Not pathogenic.
Source: Water, sewage, feces.
Habitat: Soil, water and occasionally
foodstuffs that have become contaminated
from these sources.
14. Pseudomonas pavonacea Levine
and Soppeland, 1926. (Bull. No. 77, Iowa
State Agr. College, 1926, 41.)
pa.vo.na'ce.a. L. adj. pavonaceus like a
peacock's tail, variegated.
Rods, 0.5 by 4.5 microns, with truncate
ends, occurring singly and in chains. Old
cell'- develop 2 to 4 knob-like processes.
Sluggishly motile. Gram-negative.
Gelatin stab: Crateriform liquefaction.
Medium becoming brown.
Agar colonies: Circular, raised, becoming
green, amorphous, entire.
Agar slant: Green, smooth, glistening,
viscid, medium becoming dark brown.
Broth: Turbid, with viscid sediment. Me-
dium turned brown.
Litmus milk: Slightly alkaline. Litmus
reduced. Peptonized after 10 days.
Potato: No growth.
Hydrogen sulfide produced.
Indole not produced.
Neither nitrites nor gas produced from
nitrates.
Blood serum liquefied in 5 days.
No acid or gas from glucose, lactose,
sucrose or glycerol.
Aerobic, facultative.
Optimum temperature, 22° C. Scant
growth at 37° C.
Distinctive characters: Growth on solid
media distinctly green. Not fluorescent.
Medium becomes brown.
Source: Isolated from activated sludge.
15. Pseudomonas geniculata (Wright,
1895) Chester, 1901. (Bacillus geniculatus
Wright, Memoirs Nat. Acad. Sci., 7, 1895,
459; Chester, Man. Determ. Bact., 1901,
313.)
ge.ni.cu.la'ta. L. adj. geniculatus jointed.
Medium-sized rods, occurring singly, in
pairs and in chains. Motile, possessing 1 to
4 polar flagella. Gram-negative.
Gelatin colonies: Circular, whitish, as-
sume a greenish shimmer, translucent. Deep
colonies yellowish.
Gelatin stab: Infundibuliform liquefac
tion. Sediment light pink.
106
ORDER I. PSEUDOMONADALES
Agar slant: Grayish, glistening, translu-
cent, limited. Agar becomes brownish green.
Broth: Turbid, with slight gray pellicle
and sediment. Broth becomes green.
Litmus milk: Alkaline; reduction of lit-
mus; slight coagulation. Serum becomes
green.
Potato: Thin, brownish, moist, glistening,
viscid.
Indole not produced.
Aerobic, facultative.
Optimum temperature, between 20° and
25° C. No growth at 35° C.
Source: From water from the Schuylkill
River.
Habitat: Water.
16. Pseudomonas septica Bergey et al.,
1930. (Bacilhis fluorescens septicus Stutzer
and Wsorow, Cent. f. Bakt., II Abt., 71,
1927, 113; Bergey et al., Manual, 3rd ed.,
1930, 169.)
sep'ti.ca. Gr. adj. septicus putrefactive,
septic.
Rods, 0.6 to 0.8 by 0.8 to 2.0 microns, oc-
curring singly. Motile with a polar flagel-
lum. Gram-negative.
Gelatin stab: Infundibuliform liquefac-
tion.
Agar colonies: Circular with opalescent
center and transparent periphery.
Agar slant: Moderate, undulate margin.
Broth: Turbid with fragile pellicle, green-
ish in upper portion.
Litmus milk: Alkaline, coagulated.
Blood serum not liquefied.
Acid from glucose.
Aerobic, facultative.
Optimum temperature, 20° C.
Source: Isolated from diseased caterpil-
lars.
Habitat: From infected caterpillars so
far as known.
17. Pseudomonas syncyanea (Ehren-
berg, 1840). Migula, 1895. (Vibrio syncyaneus
Ehrenberg, Berichte ii.d. Verh. d. k. Preuss.
Akad. d. Wissensch. z. Berlin, 5, 1840, 202;
Migula, in Engler and Prantl, Die natiirl.
Pflanzenfam., 1, la, 1895, 29.)
syn.cy.a'ne.a. Gr. srjn- along w^ith, en-
tirely; Gr. cyaneus dark blue, dark; M.L.
adj. syncyaneus entirely blue.
Rods with rounded ends, occurring singly,
occasionally in chains, 0.7 by 2.0 to 4.0
microns. Motile with two to four polar fla-
gella. Gram-negative.
Gelatin colonies: Flat, bluish, translu-
cent.
Gelatin stab: Surface growth shiny,
grayish blue. The medium is colored steel-
blue with greenish fluorescence. Gelatin is
liquefied. Some strains do not liquefy.
Agar slant: Grayish white streak. The
medium takes on a bluish gray color with
slight fluorescence.
Broth: Turbid with marked fluorescence.
Litmus milk: Unchanged. In association
with lactic-acid bacteria the milk takes on a
deep blue color.
Potato: Yellowish gray, shiny layer, be-
coming bluish gray. The tissue becomes
bluish gray.
Indole not produced.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 25° C.
Source: From milk that was bluish in
color.
Habitat: The cause of blue milk.
18. Pseudomonas iodinum (Davis,
1939) Tobie, 1939. (Chromobacierium io-
dinum Davis, Zent. f. Bakt., II Abt., 100,
1939, 273; Tobie, Bull. Assoc. Diplomes
Microbiol. Fac, Nancy, No. 18, 1939, 16.)
i.o.di'num. M.L. neut.noun iodinum
iodine.
Rods, 0.5 by 1.0 to 2.0 microns, occurring
singly. Non-motile. Gram-negative.
Gelatin stab: Stratiform liquefaction.
Crystals of iodinin form.
Agar colonies: Round, smooth, gray-
white, moist, glistening. Dark purple crys-
tals having the appearance of iodine crystals
form in the growth and in the adjacent
medium. This pigment is actually a phena-
zine di-N-oxide, there being no iodine
present (Clemo and Mcllwain, Jour. Chem.
Soc, Pt. 1, 1938, 479; Clemo and Daglish,
Jour. Chem. Soc, Pt. 1, 1950, 1481).
Broth: Turbid. Crystals of iodinin form
on bottom of tube.
Litmus milk: Alkaline; slow reduction of
litmus.
FAMILY IV. PSEUDOMONADACEAE
107
Potato: Viscous, creamy, spreading, be-
coming dark.
Catalase-positive.
Indole not produced.
Nitrites produced from nitrates.
No acid from carbohj-drates.
Acetylmethylcarbinol not produced.
Aerobic.
Optimum temperature, 28° C. Grows at
37° C.
Distinguishing character: The pigment,
iodinin, is readih' formed in any medium
containing soluble nitrogenous compounds.
Potassium and sodium citrates markedly
stimulate pigment production. Yeast ex-
tract is inhibitory to formation of iodinin,
which is soluble in benzene, toluene, xylene,
chloroform, carbon disulfide and ethyl ace-
tate. Such solutions are ruby red. The pig-
ment, like pyocyanin and chlororaphine, is a
phenazine derivative.
Source: Isolated from milk.
Habitat: Unknown.
19. Pseudomonas smaragdina (Mez,
1898) Migula, 1900. {Bacillus smaragdinus
foetidus Reiman, Inaug. Dissertation,
Wiirzburg, 1887; Bacterium smaragdinum
Mez, Mikroskopische Wasseranalyse, Ber-
lin, 1898, 49; Migula, Svst. d. Bakt., £, 1900,
890.)
sma.rag'di.na. Gr. adj. smaragdinus of
smaragdus, emerald -green.
Small rods, occurring singly. Non -motile.
Gram-negative.
Gelatin colonies: Small, convex, irregular,
whitish with greenish shimmer.
Gelatin stab: Slight surface growth. In-
fundibuliform liquefaction. The liquefied
medium becomes light emerald-green in
color.
Agar colonies: Small, brownish yellow,
convex.
Agar slant: Abundant growth with green-
ish fluorescence.
Broth: Turbid.
Litmus milk: Not coagulated.
Potato: Dark brown, becoming chocolate-
brown.
Indole not produced.
Nitrates not reduced.
The cultures give off an odor resembling
jasmine.
Aerobic, facultative.
Optimum temperature, 37° C.
Subcutaneous and intravenous inocula-
tions into rabbits cause death in 36 to 48
hours.
Source: Isolated from nasal secretions in
ozena.
Habitat: Unknown.
20. Pseudomonas putida (Trevisan,
1889) Migula, 1895. {Bacillus fluorescens
putidus Fliigge, Die Mikroorganismen, 2
Aufl., 1886, 288; Bacillus putidus Trevisan,
I gen. e le specie d. Batteriacee, 1889, 18;
Migula, in Engler and Prantl, Die natiir.
Pflanzenfam., 1, la, 1895, 29.)
])u'ti.da. L. adj. putidus stinking, fetid.
Rods with rounded ends. Motile, posses-
sing polar flagella. Gram-negative.
Gelatin colonies: Small, finely granular,
fluorescent with dark center, surrounded by
a yellow zone, with pale gray margin.
Gelatin stab: Dirty white surface growth,
becoming greenish, fluorescent. No liquefac-
tion.
Agar colonies: Circular, raised, smooth,
amorphous, entire, with fluorescent zone
around the periphery.
Agar slant: Yellowish green layer, be-
coming fluorescent.
Broth: Turbid, fluorescent.
Litmus milk: Unchanged.
Potato: Thin, gray to brownish, slim}-
layer.
Cultures give off odor of trimethylamine.
Indole not produced.
Nitrites produced from nitrates.
Aerobic, facultative.
Temperature relations: Optimum, 25° C.
Will grow at 37° C. (Reid et al., Penn. Agr.
Exp. Sta., Bull. 422, 1942,9).
Relationship to other species: Identical
with Pseudomonas flxiorescens Migula ac-
cording to Lehmann and Neumann (Bact.
Diag., 1, Aufl., 2, 1896, 271) except that it
does not liquefy gelatin. See Pseudomonas
eisenbergii Migula.
Source: Isolated from putrid materials.
Habitat: Putrefying materials; water.
21. Pseudomonas striata Chester, 1901.
{Bacillus striatus viridis Ravenel, Memoirs
108
ORDER I. PSEUDOMONADALES
Nat. Acad. Sci., 8, 1896, 22; Chester, Man.
Determ. Bact., 1901, 325.)
stri.a'ta. L. v. strio to groove; L. part. adj.
striatus grooved.
Slender rods, of variable lengths, staining
irregularl}^ occurring singly and in pairs.
Motile, possessing polar flagella. Gram-nega-
tive.
Gelatin colonies: Circular, yellowish, with
filamentous border.
Gelatin stab: No liquefaction.
Agar slant: Smooth, glistening, irregular,
spreading. Agar becomes j-ellowish green.
Broth: Turbid, becoming slightly greenish.
Litmus milk: No coagulation; becoming
alkaline; litmus reduced.
Potato: Moist, glistening, spreading, be-
coming chocolate-brown.
Indole not produced.
Aerobic.
Grows well at 25° and 36° C.
Source: Isolated from soil.
Habitat: Soil.
22. Pseudomonas ovalis Chester, 1901.
(Bacillus fltmrescens ovalis Ravenel, Mem-
oirs Nat. Acad. Sci., 8, 1896, 9; Chester,
Man. Determ. Bact., 1901, 325.)
o.va'lis. L. n. ovum an egg; M.L. adj.
ovalis oval.
Rods, short with rounded ends, occurring
singly. Motile, possessing polar flagella.
Gram-negative.
Gelatin colonies: Irregular, lobate,
slightly granular, translucent, grayish be-
coming bluish.
Gelatin stab: No liquefaction. Faintly
green near surface.
Agar colonies: Circular, opaque, entire,
greenish fluorescence.
Agar slant: Thin, spreading, greenish
white. Agar becomes j'ellowish.
Broth: Turbid, with pellicle and white
sediment; faintly green.
Potato: Scant, yellowish brown growth.
Indole not produced.
Aerobic, facultative.
Grows well at 25° and 36° C.
Source: Isolated from soil.
Habitat: Soil.
and Anderson, Jour. Bact., 23, 1932, 343;
not Pseudomonas graveolens Migula, Syst. d.
Bakt., 2, 1900, 934.)
taet'ro.lens. L. adj. taeter offensive; L.
part, olens having an odor; M.L. part. adj.
taetrolens foul-smelling.
Short rods with rounded ends, occurring
singly, in pairs and in short chains. Motile
(Levine and Anderson). One to five polar
flagella (found on retest of cultures bj-
Haynes, 1953). Gram-negative.
Gelatin stab: Not liquefied.
Agar colonies: Circular, slightly raised,
smooth, entire, amorphous internal struc-
ture.
Agar slant: Growth abundant and tan-
colored; medium darkened. Penetrating
odor of must.
Broth: Turbid; thin, oily pellicle and
sediment. Odor of must.
Litmus milk: Acid, coagulated; litmus
reduced.
Indole not produced.
Nitrites not produced from nitrates.
Hj'drogen sulfide produced.
Starch not hydrolyzed.
Acid but no gas produced from glucose,
lactose, galactose, mannose, fructose,
rhamnose and xjdose. Slight acidity in
glycerol and mannitol. No acid or gas from
aesculin, amygdalin, arabitol, dextrin, dul-
citol, glycogen, inulin, maltose, melizitose,
pectin, raffinose, salicin, sorbitol, starch,
sucrose, xylan, arabinose, erythritol or
trehalose.
Aerobic.
Catalase-positive.
Optimum temperature, between 23° and
25° C. Scant growth at 33° and 10° C.
Distinctive character: A strong musty-
odor develops in media in which this or-
ganism grows. In this respect it resembles
P. perolens which, however, liquefies gelatin
and reduces nitrates.
Source: Isolated from musty eggs; also
from milk by Olsen and Hammer (Iowa
State College Jour. Sci., 9, 1934, 125).
Habitat : Found in various foods that have
a musty odor; presumably widely dis-
tributed.
23. Pseudomonas laelrolens Haynes,
7iom. nov. (Pseudomonas graveolens Levine
24. Pseudomonas incognita Chester,
1901. (Bacillus fluorescensincognitus'WTight,
FAMILY IV. PSEUDOMONADACEAE
109
Memoirs Nat. Acad. Sci., 7, 1895, 436;
Chester, Man. Determ. Bact., 1901, 323.)
in.cog'ni.ta. L. adj. incognitus not ex-
amined, unknown.
Short rods, with rounded ends, occurring
singly, in pairs and in chains. Motile, pos-
sessing a polar flagellum. Gram-negative.
Gelatin colonies: Thin, translucent,
slightly granular, becoming greenish. Mar-
gin undulate. The medium assumes a blue-
green fluorescence.
Gelatin stab: No liquefaction.
Agar slant: Thin, moist, translucent. Agar
becomes greenish.
Broth: Turbid, becoming greenish. Pel-
licle and whitish sediment form.
Litmus milk: Slightly acid in a month.
Litmus slowly reduced.
Potato: Moist, glistening, spreading,
brown.
Indole is produced (trace).
Aerobic, facultative.
No growth at 35° to 36° C.
Comment: Wright (op. cit., 1895, 441)
described an organism that is very similar
to this species except that it may produce a
faint brownish green coloration in a gelatin
stab; Wright named the organism Bacillus
nexibilis {Bacterium nexibilis Chester, Ann.
Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 74;
Pseudomonas nexibilis Chester, op. cit.,
1901, 309).
Source: Isolated from water from the
Schujdkill River.
Habitat: Water.
25. Pseudomonas rugosa (Wright, 1895)
Chester, 1901. {Bacillus rugosus Wright,
Memoirs Nat. Acad. Sci., 7, 1895, 438;
Chester, Man. Determ. Bact., 1901, 323.)
ru.go'sa. L. adj. rugosus full of wrinkles.
Small rods, with rounded ends, occurring
singl}^ in pairs and in chains. Motile, pos-
sessing 1 to 4 polar flagella. Gram-negative.
Gelatin colonies: Grayish, translucent,
slightly raised, irregular, sinuous, radiately
arose to entire.
Gelatin stab : Dense grayish green, lim-
ited, wrinkled, reticulate surface growth.
No liquefaction. Medium becomes green.
Agar slant: Grayish white, limited,
slightl}^ wrinkled, translucent. Agar be-
comes green.
Broth: Turbid, with thin whitish pellicle
and sediment.
Litmus milk: Acid, coagulated, partly re-
duced.
Potato: Moist, glistening, brown, spread-
ing.
Indole is produced (trace).
Aerobic.
Optimum temperature, 30° C. Does not
grow at 35° C.
Source: From water from the Schuylkill
River.
Habitat: Water.
26. Pseudomonas mildenbergii Bergey
et al., 1930. (Der Blaubacillus, Mildenberg,
Cent. f. Bakt., II Abt., 56, 1922, 309; Bergey
et al., Manual, 3rd ed., 1930, 172.)
mil.den.ber'gi.i. Mildenberg, a patro-
nymic; M.L. gen. noun mildenbergii of Mil-
denberg.
Rods, 0.3 to 0.5 by 1.0 to 3.5 microns,
with rounded ends, occurring singly. Mo-
tile, possessing polar flagella. Gram-nega-
tive.
Gelatin colonies: Circular, lobed, smooth,
glistening, slightly raised, steel-blue, entire.
Gelatin stab: No liquefaction.
Agar colonies: Small, circular, yellowish
or reddish yellow, entire, becoming lobed,
grayish green, iridescent. The medium be-
comes dirty grayish green.
Agar slant: Smooth, spreading, slimy,
glistening, grayish green to dark green,
fluorescent.
Broth: Turbid green, iridescent to opales-
cent with slimy sediment.
Litmus milk: Not coagulated, blue ring.
Potato: Slimj^, glistening, spreading, steel
blue.
Indole not produced.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 25° C.
Source : Isolated from air.
27. Pseudomonas eonvexa Chester,
1901. {Bacillus fluorescens convexus Wright,
Memoirs Nat. Acad. Sci., 7, 1895, 438;
Chester, Man. Determ. Bact., 1901, 325.)
con.vex'a. L. adj. convexus vaulted, con-
vex.
Short, thick rods, with rounded ends.
no
PSEUDOMONADALES
Motile, possessing a polar flagellum. Gram-
negative.
Gelatin colonies: Circular, convex, glis-
tening, greenish, translucent. The medium
becomes blue-green, fluorescent.
Gelatin stab: Light green, raised, glisten-
ing surface growth. No liquefaction. Me-
dium becomes blue-green fluorescent.
Agar slant: Moist, translucent, glisten-
ing, light greenish. The medium assumes a
greenish color.
Broth: Turbid, becoming greenish.
Litmus milk: No coagulation; alkaline.
Potato: Pale brown, spreading.
Indole not produced.
Aerobic, facultative.
Little or no growth at 35° to 36° C.
Source: From water from the Schuylkill
River.
Habitat: Water.
28. Pseudomonas eisenbergii Migula,
1900. (Fluorescirender Bacillus No. 18,
Eisenberg, Bakt. Diag., 1 Aufl., 1886, Taf.
7; Bacillus fluorescens non liquefaciens Eis-
enberg, Bakt. Diag., 3 Aufl., 1891, 145; Mig-
ula, Syst. d. Bakt., 3, 1900, 913; Pseud-
omonas non-liquefaciens Bergey et al..
Manual, 1st ed., 1923, 132.)
eis.en.ber'gi.i. Named for James Eisen-
berg, the bacteriologist who first described
this species; M.L. gen.n. eisenbergii of
Eisenberg.
Short, slender rods with rounded ends.
Non-motile. Gram-negative.
Gelatin colonies: Fern-like surface col-
onies. Medium around colonies has a pearly
luster.
Gelatin stab: Surface growth has fluores-
cent shimmer. Scant growth along stab. No
liquefaction.
Agar slant: Greenish growth.
Broth: Turbid, fluorescent.
Litmus milk: Unchanged.
Potato: Diffuse, brownish laj-er. The sur-
face acquires a grayish blue color.
Indole not produced.
Nitrites produced from nitrates.
Acid from glucose.
Blood serum liquefied.
Aerobic.
Optimum temperature, 25° C.
Not pathogenic.
Habitat : Water.
29. Pseudomonas erythra Fuller and
Norman, 1943. (Jour. Bact., 46, 1943, 276.)
e'ry.thra. Gr. adj. erythrus red.
Rods, 0.2 to 0.4 by 1.2 to 1.5 microns, with
rounded ends, usually arranged singly.
Motile with a single polar flagellum. Encap-
sulated. Gram-negative.
Gelatin stab: No growth.
Starch agar: No growth.
Water-insoluble dextrin agar: Scant
growth. Subsurface colonies appear after
8 to 10 days. Colonies are angular, small,
surrounded by a clear zone 2 to 5 mm in
diameter. Buff or reddish brown.
Litmus milk: No growth.
Indole not produced.
Nitrites not produced from nitrates.
Starch not hydrolyzed.
No growth in media containing the usual
carbohydrates. Cellulose and water-in-
soluble dextrins are utilized. Filter paper
strips in mineral solution develop reddish
brown spots above the surface of the liquid.
Solution becomes cloudy. Colonies enlarge
and become viscous, and the paper becomes
reddish. The filter paper does not break
with moderate shaking but may be wound
up in a slimy mass. In cellulose media a
reddish, water-soluble pigment is produced.
Yeast extract and nitrate are suitable
nitrogen sources.
Aerobic.
Grows in a range from 22° to 35° C.
Source: Isolated from soil.
Habitat: Soil.
30. Pseudomonas fragi (Eichholz, 1902)
Huss, 1907, emend. Hussong et al., 1937.
{Bacterium fragi Eichholz, Cent. f. Bakt.,
II Abt., 9, 1902, 425; Huss, Cent. f. Bakt.,
II Abt., 19, 1907, 661; Hussong, Long and
Hammer, Iowa Agr. Exp. Sta. Res. Bull.
225, 1937, 122.)
fra'gi. L. neut.n. fragum strawberry; L.
gen.n. fragi of the strawberry.
Description from Hussong, Long and
Hammer {loc. cit.).
Rods, 0.5 to 1.0 by 0.75 to 4.0 microns,
occurring singly, in pairs and in chains.
FAMILY IV. PSEUDOMONADACEAE
111
Motile with a polar flagellum. Gram-nega-
tive.
Gelatin: Crateriform to stratiform lique-
faction in 3 to 4 daj's.
Agar colonies: Convex, glistening, gen-
erally butyrous, occasionally viscid. Rough,
smooth and intermediate forms are recog-
nized in the description quoted. The rough
forms are less proteolytic and less active
in the hydrolysis of fats.
Agar slant: Growth abundant, spreading,
raised, white, shiny, generally butyrous.
Sweet ester-like odor resembling that of the
flower of the May apple.
Broth: Turbidit}^ and sediment with a
thin pellicle.
Litmus milk: Acid ring followed by acid
coagulum at surface. Complete coagulation
in 2 to 3 weeks, some digestion. Characteris-
tic Maj'-apple or strawberry odor.
Potato : Growth echinulate to arborescent,
raised, glistening, white, becoming brown-
ish.
Indole not produced.
Nitrites not produced from nitrates.
Ammonia produced from peptone.
Hydrogen sulfide not produced.
Acid from glucose and galactose, some-
times arabinose. No acid from glycerol,
inulin, lactose, fructose, maltose, mannitol,
raffinose, salicin or sucrose.
No acetylmeth3-lcarbinol produced.
Fat is generally hydrolyzed (Nashif and
Nelson, Jour. Dairy Sci., 36, 1953, 459-488).
Aerobic.
Grows from 10° to 30° C. No growth at
37° C. Very sensitive to heat.
Comment: Various names have been
given this species. Hussong (Thesis, Iowa
State College, 1932) thinks that these varie-
ties are the result of dissociative action.
Source: Isolated from milk and other
dairy products, dairy utensils, water, etc.
Habitat: Soil and water. Widely dis-
tributed (Morrison and Hammer, Jour.
Dairy Sci., ^4, 1941,9).
31. Pseudomonas perolens (Turner,
1927) Szybalski, 1950. {Achromobacter pero-
lens Turner, Austral. Jour. Exp. Biol, and
Med. Sci., 4, 1927, 57; Szybalski, Nature,
165, 1950, 733.)
pe.ro 'lens. L. v. perolere to emit a pene-
trating odor; L. part. adj. perolens emitting
an odor.
Small, imperfect spheres and coccoid rods;
occasionallj' longer rods with rounded ends;
occur singly and in short chains. 0.3 by 0.4
to 2.55 microns. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied.
Agar slants: Growth moderate, glisten-
ing, raised, butyrous, spreading, with un-
dulate border; whitish by reflected and
semi-translucent by transmitted light.
Broth: Turbid, with a flocculent sediment
and a slight pellicle.
Litmus milk: Acid, gradually decolorized,
partial clotting.
Blood serum: Liquefied.
Potato: Growth thick, glistening, raised,
brownish.
Nitrites and ammonia produced from ni-
trates.
Indole not produced.
Acid but no gas from glucose, fructose,
galactose, glycerol, mannitol and arabinose.
Sucrose, maltose, lactose, raffinose, dulcitol,
salicin and inulin not utilized.
Aerobic, facultative.
Grows well at room temperature. No
growth at 37° C.
Distinctive characters : Produces a musty
odor in eggs. Other varieties and species of
Pseudomonas that produce the same odor
have been described (Szybalski, loc. cit.).
Resembles Pseudomonas fragi but produces
a musty rather than a May-apple odor in
media.
Source: Isolated from eggs with a musty
odor.
Habitat: Musty eggs.
32. Pseudomonas mephitica Claydon
and Hammer, 1939. (Jour. Bact., 37, 1939,
254.)
me.phi'ti.ca. L. adj. mephiticus pestilen-
tial (skunk-like) odor.
Rods, 0.5 to 1.0 by 1.5 to 14.0 microns,
occurring singly, in pairs and in chains.
Actively motile with a polar flagellum.
Gram-negative.
Gelatin: Slow liquefaction.
Agar colonies: Convex, circular, about 3
112
ORDER I. PSEUDOMONADALES
mm in diameter, shiny, grajish white, en-
tire, of the consistency of bread dough.
Agar slant: Growth grayish white,
wrinkled, echinulate. After 1 or 2 days a
skunk-like odor develops.
Broth: Turbid. Sediment. White pellicle.
Potato: Growth echinulate, shiny, brown-
ish.
Litmus milk: A skunk-like odor develops
in 1 to 2 days. Grayish blue surface ring in
about 3 days. Alkaline in 7 to 10 daj^s. In
two weeks complete reduction. Slight pro-
teolysis and viscosity.
Hydrogen sulfide not produced.
Indole not produced.
Nitrites produced from nitrates.
Acid but no gas produced slowly from
glucose, fructose, maltose and sucrose. No
acid from arabinose, dextrin, galactose,
glycerol, lactose, mannitol, raffinose or
salicin.
Aerobic, facultative.
Optimum temperature, 21° C. Growth
slight at 5° and 30°C. No growth at 37° C.
Source: Several cultures isolated from
butter having a skunk-like odor.
Habitat: Presumably derived from the
rinse water.
33. Pseudomonas putrefaciens (Derby
and Hammer, 1931) Long and Hammer,
1941. (Achromobacter putrefaciens Derby
and Hammer, Iowa Agr. Exp. Sta., Res.
Bull. 145, 1931, 401; Long and Hammer,
Jour. Bact., 41, 1941, 100.)
pu.tre.fa'ci.ens. L. v. ptdrefacio to make
rotten; L. part. adj. putrefaciens making
rotten.
Rods, 0.5 to 1.0 by 1.1 to 4.0 microns,
occurring singly and in pairs. Motile with
a single flagellum. Gram-negative.
Gelatin stab: Rapid, saccate to strati-
form liquefaction, with reddish brown sedi-
ment in the liquefied portion.
Agar colony: Circular, smooth, glisten-
ing, slightly raised, somewhat transparent,
with brownish tinge.
Agar slant: Echinulate, slightly reddish
brown, viscous.
Broth: Turbid, with thin, gray pellicle
and reddish brown sediment.
Litmus milk: Rapid reduction and pro-
teolysis with odor of putrefaction.
Potato: Echinulate, smooth, glistening,
viscous, reddish brown.
Indole not produced.
Nitrites are produced from nitrates.
Acid from maltose and sucrose. No action
on glucose, fructose, galactose, arabinose,
lactose, raffinose, dextrin, inulin, salicin,
amj'gdalin, glycerol, mannitol or sorbitol.
Ammonia is formed.
Aerobic, facultative.
Optimum temperature, 21° C. No growth
at 37° C.
Source: Isolated from tainted butter.
Habitat: Milk, cream, butter, water, soil
and creamery equipment (Long and Ham-
mer, loc. cit.; Claj'don and Hammer, op.
cit., Res. Bull. 267^ 1939).
34. Pseudomonas cohaerens (Wright,
1895) Chester, 1901. {Bacillus cohaerens
Wright, Mem. Nat. Acad. Sci., 7, 1895, 464;
Pseudomonas cohaerea (sic) Chester, Man.
Determ. Bact., 1901, 312.)
co.hae'rens. L. part. adj. cohaerens co-
hering, uniting together.
Rods, occurring singly and in pairs, some-
times in chains. Motile, possessing a polar
flagellum. Gram-negative.
Gelatin colonies : Circular, elevated, gray-
ish, translucent, entire. Become white with
an elevated, brownish, central nodule.
Gelatin stab: Slow liquefaction.
Agar slant: Elevated, grayish white,
translucent, glistening, with irregular mar-
gins.
Broth: Turbid; coherent, wrinkled pel-
licle which adheres to the walls of the con-
tainer.
Litmus milk: Alkaline, coagulated, slowly
peptonized, litmus reduced.
Potato: Thick, granular, translucent,
spreading.
Indole not produced.
Grows at 25° C.
Aerobic.
Source: Isolated from water from the
Schuylkill River.
Habitat: Water.
35. Pseudomonas ambigua (Wright,
1895) Chester, 1901. (Bacillus amhiguus
Wright, Memoirs Nat. Acad. Sci., 7, 1895,
FAMILY IV. PSEUDOMONADACEAE
113
439; Chester, Man. Determ. Bact., 1901,
308.)
am.bi'gu.a. L. adj. anibiguus going
about, hence uncertain.
Small rods, with rounded ends, occurring
singly, in pairs and in chains. Motile, pos-
sessing a polar flagellum. Gram-negative.
Gelatin colonies: Gray, translucent,
slightly raised, irregular, radiate, with
transparent margin.
Gelatin stab: No liquefaction.
Agar slant: Graj^, limited, entire.
Broth: Turbid, with gray sediment.
Litmus milk: Acid, slowly coagulated.
Litmus reduced.
Potato: Gray to creamj-, viscid, spread-
ing.
Indole produced.
Aerobic, facultative.
Optimum temperature, between 30° and
35° C.
Source: Isolated from water from the
Schuylkill River.
Habitat: Water.
36. Pseudonionas oleovorans Lee and
Chandler, 1941. (Jour. Bact., 4^, 1941, 378.)
o.le.o'vor.ans. L. ole^^moil■, L. v. voro to
destro}', consume; M.L. part. adj. oleovorans
oil-consuming.
Short rods, 0.5 by 0.8 to 1.5 microns,
occurring singly and in pairs. Motile. Gram-
negative.
Gelatin stab: No liquefaction after 6
weeks.
Gelatin colonies : Up to 1 mm in diameter,
fluorescent; similar to agar colonies.
Surface agar colonies: After 24 hours 1 to
2 mm in diameter, smooth, convex, shiny,
opaque, creamy, fluorescent bj^ transmitted
light. Edge entire in young colonies.
Deep agar colonies: 0.5 by 1.0 to 1.5 mm,
lens-shaped, buff-colored, not fluorescent.
Agar slant: Growth raised, smooth, fluo-
rescent, edge erose.
Broth : After 24 hours, moderate turbidity
with slight yellowish, viscid sediment. No
pellicle or ring. No soluble pigment pro-
duced.
Litmus milk: No change.
Indole not produced.
Potato: Good growth.
Nitrites are produced from nitrates.
Starch is hydrol3'zed.
No acid from glucose, lactose, sucrose,
galactose, xylose, mannitol, salicin or
glycerol.
Equally good growth at 25° and 37° C.
Aerobic.
Distinctive character: The fluorescent
quality of the colonies is not imparted to
any of the artificial media used.
Source: Isolated from cutting compound
(oil-water emulsion) circulating in a ma-
chine shop. The oil in this compound may
be utilized as a sole source of energj'.
Habitat: Probably oil-soaked soils. Abun-
dant in cutting compounds.
37. Pseudonionas arvilla Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 90.)
ar.vil'.la. L. arvum a field; M.L. dim. noun
arvilla a small field.
Rods, 0.5 to 0.7 by 2.0 to 3.0 microns.
Motile with one to five polar flagella. Gram-
negative.
Gelatin colonies: Circular, whitish, con-
vex, smooth, glistening, lobate.
Gelatin stab: No liquefaction.
Agar colonies: Circular or amoeboid,
white to buff, flat to convex, smooth, glis-
tening, opaque, entire.
Agar slant: Filiform, whitish, convex,
smooth, ringed, entire.
Broth: Turbid.
Nitrites not produced from nitrates.
Starch not hydrolj'zed.
Acid from glucose.
Attacks naphthalene.
Aerobic, facultative.
Grows at 37° C.
Source: Isolated from soil.
Habitat: Soil.
38. Pseudomonas daciinhae Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 90.)
da.cun'hae. d'Acunha, place name. Island;
M.L. gen. noun dacimhae of d'Acunha.
Rods 0.5 to 0.8 by 1.5 to 3.0 microns.
Motile with one to six polar flagella. Gram-
negative.
Gelatin colonies: Circular, whitish,
raised, smooth, glistening, entire.
Gelatin stab: No liquefaction.
114
ORDER I. PSEUDOMONADALES
Agar colonies: Circular to amoeboid,
white, flat, glistening, opaque, entire.
Agar slant: Filiform, pale buff, raised,
smooth, glistening, undulate.
Broth: Turbid.
Nitrites not produced from nitrates.
Starch not hydrolyzed.
No acid from carbohydrate media.
Attacks phenol.
Aerobic, facultative.
Grows at 37° C.
Source: Isolated from soil.
Habitat: Soil.
39. Pseudomonas desmolytica Gray
and Thornton, 1928. (Cent. f. Bakt., II Abt.,
73, 1928, 90.)
des.mo.ly'ti.ca. Gr. desmus bond; Gr.
lyticus able to loose; M.L. adj. desmolyiiciis
bond-loosening.
Rods, 0.7 to 0.8 by 2.0 to 3.0 microns,
occurring singly and in pairs. Motile, with
one to five polar flagella. Gram-negative.
Gelatin colonies: Circular, gray to buff,
raised or umbonate. Smooth, glistening,
entire.
Gelatin stab: No liquefaction.
Agar colonies: Circular or amoeboid,
whitish, flat or convex, smooth, translucent
to opaque, entire.
Agar slant: Filiform, pale buff, raised,
smooth, undulate.
Broth: Turbid.
Nitrites often produced from nitrates.
Starch not hydrolyzed.
Acid usually produced from glucose.
Attacks naphthalene.
Aerobic, facultative.
Grows at 35° C.
Source: Isolated from soil.
Habitat: Soil.
40. Pseudomonas rathonis Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 90.)
ra.tho'nis. RathoPark, place name; M.L.
gen. noun rathonis of Ratho.
Small rods, 0.5 to 1.0 by 1.0 to 3.0 microns,
occurring singly and in pairs. Motile, with
polar flagella. Gram-negative.
Gelatin colonies: Circular, white, raised,
smooth, glistening, undulate.
Gelatin stab: No liquefaction.
Agar colonies : Circular, buff, flat, smooth,
glistening, entire.
Agar slant: Filiform, pale buff, convex,
smooth, glistening, undulate.
Broth: Turbid; pellicle may form.
Nitrites may be produced from nitrates.
Starch may be hydrolyzed.
Acid may be produced from glucose and
glycerol.
Attacks phenol and cresol at times, also
naphthalene.
Aerobic, facultative.
Grows at 35° C.
Source: Isolated from manure and soil.
Habitat: Manure and soil.
41. Pseudomonas salopia Gray and
Thornton, 1928. (Pseudomonas salopium
(sic) Gray and Thornton, Cent. f. Bakt., II
Abt., 73, 1928, 91.)
sa.lo'pi.a. Med.L. Salop Shropshire;
M.L. adj. salopius of Shropshire.
Rods, 0.7 to 1.0 by 1.0 to 3.0 microns,
occurring singly and in pairs. Motile with
one to six polar flagella. Gram-negative.
Gelatin colonies: Circular, grayish buff',
flat, rugose or ringed, translucent border.
Gelatin stab: No liquefaction.
Agar colonies: Circular or amoeboid,
white to buff, flat to convex, smooth, glis-
tening, translucent border, entire.
Agar slant: Filiform, whitish, raised,
smooth, glistening, lobate.
Broth: Turbid with pellicle.
Nitrites not produced from nitrates.
Starch not hydrolyzed.
Acid from glucose and sucrose.
Attacks naphthalene.
Aerobic, facultative.
Grows at 35° C.
Source: Isolated from soil.
Habitat: Soil.
42. Pseudomonas cruciviae Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 91.)
cru.ci'vi.ae. L. crux, cruets a cross; L.
via a way; M.L. Crucivia Waycross, a place
name.
Rods, 1.0 by 1.0 to 3.0 microns, occurring
singly and in pairs. Motile with one to five
polar flagella. Gram-negative.
FAMILY IV. PSEUDOMONADACEAE
115
Gelatin colonies: Circular, white, convex,
smooth, undulate.
Gelatin stab: No liquefaction.
Agar colonies: Circular or amoeboid,
white to buff, flat to convex, smooth, entire.
Agar slant: Filiform, pale buff, raised,
smooth, undulate.
Broth: Turbid.
Nitrites not produced from nitrates.
Starch not hj^droh-zed.
No acid in carbohydrate media.
Attacks phenol and m-cresol.
Aerobic, facultative.
Optimum temperature, between 30° and
35° C.
Source: Isolated from soil.
Habitat: Soil.
43. Pseudonionas stutzeri (Lehmann
and Neumann, 1896) Kluyver, 1942.
{Bacillus denitrificans II Burri and Stutzer,
Cent. f. Bakt., II Abt., 1, 1895, 392; Bac-
terium stutzeri Lehmann and Neumann,
Bakt. Diag., 1 Aufl., 2, 1896, 237; Bacillus
nitrogenes Migula, Syst. d. Bakt., 2, 1900,
793; Pseudonionas stutzeri Kluyver, in
Koningsberger, Leerb. d. algem. Plant-
kunde, Scheltema and Holkema, Amster-
dam, 2, 1942, 198; not Pseudonionas stutzeri
Migula, Syst. d. Bakt., 2, 1900, 929.)
stut'ze.ri. Named for Dr. A. Stutzer, one
of the bacteriologists who originally de-
scribed this species; M.L. gen. noun stutzeri
of Stutzer.
Description taken from van Niel and
Allen (Jour. Bact., 64, 1952, 421).
Rods, 0.5 to 0.8 by 1.0 to 3.0 microns. Mo-
tile, possessing a single polar flagellum.
Gram-negative.
Gelatin and agar colonies: Strongly co-
herent to media, dry consistency later be-
coming mucoid, resemble craters with ele-
vated ridges which often branch and merge,
concentric zones, polygonal elements,
granular.
Gelatin: No liquefaction.
Peptone and yeast agar: Good growth.
Broth: Surface film on nitrate- or nitrite-
free media which readily breaks up and pre-
cipitates.
Potato: Luxuriant, wrinkled, slimy, flesh-
to peach-colored growth.
Nitrates, nitrites, nitramines and N2O
reduced to elemental nitrogen.
Carbohj^drates : No growth when used as
a carbon source in mineral media.
Aerobic, facultative.
Optimum pH, 7.0; growth even at pH 9.0.
Optimum temperature, 35° C.
Distinctive characters: Colony shape
and consistency, mode and color of potato
growth, ability to grow anaerobically in
media with nitrate, nitrite, nitramine or
N2O, producing foam.
Source: Isolated from soil.
Habitat: Found widely distributed in soil,
manure, mud and stagnant water.
44. Pseudomoiias tralucida Kellerman
et al., 1913. (Kellerman, McBeth, Scales
and Smith, Cent. f. Bakt., II Abt., 39, 1913,
37.)
tra.lu'ci.da. L. adj. frahicidus trans-
parent.
Rods 0.6 by 1.2 microns. Motile with one
or two polar flagella. Gram-negative.
Gelatin stab: No liquefaction.
Agar slant: Moderate, flat, glistening,
grayish growth.
Broth: Turbid; granular sediment.
Litmus milk: Acid, no coagulation.
Potato: No growth.
Indole not produced.
Nitrites produced from nitrates.
Ammonia not produced.
Starch hydrolysis slight.
Acid from glucose, maltose, lactose, su-
crose, starch, glycerol and mannitol.
Attacks cellulose.
Aerobic, facultative.
Optimum temperature, 37° C. Grows also
at 20° C.
Habitat: Soil.
45. Pseudonionas lasia Fuller and Nor-
man, 1943. (Jour. Bact., 46, 1943, 275.)
la'si.a. Gr. adj. lasius hairy, rough,
shaggy, woolly.
Rods, 0.5 to 0.6 by 1.2 to 2.0 microns,
usually occurring singly but sometimes in
chains. Motile with a single polar flagellum.
Gram-negative.
Gelatin stab: No liquefaction.
Starch agar colonies : Convex, pale 3'ellow,
becoming cream color, entire, round. Sub-
116
ORDER I. PSEUDOMONADALES
surface colonies look like small, woolly
balls.
Water-insoluble dextrin colonies: Col-
onies grow below the surface and have a
woolly appearance. Colonies are surrounded
by clear zones. Become cream to pale yellow
in color.
Litmus milk : Unchanged except for reduc-
tion of litmus at bottom of the tube.
Indole not produced.
Nitrites produced from nitrates.
Starch hydrolyzed.
Glucose, .xylose, maltose and starch read-
ily utilized. Arabinose, galactose and gum
arable feebly attacked. No acid formed in
any of the above-mentioned substrates.
Cellulose, cellulosan, water-soluble and
water-insoluble cellulose, dextrins, hemi-
cellulose and pectin readily attacked. Filter
paper strips become pale yellowish in the
area attacked.
Peptone, yeast extract, nitrate and am-
monia are suitable nitrogen sources.
Aerobic.
Grows between 22° and 35° C.
Source: Isolated from soil.
Habitat: Soil.
46. Pseudonionas riboflavina Foster,
1944. (Pseudotnonas riboflavinus (sic) Foster,
Jour.Bact.,47, 1944,27; also see Jour. Bact.,
48, 1944, 97.)
ri.bo.fla'vi.na. M.L. adj. riboflavinus
pertaining to riboflavin.
Thin rods of variable length. Motile.
Gram-negative.
Gelatin stab: No liquefaction.
Yeast-extract agar colonies: Small, con-
vex, smooth, transparent; slightly dentate
edges. If glucose is added to the agar,
copious quantities of polj'saccharides are
formed. Presence of fructose, mannitol, su-
crose, maltose, lactose, xylose and galactose
also lead to polysaccharide formation.
Yeast-extract glucose broth: Becomes so
viscid that it scarcely flows.
Milk: Soft curd forms. Slowly peptonized.
Nitrites produced from nitrates.
No acid or gas from fructose, mannitol,
sucrose, maltose, lactose, xylose or galac-
tose. Acetic acid oxidized.
Acetylmethylcarbinol not produced.
Urea, glycine, ammonium chloride or
sodium nitrate cannot be used as substi-
tutes for organic-nitrogen sources. Neither
could 20 water-soluble accessory factors
substitute for yeast extract in a synthetic
mineral salts-glucose medium.
No pigment produced in any medium.
Starch not hydrolyzed.
Optimum temperature, between 30° and
33° C.
Distinctive characters: In organic media
containing a small amount of organic matter
such as yeast extract or peptone and 0.05 to
0.2 per cent riboflavin, the riboflavin is
attacked and converted to lumichrome,
which accumulates in the culture as lemon-
yellow crystals. If riboflavin is not provided
in the medium, appreciable quantities of it
are synthesized by this organism.
Source: Isolated from soil rich in ribo-
flavin.
Habitat: Unknown.
47. Pseudomonas denitrificans Bergey
et al., 1923. (Bacillus denitrificans fluorescens
Christensen, Cent. f. Bakt., II Abt., 11,
1903, 190; Bergey et al., Manual, 1st ed.,
1923, 131.)
de.ni.tri'fi.cans. L. de away, from; L.
nitruni soda; M.L. nitrate, niter; M.L.
denitrifico to denitrify; M.L. part. adj.
denitrificans denitrifying.
Rods, 0.5 to 0.7 by 0.5 to 1.25 microns,
occurring singly and in pairs in large, slimy
masses. Motile. Gram-negative.
Gelatin colonies: Small, circular, con-
toured, raised, moist, pearly gray, glisten-
ing.
Gelatin stab: Whitish, lobed surface
growth. Yellowish green growth in stab. No
liquefaction.
Agar colonies: Pearly white, circular,
entire.
Agar slant: Broad, whitish, contoured,
moist, entire.
Broth: Turbid, with thick, wrinkled
pellicle.
Litmus milk: Not coagulated.
Potato: Reddish gray layer.
Indole not produced.
Nitrates reduced with production of
nitrogen.
Aerobic, facultative.
Optimum temperature, 25° C.
FAMILY IV. PSEUDOMONADACEAE
117
Source: Isolated from soil.
Habitat: Soil.
48. Pseudomonas indoloxidans Gray,
1928. (Proc. Roy. Soc. London, B, 102, 1928,
263.)
in.dol.o'xi.dans. M.L. neiit.n. indolum
indole; M.L. part. adj. oxidans oxidizing;
from Gr. adj. oxys sharp, acid; M.L. part,
adj. indoloxidans indole-oxidizing.
Rods 1.0 by 3.0 microns. Motile with one
to four polar flagella. Gram-negative
Gelatin colonies: Round, convex, buff,
smooth, glistening, erose.
Gelatin stab: No liquefaction.
Agar colonies: Round, convex, white,
watery; transparent border, erose.
Agar slant: Filiform, convex, whitish,
smooth, glistening, undulate.
Broth: Cloudy.
Indole not produced.
Nitrites produced from nitrates. No gas.
No acid or gas from glucose, sucrose, lac-
tose, maltose or glycerol.
Starch not hydrolyzed.
Phenol and m-cresol not attacked.
Distinctive character: Indole decomposed
in mineral salts agar medium with the for-
mation of blue crystals of indigotin.
Aerobic.
Optimum temperature, between 25° and
28° C.
Source: Isolated from soil from Italian
Tyrol.
Habitat: Soil.
49. Pseudomonas niira McBeth, 1916.
(Soil Science, 1, 1916, 467.)
mi'ra. L. adj. minis extraordinary.
Rods 0.4 by 1.6 microns. Motile with a
single polar flagellum. Gram-negative.
Gelatin stab: Good growth. No liquefac-
tion.
Agar colonies : Circular, convex, grayish
white, granular, lacerate.
Agar slant: Moderate, flat, grayish white,
somewhat iridescent.
Broth: Turbid.
Litmus milk: Alkaline.
Potato : Moderate, grayish white, leathery
growth.
Indole not produced.
Nitrites produced from nitrates.
Ammonia is produced.
No acid from glucose, maltose, lactose,
sucrose, starch, glycerol or mannitol.
Cellulose decomposed. Pllter paper strips
disintegrated at surface of liquid medium.
Aerobic, facultative.
Optimum temperature, 20° C.
Source: Isolated from soil.
Habitat: Soil.
50. Pseudomonas nigrifaciens White,
1940. (Scientific Agriculture, 20, 1940, 643.)
ni.gri.fa'ci.ens. L. niger black; L. v.
Jacio to make; M.L. part. adj. nigrofaciens
blackening.
Rods, 0.5 by 1.5 to 2.0 microns, occurring
singly or in pairs and having rounded ends.
Actively motile with a single polar flagel-
lum. Gram-negative.
Gelatin stab: Pigmented surface growth
after 24 hours. Slight crateriform liquefac-
tion changing to saccate.
Agar colonies: Circular, convex, smooth,
glistening, entire, 2 to 4 mm in diameter.
Slight fluorescence in early stages. The me-
dium assumes a brownish color.
Agar slant: Growth filiform, smooth,
moist, glistening, with blackish pigmenta-
tion at 4° and 15° C. in 48 hours, the medium
turning brownish. Slight fluorescence in
early stages.
Broth: Turbid after 24 hours. After 5 to 6
days a black ring and then a pellicle forms,
later a black sediment. Medium turns
brown.
Litmus milk: A black ring appears after 3
days at 15° C. followed by a pellicle. Litmus
is reduced. Alkaline reaction. No coagula-
tion. Digested with a putrid odor.
Potato: No growth, even in presence of
1.5 per cent salt.
Nitrites not produced from nitrates in 7
days. No gas produced.
Starch is hydrolyzed. Natural fats not
hydrolj'zed.
Alkaline reaction produced in sucrose,
maltose, lactose, glucose, mannitol and
raffinose broth (pH 8.2). No gas produced.
Ammonia produced in peptone broth.
Aerobic.
Optimum pH, 6.8 to 8.4.
Temperature relations: Minimum, 4° C.
Optimum, 25° C. Maximum, 33° to 35° C.
118
ORDER I. PSEUDOMONADALES
Distinctive characters : No or slow growth
in culture media in the absence of salt.
Maximum growth and pigmentation ap-
pears with 1.5 and 2.5 per cent salt. Opti-
mum pigmentation occurs at 4° and 15° C.
Pigment insoluble in chloroform.
Source: Several cultures isolated from
samples of discolored butter.
Habitat : Causes a black to reddish brown
discoloration of print butter. Evidently
widely distributed in nature.
51. Pseudonionas ichthyodermis (Wells
and ZoBell, 1934) ZoBell and Upham, 1944.
(Achromobacter ichthyodermis (sic) Wells and
ZoBell, Proc. Nat. Acad. Sci., £0, 1934, 123;
ZoBell and Upham, Bull. Scripps Inst.
Oceanography, 5, 1944, 246 and 253.)
ich.thy.o.der'mis. Gr. ichthys fish; Gr.
derma skin; M.L. ichthyodermis fish skin.
Small rods, 0.9 to 1.3 by 3.0 to 5.0 microns,
occurring singlj^ and in pairs. Motile, with
a tuft of polar flagella. Pleomorphic forms
predominate in old cultures. Encapsulated.
Gram-negative.
Requires sea water following initial iso-
lation. The following differential media are
prepared with sea water:
Agar colonies: Glistening, colorless, con-
vex, circular, 2 to 4 mm in diameter.
Agar slants: Abundant, filiform, raised,
smooth, opalescent growth.
Gelatin tube : Rapid crateriform liquefac-
tion complete in 5 days at 18° C.
Sea-water broth: Turbidity, with pellicle,
little granular sediment and no odor.
Milk: No growth.
Potato : No growth unless dialyzed in sea
water. Then fair growth with no pigment.
Acid from glucose, sucrose and mannitol
but not from lactose or glycerol.
Starch hydrolyzed.
Ammonia liberated from peptone, but no
hydrogen sulfide produced.
Indole formed in tryptophane sea-water
broth.
Nitrites produced from nitrates.
Optimum temperature, between 25° and
30° C.; 37° C. incubation will kill recently
isolated organisms.
Aerobic, facultative.
Source: Isolated from diseased kilifish
{Fundnlus parvipinnis) .
Habitat: Skin lesions and muscle tissue of
infected marine fish.
52. Pseudonionas niarinoglutinosa
(ZoBell and Allen, 1935) ZoBell, 1943.
{Achromobacter viarinoglutinosus (sic) Zo-
Bell and Allen, Jour. Bact., 29, 1935, 246;
ZoBell, Jour. Bact., ^6, 1943, 45.)
ma.ri.no.glu.ti.no'sa. L. marinus ma-
rine; L. glutinosus full of glue, viscous; M.L.
adj. marinoglutinosus . Meaning obscure.
Short rods, 0.7 to 1.0 by 1.8 to 2.4 microns,
with rounded ends, occurring singly, in pairs
and in clumps. Motile with polar flagella.
Staining granular. Encapsulated. Gram-
negative.
Gelatin stab: Moderate filiform growth f;
with slight napiform liquefaction. No pig-
ment.
Agar colonies: Round with concentric
circles and crinkled radial lines, 1.5 to 5.0
mm in diameter. No pigment.
Agar slant: Moderate, filiform, flat.
Butyrous consistency.
Broth: Moderate clouding, marked ring,
adherent film of growth on test tube wall,
flaky sediment.
Milk: No growth.
Potato: No growth.
Indole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide and ammonia produced
from Bacto-tryptone.
Acid but no gas from xylose and dextrin.
No acid from glucose, lactose, sucrose or
mannitol.
Starch is hydrolj'zed.
Optimum temperature, between 20° and
25° C.
Aerobic, facultative.
Source : Isolated from sea water.
Habitat: Sea water.
53. Pseudonionas nienibranoforniis
(ZoBell and Allen, 1935) ZoBell, 1943.
{Achromobacter membranoformis ZoBell and
Allen, Jour. Bact., £9, 1935, 246; ZoBell,
Jour. Bact., 46, 1943, 45.)
mem. bra. no. for 'mis. L. membrana a
membrane; L. forma appearance; M.L. adj.
membranoform is membranous .
Reds, 0.9 to 1.2 by 3.5 to 4.8 microns, oc-
curring singly and in pairs. Motile with
FAMILY IV. PSEUDOMONADACEAE
119
lophotrichous flagella. Encapsulated. Gram-
negative.
Gelatin stab : Growth filiform, best at top,
1 with slow crateriform liquefaction.
Agar colonies: Circular, 1.0 to 2.5 mm,
with crinkled surface.
Agar slant: Moderate, beaded, raised
growth. ]\Iembranous consistency. Becomes
browned with age.
Broth: Slight turbidity, flocculent sedi-
ment, film of growth on walls of test tube.
IMilk: No growth.
Potato: No growth.
Indole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide not produced.
Acid but no gas from glucose, sucrose,
dextrin and mannitol. No acid from lactose
or xylose.
Starch not hydrolyzed.
Optimum temperature, between 20° and
25° C.
Aerobic.
Source: Isolated from sea water.
Habitat: Sea water.
54. Pseudonionas gelatica (Gran, 1902)
Bergey et al., 1930. {Bacillus gelaticus Gran,
Bergens Museums Aarbog., 1902, 14; Bergey
et al.. Manual, 3rd ed., 1930, 175.)
ge.la'ti.ca. L. part. adj. gelatns frozen,
congealed, jellied; M.L. adj. gelaticus re-
sembling hardened gelatin.
Rods, with rounded ends, 0.6 to 1.2 by 1.2
to 2.6 microns, occurring singly, in pairs,
and sometimes in short chains. Motile, type
of flagellation not recorded. Gram -negative.
All media prepared with 3 per cent salt.
Fish-gelatin colonies: Circular, trans-
parent, glistening, becoming brownish in
color.
Fish-gelatin stab: Liquefaction infun-
dibuliform. Two varieties are recognized:
one produces a green fluorescence; the other
does not produce a water-soluble pigment.
Sea-weed agar colonies: Circular, flat,
entire, glistening, grayish blue center with
reddish brown periphery. Liquefied.
Fish-agar slant: Flat, transparent streak,
with undulate margin, reddish brown to
grayish white.
Broth: Turbid with flocculent pellicle
and grayish yellow sediment, viscid.
Indole not produced.
Nitrites not produced from nitrates.
Starch hj^drolyzed.
No action on sugars.
Aerobic, facultative.
Temperature relations : Optimum temper-
ature, between 20° and 25° C. Maximum,
between 30° and 32° C. Minimum, 0° C.
Distinctive character: Requires 3 to 4 per
cent salt for growth.
Source : Isolated from sea water from the
Norwegian coast.
Habitat: Probably associated with the de-
composition of algae in coastal waters.
55. Pseudomonas calcis (Drew, 1912)
Kellerman and Smith, 1914. (Bacterium
calcis Drew, Yearbook Carnegie Inst.
Wash., 11, 1912, 136; Kellerman and Smith,
Jour. Wash. Acad. Sci., 4, 1914, 400.)
cal'cis. L. fem.noun calx, calcis limestone,
chalk; L. calcis of limestone.
Rods, 1.1 by 1.5 to 3.0 microns, usually
single but may form long chains. Actively
motile with one polar flagellum. Gram-
negative.
Grows best in sea water or 3 per cent salt
media. Deposits CaCOs .
Agar colonies: Circular, with finely ir-
regular outline, granular appearance, ele-
vated, spreading; old colonies having
brownish tinge in center.
Gelatin stab: Infundibuliform liquefac-
tion.
Gelatin colonies : Small, with liquefaction.
Broth: Good growth especially in pres-
ence of potassium nitrate, peptone or cal-
cium malate.
Acid from glucose, mannitol and sucrose
but not from lactose.
Nitrates reduced to nitrites and am-
monia.
Aerobic, facultative.
Optimum temperature, between 20° and
28° C.
Habitat: Sea water and marine mud.
56. Pseudomonas calciprecipitans
Molisch, 1925. (Cent. f. Bakt., II Abt., 65,
1925, 130.)
cal.ci. pre. ci'pi. tans. L. calx, calcis lime;
L. praecipito to throw down; M.L. part. adj.
calciprecipitans lime-precipitating.
120
ORDER I. PSEUDOMONADALES
Thin rods, 0.5 to 0.8 by 1.5 to 3.6 microns,
with rounded ends, often staining irregu-
larly. Motile, with one polar flagellum.
Gram-negative.
Gelatin colonies: Circular, light brown in
color (large colonies show CaCOg crystals).
Gelatin stab : Surface growth with filiform
growth in depth. Liquefaction starts at
bottom.
Agar colonies (sea water) ; Grayish white,
glistening. In two to three weeks crystals
of calcium carbonate form in the agar.
Agar slant: Slight, whitish surface
growth becoming thick, spreading, glisten-
ing, with abundant CaCOs crystals in
medium.
Ammonia formed.
Aerobic, facultative.
Optimum temperature, 20° C.
Source : Isolated from sea water.
Habitat: Sea water.
56a. Pseudomonas halestorga Elazari-
Volcani, 1940. (Pse2idomonas halestorgus (sic)
Elazari-Volcani, Studies on the Microflora
of the Dead Sea. Thesis, Hebrew University,
Jerusalem, 1940, VIII and 82.)
hal.e'stor.ga. Gr. noun hale salt water;
Gr. adj. storgus loving; M.L. adj. halestorgus
salt-water-loving.
Rods, the length of which varies greatly
depending on the concentrations of salt : at
3 to 24 per cent, they are usually 0.5 bj^ 1.3
to 4.0 microns, occurring singly and in pairs;
in 0.5 and 30 per cent salt and in Dead Sea
water, the rods are usually very long,
twisted threads. Motile by means of a single,
polar flagellum. Gram-negative.
Gelatin stab (12 per cent salt, 1 per cent
proteose peptone, 15 per cent gelatin) : Fili-
form, very slight infundibuliform liquefac-
tion after six weeks.
Agar colonies (12 per cent salt, 1 per cent
proteose peptone, 2 per cent KNO3) : Circu-
lar, smooth, entire, slightly convex, glisten-
ing, slightly transparent, grayish.
Agar slant (12 per cent salt, 1 per cent
proteose peptone, 2 per centKNOa) : Moder-
ate, filiform, raised, smooth, slightly trans-
parent, grayish growth.
Broth (12 per cent salt, 1 per cent pej)-
tone) : Very turbid; whitish pellicle is
formed.
Indole not produced.
No acid or gas from glucose, fructose,
galactose, mannose, lactose, sucrose, mal-
tose, arabinose, xylose, rafEnose, inulin,
dextrin, glycerol, mannitol or salicin.
Starch not hydrolyzed.
Nitrites are produced from nitrates; no
gas is produced.
Aerobic.
Optimum temperature, 30° C.
Salt tolerance: Halotolerant, growing
slightly in 0.5 per cent salt, strongly in 3 to
30 per cent salt and moderately in Dead Sea
water.
Source: Isolated from the water of the
Dead Sea.
Habitat: Found in places where the salt
content of water is high.
57. Pseudomonas iridescens Stanier,
1941. (Jour. Bact., 4S, 1941, 542.)
ir.id.es'cens. Gr. fem.noun iris, -idis the
rainbow; M.L. part. adj. iridescens showing
colors of the rainbow.
Rods, 0.2 to 0.3 by 1.5 to 7.0 microns,
average length 5.0 to 6.0 microns, occurring
singly. Non-motile. Gram-negative.
Sea water gelatin stab: Filiform growth.
Liquefaction by some strains.
Sea water agar colonies: Concave, 2 to 3
mm in diameter, smooth, glistening, trans-
lucent, pale j^ellow, edge irregular. After 2
to 3 days a marked iridescence. Later
colonies rough, opaque, bright yellow,
sunken central portion with translucent
peripher3^
Sea water agar slant: Growth spreading,
smooth, glistening, translucent, pale yel-
low, iridescent, butyrous.
Sea water broth: Turbid, light yellow,
granular pellicle.
Indole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide not produced.
Catalase-positive.
Urease-negative.
Acid from xylose, glucose, galactose,
lactose, maltose, sucrose and cellobiose. No
acid from arabinose. Starch and cellulose
are attacked.
Aerobic.
Temperature relations: Optimum, 23° C.
Minimum, 5° C. Maximum, 30° C.
FAMILY IV. PSEUDOMONADACEAE
121
Salt range: 0.25 to 6.0 percent. Optim\im,
1.0 to 4.0 per cent.
Source: Sea water.
Habitat: Common along the coast of the
North Pacific.
58. Pseudomonas beijerinckii Hof,
1935. (Travaux botaniques neerlandais, 32,
1935, 152.)
bei.jer.inck'i.i. M.L. gen. noun heije-
rinckii of Beijerinck; named for Prof . M. W.
Beijerinck of Delft, Holland.
Small rods. Motile with polar flagelhi.
Gelatin: No liquefaction.
Indole not produced.
Nitrites produced from nitrates by four
out of six strains.
Cellulose not decomposed.
Acid from glucose. In yeast-water with 2
per cent glucose and 12 per cent NaCI, no gas
is produced.
Pigment production: Insoluble purple pig-
ment produced but not in all media; is lo-
calized markedl}^; reduced oxygen tension
necessary; optimum pH, 8.0; not produced
in yeast-water or in peptone-water; pro-
duced only when grown in extracts of beans
or some other vegetable.
Aerobic.
Source: Six strains isolated from beans
preserved with salt.
Habitat: Causes purple discoloration of
salted beans.
59. Pseudomonas aceris (Ark, 1939)
Starr and Burkholder, 1942. (Phytomonas
aceris Ark, Phytopath., 29, 1939, 969; Starr
and Burkholder, Phytopath., 32, 1942, 601.)
a'ce.ris. L. acei- the maple; L. neut.
gen. noun aceris of the maple.
Rods 0.3 to 0.8 by 0.8 to 2.5 microns.
Motile, with 1 to 2 polar flagella. Gram-
negative.
Green fluorescent pigment produced.
Gelatin: Liquefied.
Beef -extract-peptone agar: Colonies are
grayish white, appearing in 24 hours.
Broth: Turbid.
Milk: Clearing with no coagulation.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Acid from glucose, fructose, galactose,
arabinose, xylose, sucrose, maltose, lactose,
raffinose, mannitol, glycerol and dulcitol.
Slight growth in broth plus 6 per cent salt
(Burkholder).
Optimum temperature, between 13° and
31° C.
Source : From diseased leaves of the large
leaf maple, Acer niacrophyllum.
Habitat: Causes a disease of Acer spp.
60. Pseudomonas angulata (Fromme
and Murray, 1919) Holland, 1920. (Bac-
terium angulatum Fromme and Murray,
Jour. Agr. Res., 16, 1919, 219; Holland, Jour.
Bact.,5, 1920,224.)
ang.u.la'ta. L. part. adj. angulatus wdth
angles, angular.
Description from Clara (Cornell Agr.
Exp. Sta. Mem. 159, 1934, 24).
Rods 0.75 to 1.5 by 1.5 to 3.0 microns.
Motile, with 1 to 6 polar flagella. Gram-
negative.
Gelatin: Liquefied.
Green fluorescent pigment produced.
Beef-extract agar colonies: Dull white,
circular, raised, smooth and glistening.
Broth: Turbid and greenish in 36 hours.
Milk: Alkaline.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Lipolytic action negative (Starr and
Burkholder, Phytopath., 32, 1942, 601).
Acid but no gas from glucose, galactose,
fructose, mannose, arabinose, xylose, su-
crose and mannitol. Alkaline reaction from
salts of citric, malic, succinic and tartaric
acids. Rhamnose, maltose, lactose, raffinose,
glycerol, salicin, and acetic, lactic and
formic acids are not fermented.
Starch not hydrolyzed.
Slight growth in broth plus 5 to 6 per cent
salt (Burkholder).
Aerobic, facultative.
Relationship to other species: Braun
(Phytopath., 27, 1937, 283) considers this
species to be identical in culture with Pseu-
domonas tabaci, but they differ in the type of
disease they produce.
Source : Isolated by Fromme and Murray
from small angular leaf spots on tobacco.
122
ORDER I. PSEUDOMONADALES
Habitat : Causes the angular leaf spot of
tobacco {Nicotiana tahacum).
61. Pseudomonas aptata (Brown and
Jamieson, 1913) Stevens, 1925. {Bacterium
aptatum Brown and Jamieson, Jour. Agr.
Res., 1, 1913, 206; Stevens, Plant Disease
Fungi, New York, 1925, 22.)
ap.ta'ta. L. part. adj. aptahis adapted.
Rods 0.6 by 1.2 microns. Motile, with bi-
polar fiagella. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar slants: Moderate growth along
streak, filiform, whitish, glistening.
Broth: Turbid, with formation of a pel-
licle.
Milk: Becomes alkaline and clears.
Nitrites not produced from nitrates.
Indole not produced in 10 daj's. Slight
amount found later.
Hydrogen sulfide not produced.
Acid from glucose, galactose and sucrose.
No acid from lactose, maltose or mannitol
(Paine and Banfoot, Ann. Appl. Biol., 11,
1924, 312).
Starch not hydrolyzed.
Slight growth in broth plus 7 per cent salt
(Burkholder) .
Temperature relations: Optimum, be-
tween 27° and 28° C. Minimum, below 1° C.
Maximum, between 34° and 35° C.
Aerobic.
Source : Isolated from diseased nasturtium
leaves from Virginia and diseased beet
leaves from Utah.
Habitat: Pathogenic on sugar beets, nas-
turtiums and lettuce.
62. Pseudomonas primulae (Ark and
Gardner, 1936) Starr and Burkholder, 1942.
{Phytomonas primnlae Ark and Gardner,
Phytopath., S6, 1936, 1053; Starr and Burk-
holder, Phytopath., 32, 1942, 601.)
pri'mu.lae. L. dim. adj. prinmlus the first;
M.L. fem.noun Primula generic name;
M.L. gen. noun primulae of Primula.
Rods 0.51 to 0.73 by 1.0 to 3.16 microns.
Motile, with a polar flagellum. Gram-nega-
tive.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar colonies: Round, convex, smooth,
glistening, j-ellowish.
Milk: Coagulated.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolj'tic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid but no gas from glucose, lactose, su-
crose, maltose, galactose, arabinose, glyc-
erol, dulcitol and mannitol. Starch not
hydrolyzed.
Growth in broth plus 5 per cent salt.
Temperature relations : Optimum between
19° and 22° C. Minimum, 10° C. Maximum,
34° C.
Optimum pH between 6.8 and 7.0. Mini-
mum, between 4.5 and 5.0.
Aerobic, facultative.
Source : Isolated from leaf -spot of Primula
polyantha.
Habitat: Pathogenic on Primula spp.
63. Pseudomonas viridilivida (Brown,
1915) Holland, 1920. {Bacterium viridilivi-
dum Brown, Jour. Agr. Res., 4, 1915, 475;
Holland, Jour. Bact., 5, 1920, 225.)
vi.ri.di.li'vi.da. L. viridis green; L. livi-
dus blue; M.L. adj. viridilividus greenish
blue.
Rods 1.0 to 1.25 by 1.25 to 3.0 microns.
Motile, with 1 to 3 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Slow liquefaction.
Beef agar colonies: Cream- white, round,
smooth, translucent, edges entire.
Broth: Turbid, becomes lime-green.
Milk: Alkaline and clears.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole produced.
Not lipolytic (Starr and Burkholder,
Phytopath., S^, 1942,601).
Acid from glucose and sucrose (Burk-
holder).
Grows well in 4.5 per cent salt. Grows in 7
per cent salt (Burkholder).
Temperature relations: Minimum, 1.5° C.
Maximum, 34.5° C.
Aerobic.
FAMILY IV. PSEUDOMONADACEAE
123
Source: Isolated from diseased lettuce
from Louisiana.
Habitat: Pathogenic on lettuce, Lactuca
saliva.
64. Pseudonionas delphinii (Smith,
1904) Stapp, 1928. {Bacillus delphinii Smith,
Science, 19, 1904, 417; Stapp, in Sorauer,
Handbuch der Pflanzenkrankheiten, 2, 5
Aufl., 1928, 106.)
del.phi'ni.i. Gr. delphinium the larkspur;
ALL. dim. neut. noun Delphinium generic
name; M.L. gen. noun delphinii of larkspur.
Rods 0.6 to 0.8 by 1.5 to 2.0 microns.
Chains present. Motile, with 1 to 6 polar
flagella. Encapsulated. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef agar slants: Growth thin, smooth,
shining, transparent, margins entire, crys-
tals. Agar becomes dark brown.
Broth: Turbid in 24 hours with delicate
pellicle.
Milk: Becomes alkaline and clears.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Lidole not produced.
Hydrogen sulfide not produced.
Lipolytic action negative (Starr and
Burkholder, Phytopath., 32, 1942, 601).
Acid from glucose, galactose and fructose;
slight acid from sucrose. No acid from lac-
tose, maltose, glj'cerol or mannitol.
Starch: Hydrolysis feeble.
Weak growth in broth plus 4 per cent salt.
Optimum pH, 6.7 to 7.1. pH range, 5.6 to
8.6.
Temperature relations: Optimum, 25° C.
Minimum, 1° C. or less. Maximum, 30° C.
Source: Isolated from black spot of del-
phinium.
Habitat: Pathogenic on delphinium caus-
ing a black spot in the leaves.
65. Pseudonionas cepacia Burkholder,
1950. (Phytopath., J^0, 1950, 116.)
ce.pa'ci.a. L. fem.noun caepa or cepa
onion; M.L. adj. cepacius of or like onion.
Rods, 0.8 by 1.0 to 2.8 microns, occurring
singly or in pairs. Motile, with 1 to 3 polar
flagella. Gram-negative.
Gelatin: Slow liquefaction.
Beef -extract-peptone agar: Slants sulfur-
yellow, filiform, butyrous to slightly viscid.
Most cultures appear rough. Yellow to
yellow-green pigment diffuses into medium
about the colony.
Potato dextrose agar: Pale yellow. No
change in medium.
Broth: Turbid in 24 hours; yellow pellicle.
Milk: Litmus reduced. Medium clears and
becomes tan with a yellow pellicle.
Krumwiede's Triple sugar agar: Growth
very abundant, j-ellow-green and extremely
wrinkled; medium becomes red.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, fructose,
lactose, maltose, sucrose, arabinose, xylose,
glycerol, mannitol and salicin; alkaline re-
action from sodium salts of citric, hippuric,
malonic and tartaric acids. Growth is slight
in rhamnose. 2 per cent ethyl alcohol jiot
utilized.
Starch not hydrolj^zed.
Sodium ammonium pectate medium not
liquefied.
Methyl red test negative; acetylmethyl-
carbinol not produced.
Growth in 3 per cent but not in 5 per cent
salt.
Temperature relations: Optimum, 30° C.
Minimum, between 6° and 9° C. Maximum,
42° C.
Aerobic.
Source: Seven isolates from different
onion bulbs collected in New York State.
Habitat: Pathogenic on onions. Allium
cepa.
66. Pseudonionas apii Jagger, 1921.
(Jour. Agr. Res., 21, 1921, 186.)
a'pi.i. L. apium celery; M.L. neut. noun
Apium generic name of celery; M.L. neut.
gen. noun apii of celery.
Description from Clara (Cornell Agr.
Exp. Sta. Mem. 159, 1934, 24).
Rods 0.75 to 1.5 by 1.5 to 3.0 microns.
Motile with a polar flagellum. Gram-nega-
tive.
Green fluorescent pigment produced in
various media.
Gelatin: Liquefied.
Beef -extract agar colonies: Circular,
124
ORDER I. PSEUDOMONADALES
glistening, smooth, edges entire. Grayish
white with bluish tinge.
Broth: Turbid in 36 hours. Pellicle
formed.
Milk: Becomes alkaline. No curd.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, galactose,
fructose, mannose, arabinose, xylose, su-
crose, mannitol and glycerol. Alkaline reac-
tion from salts of acetic, citric, malic and
succinic acids. Rhamnose, maltose, lactose,
raffinose salicin, and formic, lactic and
artaric acids are not utilized.
Starch not hydrolyzed.
Aerobic, facultative.
Distinctive character: Pathogenicity ap-
pears limited to celery.
Source: Jagger isolated this repeatedly
from diseased celery leaves.
Habitat: Pathogenic on celery, Apiutn
graveolens.
67. Pseudomonas asplenii (Ark and
Tompkins, 1946) Savulescu, 1947. (Phj/to-
monas asplenii Ark and Tompkins, Phyto-
path., 36, 1946, 760; Savulescu, Anal. Acad.
Romane, III, 2^, 1947, 11.)
a.sple'ni.i. Gr. neut.noun asplenum
spleenwort; M.L. neut.noun Asplenium
generic name; M.L. gen.noun asplenii of
Asplenium.
Rods 0.3 to 0.5 by 1.2 to 2.4 microns.
Motile, with 1 to 3 polar flagella. Gram-
negative.
Gelatin: Liquefied.
Beef -extract-peptone agar slants : Grayish
white with fluorescence in the medium.
Potato-dextrose-peptone agar: Growth
rapid, heavy, strongly grayish white,
butyrous; medium darkens with age.
Nutrient broth: Turbid in 24 hours; no
pellicle.
Milk: No curd.
Indole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide not produced.
Acid but no gas from glucose, galactose,
fructose, arabinose, xylose, maltose and
sucrose. Slight acidity in lactose after long
incubation; no acid in rafiinose.
Starch not hydrolyzed.
Growth good in Fermi's, Cohn's and
Uschinsky's solutions.
Temperature relations : Optimum between
22° and 30° C. Minimum, 1° C. Maximum,
34° C.
Source : Six isolates and 3 reisolates from
lesions on the bird's nest fern.
Habitat: Pathogenic on the fern, Asple-
nium nidus.
68. Pseudomonas berberidis (Thorn-
berry and Anderson, 1931) Stapp, 1935.
(Phytomonas berberidis Thornberry and An-
derson, Jour. Agr. Res., 43, 1931, 36; Stapp,
Bot. Rev., 1, 1935,407.)
ber.be'ri.dis. M.L. Berberis generic name
of barberry; M.L. fem. gen. noun berberidis
of barberry.
Rods, 0.5 to 1.0 by 1.5 to 2.5 microns, oc-
curring singly or in pairs. Motile with 2 to 4
polar flagella. Encapsulated. Gram-negative
(Burkholder); not Gram-positive as stated
in original description.
Green fluorescent pigment produced in
culture (Burkholder).
Gelatin: Not liquefied.
Glucose agar slants: Growth moderate,
filiform at first, later beaded, raised,
smooth, white. Butyrous in consistency.
Milk: Becomes alkaline. No other change.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 3^, 1942, 601).
Acid from glucose, galactose and sucrose.
Maltose and rhamnose not utilized (Burk-
holder).
No gas from carbohydrates.
Starch not hydrolyzed.
Temperature relations: Optimum, 18° C.
]\Iinimum, 7° C. Maximum, 30° C.
Aerobic.
Source: Repeated isolations from leaves
and twigs of barberry.
Habitat: Pathogenic on barberry, Berberis
thnnbergerii and B. vulgaris.
69. Pseudomonas coronafaeiens (El-
liott, 1920) Stevens, 1925. {Bacterium corona-
FAMILY IV. PSEUDOMONADACEAE
125
faciens Elliott, Jour. Agr. Res., 19, 1920, 153;
Steven.s, Plant Disease Fungi, 1925, 27.)
co.ro.na.fa'ci.ens. L. corona crown; L.
facio to make; M.L. part. adj. coronafaciens
halo-producing.
Rods, 0.65 by 2.3 microns, occurring in
chains. Motile with polar flagella. Encapsu-
lated. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Slow liquefaction.
Nutrient agar colonies: White, becoming
irregularly circular, flat with raised mar-
gins.
Broth : Slight turbidity in 24 hours. Heavy
pellicle formed.
Milk: Alkaline. A soft curd formed fol-
lowed by clearing. Curd sometimes absent.
Test for nitrites produced in nitrate broth
negative or faint (Burkholder and Starr,
Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid but no gas from glucose and sucrose.
Starch hj^drolysis slight.
Slight growth in broth plus 2 per cent salt.
Temperature relations: Optimum between
24° and 25° C. Minimum, 1° C. Maximum,
31° C.
Comment: A variety pathogenic on
brome-grass, Bromus inermis, has been de-
scribed by Reddy and Godkin (Phytopath.,
IS, 1923, 81). Produces water-soaked spots
which are dark purple in color. Has been
artificially inoculated on oats {Avena saliva) .
Also pathogenic on Agropyron repens.
Source: Numerous isolations from
blighted blades of oats.
Habitat: Causes a halo spot on oats
{Avena saliva). Artificial inoculations show
barley (Hordeum vulgare), rye (Secale
cereale) and wheat {Trilicum aesHvum) to be
susceptible.
70. Pseudotnonas lachrymans (Smith
and Bryan, 1915) Carsner, 1918. {Bacterium
lachrymans Smith and Bryan, Jour. Agr.
Res., 5, 1915, 466; Carsner, Jour. Agr. Res.,
15, 1918, 201.)
lach'ry.mans. L. lacrimo to shed tears;
M.L. part. adj. lachrymans shedding tears.
Description from Smith and Bryan {op.
cit., 1915, 466) and Clara (Cornell Agr. Exp.
Sta. Mem. 159, 1934,26).
Rods 0.8 by 1.0 to 2.0 micron.s. Motile
with 1 to 5 polar flagella. Encapsulated.
Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef-peptone agar colonies: Circular,
smooth, glistening, transparent, whitish,
entire margins.
Broth: Turbid in 24 hours. White precipi-
tate with crystals.
Milk: Turns alkaline and clears.
Nitrites not produced from nitrates.
Indole reaction weak.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder, Phj--
topath.,5j?, 1942, 601).
Acid but no gas from glucose, fructose,
mannose, arabinose, xylose, sucrose and
mannitol. Alkaline reaction from salts of
citric, malic and succinic acids. Maltose,
rhamnose, lactose, raffinose, glycerol and
salicin not fermented (Clara, op. cil., 1934,
26).
Starch partially digested. Not digested
(Clara, loc. cit.).
Growth in 3 per cent salt after 12 days. No
growth in 4 per cent salt.
Temperature relations : Optimum between
25° and 27° C. Minimum, 1° C. Maximum,
35° C.
Aerobic, facultative (Clara, loc. cit.).
Source : Isolated from diseased cucumber
leaves collected in New York, Wisconsin,
Indiana and in Ontario, Canada.
Habitat: Pathogenic on cucumber, Cucu-
mis sativus, and related plants.
71. Pseudomonas niaculicola (McCul-
loch, 1911) Stevens, 1913. {Baclerium macu-
licolum McCulloch, U. S. Dept. Agr., Bur.
Plant Ind. Bui., 225, 1911, 14; Stevens, The
Fungi which cause Plant Diseases, 1913, 28.)
ma.cu.li'co.la. L. macula spot; L. -cola a
dweller; M.L. noun maculicola spot dweller.
Rods 0.9 by 1.5 to 3.0 microns. Filaments
present. Motile with 1 to 5 polar flagella.
Gram-negative.
126
ORDER I. PSEUDOMONADALES
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef -peptone agar colonies: Whitish, cir-
cular, shining, translucent, edges entire.
Broth: Turbid. No ring or pellicle.
Milk: Becomes alkaline and clears.
Nitrites not produced from nitrates.
Indole production feeble.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid from glucose, galactose, xylose, su-
crose, glycerol and mannitol. Alkaline reac-
tion from salts of citric, malic, malonic and
succinic acids. Salicin, maltose and salts of
hippuric and tartaric acids not utilized
(Burkholder).
Slight growth in broth plus 4 per cent salt
(Erw. Smith, Bact. Plant Diseases, 1920,
306).
Aerobic.
Temperature relations : Optimum between
24° and 25° C. Minimum, 0° C. Maximum,
29° C.
Source: Isolated from diseased cauliflower
leaves from Virginia.
Habitat: Pathogenic on cauliflower and
cabbage.
72. Pseudonionas mangiferaeindicae
Patel et al., 1948. (Pseudonionas mangijerae-
indicae (sic) Patel, Moniz and Kulkarni,
Curr. Sci., 17, 1948, 189; Indian Phytopath.,
1, 1948, 147.)
man.gi'fe.rae.in"di.cae. M.L. fem.noun
Mangifera mango bearer; L. adj. indicus of
Indm; mangiferaeindicae of Mangifera indica.
Rods, 0.36 to 0.54 by 0.45 to 1.44 microns,
occurring singly or in chains of 2 to 4. Motile
with 1 or 2 polar flagella. Gram-negative.
Gelatin: Liquefied.
Nutrient agar colonies: Flat, smooth,
glistening, round with entire margins, white
to creamy, border deeper in color.
Potato glucose agar slants: Growth
copious, raised, smooth, glistening, filiform,
opalescent, butyrous, white.
Broth: Turbid with pellicle in 7 days.
Slight sediment.
Milk: Litmus reduced in 7 days. Cleared
with gelatinous sediment. Casein digested.
Loeffler's blood serum: Liquefied.
Uschinsky's solution: Good growth.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Acid but no gas from glucose, lactose and
sucrose. Slight growth and acid in mannitol.
L-arabinose, maltose, fructose, inulin,
glycerol, salicin, sodium tartrate and
asparagine not utilized.
Starch hydrolyzed.
Lipase not produced.
Growth in 2 per cent salt.
Temperature relations : Optimum between
20° and 25° C. Minimum, 5° C. Maximum,
35° C.
Aerobic.
Source: Isolated from diseased leaves of
mangoes.
Habitat : Pathogenic on Mangifera indica,
Spondias mangiferae and Anacardium occi-
dentale.
73. Pseudonionas marginata (McCul-
loch, 1921) Stapp, 1928. {Bacterium margi-
natum McCulloch, Science, 54, 1921, 115;
Jour. Agr. Res., 29, 1924, 174; Stapp, in
Sorauer, Handbuch der Pflanzenkrank-
heiten, 2, 5 Aufl., 1928, 56.)
mar.gi.na'ta. L. margino to furnish with
a border; L. part. adj. marginatus margined.
Rods 0.5 to 0.6 by 0.8 to 1.8 microns. Mo-
tile with 1 to 4 bipolar flagella. Encapsu-
lated. Gram-negative.
Green fluorescent pigment produced in
Uschinsky's and Fermi's solutions.
Gelatin: Liquefied.
Agar colonies: White, circular, smooth,
translucent, viscid, with definite margins at
first thin but later thick and contoured.
Surface wrinkled.
Milk: At first slightly acid, then alkaline.
Casein digested.
Nitrites not produced from nitrates.
Indole production slight.
Hydrogen sulfide production slight.
Lipolytic (Starr and Burkholder, Phyto-
path., 32, 1942, 601).
Acid but no gas from glucose, lactose, su-
crose and glycerol.
Starch hydrolysis feeble.
Growth in 3.5 per cent salt. No growth in
4 per cent salt.
Temperature relations : Optimum between
FAMILY IV. PSEUDOMONADACEAE
127
30° and 32° C. Minimum between 8° and
9° C. Maximum, 40° C.
pH range, 4.6 to 9.1.
Source : Repeatedly isolated from diseased
gladiolus.
Habitat: Pathogenic onGladiolus spp. and
7m spp.
74. Pseudonionas inedicaginis Sackett,
1910. (Science, 31, 1910, 553; also Colorado
Agr. Exp. Sta., Bull. 158, 1910, 11.)
me.di.ca'gi.nis. Gr. medice the Median
grass, alfalfa, lucerne, medic; M.L. fern,
noun Medicago generic name of alfalfa; M.L.
fem. gen. noun medicaginis of lucerne or
alfalfa.
Rods 0.7 by 1.2 microns. Motile with 1 to
4 flagella. Filaments present. Gram-nega-
tive.
Green fluorescent pigment produced in
culture.
Gelatin: Not liquefied.
Nutrient agar colonies: Growth in 24
hours whitish, glistening.
Broth: Turbid in 24 hours. Pellicle
formed. Viscid sediment.
Milk: Becomes alkaline. No change.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 3£, 1942, 601).
Starch not hydrolyzed.
No gas from carbohydrates. Acid from
sucrose.
Slight growth in broth plus 3.75 per cent
salt.
Temperature relations : Optimum between
28° and 30° C. Maximum, 37.5°C.
Aerobic.
Source: Isolated from brown lesions on
leaves and stems of alfalfa.
Habitat: Pathogenic on alfalfa, Medicago
sp.
75. Pseudomonas phaseolicola (Burk-
holder, 1926) Dowson, 1943. (Phytomonas
medicaginis var. phaseolicola Burkholder,
Phytopath., 16, 1926, 915; Dowson, Trans.
Brit. Mycol. Soc, 26, 1943, 10.)
pha.se.o.li'co.la. Gr. phaseolus the kid-
ney bean; L. dim. mas. noun phaseolus the
kidney bean; L.mas.gen.noun phaseoli of the
bean; L. cola dweller; M.L. fem. noun
phaseolicola the bean dweller.
Description from Burkholder and Zaleski
(Phytopath., ^^, 1932, 85).
Rods 1.0 by 2.0 microns, sometimes
slightly curved; filaments present. Motile
with a polar flagellum. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin stab: Slow liquefaction.
Beef extract agar: Whitish, circular
colonies, 2 mm in diameter. Edges entire.
Broth: Turbid.
Milk: Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 82, 1942, 601).
Acid but no gas from glucose, fructose,
mannose, arabinose, xylose, sucrose and
glycerol. No acid from rhamnose, lactose,
maltose, mannitol or salicin. Alkali from
salts of citric and malic acids, but not from
acetic, formic, lactic or tartaric acids.
Starch and cellulose not hydrolyzed.
Slight growth in broth plus 4 per cent salt.
Temperature relations : Optimum between
20° and 23° C. Minimum, 2.5° C. Maximum,
33° C. (Hedges, Jour. Agr. Res., 36, 1928,
428).
Chemical tolerance: Optimum pH be-
tween 6.7 and 7.3. Minimum between 5.0
and 5.3. Maximum between 8.8 and 9.2
(Kotte, Phyt. Zeitsch., 2, 1930 453).
Microaerophilic.
Source: Isolated from leaves, pod and
stem of beans showing halo blight.
Habitat: Pathogenic on beans (Phaseo-
lus vidgaris), the kudzu vine (Pueraria
hirsuta) and related plants.
76. Pseudonionas pisi Sackett, 1916.
(Colorado Agr. Exp. Sta., Bull. 218, 1916,
19.)
pi'si. Gr. pisus or pisum the pea; M.L.
neut.noun Pisum generic name of the pea;
M.L. neut. gen. noun pisi of the pea.
Rods 0.68 to 2.26 microns. Motile with a
polar flagellum. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
128
ORDER I. PSEUDOMONADALES
Agar slants : Moderate growth in 24 hours,
filiform, glistening, grayish white.
Broth: Turbid with a scum in 5 days.
Milk: Alkaline, soft curd, clears.
Nitrites not produced from nitrates.
Indole not produced.
Hj'drogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid but no gas from glucose, galactose
and sucrose.
Starch not hydrolyzed.
Temperature relations : Optimum between
27° and 28° C. Minimum, 7° C. Maximum,
37.5° C.
Aerobic.
Source: Ten cultures isolated from 5 col-
lections of diseased peas showing water-
soaked lesions on stems and petioles.
Habitat: Pathogenic on garden peas,
Pisum sativum, and field peas, P. sativum
var. arvense.
77. Pseudonionas syringae van Hall,
1902. (Kennis der Bakt. Pflanzenziekte,
Inaug. Diss., Amsterdam, 1902, 191.)
sy.rin'gae. Gr. syrinx, syringis a pipe or
tube; M.L. fem.noun Syringa generic name
of syringa or lilac; M.L. fem. gen. noun.
syringae of the lilac.
Description from Clara (Cornell Agr.
Exp. Sta. Mem. 159, 1934, 29).
Rods 0.75 to 1.5 by 1.5 to 3.0 microns.
Motile with 1 or 2 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef -extract agar colonies: Circular,
grayish white with bluish tinge. Surface
smooth. Edges entire or irregular.
Broth: Turbid in 36 hours. No pellicle.
Milk: Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 3^, 1942, 601).
Slight growth in broth plus 4 per cent salt.
Acid but no gas from glucose, galactose,
mannose, arabinose, xylose, sucrose, man-
nitol and glycerol. Alkaline reaction from
salts of citric, malic, succinic and lactic
acids. Rhamnose, maltose, lactose, raffinose,
salicin, and acetic, formic and tartaric acids
not fermented.
Starch not hydrolj'zed.
Aerobic, facultative.
Comment: Orsini (Intern. Bull. Plant
Protect., S3, 1942, 33) reports that a variety
of this species is pathogenic on the pepper
plant {Capsicum).
Source: Van Hall originally isolated this
pathogen from lilac.
Habitat: Pathogenic on lilac, citrus, cow
peas, beans, lemons, cherries and many un-
related plants.
78. Pseudonionas tomato (Okabe, 1933)
Alstatt, 1944. {Bacterium tomato Okabe,
Jour. Soc. Trop. Agr. Formosa, 5, 1933, 32; .„
Alstatt, U. S. Dept. Agr., Plant Dis. Rept.,
28, 1944, 530.)
to.ma'to. Am.Ind. tomatl; Sp. tornate;
Eng. tomato; M.L. noun tomato.
Rods 0.69 to 0.97 by 1.8 to 6.8 microns.
Motile with 1 to 3 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Slow liquefaction.
Beef -extract agar colonies: White, circu-
lar, flat and glistening.
Broth: Turbid in 24 hours. Pellicle.
Milk: Becomes alkaline and clears.
Nitrites are usually produced from ni-
trates.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, sucrose and
lactose. No acid from maltose or glycerol.
Starch hydrolysis feeble.
Slight growth in 3 per cent salt.
Temperature relations : Optimum between
20° and 25° C. Maximum, 33° C.
Aerobic.
Source: Isolated from diseased tomato
leaves.
Habitat: Pathogenic on tomato, Lyco-
persicon esculenium.
79. Pseudomonas atrofaciens (McCul-
loch, 1920) Stevens, 1925. {Bacterium atro-
faciens McCulloch, Jour. Agr. Res., 18, 1920,
549; Stevens, Plant Disease Fungi, New
York, 1925, 22.)
FAMILY IV. PSEUDOMONADACEAE
129
at.ro.fa'ci.ens. L. ater black; L. facio to
make;M.L. part. adj. atrofaciens blackening.
Rods 0.6 by 1.0 to 2.7 microns. Long
chains formed in culture. Encapsulated.
Motile with 1 to 4 polar or bipolar flagella.
Gram-negative
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef-peptone-agar colonies: Circular,
shining, translucent, white.
Broth: Growth never heavy, slight rim,
and a delicate pellicle.
Milk: Becomes alkaline and clears.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole: Slight production.
Hydrogen sulfide: Slight production.
Acid but no gas from glucose, galactose
and sucrose.
Starch is slightly hydrolyzed.
Temperature relations : Optimum between
25° and 28° C. Minimum below 2° C. Maxi-
mum between 36° and 37° C.
Aerobic.
Source: Isolated from diseased wheat
grains collected throughout the United
States and Canada.
Habitat: Causes a basal glume-rot of
wheat.
80. Pseudomonas cuniini (Kovachev-
ski, 1936) Dowson, 1943. {Phytomonas cumini
Kovachevski, Bull. Soc. Bot. Bulgarie, 7,
1936, 27; Dowson, Trans. Brit. Mycol. Soc,
m, 1943, 10.)
cu'mi.ni. Gr. cuminum cumin; M.L.
neut.noun Cuminum generic name of cu-
min; M.L. neut. gen. noun cumini of cumin.
Rods, 0.5 to 0.7 by 1.0 to 3.0 microns, oc-
curring in chains and filaments. Motile with
1 to 3 polar fiagella. Gram-negative.
Green fluorescent pigment formed in cul-
ture.
Gelatin: Rapidly liquefied.
Potato agar colonies: Grayish white, cir-
cular, glistening, smooth, butyrous.
Broth: Moderate turbidity. Pseudozoo-
gloea.
Milk: Not coagulated. Casein peptonized.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose and sucrose.
No acid from lactose or glycerol. Starch not
hydrolyzed.
Temperature range, 5° to 31° C.
Aerobic.
Source: Isolated from blighted cumin
(Cuminian).
Habitat: Pathogenic on cumin and dill.
81. Pseudomonas desaiana (Burk-
holder, 1939) Savulescu, 1947. (B. pyo-
cyaneiis saccharum Desai, Ind. Jour. Agr.
Sci., 5, 1935, 391; Phytomonas desaiana Burk-
holder, in Bergey et al., Manual, 5th ed.,
1939, 174; Savulescu, Anal. Acad. Romane,
III, 22, 1947 11.)
de.sai.a'na. M.L. adj. desaianus. Named
for Prof. Desai of India.
Rods 0.6 to 1.2 by 1.2 to 2.2 microns. Mo-
tile with a polar flagellum. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar colonies: Grayish blue. Raised.
Broth: Light clouding. Pellicle.
Milk: Peptonized without coagulation.
Nitrites not produced from nitrates.
Indole not produced.
Glucose, sucrose, lactose and gh'cerol fer-
mented without gas.
Starch: Hydrolysis present.
Optimum temperature, 30° C.
Aerobic.
Source: Isolated from stinking rot of
sugar cane in India and associated with a
white non-pathogenic bacterium.
Habitat: Pathogenic on sugar cane, Sac-
char u m officinaru tn .
82. Pseudomonas erodii Lewis, 1914.
(Phytopath.,.^, 1914,231.)
e.ro'di.i. Gr. erodius the heron; M.L.
neut.noun Erodium generic name of heron-
bill; M.L. neut. gen. noun erodii of Erodium.
Rods 0.6 to 0.8 by 1.2 to 1.8 microns.
Motile with 1 to 3 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar streak: Heavy, smooth, cream-
colored growth in 24 hours.
Broth: Dense clouding in 24 hours.
130
ORDER I. PSEUDOMONADALES
Milk: Turns alkaline and clears, litmus
reduced.
Nitrites not produced from nitrates.
Indole produced in 14 days.
Hydrogen sulfide not produced.
Acid but no gas from glucose, sucrose, lac-
tose and glycerol.
Temperature: No growth at 35° C.
Aerobic, obligate.
Source: Isolated from Erodium texanum
and 4 varieties of Pelargonium.
Habitat: Causes a leaf spot of Erodium
texanum and Pelargonium spp.
83. Pseudomonas lapsa (Ark, 1940)
Starr and Burkholder, 1942. {Phytomonas
lapsa Ark, Phytopath., 30, 1940, 1 ; Starr and
Burkholder, Phytopath., SS, 1942, 601.)
lap'sa. L. V. labor to fall down, slip; L.
part. adj. lapsus fallen down.
Rods 0.56 by 1.55 microns. Motile, with
1 to 4 polar flagella. Gram reaction not re-
ported; presumably Gram-negative.
Produces fluorescence in Uschinsky's,
Fermi's and Cohn's solutions.
Gelatin: Liquefied (Burkholder).
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Acid but no gas from glucose, sucrose,
maltose, lactose, glycerol, arabinose, xy-
lose, galactose, raffinose and mannitol.
Slight grow^th in broth plus 5 per cent salt
(Burkholder).
Relationship to other species: Resembles
Pseudomonas desaiana.
Source: Isolated from stalk rot of field
corn in California; also from Diabrotica
beetles.
Habitat: Pathogenic on corn and sugar
cane.
84. Pseudomonas martyniae (Elliott,
1924) Stapp, 1928. {Bacterium martyniae
Elliott, Jour. Agr. Res., 29, 1924, 490; Stapp,
in Sorauer, Handbuch der Pflanzenkrank-
heiten,^, 5 Aufl., 1928,278.)
mar.tyn'i.ae. M.L. Martynia genus of
flowering plants; M.L. gen.fem. martyniae of
Martynia.
Rods, 0.59 to 1.68 microns, occurring in
chains. Encapsulated. Motile with one to
several bipolar flagella. Gram-negative.
Green fluorescent pigment produced.
Gelatin: Liquefied.
Beef agar colonies: White, round, smooth,
glistening, raised.
Broth: Clouding in bands. Thin pellicle.
Small crystals.
Milk: Soft acid curd with peptonization.
Nitrites jiroduced from nitrates after 2
weeks.
Indole not produced.
Hydrogen sulfide production slight.
Acid but no gas from glucose, galactose,
arabinose and sucrose. No acid from rham-
nose, lactose, maltose, raffinose, mannitol
or glycerol.
Starch hydrolysis none or feeble.
Temperature relations: Optimum, 26° C.
Minimum, 1.5° C. Maximum, 37° C.
Chemical tolerance: Optimum pH, 6.0 to
6.7. pH range, 5.4 to 8.9.
Aerobic.
Source: Isolated from diseased leaves of
the unicorn plant from Kansas.
Habitat: Pathogenic on Martynia louisi-
ana.
85. Pseudomonas matthiolae (Briosi and
Pavarino, 1912) Dowson, 1943. {Bacterium
matthiolae Briosi and Pavarino, Atti della
Reale Accad. dei Lincei Rend.,^i, 1912, 216;
Dowson, Trans. Brit. Mycol. Soc, 26, 1943,
10.)
mat.thi'o.lae. Mattioli patronymic; M.L.
fern. noun Matthiola generic name of stock;
M.L. fern. gen. noun matthiolae of Matthiola.
Rods 0.4 to 0.6 by 2.0 to 4.0 microns.
Gram-positive. Gram-negative (Mushin,
Proc. Roy. Soc. Victoria, 53, 1941, 201).
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef agar colonies: White, circular,
slightly elevated, margins smooth.
Broth: Slightly turbid. Becomes pale
green.
Milk: Coagulation with acid reaction.
Nitrites produced from nitrates (Mushin,
Proc. Trans. Brit. Mycol. Soc, 26, 1943, 10).
Hydrogen sulfide not produced.
Acid from glucose, galactose, fructose,
mannose, rhamnose, glycerol, mannitol,
acetic acid, citric acid, formic acid, lactic
acid, malic acid and succinic acid. Feeble
acid from maltose. No acid or gas from lac-
FAMILY IV. PSEUDOMONADACEAE
131
tose, sucrose, raffinose, starch, salicin or
tartaric acid (Mushin).
Temperature relations : Optimum, between
20° and 24° C. Minimum, below 0° C. Maxi-
mum, 38.5° C. (Mushin).
Limits of growth in broth are pH 4.4 to
pH 9.5 (Mushin).
Aerobic.
Source: Isolated from vascular and
parenchymatic disease of stocks, Matthiola
incana var. annua.
Habitat: Pathogenic on stocks.
86. Pseudonionas inorsprunoruin Wor-
mald, 1931. {Pseudomonas mors-pnmorum
(sic) Wormald, Jour. Pom. and Hort. Sci.,
9, 1931, 251.)
mors'pru.no.rum. L. mors death; L.
prunus plum; M.L. fem.noun morspruno-
rum plum death.
Rods. Motile with a polar flagellum.
Gram-positive (1931). Gram-negative
(1932).
Gelatin: Liquefied.
Agar colonies: White.
Broth plus 5 per cent sucrose : White and
cloudy.
Nitrites not produced from nitrates.
Acid but no gas from glucose, lactose,
sucrose and glycerol.
Starch not hydrolyzed.
Strict aerobe.
Comment: Possibly a green fluorescent
organism since it produces a faint yellow
color in Uschinsky's solution.
Distinctive characters: Differs from
Pseudomonas prunicola (Pseudomonas syrin-
gae) in that it produces a white cloudy
growth in broth plus 5 per cent sucrose, a
rapid acid production in nutrient agar plus
5 per cent sucrose, and a faint yellow or no
color in Uschinsky's solution.
Source: Isolated from cankers on plum
trees in England.
Habitat: Pathogenic on Prunus spp.
87. Pseudomonas papulans Rose, 1917.
(Phytopath., 7, 1917, 198.)
pa'pu.lans. L. v. papulo to produce pus-
tules; L. part. adj. papulans producing
pustules.
Rods 0.8 by 0.8 to 2.5 microns. Motile
with 1 to 6 polar flagella. Gram-negative.
Green, fluorescent pigment produced in
various media.
Gelatin: Liquefied.
Broth: Turbid with pellicle.
Fermi's and Uschinsky's solutions: Good
growth.
Milk: Litmus reduced; no acid.
Nitrites not produced from nitrates. Ni-
trites produced from nitrates (Burkholder
and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, fructose,
galactose, mannose, arabinose, xylose, su-
crose, glycerol, mannitol, sorbitol, salicin
and esculin.
No acid or gas from rhamnose, lactose,
maltose, raffinose, trehalose, melizitose,
starch, inulin, dextrin, dulcitol or arbutin.
Alkaline reaction produced in glycogen
and in acetic, citric, formic, lactic, malic
and succinic acids.
Temperature relations: Optimum, 27° C.
Minimum, 3.5° C. Maximum, 34.5° C.
Chemical tolerance: Optimum pH, 7.0.
Minimum, 5.0. Maximum, 9.4.
Source: Twenty-five cultures isolated
from blisters on apples and from rough bark.
Habitat: Pathogenic on apple trees.
88. Pseudomoiias pseudozoogloeae
(Honing, 1914) Stapp, 1928. (Bacterium
pseudozoogloeae Honing, Bull, van Het. Deli
Proefstation, Medan, 1, 1914, 7; Stapp, in
Sorauer, Handbuch der Pflanzenkrank-
heiten, ^, 5 Aufl., 1928,274.)
pseu.do.zo.o.gloe'ae. Gr. pseudes false;
Gr. zoum animal; Gr. gloea glue; M.L.
fem.noun Zoogloea bacterial generic name;
M.L. fem. gen. noun pseudozoogloeae of a
false zoogloea.
Rods 0.7 to 1.5 by 0.9 to 2.5 microns.
Chains. Motile with 1 or 2 polar flagella.
Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar colonies: Round, flat, yellow-gray.
Broth: Moderate turbidity with pseudo-
zoogloeae in the pellicle.
Milk: Coagulation. No clearing.
Nitrites not produced from nitrates.
Indole not produced.
132
ORDER I. PSEUDOMONADALES
Hydrogen sulfide produced.
Acid but no gas from glucose, lactose, mal-
tose, sucrose and mannitol.
Aerobic, facultative.
Source: Isolated from the black rust of
tobacco.
Habitat: Pathogenic on tobacco, Nico-
tiana tnbacum.
89. Pseudomoiia.s riniaefaciens Koning,
1938. (Chron. Bot., 4, 1938, 11; Meded.
Phj-top. Labor, Willie Comm. Scholt., 14,
1938, 24.)
ri.mae.fa'ci.ens. L. rima a crack; L. v.
facio to make; M.L. part. adj. riniaefaciens
making cracks.
Rods 0.6 to 2.4 microns in length. Motile
with 1 to 3 flagella. Gram-negative.
Yellow-green, fluorescent, water-soluble
pigment produced in culture.
Gelatin: Liquefied.
Agar colonies: Round, conve.x, smooth,
somewhat granular with hyaline edge.
Broth: Turbid. Surface growth with a
sediment in a few days.
Milk: Alkaline and clears.
Nitrites not produced from nitrates. Pep-
tone, asparagin, urea, gelatin, nitrates and
ammonium salts are sources of nitrogen.
Hydrogen sulfide not produced.
Indole production slight.
Growth with the following carbon sources
plus NO3 : glucose, sucrose, glycerol, suc-
cinates, malates, citrates and oxalates. Less
growth with mannitol, fructose, galactose,
lactose and salicylate. Acid is produced
from the sugars. No growth with dextrin,
inulin, maltose, lactose, rhamnose, salicin,
tartrates, acetates or formates.
Starch not hydrolyzed.
Aerobic.
Temperature relations: Optimum, 25° C.
Very slow growth at 14° C. Maximum, about
37° C. Thermal death point between 42° and
48° C.
Relationship to other species: This may
be Pseudomonas syringae since the charac-
ters are the same and both organisms can
infect Impatiens sp. Pseudomonas syringae
infects poplars (Elliott, Bacterial Plant
Pathogens, 1930, 218).
Source: Strains of the pathogen isolated
from poplar cankers in France and in the
Netherlands.
Habitat: Pathogenic on Populus braban-
tica, P. trichocarpa and P. candicans.
90. Pseudomonas striafaciens (Elliott,
1927) Starr and Burkholder, 1942. (Bac- y
terium striafaciens Elliott, Jour. Agr. Res.,
35, 1927, 823; Starr and Burkholder, Phyto-
path., 32, 1942, 601.)
stri.a.fa'ci.ens. L. stria a furrow; M.L.
part. adj. striafaciens furrowing.
Rods 0.66 by 1.76 microns. Motile with
one to several flagella. Encapsulated. Gram-
negative.
Green fluorescent pigment produced.
Gelatin: Liquefied.
Beef -peptone agar colonies: White,
raised, margins entire or slighth' undulat-
ing.
Broth: Clouding in layers. Ring and slight
pellicle.
Milk: Alkaline, sometimes a soft curd
w^hich digests or clears.
Slight production of nitrites from ni-
trates.
Indole not produced.
Acid but no gas from glucose, fructose
and sucrose. No acid from lactose, maltose,
glycerol or mannitol.
Starch: Hydrolysis slight.
Optimum temperature, 22° C.
Optimum pH, between 6.5 and 7.0.
Aerobic.
Distinctive characters: Differs from
Pseudomonas coronafaciens in that the cells
are somewhat smaller and the pathogen pro-
duces a streak on oat blades instead of a
halo spot.
Source : Forty cultures isolated from oats
gathered in various parts of America.
Habitat: Pathogenic on cultivated oats
and, to a slight degree, on barley.
91. Pseudomonas tabaci (Wolf and
Foster, 1917) Stevens, 1925. {Bacterium ta-
bacum (sic) Wolf and Foster, Science, 46,
1917, 362; also Jour. Agr. Res., 12, 1918, 449;
Stevens, Plant Disease Fungi, New York,
1925, 36.)
ta.ba'ci. M.L. noun iabacum tobacco;
M.L. gen. noun tabaci of tobacco.
FAMILY IV. PSEUDOMONADACEAE
133
Rods 1.2 by 3.3 microns. Motile with a
polar flagellum. Gram-negative.
Gelatin: Liquefied.
Potato agar colonies: Grayish white, cir-
cular, raised, wet-shining, smooth.
Milk: Alkaline; clears.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole not produced.
Acid from glucose, galactose, fructose,
1-arabinose, xj'lose, sucrose, pectin, man-
nitol and glycerol (Braun, Phytopath., 27,
1937, 289).
Ammonium sulfate, potassium nitrate,
cystine, glutamic acid, glycine, succin-
imide, oxamide, acetamide and urea can be
used as nitrogen sources (Braun).
Starch not hydrolyzed.
Aerobic.
Relationship to other species: Braun {loc.
cit.) states that Pseudomonas tabaci and
Pseiidomonas angulata are identical in
culture.
Source: Isolated from wildfire lesions and
tobacco leaves in North Carolina.
Habitat: Pathogenic on tobacco, Nico-
tinna fabacvm.
92. Pseudoniona.s poly color Clara, 1930.
(Phytopath., 20, 1930, 704.)
po.lj^'co.lor. Gr. poly- many; L. color
color; M.L. adj. polycolor many colored.
Description taken from Clara (Cornell
Agr. E.xp. Sta. Mem. 159, 1934, 28).
Rods 0.75 to 1.2 by 1.05 to 3.0 microns.
Motile with 1 or 2 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef -extract agar colonies: Graj^ish
white, circular, raised; thin, transparent
margins.
Broth: Turbid in 36 hours with thin pel-
licle.
Milk: Alkaline; no curd.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Lipolytic (Starr and Burkholder, Phyto-
path., 32, 1942, 601).
Acid l)Ut no gas from glucose, galactose.
fructose, mannose, arabino.se, xylose, man-
nitol and glycerol. Alkaline reaction from
salts of acetic, citric, malic, lactic and for-
mic acids. Rhamnose, sucrose, maltose, lac-
tose, raffinose and salicin not fermented.
Starch not hydrolyzed.
Aerobic, facultative.
Good growth in broth plus 7 per cent salt.
Temperature relations: Optimum be-
tween 25° and 30° C. Maximum between 37°
and 39° C.
Distinctive characters: Differs from
Pseudomonas mellea in type of lesion pro-
duced; does not digest starch nor reduce
nitrates and does not form acid from lactose
nor sucrose. Pathogenic for laboratory ani-
mals (Elrod and Braun, Sci., 94, 1941, 520).
Cultural characters differ from those of
Pseudomonas aeruginosa Migula.
Source: Repeatedly isolated from leaf
spot of tobacco in the Philippines.
Habitat: Pathogenic on tobacco.
93. Pseudomonas viridiflava (Burk-
holder, 1930) Clara, 1934. (Phytomonas viri-
diflava Burkholder, Cornell Agr. Exp. Sta.
Mem. 127, 1930, 63; Clara, Science, 75, 1934,
111.)
vi.ri.di.fla'va. L. viridis green; L. flavus
yellow; M.L. adj. viridiflavus greenish
yellow.
Description from Clara (Cornell Agr.
Exp. Sta. Mem. 139, 1934, 30).
Rods 0.75 to 1.5 by 1.5 to 3.15 microns.
Motile with 1 or 2 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Beef -extract agar colonies: Grayish
white, margins corrugated, edges irregular.
Broth: Turbid in 36 hours.
Milk: Becomes alkaline and clears.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath.,3^, 1942, 601).
Acid but no gas from glucose, fructose,
mannose, arabinose, xjdose, mannitol and
glycerol. Alkaline reaction from salts of
acetic, citric, malic, lactic and succinic
134
ORDER I. PSEUDOMONADALES
acids. Sucrose, lactose, maltose, raffinose,
salicin, and salts of formic and tartaric acids
not fermented.
Starch: No hydrolj'sis.
Growth in broth plus 5 per cent NaCl.
Aerobic, facultative.
Comment: A variety that does not grow
in Uschinsky's solution and that produces
colonies of an unusual shape has been iso-
lated from the stems and leaves of blighted
beans in Denmark. See Petersen (Tidsskr.
f. Planteavl., 38, 1932, 851).
Source: Two cultures isolated from
spotted beans, one from England and one
from Switzerland.
Habitat: Pathogenic on bean, Phaseolus
vulgaris.
94. Pseudoitionas ananas Serrano, 1934.
(Philipp. Jour. Sci., 55, 1934, 355.)
a'na.nas. Braz.Ind. ananas pineapple;
M.L. indecl.neut.noun ananas.
Rods 0.6 by 1.8 microns. Motile with 1 to
4 polar flagella. Gram-negative.
Green fluorescent pigment produced in
certain media.
Gelatin: Liquefied.
Beef -extract glucose agar colonies: White,
with undulating edges, smooth to rugose,
glistening to dull.
Beef-extract agar: Growth scant.
Broth: Feeble growth.
Milk: Becomes alkaline with curd.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, xylose and
mannitol. Feeble with lactose. No acid from
sucrose.
Starch not hydrolyzed.
Temperature relations : Optimum between
30° and 31° C. Minimum between 7° and
10° C. Maximum, 45° C.
Aerobic.
Source: Isolated from rotted pineapples.
Habitat: Causes a rot of pineapples.
Ananas comosus.
95. Pseudomonas bowlesiae (Lewis and
Watson, 1927) Dowson, 1943.* (Phytomonas
bowlesii (sic) Lewis and Watson, Phyto-
path., 17, 1927, 511; Pseudomonas bowlesiae
Dowson, Trans. Brit. Mycol. Soc, 26, 1943,
9.)
bow.le'si.ae. M.L. fem.n. Bowlesia ge-
neric name; M.L. gen. noun bowlesiae of
Bowlesia.
Rods, 0.5 to 0.7 by 1.2 to 1.6 microns,
occurring singly, in pairs or in short chains.
Motile with bipolar flagella. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar slants: Yellowish, moist, glistening
and viscid.
Broth: Uniform turbidity throughout.
Heavy viscous sediment in old cultures.
Milk: Alkaline; coagulation with a slow
peptonization.
Nitrites produced from nitrates.
Indole produced.
Hj'drogen sulfide produced.
Acid from glucose, maltose and xylose.
No acid from sucrose.
Temperature relations: Optimum, 27° C.
Minimum, —1° C. Maximum, 37° C.
Chemical tolerance: Optimum pH, 7.2.
pH range, 4.5 to 8.6.
Aerobic.
Source: Isolated from diseased, water-
soaked spots of bowlesia.
Habitat: Pathogenic on Bowlesia septen-
trionalis.
96. Pseudomonas ligustri (d'Oliveira,
1936) SSvulescu, 1947. {Bacterium ligustri
d'Oliveira, Revista Agron., 2J^, 1936, 434;
Savulescu, Anal. Acad. Romane, III, 22,
1947, 11.)
li.gus'tri. L. ligustrum the privet; M.L.
neut.noun Ligustrum generic name of
privet; ligustri of the privet.
Rods 0.5 to 0.7 by 1.3 to 3.0 microns. No
chains. Not encapsulated. Motile with 2 to
5 polar flagella. Gram-negative.
Green pigment produced on Dox agar and
in broth.
Gelatin: Liquefied.
* The authors of this binomial report (personal communication) that the original spell-
ing bowlesii used for the specific epithet is an orthographic error. The correct spelling is
"bowlesiae" .
FAMILY IV. PSEUDOMONADACEAE
135
Beef -extract agar colonies: Growth mod-
erate. Milky white, circular, convex.
Broth: Turbid in 24 hours. No pellicle.
Milk: Coagulated in 6 days and later di-
gested. Litmus slightly acid.
Nitrites not produced from nitrates.
Indole not produced.
Ammonia not produced.
No gas from carbohydrates. Acid from
glucose, galactose, arabinose and mannose.
No acid from sucrose, maltose, lactose, raf-
finose, raannitol or salicin.
Source: From diseased Japanese privet
in Lisbon, Portugal.
Habitat: Pathogenic on privet, Ligus-
trum ja'ponicum.
97. Pseudomonas marginalis (Brown,
1918) Stevens, 1925. {Bacterium marginale
Brown, Jour. Agr. Res., 13, 1918, 386;
Stevens, Plant Disease Fungi, New York,
1925, 30; Phyfomonas intyhi Swingle, Phyto-
path., 15, 1925, 730.)
mar.gi.na'lis. L. margo, marginis edge,
margin; M.L. adj. marginalis marginal.
Description from Brown (op. cit., 1918,
386) and Clara (Cornell Agr. Exp. Sta. Mem.
159, 1934, 27).
Rods. Motile with 1 to 3 polar flagella.
Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar colonies: Cream-colored to yellow-
ish.
Broth: Turbid, with pellicle.
Milk: Alkaline. Soft curd at times.
Nitrites are produced from nitrates. Not
produced (Clara).
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, galactose,
fructose, mannose, arabinose, xylose, rham-
nose, mannitol and glycerol. Alkali from
salts of acetic, citric, malic, formic, lactic,
succinic and tartaric acids. Sucrose, mal-
tose, lactose, raffinose and salicin not fer-
mented (Clara).
Starch hydrolysis feeble. None (Clara).
Temperature relations : Optimum between
25° and 26° C. Minimum, 0° C. Maximum,
38° C.
Aerobic.
Source: Isolated from marginal lesion on
lettuce from Kansas.
Habitat: Pathogenic on lettuce and re-
lated plants.
98. Pseudomonas sesami Malkoff, 1906.
(Cent. f. Bakt., II Abt., 16, 1906, 665.)
se'sa.mi. Gr. sesamum sesame; M.L.
neut.noun Sesamum generic name of sesame;
sesami of sesame.
Description from Nakata (Ann. Phyt.
Soc. Japan, .?, 1930, 242).
Rods 0.6 to 0.8 by 1.2 to 3.8 microns.
Motile with 2 to 5 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefaction rapid.
Beef -agar colonies: Circular, flat, striate,
smooth, entire margins, white.
Broth: Growth rapid. No pellicle.
Milk: Alkaline. No coagulation.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose. No acid
from lactose, sucrose or glycerol.
Starch not hj^drolyzed.
Temperature relations: Optimum, 30° C.
Minimum, 0° C. Maximum, 35° C.
Aerobic, facultative.
Source: Isolated from l)rown spots on
leaves and stems of sesame.
Habitat: Pathogenic on sesame.
99. Pseudomonas setariae (Okabe,
1934) Savulescu, 1947. (Bacterium setariae
Okabe, Jour. Soc. Trop. Agr. Formosa, 6,
1934, 63; SSvulescu, Anal. Acad. Romane,
III, 22, 1947, 11.)
se.ta'ri.ae. L. saeta a bristle; M.L.
saetarius bristle-like; M.L. fem.noun Se-
taria generic name of foxtail; setariae of
Setaria.
Rods 0.4 to 0.8 by 1.8 to 4.4 microns.
Motile with a polar, seldom bipolar, flagel-
lum. Gram-negative.
Yellowish, water-soluble pigment pro-
duced in culture.
Gelatin: Slow liquefaction.
Beef -extract agar colonies: Circular,
white, opalescent, smooth, glistening.
Broth: Tur])id after 18 hours. Pellicle.
136
ORDER I. PSEUDOMONADALES
Milk: Alkaline; clears.
Nitrites produced from nitrates.
Indole produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose, galactose
and glycerol. No acid from lactose, maltose
or sucrose.
Starch: Feeble hydrolysis.
Grows in 3 per cent salt.
Temperature relations: Optimum be-
tween 31° and 34° C. Maximum, 42° C.
Aerobic.
Source: Isolated from brown stripe of
Italian millet.
Habitat: Pathogenic on Italian millet,
Setaria italica.
100. Pseudomonas tolaasii Paine, 1919.
(Ann. Appl. Biol., 5, 1919, 210.)
to.laa'si.i. Tolaas patronymic; tolaasii
of Tolaas.
Rods 0.4 to 0.5 by 0.9 to 1.7 microns.
Motile with 1 to 5 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Bouillon agar: Streak develops in 24
hours, dirty bluish white, wet-shining and
slightly raised.
Broth: Turbid in 24 hours. Pellicle.
Milk: Becomes alkaline and clears.
Nitrites not produced from nitrates.
Indole production slight.
Acid but no gas from glucose. No acid
from lactose or sucrose.
Starch hydrolysis feeble.
Optimum temperature, 25° C.
Source : Isolated in England from brown-
spot of cultivated mushrooms.
Habitat: Pathogenic on cultivated mush-
101. Pseudomonas washingtoiiiae
(Pine 1943) Elliott 1951. {Phytomonas
washingtoniae Pine, Phytopath., 33, 1943,
1203; Elliott, Man. Bact. Plant Path., 2nd
ed., 1951, 100.)
wash.ing.to'ni.ae. M.L. fem.noun Wash-
ingtonia a generic name; washingtoniae of
Washingtonia.
Rods, 0.69 l)y 1.61 microns, occurring
singly or in short chains. Motile with 1 to 3
polar flagella. Gram-negative.
Green pigment in certain media.
Gelatin: Liquefied.
Potato dextrose agar colonies: Circular,
smooth, convex, glistening, white to cream,
butyrous, edges entire.
Milk: No curd; peptonization with a green
color in 7 days.
Indole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide produced in minute
amounts.
Acid but no gas from glucose, fructose
and L-arabinose in 24 hours; from galactose
and xylose in 48 hours. No acid from sucrose,
lactose, cellobiose, maltose, mannitol,
D-sorbitol, glycerol, salicin or raffinose.
Starch not hydrolyzed.
Aerobic.
Thermal death point between 47° and
48° C.
Source : Isolated from spots on the leaves
of the palm, Washingtonia filifera.
Habitat: Pathogenic on the Washington
palm.
102. Pseiidonioiias barker! (Berridge,
1924) Clara, 1934. (Bacillus of pear blossom
disease. Barker and Grove, Ann. Appl.
Biol., 1, 1914, 94; Barker and Grove's or-
ganism, Doidge, Ann. Appl. Biol., 4, 1917,
50; B. barkeri Berridge, Ann. Appl. Biol., 11,
1924, 73; Clara, Science, 75, 1934, 11.)
bar'ker.i. M.L. gen. noun barkeri of
Barker; named for B. T. P. Barker, one of
the two men who first described this or-
ganism.
Description from Doidge (op. cit., 1917,
50).
Rods 0.5 to 0.8 by 2.0 to 4.0 microns.
Motile with 1 to 4 polar flagella. Gram-nega-
tive (Burkholder), not Gram-positive.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Agar: Growth is white, feeble, flat, glist-
ening, smooth-edged.
Broth: Slightly turbid in 24 hours.
Milk: Slowly cleared.
Nitrites not produced from nitrates.
Indole not produced unless culture
warmed.
FAMILY IV. PSEUDOMONADACEAE
137
Starch slowly digested.
Source: Barker made many cultures from
blighted pear blossoms. Doidge received a
culture from Barker.
Habitat : Causes a blossom blight of pear.
103. Pseudonionas betle (Ragunathan,
1928) Burkholder, 1948. {Bacterium betle
Ragunathan, Ann. Roy. Gard., Peradeniya,
Ceylon, 11, 1928, 51; Burkholder, in Man-
ual, 6th ed., 1948, 130.)
bet'le. Malaj-an noun betle betel; M.L.
indeclin.noun betle.
Rods, 0.5 by 1.5 to 2.5 microns, occurring
singly or in short chains. Non-motile. Gram-
negative.
Green pigment formed in nutrient gelatin
and in broth.
Gelatin: Liquefied.
Bovril agar colonies: Honey-yellow, cir-
cular at first, later echinulate. Raised,
smooth and shiny.
Bi-oth: Surface becomes cloud.y in 2
days. Pellicle.
No gas from lactose, maltose or sucro.se.
Starch is reduced.
Aerobic.
Source: Five cultures isolated from leaf
spots on the betel vine.
Habitat: Pathogenic on the betel vine,
Piper betle.
104. Pseudomonas gladioli Severini,
1913. (Annali d. Bot., Rome, 11, 1913, 420.)
gla.di'o.li. L. gladiolus a small sword
lily; M.L. mas.n. Gladiolus generic name of
gladiolus; M.L. gen. noun gladioli of gladio-
lus.
Rods 0.6 by 2.3 to 2.8 microns. Motile
with one or more polar flagella. Gram-nega-
tive.
A pale yellow, water-soluble pigment
found, later orange.
Gelatin colonies: Cream-colored, wart-
like. Rapid liquefaction.
Milk: Coagulated and slowly peptonized.
Nitrites not produced from nitrates.
Indole not produced.
No gas.
Aerobic.
Optimum temperature between 28° and
30° C.
Habitat: Causes a corm rot of gladiolus
and other tubers.
105. Pseudomonas mellea Johnson,
1923. (Jour. Agr. Res., 23, 1923, 489.)
mel'le.a. L. adj. melleus pertaining to
honey.
Rods 0.6 by 1.8 microns. Encapsulated.
Motile with 1 to 7 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Liquefied.
Potato-glucose agar: Growth abundant,
smooth, glistening, viscid, honey-colored.
Broth: Turbid in 24 hours. Pellicle.
Milk: Alkaline; clears.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Starch hydrolysis feeble.
Growth inhibited by 4 per cent salt.
Temperature relations: Optimum be-
tween 26° and 28° C. Maximum, 36° C.
Aerobic, facultative.
Distinctive character: Differs from Pseu-
domonas pseudozoogloeae in that it produces
on tobacco a brown instead of a black spot
with a halo, is orange-yellow in culture, and
turns milk alkaline.
Source: Isolated from brown rusty spots
on tobacco in Wisconsin.
Habitat: Pathogenic on leaves of tobacco,
Nicotiana tobacum.
106. Pseudomonas panacis (Nakata
and Takimoto, 1922) Dowson, 1943. {Bac-
terium panaxi Nakata and Takimoto, Bull.
Agr. Sta. Chosen, 5, 1922, 1 ; Dowson, Trans.
Brit. Mycol. Soc, S6, 1943, 10.)
pa'na.cis. Gr. panax the plant heal-all;
M.L. neut.noun Panax a generic name;
M.L. gen. noun panacis of Panax.
Description from Elliott (Bact. Plant
Pathogens, 1930, 173).
Rods 0.5 by 1.3 to 1.5 microns. Chains.
Motile with 4 to 6 polar flagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: Slight liquefaction.
Agar colonies: White.
Milk: Coagulated.
No gas from sugars.
138
ORDER I. PSEUDOMONADALES
Habitat: Causes a root rot of ginseng,
Panax quinquefoliitm.
107. Pseudomonas ribicola Bohn and
Maloit, 1946. (Jour. Agr. Res., 73, 1946, 288.)
ri.bi'co.la. M.L. noun Ribes generic name
of currant; L. colo to dwell; M.L. fera.n.
ribicola the currant dweller.
Rods, 0.4 to 0.9 by 0.9 to 1.7 microns,
occurring singly, in pairs and in hypha-like
chains. Motile by 1 or more polar flagella.
Gram-negative.
Gelatin: Very slow liquefaction.
Beef -extract agar colonies: Punctiform,
smooth, translucent, white; edges entire.
Beef -extract agar slant: Growth scant,
filiform, glistening, translucent, white,
slightly viscid.
Broth: Slightly turbid; no ring or pellicle.
Potato dextrose slants: Growth moder-
ate, filiform, glistening, butyrous to viscid.
Medium slightly yellow. Dirty pink pig-
ment in old cultures.
Milk: Slightly darkened, becoming alka-
line.
Nitrites produced from nitrates.
Growth good in Uschinsky's and Fermi's
solutions; yellow-green pigment produced.
No growth in Cohn's and Ashby's mannitol
solutions.
Indole not produced.
Hydrogen sulfide not produced.
Acid from glucose, galactose, fructose,
xylose and mannitol.
Starch not hydrolyzed.
Asparagine utilized as a carbon-nitrogen
source. Tj^rosine oxidized.
Not lipolytic.
Temperature relations: Optimum, be-
tween 20° and 25° C. Minimum, less than
3.5° C. Maximum, between 30° and 32° C.
Source: Six single-cell isolates from leaf
spot of golden currant in Wyoming.
Habitat: Pathogenic on Ribes aureum.
108. Pseudomonas xanthochlora (Schuster,
1912) Stapp, 1928. {Bacterium xanthochlorxim
Schuster, Arbeit, a. d. Kaiserl. Biolog.
Anstalt. f. Land. u. Forstw., 8, 1912, 452;
Stapp, in Sorauer, Handbuch der Pflanzen-
krankheiten, 2, 5 Aufl., 1928, 213.)
xan.tho.chlo'ra. Gr. xanthus yellow; Gr.
chlorus green; M.L. adj. xanthochlorvs yel-
lowish green.
Description from Erw. Smith (Bacteria
in Rel. to Plant Dis., 3, 1914, 272).
Rods 0.75 to 1.5 by 3.0 microns. Motile
with 1 to 3 polar flagella. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Slow liquefaction.
Agar colonies: Circular, slightly raised,
yellow-white.
Broth: Strong clouding in 24 hours. A
white pellicle.
Milk: Slow coagulation and clearing.
Nitrites produced from nitrates.
Indole produced after 10 days.
Hydrogen sulfide produced slowly.
Acid but no gas from glucose and galac-
tose.
Temperature relations: Optimum, 27° C.
Minimum, 2° C. Maximum, 44° C.
Source: Isolated from rotting potato
tubers in Germany.
Habitat : Pathogenic on potato tubers and
a number of unrelated plants.
109. Pseudomonas aleuritidis (McCul-
loch and Demaree, 1932) Stapp, 1935. (Bac-
terium aleuritidis McCulloch and Demaree,
Jour. Agr. Res., 43, 1932, 339; Stapp, Bot.
Rev., 1, 1935,408.)
a.leu.ri'ti.dis. Gr. aleurites of wheaten
flour; M.L. fem.noun Aleurites generic
name; M.L. gen. noun aleuritidis of Aleu-
rites.
Rods 0.6 to 0.7 by 1.1 to 3.0 microns.
Motile with 1 to 5 polar, rarely bipolar,
flagella. Encapsulated. Gram-negative.
Green fluorescent pigment produced in
certain media.
Gelatin: Not liquefied.
Beef agar slants: Growth is thin, white
and viscid.
Broth: A heavy white surface growth in
24 hours. Sediment.
Milk: Becomes alkaline, but no separa-
tion.
Nitrites produced from nitrates.
Indole test feebly positive.
Hydrogen sulfide test feebly positive.
Acid but no gas from glucose, galactose
and glycerol. Slow acid production from
sucrose, maltose and lactose.
FAMILY IV. PSEUDOMONADACEAE
139
Starch hydrolysis feeble.
Temperature relations : Optimum between
27° and 28° C. Maximum, 37° C.
Chemical tolerance: Optimum pH be-
tween 6.2 and 6.8. pH range, 5.4 to 8.9.
Source: Isolations from naturally infected
timg oil trees in Georgia.
Habitat: Pathogenic on the tung oil tree
(Aleurites Jordi), on the bean {Phaseolus
vulgaris) and the castor bean (Ricinus com-
munis) .
110. Pseudoinonas glycinea Coerper,
1919. (Jour. Agr. Res., 18, 1919, 188.)
gly.ci'ne.a. Gr. glycys sweet; M.L.
f em. noun Glycine generic name of a legume ;
M.L. adj. glycinea of the soybean.
Rods 1.2 to 1.5 by 2.3 to 3.0 microns.
Motile with polar flagella. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: Not liquefied.
Beef -peptone agar colonies: Appear in
24 hours. Circular, creamy white, smooth,
shining and convex. Margins entire. Buty-
rous in consistency.
Milk: Litmus turns blue and later a sepa-
ration of the milk occurs. Casein not di-
gested.
Nitrites produced from nitrates (Burk-
holder and Starr, Phytopath., 38, 1948, 498).
Indole test feebly positive.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid from glucose and sucrose.
Starch not hydrolyzed.
Temperature relations: Optimum be-
tween 24° and 26° C. Minimum, 2° C. Maxi-
mum, 35° C.
Aerobic, facultative.
Comment: A variety of this species that
differs slightly in morphology, action in
milk and in chromogenesis has been de-
scribed by Takimoto (Jour. Plant Prot.,
Tokyo, 14, 1927 556). It was isolated from
leaf spots on soy bean in Formosa.
Source: A number of cultures isolated
from soybeans in Wisconsin.
Habitat: Pathogenic on soybean. Glycine
max {Soja niax) .
111. Pseudonionas savastanoi (Erw.
Smith, 1908) Stevens, 1913. (Bacterium
savastanoi Erw. Smith, U. S. Dept. Agr.
Plant Ind. Bull. 131, 1908, 31; Stevens, The
Fungi which Cause Plant Diseases, 1913,
33.)
sa.vas.ta'no.i. Savastano patronymic;
savastanoi of Savastano.
Description from Brown (Jour. Agr. Res.,
U, 1932, 711).
Rods 0.4 to 0.8 by 1.2 to 3.3 microns.
Motile with 1 to 4 polar flagella. Gram-
negative.
Green fluorescent pigment found in cul-
ture.
Gelatin: No liquefaction.
Beef agar colonies: White, smooth, flat,
glistening, margins erose or entire.
Broth: Turbid on the second day. No
pellicle or ring.
Milk: Becomes alkaline.
Nitrites not produced from nitrates.
Hydrogen sulfide not produced.
Acid but no gas from glucose, galactose
and sucrose.
Starch hydrolyzed.
Temperature relations: Optimum be-
tween 23° and 24° C. Minimum, 1° C. Max-
imum, 32° C.
Chemical tolerance: Optimum between
6.8 and 7.0. Minimum, 5.6. Maximum, 8.5.
Aerobic.
Comment: A variety that differs but
slightly from this species is described as
pathogenic on ash, Fraxinus excelsior and F.
americana, but not on olive. Produces a
canker on ash. See Brown (Jour. Agr. Res.,
U, 1932, 721).
Source: Smith isolated his cultures from
olive galls collected in California.
Habitat: Pathogenic on olive.
112. Pseudoinonas tonelliana (Ferra-
ris, 1926) Burkholder, 1948. {Bacterixim to-
nellianum Ferraris, Trattato di Patologia e
TerapiaVegetale,3rded.,/, 1926, 104; Burk-
holder, in Manual, 6th ed., 1948, 132.)
to.nel.li.a'na. M.L. adj. tonellianus per-
taining to Tonelli; named for A. Tonelli.
Description from C. O. Smith (Phyto-
path., 18, 1928, 503) unless otherwise noted.
Rods 0.5 to 0.6 by 1.5 to 2.5 microns.
Motile with 1 to 3 polar flagella. Gram-
negative (Adam and Pugsley, Jour. Dept.
Agr. Victoria, 32, 1934, 304).
140
ORDER I. PSEUDOMONADALES
Gelatin: No liquefaction.
Potato glucose agar colonies: Flat, circu-
lar, shining; margins somewhat undulated.
Broth: Dense clouding with partial pel-
licle.
Milk: Alkaline. No separation.
Nitrites not produced from nitrates
(Adam and Pugsley).
Indole produced. Not produced (Adam
and Pugsley).
Acid but no gas from glucose and sucrose.
No acid from lactose (Adam and Pugsley).
Starch not hydrolyzed (Adam and Pugs-
ley).
Comment: Pseudomonas savastanoi is
similar in culture but is not pathogenic on
oleanders.
Source: Both Ferraris and C. O. Smith
isolated the pathogen from galls on olean-
der.
Habitat: Pathogenic on oleander, Nerium
oleander.
113. Pseudomonas cissicola (Takimoto,
1939) Burkholder, 1948. (Aplanobacter cissi-
cola Takimoto, Ann. Phytopath. Soc. Japan,
9, 1939, 43; Burkholder, in Manual, 6th ed.,
1948, 134.)
cis.si'co.la. Gr. cissus ivy; M.L. fem.noun
Cissiis generic name of flowering plant;
L. -cola dweller; M.L. fem.noun cissicola
Cissus dweller.
Rods 0.5 to 0.9 by 1.0 to 2.0 microns.
Non-motile. Encapsulated. Gram-negative.
Green fluorescent pigment formed in
Uschinsky's solution.
Gelatin: No liquefaction.
Potato-extract agar colonies: Circular,
conve.x, smooth, dirty white.
Broth: Feeble clouding followed by pre-
cipitation of pellicle and rim.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No acid nor gas from sucrose, glucose, lac-
tose or glycerol.
Starch not hydrolyzed.
Salt toleration, 3 per cent.
Temperature relations: Optimum, 30° C.
Minimum, 10° C. Maximum, 35° C. Ther-
mal death point between 49° and 50° C.
Source: Isolated from black spots on
leaves of Japanese ivy, Cissus japonica, in
Japan.
Habitat: Pathogenic only on Cissus ja-
ponica.
114. Pseudomonas ealendulae (Taki-
moto, 1936) Dowson, 1943. (Bacterium
ealendulae Takimoto, Ann. Phytopath. Soc.
Japan, 5, 1936, 341; Dowson, Trans. Brit.
Mycol. Soc, 26, 1943, 9.)
ca.len'du.lae. L. fem.pl. noun calendae
(Kalendae) The Calends, first day of month;
M.L. fem. dim. noun Calendula generic name
of a flowering plant; M.L. gen. noun ealen-
dulae of Calendula.
Rods 0.5 by 1.0 to 2.0 microns. Motile
with 1 to 3 polar flagella. Gram-negative.
Green fluorescent pigment produced in
Uschinsky's and in Cohn's solutions.
Gelatin: Not liquefied.
Agar colonies: Circular, smooth, flat,
dirty white.
Broth: Turbid.
Milk: No coagulation.
Nitrites not produced from nitrates.
Indole produced in small amount.
Hydrogen sulfide not produced.
Acid but no gas from glucose and glycerol.
No acid from lactose or sucrose.
Starch not hydrolyzed.
Temperature relations: Optimum be-
tween 27° and 30° C. Minimum between 0°
and 7° C. Maximum, 37° C.
Habitat: Pathogenic on marigolds, Calen-
dula officinalis.
115. Pseudomonas cichorii (Swingle,
1925) Stapp, 1928. {Phjtomonas cichorii
Swingle, Phytopath., 15, 1925, 730; Stapp, in
Sorauer, Handbuch der Pfianzenkrank-
heiten, 2, 5 Aufll., 1928, 291; Pseudomonas
endiviae Kotte, Phyt. Ztschr., 1, 1930, 609;
Bacterium formosanum Okabe, Jour. Soc.
Trop. Agr., Formosa, 7, 1935, 65.)
ci.cho'ri.i. Gr. cichora (pi.) succory, chi-
cory; L. cichoriutn chicory; M.L. neut.noun
Cichoriutn generic name of flowering plant;
M.L. gen. noun cichorii of chicory.
Description from Clara (Cornell Agr.
Exp. Sta. Mem. 159, 1934, 26) which is a de-
scription of a culture of Pseudomonas endi-
viae from Kotte. Swingle's description is
very meager.
FAMILY IV. PSEUDOMONADACEAE
141
Rods 0.75 to 1.5 by 1.5 to 3.75 microns.
Motile with 1 or 2 polar fiagella. Gram-
negative.
Green fluorescent pigment produced in
culture.
Gelatin: No liquefaction.
Beef -extract agar colonies: Circular,
grayish white with bluish tinge, raised with
slightly irregular edges.
Broth: Turbid in 36 hours with a smooth,
viscous pellicle.
Milk: Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
H3'drogen sulfide not produced.
Not lipolytic (Starr and Burkholder, Phj^-
topath., 32, 1942, 601).
Acid but no gas from glucose, galactose
fructose, mannose, arabinose, xylose, man-
nitol and glycerol. Alkaline production from
salts of acetic, citric, lactic, malic, succinic
and tartaric acids. Rhamnose, maltose,
sucrose, lactose, raffinose and salicin not
utilized.
Starch not hydrolyzed.
Slight growth in broth plus 6 per cent
NaCl.
Chemical tolerance: Optimum pH, be-
tween 6.8 and 7.1. Minimum, between 5.0
and 5.3. Maximum, between 9.2 and 9.4.
(Kotte, op. cit., 2, 1930, 453).
Aerobic, facultative.
Source: Isolated from rot of French en-
dive, Cichorium intybus, by Swingle and by
Okabe, and from C. endivia by Kotte.
Habitat: Pathogenic on endive, lettuce
and larkspur.
116. Pseiidomonas nectarophila
(Doidge, 1917) Rosen and Bleeker, 1933.
{Bacterium nectar ophilum Doidge, Ann.
Appl. Biol., J^, 1917, 73; Rosen and Bleeker,
Jour. Agr. Res., ^6, 1933, 98.)
nee. ta.ro 'phi. la. Gr. nectar nectar; Gr.
adj. philus loving; M.L. adj. nectarophilns
nectar-loving.
Rods 0.5 to 0.7 by 0.6 to 1.5 microns.
Motile with 1 to 5 polar flagella. Encapsu-
lated. Gram-negative.
Green fluorescent pigment produced in
culture.
Gelatin: No liquefaction.
Nutrient agar colonies: Yellowish white,
wet-shining, smooth; margins irregular.
Broth: Heavy turbidity in 24 hours.
Sediment.
Milk: Cleared.
Nitrites not produced from nitrates.
Indole not produced.
Acid from glucose and galactose. No acid
from sucrose.
Starch hydrolysis feeble.
Optimum temperature between 25° and
30° C.
Aerobic, facultative.
Distinctive character : Differs from Pseu-
domonas barkeri in that it does not liquefy
gelatin nor produce indole. Produces cap-
sules.
Source : Isolated from blighted pear blos-
soms in South Africa.
Habitat: Pathogenic on pear blossoms.
117. Pseudomonas viburni (Thornberry
and Anderson, 1931) Stapp, 1935. (Phyto-
monas viburni Thornberry and Anderson,
Phytopath., 21, 1931, 912; Stapp, Bot. Rev.
1, 1935,407.)
vi.bur'ni. L. viburnum waj' -faring tree;
M.L. neut.noun Viburnum name of a genus
of flowering plants; M.L. gen. noun viburni
of Viburnum.
Rods 0.5 to 1.0 by 1.0 to 2.0 microns.
Encapsulated. Motile with 2 to 4 polar
flagella. Gram-negative (Burkholder); not
Gram-positive as stated in original.
Green fluorescent pigment produced in
culture (Burkholder).
Gelatin: No liquefaction.
Glucose beef-extract colonies: Dull gray,
circular, edges entire.
Broth: Turbid with pellicle.
Milk: Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 601).
Acid from glucose and galactose but not
sucrose (Burkholder).
Starch: No hydrolysis.
Slight growth in 3.5 per cent salt (Burk-
holder).
Temperature relations: Optimum, 25° C.
Minimum, 12° C. Maximum, 35° C.
142
ORDER I. PSEUDOMONADALES
Aerobic.
Source: Isolated from angular leaf spots
and stem lesions on arrow-wood, Viburnum
opulus, etc.
Habitat: Pathogenic on Viburnum spp.
118. Pseudomonas mori (Boyer and
Lambert, 1893) Stevens, 1913. {Bacterium
mori Boyer and Lambert, Compt. rend.
Acad. Sci., Paris, 117, 1893, 342; Bacterium
mori Boyer and Lambert emend. Erw. Smith,
Science, 31, 1910, 792; Stevens, The Fungi
which Cause Plant Diseases, 1913, 30.)
mo'ri. Gr. morum the black mulberry;
M.L. fem.noun Morus the generic name of
mulberry; M.L. gen. noun mori of the mul-
berry.
Description from Smith (op. cit., 1910,
792).
Rods 0.9 to 1.3 by 1.8 to 4.5 microns.
Motile with a polar flagellum. Gram-nega-
tive.
Green fluorescent pigment produced in
culture.
Gelatin: Not liquefied.
Agar colonies: White, slow-growing,
smooth, flat; edges entire, becoming undu-
late.
Milk: Becomes alkaline and clears.
Nitrites not produced from nitrates.
Indole test negative or feebly positive.
Hydrogen sulfide not produced (Okabe,
Jour. Soc. Trop. Agr., 5, 1933, 166).
No growth in broth plus 4 per cent salt
(Okabe, loc. cit.).
No gas from carbohydrates.
Temperature range, 1° C. to 35° C.
Source: Smith isolated the pathogen from
blighted shoots of mulberry from Georgia.
Also received cultures from Arkansas and
the Pacific Coast.
Habitat: Pathogenic on mulberry, Morus.
119. Pseudomonas stizolobii (Wolf, 1920)
Stapp, 1935. (Aplanobactcr stizolobii Wolf,
Phytopath., 10, 1920, 79; Stapp, Bot. Rev.,
1, 1935, 405.)
sti.zo.lo'bi.i. Gr. stizo to prick, tat-
too; Gr. dim. labium a small lobe; M.L.
neut.noun Stizolobium plant generic name;
M.L. gen. noun stizolobii of Stizolobium.
Rods 0.6 to 0.7 by 1.0 to 1.6 microns.
Non-motile (Wolf, op. cit., 1920, 79). Motile
with a short polar flagellum (McCulloch,
Phytopath., 18, 1928, 460). Encapsulated.
Gram-negative.
Gelatin: No liquefaction.
Agar colonies: Circular, smooth, white,
raised and opaque. Margins entire to
slightly undulate.
Broth: Slightly turbid throughout. No
pellicle or ring.
Milk: Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
No acid or gas in peptone broth plus
sugars.
Starch not hydrolyzed.
Optimum temperature between 25° and
28° C.
Distinctive characters: Differs from
Pseudomonas sojae (Pseudomonas glycinea)
in the smaller size of cell, the absence of a
pellicle and dense clouding of broth. The
pathogen does not infect soybean.
Source: Isolated from the leaf spot of
velvet bean.
Habitat: Pathogenic on velvet bean,
Stizolobium deeringianium.
120. Pseudomonas viciae Uyeda, 1915.
(Uyeda, in Takimoto, Jour. Plant Protect.,
Japan, 2, 1915, 845.)
vi'ci.ae. L. vicia vetch; M.L. fem.noun
Vicia generic name of vetch; M.L. gen.noim
viciae of vetch.
Rods 0.5 to 0.8 by 1.2 to 2.0 microns.
Motile with 2 to 4 polar flagella. Reported
as Gram-positive; however, probably in
error. No cultures are available for a retest
of this character.
Green fluorescent pigment produced in
culture.
Gelatin colonies: Pale white, glistening,
finally turning brown. No liquefaction.
Milk: Coagulates and clears.
Nitrites not produced from nitrates.
Hj'drogen sulfide not produced.
Aerobic, facultative.
Habitat: Pathogenic on the broad bean
(Vicia f aba), the turnip (Brassica rapa), the
carrot (Da^lcus carota) and the sweet potato
(Ipomoea batatas) .
121. Pseudomonas alliieola (Burk
holder, 1942) Starr and Burkholder, 1942.
FAMILY IV. PSEUDOMONADACEAE
143
(Phytomonas alliicola Burkholder, Phyto-
path., 32, 1942, 146; Starr and Burkholder,
Phytopath., ibid., 601.)
al.li.i'co.la. L. allium onion; L. -cola
dweller; M.L. fern. noun alliicola onion
dweller.
Rods 0.7 to 1.4 by 1.05 to 2.8 microns.
Motile with 1 to several polar flagella, at
times bipolar. Gram-negative.
Gelatin: Liquefied.
Beef -extract peptone agar streaks : Mod-
erate in growth, white at first, later dirty
in appearance, edges wavy, consistency vis-
cid. Medium deep brown.
Potato-glucose agar frequently becomes
greenish.
Broth: Turbid with light pellicle. Brown.
Milk: Cleared and litmus reduced. Neu-
tral.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Lipolytic action very strong.
Acid but no gas from 1-arabinose, d-xy-
lose, rhamnose, glucose, d-galactose, fruc-
tose, d-lactose, maltose, sucrose, glycerol,
mannitol and salicin. Alkali from salts of
acetic, citric, formic, hippuric, lactic, malic,
succinic and tartaric acids.
Starch not hydrolyzed.
Slight growth in broth plus 4 per cent
salt.
Aerobic.
Temperature relations: Optimimi, 30° C.
Minimum, 5° C. Maximum, 41° C.
Source: Seven isolates from storage rot
of onion bulbs.
Habitat: Pathogenic on onion bulbs, Al-
lixun cepa.
122. Pseiidonionas gardeniae Burk-
holder and Pirone, 1941. (Phytopath., 31,
1941, 194.)
gar.de'ni.ae. L. Garden patronymic;
M.L. fem.noun Gardenia plant generic
name; M.L. gen. noun gardeniae of Gardenia.
Rods 0.75 by 2.4 microns. Motile with
1 to 2 polar flagella. Gram-negative.
Gelatin: Liquefied.
Beef -extract peptone agar colonies:
Growth fair, white to dirty gray and viscid.
Medium becoming dark brown.
Potato-glucose agar: No brown color.
Broth: Turbid with pellicle. Dark brown.
Milk: Soft curd with pellicle. Clears in
zones. Litmus reduced.
Nitrites produced from nitrates.
Hydrogen sulfide not produced.
Indole not produced.
Acid from glucose, galactose, xylose,
rhamnose, sucrose, maltose, mannitol,
glycerol and salicin. Alkali produced from
the salts of citric, malic, malonic, succinic,
tartaric and hippuric acids. Good growth
in tyrosine and in asparagine broth.
Starch not hydrolyzed.
Aerobic.
Source: Eight isolates from leaf spots of
gardenias in New Jersey.
Habitat : Pathogenic on leaves of Gardenia
jasminoides.
123. Pseudonionas caryophylli (Burk-
holder, 1942) Starr and Burkholder, 1942.
{Phytomonas caryophylli Burkholder, Phy-
topath., 32, 1942, 143; Starr and Burk-
holder, ibid., 601.)
ca.ry.o'phyl.li. Gr. caryophyllum nut
leaf, the clover tree; M.L. mas. noun
caryophyllus specific epithet in Dianthus
caryophyllus , the clove-pink or carnation;
M.L. gen. noun caryophylli of the carnation.
Rods 0.35 to 0.95 by 1.05 to 3.18 microns.
At times slightly curved. Motile with 1 to
several polar flagella. Frequently bipolar.
Gram-negative.
Gelatin: Liquefaction after 3 to 4 weeks.
Potato glucose agar colonies: 3 to 4 mm in
diameter, circular, smooth, glistening,
edges entire. Color is tan to gray-mauve.
Old culture dark brown. Consistency buty-
rous.
Broth: Turbid with a white sediment.
Milk: Litmus slowly becomes blue. Slight
reduction at bottom of tube. No clearing.
Nitrites produced from nitrates. Also
ammonia and gas are produced in a syn-
thetic nitrate medium. Asparagine, KNO3
and NH4H2PO4 can be utilized.
Indole not produced.
Hydrogen sulfide not produced.
Lipolytic action slight to moderate.
Acid from 1-arabinose, d-xylose, rham-
nose, glucose, d-galactose, fructose, d-lac-
tose, maltose, sucrose, glj^cerol, mannitol
and salicin. Alkali with sodium salts of
144
ORDER I. PSEUDOMONADALES
acetic, citric, formic, hippuric, lactic, malic,
maleic, succinic and tartaric acids.
Starch not hydrolyzed.
Aerobic.
Temperature relations: Optimum be-
tween 30° and 33° C. Minimum, 5° C. or
less. Maximum, 46° C.
Slight growth in broth plus 3.5 per cent
salt.
Source: Isolated first by L. K. Jones and
later by W. H. Burkholder from dying car-
nation plants from Spokane, Washington.
Twelve isolates used in description.
Habitat: Pathogenic on roots and stalks
of the carnation, Dianthus caryophyllus .
35° and 37° C. Minimum, 10° C. Maximum,
41° C.
Pathogenicity readily lost in culture.
Comment: A variety that turns litmus
milk and cream red has been described by
Erw. Smith (Bact. in Relation to Plant
Diseases, S, 1914, 282). It was isolated by
J. A. Honing from diseased tobacco plants
in Medan, Sumatra.
Source: Isolated from brown-rot of
solanaceous plants.
Habitat: Soil pathogen in warm, moist
climates attacking numerous species of
plants, especially potato, tobacco and
tomato.
124. Pseudonionas solanacearuni (Erw.
Smith, 1896) Erw. Smith, 1914. {Bacillus
solanacearuni Erw. Smith, U. S. Dept. Agr.,
Div. Veg. Phys. and Path., Bull. 12, 1896,
10; Erw. Smith, Bacteria in Relation to
Plant Diseases, 3, 1914, 178.)
so.la.na.ce.a'rum. L. solarium the night-
shade; -aceae familial ending; M.L. fem.pl.
noun Solanaceae the nightshade family;
M.L. fem.pl. gen. n. solanacearuni of the
Solanaceae.
Rods 0.5 to 1.5 microns. Motile with a
polar flagellum. Gram-negative.
Gelatin: Nakata (Jour. Sci. Agr. Soc.
Tokyo, 29^, 1927, 216) states there are two
forms, one of which shows slight liquefac-
tion. The other shows no liquefaction.
Agar colonies: Small, irregular, roundish,
smooth, wet-shining, opalescent, becoming
brown.
Broth: Slight pellicle. Broth turns brown.
Milk: Cleared without precipitation of
casein.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced (Burk-
holder).
Glucose, sucrose, glycerol, sodium citrate,
peptone, tyrosine, asparagine and glutamic
acid are utilized (Mushin, Austral. Jour.
Expt. Biol, and Med., 16, 1938, 325).
Nitrogen sources utilized are ammonia,
nitrates (KNO3), asparagine, tyrosine, pep-
tone and glutamic acid, but not potassium
nitrite (Mushin, loc. cit.).
Starch not hydrolyzed.
Temperature relations: Optimum between
125. Pseudonionas castaneae (Kawa-
mura, 1934) SSvulescu, 1947. {Bacterium
castaneae Kawamura, Ann. Phytopath. Soc.
Japan, 3, 1934, 15; SSvulescu, Anal. Acad.
Romane, III, 22, 1947, 11.)
cas.ta'ne.ae. Gr. castanum the chest-
nut tree; L. castanea the chestnut; M.L.
fem.noun Castanea generic name of chest-
nut; M.L. gen. noun castaneae of the chest-
nut.
Rods 0.8 to 1.2 by 1.0 to 1.8 microns.
Motile with 1 to 5 polar flagella. Gram-nega-
tive.
Gelatin: Liquefied.
Beef agar colonies: White, circular, edges
slightly undulate, viscid.
Milk: No coagulation. Peptonized.
Acid but no gas from glucose, sucrose and
glycerol. No acid from lactose.
Temperature relations : Optimum between
25° and 27° C. Minimum, 3° C. Maximum,
35° C.
Aerobic, facultative.
Habitat: Causes water-soaked spotting
on leaves and shoots of chestnut, Castanea.
126. Pseudonionas passiflorae (Reid,
1939) Burkholder, 1948. {Phytomonas passi-
florae Reid, New Zealand Jour. Sci. and
Tech., 22, 1939, 264a; Burkholder, in Man-
ual, 6th ed., 1948, 138.)
pas.si.flo'rae. L. passio passion; L. flos,
floris a flower; M.L. fem.noun Passiflora
generic name of passion flower; M.L.
gen. noun passiflorae of the passion flower.
Rods 0.2 to 0.5 by 1.2 to 3.2 microns-
FAMILY IV. PSEUDOMO^^ADACEAE
145
Motile with 1 to 5 polar flagella. Encapsu-
lated. Gram-negative.
Gelatin: Liquefied.
Beef -peptone agar colonies: Small, flat,
.smooth, dry, shining, translucent, graj^ish
and butyrous.
Broth: Turbid in 4 days. Transient pel-
licle.
Milk: Slightly alkaline. No coagulation
nor clearing.
Nitrites not produced from nitrates. No
growth on synthetic nitrate agar.
Indole not produced.
Hydrogen sulfide not produced.
Acid reaction occurs in galactose, starch
and sucrose. No gas.
Starch not hj-drolyzed.
Source : From diseased leaves and fruit of
the passion fruit in New Zealand.
Habitat: Pathogenic on Passiflora edulis.
127. Pseudomonas seininuni Caylej',
1917. (Jour. Agr. Sci., 8, 1917, 461.)
se'mi.num. L. semen, sennnis seed; L.
gen.pl. semimim of seeds.
Rods 1.0 by 4.0 to 5.0 microns. Spore-
like bodies present. Encapsulated. Motile
with a single flagellum. Reported as Gram-
po.sitive; however, probablj^ in error. No
cultures are available for a retest of this
character.
Gelatin: Rapid liquefaction.
Agar colonies: White, more or less circu-
lar, transparent, spreading.
Broth: Turbid. Pellicle.
Litmus milk: Milk becomes clear and
apricot color.
Nitrites produced from nitrates.
Acid but no gas from glucose and sucrose.
No acid from lactose.
Starch: No hydrolysis.
Optimum temperature, 25° C.
Aerobic, facultative.
Source: Isolated from seeds, stems and
pods of diseased peas in England.
128. P.seiidonionas vitiswoodrowii Pa-
id and Kulkarni, 1951. (Pseudomonas ritis-
woodrowii (sic) Patel and Kulkai'ui, Curr.
Sci.,^0, 1951, 132.)
vi.tis.wood.ro'wi.i. L. fem.noun vitis
a vine; M.L. fem.noun Vitis a generic name;
Woodrow patronymic; M.L. gen. noun woo(/-
rowii of Woodrow.
Rods 0.8 by 1.5 microns. Motile with
a single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Potato dextrose agar colonies: Circular,
capitate with margins entire. Pale, dull
gray. 1.2 cm in diameter in 7 daj's.
Broth: Turbid.
Potato cylinders: Scant growth. Medium
dark gray.
Milk : Litmus reduced and casein digested.
Hydrogen sulfide produced.
Loefiler's blood serum: Liquefied.
Indole not produced.
Synthetic asparagine medium. No growth.
Methyl red test negative; acetylmethyl-
carbinol not produced.
Nitrites not produced from nitrates.
Acid but no gas from glucose, lactose and
sucrose, but no growth in salicin.
Salt tolerance: Up to 1 per cent.
Optimum temperature between 25° and
28° C.
Aerobic.
Source: Isolated from leaves of Vitis
woodrowii in India.
Habitat : Pathogenic on V. woodrowii but
not on V. vinifera.
129. Pseudomonas fabae (Yu, 1936)
Burkholder, 1948. (Phytomonas fabae Yu,
Bull, of the Chinese Bot. Soc, 2, 1936, 34;
Burkholder, in Manual, 6th ed., 1948, 139.)
fa'bae. L. faba the horse bean; M.L.
gen. noun /aftoe of the hor.se bean.
Rods 0.8 to 1.1 by 1.1 to 2.8 microns. Mo-
tile with 1 to 4 polar flagella. Gram-nega-
tive.
Gelatin: Liquefied.
Nutrient agar colonies: Circular, entire,
viscid, glistening, raised, smooth to
wrinkled, white to salmon. Medium amber.
Broth: Turbid after 12 hours. Pellicle.
Milk: Growth slow. Clears.
Nitrites produced from nitrates.
Indole production slight.
Hydrogen sulfide not produced.
Acid but no gas from glucose. No acid
nor gas developed from arabinose, xylose,
fructose, galactose, sucrose, lactose, mal-
tose, raffinose, dextrin, inulin, mannitol or
146
ORDER I. PSEUDOMONADALES
adonitol in a 1 per cent Bacto-peptone
broth.
Starch: Very weak diastatic action.
Temperature relations: Optimum, 35° C.
Minimum, 4° C. Maximum between 37° and
38° C. Thermal death point between 52° and
53° C.
Aerobic.
Growth retarded in 2 per cent salt. Very
slight growth in 3 per cent salt.
Source: From diseased broad beans at
Nanking, China.
Habitat : Pathogenic on broad or Windsor
bean, Vicia faba.
130. Pseudomonas astragali (Taki-
moto, 1930) Savulescu, 1947. {Bacterium
astragali Takimoto, Jour. Plant Protect.,
17, 1930, 732; SSvulescu, Anal. Acad. Ro-
mane. III, ££, 1947, 11.)
as.tra'ga.li. Gr. astragalus a vertebra,
also a leguminous plant; M.L. mas. noun
Astragalus a generic name; M.L. gen. noun
astragali of Astragalus.
Description translated by Dr. K. To-
gashi.
Rods 0.7 to 0.8 by 1.2 to 2.2 microns.
Motile, with 1 or 2 flagella. Gram-negative.
Gelatin: Liquefied.
Agar plates: Growth somewhat slow,
colorless or grayish white, entire margins,
more or less aqueous, butyrous.
Uschinsky's medium: Growth vigorous,
turbid, not viscid; ring and sediment.
Milk: No coagulation of casein, slow di-
gestion. Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced in small
amount.
No acid or gas from glucose, sucrose,
lactose or glycerol in broth.
Starch not hydrolyzed.
Temperature relations: Minimum, below
5° C. Maximum, 33° C. Thermal death
point between 50° and 51° C.
Aerobic.
Source: Species isolated from Astragalus
sp.
Habitat: Causes a black leaf -spot of
Astragalus sp.
131. Psevidomonas colurnae (Thorn-
berry and Anderson, 1937) Burkholder,
1948. {Phytomonas colurnae Thornberry and
Anderson, Phytopath., 37, 1937, 948; Burk-
holder, in Manual, 6th ed., 1948, 139.)
co.lur'nae. L. fem.noun corylus the hazel
or filbert; L. adj. colurnus (transposition of
corulnus) pertaining to hazel.
Rods 0.8 to 1.0 by 1.0 to 1.8 microns.
Single, in pairs or chains. Encapsulated.
Motile with 1 to 2 polar flagella. Gram-
negative.
Gelatin: Liquefied.
Glucose agar slants: Growth filiform,
raised, dull, smooth, opaque and viscid.
Broth: Moderate turbidity. Ring.
Milk: Peptonization complete with acid
production. No reduction of litmus nor
coagulation.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No appreciable amount of gas from
xylose, glucose, sucrose or glycerol.
Starch hydrolyzed.
Temperature relations: Optimum, 21° C.
Minimum, 5° C. Maximum, 35° C. Thermal
death point, 50° C.
Aerobic.
Source: From leaves and young stems of
the Turkish hazelnut in Illinois.
Habitat: Pathogenic on the Turkish
hazelnut, Corylus colurna.
132. Pseudomonas iridicola (Taki-
moto, 1931) Stapp, 1935. {Bacterium iridicola
Takimoto, Fungi, Nippon Fungological
Soc, 1, 1931, 24; Stapp, Bot. Rev., 1, 1935,
408.)
i.ri.di'co.la. Gr. iris, iridis the rainbow,
the plant iris; M.L. fem.noun 7ns generic
name; L. -cola dweller; M.L. fem.noun
iridicola iris dweller.
Rods 0.7 to 0.8 by 1.2 to 2 microns.
Motile with 1 to 3 polar flagella. Gram-
negative.
Gelatin: Liquefied.
Beef agar colonies: White, circular,
raised or convex.
Milk: Clears without coagulation.
No acid or gas from carbohydrates.
Starch digested.
Temperature relations: Optimum, 38° C.
Minimum, 4° C.
FAMILY IV. PSEUDOMONADACEAE
147
Source : Isolated from a brown leaf spot of
iris.
Habitat : Pathogenic on Iris tectorum and
Iris japo7iica.
133. Pseudomonas levistici Oster-
walder, 1909. (Cent. f. Bakt., II Abt., 25,
1909, 260.)
le.vis'ti.ci. L. neut.noun ligusticum a
Ligurian plant, lovage; L. neut.noun le-
visticum a corruption of ligusticum; M.L.
neut.noun Levisticum generic name of
lovage; M.L. gen. noun levistici of Levisti-
cum.
Rods 0.5 to 0.7 by 1.1 to 1.5 microns.
Motile with a polar flagellum. Gram-nega-
tive.
Gelatin: Colonies greenish white. Li-
quefied.
Nutrient agar: Good growth at room
temperature. Yellowish white.
Broth: Pellicle.
Indole produced.
Hydrogen sulfide not produced.
Source : Isolated from spots on the leaves
of lovage.
Habitat: Pathogenic on lovage, Levisti-
cum officinale.
134. Pseudomonas niaublaucii (Foex
and Lansade, 1936) SHvulescu, 1947. {Bac-
terium maublancii Foex and Lansade,
Compt. rend. Acad. Sci., Paris, 202,
1936,2174; Savulescu, Anal. Acad. Romane,
III, 22, 1947, 11.)
mau.blan'ci.i. Maublanc, patronymic;
M.L. gen. noun maublancii of Maublanc.
Rods 0.4 by 1.3 microns. Motile with 1
to 3 polar flagella. Gram-negative.
Gelatin: Liquefied.
Gelatin colonies: Round, translucent,
margins entire.
Broth: Thin pellicle.
Milk: Not coagulated; clears.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Carbohydrates not fermented.
Ammonia produced.
Growth in Fermi's solution, not in Us-
chinsky's solution.
Source: Isolated from rotting vascular
and parenchymatic tissue of banana stalks.
Habitat: Causes a disease of the banana
plant.
135. Pseudomonas polygoni (Thorn-
berry and Anderson, 1937) Burkholder,
1948. (Phytomonas polygoni Thornberry
and Anderson, Phytopath., 27, 1937, 947;
Burkholder, in Manual, 6th ed., 1948, 140.)
po.ly'go.ni. Gr. polygonum knot weed;
M.L. neut.noun Polygonum generic name;
M.L. gen. noun polygoni of Polygonum.
Rods 0.5 to 1.5 by 1.5 to 2.5 microns.
Motile with 2 to 8 bipolar flagella. Encap-
sulated. Gram-positive (?). Other species
reported by these investigators as Gram-
positive have proved to be Gram-negative
on a retest (Burkholder).
Gelatin: Liquefied. Brown.
Glucose agar slant: Abundant, filiform,
flat, dull, smooth, pale olive-gray, bvity-
rous. Medium turns brown.
Broth: Turbid. Pellicle.
Milk: Alkaline and clears. Litmus not re-
duced.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No appreciable amount of gas from car-
bohydrates.
Starch: No hydrolysis.
Temperature relations: Optimum, 18° C.
Minimum, 7° C. Maximum, 35° C.
Aerobic.
Source: From diseased leaves of Poly-
gomim convolvulus in Illinois.
Habitat: Pathogenic on black bindweed,
Polygonum convolvulus.
136. Pseudomonas radiciperda (Javo-
ronkova, 1932) Savulescu, 1947. (Bacterium
radiciperda Javoronkova, Bull. Plant Pro-
tect., Leningrad, Ser. II, 5, no. 1, 1932, 161;
savulescu. Anal. Acad. Romane, III, 22,
1947, 11.)
ra.di.ci.per'da. L. radix, radicis root; L.
perdo to destroy; M.L. fem.noun radiciperda
the root destroyer.
Description from Javoronkova (Rev. App.
Myc, 11, 1932, 652).
Rods 0.8 by 1.0 to 2.0 microns. Encapsu-
lated. Motile by means of 1 or 2 polar fla-
gella. Gram-negative.
Gelatin: Liquefied.
148
ORDER I. PSEUDOMONADALES
Beef-peptone agar colonies: Round,
smooth, shining, white to pale yellow.
Milk: Peptonized.
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from carbohydrates.
Optimum temperature between 23° and
25° C.
Aerobic.
Habitat: Causes a root rot of red clover
{Trifolium praiense) , lentils (Lens escu-
lenta) and lucerne.
137. Pseudomonas cattleyae (Pava-
rino, 1911) SS,vulescu, 1947. {Bacterium
cattleyae Pavarino, Atti R. Acad. Naz.
Lincei Rend. CI. Sci. Fis., Mat. e Nat., 20,
1911, 233; Savulescu, Anal. Acad. Romane,
III, 22, 1947, 11.)
catt'ley.ae. M.L.fem.noun Cattleya a
generic name; M.L. gen. noun cattleyae of
Cattleya.
Description from Ark and Thomas (Phy-
topath., 36, 1946, 697).
Rods, 0.4 to 0.6 by 2.4 microns, occurring
singly or in pairs. Motile by means of 1 or
2 bipolar flagella. Gram-negative.
Gelatin: No liquefaction.
Beef -extract peptone agar colonies:
Large, entire, smooth, with criss-cross
markings. Grayish white and butyrous.
Broth: Turbid in 24 hours with very deli-
cate pellicle.
Fermi's, Cohn's, and Uschinsky's solu-
tions: Good growth.
Hydrogen sulfide not produced.
Indole not produced.
Nitrites produced from nitrates.
Litmus milk: Unchanged after 2 weeks.
Acid but no gas from glucose, galactose,
fructose, arabinose, xylose, lactose, sucrose,
dulcitol, glycerol and mannitol. No acid or
gas from raffinose.
Starch: Slight hydrolj'sis.
Optimum temperature between 25° and
35° C.
Source : Four isolates and 4 reisolates from
leaf spots of orchids by Ark and Thomas.
Habitat: Pathogenic on Cattleya sp. and
Phalaenopsis sp.
138. Pseudomonas dysoxyli Hutchin-
son, 1949. (New Zealand Jour. Sci. and
Tech., Sec. B, 30, 1949, 275.)
dy.so'xy.li. M.L. neut.noun Dysoxylum
generic name of a forest tree; M.L. gen. noun
dysoxyli of Dysoxylum.
Rods 0.4 to 0.6 by 0.6 to 1.0 micron.
Motile with 1 to 2 polar flagella. Gram-nega-
tive.
Gelatin: No liquefaction.
Beef -peptone agar colonies: Circular,
punctiform to 8 mm in diameter, gray,
translucent. Surface smooth and edges en-
tire to undulate. Medium brown in 1 week.
Nutrient broth: Dense flocculent pellicle
at surface. Strong clouding in 3 days.
Litmus milk: No change in 4 days. After
12 days amber whey at top and 1 inch of pink
precipitate at base.
Indole not produced.
Hydrogen sulfide not produced.
Methyl red test negative; acetylmethyl-
carbinol not produced.
Nitrites not produced from nitrates.
Synthetic medium: Acid but no gas from
glucose, fructose, lactose, sucrose, raffinose
and mannitol within 3 weeks. No acid from
arabinose, maltose, melizitose, starch,
inulin, dextrin, glycerol or salicin.
Starch hydrolj^zed.
Temperature relations: Optimum, 25° C.
Minimum, 4° C. Maximum, 36° C.
Source: Many cultures isolated from
diseased leaves of Dysoxylum spectabile in
New Zealand.
Habitat: Pathogenic on leaves of the
forest tree Dysoxylum spectabile.
139. Pseudomonas helianthi (Kawa-
mura, 1934) SSvulescu, 1947. {Bacterium
helianthi Kawamura, Ann. Phyt. Soc.
Japan, 4, 1934, 27; Savulescu, Anal. Acad.
Romane,III,^^, 1947, 11.)
he.li.an'thi. Gr. heliiis the sun; Gr.
anthus a flower; M.L. mas. noun Helianthus
generic name of sunflower; M.L. gen. noun
helianthi of the sunflower.
Rods 1.0 to 1.4 by 1.6 to 2.4 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: No liquefaction.
Beef agar colonies: White, circular, edges
entire.
Broth: Turbid. Pellicle.
Milk: Peptonized. Litmus reduced.
Nitrates: Gas production.
FAMILY IV. PSEUDOMONADACEAE
149
Indole not produced.
Hydrogen sulfide not produced.
Acid but no gas from sucrose and glycerol.
No acid from lactose and maltose.
Starch hydrolyzed.
Temperature relations : Optimum between
27° and 28° C. Minimum, 12° C. Maximum,
35.5° C.
Chemical tolerance: Good growth at pH
6.4. No growth at pH 5.4 and pH 8.8
Habitat: Pathogenic on sunflower, Heli-
anthus debilis.
140. Pseudonionas melophthora Allen
and Riker, 1932. (Phytopath., 22, 1932, 557.)
me.loph'tho.ra. Gr. melum apple; Gr.
phthora decaj^ destruction; M.L. adj.
rnelophthonis apple-destroying.
Rods 0.68 by 1.32 microns. Motile with 2
polar flagella. Gram-negative; Gram-posi-
tive cells appear in old cultures.
Gelatin : No liquefaction.
Nutrient agar plus 2 per cent glucose:
Colonies appear in 36 hours. After 3 days
colonies circular, smooth, glistening, con-
ve.x; edges entire; light pink, but not con-
stant.
Broth: Good growth. Pellicle and sedi-
ment.
Milk: Little change, if any.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Acid from arabinose, glucose, galactose,
fructose, sucrose and glycerol. No acid from
lactose, maltose, dextrin or inulin.
Starch not hydrolyzed.
Optimum temperature between 21° and
25° C.
Source: Description based on 7 cultures
isolated from rotting apples and from apple
maggots.
Habitat : Pathogenic on apples and found
with the apple maggot , Rhagoletis ponionella.
141. Pseudonionas alboprecipitans
Rosen, 1922. (Ann. Missouri Bot. Garden, 9,
1922, 383.)
al.bo.pre.ci'pi.tans. L. album the color
white; L. v. praecipito to precipitate; M.L.
part. adj. alboprecipitans forming a white
sediment.
Description revised in accordance with
Johnson, Roberts and Cash (Jour. Agr. Res.,
78, 1949, 723).
Rods, 0.6 by 1.8 microns, occurring singly
or in pairs. Encapsulated. Motile with a
polar flagellum. Gram-negative.
Gelatin: Liquefied
Nutrient agar colonies: White, circular,
raised, smooth, sticky, with margins entire.
Whitish discoloration of the medium.
Broth: Turbid in 24 hours. Heavy sedi-
ment in old cultures.
Uschinsky's solution: Turbid in 24 hours;
pellicle formed.
Cohn's and Fermi's solutions: No growth.
Milk: Becomes alkaline and slowly clears.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide production slight.
Acid but no gas from glucose, galactose,
fructose, sucrose, lactose, raflfinose, glycerol
and mannitol. No acid from maltose.
Starch hydrolyzed.
Temperature relations : Optimum between
30° and 35° C. Minimum, 0° C. Maximum,
40° C.
Aerobic.
Distinctive character: White precipitate
in culture media.
Source: Isolated a number of times from
foxtail grass.
Habitat: Pathogenic on foxtail, Chacto-
chloa hdescens, and other grasses.
142. Pseudonionas andropogonis (Erw.
Smith, 1911) Stapp, 1928. (Bacterium andro-
pogoni (sic) Erw. Smith, Bacteria in Relation
to Plant Diseases, 2, 1911, 63; Elliott and
Smith, Jour. Agr. Res., 38, 1929, 4; Pseudo-
nionas andropogoni (sic) Stapp, in Sorauer,
Handbuch der Pflanzenkrankheiten, 2, 5
Aufl., 1928,27.)
an.dro.po.go'nis. Gr. aner, andris a man;
Gr. mas. noun pogon, pogonis beard; M.L.
mas. noun Andropogon, -onis man's beard,
generic name; M.L. gen. noun andropogonis
of Andropogon.
Description from Elliott and Smith (op.
cit., 1929, 4).
Rods 0.64 by 1.76 microns. Motile with
one to several bipolar flagella. Encapsu-
lated. Gram-negative.
Gelatin: Feeble liquefaction or none.
Beef -extract agar colonies : Slow growing,
round, smooth, glistening, viscid, white.
150
ORDER I. PSEUDOMONADALES
Broth: Growth slow with moderate tur-
bidity in 48 hours. A thin pellicle.
Milk: Alkaline and clears.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 3£, 1942,601).
Acid but no gas from glucose, arabinose,
fructose and xylose. No acid from sucrose,
maltose, lactose, raffinose, glycerol or
mannitol.
Starch partially digested.
Temperature relations : Optimum between
22° and 30° C. Minimum, 1.5° C. Maximum
between 37° and 38° C.
Chemical tolerance: Optimum pH be-
tween 6.0 and 6.6. Minimum, 5.0. Maximum
between 8.3 and 8.6.
Source : Elliott used for her description 4
cultures isolated from lesions on sorgo,
sorghum and broom-corn.
Habitat: Pathogenic on sorghum, Holcus
sorghum.
143. Pseudomonas lignicola Westerdijk
and Buisman, 1929. (De lepenziekte, Arn-
hem, 1929, 51.)
lig.ni'co.la. L. lignum wood; L. -cola
dweller; M.L. fem.noun lignicola wood
dweller or inhabitant.
Rods. Single or short chains. Motile with
1 to several polar flagella. Gram-negative.
Gelatin: No liquefaction.
Malt agar streaks: Milk-w^hite with a
colorless edge.
Broth: Turbid with light pellicle.
Milk: No coagulation. No acid.
Nitrites not produced from nitrates.
Indole not produced.
Starch hydrolysis slight.
Optimum temperature, ±25° C.
Source : From vessels of elm wood showing
dark discoloration, in Holland.
Habitat: Pathogenic on elm wood.
144. Pseudomonas petasitis (Takimoto,
1927) Siivulescu, 1947. {Bacterium petasitis
(sic) Takimoto, Ann. Phyt. Soc. Japan,
2, 1927, 55; Savulescu, Anal. Acad. Romane,
III,^^, 1947, 11.)
pe.ta'si.tis. Gr. mas. noun petasus a
sombrero, a broad-brimmed felt hat; Gr.
mas. noun petasites a broad-leafed plant,
colt's foot; M.L. mas. noun Petasites generic
name; M.L. gen. noun petasitis of Petasites.
Rods 0.8 to 1.1 by 1.1 to 1.7 microns.
Motile with a polar flagellum. Gram-
negative.
Gelatin: No liquefaction.
Beef agar colonies: White, circular or
amoeboid, butyrous.
Broth: Strong turbidity. Pellicle.
Milk: Coagulated in 30 days.
Nitrites produced from nitrates with gas
formation.
Indole not produced.
Hydrogen sulfide not produced.
No evident acid in peptone broth, but gas
from glucose, lactose and sucrose. Acid but
no gas from glycerol.
Weak growth in broth plus 6 per cent salt.
Temperature relations : Optimum between
27° and 30° C. Minimum, approximately
5° C. Maximum, 47° C.
Source: Isolated from brown to black
lesions on Petasites japonicus in Japan.
Habitat : Pathogenic on leaves of Petasites
japonicus.
145. Pseudomonas woodsii (Erw. Smith,
1911) Stevens, 1925. {Bacterium woodsii
Erw. Smith, Bacteria in Relation to Plant
Diseases, 2, 1911, 62; Stevens, Plant Disease
Fungi, New York, 1925, 39.)
wood 'si. i. Named for A. F. Woods, an
American plant pathologist; M.L. gen. noun
woodsii of Woods.
Description from Burkholder and Guter-
man (Phytopath., 25, 1935, 118).
Rods 0.67 by 1.56 microns. Motile with a
polar flagellum. Gram-negative.
Gelatin: No liquefaction.
Beef -extract agar slants : Growth slow and
scant, filiform, creamy, butyrous.
Broth: Turbid.
Milk: Becomes alkaline but otherwise
little changed.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 3^, 1942,601).
Acid but no gas from glucose, fructose,
galactose, arabinose, xylose, rhamnose, lac-
tose, glycerol and mannitol. Alkaline reac-
FAMILY IV. PSEUDOMONADACEAE
151
tion from salts of acetic, citric, malic and
succinic acids. Sucrose, maltose, salicin, and
lactic and formic acids not fermented.
Starch not hydrol3-zed.
Slight growth in broth plus 3 per cent salt.
Aerobic.
Source: Isolated from water-soaked
lesions on carnation leaves.
Habitat: Pathogenic on carnation, Dian-
tfius canjophylbis.
146. Pseudonionas eriobotryae (Taki-
moto, 1931) Dowson, 1943. {Bacterium erio-
botryae Takimoto, Jour. Plant Protect., 18,
1931, 354; Dowson, Trans. Brit. Mycol. Soc,
26, 1943, 10.)
e.ri.o.bo'trj^.ae. Gr. erium wool; Gr.
hotrys grape cluster; M.L. fem.noun Erioho-
trya woolly grape, a generic name; M.L.
gen. noun eriobotryae of Eriobotrya.
Translated by Dr. K. Togashi.
Rods 0.7 to 0.9 by 2.2 to 3.0 microns.
Motile, with 1 or 2 flagella. Gram-negative.
Gelatin: Not liquefied.
Agar-plates : Colonies appear after 3 days,
white or hyaline, butyrous, margins entire.
Broth: Moderately turbid; pellicle pow-
dery; ring formed.
Milk: No coagulation, peptonized slowly.
Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No acid or gas from glucose, sucrose, lac-
tose or glycerol in broth.
Starch not hj^lrolyzed.
Temperature relations : Optimum between
25° and 26° C. Minimum, below 4° C. Maxi-
mum, 32° C. Thermal death point, 51° C.
Aerobic.
Source: Species isolated from loquat,
Eiiobotrya japonica.
Habitat: Causes a bud rot of Eriobotrya
japonica.
147. Pseudonionas panicimiliacei
(Ikata and Yamauchi, 1931) SSvulescu,
1947. {Bacterium panici-miliacei Ikata and
Yamauchi, Jour. Plant Protect., 18, 1931,
35; Pseudomonas panici-miliacei (sic) Siivu-
lescu. Anal. Acad. Romane, III, 22, 1947,
11.)
pa.ni.ci.mi.li.a'ce.i. L. panicum panic
grass; L. adj. miliaceus pertaining to millet;
Panicum miliaceum millet.
Description translated by Dr. K. Togashi.
Rods 0.8 to 1.1 by 1.8 to 2.6 microns.
Motile, with a single flagellum. Gram-
negative.
Gelatin: Not liquefied.
Potato-agar plates: Growth moderate,
whitish, then tinged with light orange, un-
dulating margins.
Broth: Turbid; white pellicle formed.
Milk: No coagulation and slow digestion.
Alkaline.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No acid and no gas from sucrose, glucose,
lactose, glycerol or sodium nitrate.
Starch not hydroh'zed.
Optimum temperature between 30° and
35° C.
Aerobic, facultative.
Source : Species first isolated from millet,
Panicum, miliaceum.
Habitat: Causes a leaf stripe of Panicum
miliaceum.
148. Pseudonionas saliciperda Lin-
deijer, 1932. (Inaug. Diss., Univ. Amster-
dam, 1932; Phytopath. Ztschr., 6, 1933, 373.)
sa.li.ci.per'da. L. salix, salicis willow; L.
perdo to destroy; M.L. fem.noun saliciperda
willow^ destroyer.
Rods 1.2 to 2.1 microns in length. Motile
with a polar flagellum. Gram-negative.
Gelatin: No liquefaction.
Beef wort agar colonies: Gray-white.
Milk: No acid nor coagulation.
Nitrites produced (small amount) from
nitrates.
Indole production slight.
No gas from carbohydrates.
Starch not hydrolyzed.
Aerobic, facultative.
Source: Isolated from wilted branches of
willow.
Habitat: Pathogenic on willow, Salix spp.
149. Pseudomonas wieringae (Elliott,
1930) Savulescu, 1947. {Phytomonas betae
Wieringa, Nederl. Tijdschr. Hyg., Micro-
biol, en Serol., Leiden, 2, 1927, 148; Bac-
152 ORDER I. PSEUDOMONADALES
terium wieringae Elliott, Man. Bact. Plant Rods 0.5 by 2.0 microns. Motile with 1 to
Pathogens, 1930, 264; Saviilescu, Anal. 5 polar flagella. Gram-negative.
Acad. Romane, III, 22, 1947, 11.) Beef -agar colonies: Smooth, round, white
wie'ring.ae. Named for Dr. K. T. Wie- to grayish, fluorescent,
ringa, the bacteriologist who first described Milk: Cleared in 5 days. Not coagulated,
the species; M.L. gen. noun wieringae of Nitrites not produced from nitrates.
Wieringa. No gas from sugars.
Because Bacterium betae Chester (Ann. Temperature relations : Optimum between
Rept. Del. Col. Agr. Exp. Sta., 9, 1897, 53) 28° and 30° C. Minimum, 4° C. Maximum,
may be a pseudomonad, the more distinctive 37° C.
species name proposed by Elliott has been Source: Isolated from vascular rot of
retained. beets in Holland.
Description from Elliott (op. cit., 1930, Habitat: Pathogenic on beets, Beta vul-
264). garis.
Genus II. Xanthomonas Dowson, 1939 *
{Phijtomonas Bergey et al., Manual, 1st ed., 1923, 174; not Phytomonas Donovan, Lancet,
177, 1909, 1495 (type species (monotypy) Phytomonas davidi Donovan, a flagellate);
Dowson, Zent. f. Bakt., II Abt., 100, 1939, 187.)
Xan.tho'mo.nas or Xan.tho.mo'nasf. Gr. adj. xanthus yellow; Gr. fem.n. monas unit,
monad; M.L. fem.n. Xanthomonas yellow monad.
Cells usually monotrichous. A yellow, non-water-soluble pigment is produced on agar. A
diffusible, brown color infrequently occurs in beef extract agar. Proteins are usually readily
digested. Milk usually becomes alkaline. Hydrogen sulfide is produced. Asparagine is not
sufficient as an only source of carbon and nitrogen. Acid (and also gas in one species, No.
19) produced from mono- and disaccharides. Some species liquefy a pectin medium, others
do not (Burkholder and Starr, Phytopath., 38, 1948, 500). Mostly plant pathogens causing
necroses.
The type species is Xanthomonas hyacinthi (Wakker) Dowson.
Key to the species of genus Xanthomonas.
I. Colonies yellow; pigment non -water-soluble.
A. Gelatin liquefied.
1. Starch hydrolysis feeble.
a. Nitrites not produced from nitrates.
1. Xanthomonas hyacinthi.
2. Xanthomonas pruni.
3. Xanthomonas vitians.
aa. Nitrites produced from nitrates.
4. Xanthomonas beticola.
5. Xanthomonas rubrilineans .
2. Starch hydrolysis strong.
a. Nitrites not produced from nitrates.
b. No brown pigment in beef -extract agar.
6. Xanthomonas barbareae.
7. Xanthomonas begoniae.
8. Xanthomonas betlicola.
9. Xanthomonas campestris.
* Prepared by Prof. Walter H. Burkholder, Cornell University, Ithaca, N. Y., June,
1943; revised November, 1953.
t The former accords with the Latin rules of accentuation; the latter is in common usage.
FAMILY IV. PSEUDOMONADACEAE 153
10. Xanthomonas cassiae.
11. Xanthomonas cajani.
12. Xanthomonas citri.
13. Xanthomonas derodendri.
14. Xanthomonas corylina.
15. Xanthomonas cucurbitae.
16. Xanthomonas desmodii.
17. Xanthomonas desmodiigangeticii.
18. Xanthomonas dieffenbachiae .
19. Xanthomonas hemmiana.
20. Xanthomonas holcicola.
21. Xanthomonas incanae.
22. Xanthomonas juglandis.
23. Xanthomonas lespedezae.
24. Xanthomonas macidifolngardeniae.
25. Xanthomonas malvacearum.
26. Xanthomonas pelargonii.
27. Xanthomonas phaseoli.
28. Xanthomonas plantaginis .
29. Xanthomonas ricinicola.
30. Xanthomonas sesbaniae.
31. Xanthomonas stizolobiicola.
32. Xanthomonas taraxaci.
33. Xanthomonas translucens.
34. Xanthomonas uppalii.
35. Xanthomonas vasculorum.
36. Xanthomonas vesicatoria.
37. Xanthomonas vignicola.
bb. Brown pigment produced in beef-extract media.
38. Xanthomonas nakatae.
27. Xanthomonas phaseoli.
aa. Nitrites produced from nitrates.
39. Xanthomonas papavericola.
aaa. Ammonia produced from nitrates.
40. Xanthomonas alfalfae.
3. Starch not hydrolyzed.
a. Nitrites produced from nitrates.
41. Xanthomonas acernea.
aa. Nitrites not produced from nitrates.
42. Xanthomonas carotae.
43. Xanthomonas hederae.
44. Xanthomonas phormicola.
36. Xanthomonas vesicatoria.
aaa. Ammonia produced in nitrate media.
45. Xanthomonas geranii.
4. Starch hydrolysis not reported.
a. Nitrites produced from nitrates.
46. Xanthomonas antirrhini.
47. Xanthomonas heterocea.
aa. Nitrites not produced from nitrates.
48. Xanthomonas badrii.
49. Xanthomonas gummisudans.
50. Xanthomonas nigromacidans.
154 ORDER I. PSEUDOMONADALES
B. Gelatin not liquefied.
1. Starch hydrolyzed.
2. Starch not hydrolyzed.
C. Gelatin not reported.
1. Starch hydrolyzed.
51. Xanthomonas axonopodis.
52. Xanthomonas oryzae.
53. Xanthomonas celebensis.
II. Colonies whitish to cream; pigment non-water-soluble.
A. Gelatin liquefied.
1. Starch hydrolyzed.
a. Nitrites produced from nitrates.
54. Xanthomonas panici.
aa. Nitrites not reported.
2. Starch not reported.
55. Xanthomonas proteamaculans .
56. Xanthomonas manihotis .
57. Xanthomonas ruhrisuhalhicans
58. Xanthomonas cannae.
59. Xanthomonas conjac.
60. Xanthomonas zingiberi.
HOST PLANT KEY
The following key will be found useful for purposes of identification where the bacterium
is isolated from a known host plant.
I. Cause of leaf, stem and fruit spots and occasional blights of monocotyledonous plants.
A. Attack members of the family Araceae.
59. Xanthomonas conjac.
18. Xanthomonas dieffenbachiae.
B. Attacks members of the family Cannaceae.
58. Xarithomonas cannae.
C. Attack members of the family Gramineae.
51. Xanthomonas axonopodis.
20. Xanthomonas holcicola.
52. Xanthomonas oryzae.
53. Xanthomonas panici.
5. Xanthomonas rubrilineans .
57. Xanthomonas rubrisubalbicans .
33. Xanthomonas translucens.
35. Xanthomonas vasculorum,.
D. Attacks members of the family Iridaceae.
49. Xanthomonas gummisudans.
E. Attack members of the family Liliaceae.
1. Xanthomonas hyacinthi.
44. Xanthomonas phormicola.
F. Attacks members of the family Musaceae.
52. Xanthomonas celebensis.
G. Attacks members of the family Zingiberaceae.
60. Xanthomonas zingiberi.
II. Cause of leaf, stem and fruit spots and occasional blights of dicotyledonous plants.
A. Attacks members of the family Aceraceae.
41. Xanthomonas acernea.
FAMILY IV. PSEUDOMONADACEAE 155
B. Attacks members of the family Araliaceae.
43. Xanlhomonas hederae.
C. Attacks members of the family Bcgoniaceae.
7. Xanthovwnas begomae.
D. Attacks members of the family Befulaceae.
14. Xanthomonas corylina.
E. Attacks members of the family Chenopodiaceae.
4. Xanthomonas beticola.
F. Attack members of the family Compositae.
48. Xanthomonas hadrii.
50. Xanthomonas nigromaculans.
32. Xanthomonas taraxaci.
3. Xanthomonas vitians.
G. Attacks members of the family Convolvulaceae.
34. Xanthomonas uppalii.
H. Attack members of the family Cruciferae.
6. Xanthomonas barbareae.
9. Xanthomonas campestris.
21. Xanthomonas incanae.
36. Xanthomonas vesicatoria.
I. Attacks members of the family Cucurbitaceae.
15. Xanthomonas cucurbitae.
J. Attack members of the family Euphorbiaceae.
56. Xanthomonas manihotis.
29. Xanthomonas ricinicola.
K. Attack members of the family Geraniaceae.
45. Xanthomonas geranii.
26. Xanthomonas pelargonii.
L. Attacks members of the family Jnglandaceae.
22. Xanthomonas juglandis.
M. Attack members of the family Leguminosae.
40. Xanthomonas alfalfae.
11. Xanthomonas cajani.
10. Xanthomonas cassiae.
16. Xanthomonas desmodii.
17. Xanthomonas desmodiigangeticii .
23. Xanthomonas lespedezae.
27. Xanthomonas phaseoU.
30. Xanthomonas sesbaniae.
31. Xanthomonas stizolobiicola.
37. Xanthomonas vignicola.
N. Attacks members of the family Malvaceae.
25. Xanthomonas malvacearum.
O. Attacks members of the family Papaveraceae.
39. Xanthomonas papavericola.
P. Attacks members of the family Piper aceae.
8. Xanthomonas betUcola.
Q. Attacks members of the family Plantaginaceae .
28. Xanthomonas plantaginis.
R. Attacks members of the family Proteaceae.
55. Xanthomonas proteamaculans .
S. Attacks members of the family Rosaceae.
2. Xanthomonas pruni.
156
ORDER I. PSEUDOMONADALES
T. Attacks members of the family Rubiaceae.
24. Xanthomonas maculifoliigardeniae.
U. Attacks members of the family Rutaceae.
12. Xanthomonas citri.
V. Attacks members of the family Scrophulariaceae.
46. Xanthomonas antirrhini.
W. Attack members of the family Solanaceae.
19. Xanthomonas hemmiana.
47. Xanthomonas heterocea.
36. Xanthomonas vesicatoria.
X. Attacks members of the family Tiliaceae.
38. Xanthomonas nakatae.
Y. Attacks members of the family Umbelliferae.
42. Xanthomonas carotae.
Z. Attacks members of the family Verbenaceae.
13. Xanthomonas clerodendri .
1. Xanthomonas hyacinth! (Wakker,
1883) Dowson, 1939. {Bacterium hyacinthi
Wakker, Botan. Centralblatt, 14, 1883, 315;
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
188.)
hy.a.cin' thi. Gr. hyacinthus the hyacinth;
M.L. mas.n. Hyacinthus generic name; M.
L. gen. noun hyacinthi of hyacinth.
Description from Smith (Div. Veg. Phys.
and Path., U. S. D. A. Bui. 26, 1901, 40);
additional characters determined by Burk-
holder.
Rods 0.4 to 0.6 by 0.8 to 2.0 microns.
Motile with a polar flagellum. Filaments
present. Gram-negative.
Gelatin: Slow liquefaction.
Agar colonies: Circular, flat, moist,
shining, bright yellow. Media stained
brown.
Milk: Casein is precipitated and digested.
Tyrosine crystals produced.
Nitrites not produced from nitrates.
Indole: Slight production.
Hydrogen sulfide produced.
Acid but no gas from glucose, fructose,
lactose, sucrose, galactose, maltose, salicin
and ethyl alcohol. Slight acid from xylose.
Alkaline reaction in citrate. Mannitol,
dulcitol and malonate not utilized.
Starch: Hydrolysis slight.
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 28° and 30° C. Minimum, 4° C. Ma.xi-
mum, between 34° and 35° C.
Aerobic, with the exception of maltose,
where it is facultatively anaerobic.
Habitat: Produces a yellow rot of hj-a-
cinth bulbs, Hyacinthus ovientalis.
2. Xanthomonas pruni (Erw. Smith,
1903) Dowson, 1939. {Pseudomonas pruni
Erw. Smith, Science, N. S. 17, 1903, 456;
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
190.)
pru'ni. L. prunus plum; M.L. fem.n.
Prunus generic name; M.L. gen. noun
pruni of Prunus.
Description from Dunegan (U. S. Dept.
Agr., Tech. Bull. 273, 1932, 23).
Rods 0.2 to 0.4 by 0.8 to 1.0 micron.
Encapsulated. INIotile with a polar flagellum.
Gram-negative.
Gelatin: Liquefied.
Beef -extract agar colonies: Yellow, circu-
lar, smooth, convex, edges entire.
Broth: Turbid becoming viscid.
Milk: Precipitation of casein; digestion.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced. Hydrogen
sulfide produced (Burkholder).
Lipolytic (Starr and Burkholder, Phj^-
topath., 32, 1942, 600).
Acid from arabinose, .xylose, glucose,
fructose, galactose, manno.se, maltose, lac-
tose, sucrose, rafRnose and melezitose.
Starch is hydrolyzed (slight).
Pectate medium not liquefied.
Aerobic.
Temperature relations: Optimum, be-
tween 24° and 29° C. Maximum, 37° C.
FAMILY IV. PSEUDOMONADACEAE
157
Source: Smith isolated this ])uthogeii from
Japanese plimis.
Habitat: Pathogenic on plum {Prunus
salicina), peach (P. persica), apricot (P.
(irmeniaca), etc.
3. Xaiilhonionas vilians (Brown, 1918)
Dowson, 1943. {Bacterium vitians Brown,
Jour. Agr. Res., 13, 1918, 379; Dowson,
Trans. Brit. Mycol. Soc, 26, 1943, 13.)
vi'ti.ans. L. vitio to injure; L. part. adj.
vitians injuring.
Rods. Motile with bipolar flagella. Gram-
negative.
Gelatin: Slow liquefaction.
Beef-extract agar colonies: Circular,
smooth, thin, cream to cream-yellow.
Broth: Turbid with yellow ring.
Milk: Clears and turns alkaline.
Nitrites not produced from nitrates.
Indole: Feeble production.
Hydrogen sulfide: Feeble production.
Acid but no gas from glucose.
Starch: Feeble hydrolysis.
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 26° and 28° C. Minimum, 0° C. Maxi-
mum, 35° C.
Aerobic.
Source : Isolated from the stem of diseased
lettuce plants from South Carolina.
Habitat: Pathogenic on lettuce, Lactuca
sativa.
4. Xanthonionas beticola (Smith
et al., 1911) Savulescu, 1947. {Bacterium
beticolum Smith, Brown and Townsend, U.
S. Dept. Agr., Bur. Plant Ind., Bui. 213,
1911, 194; S&vulescu, Anal. Acad. Romane,
III, 22, 1947, 12).
be.ti'co.la. L. beta the beet; L. v. colo to
inhabit; M.L. noun beticola the beet dweller.
Description from Brown, Jour. Agr. Res.,
37, 1928, 167, where the species is referred to
as Bacterium beticola (Smith, Brown and
Townsend) Potebnia.
Rods 0.4 to 0.8 by 0.6 to 2.0 microns.
Motile with 1 to 4 polar flagella. Encapsu-
lated. Presumably Gram-negative although
originally reported as Gram-variable.
Gelatin: Liquefied.
Beef-agar slants: Growth moderate,
filiform, flat, glistening, yellow.
Broth: Turbid, yellow ring, abundant
sediment.
Milk: Coagulation and peptonization.
Indole not produced.
Hydrogen sulfide produced.
Nitrites produced from nitrates.
Acid from glucose, sucrose, maltose and
mannitol. No acid from lactose.
Starch hydrolysis feeble.
Temperature relations: Optimum, 29° C.
Minimum, 1.5° C. Maximum, 39° C.
Chemical tolerance: Optimum pH, 6.5.
Minimum, between 4.5 and 4.8. Maximum,
between 9.0 and 9.5.
Tolerates salt up to 9 per cent.
Aerobic.
Comment: It is doubtful whether this
species belongs in this genus.
Source: Isolated from galls on sugar beets
collected in Colorado, Kansas and Virginia.
Habitat: Produces galls on sugar beets
and on garden beets.
5. Xanthomonas rubrilineans (Lee et
al., 1925) Starr and Burkholder, 1942.
{Phytomonas rubrilineans Lee, Purdy, Bar-
num and Martin, Hawaiian Sugar Planters'
Assoc. Bui., 1925, 25; Starr and Burkholder,
Phytopath., 32, 1942, 600.)
ru.bri.li'ne.ans. L. ruber red; lineo to
make a straight line; rubrilineans making
red stripes.
Rods 0.7 by 1.67 microns. Motile with 1
or seldom more polar flagella. Gram-nega-
tive.
Gelatin: Liquefied.
Agar (Beef-extract + glucose) colonies:
Small, smooth, glistening, buff to yellow.
Broth: Turbid with pellicle. Sediment.
Milk: Casein precipitated and digested.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 600).
Acid from glucose, fructose, arabinose,
xylose, lactose, sucrose, raffinose and man-
nitol.
Starch: Slight hydrolysis.
Pectate medium not liquefied.
Growth range, pH 5.4 to pH 7.3.
Aerobic, facultative.
158
ORDER I. PSEUDOMONADALES
Source: Isolated from red stripe lesions in
sugar cane.
Habitat: Pathogenic on sugar cane.
6. Xanthoinonas barbareae Burkholder,
1941. (Phytopath., 31, 1941, 348.)
bar.ba're.ae. M.L. fem.n. Barbarea ge-
neric name of cress; M.L. gen. noun barbareae
of Barbarea.
Rods 0.4 to 0.95 by 1.0 to 3.15 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: Liquefied.
Beef-extract peptone colonies: Circular,
yellow, smooth, butyrous, growth moderate.
Potato glucose agar: Growth abundant,
pale 3'ellow. Mucoid.
Broth: Turbid, j^ellow granular ring.
Milk: Soft curd, with clearing and produc-
tion of tyrosine crystals. Litmus reduced.
Nitrates utilized but no nitrites produced.
Asparagine and nitrites not utilized.
Hydrogen sulfide produced.
Indole not produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid from glucose, galactose, xylose,
maltose, sucrose and glycerol. Alkali pro-
duced from salts of malonic, citric, malic
and succinic acids. Rhamnose, salicin and
hippuric acid salts not utilized.
Starch hydrolyzed.
Pectate medium liquefied.
Aerobic.
Distinctive characters: Similar to
Xanthomonas campestris but does not infect
cabbage, cauliflower or horseradish.
Source: From black rot of winter cress,
Barbarea vulgaris.
Habitat: Pathogenic on leaves and stems
of Barbarea vulgaris.
7. Xanthoinonas begoniae (Takimoto,
1934) Dowson, 1939. (Bacterium begoniae
Takimoto, Jour. Plant Protect., 21, 1934,
262; Dowson, Zent. f. Bakt., II Abt., 100,
1939, 190.)
be.go'ni.ae. Named for B6gon; M.L.
fem.n. Begonia generic name; M.L. gen.
noun begoniae of Begonia.
Translated by Dr. K. Togashi.
Rods 0.5 to 0.6 by 1.2 to 2.0 microns.
Motile with a polar flagellum. Gram-
negative.
Gelatin: No liquefaction. Liquefaction
(Wieringa, Tidschr. Plantziekt., 4I, 1935,
312; McCulloch, Jour. Agr. Res., 54, 1937,
859; Dowson, op. cit., 1939, 190; Stapp,
Arbeiten Biol. Reichsanst. f. Land- u.
Forstw., 22, 1938, 392).
Potato agar colonies: Circular, convex,
smooth, moist, shining, yellow.
Broth: Turbid. Yellow pellicle and pre-
cipitation.
Milk: No coagulation. Casein digested.
Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
No acid or gas in peptone broth from
glucose, sucrose, lactose or glycerol. Acid
from glucose, sucrose, lactose, mannitol and
glycerol in peptone-free medium (McCul-
loch, op. cit., 1937,859).
Starch hydrolyzed (Dowson, Jour. Roy.
Hort. Soc.,6S, 1938,289).
Pectate medium not liquefied.
Temperature relations: Optimum, 27°C.
Minimum, between 1° and 3° C. Maximum,
37° C.
Source: Isolated from leaf spot of tu-
berous begonia.
Habitat: Pathogenic on Begonia spp.
8. Xanthomonas betlicola Patel et al.,
1951. (Patel, Kulkarni and Dhande, Curr.
Sci.,^0, 1951, 106.)
bet.li'co.la. East Indian betle, the name
of the betel, a shrubby vine; L. v. colo to
dwell; M.L. fem.n. betlicola the betel-
dweller.
Rods slender, occurring singly or in
pairs. Motile. Encapsulated. Gram-nega-
tive.
Gelatin: Liquefied.
Potato-glucose agar colonies: 11 mm in
diameter in 7 days, baryta-yellow, lobate,
striations at periphery.
Broth: Turbid; yellow growth.
Milk: Litmus reduced. Casein digested.
Loeffler's blood serum liquefied.
Indole not produced.
Hydrogen sulfide produced.
FAMILY IV. PSEUDOMONADACEAE
159
Nitrites not produced from nitrates.
Methyl red negative: acetylmethylcarbi-
nol not produced.
Synthetic asparagine medium: No growth.
Acid but no gas from glucose, lactose and
sucrose. Salicin not attacked.
Starch hydrolyzed.
Salt tolerance: Up to 3 per cent.
Optimum temperature, between 25° and
28° C.
Aerobic.
Source: Isolated from leaves, stems and
petioles of Piper betle in India.
Habitat: Pathogenic on Piper betle.
Aerobic.
Distinctive characters : Causes a vascular
infection in cabbage, cauliflower and ruta-
bagas.
Comment : A variety pathogenic on horse-
radish and related species has been de-
scribed by McCulloch (Jour. Agr. Res., 38,
1929, 269). Causes a leaf spot. Does not
liquefy pectate medium.
Source: Pammel {op. cit., 1895, 130) first
isolated the pathogen from diseased ruta-
bagas.
Habitat: Pathogenic on cabbage, cauli-
flower and other related species.
9. Xanthoiuonas campestris (Pammel,
1895) Dowson, 1939. (Bacillus campestris
Pammel, Iowa Agr. Exp. Sta. Bull. 27,
1895, 130; Dowson, Zent. f. Bakt., II Abt.,
100, 1939, 190.)
cam.pes'tris. L. campestris of a level
field this specific epithet is also that of
Brassica campestris , a host.
Description from McCulloch (Jour. Agr.
Res., 38, 1929, 278). Species is probably
composed of several varieties. See descrip-
tions by Mekta, Ann. Appl. Biol., 12, 1925,
330; Paine and Nirula, Ann. Appl. Biol.,
15, 1928, 46; Wormald and Frampton, Ann.
Rept. East. Mall. Res. Sta., 1926 and 1927,
II Supplement, 1928, 108; and others.
Rods 0.3 to 0.5 by 0.7 to 2.0 microns.
Motile with a polar flagellum. Encapsu-
lated. Gram-negative.
Gelatin: Liquefied.
Beef agar colonies: Wax-j'ellow, round,
smooth, shining, translucent, margins
entire.
Broth: Turbid with yellow rim and some-
times a pellicle.
Milk: Casein digested with the formation
of tyrosine crystals. Alkaline.
Nitrites not produced from nitrates.
Indole production weak.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid but no gas from glucose, sucrose,
lactose, glycerol and mannitol.
Starch hydrolyzed.
Pectate medium liquefied.
Temperature relations: Optimum, be-
tween 28° and 30° C. Maximum, 36° C.
10. Xanthonionas cassiae Kulkarni et
al., 1951. (Kulkarni, Patel and Dhande,
Curr. Sci., 20, 1951, 47.)
cas'si.ae. M.L. fem.n. Cassia generic name
of host; M.L. gen. noun cassiae of Cassia.
Rods 0.8 to 1.0 by 1.2 to 2.1 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: Liquefied.
Potato-glucose agar colonies: 1.2 cm in
diameter after 7 days, smooth, circular,
lobate, glistening, convex, butyrous, pinard-
yellow.
Milk: Litmus reduced. Medium pepto-
nized.
Hydrogen sulfide produced.
Nitrites not produced from nitrates.
Loeffler's blood serum: Liquefied.
Methyl red negative; acetylmethylcar-
binol not produced.
Acid but no gas from glucose, lactose and
sucrose. Arabinose, glycerol and salicin not
attacked.
Starch hydrolyzed.
Koser's citrate medium: Growth.
Synthetic asparagine medium: Slight
growth.
Non-lipolytic.
Salt tolerance: Up to 3 per cent.
Optimum temperature, 27° C.
Aerobic.
Source: Isolated from leaves, stems and
petioles of Cassia tora in India.
Habitat: Pathogenic on Cassia tora.
11. Xanthonionas cajani Kulkarni et
al., 1950. (Kulkarni, Patel and Abhyankar,
Curr. Sci., 19, 1950, 384.)
160
PSEUDOMONADALES
ca'ja.ni. M.L. mas.n. Cajanus generic
name of host; M.L. gen. noun cajani of
Cajanus.
Rods 0.9 to 1.4 by 1.3 to 2.2 microns.
Encapsulated. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied.
Potato-glucose agar colonies: 1.5 cm in
diameter after 7 days, smooth, glistening,
entire, pulvinate, naphthalene-yellow.
Milk: Litmus reduced. Casein digested.
Loeffler's blood serum: Liquefied in 10
days.
Hydrogen sulfide produced.
Nitrites not produced from nitrates.
Acid but no gas from glucose, lactose and
sucrose. Salicin not attacked. Citrates
utilized.
Starch hydrolyzed.
Methyl red negative; acetylmethylcar-
binol not produced.
Synthetic asparagine medium: No growth.
Salt tolerance: Up to 3 per cent.
Optimum temperature, 30° C.
Aerobic.
Relationship to other species: Similar to
Xanthomonas phaseoli, which also infects
various legumes.
Source: Isolated from the pigeon pea,
Cajanus cajan, in India.
Habitat: Pathogenic on Cajanus cajan.
12. Xanthomonas citri (Hasse, 1915)
Dowson, 1939. {Pseudomonas citri Hasse,
Jour. Agr. Res., 4, 1915, 97; Dowson, Zent. f.
Bakt., II Abt., 100, 1939, 190.)
cit'ri. L. citrus the citrus; M.L. fem.n.
Citrus generic name; M.L. gen. noun citri of
Citrus.
Rods, 0.5 to 0.75 by 1.5 to 2.0 microns,
occurring in chains. Motile with a single
polar flagellum. Gram-negative.
Gelatin: Liquefied.
Beef agar colonies: Appear in 36 to 48
hours; circular, smooth, raised, dull yellow.
Broth: Turbid in 24 hours. A yellow ring
forms.
Milk: Casein is precipitated.
Nitrites not produced from nitrates.
Hydrogen sulfide produced (Reid, New
Zealand Jour. Sci. and Tech., 22, 1938, 60).
Indole not produced.
No gas from glucose, lactose or mannitol.
Starch hydrolyzed (Reid, loc. cit.).
Aerobic.
Temperature relations: Optimum, be-
tween 25° and 34° C. Minimum, 10° C.
Maximum, 38° C. (Okabe, Jour. Soc. Trop.
Agr., J^, 1932,476).
Source: Isolated from canker on orange.
Habitat: Produces a canker on man.y
species of Citrus and related plants.
13. Xanthomonas clerodendri Patel
et al., 1952. (Xanthomonas clerodendroni (sic)
Patel, Kulkarni and Dhande, Curr. Sci., 21,
1952, 74.)
cle.ro. den'dri. M.L. neut.n. Clerodendron
generic name of the plant host; M.L. gen.
noun clerodendri of Clerodendron.
Rods, 0.5 by 1.1 microns, occurring singly
or in chains. Encapsulated. Gram-negative.
Gelatin: Liquefied.
Potato-glucose agar colonies: Circular,
1.8 cm in diameter in 7 days, margins entire.
Pale lemon-yellow.
Litmus milk: Casein digested. Litmus re-
duced and milk peptonized.
Hydrogen sulfide produced.
Nitrites not produced from nitrates.
Acid but no gas from glucose, sucrose and
lactose. No growth in salicin.
Starch h3^drolyzed.
Optimum temperature, about 31° C.
Thermal death point, 51° C.
Source: From a leaf spot on Clerodendron
phlornoides .
Habitat: Pathogenic on Clerodendron
phlomoides.
14. Xanthomonas corylina (Miller et
al., 1940) Starr and Burkholder, 1942. (Mil-
ler, Bollen, Simmons, Gross and Barss,
Phytopath., 30, 1940, 731; Starr and Burk-
holder, Phytopath., 32, 1942, 598.)
co.ry.li'na. Gr. conjlus the hazel; IVI.L.
adj. corylinus pertaining to hazel.
Rods 0.5 to 0.7 by 1.1 to 3.8 microns.
Motile with a polar flagellum. Encapsu-
lated. Gram-negative.
Gelatin: Liquefied.
Nutrient glucose-agar streaks: Growth
abundant, filiform, convex, glistening,
smooth, opaque, pale lemon-j^ellow, viscid.
Broth: Turbid. Ring formed in 2 to 5
days.
Milk: Enzymatic curd that is slowly di-
FAMILY IV. PSEUDOMONADACEAE
161
gested. Litmus reduced. Crystal formation
(Burkholder).
Nitrites not produced from nitrates.
Nitrogen sources utilized are peptone,
aspartic acid, alanine, leucine, sodium
ammonium phosphate, allantoin, t.yrosine,
uric acid and brucine.
Indole not produced.
Hydrogen sulfide not produced on lead
acetate agar. HoS produced after ZoBell
and Feltham's method (Burkholder).
Selenium dioxide reduced.
Lipolytic (Starr and Burkholder, ibid.,
600).
Acid but no gas from glucose, fructose,
galactose, lactose, sucrose, maltose, xylose,
raffinose, mannitol, glycerol and starch.
Alkali from salts of citric, lactic, malic and
succinic acids. Arabinose, rhamnose, dulci-
tol, salicin, inulin and cellulose not utilized.
Starch hydrolj^zed.
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 28° and 32° C. Minimum, between 5°
and 7° C. Maximum, 37° C. Thermal death
point between 53° and 55° C.
pH range for growth: pH 5.2 to 10.5. Opti-
mum pH, between 6 and 8.
Strict aerobe.
Distinctive characters: Cultural charac-
ters the same or similar to those of Xantho-
monas juglandis. The two species do not
cross-infect.
Source: 26 isolates from widely scattered
filbert orchards in Oregon and Washington.
Habitat: Pathogenic on filberts {Corylus
avellana and C. maxima).
15. Xanthomonas cucurbitae (Bryan,
1926) Dowson, 1939. (Bacterium cucurbitae
Bryan, Science, 63, 1926, 165; Bryan, Jour.
Agr. Res., 40, 1930, 389; Dowson, Zent. f.
Bakt., II Abt., 100, 1939, 190.)
cu.cur'bi.tae. L. cucurbita a gourd; M.L.
fem.n. Cucurbita generic name; M.L. gen.
noun cucurbitae of Cucurbita.
Rods 0.45 to 0.6 by 0.5 to 1.3 microns.
Motile, usually with a single polar flagellum.
Gram-negative.
Gelatin: Liquefied.
Beef -agar slants: Growth moderate, mus-
tard-yellow, undulating margins, viscid to
butyrous.
Broth: Moderately turbid. Ring and j-el-
low sediment.
Milk: Precipitation of casein; digestion.
Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Acid from glucose, galactose, fructose,
lactose, maltose, sucrose and glycerol. No
acid from mannitol.
Starch hydrolyzed.
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 25° and 30° C. Maximum, 35° C.
pH range for growth: pH 5.8 to 9.0. Opti-
mum pH, between 6.5 and 7.0.
Slight growth in 5 per cent salt.
Aerobic.
Source: Species first isolated from squash.
Habitat : Causes a leaf spot of squash and
related plants.
16. Xanthomonas desniodii Uppal and
Patel, 1949. (Uppal and Patel, in Patel,
Curr. Sci., 18, 1949, 213; also see Patel,
Indian Phytopath., 2, 1949, 5.)
des.mo'di.i. M.L. neut.n. Desmodium
generic name of host; M.L. gen. noun
desmodii of Desmodium.
Rods, 0.4 to 0.8 by 1.6 to 2.4 microns, oc-
curring singly or in pairs. Motile with a
single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Nutrient agar slants: Growth fair, filiform,
flat, dull, smooth, opaque and pinard-
yellow.
Potato-glucose agar (neutral) colonies :
Yellowish amber with colorless margins,
circular, viscid, smooth and wet.
Beef broth: Growth slow. Moderate in 48
hours and good in 4 daj's.
Milk: Litmus turns red in 10 days. Reduc-
tion slow.
Indole not produced.
Hydrogen sulfide production fair.
Nitrites not produced from nitrates.
No growth in Cohn's, Uschinsky's or
Fermi's solution.
Acid but no gas from glucose, galactose,
lactose, mannitol, maltose and sucrose in
synthetic medium. Poor growth in salicin,
rafhnose, fructose, arabinose, xylose, dulci-
tol and glycerol, and no growth in tartaric,
162
ORDER I. PSEUDOMONADALES
citric, acetic or formic acids. No growth
when asparagine is used as carbon-nitrogen
source.
Starch hydrolyzed.
Temperature relations: Optimum, be-
tween 25° and 30° C. Slight growth at 11° C.
No growth at 38° C.
Chemical tolerance: Optimum pH, be-
tween 6.8 and 7.3. No growth at pH 8.5;
slight growth at pH 3.2.
Aerobic.
Source: From diseased Desmodium dif-
fusum in India.
Habitat: Pathogenic on Desmodium dif-
fusum, not on D. gangeticum.
17. Xanthonionas desmodiigangeticii
Uppal et al., 1948. {Xanthomonas desmodii-
gangeticii (sic) Uppal, Patel and Moniz, in
Patel and Moniz, Indian Phytopath., 1,
1948, 140; also see Patel and Moniz, Curr.
Sci., 17, 1948, 268.)
des.mo'di.i.gan.ge'ti.ci.i. M.L. neut.n.
Desmodium gangeticum name of host species;
M.L. gen. noun desmodiigangeticii of Des-
)iiodium gangeticum.
Rods 0.7 to 1.4 by 1.5 to 2.5 microns.
Motile with a single flagellum. Gram-nega-
tive.
Gelatin: Liquefied.
Nutrient agar slants: Growth fair, dull,
flat, opalescent, lemon-chrome.
Nutrient broth: Moderately turbid. No
pellicle.
Milk: Litmus reduced. No tyrosine.
Nitrites not produced from nitrates.
Hydrogen sulfide produced.
Indole not produced.
Non-lipolytic.
Uschinsky's solution: Growth.
Acetylmethjdcarbinol not produced.
Arabinose, xylose, glucose, galactose,
fructose, maltose, sucrose, raffinose, manni-
tol, salicin and sodium citrate are utilized.
Asparagine utilized as carbon-nitrogen
source.
Starch hydrolyzed.
Salt tolerance: Growth retarded by 3 per
cent salt; inhibited by 4 per cent salt.
Temperature relations: Optimum, be-
tween 20° and 25° C. Minimum, 5°C. Maxi-
mum, 35° C.
Aerobic.
Source: From a disease of Desmodium
gangeticum found in India.
Habitat: Pathogenic on Desmodium gange-
ticum but not on D. diffusum.
18. Xanthomonas diefFenbachiae (Mc-
Culloch and Pirone, 1939) Dowson, 1943.
{Phytomonas dieffenbachiae McCulloch and
Pirone, Phytopath., 29, 1939, 962; Dowson,
Trans. Brit. Mycol. Soc, 26, 1943, 12.)
dief .fen.bach'i.ae. Dieffenbach patro-
nymic; M.L. fem.n. Dieffenbachia generic
name; M.L. gen. noun dieffenbachiae oi Dief-
fenbachia.
Rods 0.3 to 0.4 by 1.0 to 1.5 microns.
Encapsulated. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied.
Beef -infusion peptone agar colonies:
Slow growing, circular, flat, smooth, trans-
lucent, butyrous, massicot- to Naples-
yellow.
Broth: Turbid. Yellow rim or slight
pellicle.
Milk: Slow peptonization and formation
of tyrosine crystals. Litmus reduced.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Acid from glucose, sucrose, lactose,
galactose, fructose and glycerol. Growth
but no acid in maltose and mannitol.
Starch moderately hydrolyzed.
Temperature relations: Optimum, be-
tween 30° and 31° C. Minimum, 5° C. Maxi-
mum, between 37° and 38° C.
Aerobic.
Source: Seven isolates from diseased
leaves of Dieffenbachia picta.
Habitat: Pathogenic on Dieffenbachia
picta. Artificial infection of Dracaena fra-
grans.
19. Xanthomonas hemmiana (Yama-
moto, 1951) Burkholder, comb. nov. (Phyto-
monas hemmianus (sic) Yamamoto, Forsch.
auf dem Gebiet d. Pflanzenkr., 4, 1951, 163.)
hem.mi.a'na. Named for T. Hemmi, a
Japanese plant pathologist; M.L. adj.
hemmianus of Hemmi.
Rods, 0.3 to 0.7 by 1.3 to 2.2 microns,
occurring singly or in pairs. Motile with 1 to
3 polar flagella. Gram-negative.
FAMILY IV. PSEUDOMONADACEAE
163
Gelatin: Liquefied.
Beef extract agar colonies: Small, circu-
lar, smooth, flat or raised with regular
margins, white to pale yellow.
Beef broth: Moderate clouding.
Milk: Clearing after coagulation. Litmus
red.
Uschinsky's solution: Good growth.
Cohn's solution: Poor growth.
Potato: Growthsmooth, copious andolive-
bufT.
Nitrites produced from nitrates.
Indole produced.
Hydrogen sulfide produced.
Acid and gas from glucose, sucrose and
glycerol. Acid from lactose.
Starch hydrolyzed.
Temperature relations: Optimum, 32° C.
Growth above 36° C. and below 2° to 8° C.
Optimum pH, between 6 and 7; no growth
below pH 3.
Aerobic.
Relationship to other species: This spe-
cies closely resembles the specie.s placed in
Aeromonas Kluyver and van Niel.
Source : Isolated from leaf spot of Jimson
weed. Datura spp.
Habitat: Pathogenic on Datura metel, D.
meteloides, D. inermis, tomato and petunia.
20. Xaiithomona.s holcicola (Elliott,
1930) Starr and Burkholder, 1942. {Bac-
terium holcicola Elliott, Jour. Agr. Res., 40,
1930, 972; Starr and Burkholder, Phytopath.
32, 1942, 600.)
hol.ci'co.la. Gr. holcus kind of grass;
M.L. mas.n. Holcus generic name of velvet
grass and sorghum; L. v. cola to dwell ; M.L.
fem.n. holcicola, Holcus dweller.
Rods 0.75 by L58 microns. Motile with
1 or 2 polar flagella. Encapsulated. Gram-
negative.
Gelatin: Liquefied.
Beef -infusion peptone agar colonies:
Round, umbonate, glistening, smooth,
translucent to opaque, wax-yellow, buty-
rous.
Broth: Trace of growth in 24 hours. Later
turbid with a slight ring.
Milk: Casein precipitated and peptonized.
Alkaline.
Nitrite production doubtful.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, loc.
cit.).
Acid but no gas from sucrose.
Starch hydrolyzed.
Temperature relations: Optimum, be-
tween 28° and 30° C. Minimum, 4° C. Maxi-
mum, between 36° and 37° C.
pH range for growth: pH 5.5 to 9.0. Opti-
mum pH, between 7.0 and 7.5.
Source: Isolated from many collections of
sorghum leaves showing a streak disease.
Habitat: Pathogenic on leaves of Holcus
sorghum and H. halepensis.
21. Xanthomonas incanae (Kendrick
and Baker, 1942) Starr and Weiss, 1943.
(Phytomonas incanae Kendrick and Baker,
California Bull. 665, 1942, 10; Starr and
Weiss, Phytopath., 33, 1943, 316.)
in.ca'nae. L. adj. incanus hoary, gray;
from host Matthiola incana.
Rods 0.4 to 0.8 by 0.6 to 2.5 microns.
Motile with a polar flagellum. Gram-nega-
tive.
Gelatin: Liquefied.
Beef extract agar colonies: Round,
smooth, convex or pulvinate, glistening,
margin entire, picric-yellow to amber color.
Broth: Turbid.
Milk: No coagulation. A clearing of
the medium.
Nitrites not produced from nitrates.
Indole not produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid but no gas from glucose, lactose,
sucrose, mannitol, d-galactose, xylose,
d-mannose, raffinose, trehalose and glycerol.
No acid from maltose, 1-arabinose or
rhamnose.
Starch not hydrolyzed. Starch hydrolyzed
(Burkholder).
Pectate medium liquefied.
Tolerates 3 per cent salt.
Growth in beef broth at pH 4.4.
Aerobic.
Distinctive characters: Causes a disease
of flowering stock but not of cabbage. Dif-
fers from Xanthomonas campestris in that it
does not utilize 1-arabinose or maltose.
Source: Four isolates from diseased plants
of Matthiola incana.
164
ORDER I. PSEUDOMONADALES
Habitat: Pathogenic on flowering stocks.
22. Xanthoinouas juglandis (Pierce,
1901) Dowson, 1939. (Pseudomonas juglandis
Pierce, Bot. Gaz., 31, 1901, 272; Dowson,
Zent. f. Bakt., II Abt., 100, 1939, 190.)
jug.lan'dis. L. juglans, juglandis the wal-
nut; M.L. fem.n. Juglans generic name of
walnut; M.L. gen.noun juglandis of the wal-
nut.
Description from Miller, Bollen, Sim-
mons, Gross and Barss (Phytopath., 30,
1940, 731).
Rods 0.5 to 0.7 by 1.1 to 3.8 microns.
Motile with a polar flagellum. Encapsu-
lated. Gram-negative.
Gelatin: Liquefied.
Nutrient glucose-agar streaks: Growth
abundant, filiform, convex, glistening,
smooth, opaque, pale lemon-yellow, viscid.
Broth : Turbid. Ring formed in 2 to 5 days.
Milk: Enzjmiatic curd that is slowly di-
gested. Litmus reduced. Crystal formation
(Burkholder) .
Nitrites not produced from nitrates.
Nitrogen sources utilized are peptone,
aspartic acid, alanine, leucine, sodium
ammonium phosphate, allantoin, tyrosine,
uric acid and brucine.
Indole not produced.
Hydrogen sulfide not produced on lead
acetate agar. H2S produced after ZoBell
and Feltham's method (Burkholder).
Selenium dioxide reduced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid but no gas from glucose, fructose,
galactose, lactose, sucrose, maltose, xylose,
raffinose, mannitol, glycerol and starch.
Alkali from salts of citric, lactic, malic and
succinic acids. Arabinose, rhamnose, dulci-
tol, salicin, inulin and cellulose not utilized.
Starch hydrolyzed.
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 28° and 32° C. Minimum, between 5°
and 7° C. Maximum, 37° C. Thermal death
point, between 53° and 55° C.
pH range for growth, 5.2 to 10.5. Optimum
pH, between 6 and 8.
Source: Isolated from black spots on the
leaves and nuts of English walnuts, Juglans
regia.
Habitat: Pathogenic on the walnut, Jug-
lans spp.
23. Xanthomonas lespedezae (Ayers
et al., 1939) Starr, 1946. (Phytomonas lespe-
dezae Ayers, Lefebvre and Johnson, U. S.
Dept. Agr. Tech. Bull. 704, 1939, 19; Starr,
Jour. Bact., 51, 1946, 136.)
les.pe.de'zae. Named after Lespedez;
M.L. fem.n. Lespedeza generic name; M.L.
gen.noun lespedezae of Lespedeza.
Rods, 0.56 by 1.62 microns, occurring
singly, in pairs, or occasionally in short
chains. Encapsulated. Motile with a single
polar flagellum. Gram-negative.
Gelatin: Liquefied.
Nutrient agar colonies: Circular, raised,
glistening, translucent, viscid, yellow.
Broth: Turbid in 48 hours.
Milk: Peptonized; becomes alkaline.
Blood serum and egg albumin: Liquefied.
Nitrites not produced from nitrates.
Indole produced after 11 days.
Hydrogen sulfide produced.
No gas from carbohydrates.
Starch hydrolyzed.
Pectate medium liquefied.
Aerobic.
Temperature relations: Optimum, near
35°C. No growth at 5°C. or at 40°C.
Source: Isolated from diseased Lespedeza
spp. collected in Virginia, New York and
Illinois.
Habitat: Pathogenic on Lespedeza spp.
24. Xanthomonas maculifoliigarden-
iae (Ark, 1946) Elrod and Braun, 1947.
(Phytomonas maculifolium-gardeniae (sic)
Ark, Phytopath., 36, 1946, 867; Xantho-
monas maculajoliumgardeniae (sic) Elrod
and Braun, Jour. Bact., 53, 1947, 515.)
ma.cu.li.fo'li.i.gar.de'ni.ae. L. fem.n.
macula a spot; L. neut.n. folium a leaf; M.L.
neut.n. maculijolium a leaf spot; M.L.
fem.n. Gardenia the generic name of the
host; gardeniae of gardenia; M.L. gen.noun
maculifoliigardeniae of leaf spot of gar-
denia.
Rods 0.3 to 0.5 by 1.6 to 2.0 microns.
Encapsulated. Motile with 1 to 2 polar
flagella. Gram-negative.
Gelatin: Slow liquefaction.
Beef-peptone agar colonies: Growth
FAMILY IV. PSEUDOMONADACEAE
165
rapid. Slightly raised, yellow, butyrous in
young cultures, difficult to pick up in old
cultures.
Broth: Turbid in 24 hours.
Milk: White curd in bottom. Litmus a
dirty wine color in supernatant liquid.
Uschinsky's medium: Good growth.
Fermi's solution: Scant growth.
Indole not produced.
Hydrogen sulfide not produced.
Nitrites not produced from nitrates.
Ammonia produced from peptone.
Acid but no gas from arabinose, glucose,
fructose, galactose, lactose, maltose, man-
nitol, raffinose, sucrose and xylose. Glycerol
not attacked. Tartrate utilized.
Starch hydrolj^zed.
Temperature relations: Optimum, be-
tween 22° and 28° C. Minimum, 10° C. Maxi-
mum, 37° C. Thermal death point, 50° C.
Source: Six isolates from gardenia leaf
spots.
Habitat : Causes a spot on young leaves of
gardenias.
25. Xanthomonas nialvacearum (Erw.
f Smith, 1901) Dowson, 1939. (Pseudomonas
nialvacearum Erw. Smith, U. S. Dept. Agr.,
Div. Veg. Phys. and Path., Bull. 28, 1901,
153; Dowson, Zent. f. Bakt., II Abt., 100,
1939, 190.)
mal.va.ce.a'rum. L. malva the mallow;
M.L. fem.pl.n. Malvaceae the mallow family;
M.L. fem.pl. gen. n. malvacearum of the mal-
lows.
Description from Elliott (Man. Bact.
Plant Pathogens, 1930, 153) and Lewis
(Phytopath., 20, 1930, 723).
Rods. Motile with a single polar fiagellum.
Gram-negative.
Gelatin: Liquefied.
Agar slants: Growth moderate, convex,
smooth, glistening, pale yellow, wavy to
irregular margins.
Broth: Slight to moderate turbidity.
Sediment.
Milk: Casein precipitated and slowly di-
gested.
Nitrites not produced from nitrates.
Hydrogen sulfide produced (Burkholder) .
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 600).
Acid but no gas from glucose, galactose.
fructose, xylose, lactose, maltose, sucrose,
raffinose, glycerol, inulin and glycogen.
Alkaline reaction from salts of acetic, citric,
lactic and succinic acids. No fermentation
of arabinose, mannitol, dulcitol, salicin, and
salts of formic, oxalic and tartaric acids
(Lewis).
Starch hydrolyzed (Lewis).
Pectate medium not liquefied.
Temperature relations: Optimum, be-
tween 25° and 30° C. Maximum, between
36" and 38° C. (Elliott).
Source: Isolated from angular leaf spot
of cotton.
Habitat: Pathogenic on cotton, where-
ever it is grown, causing a leaf spot, a stem
lesion and a boll lesion.
26. Xanthomonas pelargonii (Brown,
1923) Starr and Burkholder, 1942. (Bac-
terium pelargoni (sic) Brown, Jour. Agr.
Res., 23, 1923, 372; Starr and Burkholder,
Phytopath., 32, 1942, 600.)
pe.lar.go'ni.i. Gr. pelargus the stork:
M.L. neut.n. pelargonium generic name of
stork's bill; M.L. gen. noun pelargonii of
Pelargonium.
Rods 0.67 by 1.02 microns. Encapsulated.
Motile with a single polar fiagellum. Gram-
negative.
Gelatin: Slow liquefaction.
Beef -agar colonies: Cream-colored, glis-
tening, round, with delicate internal mark-
ings.
Broth: Turbid in 24 hours. Incomplete
pellicle.
Milk: Alkaline. Clearing in bands.
Nitrites not produced from nitrates.
Indole production slight.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Slight acid but no gas from glucose, su-
crose and glycerol.
Starch hydrolysis feebly positive.
Pectate medium liquefied.
Temperature relations: Optimum, 27° C.
Maximum, 35° C.
No growth in broth plus 3.5 per cent salt.
Aerobic.
Source: Isolated from spots on leaves of
Pelargonium from District of Columbia,
Maryland and New Jersey.
166
ORDER I, PSEUDOMONADALES
Habitat: Pathogenic on Pelargonium spp.
and Geranium spp.
27. Xanthomonas phaseoli (Erw.
Smith, 1897) Dowson, 1939. {Bacillus
phaseoli Erw. Smith, Bot. Gaz., U, 1897,
192; A. A. A. S. Proc, 46 1898, 288; Dow-
son, Zent. f. Bakt., II Abt., 100, 1939, 190.)
pha.se'o.li. Gr. phaselus the kidnej^ bean;
L. phaseolus kidney bean; M.L. mas.n.
Phaseolus generic name of bean; M.L. gen.
noun phaseoli of the bean.
Description from Burkholder (Cornell
Agr. Exp. Sta. Mem. 127, 1930, 18; Phyto-
path.,^^, 1932,609).
Rods 0.87 by 1.9 microns. Motile with a
single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Beef -extract agar colonies: Circular,
amber-yellow, smooth, butyrous, edges
entire.
Broth: Turbid in 24 hours. Yellow ring.
Milk: Casein precipitated and digested.
Alkaline. Tyrosine crystals formed.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 3^, 1942,600).
Acid but no gas from glucose, galactose,
fructose, arabinose, xjdose, maltose, lactose,
sucrose, raffinose and glycerol. Alkaline
reaction from salts of acetic, malic, citric
and succinic acids. Mannitol, dulcitol, sali-
cin and formic and tartaric acids not fer-
mented.
Starch hydrolj^zed.
Pectate medium not liquefied.
Aerobic.
Very slight growth in beef broth plus 4
per cent salt (Hedges, Jour. Agr. Res., 29,
1924, 243).
Distinctive character: Similar in culture
to Xanthomonas campestris, X.juglandis, X.
vesicatoria, etc., but they do not cross
infect.
Comments: A variety that produces pus-
tules on the leaves and pod of soy bean,
Glycine max, both in America and Japan,
has been described by Hedges (Science, 66,
1922, 11). Liquefies pectate medium.
Two additional varieties have been de-
scribed which produce a dark brown color
in a beef extract peptone medium and also in
tj^rosine medium. The first of these is patho-
genic on beans (Phaseolus vulgaris) and re-
lated plants. The second was isolated from
white kidney beans in India and is patho-
genic on Phaseolus vulgaris, P. lunatus, P.
coccineus and Dolichos lablab.
Habitat: Pathogenic on the bean {Phase-
olus vulgaris), the hyacinth bean {Dolichos
lablab) , the lupine {Lupinus polyphyllus) , etc.
Not pathogenic on the soy bean {Glycine
sp.) nor cowpea {Vigna sp.).
28. Xanthomonas plantaginis (Thorn-
berry and Anderson, 1937) Burkholder, 1948.
{Phytomonas plantaginis Thornberry and
Anderson, Phytopath., 27, 1937, 947; Burk-
holder, in Manual, 6th ed., 1948, 161.)
plan.ta'gi.nis. L. plantago, plantaginis
the plantain; M.L. fem.n. Plantago generic
name of plantain; M.L. gen. noun plantaginis
of plantain.
Rods, 0.6 to 1.0 by 1.0 to 1.8 microns, oc-
curring singly or in chains. Encapsulated.
Motile with 1 to 2 polar flagella. Gram-nega-
tive.
Gelatin: Slight liquefaction.
Glucose agar slant: Growth moderate,
filiform, raised, opaque, yellow and viscid.
Broth: Moderately turbid with ring.
Milk: Slight acidity, no reduction of lit-
mus. Peptonization.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced.
No appreciable amount of gas from carbo-
hydrates.
Starch hydrolyzed.
Temperature relations: Optimum, 25° C.
Minimum, 12° C. Maximum, 35° C. Thermal
death point, 50° C.
Aerobic.
Source: From diseased leaves of Plantago
lanceolata in Illinois.
Habitat: Pathogenic on Plantago spp.
29. Xanthomonas ricinicola (Elliott,
1930) Dowson, 1939. {Bacterium ricini Yoshi
and Takimoto, Jour. Plant Protect., Tokyo,
16, 1928, 12; Bacterium ricinicola Elliott,
Man. Bact. Plant Path., 1930, 193; Dowson,
Zent. f. Bakt., II Abt., 100, 1939, 190.)
ri.ci.ni'co.la. L. ricinus the castor oil
FAMILY IV. PSEUDOMONADACEAE
167
plant; M.L. mas.n. Kicinus generic name of
the castor bean; L. v. colo to dwell; M.L.
fem.n. ricinicola, Ricinus dweller.
Rods, 0.4 to 0.9 by 1.3 to 2.6 microns,
occurring in short chains. Encapsulated.
Motile with polar flagella. Gram-negative.
Gelatin: Liquefied.
Nutrient agar colonies: Lemon-yellow,
changing to brown.
Milk: Slightly acid. No coagulation. Pep-
tonization.
Nitrites not produced from nitrates.
Acid but no gas from lactose.
Starch hydrolyzed.
Temperature relations: Optimum, be-
tween 29° and 30° C. Minimum, 2.5° C. Maxi-
mum, 39° C.
Aerobic.
Comment: Elliott (loc. cit.) renamed this
species to avoid confusion with Phytomonas
ricini Archibald.
Source: Isolated from leaf -spot of castor
bean.
Habitat: Pathogenic on Ricinus com-
30. Xanthonionas sesbaniae Patel
et al., 1952. (Patel, Kulkarni and Dhande,
Curr. Sci.,^^, 1952,74.)
ses.ba'ni.ae. M.L. fem.n. Sesbania generic
name of the plant host; M.L. gen. noun
sesbaniae of Sesbania.
Rods, 0.7 by 1.3 microns, occurring singly
or in chains. Encapsulated. Gram-negative.
Gelatin: Liquefied.
Potato-glucose agar colonies: Circular, 2
cm in diameter in 7 daj^s, with striations
starting 5 mm awaj" from the center up to
the periphery. Barium-yellow.
Litmus milk: Slightly peptonized with
casein digested. Litmus slowly reduced.
Hydrogen sulfide produced.
Nitrites not produced from nitrates.
Acid but no gas from glucose, sucrose and
lactose. Salicin not attacked.
Starch hydrolyzed.
Temperature relations: Optimum, 31° C.
Thermal death point, 51° C.
Source: Isolated from leaf spots on jSes-
bania aegyptiaca.
Habitat: Pathogenic on Sesbania aegyp-
tiaca.
31. Xanthonionas stizolobiicola Patel
et al., 1951. (Patel, Kulkarni and Dhande,
Curr. Sci., 20, 1951, 106.)
sti.zo.Io.bi.i'co.la. M. L. neut.n. Stizolo-
biutn generic name of host; L. v. colo to
inhabit; M.L. fem.n. stizolobiicola the
Stizolobium dweller.
Rods. Mostly single. Encapsulated. Mo-
tile. Gram-negative.
Gelatin: Liquefied.
Nutrient agar colonies : 8 mm in diameter
in 4 days, flat, entire, glistening, creamy to
pinard-yellow.
Broth: Good growth.
Synthetic asparagine medium: No growth.
Loeffller's blood serum: Liquefied in 10
days.
Hydrogen sulfide produced.
Nitrites not produced from nitrates.
Methyl red negative; acetylmethylcar-
binol not produced.
Indole not produced.
Acid but no gas from glucose, lactose and
sucrose. Salicin not attacked.
Starch and casein hydrolyzed.
Lipolytic.
Salt tolerance: Up to 3 per cent.
Optimum temperature, between 28° and
30° C.
Aerobic.
Relationship to other species: Elliott
(Man. Bact. Plant Path., 2nd ed., 1951, 129)
lists Xanthomonas phaseoli on Stizolobium
deeringianum. The two pathogens appear
similar.
Source: Isolated from leaves, stems and
petioles of Stizolobium deeringianum in
India.
Habitat: Pathogenic on Stizolobium
deeringianum.
32. Xanthonionas taraxaci Nieder-
hauser, 1943. (Phytopath., 33, 1943, 961.)
ta.ra.x'a.ci. M.L. neut.n. Taraxacum ge-
neric name of host; M.L. gen. noun taraxaci
of Taraxacum.
Rods, 0.7 to 1.2 by 1.4 to 3.3 microns,
occurring singly or in pairs. Motile with a
single polar flagellum. Gram-negative.
Gelatin: Rapid liquefaction.
Beef-extract peptone agar colonies: Cir-
cular, smooth, bright yellow. Growth mod-
erate.
168
ORDER I. PSEUDOMONADALES
Broth: Turbid with thin ring.
Milk: Litmus reduced. Soft curd precipi-
tated and slowly digested. Liquid gradually
clears. Tyrosine crystals produced.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Lipase produced.
Acid from glucose, xylose, galactose,
fructose, lactose, sucrose and glycerol.
Arabinose, maltose, raffinose, inulin, manni-
tol, ethanol and salicin not attacked. Salts
of acetic, citric, lactic, malic and succinic
acids utilized with an increase in pH. Salts
of tartaric, formic, salicylic and benzoic
acids not utilized.
Starch hydrolyzed.
Pectate medium liquefied.
Salt tolerance: 3.25 to 3.5 per cent.
Temperature relations: Optimum, 30° C.
Minimum, between 0° and 3° C. Maximum,
38° C.
Aerobic.
Source: Seven isolates from diseased Rus-
sian dandelions grown at Ithaca, New York.
Habitat: Pathogenic on Taraxacum kok-
Russian dandelion.
33. Xanthoinonas translucens (Jones
et al., 1917) Dowson, 1939. (Bacterium trans-
lucens Jones, Johnson and Reddy, Jour.
Agr. Res., 11, 1917, 637; Dowson, Zent. f.
Bakt., II Abt., 100, 1939, 190.)
trans. lu'cens. L. transluceo to be trans-
lucent; L. part, translucens being translu-
cent.
Rods 0.5 to 0.8 by 1.0 to 2.5 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: Liquefied.
Beef-peptone agar colonies: Round,
smooth, shining, amorphous except for
inconspicuous, somewhat irregular con-
centric striations within, wax-yellow tinged
with old gold; margin entire.
Broth: Turbidity becomes rather strong.
Pellicle.
Milk: Soft coagulum and digestion. Milk
clears. Tyrosine crystals produced.
Nitrites not produced from nitrates.
Indole: Slight production.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Ammonia from peptone.
Acid but no gas from glucose, d-fructose,
d-mannose, d-galactose, sucrose, lactose
and sometimes salicin. No utilization of
1-rhamnose, inositol, maltose, raffinose,
inulin, d-mannitol or dulcitol.
Starch hydrolyzed.
Pectate medium not liquefied.
Temperature relations: Optimum, 26° C.
Minimum, 6° C. Maximum, 36° C.
Aerobic.
Distinctive characters: Many forms of
Xanthomonas translucens have been de-
scribed, all of which have the same cultural
characters; they differ mainly in patho-
genicity.
Comment: Various varieties, formae spe-
ciales and races of this species have been
described. See Elliott (Man. Bact. Plant
Path., 2nd ed., 1951, 142-146) for details.
Source : Originally isolated from bacterial
blight of barley.
Habitat: Causes water-soaked stripes,
streaks or other lesions on leaves, culms
or glumes of grain and related plants.
Occurs naturally on Triticum spp., Hor-
deum spp., Bromus spp., Secale cereale,
Phleum pratense and, by inoculation, on
Avena spp.
34. Xanthomonas uppalii Patel, 1948-
(Indian Phytopath., 1, 1948, 67.)
up.pa'li.i. Named for B. N. Uppal, an
Indian plant pathologist; M.L. gen. noun
uppalii of Uppal.
Rods, 0.7 to 1.0 by 2.0 to 2.4 microns,
mostly single. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Rapid liquefaction.
Nutrient agar slants: Growth smooth,
slightly raised, dull, filiform, opalescent,
lemon-chrome.
Potato-glucose agar colonies: Growth
copious, glistening, butyrous, empire-
yellow.
Broth: Turbid. No pellicle. Sediment and
floccules in 4 days.
Milk: Growth good. Litmus reduced.
Indole not produced.
Hydrogen sulfide produced.
FAMILY IV. PSEUDOMONADACEAE
169
Nitrites and ammonia not produced.
Acetylmethylcarbinol not produced.
No growth in Uschinsky's, Cohn's or
Koser's uric acid medium.
Acid but no gas from glucose, lactose,
sucrose, mannitol, raffinose, salicin and
-xylose. Fructose, arabinose and rhamnose
not attacked.
Starch hydrolyzed.
Temperature relations: Optimum, 30°C.
Minimum, 10°C. Maximum, 40°C.
pH range for growth, pH 5.3 to 9.2. Opti-
mum pH, 7.0.
Source: Isolated from Iponioea niuricata
in India.
Habitat: Pathogenic on Ipomoea nmricata.
35. Xanthoiiionas vasculoruni (Cobb,
1893) Dowson, 1939. {Bacillus vascidarum
(sic) Cobb, Agr. Gaz. of New South Wales,
Jf, 1893, 777; abst. in Cent. f. Bakt., II Abt.,
1, 1895, 41; Xanthomonas vascularum (sic)
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
190.)
vas.cu.lo'rum. L. vascnlum a small
vessel; M.L. neut.pl. gen. n. vasculorum of
small vessels.
Description from Erw. Smith (Bact. in
Rel. to Plant Dis., 3, 1914, 54).
Rods 0.4 by 1.0 micron. Motile with a
single polar flagellum. Originally reported
as Gram-variable but later found to be
Gram-negative (Elliott, Man. Bact. Plant
Path., 2nded., 1951, 147).
Gelatin: Liquefaction feeble. Liquefac-
tion good (Burkholder).
Beef-e.xtract agar colonies: Pale yellow,
smooth, glistening, not noticeablj^ viscid.
Broth: Good growth.
Milk: Alkaline.
Nitrites not produced from nitrates.
Lipolytic (Starr and Burkholder, Phy-
topath., 3^, 1942, 600).
Acid but no gas from glucose, fructose
and glycerol. No acid from lactose.
Starch hydrolyzed (Burkholder).
Pectate medium liquefied.
Temperature relations: Optimum, 28° C.
Thermal death point, about 50° C. (Elliott,
op. cit., 1951, 147).
Source: Isolated from diseased sugar
cane.
Habitat: Pathogenic on sugar cane,
Saccharum officinarum, causing a bacterial
gummosis.
36. Xanthomonas vesica torla (Doidge,
1920) Dowson, 1939. {Bacterium vesicatorium
Doidge, Jour. Dept. Agr., S. Africa, 1, 1920,
718; also Ann. Appl. Biol., 7, 1921, 428;
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
190.)
ve.si.ca.to'ri.a. L. vesica a blister;
M.L. adj. vesicatorius causing a blister.
Rods 0.6 to 0.7 by 1.0 to 1.5 microns. En-
capsulated. Motile with a single polar flagel-
lum. Originally reported as Gram-positive
but later found to be Gram-negative by
Gardner and Kendrick (Phytopath., 13,
1923, 307) and Higgins (Phytopath., 12,
1922, 513).
Gelatin: Liquefied.
Nutrient agar colonies: Good growth.
Circular, wet-shining, Naples-yellow, edges
entire.
Milk: Casein precipitated and slowly
digested. Tyrosine crystals.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced (Burkholder).
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid but no gas from glucose, fructose,
sucrose, lactose, galactose, glycerol and
dextrin.
Certain strains hydrolyze starch, others
do not (Burkholder and Li, Phytopath., 31,
1941, 753).
Pectate medium liquefied.
Optimum temperature, 30° C.
Distinctive character: Xanthomonas vesi-
catoria is reported as pathogenic on toma-
toes and peppers. However Burkholder and
Li {loc. cit.) report that there are sufficient
cultural and pathogenic differences between
the organism infecting tomatoes and the
organism infecting peppers to warrant their
separation into distinct species.
Comment: A variety pathogenic on
radishes, turnips and other crucifers, and
on tomato and peppers, has been described
by White (Phytopath., 20, 1930, 653). Differs
from Xanthomonas campestris in that it
does not cause a vascular disease. Unlike a
170
ORDER I. PSEUDOMONADALES
variety of the latter species, it is not patho-
genic on horseradish. Originally isolated
from leaf spots of radishes and turnips in
Indiana.
Source: Isolated from spotted tomato
fruits in South Africa.
Habitat: Pathogenic on tomatoes, Lyco-
persicon esculenhim, and peppers, Capsicum
annuum.
37. Xanthomonas vignicola Burk-
holder, 1944. (Phytopath., 34, 1944, 431.)
vig.ni'co .la. M.L. fem.n. V igna gen&vic
name of host; L. v. colo to dwell; M.L.
fem.n. vignicola the Vigna dweller.
Rods 0.7 (0.46 to 0.92) by 1.76 (1.0 to
2.8) microns. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied.
Beef -extract peptone agar slant: Fili-
form, glistening, edges entire, primuline-
yellow, butyrous.
Broth: Turbid in 48 hours; heavy ring;
no pellicle.
Litmus milk: Light curd becoming solid.
Slow peptonization with crystal formation.
Litmus reduced. Brownish syrup at end of
6 weeks.
Hydrogen sulfide produced.
Indole not produced.
Nitrites not produced from nitrates.
Asparagine and tyrosine not utilized as
carbon-nitrogen sources. Tyrosine broken
down to a brownish pigment in other media.
Lipolytic.
Salt tolerance: 2 per cent retards and 3
per cent inhibits growth.
Acid but no gas from glucose, galactose,
lactose, maltose, sucrose and raffinose.
Alkaline reactions with salts of citric and
malic acids. Fructose, 1-arabinose, xylose,
rhamnose, glycerol, salicin and the sodium
salts of lactic, formic, succinic, tartaric and
hippuric acids not attacked.
Starch hydrolyzed.
Pectate medium liquefied.
Aerobic.
Temperature relations: Optimum, be-
tween 27° and 30° C. Minimum, between
6° and 9° C. Maximum, 37° C.
Source : Six isolates from cankers of cow-
pea stems.
Habitat: Causes canker disease of cow-
peas, Vigna spp., and disease of the red
kidney bean, Phaseolus vulgaris.
38. Xanthomonas nakatae (Okabe,
1933) Dowson, 1943. {Bacterium nakatae
Type B, Okabe, Jour. Soc. Trop. Agr.,
Formosa, 5, 1933, 161; Dowson, Trans.
Brit. Mycol. Soc, 26, 1943, 12.)
na'ka.tae. Named for K. Nakata, a
Japanese plant pathologist; M.L. gen. noun
nakatae of Nakata.
Rods 0.3 to 0.4 by 1.1 to 2.5 microns.
Encapsulated. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied. Brown color.
Beef -extract agar colonies: Amber-yellow,
circular, smooth, glistening, margins entire.
Medium turns brown.
Broth: Moderate turbidity with yellow
ring. Medium turns brown.
Milk: Casein precipitated and digested.
Tyrosine crystals produced. Brown color.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced (slight) .
Acid but no gas from glucose, sucrose,
maltose and lactose.
Starch: Active hydrolysis.
Temperature relations: Optimum, be-
tween 30° and 32° C. Minimum, 10° C.
Maximum, 39° C.
No growth in beef extract broth plus 2
per cent salt.
Aerobic.
Distinctive character: Differs from Type
A in that it produces a brown pigment in
culture. (Description of Type A not seen.)
Source: Isolated from water-soaked to
brown leaf spots on jute.
Habitat: Pathogenic on jute, Corchorus
capsularis.
39. Xanthomonas papavericola (Bryan
and McWhorter, 1930) Dowson, 1939.
(Bacterium papavericola Bryan and Mc-
Whorter, Jour. Agr. Res., 40, 1930, 9; Dow-
son, Zent. f. Bakt., II Abt., 100, 1939, 190.)
pa.pa.ve.ri'co.la. L. papaver the poppy;
M.L. neut.n. Papaver generic name of
poppy; L. V. colo to dwell; M.L. fem.n.
papavericola poppy dweller.
Rods, 0.6 to 0.7 by 1.0 to 1.7 microns,
occurring in chains. Encapsulated. Motile
FAMILY IV. PSEUDOMONADACEAE
171
with a single polar flagellum. Gram-nega-
tive.
Gelatin: Liquefied.
Beef agar colonies: Mustard-yellow to
primuline-yellow, circular, margins entire.
Broth: Turbidity prompt with a yellow
ring and an incomplete pellicle.
Milk: Soft coagulation; peptonization
and production of tyrosine crystals.
Nitrates: A weak reaction for nitrites
after 10 days.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid but no gas from glucose, galactose,
fructose, sucrose, lactose, maltose, glycerol
and mannitol.
Starch hydrolyzed.
Pectate medium liquefied.
Temperature relations: Optimum, be-
tween 25° and 30° C. Maximum, 35° C.
No growth in broth plus 5 per cent salt.
Aerobic.
Source: Isolated from black spots on
leaves, buds and pods of poppy.
Habitat: Pathogenic on poppy, Papaver
rhoeas .
40. Xaiithomonas alfalfae (Riker
et al., 1935) Dowson, 1943. (Bacterium al-
falfae Riker, Jones and Davis, Jour. Agr.
Res., 51, 1935, 177; Dowson, Trans. Brit.
Mycol. Soc, £6, 1943, 11.)
al.fal'fae. Spanish alfalfa (lucerne);
M.L. gen. noun alfalfae of alfalfa.
Rods 0.45 by 2.4 microns. Motile with
a single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Nutrient agar slant: Growth abundant,
filiform, smooth, glistening, butyrous, pale
yellow.
Broth: Turbid in 24 hours. Light sedi-
ment.
Milk: Casein precipitated and digested.
Ammonia produced slowly in a nitrate
medium.
Acid but no gas from glucose, maltose,
lactose, arabinose and salicin (Patel, Kul-
karni and Dhande, Indian Phytopath., 2,
1949, 166). No acid in yeast broth plus
sugars.
Starch hydrolyzed.
Aerobic.
Temperature relations: Optimum, be-
tween 24° and 32° C. Minimum, below 4° C.
Maximum, below 36° C.
Source: Six single-cell cultures isolated
from diseased alfalfa.
Habitat: Pathogenic on the leaves of
alfalfa, Medicago saliva.
41. Xanthomonas acernea (Ogawa, 1937)
Burkholder, 1948. (Pseudomonas acernea
Ogawa, Ann. Phyt. Soc. Japan, 7, 1937,
123; Burkholder, in Manual, 6th ed., 1948,
165.)
a.cer'ne.a. L. acerneus made of maple.
Rods 0.2 to 0.6 by 0.5 to 1.2 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: Liquefied.
Agar colonies: Round, smooth, convex,
white to citron-yellow, glistening, trans-
lucent with amorphous structure.
Broth: Turbid.
Milk: Slowly cleared, slightly acid. No
coagulation.
Nitrites produced from nitrates.
Hydrogen sulfide produced.
No gas produced in peptone water plus
sugars.
Starch not hydrolyzed.
Temperature relations: Optimum, about
32° C. Thermal death point, 59° C.
Aerobic.
Source: From diseased leaves of Acer
trifidum in Japan.
Habitat: Causes a disease in Acer spp.
and in Aesculus turhinata and Koelrenteria
paniculata.
42. Xanthomonas carotae (Kendrick,
1934) Dowson, 1939. (Phytomonas carotae
Kendrick, Jour. Agr. Res., 49, 1934, 504;
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
190.)
ca.ro 'tae. L. carota the carrot; M.L.
gen. noun carotae of the carrot.
Rods 0.42 to 0.85 by 1.38 to 2.75 microns.
Motile with 1 or 2 polar flagella. Gram-nega-
tive.
Gelatin: Liquefied.
Potato-glucose agar colonies: Circular,
smooth, glistening, entire, straw-j^ellow in
color.
172
ORDER I. PSEUDOMONADALES
Milk: Casein precipitated and milk
cleared; alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Acid but no gas from glucose, d-galactose,
xylose, d-mannose, 1-arabinose, sucrose,
lactose, raffinose, trehalose, d-mannitol
and glycerol. No acid from maltose or
rhamnose.
Starch not hydrolyzed.
Pectate medium liquefied.
Optimum temperature, between 25° and
30° C.
Tolerates 4 per cent salt at pH 7.
Aerobic.
Source: Two original isolations from
diseased carrots and a reisolation from
inoculated carrots were used for the descrip-
tion.
Habitat : Pathogenic on leaves of Daucvs
carota var. saliva.
43. Xanthonionas hederae (Arnaud,
1920) Dowson, 1939. {Bacterium hederae
Arnaud, Compt. rend. Acad. Sci., Paris,
171, 1920, 121; Dowson, Zent. f. Bakt., II
Abt., 100, 1939, 190.)
he'de.rae. L. Aedera the ivy; M.L. fem.n.
Hedera generic name of ivy; M.L. gen. noun.
hederae of ivy.
Description taken from Burkholder and
Guterman (Phytopath., £2, 1932, 783).
Rods 0.6 by 2.13 microns. Motile with a
single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Beef-extract-agar slants: Growth good,
filiform, amber-yellow, butyrous.
Broth: Turbid.
Milk: Casein precipitated and digested.
Milk becomes alkaline.
Nitrites not produced from nitrates.
Hydrogen sulfide produced.
Indole not produced.
Not lipolytic (Starr and Burkholder,
Phytopath., 32, 1942, 600).
Acid from glucose, fructose, galactose,
.xylose, sucrose, lactose and glycerol. Alkali
from salts of acetic, citric, lactic, malic
and succinic acids. The following are not
utilized: arabinose, rhamnose, maltose,
salicin, cellulose and formic acid.
Starch not hydrol3^zed.
Pectate medium not liquefied.
Aerobic, facultative.
Source : Isolated from diseased ivy leaves.
Habitat: Pathogenic on ivy, Hedera
helix.
44. Xanthonionas phorniicola (Taki-
moto, 1933) Dowson, 1943. {Bacterium
phorniicola Takimoto, Jour. Plant Protect.,
20, 1933, 777; Dowson, Trans. Brit. Mycol.
Soc, 26, 1943, 12.)
phor.mi'co.la. Gr. dim. phormi^im the
name of a plant; M.L. neut.n. Phormium
generic name of New Zealand flax; L. v.
colo to dwell; M.L. fem.n. phormicola the
Phormium dweller.
Description translated by Dr. K. Togashi.
Rods 0.5 to 0.6 by 1.0 to 2.0 microns.
Motile with a single flagellum. Gram-nega-
tive.
Gelatin: Liquefied.
Agar colonies: Light j'ellow, then waxy
yellow; butyrous, then viscid.
Broth: Turbid; pellicle formed.
Milk: Casein coagulated slowly and pre-
cipitated, then digested. Alkaline.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
No gas from sucrose, glucose, lactose or
glycerol.
No acid from various sugars in broth.
Temperature relations: Optimum, about
29° C. Minimum, about 0° C. Maximum,
39° C.
Aerobic.
Source: Species isolated from New Zea-
land flax, Phormixim tenax.
Habitat: Causes a leaf stripe of Phor-
mium tenax.
45. Xanthonionas geranii (Burkholder,
1937) Dowson, 1939. {Phytomonas geranii
Burkholder, Phytopath., 27, 1937, 560;
Dowson, Zent. f. Bakt., II Abt., 100, 1939,
190.)
ge . ra'ni . i. Gr. geranium geranium, crane's
bill; M.L. neut.n. Geranium generic name;
M.L. gen. noun geranii of Geranium.
Rods 0.75 to 2.0 microns. Motile with a
single polar flagellum. Gram-negative.
Gelatin: Liquefied.
Beef -extract agar slants: Moderate to
FAMILY IV. PSEUDOMONADACEAE
173
good filiform growth, glistening, primuline-
yellow. Develops in 24 hours.
Broth: Turbid in 24 hours. No pellicle
but a moderate sediment.
Milk: Becomes clear with a heavy casein
precipitate. Peptonization with crystal
formation.
Nitrates reduced to ammonia.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, Phy-
topath., 32, 1942, 600).
Acid from glucose, galactose, fructose,
xylose, rhamnose, lactose, sucrose, raffinose
and glycerol. Alkaline reaction from salts
of citric, malic, malonic and succinic acids.
No growth in arabinose or formic, hippuric,
maleic or tartaric acids.
Starch not hydrolyzed.
Pectate medium liquefied.
Aerobic.
Distinctive characters: Pathogenic on
Geranium spp., not on the house geranium,
Pelargonium hortorum.
Relationship to other species: Similar in
culture to Xanthomonas pelargonii.
Source: Three cultures isolated from
Geranium sanguineum.
Habitat: Pathogenic on Geranium san-
guineum, G. maculatum, G. praiense and G.
sylvaticum.
46. Xanthomonas antirrhini (Taki-
moto, 1920) Dowson, 1943. (Pseudomonas
antirrhini Takimoto, Bot. Mag. Tokyo, 34,
1920, 257; Dowson, Trans. Brit. Mycol. Soc,
26, 1943, 11.)
an.tir.rhi'ni. Gr. antirrhinum the plant
snapdragon; M.L. gen. noun antirrhini of
the snapdragon.
Description from Elliott (Man. Bact.
Plant Path., 1930, 93).
Rods 0.3 to 0.4 by 0.8 to 1.2 microns. En-
capsulated. Motile with polar flagella.
Gram-negative.
Gelatin: Liquefied.
Agar colonies: Circular, glistening, white,
later yellow.
Milk: Coagulated and casein digested.
Nitrites produced from nitrates.
No gas produced.
Aerobic.
Temperature relations: Optimum, be-
tween 26° and 27° C. Maximum, 34° C.
Habitat: Causes a leaf spot of Aniirr
hinum niajus.
47. Xanthomonas heterocea (Vzoroff,
1930) Sa.vulescu, 1947. {Phytomonas heterocea
Vzoroff, Bull. North Caucasian Plant
Prot. Sta. Roztoff-on-Don, 6-7, 1930, 263;
Sa,vulescu, Anal. Acad. Romane, III, 22,
1947, 11.)
he.te.ro'ce.a. Gr. adj. heterus another,
different.
Description taken from Rev. App. Myc,
10, 1931, 628.
Rods 0.4 to 0.6 by 1.0 to 2.0 microns.
Motile. Gram-negative.
Gelatin: Slow liquefaction.
Agar colonies : Circular, 2 mm in diameter,
convex, smooth, semi-transparent, glisten-
ing, yellow to amber. Pitted surface.
Milk: No coagulation. At first acid, later
alkaline.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Acid from glucose, galactose, arabinose,
xylose, sucrose, maltose, salicin, glycerol
and mannitol. Does not ferment lactose,
inulin, ethyl alcohol, esculin, adonitol or
dulcitol.
Optimum temperature, between 25° and
30° C.
Source: Isolated from diseased tobacco in
the North Caucasus.
Habitat: Pathogenic on Nicoliana ta-
bacum.
48. Xanthomonas badrii Patel et al.,
1950. (Patel, Kulkarni and Dhande, Indian
Phytopath.,3, 1950, 104.)
bad'ri.i. From the given name of Badri
Uppal, Indian plant pathologist; M.L.
gen. noun badrii of Badri.
Rods, 0.7 to 1.0 by 1.4 to 1.8 microns,
occurring singly and rarely in chains. Motile
with a single polar flagellum. Gram-nega-
tive.
Gelatin: Liquefied.
Nutrient agar colonies: Smooth, glisten-
ing, entire, empire-yellow; growth slow.
Milk: Cleared in 8 days. Litmus reduced.
Loeffler's blood serum: Liquefied.
174
ORDER I. PSEUDOMONADALES
Nitrites not produced from nitrates.
Indole not produced.
Ammonia produced.
Methjd red negative; acetylmethylcarbi-
nol not produced.
Acid but no gas from glucose, lactose,
sucrose, mannitol and salicin.
Optimum temperature, 31° C.
Source: Isolated from leaf spot of Xan-
thium strumarium in India.
Habitat: Pathogenic on Xanthium stru-
marium.
49. Xanthoinonas guniniisudans (Mc-
Culloch, 1924) Starr and Burkholder, 1942.
{Bacterium giunmisudans McCulloch, Phy-
topath., 14-, 1924, 63; also see Jour. Agr.
Res., 27, 1924, 229; Starr and Burkholder,
Phytopath., 32, 1942, 600.)
gum. mi.su 'dans. L. gummi gum; L. v.
sudo to sweat, exude; M.L. part. adj. gum-
misudans exuding gum.
Rods 0.6 to 0.8 by 1.0 to 2.8 microns.
Encapsulated. Motile with a single polar
flagellum. Gram-negative.
Gelatin: Liquefied.
Beef-peptone agar colonies: Amber-
yellow, circular, transparent, smooth, with
definite margins.
Broth: Moderately turbid with a yellow
ring.
Milk: Soft curd which is digested with
formation of tyrosine crystals.
Nitrites not produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
Lipolytic (Starr and Burkholder, loc. cit.).
Acid from glucose and sucrose.
Temperature relations: Optimum, 30° C.
Minimum, 2° C. Maximum, 36° C.
Aerobic.
Source: From gummy lesions on gladiolus
leaves.
Habitat: Pathogenic on leaves of gladioli.
50. Xanthonionas nigromaculans (Ta-
kimoto, 1927) Dowson, 1943. {Bacterium
nigromaculans Takimoto, Jour. Plant Pro-
tect., Tokyo, ^, 1927, 522; Dowson, Trans.
Brit. Mycol. Soc, 26, 1943, 12.)
ni.gro.ma'cu.lans. L. niger black; L. v.
maculo to spot; M.L. part. adj. nigromacu-
lans spotting with black.
Description translated by Dr. K. Togashi.
Rods 0.6 to 0.9 by 1.5 to 2.8 microns.
Motile with 1 or 2 polar fiagella. Gram-
negative.
Gelatin: Liquefied.
Agar colonies: Yellow, circular, entire,
smooth, glistening.
Broth: Growth moderate with yellow
pellicle.
Milk: Coagulation and digestion of the
casein.
Nitrites not produced from nitrates.
Indole not produced.
No acid or gas from glucose, sucrose,
lactose, mannitol or glycerol in peptone
water.
Temperature relations: Optimum, be-
tween 27° and 28° C. Minimum, 0° C. Maxi-
mum, 33° C.
Aerobic.
Comment : A forma specialis that is path-
ogenic on Zinnia spp. has been described
(Hopkins and Dowson, Trans. Brit. Mycol.
Soc, 32, 1949, 253).
Source: Isolated from lesions on leaf and
petioles of burdock.
Habitat: Pathogenic on leaves and peti-
oles of Arctium lappa, the burdock.
51. Xanthoinonas axonopodis Starr
and Garces, 1950. {Xanthomonas axonoperis
(sic) Starr and Garces, Rev. Fac. Nal. de
Agron. de Medellin, 12, 1950, 75.)
ax.on.o'pod.is. Gr. noun axon axis; Gr.
noun pous foot; M.L. mas.n. Axonopus
generic name of a grass; M.L. gen. noun
axonopodis of Axonopus.
Rods 0.4 by 1.0 to 3.0 microns. Encapsu-
lated. Motile by means of a single polar
flagellum. Gram-negative.
Gelatin: Not liquefied.
Yeast extract agar: Growth slow; small,
yellow colonies appear in 7 days.
Peptone sucrose agar: Growth slow; yel-
low colonies 1 mm in diameter appear in 7
days.
Potato-glucose agar: No growth.
Broth: Slight turbidity in two days;
slimy pellicle in 2 weeks. Yellowish ring
produced.
Milk: Litmus not reduced.
Nitrites not produced from nitrates.
Indole not produced.
FAMILY IV. PSEUDOMONADACEAE
175
Hydrogen sulfide not produced.
Non-Iipolytic.
Tyrosine in a caseinate medium: Growtli
slight; no color reaction.
Carbohj'drate utilization difficult to
determine because of meager growth. Glu-
cose, sucrose and trehalose probably uti-
lized. Lactose, maltose, raffinose, dulcitol,
gl,ycerol, mannitol, sorbitol, dextrin, inulin,
aesculin and salicin utilization doubtful.
Starch hydrolyzed.
Pectate medium not liquefied.
Temperature relations: Optimum, 30° C.
Minimum, 5° C. Maximum, 37° C.
Moderate growth in broth plus 1 per cent
NaCl; no growth with 1.5 per cent NaCl.
Chemical tolerance: Optimum pH be-
tween 6.6 and 7.6. Minimum, 5.8.
Source: Isolated from diseased grass,
Axonopus scoparius, in Colombia.
Habitat: Pathogenic on Axonopvs spp.
52. Xanthonionas oryzae (Uyeda and
Ishiyama, 1926) Dowson, 1943. (Pseudo-
monas oryzae Uyeda and Ishiyama, Proc.
Third Pan-Pacific Sci. Congr., Tokyo, 2,
1926, 2112; Dowson, Trans. Brit. Mycol.
Soc, 26, 1943, 12.)
o.ry'zae. Gr. oryza rice; M.L. fem.n.
Oryza generic name of rice; M.L. gen. noun
oryzae of Oryza.
Rods 0.5 to 0.8 by 1.0 to 2.0 microns.
Motile with a single polar flagellum. Gram-
negative.
Gelatin: No liquefaction.
Nutrient agar colonies: Circular, smooth,
glistening, wax-j-ellow.
Milk: Slightly acid.
Nitrites not produced from nitrates.
Hydrogen sulfide produced.
Acid but no gas from glucose, lactose and
sucrose.
Optimum temperature, between 26° and
30° C.
Strict aerobe.
Source: Isolated from a leaf blight of rice.
Habitat: Pathogenic on rice, Oryza saliva.
53. Xanthonionas celebensis (Giiu-
mann, 1923) Dowson, 1943. {Fseiidomonas
celebensis Gaumann, Ztschr. f. Pflanzen-
krank., SS, 1923, 11; Meded. Inst, voor
Plantenziek., Buitenzorg, 59, 1923, 17;
Dowson, Trans. Brit. Mycol. Soc, 26, 1943,
11.)
ce.le.ben'sis. Celebes, an island name;
M.L. adj. celebensis of Celebes.
Rods 0.9 by 1.5 microns. Motile by a
single polar flagellum. Gram-negative.
Agar colonies: Graj'ish yellow.
Broth: Thin pellicle.
Milk: Coagulated and cleared.
Nitrites not produced from nitrates.
Sodium selenite: Brick red.
Starch hydrolyzed.
Source: From vascular bundles of dis-
eased bananas from the Celebes.
Habitat: Causes the blood disease of
banana.
54. Xanthonionas panici (Elliott, 1923)
Savulescu, 1947. (Bacterium panici Elliott,
Jour. Agr. Res., 26, 1923, 157; Sivulescu,
Anal. Acad. Romane, III, 22, 1947, 11.)
pa'ni.ci. L. panicum Italian panic grass;
M.L. neut.n. Panicum generic name; M.L.
gen. noun panici of Panicum.
Rods 0.69 by 1.66 microns. Encapsu-
lated. Motile with 1, rarely 2, polar flagella.
Gram-negative.
Gelatin: Liquefaction slow.
Beef agar colonies: Circular, white,
smooth, glistening, margins at first entire,
later undulate.
Broth: Moderate turbidity in 24 hours.
Thin pellicle. Medium brownish.
Milk: Alkaline and clears.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
No acid or gas from carbohydrates.
Starch: Hydrolysis moderate.
Temperature relations: Optimum, 33° C.
Minimum, 5° C. Maximum, 45° C.
pH range for growth, pH 5.4 to 10.0.
Optimum pH, between 6.15 and 6.3.
Aerobic.
Distinctive characters: Differs from
Pseudomonas andropogoni in that it liquefies
gelatin, produces nitrites from nitrates and
does not infect sorghum and broom corn.
Source: Isolation from water-soaked le-
sions on leaves, sheaths and culms of millet
collected in Wisconsin and in S. Dakota.
Habitat: Pathogenic on i)roso millet,
Panicum miliaceum.
176
ORDER I. PSEUDOMONADALES
55. Xanthomonas proteamaculans
(Paine and Stansfield, 1919) Burkholder,
1948. (Pseudomonas proteamaculans Paine
and Stansfield, Ann. Appl. Biol., 6, 1919,
38; Burkholder, in Manual, 6th ed., 1948,
169.)
pro.te.a.ma'cu.lans. Gr. Proteus a god;
M.L. noun Protea a plant generic name; L.
V. maculo to spot; M.L. part. adj. 'proteama-
culans spotting Protea.
Rods 0.6 to 0.8 by 0.8 to 1.6 microns.
Motile with 1 to 3 polar flagella. Gram-
positive (Paine and Stansfield). Gram-nega-
tive (Dowson, personal communication,
August, 1953).
Gelatin: Liquefied.
Agar slant: Growth wet-shining, dirty
white with a faint yellow tinge.
Broth: Turbid in 24 hours. Slight ring.
Milk: Acid with soft curd after 2 days.
Later a separation of whey.
Nitrites produced from nitrates.
Acid and gas from glucose, sucrose and
mannitol. No acid or gas from lactose.
Starch: Slight hydrolysis.
Source: Repeated isolation from a leaf-
spot of Protea in England.
Habitat: Pathogenic on Protea cynar-
oides .
56. Xanthomonas manihotis (Ar-
thaud-Berthet, 1912) Starr, 1946. {Bacillus
manihotus (sic) Arthaud-Berthet, in Bon-
dar, Chacaras and Quintaes, 5 (4), 1912,
15; Starr, Jour. Bact., 51, 1946, 136.)
ma.ni.ho'tis. M.L. Manihot a plant
generic name; M.L. gen. noun manihotis of
Manihot.
Description from Burkholder (Phyto-
path.,S^, 1942, 147).
Rods 0.35 to 0.93 by 1.4 to 2.8 microns.
Mostly non-motile. One isolate showed a
few cells with a single polar flagellum.
Amaral (Instit. Biol., Sao Paulo, Arq.,
IS, 1942, 120) states that the species is mo-
tile with a single polar flagellum. Gram-
negative.
Gelatin: Liquefied.
Beef -extract-peptone agar slant: Growth
raised, ivory-colored, smooth, shiny, with
edges entire.
Potato-glucose agar: Growth abundant,
white to hyaline, very mucoid.
Broth: Turbid with a whitish granular
ring.
Litmus milk: Litmus reduced and milk
clears. With return of color, litmus is purple.
Indole not produced.
Hydrogen sulfide produced.
Nitrites produced from nitrates (Drum-
mond and Hipolito, Ceres, 2, 1941, 298).
Asparagine not used as a nitrogen and
carbon source. No growth in nitrate syn-
thetic broth.
Weak growth but slight acid production
in synthetic medium plus glucose, d-galac-
tose, d-fructose, d-xylose, maltose or su-
crose. No growth in rhamnose, 1-arabinose,
d-lactose, glycerol, mannitol or salicin.
Good growth with alkaline reaction in same
medium plus salts of the following acids:
acetic, citric, malic, maleic or succinic.
The salts of formic, hippuric, lactic and
tartaric acids were not utilized.
Starch hydrolyzed.
Pectate medium liquefied.
Lipolytic action slight.
Aerobic.
Temperature relations: Optimum, 30° C.
Minimum, 5° C. Maximum, 38° C.
Source: First isolated from the cassava.
Manihotus utilissima, in Brazil.
Habitat: Produces a wilt disease on
various species of Manihotus.
57. Xanthomonas rubrisubalbioans
(Christopher and Edgerton, 1930) Savu-
lescu, 1947. {Phytomonas rubrisubalbicans
Christopher and Edgerton, Jour. Agr. Res.,
41, 1930, 266; Sivulescu, Anal. Acad. Ro-
mane. III, 22, 1947, 11.)
ru.bri. sub. al'bi. cans. L. ruber red; L.
subalbicans whitish; M.L. adj. rubrisubalbi-
cans red whitish.
Short rods with polar flagella. En-
capsulated. Gram-negative.
Gelatin: No liquefaction.
Bacto-glucose agar colonies: Circular,
glistening, viscid, milky gray to buff.
Margins translucent, entire.
Broth: Turbid after 24 hours. Pellicle
and a ropy sediment.
Indole produced.
Hydrogen sulfide produced.
No acid or gas from carbohydrates.
Starch hydrolyzed.
FAMILY IV. PSEUDOMONADACEAE
177
Optimum temperature, 30° C.
Optimum pH, 6.8 to 8.0.
Source: Isolated many times from mot-
tled stripe of sugar cane in Louisiana.
Habitat: Pathogenic on sugar cane.
Johnson's grass and sorghum
58. Xanthomonas cannae (Bryan, 1921)
Savulescu, 1947. (Bacterium cannae Bryan,
Jour. Agr. Res., 21, 1921, 152; Sivulescu,
Anal. Acad. Romane, III, 22, 1947, 12.)
can'nae. Gr. carina a reed; M.L. fem.n.
Canna generic name; M.L. gen. noun cannae
of Canna.
Rods 0.5 to 0.7 by 1.0 to 2.0 microns.
Encapsulated. Motile with 1 to 3 polar
flagella. Gram-negative.
Gelatin: Slow liquefaction.
Agar slants: Growth filiform, white,
moist, with thin margins and granular
centers.
Broth: Turbid; heavy sediment.
Milk: Alkaline and clears.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
No acid produced from carbohydrates.
Temperature relations: Optimum, 35° C.
Minimum, 5° C. Maximum, 40° C.
Aerobic.
Source: Isolated from diseased canna
leaves collected in Washington, D. C. and
in Illinois.
Habitat: Causes a disease in Canna
indica.
59. Xanthomonas conjac (Uyeda,
1910) Burkholder, 1948. {Pseudomonas
conjac Uyeda, Bot. Mag. Tokyo, 24, 1910,
182; Burkholder, in Manual, 6th ed., 1948,
171.)
con'jac. M.L. conjac the specific epithet
of Amorphophallus konjac, the host.
Description from Elliott (Man. Bact.
Plant Path., 19,30, 121).
Rods 0.75 to 1.0 by 1.5 microns. Motile
with 1 to 4 polar flagella. Presumably Gram-
negative although the original description
records this species as Gram-positive (Burk-
holder).
Gelatin colonies: Circular to irregular,
light yellow.
Broth : Pellicle formed.
Milk: Coagulated.
Conjac: Liquefied.
Nitrites produced from nitrates.
Indole produced.
Hydrogen sulfide produced.
Gas from glucose.
Optimum temperature, 24° C.
Habitat: Pathogenic on Amorphophallus
konjac.
60. Xanthomonas zingiber! (Uyeda,
1908) SSvulescu, 1947. (Eine neue species,
Uyeda, Cent. f. Bakt., II Abt., 17, 1907,
383; Pseudomonas zingiheri Uyeda, Rept.
Imp. Agr. Exp. Sta., Japan, No. 35, 1908,
114; Savulescu, Anal. Acad. Romane, III,
22, 1947, 13.)
zin.gi'be.ri. Gr. indecl. zingiheri ginger.
Description from Stapp (in Sorauer,
Handb. d. Pflanzenkrank., 2, 5 Aufl., 1928,
65).
Rods 0.5 to 1.1 by 0.75 to 1.8 microns.
Non-motile at first, later a polar flagellum
develops. Gram-negative.
Gelatin: Liquefied.
Agar colonies: White.
Milk: Coagulation and peptonization of
the casein.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide produced.
No gas from glucose.
Temperature relations: Optimum, 28° C.
Minimum, 5° C. Maximum, 40° C.
Source: Isolated from ginger plant show-
ing a rot at the base of the sprouts.
Habitat: Pathogenic on ginger. Zingiber
officinale.
Addendum: Species inceriae sedis. Two
additional groups of yellow, polar-flagellate
species are described in this addendum al-
though they are not typical of the genus
Xanthomonas in all respects. The first is a
group of three species of plant pathogens.
One of these species is non-motile, but it ap-
pears to be closely related to the two polar-
flagellate species with which it is associated.
Plant pathologists have placed these three
species in Xanthomonas even though they do
not possess all of the characteristics of the
species in this genus senszi stricto. The
non-water-soluble, yellow pigment differs
17i
ORDER I. PSEUDOMONADALES
from that found in true xanthomonads.
Likewise none of the three species liquefies
gelatin. Neither do they show the gummy
growth of true xanthomonads, and they
differ in other important characteristics.
The second group comprises eleven species
which are not pathogenic to plants so far
as is known. They have been isolated from
the surface of leaves, soil and similar ma-
terials. All produce a non-water-soluble,
yellow pigment, but no one has as yet under-
taken a comparative study of cultures to
determine which of these species, if any,
are true xanthomonads.
Key to Xanthomonas Addendum.
I. Plant pathogens.
A. Non-motile.
1. Xanthomonas stewartii.
B. Polar flagellate.
1. Litmus milk alkaline. Pathogenic on 7ns spp.
2. Xanthomonas tardicrescens.
2. No change in litmus milk. Pathogenic on sugar cane, Saccharum officinarum.
3. Xanthomonas albilineans.
II. Saprophytic species.
A. Gelatin liquefied.
1. Nitrites produced from nitrates.
a. Acid but no gas from glucose.
4. Pseudomonas trifolii.
5. Pseudomonas xanthe.
aa. Action on glucose not recorded.
6. Pseudomonas caudata.
2. Nitrites not produced from nitrates.
a. Litmus milk acid; ferments lactose.
7. Pseudomonas perlurida.
aa. Litmus milk slimy, alkaline.
8. Pseudomonas ochracea.
B. Gelatin not liquefied.
1. Nitrites produced from nitrates.
a. Do not attack cellulose.
b. Does not attack phenol.
9. Pseudomonas cerevisiae.
bb. Attacks phenol.
10. Pseudomonas pictorum.
aa. Attack cellulose.
b. Litmus milk acid but no digestion.
11. Pseudomonas arguta.
bb. No growth in litmus milk.
12. Pseudomonas subcreta.
2. Nitrites not produced from nitrates; may or may not hydrolyze agar.
a. Butter-colored pellicle on litmus milk.
13. Pseudomonas lacunogenes.
aa. No surface pellicle.
14. Pseudomonas segnis.
Group I. -Plant pathogens.
1. Xanthomonas stewartii (Erw.
Smith, 1914) Dowson, 1939. (Sweet corn
bacillus, Stewart, N. Y. Agr. Exp. Sta. Bull.
130, 1897 , 423; Bacterium stewarti (sic) Smith,
Bacteria in Relation to Plant Diseases, S,
1914, 89; Xanthomonas stewarti (sic) Dow-
son, Zent. f. Bakt., II Abt., 100, 1939, 190.)
FAMILY IV. PSEUDOMONADACEAE
179
ste.war'ti.i. Stewart patronymic; JVI.L.
gen. noun stewartii of Stewart.
Description from Smith (U. S. Dept.
Agr., Div. Veg. Phys. and Path., Bull. 28,
1901).
Rods 0.4 to 0.7 by 0.9 to 2.0 microns.
Encapsulated. Non- motile (McCulloch,
Phytopath., 8, 1918, 440). Gram-negative.
Gelatin: No liquefaction.
Nutrient agar colonies: Small, round,
yellow.
Broth: Growth feeble with whitish ring
and yellow precipitate.
Milk: Yellow ring but no visible action
on the milk. Slightly acid.
Nitrites not produced from nitrates.
McNew (Phytopath., 28, 1938, 773) states
that less virulent strains assimilate only
organic nitrogen; those of intermediate
virulence assimilate nitrogen from inorganic
salts without reduction of nitrates to ni-
trites; virulent strains reduce nitrates to
nitrites.
Hydrogen sulfide not produced.
Indole production slight or none.
Reduction of methylene blue in Dun-
ham's solution feeble or doubtful.
Acid but no gas from glucose, galactose,
sucrose, mannitol and glycerol. No acid
from maltose. Acid from fructose, arabinose
and xylose (McNew, loc. cit.); also from
lactose and mannose (Dowson, op. cit.,
100, 1939, 190).
Starch not hydrolyzed.
Temperature relations: Optimum, 30° C.
Maximum, 39° C. Minimum, 8° C.
Chemical tolerance: Optimum pH be-
tween 6.0 and 8.0. Limits, about pH 4.5 to
8.5.
8 per cent salt restricts growth.
Strict aerobe.
Source: From wilted sweet corn.
Habitat: Pathogenic on corn, Zea mays.
Sweet corn very susceptible and field corn
slightly so.
2. Xanthomouas tardicrescens (Mc-
Culloch, 1937) Dowson, 1943. (Bacterium
tardicrescens McCulloch, Phytopath., £7,
1937, 135; Dowson, Trans. Brit. Mycol. Soc,
26, 1943, 12.)
tar.di.cres'cens. L. adj. tardus slow; L.
part. adj. crescens growing; M.L. adj. tardi-
crescens slow growing.
Rods 0.6 to 0.8 by 1.58 microns. Motile
with a polar flagellum. Gram-negative.
Gelatin: No liquefaction.
Beef -extract agar colonies: Circular,
mustard -yellow, edges entire, 1.0 to 1.5 mm
in diameter.
Broth: Light clouding.
Milk: Slightly alkaline. Clearing after 5
to 6 weeks.
Nitrites produced from nitrates.
Indole not produced.
Hydrogen sulfide not produced, or feebly
so.
Acid but no gas from glucose, fructose,
galactose, arabinose, xylose and rhamnose.
Alkaline reaction from salts of citric, malic
and succinic acids.
Starch not hydrolyzed.
Not lipolytic (Starr and Burkholder,
Phytopath., 82, 1942, 603).
Temperature relations: Optimum, 26° C.
Maximum, 32° C. Minimum, 5° C. (McCul-
loch, Phytopath., 28, 1938, 648).
Chemical tolerance: Optimum pH be-
tween 6.5 and 7.5. Growth slight at 5.8 and
8.0 (McCulloch, loc. cit.).
No growth with 3 per cent salt (McCul-
loch, loc. cit.).
Aerobic.
Distinctive character: Very slow grower.
Source: Isolated by McCulloch and by
Burkholder from blighted iris leaves.
Habitat: Pathogenic on Iris spp.
3. Xanthomonas albilineans (Ashby,
1929) Dowson, 1943. {Bacterium albilineans
Ashby, Trop. Agr., Trinidad, 6, 1929, 135;
Dowson, Trans. Brit. Mycol. Soc, .^5,1943,
11.)
al.bi.li'ne.ans. L. adj. albus white; L.
part. adj. lineans striping; M.L. adj. albili-
neans white-striping.
Description from Martin, Carpenter and
Weller (The Hawaiian Planters' Record,
36, 1932, 184).
Rods, 0.25 to 0.3 by 0.6 to 1.0 micron,
occurring singly or in chains. Motile with a
polar flagellum. Gram-negative.
Agar colonies: After 7 to 10 days, minute
transparent drops, moist, shining. Honey-
yellow to Naples -yellow.
180
ORDER I. PSEUDOMONADALES
Gelatin: No liquefaction.
Milk: Growth, but no visible change in
the milk.
No growth with ammonium salts, nitrates
or asparagine as a source of nitrogen.
No growth in peptone water without
carbohydrates. Invertase secreted.
Starch not hj'drolyzed.
Temperature relations: Optimum, about
25° C. Maximum, 37° C.
Distinctive characters: Differs from
Xanthomonas vasculorum, which produces a
large gummy type of colony and which is
a very active organism biochemically. The
two pathogens also differ in the type of
lesion they produce on sugar cane.
Source : Isolated by D. S. North (Colonial
Sugar Ref. Co., Sydney, N. S. Wales, Agr.
Rept., 8, 1926, 1) from white stripe and leaf
scald of sugar cane in Australia.
Habitat: Vascular pathogen of sugar
cane, Saccharum officinarum.
Group II. — Saprophytic species.
4. Pseudomonas trifolii Huss, 1907.
(Huss, Cent. f. Bakt., II Abt., 19, 1907, 68;
Xanthomonas trifolii James, Canadian Jour.
Microbiol., 1, 1955, 479.)
tri.fo'li.i. L. trifoUum trefoil, clover;
M.L. neut.n. TrifoUum generic name of
clover; M.L. gen. noun trifolii of TrifoUum.
Rods, 0.5 to 0.7 by 0.75 to 2.0 microns,
occurring singly, in pairs and in chains.
Motile, possessing a single polar flagellum.
Gram-negative.
Gelatin colonies: Convex, smooth, moist,
glistening, grayish yellow.
Gelatin stab: Napiform liquefaction.
Agar colonies: Small, circular, grayish,
becoming brownish yellow.
Agar slant: Yellowish, becoming brown-
ish yellow streak, lacerate margin.
Broth: Turbid, with grayish yellow pel-
licle and sediment.
Litmus milk: Slowly coagulated; alkaline;
with yellow ring.
Potato: Thick, yellowish, fiat, smooth,
glistening.
Hydrogen sulfide produced.
Indole produced.
Acid from glucose, sucrose, xylose, arabi-
nose and mannitol. No acid from lactose.
Nitrites produced from nitrates.
Cultures have an agreeable odor.
Volutin formed.
Aerobic, facultative.
Optimum temperature, between 33° and
35° C.
Source: Isolated from clover hay.
Habitat: Evidently a common organism
on the leaves of plants.
5. Pseudomonas xanthe Zettnow, 1916.
Cent. f. Bakt., I Abt., Orig., 77, 1916, 220.)
xan'tha. Gr. adj. xanthus yellow.
Rods 0.5 to 0.6 by 0.4 to 1.4 microns.
Motile, possessing a single or occasionally
two or more very long (20 microns) polar
flagella. Gram-negative.
Gelatin colonies: Circular, yellow, granu-
lar.
Gelatin stab: Pale yellow surface growth.
Brownish j^ellow under surface colonies.
Saccate liquefaction.
Agar slant: Dark yellow, glistening, with
dark yellow sediment in water of condensa-
tion. Pigment not water-soluble.
Broth: Turbid.
Milk becomes rose-yellow in 4 weeks with-
out anj^ other change.
Potato: Grayish yellow to brownish
growth.
Indole produced.
Nitrites produced from nitrates.
Acid produced from glucose, sucrose and
maltose.
Starch hydrolyzed.
Blood serum not liquefied.
Aerobic, facultative.
Optimum temperature, 30° C.
Source: Air contamination.
6. Pseudomonas caudata (Wright,
1895) Conn, 1919. (Bacillus caudatus Wright,
Memoirs Nat. Acad. Sci., 7, 1895, 444;
Conn, Jour. Agr. Res., 16, 1919, 313.)
cau.da'ta. L. noun canda a tail; M.L. adj.
caudatus having a tail.
Rods long, granular, slender, occurring
singly, in pairs and in chains. Appear like
cocci in old cultures. Motile, possessing a
polar flagellum (Conn). Gram-negative.
Gelatin colonies: Yellow, translucent,
smooth, undulate.
Gelatin stab: Villous growth in stab.
Crateriform liquefaction.
FAMILY IV. PSEUDOMONADACEAE
181
Agar slant: Yellow to orange, glistening,
translucent, slightly spreading. May lose
power to form pigment.
Broth: Turbid, with yellow sediment.
Litmus milk: Unchanged.
Potato: Dark yellow, raised, rough,
spreading.
Indole not produced.
Nitrites and ammonia produced from
nitrates.
Ammonia produced from peptone.
Starch digested.
Aerobic, facultative.
Optimum temperature, 25° C.
Habitat: Water and soil.
7. Pseudonionas perlurida Kellerman
et al., 1913. (Kellerman, McBeth, Scales
and Smith, Cent, f . Bakt., II Abt., S9, 1913,
516; also see McBeth, Soil Sci., /, 1916,
472.)
per.lu'ri.da. L. prefi.x per very; L. luridus
pale yellow, sallow; M.L. adj. perluridus
verj' sallow.
Rods 0.4 by 1.0 micron. Motile with one
to three polar flagella. Gram-negative.
Gelatin stab: Liquefied.
Agar slant: Moderate, flat, faint yellow
growth.
Broth: Turbid in 5 days.
Litmus milk: Acid. Peptonization after
16 days.
Potato: Scant, 3'ellow growth with
bleaching along line of growth.
Indole not produced.
Nitrites not produced from nitrates.
Ammonia produced.
Acid from glucose, maltose, lactose,
sucrose, starch, glycerol and mannitol.
Aerobic, facultative.
Optimum temperature, 20° C.
Source: Soil from Virginia, Louisiana
and Missouri.
Habitat: Soil.
8. Pseudonionas ochracea (Zimmer-
mann, 1890) Chester, 1901. {Bacillus ochra-
ceus Zimmermann, Bakt. unserer Trink-
und Nutzwasser, Chemnitz, /, 1890, 60;
Chester, Man. Determ. Bact., 1901, 316.)
och.ra'ce.a. Gr. noun ochra ochre; M.L.
adj. ochraceus of the color of ochre.
Rods, 0.7 to 0.8 by 1.2 to 4.5 microns,
occurring in pairs and longer chains. Slow,
undulatory motion (Zimmermann). Polar
flagella (Lehmann and Neumann, Bakt.
Diag., 1 Aufl., 2, 1896, 255). Gram-negative.
Gelatin colonies: Pale yellow to golden,
ochre-yellow, slightly raised, with slightly
fringed margin, granular.
Gelatin stab: Yellowish to yellow -gray
surface growth. Infundibuliform lique-
faction. Pale yellow to ochre-yellow sedi-
ment.
Agar colonies: Thin, flat, yellowish,
smooth.
Agar slant: Thin, yellowish gray to
ochraceous growth.
Broth: Slightly turbid, with pale yellow
sediment.
Litmus milk: Medium becomes slimv:
alkaline.
Potato: Ochre-yellow streak.
Indole produced.
Nitrites not produced from nitrates.
Hydrogen sulfide produced.
Aerobic, facultative.
Optimum temperature, 35° C.
Source: Chemnitz tap water.
Habitat: Water.
9. Pseudonionas cerevisiae Fuhrmann,
1906. (Cent. f. Bakt., II Abt., 16, 1906, 309.)
ce.re.vi'si.ae. L. cerevisia beer; M.L.
gen. noun cerevisiae of beer.
Rods straight or slightly curved, 0.6
by 1.5 to 2.0 microns, occurring singly and
in chains. Motile, possessing a tuft of four
to six polar flagella. Gram-negative.
Gelatin colonies: Circular, white, slightly
contoured, becoming brownish yellow.
Gelatin stab: Slight yellowish growth in
stab. No liquefaction.
Agar colonies: Thin, spreading, con-
toured.
Agar slant: Moist, glistening, thin, pale
yellow, spreading, contoured.
Litmus milk: Slow coagulation.
Potato: Yellowish brown, spreading
growth.
Indole not produced.
Nitrites produced from nitrates.
No gas from glucose.
Aerobic, facultative.
Optimum temperature, 30° C.
Source: Isolated from beer.
Habitat: Unknown.
182
ORDER I. PSEUDOMONADALES
10. Pseudomonas pictorum Gray and
Thornton, 1928. (Cent. f. Bakt., II Abt., 73,
1928, 89.)
pic.to'rum. Named for the Picts, a Scot-
tish tribe; M.L. neut.pl. gen. n. 'pictorum of
the Picts.
Rods 0.5 to 0.8 by 1.5 to 5.0 microns.
Motile, usually with a single polar flagel-
lum. Gram-negative.
Gelatin colonies: Circular, greenish
yellow, convex, smooth, glistening, entire.
Gelatin stab: No liquefaction.
Agar colonies: Circular, yellow, conve.x,
smooth, glistening, entire.
Agar slant: Filiform, yellow, convex,
smooth, glistening, entire.
Broth: Turbid.
Nitrites produced from nitrates.
Starch not hydrolyzed.
Acid from glucose and maltose.
Attacks phenol.
Aerobic, facultative.
Optimum temperature, 25°C.
Source: One culture from soil.
Habitat: Soil.
11. Pseudomonas argiita McBeth, 1916.
(Soil Science, 1, 1916, 465.)
ar.gu'ta. L. part. adj. argutus clear,
bright.
Rods 0.3 by 0.8 micron. Motile with one
or two polar flagella. Gram-negative.
Gelatin stab : Moderate, yellowish growth.
No liquefaction in 30 daj^s.
Agar colonies: Circular, slightly convex,
soft, grayish white, granular, entire.
Agar slant: Scant, grayish white growth.
Potato agar slant: Moderate, yellowish,
glistening growth.
Broth: Turbid.
Ammonia cellulose agar: Enzymatic zone
2 to 3 mm in 30 days.
Filter paper broth: Paper is reduced to
loose flocculent mass which disintegrates
very readily on slight agitation. More rapid
decomposition when the broth contains
ammonium sulfate, potassium nitrate, pep-
tone or casein as source of nitrogen.
Litmus milk: Acid, not digested.
Potato: No growth.
Indole not produced.
Nitrites produced from nitrates.
Ammonia not produced.
Acid from glucose, maltose, lactose and
starch. No acid from glycerol, mannitol or
sucrose.
Aerobic, facultative.
Optimum temperature, 20° C.
Source: Isolated twice from California
soils.
Habitat: Soil.
12. Pseudomonas subcreta McBeth and
Scales, 1913. (Bur. Plant Industry, U. S.
Dept. Agr., Bull. 266, 1913, 37.)
sub.cre'ta. L. pref. sub- somewhat; L.
creta chalk; M.L. adj. subcretus somewhat
chalky.
Rods 0.3 by 1.4 microns. Motile with one
to five polar flagella. Gram-negative.
Gelatin stab: Filiform growth, no lique-
faction.
Cellulose agar: No surface growth.
Moderate, generally faint yellow growth
in medium, area of growth sunken.
Agar slant: Glistening, smooth, moist,
vitreous to faint yellow growth.
Starch agar: Enzymatic zone 2 to 4 mm.
Broth: No growth.
Litmus milk: No growth.
Potato: Growth scant, concave due to
slight liquefaction, white to faint yellow.
Bleached around growth.
Indole not produced.
Trace of nitrites produced from nitrates.
Ammonia not produced.
Acid from glucose, lactose, maltose,
sucrose and starch. No acid from glycerol
or mannitol.
Aerobic, facultative.
Optimum temperature, 20° C.
Habitat: Soil.
13. Pseudomonas lacunogenes Gores-
line, 1933. (Jour. Bact., £6, 1933, 447.)
la.cu.no'ge.nes. L. lacuna a hollow; L.
gicjno to produce; M.L. adj. lacunogenes pro-
ducing hollows.
Short rods, 0.2 to 0.3 by 1.0 to 1.2 microns,
with pointed ends, occurring singly or in
pairs. Motile with a single polar flagellum
from 2 to 15 microns in length. Gram-nega-
tive.
Plain gelatin stab: No growth.
Nutrient gelatin stab: Growth brownish
yellow, half-way down stab, heavier at
surface. No liquefaction.
FAMILY IV. PSEUDOMONADACEAE
183
Nutrient agar colonies: Small, yellow;
surface of the agar pitted or dimpled. After
5 days colonies 5 to 7 mm in diameter,
orange-yellow, slightly raised, surrounded
b.y a depression.
Nutrient agar slant: Growth heavy, light
orange-yellow; consistency of warm butter;
edge entire, slightly raised. Shallow de-
pression formed on each side of streak.
Agar softened beneath growth.
Nutrient broth: Turbid in 48 hours. Light
orange-j'ellow pellicle; considerable, viscous
sediment.
Litmus milk: Alkaline; butter-colored
pellicle. Reduction in bottom of tube after
10 daj's. No curd. No digestion.
Potato: Growth moderate, orange-yellow,
smooth. No darkening.
Indole not produced.
Nitrites not produced from nitrates.
Starch agar plates not hydrolyzed.
Utilizes arabinose, galactose, lactose,
fructose, maltose, melezitose, raffinose,
starch, xj-lose, glucose, mannose, sucrose,
pectin, rhamnose, salicin and dextrin. No
growth in dulcitol, erythritol, glycerol,
sorbitol, mannitol or inulin.
Temperature relations: Optimum, 28° C.
Good growth at 25° C. Moderate growth at
20° and at 37° C. No growth at 10° and at
42° C.
Limits of pH: 5.4 to 10.0.
Aerobic, facultative.
Distinctive characters: Softens agar;
considerable change in viscosity of agar due
to this digestion; utilization of ammonium
sulfate as nitrogen source.
Source: Three cultures isolated from an
experimental trickling filter receiving
creamery wastes.
Habitat: Probably widely distributed in
nature.
14. Pseudonionas segnis Goresline,
1933. (Jour. Bact., 26, 1933, 452.)
seg'nis. L. adj. segnis slow, tardy.
Short rods, 0.2 to 0.3 by 1.0 to 1.2 microns,
with pointed ends, occurring singly or in
pairs. Motile with a single polar flagellum.
Gram-negative.
Plain gelatin stab: No growth.
Nutrient gelatin stab: Growth yellow,
half-way down stab, best at surface. No
liquefaction.
Nutrient agar colonies: Very small, light
yellow surface pitted. After 5 days colonies
5 mm in diameter.
Nutrient agar slant: Growth heavy,
orange -yellow, consistency of warm butter;
edge entire, slightly raised; slight de-
pression formed on each side of growth.
Agar softened beneath growth.
Nutrient broth: Turbid in 48 hours. No
pellicle or surface growth. Moderate amount
of sediment. Old cultures with a yellow ring
at surface and occasionally a loose mem-
brane.
Litmus milk: Slightly alkaline after 10
daj's. No reduction. No surface growth.
Potato: Scant j'ellow-orange growth. No
darkening.
hidole not produced.
Nitrites not produced from nitrates.
Hydrogen sulfide not produced.
Starch not hydrolyzed.
Arabinose, glucose, galactose, lactose,
fructose, maltose, mannose, xylose, sucrose,
melezitose and raffinose utilized.
Temperature relations: Optimum, 28° C.
Good growth at 25° C. Moderate growth at
20° and at 37° C. No growth at 10° and at
42° C.
Limits of pH: 5.8 to 9.0.
Aerobic, facultative.
Distinctive characters: Softens agar; con-
siderable change in viscosity of agar due to
this digestion.
Source: Isolated from an experimental
trickling filter receiving creamery wastes.
Habitat: Probably widely distributed in
nature.
Genus III. Acetobacter Beijerinck, 1898.*
{Acetobacter Beijerinck, quoted from Krai's Sammlung v. Mikroorg., Prague, 1898, 7;
Acetobacterium in Ludwig's abstract of Hoyer, Bijdrage tot de kennis van de azijnbacterien,
* Revised by Dr. Reese H. Vaughn, Univ. of California, Berkeley, California, June,
1943, and Davis, California, March, 1954.
184 ORDER I. PSEUDOMONADALES
Thesis, Leiden, 1898, 115 pp.. Delft, in Cent. f. Bakt., II Abt., 4, 1898, 857; Acetobader
Beijerinck, Proc. Kon. Akad. v. Wetenschapp., Amsterdam, 2, 1900, 503; Acetobacter Bei-
jerinck. Arch, n^erl. d. sciences exact, et natur., S^r. II, 6, 1901, 212; Acetobacter in Fuhr-
mann, Beiheft Bot. Centralbl., Orig., 19, 1905, 8; Acetimonas Orla-Jensen, Cent. f. Bakt.,
II Abt., 22, 1909, 312; Acetobacter Winslow et al.. Jour. Bact., 5, 1920, 201; Acetomonas
Leifson, Antonie van Leeuwenhoek, 20, 1954, 109.)
A.ce.to.bac'ter. L. noun acetum vinegar; M.L. mas.n. bacter the masculine form of the
Gr. neut.n. bactrum a rod or staff; M.L. mas.n. Acetobacter vinegar (acetic) rod.
Individual cells ellipsoidal to rod-shaped, occurring singly, in pairs or in short or long
chains. Motile with polar flagella*, or non-motile. Involution forms may be spherical,
elongated, filamentous, club-shaped, swollen, curved or may even appear to be branched.
Young cells Gram-negative; old cells often Gram- variable. Obligate aerobes; as a rule
strongly catalase-positive, sometimes weakly so. Oxidize various organic compounds to
organic acids and other oxidation products which may undergo further oxidation. Com-
mon oxidation products include acetic acid from ethyl alcohol, gluconic and 5-keto-
gluconic acid from glucose, dihydroxy-acetone from glycerol, sorbose from sorbitol, etc.
Nutritional requirements vary from simple to complex. Development generally best in
yeast infusion or yeast autolysate media with added ethyl alcohol or other o.xidizable sub-
strates. Optimum temperature varies with the species. Widely distributed in nature where
they are particularly abundant in plant materials undergoing alcoholic fermentation;
of importance to man for their role in the completion of the carbon cycle and for the pro-
duction of vinegar.
It is recognized that there are marked morphological and physiological similarities be-
tween species of Acetobacter and Pseudomonas (see Vaughn, Jour. Bact., 46, 1943, 394; and
Stanier, Jour. Bact., 54, 1947, 191, among others). However, the species of Acetobacter may
be differentiated from all other Pseudomonadaceae by their unique ability to oxidize sig-
nificant quantities of ethanol under the extremely acidic conditions imposed by the
presence of from about 2 to more than 11 per cent acetic acid.
The evidence also indicates a significant difference in the end-products of hexose and di-
saccharide oxidation. The species of Acetobacter produce gluconic and 5-ketogluconic acids
from both glucose and maltose whereas species of Pseudomonas oxidize glucose to gluconic
and 2-ketogluconic acids and maltose to maltobionic acid (see Pervozvanski, Khim. Referat.
Zhur., 7, 1939, 43; Lockwood, Tabenkin and Ward, Jour. Bact., 4^, 1941, 51; Stodola and
Lockwood, Jour. Biol. Chem., 171, 1947, 213; Kluyver, Deley and Rijven, Antonie van
Leeuwenhoek, 16, 1950, 1; and Foda and Vaughn, Jour. Bact., 65, 1953, 233, among others).
The type species is Acetobacter aceti (Beijerinck) Beijerinck.
Key to the species of genus Acetobacter.
I. Oxidize acetic acid to carbon dioxide and water.
A. Utilizes ammonium salts as a sole source of nitrogen (Hoyer's solution). f
1. Acetobacter aceti.
* Leifson (Bact. Proc, 53rd Gen. Meeting Soc. Amer. Bact., 1953, 34, and Antonie van
Leeuwenhoek, 20, 1954, 102), in a study of the flagellation of cultures of Acetobacter, reports
that the species of Acetobacter that oxidize acetic acid are peritrichous, and that the species
that do not oxidize acetic acid ordinarily have four polar flagella. Further photographs such
as can be obtained with the electron microscope must, however, be obtained before the exact
point of attachment of the flagella can be determined with certaint3^
t It is not known with certainty whether Acetobacter pasteurianus and Acetobacter kuetz-
ingianus are capable of using inorganic nitrogen as a sole source of nitrogen for growth.
See Acetobacter rancens Beijerinck to which these two species are very closely related. Also
see Frateur, La Cellule, 53, 1950, 316^320.
Species Nos. 2 to 3b inclusive will, however, utilize ammonium salts if supplied with
FAMILY IV. PSEUDOMONADACEAE
185
B. Do not utilize ammonium salts as a sole source of nitrogen.
1. Forms a thick, zoogloeal, cellulose membrane on the surface of liquid media.
2. Acetobacter xylinum.
2. Do not form a thick, zoogloeal, cellulose membrane on the surface of liquid media.
3. Acetobacter rancens.
3a. Acetobacter pasteurianus .
3b. Acetobacter kuetzingianus .
II. Do not oxidize acetic acid.
A. Form pigments in glucose media.
1. Dark brown to blackish pigment.
4. Acetobacter melanogenus.
2. Pink to rose pigment.
5. Acetobacter roseus.
B. Do not form pigments.
1. Optimum temperature, between 30° and 35° C.
6. Acetobacter suboxydans.
2. Optimum temperature, between 18° and 21° C.
7. Acetobacter oxydans.
1. Acetobacter aceti (Beijerinck, 1898)
Beijerinck, 1900. (Mycodermes, Pasteur
Compt. rend. Acad. Sci., Paris, 54, 1862
265; Pasteur, ibid., 55, 1862, 28; Mycoderma
aceti Pasteur, Ann. Sci. d. Ecole Normal
superiore, 1, 1864, 103-158; Bacterium aceti
Beijerinck, Cent. f. Bakt., II Abt., 4, 15
211; Acetobacter aceti Beijerinck, published
as a synonym in Krai's Sammlung v. Mikro
org., Prague, 1898, 7; Beijerinck, Proc. Kon
Akad. V. Wetensch., Amsterdam, 2, 1900
503.)
a.ce'ti. L. noun acetum vinegar; L. gen
noun aceti of vinegar.
Beijerinck's description of this organism
which forms the basis of the description
given here, is based on Pasteur's earlier de
scription.*
Rods, 0.4 to 0.8 by 1.0 to 2.0 microns, oc
curring singly and in long chains, frequently
showing large club-shaped forms. Stain yel
low with iodine solution. Motility variable
Motile cells possess a single polar flagellum
(Vaughn, Jour. Bact., 46, 1943, 394).
Beer gelatin containing 10 per cent su-
crose: Large, shiny colonies are formed.
Liquid media: Forms slimy pellicle; may
also form a ring or turbidity without pel-
licle.
Acid from glucose, ethanol, propanol and
glycol. No acid from arabinose, fructose,
galactose, sorbose, sucrose, maltose, lactose,
raffinose, dextrin, starch, glycogen, inulin,
methanol, isopropanol, butanol, isobutanol,
pentanol, glj'cerol, erythritol, mannitol,
dulcitol or acetaldehyde (Henneberg, Die
deutsch. Essigind., 2, 1898, 147).
Distinctive characters : Marked oxidative
power causing rapid and complete oxidation
of substrate such as glucose or ethyl alcohol ;
ability to utilize inorganic nitrogen salts as
a sole source of nitrogen (Hoyer, Inaug.
Diss., Leiden, 1898, 43; Beijerinck, Cent. f.
Bakt., II Abt., 4, 1898, 215); growth and
oxidative activity in association with fer-
menting yeasts (Vaughn, Jour. Bact., 36,
1938, 360).
Optimum temperature, 30° C. Growth
occurs between 10° and 42° C.
Habitat: Vinegar, souring fruits, vege-
tables and beverages.
other required nutrients (Stokes and Karsen, Jour. Bact., 49, 1945, 495; Foda and Vaughn,
Jour. Bact., 65, 1953,79).
* Beijerinck (op. cit., 4, 1898, 211) explains the relationship of Pasteur's organism to
those described by others as follows: "Two of the many varieties of B. (Bacterium) rancens
have been described by Henneberg under the names B. oxydans and B. acetosum. Hansen
erroneously named this species B. aceti as did Brown also. Neither Hansen nor Brown
knew B. aceti of Pasteur."
186
ORDER I. PSEUDOMONADALES
2. Acetobacter xylinum (Brown,
Holland, 1920. {Bacterium xylinum Brown,
Jour. Chem. Soc, London, Jfi, 1886, 439;
Holland, Jour. Bact., 5, 1920, 216.)
xy'li.num. Gr. adj. xylinus of cotton; L.
neut.n. xylinum cotton.
Rods, about 2 microns long, occurring
singl}^ and in chains. The cells have a slimy
envelope which gives the cellulose reaction.
A zoogloeal film forms on all liquid media
in which growth occurs; the nature of the
medium influences the thickness of the film
which may vary from 2 to 250 millimeters.
The film becomes cartilagenous and falls
to the bottom if disturbed.
X-ray pattern studies made by Khouvine,
Champetier and Sutra (Compt. rend. Acad.
Sci. Paris, IH, 1932, 208) and by Barsha and
Hibbert (Can. Jour. Research, 10, 1934,
170) have shown that the cellulose contained
in the membranes formed by Acetobacter
xylinum is identical with cotton cellulose.
Acid from glucose, ethanol, propanol and
glycol. No acid from arabinose, fructose,
galactose, maltose, lactose, raffinose, dex-
trin, starch, methanol, isopropanol, bu-
tanol, isobutanol, pentanol, mannitol or
acetaldehyde (Henneberg, Die deutsch.
Essigind., 2, 1898, 147).
Distinctive character: The production of
thick, leather}'', zoogloeal, cellulosic mem-
branes on the surface of liquids.
Optimum temperature, 28° C.
Habitat: Vinegar, souring fruits, vege-
tables and beverages.
3. Acetobacter rancens Beijerinck, 1898.
{Bacterium rancens Beijerinck, Cent. f.
Bakt., II Abt., 4, 1898, 211; Beijerinck, in
Krai's Sammlung v. Mikroorg., Prague,
1898, 4.)
ran'cens. L. part. adj. rancens putrid,
stinking.
The following description is taken in part
from a study of a culture of Acetobacter
rancens received from Kluyver by Vaughn;
also see Frateur (La Cellule, 53, 1950, 339).
Rods with the usual morphological ap-
pearance of cultures of acetic-acid bacteria.
Gram-negative. Molitility variable. Motile
cells possess a single polar fiagellum
(Vaughn, Jour. Bact., ^6, 1943, 394). Involu-
tion forms commonly appear as filaments
and enlarged cells.
Wort agar slant: Growth abundant,
butyrous, pale-buff in color in one week.
Yeast infusion, glucose, calcium carbon-
ate slant: Growth abundant, butyrous and
cream-colored in one week.
With petri-dish cultures, well isolated
colonies are large, smooth and butyrous on
the above-mentioned media.
Broth cultures containing peptone or
yeast infusion form a mucilaginous, slimy
pellicle. Beijerinck {op. cit., 4, 1898, 211)
called this polysaccharide pellicle cellulose-
like and intimated that the mucilaginous
material in the pellicle was somewhat
different from that produced by Acetobacter
xylinutn. The pellicle material stained blue
when treated with iodine and hydroiodic
acid.
Minimum nutritional requirements: Pan-
tothenic acid, nicotinic acid, p-aminoben-
zoic acid, thiamine, valine, alanine, isoleu-
cine, histidine, cystine, proline, aspartic or
glutamic acid, mineral salts and an oxidiz-
able substrate such as alcohol, glucose, etc.
(Foda and Vaughn, Jour. Bact., 65, 1953,
79).
Acid from glucose, ethanol, propanol,
butanol, glycol, adonitol, mannitol and sor-
bitol. No acid from numerous other com-
pounds tested.
Distinctive character: Production of a
thin, mucilaginous, slimj^, polysaccharide
membrane on the surface of liquids as com-
pared with the thick, true cellulose mem-
brane of Acetobacter xylinum grown under
the same conditions. Beijerinck {op. cit., 4-,
1898, 211) reported the production of a
cellulose-like membrane with some cultures
of Acetobacter rancens.
Source: Isolated from shavings in the
quick- vinegar process.
Habitat: Found in fermented grain mash,
malt beverages, mother of vinegar, and
souring fruits.
Beijerinck (Cent. f. Bakt., II Abt., 4,
1898, 211) thought that the next two species
were hardly more than varieties of Acetobac-
ter rancens; also see Frateur (La Cellule,
53, 1950, 339).
FAMILY IV. PSEUDOMONADACEAE
187
3a. Acetobacter pasteurianus (Han-
sen, 1879) Beijerinck, 1916. (Mycoderma
pasteurianum Hansen, Compt. rend. d.
Trav. d. Lab. d. Carlsberg, 1, 1879, 96; Bei-
jerinck, Proc. Sect. Sci., Kon. Akad. v.
Wetenschappen, Amsterdam, 18, 1916, 1199.)
pas.teur.i.a'nus. Named for Pasteur,
French chemist and bacteriologist; M.L.
adj . pasteurianus of Pasteur.
Rods, 0.4 to 0.8 by 1.0 micron, occurring
singly and in chains, at times showing thick,
club-shaped forms. Motility variable.
Motile cells possess a single polar flagellum
(Vaughn, Jour. Bact., 46, 1943, 394). Stain
blue with iodine.
Wort gelatin colonies: Small, circular,
entire, gray, slimy.
Forms a dry, wrinkled, folded pellicle on
double beer with one per cent alcohol.
Meat infusion gelatin: Growth wide-
spread; later rosette form, toothed.
Acid from glucose, ethanol, propanol and
glycol. No acid from arabinose, fructose,
galactose, sorbose, sucrose, maltose, lactose,
raffinose, dextrin, starch, glycogen, inulin,
methanol, isopropanol, butanol, isobutanol,
pentanol, glycerol, erythritol, mannitol,
dulcitol or acetaldehyde (Henneberg, Die
deutsch. Essigind., 2, 1898, 147).
Optimum temperature, 30° C. Growth
occurs between 5° and 42° C.
Habitat: Vinegar; beer and beer wort.
3b. Acetobacter kuetzingianus (Han-
sen, 1894) Bergey et al., 1923. {Bacterium
klitzingianum (sic) Hansen, Compt. rend. d.
Trav. d. Lab. d. Carlsberg, 3, 1894, 191;
Bergey et al.. Manual, 1st ed., 1923, 35.)
kuet.zing.i.a'nus. Named for Kuetzing,
a German botanist; AL L. adj. kuetzingianus
of Kuetzing.
Short, thick rods, occurring singly, rarely
forming chains of notable length. Capsule
stains blue with iodine and with potassium
iodide. Non-motile.
Double beer gelatin colonies: Small, en-
tire, w'ith vermiform surface.
Wort gelatin colonies: Small, entire, with
surface free of wrinkles.
Double beer: Forms a rather thick, folded
pellicle. Distinguished from Acetobacter
aceti in showing a heavier growth above the
surface of the medium.
Acid from glucose, ethanol, propanol and
glycol. No acid from arabinose, fructose,
galactose, sorbose, sucrose, maltose, lactose,
rafiinose, dextrin, starch, glycogen, inulin,
methanol, isopropanol, butanol, isobutanol,
pentanol, glycerol, erythritol, mannitol,
dulcitol or acetaldehyde (Henneberg, Die
deutsch. Essigind., 2, 1898, 147).
Optimum temperature, 34° C; minimum,
between 6° and 7° C; maximum, 42° C.
Habitat: Beer. Found in double beer.
4. Acetobacter inelanogenus Beije-
rinck, 1911. (Cent. f. Bakt., II Abt., 29,
1911, 175.)
me.la.no'ge.nus. Gr. adj. melas, melanis
black; Gr. v. gennao to produce; M.L. adj.
inelanogenus black-producing.
Rods. Non-motile or motile. Motile cells
possess a single polar flagellum (Vaughn,
Jour. Bact., 46, 1943, 394).
Gelatin: Apparent liquefaction probably
caused by acid, not an enzyme. When held
on artificial media for some time, the power
of liquefying gelatin is lost, probably due to
a slower production of acid. Deep brown pig-
ment produced; gelatin becomes insoluble
in boiling water and in trypsin solution.
Beer- or wort-gelatin plates: Characteris-
tic dark brown, wide-spreading, diffuse
areas.
Tap water - agar - glucose - peptone - potas-
sium phosphate-iron citrate-chalk medium:
In 24 hours at 30° C, black, spreading, dif-
fuse areas.
Produces the pigment from peptone or
yeast autolysate if maltose or glucose is
present as a source of carbon. When grown in
glucose-peptone broth or agar with CaCOa
at 25° to 30° C, the pigment is produced
after one to several weeks.
Pigment : The pigment causing the brown
coloration is an aromatic substance which is
blackened by iron salts. Reduces alkaline
solutions of silver and mercury, blackening
them (Beijerinck, op. cit., 29, 1911, 175).
Minimum nutritional requirements : Pan-
tothenic acid, nicotinic acid, p-aminoben-
zoic acid, thiamine, mineral salts and an
oxidizable substrate such as alcohol, glu-
188
ORDER I. PSEUDOMONADALES
cose, etc. (Gray and Tatum, Proc. Nat'l.
Acad. Sci., 30, 1944, 404, and Foda and
Vaughn, Jour. Bact., 65, 1953, 79).
Acetic acid produced from alcohol. Glu-
conic and 5-ketogluconic acids produced
from glucose and maltose.
O.xidizes mannitol to fructose; sorbitol to
sorbose; and glj^cerol to dihydroxj^acetone.
Produces acid from arabinose, .xylose, glu-
cose, fructose, galactose and maltose. Some
strains do not attack maltose.
Distinctive character: The formation of
dark brown to black pigment in media con-
taining glucose.
Source: Isolated from beer.
Habitat : Causes light-colored beer to be-
come darker brown. It is a very strong beer-
vinegar bacterium. Also found in souring
fruits.
5. Acetobacter roseus Vaughn, 1942.
{Bacterium hoshigaki var. rosea Takahashi
and Asai, Zent. f. Bakt., II Abt., 82, 1930,
390; Vaughn, Wallerstein Lab. Communica-
tions, 5, No. 14, 1941,20.)
ro'se.us. L. adj roseus rose -colored.
Rods, 0.7 to 0.9 by 1.5 to 1.8 microns,
generally occurring singly, in pairs, often
in chains. Non-motile. Pellicle on fluid
media yields no starch or cellulose reaction.
Koji (a mixture of rice and mold spores
used to start fermentation of Japanese bread
and sake) extract agar colonies: Small,
granular, circular, glistening, umbonate,
becoming brownish.
Wort agar colonies: Circular, milky
white, becoming brownish in the center and
yellowish at the periphery.
Koji extract agar streak: Grayish white,
glistening with ciliate margin, becoming
purple-brown to brown.
Koji extract: Turbid, with thin film as-
cending on wall of tube.
Bouillon: Turbid with ring formation.
Yeast infusion glucose agar: Colonies
similar to those on wort agar.
Yeast infusion glucose broth: Turbid with
thin, ascending film.
Red color produced on sake-wort agar and
all media containing calcium carbonate.
Acid from glucose, fructose, galactose,
arabinose, glycerol, mannitol, ethanol and
propanol. No acid from maltose, sucrose.
lactose, raffinose, de.xtrin, starch, inulin,
sorbitol, glycogen, isodulcitol or methanol.
Optimum temperature, between 30° and
35° C; minimum, between 10° and 15° C;
maximum, between 40° and 41° C.
Thermal death point, 50° C. for 5 minutes.
Distinctive character: The formation of a
rose to red pigment in suitable media, par-
ticularly those containing glucose and cal-
cium carbonate.
Note: Vaughn {loc. cit.) has proposed the
name Acetobacter roseus to replace the name
Acetobacter hoshigaki. As originally de-
scribed, this organism was given the name
Bacterium hoshigaki var. rosea by Takahashi
and Asai {op. cit., 82, 1930, 390) without the
authors having first named and described
the species Bacterium hoshigaki. The Japa-
nese word "hoshigaki" has been used in a
confusing manner, viz. Bacterium indiis-
trium var. hoshigaki (Takahashi and Asai,
loc. cit.) and Bacterium hoshigaki var.
glucuronicum I, II and III (Takahashi and
Asai, Jour. Agr. Chem. Soc. Japan, 9, 1933,
351 and Zent. f. Bakt., II Abt., 87, 1933,
385). None of these Japanese names are in
the form of true binomials.
Source: Isolated from fermenting mash of
dried persimmons (hoshigaki) ; also from
souring figs and dates.
6. Acetobacter suboxydans Kluyver
and de Leeuw, 1923. (Paper read at the con-
vention of the Dutch Society of Micro-
biology, Utrecht, December, 1923; see
Tijdschrift v. Vergelijkende Geneeskunde,
10, Afl. 2-3, 1924.)
sub. ox'y. dans. L. pref. sub- somewhat,
slightly; Gr. adj. oxys sharp; M.L. part. adj.
oxydans oxidizing; M.L. part. adj. suboxydans
slightly o.xidizing.
Short rods. Occur singly or in chains.
Non-motile. Morphologically like Aceto-
bacter rancens.
Forms a very thin, hardlj' visible pellicle
on fluid media.
Wort agar colonies: Very small, circular,
slightly yellow.
Minimum nutritional requirements: Pan-
tothenic acid, nicotinic acid, p-aminoben-
zoic acid, valine, alanine, isoleucine,
histidine, cystine, proline, mineral salts
and an oxidizable substrate such as alcohol,
FAMILY IV. PSEUDOMONADACEAE
189
glucose, etc. (Landy and Dicken, Jour.
Biol. Chem., H6, 1942, 109; Lampen, Under-
kofler and Peterson, Jour. Biol. Chem., 1^6,
1942, 277; Underkofler, Bantz and Peterson,
Jour. Bact., 45, 1943, 183; Stokes and Lar-
sen, Jour. Bact., J^, 1945, 495).
Acid from ethanol, propanol, glycol,
glucose, glycerol and sorbitol.
Optimum temperature, 30° C.
Distinctive character: Partial o.xidation
of substrates as indicated by the formation
of calcium 5-ketogluconate crystals on the
surface of agar slants containing glucose and
calcium carbonate.
Source: Isolated from spoiled beer.
Habitat: Beer; also found in souring
fruits and wine vinegar.
7. Acetobacter oxydans (Henneberg,
1897) Bergey et al., 1923. {Bacterium oxydans
Henneberg, Cent. f. Bakt., II Abt., 3, 1897,
223; Bergey et al.. Manual, 1st ed., 1923,
36.)
ox'y.dans. Gr. adj. oxys sharp; M.L.
part. adj. oxydans oxidizing.
Rods, 0.8 to 1.2 by 2.4 to 2.7 microns,
occurring singly and in chains. Motile cells
possess a single polar flagellum (Vaughn,
Jour. Bact., 46, 1943, 394). The chains show
bud-like swellings.
Gelatin colonies: Circular, becoming
irregular in shape with peculiar ramifica-
tions.
Minimum nutritional requirements: Pan-
tothenic acid, nicotinic acid, p-amino-
benzoic acid, valine, alanine, isoleucine,
histidine, cystine, proline, mineral salts
and an o.xidizable substrate such as alcohol,
glucose, etc. (Foda and Vaughn, Jour.
Bact., 65, 1953, 79).
Acid from arabinose, fructose, glucose,
galactose, sucrose, maltose, rafiinose, dex-
trin, ethanol, propanol, erythritol, man-
nitol, glycol or glycerol. No acid from
sorbose, lactose, starch, gl3^cogen, inulin,
methanol, isopropanol, butanol, isobutanol,
pentanol, dulcitol or acetaldehyde (Hen-
neberg, Die deutsch. Essigind., 2, 1898, 147).
Optimum temperature, between 18° and
21° C.
Distinctive characters: Low optimum
temperature for growth and oxidation of
substrates; also the ability to oxidize a
large number of substrates.
Habitat: Beer, souring fruits, wine vine-
gar.
Genus IV. Aeromonas Kluyver and van Niel, 1936.*
(Zent. f. Bakt., II Abt., 94, 1936, 398.)
A.e.ro.mo'nas. Gr. mas.n. aer air, gas; Gr. fem.n. monas unit, monad; M.L. fem.n.
Aeromonas gas (-producing) monad.
Short (rarely more than 3 microns), rod-shaped cells. Motile by means of polar flagella,
usually monotrichous; occasionally non-motile. Gram-negative. Heterotrophic, o.xidizing
various organic compounds. Carbohydrates fermented with the production of Ho , CO2 and
2,3-butylene glycol. Methyl red negative. Slow or no fermentation of lactose. The majority
of species thus far described are from water or are known to be pathogenic to marine and
fresh -water animals such as fish and amphibians.
Physiologically these organisms appear to be identical with certain species found in the
family Enter ohacteriaceae. The chief differences between the species in Aeromonas and those
in Paracolobactrum Borman, Stuart and Wheeler are found in the arrangement of their
flagella, in the less active fermentation of carbohydrates by the former, and in their patho-
genicity.
The type species is Aeromonas liquefaciens (Beijerinck) Kluyver and van Niel.
Key to the species of genus Aeromonas.
I. Motile.
A. Originally isolated from water.
* Prepared by Dr. S. F. Snieszko, U. S. Fish and Wildlife Service, Leetown via Kear-
neysville, West Virginia, August, 1953.
190
ORDER I. PSEUDOMONADALES
1. Not proven to be pathogenic for fish and amphibians.
1. Aeromonas liquefaciens.
2. Generally regarded as the cause of an infectious edema of carp and other fish.
2. Aeromonas punctata.
B. Originally isolated from a septicemia in frogs (red leg).
3. Aeromonas hydrophila.
II. Non-motile. Pathogenic for fish, particularly Salmonidae.
4. Aeromonas salmonicida.
1. Aeromonas liquefaciens (Beijerinck,
1900) Kluyver and van Niel, 1936. {Aero-
bacter liquefaciens Beijerinck, Cent. f. Bakt.,
II Abt., 6, 1900, 199; Kluyver and van Niel,
Zent. f. Bakt., II Abt., 94, 1936, 399.)
li.que.fa'ci.ens. L. v. liquefacio to
liquefy; L. part. adj. liquefaciens liquefying.
Description taken from Beijerinck (op.
cit., 6, 1900, 199) and from E. M. Miles and
A. A. Miles (Jour. Gen. Microbiol., 5, 1951,
299).
Rods, 0.4 to 0.8 by 1.5 to 3.0 microns,
with parallel sides and rounded ends. Fila-
ments common in "rough" colony forms.
Motile by means of a single polar flagellum
about 5 to 6 microns long. Gram-negative.
Gelatin stab: Liquefaction marked and
commonly saccate, good growth.
Horse blood agar colonies: 2 to 3 mm in
diameter, round, entire, raised, smooth,
moist, opaque, semi-translucent, grayish
white, forming a dirty brown-yellow colora-
tion after 3 to 5 days at room temperature ;
non-hemolytic.
Broth: Growth abundant, turbid, with a
moderate, readily disintegrable sediment
and delicate pellicle.
Loeffler's serum: Growth abundant, but
no digestion.
Litmus milk: Acid; coagulated; digested.
Potato: Growth abundant, moist and
glistening, light brown.
Indole is produced.
Nitrites but not ammonia produced from
nitrates (Beijerinck); ammonia produced,
presumably from peptones (Miles and
Miles) .
Methyl red test negative.
Citric acid and salts of citric acid may be
utilized as sole sources of carbon.
Ammonium sulfate, uric acid and aspara-
gine may be utilized as sources of nitrogen.
Catalase produced.
Hydrogen sulfide produced.
Urea not attacked.
Methylene blue reduced.
Starch hydrolyzed (Miles and Miles) ;
starch not hydrolyzed (Beijerinck).
Acid and gas from glucose, galactose,
fructose, mannose, maltose, sucrose, man-
nitol, glycerol and starch. Acid from lactose,
raffinose, inositol and sorbitol. Slight acid
from salicin at 22° C. but none at 37° C.
Glucose fermented with the production of
2,3-butanediol. Arabinose, rhamnose and
dulcitol not attacked.
Aerobic, facultative.
Temperature relations: Optimum, 37° C;
good growth on ordinary laboratory media
at 20° C.
Produces a characteristic black-rot in
hen eggs.
Pathogenic to mice, also to frogs, causing
a fatal bacteriemia.
Source: Found rarely in canal mud,
generally in certain marshes and swamps.
Habitat relationships uncertain. Those
that believe this organism to be identical
with Aeromonas punctata would associate it
with a disease of carp, eels and other fishes.
2. Aeromonas punctata (Zimmermann,
1890, emend. Lehmann and Neumann, 1896)
Snieszko, comb. nov. (Bacillus punctatus
Zimmermann, Bakt. unserer Trink- und
Nutzwasser, Chemnitz, 1, 1890, 38; Bac-
terium punctatum Lehmann and Neumann,
Bakt. Diag., 1 Aufl., 2, 1896, 238; Pseudo-
monas punctata Chester, Man. Determ.
Bact., 1901, 147; also see Schaperclaus,
Ztschr. f. Fischerei, 28, 1930, 289.)
punc.ta'ta. L. noun punctum a point, a
small hole; M.L. adj. punctata full of points.
Rods, 0.7 by 1.0 to 1.5 microns, occurring
singly, in pairs and in chains. Motile with
a single polar flagellum. Gram-negative.
Gelatin colonies: Small, circular, gray,
erose to filamentous, punctiform.
FAMILY IV. PSEUDOMONADACEAE
191
Gelatin stab: Crateriform liquefaction.
No pellicle.
Agar slant: Gray, smooth, filamentous.
Broth: Turbid with delicate pellicle.
Litmus milk: Acid; coagulated; pep-
tonized.
Potato: Brownish yellow to brownish
red color.
Indole is produced.
Hydrogen sulfide is produced.
Acid and gas from glucose broth (Leh-
mann and Neumann, op. cit., 1896, 238).
Aerobic, facultative.
Optimum temperature, between 25° and
30° C.
Distinctive characters: There does not
seem to be any real difference between this
organism and Aeromonas liquefaciens Kluy-
ver and van Niel. Schaperclaus (Fisch-
krankheiten, Braunschweig, 1 Aufi., 1935,
46; Ztschr. f. Fischerei, 37, 1939, 7) recog-
nizes definite varieties of this species:
some are non-pathogenic, others are patho-
genic to carp, and still others are patho-
genic to eels.
Source: From Chemnitz tap water (Zim-
mermann). Commonly found in water of
the River Main (Lehmann and Neumann,
op. cit., 238; also see op. cit., 7 Aufl., 2, 1927,
47).
Habitat: Found in water supplies, es-
pecially those in which carp, eels and other
fishes occur. Causes an infectious edema in
carp (Cyprinus) (Schaperclaus, op. cit.,
1930, 289; see Zent. f. Bakt., II Abt., 105,
1942, 49) and other fishes.
3. Aeromonas hydrophila (Chester,
1901) Stanier, 1943. {Bacillus htjdrophilus
fuscus Sanarelli, Cent. f. Bakt., 9, 1891,
222; Bacillus hydrophilus Chester, Manual
Determ. Bact., 1901, 235; Proteus hydro-
philus Bergey et al.. Manual, 1st ed., 1923,
211; Stanier, Jour. Bact., 46, 1943, 213.)
hy.dro'phi.la. Gr. noun hydor water; Gr.
philus loving; M.L. adj. hydrophilus water-
loving.
Description taken from Emerson and
Norris (Jour. Exp. Med., 7, 1905, 32) and
from E. M. Miles and A. A. Miles (Jour.
Gen. Microbiol., 5, 1951, 299).
Rods, 0.6 by 1.3 microns, occurring singly
and in chains. Motile, with a single polar
flagellum (Kulp and Borden, Jour, of Bact.,
U, 1942, 673). Gram-negative.
Gelatin colonies: Small, circular, gray,
translucent, stippled.
Gelatin stab: Napiform liquefaction.
Agar colonies: Whitish, raised, moist,
stippled.
Horse blood agar colonies: 2 to 3 mm in
diameter, round, entire, raised, smooth,
moist, semi-translucent, grayish white,
forming a dirty brown-yellow coloration
after 3 to 5 days at room temperature;
marked hemolysis.
Agar slant : Thin, whitish, glassy, spread-
ing, becoming yellowish and opalescent.
Broth: Turbid, with heavy pellicle.
Loeffler's serum: Growth abundant, but
no digestion.
Litmus milk: Acid; coagulated; pep-
tonized.
Potato: Yellowish brown, moist, slightly
raised.
Indole is produced.
Nitrites produced from nitrates.
Ammonium sulfate, uric acid and aspara-
gine may serve as sources of nitrogen.
Catalase produced.
Hydrogen sulfide produced.
Urea not attacked.
Methylene blue reduced.
Acid and gas from glucose, galactose,
fructose, mannose, maltose, sucrose, man-
nitol, glycerol and starch. Acid and gas
from salicin at 22° C. but not at 37° C. Acid
from glycogen and dextrin. Glucose fer-
mented with the production of 2,3-butane-
diol. Lactose, arabinose, raffinose, rham-
nose, dulcitol, sorbitol and inositol not
attacked.
In the fermentation of beet molasses,
Murphy, Watson, Muirhead and Barnwell
(Canad. Jour. Tech., 29, 1951, 375) found
this organism to yield up to 96 per cent of
theoretical 2,3-butariediol and acetoin.
This is a higher yield than the same authors
found for Aerobacter aerogenes.
Starch hydrolyzed.
Gas ratio HjrCOs = 1:4.71. Methyl red
negative, acetylmethylcarbinol positive,
indole negative, citrate positive (Speck and
Stark, Jour. Bact., U, 1942, 697).
Aerobic, facultative.
Optimum temperature, 37° C.
192
ORDER I. PSEUDOMONADALES
Produces a characteristic black-rot in
hen eggs.
Pathogenic for frogs, salamanders, fish,
mice, guinea pigs and rabbits, causing
hemorrhagic septicemia. Causes a hemor-
rhagic septicemia in snakes. In this case the
disease is transmitted by mites (Camin,
Jour, of Parasitol., 34, 1948, 345).
Source: Isolated from frogs dead of
septicemia (red leg).
Habitat: Water and infected fresh-water
animals.
4. Aeroinonas salmonicida (Lehmann
and Neumann, 1896) Griffin, 1954. (Bacillus
der Forellenseuche, Emmerich and Weibel,
Arch. f. Hyg.jBl, 1894, 1; Bacterium salmoni-
cida Lehmann and Neumann, Bakt. Diag.,
1 Aufl., 2, 1896, 240; see Mackie, Arkwright,
Pryce-Tannatt, Mottram, Johnston and
Menzies, Final Rept. of the Furunculosis
Committee, H. M. Stationery Office, Edin-
burgh, 1935; Griffin, Trans. Amer. Fish.
Soc, 83, (1953) 1954, 241.)
sal.mo.ni'ci.da. L. noun salmo, salmonis
salmon; L. v. suffix -cida from L. v. caedo to
cut, kill; M.L. fem.n. salmonicida salmon-
killer.
Description taken from Griffin (Trans.
Amer. Fish. Soc, 82 (1952) 1953, 129).
Rods, 1.0 by 1.7 to 2.0 microns, with
rounded ends, occurring singly, in pairs or
in chains. Non-motile. Gram-negative.
Gelatin stab: Crateriform to infundibuli-
form liquefaction in 1 to 3 days; complete
liquefaction in 7 days. Growth filiform,
beaded, best at top. Medium turns light
brown near the surface of old cultures.
Agar colonies : Circular, punctiform in 24
hours and 1 to 2 mm in diameter in 4 to 5
days, convex, entire, semi-translucent.
Colonies and medium turn brown in old
cultures.
Agar slant: Growth abundant, butyrous,
glistening, filiform, opaque to transparent,
odorless, colorless. A soluble, brown, me-
lanin-like pigment forms in 3 to 5 days.
A bright salmon-pink color develops when
i8-2-thienylalanine is present (Griffin,
Snieszko and Friddle, Jour. Bact., 65, 1953,
658).
Colonies developed on trypticase agar
quickly turn a violet-black color after the
addition of 1 per cent aqueous p-phenyl-
enediamine (Griffin, Proc. 52nd Gen. Meet-
ing, Soc. Ajner. Bact., Boston, 1952, 53;
also see Vet. Med., 48, 1953, 280).
Broth: Moderate to strong clouding; no
ring or pellicle; moderate, flocculent sedi-
ment. Medium may clear in the upper
layers and some growth may adhere to walls
of test tubes of old cultures.
Litmus milk: Slight and temporary acidi-
fication. Complete peptonization in one
week.
Rabbit blood agar: Beta-hemolysis in 2
days.
Indole not produced.
Nitrites produced from nitrates.
Ammonia produced in tryptic digest of
casein-yeast extract medium.
Hydrogen sulfide not produced.
Methyl red negative; acetylmethylcarbi-
nol not produced; sodium citrate does not
serve as a sole source of carbon.
Urea not attacked.
Acid and gas from glucose, fructose,
maltose, galactose, arabinose, mannose,
starch, dextrin, glycogen, salicin, esculin
and mannitol. Lactose, sucrose, xylose,
rhamnose, trehalose, melibiose, cellobiose,
raffinose, melizitose, inulin, amygdalin,
methyl glucoside, glycerol, erythritol,
adonitol, sorbitol and dulcitol not attacked.
Starch hydrolyzed.
Arginine and methionine are essential
for growth; asparagine and leucine are
highly stimulative while lysine is only
moderately so (unpublished data, Griffin).
Temperature relations: Optimum, be-
tween 20° and 25° C. Minimum, 6° C. Maxi-
mum, 34.5° C.
Aerobic, facultative.
Pathogenic for most fresh-water fish,
particularly those belonging to Salmonidae.
Source: From dead fish, of the family
Salmonidae, taken from a fish hatchery in
Southern Germany.
Habitat: Found in fresh-water lakes,
streams, rivers and fish ponds throughout
Europe and also in the United States and
Canada. Causes a furunculosis in infected
fish; also occurs in apparently normal fish.
FAMILY IV. PSEUDOMONADACEAE 193
Note: Species incertae sedis. At least tons) is listed in the Manual as producing
twelve additional species that appear to acid and gas from glucose and related
be identical with or closely related to the sugars. A monographic studj' of these or-
four species described in full have been ganisms is needed. The descriptions of some
reported in the literature. These were isola- species that were found before 1900 appear
ted from water, aquatic animals (midge to be as adequate as are the early descrip-
larvae, leeches, fishes) or dairy products. tions of Aeromonas liquefaciens. No attempt
It should also be noted that at least one has been made to determine which specific
plant pathogen {Xanthomonas proteamacu- epithet or epithets have priority.
Genus V. Photobacteriuni Beijerinck, 1889, emend. Breed and Lessel, 1954.*
(Beijerinck, Arch, neerl. d. Sci. exact, et natur., 23, 1889, 401; Photobader Beijerinck,
Troc. Sect. Sci., Kon. Akad. v. Wetensch., Amsterdam, 3, 1900, 352; ? Photomonas Orla-
Jensen {nomen nudum). Jour. Bact., 6, 1921, 271; Breed and Lessel, Antonie van Leeuwen-
hoek, 20, 1954, 60.)
Pho.to.bac.te'ri.um. Gr. noun phos light; Gr. ueut.dim.n. bacterium a small rod; M.L.
neut.n. Photobacterium light (-producing) bacterium.
Coccobacilli and occasional rods which, in the presence of glucose and asparagine, tend to
ramify in a manner analogous to that of bacteroids. Polar flagellate when motile. The type
species is normally non-motile but shows motility in young cultures (Kluyver). May or may
not liquefy gelatin. Produce acid, or acid and visible gas (H2 and CO2), from glucose and
other carbohydrates but not from lactose. Luminescent. Growth and luminescence best, or
even exclusively, on salt-water media containing 3 to 5 per cent salt. Found on dead fish
and other salt-water animals and in sea water. Reports by various authors indicate that the
luminescent^ coccoid and rod-shaped bacteria found living symbiotically in the tissues of
the phosphorescent organs of various cephalopods and deep-sea fishes also belong to this
genus. Other coccoid and rod-shaped luminescent bacteria found in the blood of Crustacea
and caterpillars appear to be parasitic or even pathogenic.
The type species is Photobacterium phosphoreum (Cohn) Ford.
Key to the species of genus Photobacterium. t
I. Coccobacilli which produce acid and gas from glucose.
A. Saprophytic on dead fish, Crustacea, meat and similar products.
1. Photobacterium phosphoreum.
B. Symbiotic, found in the photogenic organ of a cephalopod.
2. Photobacterium pierantonii .
IL Short rods which produce acid but no gas from glucose.
A. No growth at 37° C.
3. Photobacterium fischeri.
B. Grows well at 37° C.
4. Photobacterium harveyi.
1. Photobacterium phosphoreum fende het geneeskundig staatstoezicht in
(Cohn, 1878) Ford, 1927. {Micrococcus Nederland, 1878, 126; Bacterium phos-
phosphoreus Cohn, see letter addressed to phorescens Fischer, Cent. f. Bakt., 3, 1888,
J. Penn, Verzameling van stukken betref- 107; Photobacterium phosphorescens Bei-
* Prepared by Prof. Robert S. Breed and Mr. Erwin F. Lessel, Jr., Cornell University,
Geneva, New York, February, 1954.
t See Speucer (Jour. Gen. Microbiol., 13, 1955, 111) for a recent discussion of the classifica-
tion of this group.
194
ORDER I. PSEUDOMONADALES
jerinck, Arch, neerl. d. Sci. exact, et natur.,
£3, 1889, 401; Photobacter phosphoreum
Beijerinck, Folia Microbiologica, Delft,
4, 1916, 15; Ford, Textb. of Bact., 1927,
615.)
phos.pho're.um. Gr. v. phosphor eo to
bring light; M.L. adj. phosphoreus light-
bearing.
Description taken from Fischer (op. cit.,
1888, 107) and Beijerinck {op. cit., 1889,
401).
Coccobacilli, 0.5 to 2.0 microns; oc-
casional rods are 0.5 to 1.0 micron. In the
presence of glucose, especially glucose and
asparagine combined, some of the cells tend
to branch and to take the form of bac-
teroids. Frequently occur as zoogloeae.
Non-motile (Fischer); some cells show a
sluggish motility (Beijerinck); (Johnson,
personal communication, 1953, stated that
even electron micrographs failed to reveal
flagella) ; actively motile on suitable media
(Kluyver, personal communication, June,
1953) ; Leifson (personal communication,
July, 1953) reports that an occasional cell
of culture L342 from Delft shows mono-
trichous flagellation. Stain lightly with
aniline dyes. Gram-negative (Manual, 3rd
ed., 1930, 178).
Gelatin: No liquefaction.
Agar slant: Grayish white layer (Manual,
loc. cit.).
Broth: Slightly turbid with thin pellicle
(Manual, loc. cit.).
Potato: Ordinary acid potato, no growth;
neutralized with sodium phosphate, thin
brownish growth (Chester, Ann. Rept. Del.
Col. Agr. Exp. Sta., 9, 1897, 124).
Proteolytic enzymes not secreted.
Glucose, fructose, maltose and galactose
are anaerobically fermented with the pro-
duction of gas. This is a butanediol fermen-
tation that produces H2 and CO2 (Kluyver,
personal communication, 1953).
Aerobic, facultatively anaerobic.
Minimum temperature, between 5° and
10° C.
Quality of luminescence: Bluish green.
Salt tolerance: To assure phosphorescence
and good growth, the osmotic tension of
inorganic salt solutions used for cultivation
should be equivalent to that produced in a
3 per cent sodium chloride solution.
Distinctive characters: Coccoid bacteria
which do not liquefy gelatin and which
produce acid and gas from glucose but not
from lactose. In the presence of glucose,
especially when combined with asparagine,
the cells swell up greatly and lose their
luminescent property. Luminescence on
organic matter occurs only when there is a
sufficient proportion of inorganic salt.
Comments: Several publications which
antedate that of Fischer (op. cit., 1888, 107)
allude to the fact that the binomial Bac-
terium phosphorescens might have been
effectively published earlier than 1888. One
reference (Anonymous, Nature, 35, 1886-
1887, 377) cites Hermes, the Director of the
Berlin Aquarium, as having published an
article in which he describes and names as
Bacteriian phosphorescens a luminescent
organism obtained from a specimen of cod
(Gadus callarias) at the Berlin Aquarium;
this was the same organism which Fischer
secured from the Berlin Aquarium and
which he named Bacterium phosphorescens.
A second reference (Ludwig, Cent. f. Bakt.,
2, 1887, 404) states that Hermes demon-
strated before the Berlin Society the phos-
phorescent bacterium from the Berlin
Aquarium under the name Bacterium phos-
phorescens. Other references (Anonymous,
Gesell. deutsch. Naturforsch. u. Aerzte,
Tageblatt, 60, 1887, 77 and 254) showed
that Hermes used this organism several
times for demonstration purposes in the
Aquarium and before the Society. As
Hermes' publication has not been found, and
as the binomial Bacterium phosphorescens is
not effectively published in anj- of the three
references given directly above, Fischer is
credited here as the author of this binomial.
Considerable confusion exists in the
literature concerning this species, most of
which can be elucidated by the following:
(1) Fischer (Ztschr. f. Hyg., 2, 1887, 54-92)
described an organism, isolated from sea
water from the West Indies, which he named
Bacillus phosphorescens; (2) a second species
of phosphorescent bacteria, obtained from
the Berlin Aquarium, was described, but
not named, by Fischer in a supplement to
the work cited above (ibid., 92-95); Leh-
mann (Cent. f. Bakt., 5, 1889, 785) also
described an organism obtained from the
FAMILY IV. PSEUDOMONADACEAE
195
Berlin Aquarium, and he states that it is
identical with the one which Fischer ob-
tained from this same source; (3) in a later
paper (Cent. f. Bakt., S, 1888, 107), Fischer
identified the second species, as well as
phosphorescent bacteria that he isolated
from dead fish from the Baltic and North
Seas, as Bacterium phosphorescens. Some
authors, e.g. Lehmann and Neumann (Bakt.
Diag., 1 Aufl., 2, 1896, 198; and other edi-
tions), Migula (Syst. d. Bakt., 2, 1900, 433)
and Chester (Man. Determ. Bact., 1901,
181), when referring to Bacterium phos-
phorescens Fischer, quote the supplement
to Fischer's paper in the Ztschr. f. Hyg., 2,
1887, 92, as the source of the name Bacterium
phosphorescens, whereas the first use of this
binomial by Fischer was in the Cent. f.
Bakt., 3, 1888, 107. This failure to give an
exact reference has caused confusion in
later publications, especially since Bacillus
phosphorescens is the only binomial pro-
posed, or even used, by Fischer in his paper
published in the Ztschr. f. Hyg., 2, 1887,
54-95, which also contains a description of
the organism he later identified as Bacterium
phosphorescens. Still other writers (Gorham,
in Dahlgren, Jour. Franklin Inst., 180, 1915,
517 and insert following 714) have used the
name Bacillus phosphorescens in lieu of
Bacterium phosphorescens, thus augmenting
the confusion.
Relationships to other species of bacteria:
Beijerinck regards Photobacterium phos-
phorescens Beijerinck as identical with
Micrococcus phosphoreus Cohn (Folia Micro-
biologica, Delft, 4, 1916, 15, footnote 4)
but different from Photobacterium pfleugeri
Ludwig (Arch, neerl. d. Sci. exact, et natur.,
24, 1891, 369).
Source: Isolated from cod (Gadus cal-
larias) from the Baltic Sea; also found on
haddock {Melanogrammus aeglifinus) and on
lobster (Homarus sp.).
Habitat: Found on dead fish and in sea
water, so far as known.
2. Photobacterium pierantonii (Zir-
polo, 1918) Krassilnikov, 1949. {Micrococcus
pierantonii Zirpolo, Boll. del. Societa dei
Natural, in Napoli, 31, (1918) 1919, 75;
Photobacterium pierantonii, incorrectly
ascribed to Bergey et al. by Krassilnikov,
Guide to the Bacteria and Actinomycetes,
Izd. Akad. Nauk, U.S.S.R., Moskau, 1949,
514.)
pie.ran.to'ni.i. M.L. gen. noun pier-
antonii of Pierantoni; named for Prof. U.
Pierantoni, an Italian scientist.
Original description supplemented by
material taken from Meissner (Cent. f.
Bakt., II Abt., 67, 1926, 204).
Cocci, 0.8 micron in diameter, and short
rods, 0.8 by 1.0 to 2.0 microns. Occasionally
vacuolated. Motile or non-motile, the
motile cells possessing a single flagellum or
a tuft of 2 to 4 flagella. Gram-negative.
Gelatin colonies: Circular, luminous.
Gelatin stab: No liquefaction.
Sepia agar colonies: Circular, white,
convex, smooth and serrate with an intense,
greenish luminescence.
Egg glj'cerol agar slant: Yellowish green,
luminous streak.
Broth: Turbid.
Indole not produced.
Acid and gas from glucose and maltose.
Some strains produce acid but no gas from
lactose and sucrose.
Aerobic.
Optimum temperature, 33° C.
Optimum pH for growth, 9.0. No growth
at pH 5.0.
Optimum pH for luminescence, 8.0. No
luminescence at pH 5.0.
Quality of luminescence: Greenish.
Source: Isolated from the photogenic
organ of the cephalopod Rondeletia minor.
Habitat: Apparently found only in
Rondeletia minor but may also be found in
closely related species.
3. Photobacterium fischeri Beijerinck,
1889. (Einheimischer Leuchtbacillus, Fis-
cher, Cent. f. Bakt., 3, 1888, 107; Beijerinck,
Arch, nderl. d. Sci. exact, et natur., 23, 1889,
401; Vibrio fischeri Lehmann and Neumann,
Bakt. Diag., 1 Aufl., 2, 1896, 342; Achromo-
bacter fischeri Bergey et al.. Manual, 3rd ed.,
1930, 220.)
fisch'er.i. M.L. gen. noun ^sc^e/z of Fis-
cher; named for Prof. Bernhard Fischer,
one of the earliest students of luminescent
bacteria.
Description taken from Fischer (op. cit.,
1888, 107), Beijerinck (op. cit., 1889, 401) and
196
ORDER I. PSEUDOMONADALES
Johnson and Shiink (Jour. Bact., 31, 1936,
589).
Short, thick rods, 0.4 to 0.8 by 1.0 to 2.5
microns, with rounded ends, occurring
singly and in pairs. Occasional rods slightly
curved, ends slightly pointed. Not encap-
sulated. Motile. Johnson, Zworykin and
Warren (Jour. Bact., 46, 1943, 167) made
pictures with the electron microscope of
a culture which they identified with this
species; the organism showed a tuft of polar
flagella. Gram-negative.
Sea-water gelatin colonies : After 48 hours,
colonies small (less than 0.5 mm in diam-
eter), circular, entire, homogeneous, with
slight liquefaction.
Sea-water gelatin stab: Slight, infundi-
buliform liquefaction, sometimes slightly
beaded, tending to become crateriform in
old cultures.
Nutrient sea-water agar colonies: Small,
circular, smooth, entire, slightly raised,
homogeneous, iridescent. Old colonies be-
come yellowish with margins slightly ser-
rate.
Sea-water agar slant: Growth abundant,
grayish to jellowish, smooth, viscous,
homogeneous, iridescent.
Growth on autoclaved fish: Moderate,
grayish to yellowish, smooth, glistening,
luminescent, no odor of putrefaction.
Sea water containing 0.2 per cent peptone :
Moderate growth, mostly near the surface;
very thin pellicle; sediment found in old
tubes.
Milk: No growth. Milk with 2.8 per cent
sodium chloride : Slight growth and lumines-
cence, but no action on the milk.
Potato plugs resting on cotton saturated
in sea water: Growth fairly abundant,
spreading, slightly brownish, luminous.
Blood serum: No growth.
Indole not produced.
Hydrogen sulfide is produced.
Acid but no gas from glucose (Gorham,
in Dahlgren, Jour. Franklin Inst., 180,
1915, 517 and insert following 714). Acid
from glycerol, fructose, galactose, mannose,
maltose, cellobiose, dextrin and salicin.
No acid or gas from lactose, sucrose, arabi-
nose, xylose, fucose, rhamnose, trehalose,
raffinose, glycogen, inulin, adonitol, dulci-
tol, inositol, sorbitol, erythritol, arabitol
or alpha-methyl-glycoside.
Starch hydrolysis is doubtful or verj'
slight.
Decarboxjlates glutamic acid to form
7-aminobutyric acid and CO2 ; decarboxjd-
ates lysine (Pearson, Jour. Cell, and Comp.
Physiol., 41, 1953, 65).
Alanine, arginine, aspartic acid, glutamic
acid and threonine are capable of serving
as sole nitrogen sources for this organism
(Pearson, Jour. Tenn. Acad. Sci., 27, 1952,
229).
Nitrites produced from nitrates.
Ammonia produced in peptone media.
Aerobic, facultatively anaerobic.
Temperature relations: Optimum, be-
tween 25° and 28° C. Minimum, between 5°
and 10° C. No growth at 37° C.
Optimum temperature for luminescence,
28° C. Weak at 10° C., none at 5° nor at 37° C.
Optimum pH for luminescence, between
7.4 and 7.8; less intense at 7.0 and 8.2.
Fischer (Erg. d. Plankton Expedition d.
Humboldt-Stiftung, 4, 1894) noted that
this organism grows best in alkaline rnedia.
Quality of luminescence: Orangish, main-
tained for 5 to 8 weeks (Beijerinck); green-
ish (Johnson and Shunk). Luminescence
favored by the presence of glycerol in the
medium.
Salt tolerance: The osmotic tension of
inorganic salt solutions used as media for
this species must be equivalent to that
produced in a 2.8 to 3.0 per cent sodium
chloride solution to assure luminescence and
good growth.
Not pathogenic for white rats.
Distinctive character: Luminescence on
organic matter occurs only when there is a
suflftcient proportion of inorganic salt pres-
ent.
Source: Isolated from sea water at Kiel
and from herring.
Habitat: Frequentlj^ found on dead fish,
Crustacea and other salt-water animals and
in coastal sea water. Phosphorescent bac-
teria also occur on meat and even on sol-
dier's wounds where they produce no known
harmful effects. No food poisoning has been
traced to meat on which these organisms
have grown (Niven, Circular No. 2, Ameri-
can Meat Inst. Foundation, 1951, 1-11).
FAMILY IV. PSEUDOMONADACEAE
197
4. Photobacteriuni harveyi (Johnson
and Shunk, 1936) Breed and Lessel, 1954.
(Achro)nobacter harveyi Johnson and Shunk,
Jour. Bact., 31, 1936, 587; Breed and Lessel,
Antonie van Leeuwenhoek, 20, 1954, 61.)
har'vey.i. M.L. gen. noun harveyi of
Harvey; named for E. N. HarveJ^
Description taken from Johnson and
Shunk (op. cif., 1936, 587).
Rods, .0.5 to 1.0 by 1.2 to 2.5 microns,
occurring singly or in pairs, with rounded
ends. Occasionally slightly curved; ends
occasionally slightly pointed. Non-spore-
forming. Not encapsulated. Motile by
means of a single, polar flagellum 2 to 3
times the length of the cell. Gram-negative.
Sea-water gelatin colonies: After 24
hours at 20° C, circular, about 1.5 mm in
diameter or larger, margin slightly undu-
late, sunken due to the beginning of lique-
faction, interior somewhat zonate; colonies
surrounded by a halo of numerous small
secondary colonies, circular and finely
granular. In crowded plates a large number
of gas bubbles are formed. Luminescent.
Sea-water gelatin stab: Rapid saccate
liquefaction complete in 5 daj^s at 22° C.
Abundant flocculent sediment.
Sea-water agar colonies: Mostly very
large, 6 to 8 cm in diameter in 24 hours, flat,
highly iridescent, circular with undulate
margin, or composed of narrow and close or
wide filamentous growth. Occasionally
small colonies appear that are circular, with
entire or slightly undulate margin, often
producing irregular secondary growth, sur-
face always smooth. Luminescent.
Sea-water agar slant: Growth abundant,
spreading, grayishly viscous, homogeneous,
iridescent, the medium becoming rapidly
alkaline w^hen inoculated at an initial pH
of 7.0. With fish decoctions added to the
medium, luminescence is much brighter
and growth becomes brownish after several
days.
Growth on autoclaved fish: Abundant,
smooth, glistening, yellowish, becoming
dirty brown after several days. Mild putre-
factive odor. Luminescence very brilliant.
Sea water containing 0.2 per cent peptone :
Abundant uniform turbidity, thin pellicle,
sediment accumulating over a period of
several days. Luminescence at surface only
unless the tube is shaken.
Milk, with or without the addition of 2.8
per cent salt: No growth.
Potato plugs resting on cotton saturated
with sea water: Growth slight, somewhat
spreading, slightly brownish. Luminous.
Indole produced (Gore's method).
Hydrogen sulfide is produced (ZoBell
and Fantham method).
Fixed acid from glucose, fructose, man-
nose, galactose, sucrose, maltose, mannitol,
dextrin, glycogen, trehalose, cellobiose;
slowly from salicin. Non-fixed acid from
melezitose; slight acid from sorbitol, disap-
pearing in 24 hours. No acid from glycerol,
xylose, arabinose, dulcitol, inositol, adoni-
tol, erythritol, arabitol, lactose, raffinose,
rhamnose, fucose or alpha methyl glucoside.
Starch agar: Wide zone of hydrolysis.
Nitrites produced from nitrates.
Ammonia produced in peptone media
(Hansen method).
Aerobic, facultatively anaerobic.
Temperature relations: Optimum, be-
tween 35° and 39° C. Abundant growth be-
tween 22° and 25° C.
Optimum temperature for luminescence,
between 20° and 40° C.
Optimum pH for luminescence, between
pH 7.4 and 7.8.
Quality of luminescence (to completely
dark-adapted eyes) : Yellowish green to
green on fish and typically green on sea-
water agar or gelatin.
Not pathogenic for white rats or amphi-
pods.
Distinctive character: Luminescence not
favored by the presence of glycerol in the
medium.
Source: Isolated from a dead amphipod
(Talorchesda sp.) at Woods Hole, Massa-
chusetts.
Habitat: Sea water.
Note: Species incertae sedis. Additional
luminescent bacteria which probably be-
long in this genus have been reported in
the literature. However many of the de-
scriptions are not adequate enough to
permit the determination of the identity
and relationships of these organisms.
ORDER I. PSEUDOMONADALES
Genus VI. Azotonionas Stapp, 1940.*
(Stapp, Zent. f. Bakt., II Abt., 102, 1940, 18; not Azotomonas Orla-Jensen,
Cent. f. Bakt., II Abt., U, 1909, 484.)
A.zo.to.mo'nas. Gr. azous without life; Fr. noun azote nitrogen; Gr. fem.n. monas unit,
monad; M.L. fem.n. Azotomonas nitrogen (-fixing) monad.
Rod- to coccus-shaped cells. Motile by means of 1 to 3 polar fiagella. No fat-like reserve
food granules in the cells. Chemo-heterotrophic. Produce acid and sometimes gas from glu-
cose and other sugars and alcohols. Many carbon compounds other than sugars are used as
sources of energj-. Indole is produced. Aerobic. Active in the fixation of atmospheric nitro-
gen. Found in soil.
The type species is Azotomonas insolita Stapp.
Key to the species of genus Azotonionas.
I. Acid and gas from glucose.
II. Acid but no gas from glucose.
1. Azotomonas insolita.
2. Azotomonas fluorescens.
1. Azotomonas insolita Stapp, 1940.
(Abstracts of Communications, Third In-
ternat. Congr. for Microbiol., Sect. VIII,
1939, 306; abst. in Proc. Soil Sci. Soc. of
America, 4, 1939, 244; Zent. f. Bakt., II
Abt., 102, 1940, 1.)
in.so'li.ta. L. adj. insolitus unusual.
Coccoid rods 0.6 to 1.2 by 0.6 to 1.8 mi-
crons. Motile by means of 1 to 3 polar
flagella. Gram-negative.
Gelatin: No liquefaction.
Agar colonies: Flat, whitish, entire,
weakly fluorescent.
Agar slant: Glistening, white growth.
Broth: Strongly turbid; sediment; pel-
licle.
Milk: Unchanged.
Potato: Growth somewhat dry, not slimy,
dirty gray, spreading.
Hydrogen sulfide is produced.
Acid and gas from adonitol, arabinose,
dextrin, glucose, galactose, glycerol,
inositol, lactose, fructose, maltose, man-
nose, mannitol, raffinose, rhamnose, salicin,
sorbitol, starch, sucrose and xylose.
Starch is hydrolyzed.
Nitrites produced from nitrates.
Atmospheric nitrogen is fixed.
Ammonium salts are utilized.
Aerobic.
Temperature relations: Optimum, be-
tween 25° and 30° C. Minimum, between
7.0° and 9.5° C. Maximum, 48° C. Good
growth at 37° C. Thermal death point,
60° C.
Limits of pH, 3.3 to 9.5.
Source: Isolated from a mixture of chop-
ped cotton husks and rice hulls.
Habitat: Soil.
2. Azotonionas fluorescens Krassilni-
kov, 1947. (Quoted from Krassilnikov, Guide
to the Bacteria and Actinomycetes, Izd.
Akad. Nauk, U.S.S.R., Moskau, 1949, 420.)
flu.o.res'cens. L. noun fluor a flux; M.L.
v. fluoresco to fluoresce; fluor-spar, a flux-
ing mineral which is fluorescent; M.L.
part. adj. fluorescens fluorescing.
Translated by Dr. A. Petraitis, New York
State Experiment Station, Geneva, New-
York.
Rod-shaped cells, 0.5 to 0.8 by 2.0 to 5.0
microns, which become shorter in old cul-
tures. Motile by means of one to three
polar flagella. Gram-negative.
Gelatin: Slow liquefaction.
Inorganic media with or without nitro-
gen: Good growth.
Colonies are wide, smooth and glistening.
A slightly yellowish or violet fluorescent
pigment is produced which diffuses through
the medium.
Milk: Peptonized.
* Rearranged by Dr. A. W. Hofer, New York State Experiment Station, Cornell Univer-
sity, Geneva, New York, November, 1953.
FAMILY IV. PSEUDOMONADACEAE
199
Acid but no gas from various sugars and
alcohols.
Starch is hj'drolyzed.
Fixes nitrogen.
Nitrites not produced from nitrates.
Aerobic.
Source: Isolated from soil.
Habitat: Soil.
Genus VII. Zymomonas Kluyver and van Niel, 1936*
(Kluj'ver and van Niel, Zent. f. Bakt., II Abt., H, 1936, 399; Saccharomonas
Shimwell, Jour. Inst. Brewing, 56 (N.S. 47), 1950, 179.)
Zy.mo'mo.nas or Zy.mo.mo'nas. Gr. noun zyme leaven, ferment; Gr. noun monas a
unit, monad; M.L. fem.n. Zymomonas fermenting monad.
Rod-shaped cells, occasionally ellipsoidal. Motile cells are lophotrichous. Anaerobically
ferment glucose with the production of carbon dioxide, ethyl alcohol and some lactic acid.
Found in fermenting beverages such as pulque, palm juice and beer.
The type species is Zymomonas mobilis (Lindner) Kluyver and van Niel.
1. Zymomonas mobilis (Lindner, 1928)
Kluyver and van Niel, 1936. (Termobac-
terium mobile Lindner, Atlas d. Mikrosk
Grundl. d. Garungsk., 3 Aufl., 2, 1928, Taf
68; also see Lindner, 50 Jubilaumsber
Westpreuss. Bot.-Zool. Vereins, 1928, 253
Pseudomonas lindneri Kluyver and Hoppen
brouwers, Arch. f. Mikrobiol., £, 1931, 259
Zymomonas mobile (sic), Kluyver and van
Niel, Zent. f. Bakt., II Abt., 94, 1936, 399
Saccharomonas lindneri Shimwell, Jour.
Inst. Brewing, 56 (N. S. 47), 1950, 179.)
mo'bi.lis. L. adj. mobilis movable, motile.
Short rods with rounded ends, 1.4 to 2.0
by 4.0 to 5.0 microns. Occur usually as pairs
with a central constriction and rarely as
short chains. Motile with polar flagella.
Gram-negative.
Peptone gelatin: Poor growth.
Peptone agar: Poor growth.
Wort agar: White, round, raised colonies
1 mm in diameter. Good growth. Still better
growth where 2 per cent sucrose or yeast
extract with sucrose is added. Chalk may be
added to neutralize acid.
Wort gelatin stab: Uniform growth in
stab; no surface growth. No liquefaction.
Broth: Poor growth in peptone or yeast
extract broth unless sugars are added.
Carbon dioxide, ethyl alcohol and some
lactic acid produced from glucose and
fructose but not from mannose. Ferments
sucrose usually after a somewhat prolonged
lag period. May produce as much as 10 per
cent alcohol.
Catalase-positive.
Anaerobic, although with a certain
oxygen tolerance of aerobic growth in the
presence of fermentable sugars.
Optimum temperature, 30° C.
Distinctive character: Apart from the
production of some lactic acid, the fermen-
tation resembles the alcoholic fermentation
produced by j'easts.
Source : Isolated from the fermenting sap
(pulque) of Agave americana in Mexico and
from fermenting palm juice {Arenga sac-
charifera) in Sumatra and Java (Roelofsen,
Natuurwetenschappelijk Tijdschrift voor
Ned-. Indie, 101, 1941, 374).
Habitat: Found in fermenting plant
juices in tropical countries (Mexico and
Indonesia).
2. Zymomonas anaerobia (Shimwell,
1937) Kluyver, comb. nov. (Achromobacier
anaerobium Shimwell, Jour. Inst. Brewing,
43, 1937, 507; Saccharomonas anaerobia Shim-
well, op. cit., 56 (N. S. 47), 1950, 179 (type
species of genus Saccharomonas Shimwell).)
an.a.e.ro'bi.a. Gr. pref. an not; Gr.
noun aer air; Gr. noun bins life; M.L. adj.
anaerobius not living in air.
Rods, 1.0 to 1.5 by 2.0 to 3.0 microns,
plump with rounded ends. Cells occasionally
clump together to form rosette-like clusters.
Young cells actively motile with lophotri-
chous flagella, old cells become non-motile.
Not encapsulated. Gram-negative.
Glucose-beer-gelatin stab: Dense, fili-
form to beaded growth in stab; no surface
growth. No liquefaction.
* Prepared by Prof. A. J. Kluyver, Technische Hogeschool, Delft, Holland, December,
1953.
200
ORDER I. PSEUDOMONADALES
Glucose-beer-agar colonies: When in-
cubated in CO2 , irregularly circular, entire,
convex, about 1 mm in diameter, cream-
colored by reflected light, brown by trans-
mitted light, thinly butyrous, granular.
Glucose-beer-agar slant: Normally there
is no growth although there may be a slight
growth after prolonged incubation. Fili-
form or beaded, creamy white, thinly
butyrous, non-adherent growth when incu-
bated in CO2 .
Glucose-beer-agar stab: Dense, filiform
to beaded growth in stab; no surface growth.
Yeast extract, sugar-free beer: No
growth.
Beer, 2 per cent glucose: Densely turbid,
later becoming clear with a heavy sediment.
Yeast extract glucose broth: Growth only
in deep medium; slight deposit on walls of
tube; dense sediment at bottom.
Indole not produced.
Glucose and fructose readily fermented;
maltose, sucrose, lactose and ethyl alcohol
not attacked.
Acetylmethylcarbinol and diacetyl not
produced.
Nitrites not produced from nitrates.
Anaerobic, microaeroduric (not micro-
aerophilic) .
Temperature relations: Optimum, 30° C.;
thermal death point, 60° C. for 5 minutes.
pH range for growth, 3.4 to 7.5.
Distinctive characters: Does not grow in
any medium unless glucose or fructose is
present. 8himwell {loc. cit.) recognizes a
non-motile variety of this species. A related
or perhaps identical species has been de-
scribed as the cause of "cider sickness" in
England (see Barker, Ann. Rept. Nat.
Fruit and Cider Inst. Long Ashton, 1948).
A comparative study of cultures of Zymo-
monas mobilis, Z. anaerobia and the cider
organism made in 1951 shows that these
organisms are closely related. Z. anaerobia
did show fermentation of sucrose although
the cider organism did not show this fer-
mentation (Kluyver, personal communica-
tion).
Source: Isolated from beer, from the
surface of brewery yards and from the
brushes of cask-washing machines.
Habitat: Plant juices or extracts con-
taining glucose.
Note: Species incertae sedis. Additional
species which probably belong in this genus
but which have not been well described have
been reported from beer and cider.
Genus VIII. Protaminobacter den Dooren de Jong, 1926 *
(Bijdrage tot de kennis van het mineralisatieproces. Thesis, Rotterdam, 1926, 159.)
Pro.ta.mi.no.bac'ter. Gr. sup.adj. protus first; M.L. noun aminum an amine; M.L.
mas.n. bacter masculine form of Gr. neut.n. bactrum rod or staff; M.L. mas.n. Protaminobacter
protamine rod.
Cells motile or non-motile. Capable of dissimilating alkylamines. Pigmentation frequent.
Soil or water forms.
Recently Slepecky and Doetsch (Bact. Proc, 54th Gen. Meeting, Soc. of Amer. Bact.,
1954, 44) have isolated 23 fresh cultures of polar flagellate organisms that utilize alkyl-
amines. Of these, one resembled a known species of Protaminobacter, but all showed the
general characters of organisms placed in the genus Pseudomonas. The authors question
the recognition of the genus Protaminobacter on a biochemical basis only.
The type species is Protaminobacter albofiavus den Dooren de Jong.
Key to the species of genus Protaminobacter.
I. Non-motile. Gelatin colonies light yellow to colorless.
1. Protaminobacter albofiavus.
II. Motile. Gelatin colonies red.
2. Protaminobacter ruber.
* Prepared by Prof. D. H. Bergey, Philadelphia, Pennsylvania, June, 1929; further re-
vision by Prof. Robert S. Breed, New York State Experiment Station, Geneva, New York,
July, 1953.
FAMILY IV. PSEUDOMONADACEAE
201
1. Protaminobacter alboflavus den
Dooren de Jong, 1926. (Thesis, Rotterdam,
1926, 159; also see Cent. f. Bakt., II Abt.,
71, 1927, 218.)
al.bo.fla'vus. L. adj. alhus white; L.
adj. flavus j-ellow; M.L. adj. alboflavus
whitish yellow.
Rods. Non-motile. Gram-negative.
Gelatin colonies: Circular, dry, light
yellow or colorless.
Gelatin stab: No liquefaction.
Agar colonies: Circular, opaque, pigment
bright red, yellow, light gray or colorless.
Amine agar colonies: Circular, white to
dark yellow.
See Table I for list of organic substances
utilized.
Catalase produced.
Aerobic, facultative.
Optimum temperature, 30° C.
Distinctive characters : The author recog-
nizes four varieties of this species which he
differentiates on the basis of organic sub-
stances attacked (see Table) and pigment
produced. Variety a shows light yellow
growth on gelatin, bright red on agar and
yellow on amine agar. Variety' /3 is light 3'el-
low on gelatin, yellow on agar and dark
yellow on amine agar. Variety y is light
yellow on gelatin, light gray on agar and
yellow on amine agar. Variety 5 is colorless
on gelatin and agar and white on amine
agar.
Habitat: Soil and water.
2. Protaminobacter ruber den Dooren
de Jong, 1926. (Thesis, Rotterdam, 1926,
159; also see Cent. f. Bakt., II Abt., 71, 1927,
218.)
ru'ber. L. ruber red.
Rods. Motile with a single polar flagellum
(Weaver, Samuels and Sherago, Jour. Bact.,
35, 1938, 59). Gram-negative.
Gelatin colonies: Circular, red, dry.
Gelatin stab: No liquefaction.
Agar colonies: Circular, red, opaque.
Amine agar colonies: Circular, dark red.
The following organic acids are attacked:
Acetic, lactic, |3-oxybutyric, glycerinic,
succinic, malonic, formic, methyl formic,
glutaric, maleinic, fumaric, malic, tartaric,
citric and quinic.
The following amino compounds are
Table I. — Organic Substances Utilized as a
Source of Carbon by Varieties (biotypes) of
Protaminobacter alboflavus
Organic acids:
Acetic
Valerianic
a-crotonic
Undecyclic
Lactic
/3-oxybutyric
Succinic
Formic
Glutaric
Adipic
Fumaric
Malic
Tartaric
Citric
/3-phenj'lpropionic
Quinic
Amino compounds:
a-alanin
a-aminocapronic acid
Leucin
Propionamid
Capronamid
Uric acid
Hippuric acid
Alcohol :
Ethyl
Sugar :
Glucose
Amines :
Ethyl
Diethyl
Propyl
Isopropyl
Dipropyl
Tripropyl
Butyl
Isobutyl
Diisobutyl
Amyl
Diamyl
Ethanol
Glucosamin
Benzyl
«
7
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4-
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202 ORDER I. PSEUDOMONADALES
attacked: Sarcosiu, betain, hippuric acid, Catalase produced,
asparagine, propionamid, capronamid, lac- Aerobic, facultative,
tainid, succinamid, allantoin and uric acid. Optimum temperature, 30° C.
Glucose fermented. Habitat: Soil and v/ater.
Genus IX. Alginomonas Thj^tta and Kdss, 1945.*
(Thj0tta and K&ss, Norske Videnskaps-Akad., Oslo, I Mat.- Naturv. Klasse, No. 5, 1945,
17; also see lass. Lid and MoUand, ibid., No. 11, 1945, 8.)
Al. gi.no. mo 'nas. L. fem.n. alga seaweed; M.L. adj. alginicus pertaining to alginic acid
from seaweed; Gr. noun nionas a unit, monad; M.L. fem.n. Alginomonas alginic-acid (-de-
composing) monad.
Coccoid rods which are motile with one to four polar fiagella. Gram-negative. Fluorescent.
Gelatin is usually liquefied. Carbohydrates are not utilized. Citric acid is not used as a sole
source of carbon. Alginic acid is decomposed. Found on algae and in sea water and soil.
As the type of flagellation has not been determined for all of the species here included in
the genus, it may be found later that some of these species do not belong in Alginomonas
as here defined.
The type species is Alginomonas nonjermentans K&ss et al.
Key to the species of genus Alginomonas.
I. Gelatin is liquefied.
A. Grow on potato.
1. Gray to grayish brown growth on potato.
1. Alginomonas nonjermentans.
2. Pinkish or ivory-colored growth on potato.
a. Pink growth on potato.
2. Alginomonas terrestralginica.
aa. Ivory-colored growth on potato.
3. Alginomonas alginovora.
B. No growth on potato.
4. Alginomonas fucicola.
II. Gelatin not liquefied in seven days.
5. Alginomonas alginica.
1. Alginomonas nonfermentans K&ss Potato: Abundant, grayish brown
et al., 1945. (K&ss, Lid and Molland, Norske growth.
Videnskaps-Akad., Oslo, 1 Mat. -Naturv. Indole not produced.
Klasse, No. 11, 1945, 9.) Hydrogen sulfide is produced.
non.fer.men'tans. L. prefix non- not, Alginic acid is decomposed without the
non-; L. y. fermento to ferment; M.L. part. production of acid or gas.
adj. nonjermentans non-fermenting. Carbohydrates are not utilized.
Small, coccoid rods. Motile with one to (.^^^.j^ ^^j^ ^^^^^^ ^e used as a sole source
four polar fiagella. Gram-negative. , oarbon
Good growth on ordinary media. ^t-, .. i i j- -^ j.
^ . ^. -r ■ r ^- Nitrites produced from nitrates.
Gelatin: Liquefaction.
Agar colonies: Smooth, fluorescent. Aerobic.
Broth: Turbid; sediment; no pellicle. Grows at 37° C.
Litmus milk: Coagulated; peptonized; Chemical tolerance: No growth at pH
reduced. 9.6.
♦Prepared by Prof. Th. Thj0tta, Microbiological Institute, University of Oslo, Oslo,
Norway, January, 1955.
FAMILY IV. PSEUDOMONADACEAE
203
No growth in 6 per cent sodium chloride
broth.
Source: Five strains were isolated from
soil.
Habitat: Presumably soil.
2. Alginomonas terrestralginica
(Waksman et al., 1934) K^ss et al., 1945.
{Bacterium terrestralginicum Waksman,
Carey and Allen, Jour. Bact., 28, 1934, 217;
Kiss, Lid and MoUand, Norske Viden-
skaps-Akad., Oslo, 1 Mat.-Naturv. Klasse,
No. 11, 1945, 9.)
ter.res.tral.gi'ni.ca. L. noun terrestris
land, earth; M.L. adj. alginicus pertaining
to alginic acid from seaweed; M.L. adj.
terrestralginicus land-alginic; presumably
intended to mean an alginic bacterium from
the soil.
Long rods, 1.0 to 1.5 by 1.5 to 2.5 microns,
with somewhat rounded ends, usually oc-
curring singly but also in pairs, occasionally
in chains of shorter rods. Motile. Granular.
Gram-negative.
Alginic acid plate: Colonies small, whitish
in appearance with a slight metallic sheen.
Alginic acid liquid medium: Medium at
first clouded. Later a pellicle is formed on
the surface of the medium; it is soon broken
up due to active gas formation. Reaction of
medium becomes slightly alkaline.
Gelatin medium: Slow growth throughout
stab, slow liquefaction at surface of medium
at 18° C.
Agar liquefaction: None.
Glucose broth: Abundant turbidity; some
sediment; no pellicle; slightly fluorescent.
Litmus milk: Acid; milk coagulated;
only limited digestion of coagulum.
Potato: Abundant, pinkish, compact,
dry growth on surface of plug, the rest of
the plug becoming gray with a tendency to
darkening.
Starch plate: Limited growth along
streak; no diastase.
Aerobic to facultatively anaerobic.
Optimum temperature, 30° C.
Source: Isolated from New Jersey soil.
Habitat: Soil.
3. Alginomonas alginovora (Waks-
man et al., 1934) Kiss et al., 1945. {Bac-
terium alginovorum Waksman, Carey and
Allen, Jour. Bact., 28, 1934, 215; K&ss, Lid
and Molland, Norske Videnskaps-Akad.,
Oslo, I Mat.-Naturv. Klasse, No. 11, 1945,
9.)
al.gi.no'vo.ra. L. fem.n. alga seaweed;
M.L. noun acidum alginicum alginic acid
(derived from seaweed); L. v. voro to de-
vour; M.L. adj. alginovorus alginic acid-
destroying.
Rods, 0.75 to 1.2 by 1.5 to 2.0 microns,
with rounded to almost elliptical ends,
especially when single, occurring frequently
in pairs and even in chains. Encapsulated.
Actively motile. Gram-negative.
Alginic acid plate: Colony large, white
in appearance, with coarse, granular center,
entire margin. Clears up turbidity caused
by alginic acid on the plate. No odor.
Alginic acid liquid medium: Heavy pel-
licle formation. Active production of an
enzyme, alginase, which brings about the
disappearance of alginic precipitate in sea-
water medium.
Salt-water medium: A slimy pellicle of a
highly tenacious nature is produced, the
whole medium later turning to a soft jelly.
Sea-water gelatin : Active and rapid lique-
faction in two to six days at 18° C; highly
turbid throughout the liquefied zone.
Agar liquefaction: Extensive softening of
agar, no free liquid.
Sea-water glucose broth: Abundant,
uniform turbidity with surface pellicle;
some strains give heavier turbidity, and
others heavier pellicle.
Litmus milk containing 3.5 per cent
salt: No apparent growth.
Potato moistened with sea water: Moist,
spreading, ivory-colored growth; heavy
sediment in free liquid at the bottom.
Starch plate: Abundant, cream-colored,
slimy growth; extensive diastase produc-
tion.
Aerobic to microaerophilic.
Optimum temperature, 20° C.
Source: Isolated from sea water, sea-
bottom sediments and from the surface of
algal growth in the sea.
Habitat: Very common in the sea.
4. Alginomonas fucicola (Waksman et
al., 1934) Kiss et al., 1945. {Bacterium fuci-
cola Waksman, Carey and Allen, Jour. Bact.,
204
ORDER I. PSEUDOMONADALES
28, 1934, 213; K&ss, Lid and MoUand, Norske
Videnskaps-Akad., Oslo, I Mat.-Naturv.
Klasse, No. 11, 1945, 9.)
fu.ci'co.la. L. mas.n./tiCMS a seaweed; M.
L. noun Fucus a genus of brown seaweeds;
L.v. colo to inhabit; M.L. noun fucicola the
Fucus dweller.
Short rods, 0.6 to 1.0 by 1.0 to 1.5 microns,
with ends rounded to almost coccoid;
slightly curved. Actively motile with twirl-
ing motion. Gram-negative.
Alginic acid plate : Colonies finely granu-
lar, entire; at first whitish, turning brown in
three to five days, and later almost black,
producing a deep brown, soluble pigment.
Alginic acid liquid medium: Limited
growth on surface in the form of a pellicle.
Frequently produces no growth at all.
Sea-water gelatin: Active liquefaction;
no growth on stab; thin, fluorescent growth
throughout liquefied zone.
Agar liquefaction: Positive, although lim-
ited; only softening of agar.
Sea-water glucose broth: Faint turbidity;
no pellicle; no sediment.
Litmus milk containing salt: No apparent
growth.
Potato moistened with sea water: No
growth.
Starch plate: No growth.
Aerobic.
Optimum temperature, 20° C.
Source : Isolated from sea water near the
surface of the sand bottom.
Habitat: Rare in sea water.
5. Alginomonas alginica (Waksman et
al., 1934) Kiss et al., 1945. {Bacterium algini-
cum Waksman, Carey and Allen, Jour.
Bact., 28, 1934, 213; K&ss, Lid and Molland,
Norske Videnskaps-Akad., Oslo, I Mat.-
Naturv. Klasse, No. 11, 1945, 9.)
al.gi'ni.ca. L. fem.n. alga seaweed; M.L.
adj . alginicus pertaining to alginic acid from
seaweed.
Rods short to almost spherical, 0.6 to 1.0
micron in diameter. Encapsulated. Slug-
gishly motile. Gram-negative.
Alginic acid plate: White, finely granu-
lated colonies with entire margin. Does not
clear up the turbidity in plate. Odor pro-
duced resembles that of old potatoes.
Alginic acid liquid medium: Thin pellicle;
weak alginase formation.
Sea-water gelatin: Thin growth through-
out gelatin stab; no liquefaction in 7 days
at 18° C.
Agar liquefaction: None.
Sea-water glucose broth: Uniform but
very limited turbidity; no pellicle; no sedi-
ment.
Litmus milk containing salt : No apparent
growth.
Potato moistened with sea water: Moist,
spreading, cream-colored growth; heavy
sediment in free liquid at bottom.
Starch plate: Limited, pale blue growth;
no diastase.
Aerobic.
Optimum temperature, 20° C.
Source: Isolated from sea water and from
the surface of algal growth.
Habitat: Common in sea water.
Genus X. Mycoplana Gray and Thornton, 1928.*
(Cent. f. Bakt., II Abt., 73, 1928, 82.)
My.co.pla'na. Gr. tnyces fungus; Gr. planus a wandering; M.L. fem.n. Mycoplana fungus
wanderer.
Cells branching, especially in young cultures. Frequently banded when stained. Polar
flagellate. t Capable of using phenol or similar aromatic compounds as a sole source of
energy. Grow well on standard culture media. From soil.
The type species is Mycoplana diniorpha Gray and Tho rnton.
* Prepared by Prof. Robert S. Breed, Cornell University, Geneva, New York, January,
1954.
t The orginal statements regarding the flagellation of these species are contradictory.
The first reads "Polar, peritrichous"; the second "Polar or peritrichous".— Editors.
FAMILY IV. PSEUDOMONADACEAE
205
I. Gelatin not liquefied
II. Gelatin liquefied.
Key to the species of genus Mycoplana.
1. Mycoplana dimorpha.
2. Mycoplana bullata.
1. Mycoplana dimorpha Gray and
Thornton, 1928. (Cent. f. Bakt., II. Abt.,
73, 1928,82.)
di.mor'pha. Gr. adj. diniurphus two forms.
Short, curved and irregular rods, 0.5 to
0.7 by 1.25 to 4.5 microns, showing branch-
ing especially in young cultures. Originally
reported as "polar, peritrichous". Draw-
ings show some cells with a polar flagellum
and others where the several flagella shown
could represent a tuft of polar flagella.
Cultures preserved in the American Type
Culture Collection have been retested (T.
H. Lord, Manhattan, Kansas; F. E. Clark,
Beltsville, Maryland) and show typical
pseudomonad cells, i.e., straight rods with a
single polar flagellum. Meanwhile P. H. H.
Gray (Macdonald College, Quebec) reports
that his cultures still show branching cells
on the media he uses. Gram-negative.
Gelatin colonies: Circular, bufi", smooth,
resinous, entire.
Gelatin stab: No liquefaction. Growth
filiform.
Agar colonies: Circular, buff, convex,
smooth, glistening, entire.
Agar slant: Filiform, white, convex,
glistening, entire.
Broth: Turbid, with surface ring.
Nitrites not produced from nitrates, but
gas evolved in fermentation tubes.
Starch hydrolyzed.
No acid from carbohydrate media.
Attacks phenol.
Aerobic.
Optimum temperature, below 30° C.
Source : Only one strain was found in soil
by Gray and Thornton {loc. cit.). Wood
(Aust. Jour. Marine and Freshwater Res.,
4, 1953, 184) identifies 1010 cultures out of
2969 cultures isolated from Australian
marine habitats as belonging to this species.
Some appeared on svibmerged glass slides as
attached forms. A diversity of characters
was found in these cultures, indicating that
many of them should not have been identi-
fied as belonging to this species or even to
this genus. For example, while it is stated in
one place that carbohydrate fermentation
is feeble, it is stated in another place that
about 50 per cent of the cultures actively
fermented maltose and sucrose, these sugars
being fermented more actively than glucose.
Some cultures are reported as attacking
cellulose, others as attacking alginates or
even chitin. In other words many of the cul-
tures identified as Mycoplana dimorpha
possessed characters not ascribed to the
species by Gray and Thornton. Apparently
all cultures from marine habitats that were
Gram-negative branching forms were identi-
fied as Mycoplana dimorpha unless they
showed a yellow, pink or lemon-yellow chro-
mogenesis. Wood's work would indicate that
branching, polar flagellate species of very
diverse physiologies exist in marine habitats
that are as yet scarcely studied from the
standpoint of the species present. M. E.
Norris of the Pacific Fisheries Experiment
Station, Vancouver, B.C. reports (personal
communication. May, 1954) that she also
finds Gram-negative, branching, polar
flagellate organisms in sea water.
Habitat: Probably widely distributed in
soil. Possibl}^ also found in marine habitats.
2. Mycoplana bullata Gray and Thorn-
ton, 1928. (Cent. f. Bakt., II Abt., 73, 1928,
83.)
bul.la'ta. L. adj. bullatvs with a knob.
Rods curved or irregular in shape, branch-
ing, 0.8 to 1.0 by 2.25 to 4.5 microns. Origi-
nally stated to be either "polar or peri-
trichous" in its flagellation, but recent
studies show that the American Type Cul-
ture Collection culture of this organism is
polar flagellate. It resembles Mycoplana
liiinurpha in this respect. Gram-negative.
Gelatin colonies: Circular, buff, smooth,
glistening; edge diffuse. Gelatin partially
liquefied.
206
ORDER I. PSEUDOMONADALES
Gelatin stab: Saccate liquefaction.
Agar colonies: Circular, white, convex,
smooth, glistening, entire.
Agar slant: Filiform, white, convex,
smooth, glistening, entire.
Broth: Turbid.
Nitrites not produced from nitrates. Gas,
presumably N2, in fermentation tubes.
Starch not hydrolyzed.
No acid from carbohydrate media.
Attacks phenol.
Aerobic.
Optimum temperature, below 30°C.
Source: Two strains isolated from soil.
Habitat: Probably widely distributed in
soil. Possibly also found in marine habitats.
Note: Species incertae sedis. Other bac-
teria from sea water, fresh water and soil
have been described as belonging in this
genus. Their relationships to the species
described by Gray and Thornton (Cent,
f. Bakt., II Abt., 73, 1928, 82) have not yet
been definitely established.
Genus XI. Zoogloea Cohn, 1854.*
(Nov. Act. Acad. Caes. Leop. -Carol. Nat. Cur., 24, 1854, 123.)
Zo.o.gloe'a. Gr. adj. zous living; Gr. gloea glue; M.L. fem.n. Zoogloea living glue.
Rod-shaped cells embedded in a gelatinous matrix. Free-floating forms found in fresh
water that contains organic matter. Occur as compact masses or as branched forms. Cells
may become detached and motile in which case they are monotrichous.
The original description of this genus follows :
Zoogloea. Cellulae minimae bacilliformes hyalinae, gelatina hyalina in massas globosas,
uvaeformes, mox membranaceae consociatae, dein singulae elapsae, per aquam vacillantes.
This may be freely translated as follows :
Zoogloea. Transparent, very small, rod-shaped cells embedded in transparent, gelatinous,
clustered, spherical masses. Afterwards become detached as individuals swimming to and
fro in the water.
Zoogloea termo Cohn, 1854, the tj^pe species (monotypy) of this genus, is generally thought
to be unrecognizable. While awaiting further study of this problem by modern methods, it
is recommended that Zoogloea ramigera Itzigsohn be accepted as the type species of Zoogloea
Cohn.
1. Zoogloea ramigera Itzigsohn, 1867,
emend. Bloch, 1918. (Itzigsohn, Sitzungsber.
d. Gesellschaft naturf. Freunde, Berlin,
Nov. 19, 1867, 30; Bloch, Cent. f. Bakt., II
Abt., 48, 1918, 44-62.)
ra.mi'ge.ra. L. ramus a branch; L. v. gero
to bear; M.L. adj . ramigerus branch-bearing.
Description taken from Bloch (loc. cit.),
who made the first cultural studies of this
species, Butterfield (Public Health Reports,
50, 1935, 671) and Wattie (Pub. Health Re-
ports, 57, 1942, 1519).
Rods, 1 by 2 to 4 microns, with rounded
ends. Numerous cells are found embedded
in a gelatinous, branching matrix (see Koch,
Beitrage z. Biol. d. Pflanzen, 2, Heft 8, 1877,
399, Taf. XIV, and Butterfield, op. cit.,
1935, plates I-IV). Free cells are motile with
a single, long, polar flagellum. Gram-nega-
tive.
Bloch reports no growth at 25° C. on gela-
tin, poor growth on nutrient agar, good
growth in nutrient broth, weak growth in
peptone water, very good growth in hay in-
fusions, good growth in yeast extract water,
no growth in liquid manure, no growth in
beer wort, no growth on potato and no
growth on yellow sugar beet. Butterfield re-
ports that growth is best in aerated liquid
media.
Nitrites not produced from nitrates.
Ammonia produced from peptones.
Indole not produced.
Hydrogen sulfide not produced.
Methyl red negative; acetylmethylcar-
binol not produced.
* Revised by Mrs. James B. Lackey n^e Elsie Wattie, University of Florida, Gaines-
ville, Florida, March, 1954.
FAMILY IV. PSEUDOMONADACEAE
207
Bloch reports that sugars are utilized in
developing cell substances. Wattie finds that
there is evidence of slight acid production
from glucose, lactose, xylose and mannitol,
whereas Butterfield finds no action on all
sugars tested. In addition to the sugars
named above, these included sucrose,
arabinose, galactose, mannose, cellobiose,
raffinose, melizitose, dextrin and salicin.
Ptire-culture "activated sludges" formed
bj' this species have been shown to produce
a high rate of oxidation of the pollutional
material in sewage (synthetic and natural),
oxidizing about 50 per cent of the 5-day bio-
chemical oxygen demand in a 5-hour aera-
tion period and about 80 per cent in a 24-
hour interval. Nitrogenous materials are not
included in this oxidation as this species is
not capable of such action.
Temperature relations: Optimum, be-
tween 28° and 30° C. Good growth at 20° and
at 37° C. Minimum, 4° C.
Optimum pH, 7.0 to 7.4.
Strict aerobe.
Distinctive characters: Oxidizes sewage
and other organic solutions. Also see
McKinney and Horwood (Sewage and Ind.
Wastes, 2^, 1953, 117), who found other floc-
forming organisms besides Zoogloea ramigera
in activated sludge; these were identified
as Bacillus cereus, Escherichia intermedia,
Paracolohactrum aerogenoides and Nocardia
actinomorpha. A species of Flavobacterium
was also found in the floes in association
with these species.
Source: Originally (1867) found in a cul-
ture of decomposing algae. It has been re-
peatedly found in materials containing de-
composing plant materials and sewage and
is especially common in the floes formed in
the activated sludge process of purifying
sewage.
Habitat: Produces zoogloeal masses in
water containing decomposing organic mat-
ter. Common.
2. Zoogloea filipendula Beger, 1928.
(Kl. Mitt. d. Ver. f. Wasser-, Boden- und
Lufthyg., Berlin-Dahlem, 4, 1928, 143; also
see Beger, Zent. f. Bakt., I Abt., Orig., 154,
1949, 61.)
fi.li.pen'du.la. L. nounfilum a thread; L.
adj. penduhis hanging down; M.L. adj.
filipendulus thread hanging down.
Description prepared by Prof. H. Beger,
Berlin-Dahlem, Germany.
Cells coccoid (0.8 micron in diameter) to
rod-shaped (0.8 by 2.0 microns). The cells
are surrounded by a gelatinous mass which
varies in size from 1.5 to 2.0 by 4.5 cm and
which is composed of numerous, more or
less spherical masses 3 to 5 mm long. The
largest cells completely fill the newly formed
globules which lie at the end of filaments
hanging downward from zoogloeal masses
suspended from the under surface of pump
pistons and other submerged objects; the
cells in the older globules are smaller (0.4
by 0.7 micron) and are found near the sur-
face of the globule, the interior being rela-
tively free of cells.
Several other bacteria are found in asso-
ciation with this species. As a result, the
gelatinous mass appears rust-colored (cov-
ered with iron bacteria) when found in acid
waters and grayish white when isolated from
water that is neutral.
Nutrient gelatin: Only the small forms,
such as those found in older globules, are
able to grow on this medium. Substantial
growth occurs at the bottom of the stab in
48 hours.
Source: Isolated from pump pistons and
other submerged objects from a waterworks
near Berlin.
f Habitat: Found in water contaminated
with sewage or industrial wastes.
Note: Species incertae sedis: For species
that resemble those placed in the genus
Zoogloea Cohn in many important respects,
see Nevskia ramosa Famintzin and Myco-
nostoc gregarium Cohn. Additional species
have also been placed in the genus Zoogloea.
Genus XII. Halobacterium Elazari-Volcani, 1940.*
(Elazari-Volcani, Studies on the Microflora of the Dead Sea. Thesis, Hebrew Univ., Jeru-
salem, 1940, V and 59; not Halibacterium Fischer, Ergebnis.se der Plankton-E.xpedition der
* Prepared by Dr. B. Elazari-Volcani, The Weizmann Institute of Science, Rehovoth,
Israel, February, 1955.
208
ORDER I. PSEUDOMONADALES
Humboldt-Stiftung, 1894, 19; not Halophilus Sturges and Heideman (nomen nudum), Abst.
of Bact., 8, 1924, 14; not Halobacterium Schoop (nomen nudum), Zent. f. Bakt., I Abt., Orig.,
134., 1935, 26; Halobacter Anderson, Applied Microbiol., 2, 1954, 66.)
Ha.lo.bac.te'ri.um. Gr. noun hals salt; Gr. dim. noun bacterium a small rod; M.L. neut.n.
Halobacterium the salt bacterium.
Obligate halophilic, rod-shaped bacteria which are highly pleomorphic. Require at least
12 per cent salt for growth, and will live even in saturated brine solutions. Motile species
are polar flagellate; some species are non-motile. Gram-negative. Usually chromogenic,
producing non-water-soluble, carotenoid pigments which vary in shade from colorless
to orange or even brilliant red. Carbohydrates may or may not be attacked without the
production of visible gas. Nitrates are reduced, occasionally with the production of gas.
Found in tidal pools, especially in the tropics, salt ponds, salt seas or other places where
heavy brines occur naturally; also found on salted fish, salted hides and similar materials.
The type species is Halobacterium salinarium Elazari-Volcani.
Key to the species of genus Halobacterium.
I. Gas not produced from nitrates.
A. Nitrites not produced from nitrates.
1. Pale pink to scarlet chromogenesi
2. Pink to dark red chromogenesis.
B. Nitrites produced from nitrates.
II. Gas produced from nitrates.
A. Produces acid from glucose.
B. Does not produce acid from glucose.
1. Halobacterium salinarium.
2. Halobacterium cutirubrum.
3. Halobacterium halobium.
4. Halobacterium marismortui.
5. Halobacterium trapanicum.
1. Halobacterium salinarium (Harrison
and Kennedy, 1922) Elazari-Volcani, 1940.
{Pseudomonas salinaria Harrison and Ken-
nedy, Trans. Royal Soc. of Canada, 16,
1922, 121; Flavobacterium (Halobacterium)
salinarium Elazari-Volcani, Studies on
the Microflora of the Dead Sea. Thesis,
Hebrew Univ., Jerusalem, 1940, 59.)
sa.li.na'ri.um. L. adj. salinarius of salt
works.
Occurs as spheres and rods. The spheres
are 0.8 to 1.4 microns in diameter. The rods,
0.6 to 1.5 by 1.0 to 6.0 microns, occur singly
as ovoid, amoeboid, clavate, cuneate, trun-
cate, spindle- and club-shaped, pyriform
and other irregular forms. Age of culture
and nature of medium influence the size and
shape of cells. Reproduction is by means of
fission but apparently also by budding.
Motile by means of a polar flagellum at one
or both poles. Gram-negative.
Does not grow on ordinary culture media
unless supplemented with 16 to 35 per cent
sodium chloride and 2 per cent MgS04-7H20
(Katznelson and Lochhead, Jour. Bact.,
64, 1952, 97) . Grows well on salted fish and
hides.
Gelatin (salt) : Slow liquefaction (Katz-
nelson and Lochhead, loc. cit.).
Codfish agar colonies (16 to 30 per cent
salt) : In seven days punctiform, smooth,
raised, entire, granular, pale pink to scarlet
(Ridgway chart), 1.5 mm in diameter.
Milk salt agar (24 to 35 per cent salt) :
Pink colonies 4 to 5 mm in diameter, be-
coming scarlet.
Putrefactive odor. Definite proteolytic
zones develop (Lochhead, Can. Jour. Res.,
10, 1934, 275).
Codfish agar slant (16 to 35 per cent salt) :
In seven days moderate, filiform, slightly
raised, glistening, smooth, translucent,
bright red, viscid. Unpleasant odor.
Milk salt agar slants (24 to 35 per cent
FAMILY IV. PSEUDOMONADACEAE
209
salt): Filiform, slightly raised, smooth,
glistening, butyrous, bright red (Lochhead,
loc. cit).
Broth (5 to 35 per cent salt) : No growth.
Good growth when grown according to di-
rections of Katznelson and Lochhead {op.
cit., 1952,97).
Codfish broth (25 per cent salt): Turbid,
dense, pink sediment; imperfect, pink pel-
licle.
Potato immersed in brine: No growth.
Indole not produced.
Hj'drogen sulfide is produced.
No indication of action on carbohydrates.
Starch not hydrolyzed.
Cannot utilize inorganic nitrogen as a sole
source of nitrogen. Tests (Warburg respir-
ometer) show active oxidation of amino
acids (such as serine, glutamic acid and as-
partic acid); also active oxidation of glj'c-
erol.
Nitrites not produced from nitrates
(Lochhead, op. cit., 1934, 275).
Aerobic.
Optimum temperature, 37° C. Grows at
22° C.
Optimum salinity, 28 to 32 per cent (Loch-
head, loc. cit.). When the salt concentration
is reduced to 8 per cent, cells are ruptured.
Distinctive characters: See Halobacterium
cutirubrum.
Source: Isolated from cured codfish
(Harrison and Kennedy, op. cit., 1922, 121)
and salted fish (Browne, Absts. Bact., 6,
1922, 25, and Proc. Soc. Exp. Biol, and Med.,
19, 1922, 321) ; also from salted hides (Loch-
head, op. cit., 1934, 275).
Habitat: Produces a reddening of salted
fish and hides where untreated solar salt is
used. Abundant in tidal pools along shores
of tropical seas. Reddens the water in the
pools where solar salt is produced as soon as
the brine is concentrated to 18 per cent salt.
Common on untreated solar salt.
2. Halobacterium cutirubrum (Loch-
head, 1934) Elazari-Volcani, 1940. {Serratia
cutirubra Lochhead, Can. Jour. Research,
10, 1934, 275; Flavobacterium {Halobacterium)
cutirubrum Elazari-Volcani, Studies on the
Microflora of the Dead Sea. Thesis, Hebrew
Univ., Jerusalem, 1940, 59.)
cu.ti.ru'brum. L. nonn cutis the skin; L.
adj . ruber red; M.L. adj . cutirubrus skin-red.
Occurs as spheres and rods. The spheres
are 1.0 to 2.0 microns in diameter, and the
rods measure 0.7 to 4.0 by 1.5 to 8.0 microns.
Age of culture and nature of medium influ-
ence the size and shape of cells. Rod forms
are motile with a single polar flagellum;
coccoid forms are motile when young. Gram-
negative.
No growth on ordinary media.
Milk agar (20 per cent salt to saturation;
optimum 28 to 32 per cent) colonies : 3 to 4
mm in diameter, round and slightly convex,
pink to dark red (rose dorde, Ridgway
chart) .
Milk agar slants: Growth filiform, slightly
spreading, rather flat with smooth, glisten-
ing surface and membranous consistency.
Proteolytic action.
Liquid media: No or slight growth. Good
growth when grown according to directions
of Katznelson and Lochhead (Jour. Bact.,
64, 1952, 97).
Gelatin (salt): Pronounced liquefaction.
Indole not produced (Lochhead, op cit.,
1934, 275) ; faint positive (Gibbons, Jour.
Biol. Board Canada, 3, 1936, 75).
Hydrogen sulfide is produced.
Tests (Warburg respirometer) show slow
oxidation of amino acids (such as serine,
glutamic acid and aspartic acid) ; also slow
oxidation of glycerol.
Nitrites not produced from nitrates.
Diastatic action negative.
No carbohydrate fermentation.
Aerobic.
Optimum temperature, 37°C.
Salt tolerance: Halophilic, obligate. No
rupturing of cells occurs when the salt con-
centration is reduced to 8 per cent; rup-
turing occurs when the salt concentration is
4 per cent.
Distinctive characters: Resembles Halo-
bacterium salinarium. Differs from it in mor-
phology and cultural characters, particu-
larly as regards color and consistency. More
actively proteolytic. Slower oxidative ac-
tion on amino acids and glycerol. Ruptur-
ing of cells does not occur as rapidly when
the salt concentration is reduced.
210
ORDER I. PSEUDOMONADALES
Source: Isolated from salted hides which
were presumably salted with solar salt.
Habitat: Sea water and solar salt.
3. Halobacterium halobium (Petter,
1931) Elazari-Volcaui, 1940. (Microbe du
rouge de morue, Le Dantec, Ann. Inst.
Past., 5, 1891, 656; also see Le Dantec,
Compt. rend. Soc. Biol., Paris, 61, 1906, 136;
Bacillus halobius ruber Klebahn, Mitteil.
a. d. Inst. f. allg. Bot. i. Hamburg, 4, 1919,
47; abst. in Cent. f. Bakt., II Abt., 52, 1921,
123; Bacterium halobium Petter, Proc. Kon.
Acad. V. Wetensch. Amsterdam, 34, 1931,
1417; also see Petter, Over roode en andere
bacterien van gezouten visch. Thesis,
Utrecht, 1932; Flavobacterium {Halobac-
terium) halobium Elazari-Volcani, Studies on
the Microflora of the Dead Sea. Thesis,
Hebrew Univ., Jerusalem, 1940, V and 59.)
ha.lo'bi.um. Gr. noun hals salt; Gr. noun
bius life; M.L. adj. halobius living on salt.
Rods, the length of which varies greatly
with the medium and age of culture: 0.6 to
0.9 by 2.0 to 6.0 microns in young cultures
grown on agar (30 per cent NaCl + 1 per
cent peptone "Poulenc"); 2.0 to 27.0 mi-
crons long in liquid peptone media, occur-
ring singly. (Klebahn described rods up to
45 microns in liquid media; in old cultures
(horse-serum agar), irregular involution
forms appear which are round, ovoid or
coccus-like, 1.0 to 1.5 by 1.7 to 2.7 microns.)
Cells from opaque colonies contain charac-
teristic gas vacuoles. The cells are very sen-
sitive to changes in salt concentration; be-
low 12 per cent NaCl and in w^ater, they
swell and form ovoid, amoeboid and club-,
spindle-, drumstick- and pear-shaped arte-
facts. Because of these irregular forms, the
organism was described by several investi-
gators as polymorphic (Cloake, Dept. of
Scientific and Ind. Research, Food Investi-
gation Board No. 18, London, 1923). Slightly
motile with a pendulum-like movement;
flagella observed with electron microscope
(Houwink, Jour. Gen. Microlnol., 15, 1956,
146). Gram-negative.
Agar colonies (30 per cent NaCl -f 1 per
cent peptone "Poulenc"): Circular, trans-
parent or opaque; color varies from almost
white to orange, red, violet and purple; the
color of the colonj^ also changes during the
course of growth.
Broth (30 per cent NaCl + 1 per cent pep-
tone "Poulenc"): Pellicle; turbid; colored
sediment.
Asparagine broth (30 per cent NaCl -|- 3
per cent asparagine) : No growth.
Indole not produced.
No acid from glucose, sucrose or maltose
(tests made in 30 per cent salt + 1 per cent
peptone -|- 2 per cent carbohydrate).
Nitrites produced from nitrates; no gas is
produced.
Catalase-positive.
Aerobic.
Optimum temperature, 37° C.
Salt tolerance: Halophilic, obligate;
grows above 12 per cent NaCl up to satura-
tion.
Distinctive character: The pigment is
soluble in methanol, ethanol, acetone,
chloroform, carbon disulfide, benzol, petro-
leum ether, toluene and xylene. The carot-
enoids are named bacterio-ruberine a and /3.
Source: Seven different strains were iso-
lated from reddened, salted codfish and her-
ring.
Habitat: Produces a red discoloration on
salted herring and codfish.
4. Halobacterium marismortui Ela-
zari-Volcani, 1940. (Flavobacterium {Halo-
bacterium) maris-mortui (sic) Elazari-Vol-
cani, Studies on the Microflora of the Dead
Sea. Thesis, Hebrew Univ., Jerusalem, 1940,
V and 48.)
ma.ris.mor'tu.i. L. noun mare the sea;
L. gen. noun maris of the sea; L. adj. mortuus
dead; M.L. gen. noun marismortui of the
Dead Sea.
Rods, the length and shape varj'ing
greatly with the medium : in Dead Sea water
-f 1 per cent proteose peptone, the cells
occur singly and measure 0.5 by 1.6 to 3.0
microns; in 24 per cent salt + 1 per cent
peptone and on agar + peptone + salt and
on Dead Sea water + peptone + agar, the
cells are spheroids which measure 1.0 to
1.5 microns in diameter. Non-motile. When
stained, the rods burst while the spheroids
retain their shape. Gram-negative.
Gelatin stab (18 per cent salt + 1 per cent
peptone + 30 per cent gelatin) : Surface
growth. No liquefaction (2 months).
Agar colonies (24 per cent salt + 1 per
FAMILY IV. PSBUDOMONADACEAE
211
cent proteose peptoue + 2 per cent KNO3) :
Circular, smooth, entire, raised to convex,
butyrous, glistening, opaque with a slightly
transparent margin which is less colored,
orange-brown, orange-red or orange-yellow.
Agar slant (24 per cent salt + 1 per cent
peptone + 2 per cent KNO3): Growth
moderate, filiform, raised to convex, glisten-
ing, smooth, butyrous, opaque, orange-red.
Broth (24 per cent salt + 1 per cent pep-
tone) : Turbid; orange-red pellicle; slightly
viscous sediment.
Asparagine broth (24 per cent salt + 1
per cent asparagine): Turbid.
Indole not produced.
Acid from glucose, fructose, mannose and
glycerol ; slight acid from xylose and salicin
(tests made in 24 per cent salt + 1 per cent
peptone + 1 per cent carbohydrate during
3 weeks).
Starch not hj-drolyzed.
Nitrites rapidly produced from nitrates;
gas is produced (tests made in 24 per cent
salt + 1 per cent peptone + 2 per cent
KNO3).
Aerobic.
Optimum temperature, 30° C.
Salt tolerance: Halophilic, obligate;
grows in 18 per cent to saturated salt solu-
tions; slight growth in 15 per cent salt.
Distinctive character: The pigment pro-
duces a blue color with concentrated sul-
furic acid, thus suggesting a carotenoid; it
is very soluble in pyridine, less soluble in
methanol, ethanol and chloroform, slightly
soluble in acetone, very slightly so in ben-
zol and insoluble in xylene and petroleum
ether.
Source: Isolated from Dead Sea water.
Habitat: Salt lakes.
5. Halobacteriuin trapanicum (Petter,
1931) Elazari-Volcani, 1940. (Bacterium
trapanicum Petter, Proc. Kon. Acad. v.
Wetensch. Amsterdam, S4, 1931, 1417; also
see Petter, Over roode en andere bacterien
van gezouten visch. Thesis, Utrecht, 1932;
Flavohacterium (Halobacterium) trapanicum
Elazari-Volcani, Studies on the Microflora
of the Dead Sea. Thesis, Hebrew Univ.,
Jerusalem, 1940, V and 59.)
tra.pa'ni.cum. M.L. adj. trapanicus per-
taining to "Trapani" salt.
Rods, 0.6 by 1.5 to 3.5 microns. The length
and shape of the cells may vary greatly with
the medium: in Dead Sea water + 1 per cent
proteose peptone, the cells occur singly and
are 0.45 to 0.55 by 1.5 to 4.8 microns with
occasional rods measuring 8.0 to 16.0 mi-
crons in length; in 24 per cent salt + 1 per
cent peptone, the short rods predominate;
in 24 per cent salt + 1 per cent peptone + 2
per cent KNO3 agar, the cells are ovoid,
measuring 1.0 to 1.5 microns in diameter.
Non-motile. Gram-negative.
Gelatin stab (18 per cent salt -f 1 per
cent peptone -1- 30 per cent gelatin) : Surface
growth. No liquefaction (2 months).
Agar colonies (24 per cent salt + 1 per
cent proteose peptone + 2 per cent KNO3) :
Small (1 to 2 mm in diameter), circular,
smooth, entire, convex, glistening, trans-
parent, light orange or slightly colorless.
Agar slant (24 per cent salt -f 1 per cent
peptone + 2 per cent KNO3): Growth
moderate, filiform, raised, glistening,
smooth, opaque or slightly transparent,
light orange.
Broth (Dead Sea water -f 1 per cent pep-
tone) : Orange-rose pellicle; turbid; orange
sediment. In 24 per cent salt + 1 per cent
peptone: orange -rose ring; turbid; orange
sediment.
Asparagine broth (24 per cent salt + 1
per cent asparagine) : Moderately turbid.
Indole not produced.
No acid from arabinose, xylose, glucose,
fructose, galactose, mannose, lactose, su-
crose, maltose, raffinose, inulin, dextrin,
glycerol, mannitol or salicin (tests made in
24 per cent salt + 1 per cent peptone + 1
per cent carbohydrate during 3 weeks).
Starch not hydrolyzed.
Nitrites rapidly produced from nitrates;
gas is produced (tests made in 24 per cent
salt + 1 per cent peptone + 2 per cent
KNO3).
Catalase-positive.
Aerobic.
Optimum temperature, between 30° and
37° C.
Salt tolerance: Halophilic, obligate;
grows in 18 per cent to saturated salt solu-
tions ; slight growth in 15 per cent salt.
Distinctive character: The pigment pro-
212 ORDER I. PSEUDOMONADALES
duces a blue color with concentrated sul- Source: Isolated from "Trapani" salt
furic acid, thus suggesting a carotenoid. r..^^ • -d ,~.j , ,
,. 1 ui ■ -J- 1 , , , • "^"^ ^ cannery in Bergen (Norway) and
Very soluble in pyridine; less soluble in = \ j /
methanol, ethanol and chloroform; slightly
from the water of the Dead Sea.
soluble in acetone, very slightly so in ben- Habitat: Sea salt, sea-water brine and
zol; insoluble in xylene and petroleum ether. salt lakes.
FAMILY V. CAULOBACTERACEAE HENRICI AND
JOHNSON, 1935, EMEND. BREED.*
(Includes the typical families and genera of Caulobacteriales (sic) Henrici and Johnson,
Jour. Bact., 29, 1935, 4 and ibid., 30, 1935, 83. The Order Caulobacterales Henrici and Johnson
was redefined as a Sub-order, Caidobacteriineae (sic), by Breed, Murray and Kitchens, Bact.
Rev., 8, 1944, 255. The present emendation reduces the Order Caulobacterales, as originally
defined, to the status of a family in the Sub -order Pseudomonadineae Breed, Murray and
Smith.)
Cau.lo.bac.ter.a'ce.ae. M.L. neut.n. Caulobacter the type genus of the family; -aceae
ending to denote a family; M.L. fem.pl.n. Caulobacteraceae the Caulobacter famil}^
Non-filamentous, rod-shaped bacteria normally attached by branching or unbranching
stalks to a substrate. In one floating form the stalks are branched. Cells occur singly, in
pairs or in short chains. The cells are asymmetrical in that a stalk is developed at one end
of the cell or ferric hydro.xide or other material is secreted from one side of the cell to form
stalks. Cells are polar flagellate in the free-living state, non-motile in the attached forms.
Gram-negative. Multiply by transverse fission, the daughter cells remaining in place or
swimming away as swarm cells. Typically fresh- or salt-water forms.
The family Caulobacteraceae, as here defined, includes the genera Caulobacter Henrici and
Johnson, Gallionella Ehrenberg, Siderophacus Beger and Nevskia Famintzin.
The species in this family as presented here have close affinities with the species in the
family Pseudomonadaceae . In all cases where motility has been observed and stains made,
polar flagella have been found. It seems probable that when the life histories of these seden-
tary bacteria have been investigated, it will be found that practically all, if not all, of these
attached forms develop a motile stage. Such a stage permits the distribution of the species
in its environment.
The stalked bacteria studied by Henrici and Johnson (op. cit., 30, 1935, 83) were of fresh-
water origin. Bacteria of this type are found, however, equally if not more abundantly in
marine habitats where they play their part in the fouling of underwater surfaces. ZoBell
and Upham (Bull. Scripps Inst, of Oceanography, LaJolla, California, 5, 1944, 253) summa-
rize this situation as follows: "Many of the bacteria found in sea water are sessile or peri-
phytic, growing preferential!}^ or exclusivelj^ attached to solid surfaces. The sessile habit
of marine bacteria is most pronounced when they are growing in very dilute nutrient solu-
tions, such as sea water, to which nothing has been added. . . . Most sessile bacteria appear
to attach themselves tenaciously to solid surfaces by exuding a mucilaginous holdfast. A
few have stalks. Some of the sessile bacteria grow on the walls of the culture receptacle
without clouding the medium itself.".
The submerged-slide technique as employed by Henrici (Jour. Bact., 25, 1933, 277) and
* Redefined and rearranged by Prof. Robert S. Breed, Cornell University, Geneva, New
York, December, 1953. Prof. Herbert Beger, Institut fiir Wasser-, Boden- und Lufthygiene,
Berlin-Dahlem, Germany, has given this section a further revision so as to include genera
and species not previously recognized in the Manual, February, 1954.
FAMILY V. CAULOBACTERACEAE 213
by ZoBell and Allen (Proc. Soc. Exper. Biol, and Med., 30, 1933, 1409) has proved to be
most useful for studying bacteria that live attached to a substrate.
The species included in Pasteuria Metchnikoff and Blaslocmdis Henrici and Johnson
reproduce by a curious form of fission or budding. They have been transferred to a new-
Order, Hyphomicrobiales Douglas.
Key to the genera of family Caulobacteraceae.
I. Long axis of cell coincides with axis of stalk. Stalks slender.
Genus I. Caulobacter , p. 213.
II. Long axis of cell transverse to long axis of stalk. Stalks may be twisted and branched.
A. Stalks are band-shaped or rounded. Contain ferric hydroxide.
1. Stalks band-shaped and twisted. Dumb-bell-shaped in cross section.
Genus II. Gallionella, p. 214.
2. Stalks horn-shaped, not twisted. Round in cross section.
Genus III. Siderophacus, p. 216.
B. Stalks lobose, composed of gum. Forming zoogloea-like colonies. Free-floating.
Genus IV. Nevskia, p. 216.
Genus I. Caulobacter Henrici and Johnson, 1935.
(Jour. Bact., 29, 1935, 4; ihid., 30, 1935, 83.)
Cau.lo.bac'ter. L. noun caxdis a plant stem or stalk; M.L. noun bader masculine form of
Gr. neut.n. bactrum a rod; M.L. mas.n. Caulobacter stalk rod.
Stalked, curved, rod-shaped bacteria, the long axis of the elongated cells coinciding with
the long axis of the stalks. Young cells motile by means of a single polar flagellum. Old cells
attached to submerged objects by a stalk that is a continuation of the cell. A holdfast is de-
veloped at the distal end. Multiplication of cells is by transverse binary fission. Periphytic,
growing upon submerged surfaces.
The type species is Caulobacter vibrioides Henrici and Johnson emend. Bowers et al.
1. Caulobacter vibrioides Henrici and Agar colonies: Surface colonies up to 5
Johnson, 1935, emend. Bowers et al., 1954. mm in diameter, round, smooth, slightly
(Henrici and Johnson, Jour. Bact., 30, 1935, raised, glistening, finely granular in the
83; Bowers, Weaver, Grula and Edwards, center, grayish white, with center and re-
Jour. Bact., 68, 1954, 194.) verse side becoming brownish yellow. Sub-
vib.ri.oi'des. L.v. vibro to vibrate; M.L. surface colonies dense, brownish yellow,
noun Vibrio name of a genus; Gr. noun lenticular, up to 0.5 mm in diameter and 1.0
eidus shape, form; M.L. adj. vibrioides re- mm in length,
sembling a vibrio. Agar slant: Growth filiform, grayish
Cells elongated, curved, vibrio-like, with white, glistening, viscid,
rounded ends, 0.5 to 1.2 by 1.5 to 3.0 mi- Broth: Moderate turbidity with slightly
crons; filamentous forms occasionally pro- viscid sediment,
duced. Young cells actively motile with a Litmus milk: Unchanged,
single polar flagellum; older cells develop p^^^^^^ kittle or no growth,
a stalk at the flagellated end. The stalk has
a central filament or tube and a membrane
that is continuous with the cell wall. Organ-
isms attached singly or in rosettes, with
Indole not produced.
Nitrites not produced from nitrates.
No acid or gas from carbohydrates.
stalks attached to a common holdfast. Usu- Requires riboflavin, phosphates, iron and
ally surrounded by a slime layer. Gram- an organic source of energy for growth. Glu-
negative.
cose, maltose or casamino acids are used as
Gelatin: Surface growth and filiform sources of carbon and energy; sodium bi-
growth in stab without liquefaction. carbonate, sodium lactate, sodium acetate
214 ORDER I. PSEUDOMONADALES
or glycerol not utilized. Ammonium sulfate ander, Minnesota, and other fresh-water
or casamino acids used as sources of nitro- lakes (Henrici and Johnson, op. cit., 30,
gen; ammonium nitrate not utilized. 1935, 83). Also found in well-water in Ken-
Optimum temperature, 30° C. tucky (Bowers et al., op. cit.).
Aerobic, facultative. Habitat: Water, where it grows upon firm
Source: Found frequently in Lake Alex- substrates.*
Genus II. Gallionella Ehrenberg, 1838.-f
(Ehrenberg, Die Infusionsthierchen, 1838, 166; not Gaillonella Bory de St. Vincent, Diet.
Classique d'Hist. Nat., 4, 1823, 393; Didymohelix Griffith, Ann.
Mag. Nat. Hist., Ser. 2, 12, 1853, 438.)
Gal.li.o.nel'la. Named for Benjamin Gaillon, receiver of customs and zoologist (1782-
1839) in Dieppe, France; M.L. dim. ending -ella; M.L. fem.n. Gallionella a generic name.
Cells kidney-shaped or rounded. Placed at the end of the stalk with the long axis of the
cell transverse to the long axis of the stalk. Stalks secreted by the cells are slender and
twisted. Branch dichotomously or in the form of umbels. Stalks more or less dumb-bell or
bisquit-shaped in cross section. Composed of ferric hydroxide, completely dissolving in
weak acids. Two polar flagella are present when the cells are motile. Gram-negative. Multi-
plication by fission of the cells, the daughter cells remaining at first at the end of the stalk;
later they may be liberated as swarm cells. Grow only in iron-bearing waters. Do not store
manganese compounds. From both fresh and salt water. When the first species was discov-
ered the twisted stalks were thought to be a chain of diatoms.
The type species is Gallionella ferruginea Ehrenberg.
Key to the species of genus Gallionella.
I. Stalks branched.
A. Stalks dichotomously branched.
1. Stalks slender, spirally twisted,
a. Cells small, stalks very slender.
1. Gallionella ferruginea.
* The papers by Houwink (Antonie van Leeuwenhoek, 21, 1955, 29) and by Kandler,
Zehender and Huber (Arch. f. Mikrobiol., 21, 1954, 57) were received after the manuscript
covering the family Caulobacteraceae was prepared. They give further information regard-
ing the structure and function of the stalk of Caulobacter sp. Clearly the stalks developed
by species in this genus are quite different in nature from the stalks of ferric hydroxide or
gum secreted by the cells of other species placed in other genera of this family.
t Gallionella Ehrenberg is accepted and is continued in use in this edition of the Manual
although under a strict interpretation of Rules of Nomenclature it should apparently be
regarded as a homonym and therefore illegitimate. Gaillonella Bory de St. Vincent, proposed
as the name of a genus of algae, appears to have priority (see Internat. Bull. Bact. Nomen.
and Tax., 2, 1951, 96). However, Gaillonella B. de St. V. is no longer used by students of
diatoms so that Gallionella E. may be retained as a gemis conservandum in bacteriology with-
out causing confusion. Unless Gallionella E. is retained, the little used Didymohelix Griffith
must be again introduced into the Manual with the formation of a series of new combina-
tions.
The situation is complicated because the final settlement of this problem of nomenclature
requires action both by the Judicial Commission of the International Association of Micro-
biologists and the Special Committee on Diatomaceae of the International Botanical Con-
gress. The majority of the special students of iron bacteria have accepted and used Gallion-
ella E., e.g. Molisch (1910), Naumann (1921), Cholodny (1924), Butkevich (1928), Dorff
(1934), Henrici and Johnson (1934), Beger (1941) and Pringsheim (1952).
FAMILY V. CAULOBACTERACEAE
215
aa. Cells longer, stalks broader.
2. Galliunella major.
2. Stalks short, thick, not definitely in spirals.
3. GallioneUa minor.
B. Stalks branching in simple or compound umbels.
4. GallioneUa umbellata.
II. Stalks unbranched.
5. GallioneUa infiircata.
1. GallioneUa ferruginea Ehrenberg,
1836. {Gaillonella ferruginea (sic) Ehrenberg,
Vorl. Mittheil. ii. d. wirkl. Vorkommen
fossiler Infusionen u. ihre grosse Verbrei-
tung, Ann. Phys., Ser. 2, 8, 1836, 217;
GallioneUa ferruginea Ehrenberg, Die Infu-
sion thierchen, 1838, 166; Didymohelix ferru-
ginea Griffith, Ann. Mag. Nat. Hist., Ser. 2,
12, 1853, 438.)
fer.ru.gi'ne.a. L. adj. ferrugineus of the
color of iron rust.
Kidney-shaped cells. The full grown bac-
teria are 0.5 to 0.6 by 1.2 to 1.5 microns. The
cells secrete colloidal ferric hydroxide from
the concave portion of the cell, forming
band-like stalks 0.6 to 3.3 microns in width
and as much as 200 microns and more in
length. A rotatory motion of the cells gives
rise to a spiral twisting of the stalks.
In the older studies, the stalks were de-
scribed as the organism, the minute cells at
the tip having been dislodged or at least
overlooked. The cells lie at the tip of the
stalk and multiply by transverse binary
fission. This gives rise to a dichotomous
branching of the stalks. Stalks become very
long and slender, with smooth edges.
Not cultivated in artificial media.
Distribution: Usually the branched stalks
are attached separately in great numbers to
solid surfaces. They may, however, float in
irregular floes distributed throughout the
water. Less commonly they form balls up
to 3.0 microns in diameter. In these the
stalks radiate from a center and such groups
have been described as Gloeosphaera ferru-
ginea Rabenhorst. In a third variety, solid
tubercles richly encrusted with ferric com-
pounds are formed. These are found in old
pipelines or they may occur free in nature.
These tubercles have been named Sphaero-
fhrix latens Perfiliev.
Habitat: Found in cool springs and brooks
which carry reduced iron in solution; also
found in wells, in storage basins in water-
works and in pipe lines.
2. GallioneUa major Cholodny, 1927.
(Trav. Station, biolog. du Dniepre Acad,
des Sci. de I'Ukraine, Classe Sci. Phys. et
Math., 3, Livre 4, 1927.)
ma'jor. L. comp. adj. major larger.
Very similar to GallioneUa ferruginea, but
the cells are distinctly larger (1.0 by 3.0
microns). Stalks are 3.0 to 6.0 microns broad.
Some cells that fail to divide reach a length
of 7 microns or more; these form stalks of
double the normal width.
The cells contain one or more vacuoles,
apparently filled with an iron compound.
Source: Found in springs near Krassnodar
(Caucasus).
Habitat: Found in iron-bearing waters.
3. GallioneUa minor Cholodny, 1924.
(Ber. d. deutsch. Bot. Ges., 42, 1924, 42; also
see Cholodny, Die Eisenbakterien, Pflan-
zenforschung. Heft 4, 1926, 47.)
mi 'nor. L. comp. adj. minor smaller.
Cells as in GallioneUa ferruginea, but
stalks are shorter, thicker and more band-
like than twisted. After division, cells do not
separate as quickly as in GallioneUa ferrii-
ginea. The branches gradually become en-
crusted until the stalks are quite obscured.
Branches of stalks are not more than 20 to
30 microns long.
Source : Found in a small spring near the
Biological Station in Dniepre. Also found
by Beger (Ber. d. deutsch. Bot. Ges., 62,
1944, 11) in material from Camerun in
Africa.
Habitat: Found in iron-bearing waters.
4. GallioneUa umbellata Beger, 1949.
{GallioneUa ferrxiginea Palm, Svensk. Bot.
216
ORDER I. PSEUDOMONADALES
Tidskr., 27, 1933, 360; not GaUionella ferru-
ginea Ehrenberg, Die Infusionthierchen,
1838, 166; Beger, Ber. d. deutsch. Bot. Ges.,
52, 1949, 9.)
um.bel.la'ta. L. noun iimhella umbrella;
M.L. adj. umbellatus umbel-like.
Five to six cells are formed at the end of
the stalks before separation. The cells are
kidnej^-shaped and 1.0 by 2.0 microns in
size. The stalks then divide into 5 to 6
branches forming a simple umbel. This proc-
ess of cell division and growth of branches
continues until finally the whole mass ap-
pears composed of umbels.
Source: From leaf mold found in streams
in British-Gambia.
Habitat: Found in tropical, iron-bearing
streams.
5. GaUionella infurcata Beger, 1937.
(Spiro-phyUum sp., Suessenguth, Cent. f.
Bakt., II Abt., 1927, 69 and 339; Beger, Gas-
und Wasserfach, 80, 1937, 887; Spirophyllum
infurcatum Beger, ibid., 889.)
in.fur.ca'ta. L. prep, in in; L. nonn furca
fork; M.L. adj. furcatus forked; M.L. adj.
infurcatus forked.
Stalks twisted but not branched. Cells
coccoid, 1 micron in diameter. After fission
into two cells, they become detached from
the stalk.
Source: Found in water basins in the
Botanical Garden of Miinchen-Nymphen-
burg.
Habitat: Found in iron-bearing waters.
Genus. III. Siderophaciis Beger, 1944-
(Ber. d. deutsch. Bot. Ges., 61, 1944, 12.)
Si.de.ro'pha.cus. Gr. noun siderus iron; Gr. noun phacus lentil; M.L. mas.n. Sidero-
phaciis iron lentil.
The stalks are horn-shaped, without branches, and do not form twisted bands; they are
round to ovoid in transverse section. Cells biconcave or rod-like; after division they sepa-
rate from the stalk. Ferric hydroxide is stored in the stalks.
The type species is Siderophaciis corneolus (Dorff) Beger.
1. Siderophaciis corneolus (Dorff, 1934)
Beger, 1944. (GaUionella corneola Dorff, Die
Eisenorganismen, Pflanzenforschung, Heft
16, 1934, 25; Beger, Ber. d. deutsch. Bot.
Ges., 61, 1944 12.)
cor.ne'o.lus. L. adj. corneolus horny, firm.
Stalks 15 to 30 microns long, broader at
the top than at the base. Three to eight
stalks arise from a broad holdfast. Cells
0.6 to 1.0 by 2.5 to 3.0 microns.
Source: Found in an iron-bearing rivulet
near Lot-Malmby, Central Sweden; also
found near Berlin.
Habitat: Found in iron-bearing waters.
Genus IV. Nevskia Famintzin, 1892.
(Bull. Acad. Imp. Sci., St. Petersb., Ser. IV, 34 (X.S. 2), 1892, 484.)
Nev'ski.a. Neva a river at Leningrad; M.L. fem.n. Nevskia of the Neva.
Stalked bacteria, the long axis of the rod-shaped cells being set at right angles to the axis
of the stalk. Stalks lobose, dichotomously branched and composed of gum. Multiplication
of cells by transverse binary fission. Grow in zoogloea-like masses in water.
The type species is Nevskia ramosa Famintzin.
1. Nevskia ramosa Famintzin, 1892.
(Bull. Acad. Imp. Sci., St. Petersb., S^r.
IV, 34 (N. S.^), 1892,484.)
ra.mo'sa. L. adj. ramosus branched.
Globular, bush-like or plate-like colonies
of gummy consistency which float upon the
surface of water. Colonies composed of
gummy material arranged in dichotomously
branched stalks arising from a common
base, with the bacterial cells contained in
the gum, a single cell at the tip of each
stalk. At times cells are set free from the
stalks to start new colonies.
Rod-shaped cells set with their long axis
FAMILY VI. SIDEROCAPSACEAE 217
at right angles to the axis of the broad, lobe- Source: Found in the aquarium in the
like stalk. Cells 2 by 6 to 12 microns, con-
taining a number of highly refractile glob-
ules of fat or sulfur. Multiplication by
Botanical Garden, St. Petersburg. Similar
but smaller organisms found b}' Henrici
binary fission. and Johnson (Jour. Bact., SO, 1935, 63) in a
Not cultivated on artificial media.
Note: Nevskia pediculata Henrici and
Johnson is now regarded as a Lactobacillus.
See Lactobacillis brevis Bergey et al., syn. apolis.
Betabacterium vermijonne IMayer. Habitat : Found in water
jar of water from the lily pond of the Univer-
sity of Minnesota greenhouse in Minne-
FAJVIILY VI. SIDEROCAPSACEAE PRIBRAM, 1929.*
(Tribe Siderocapseae Buchanan, Jour. Bact., 3, 1915, 615; Pribram, Jour.
Bact., 18, 1929, 377.)
Si. de.ro. cap. sa'ce.ae. M.L. fem.n. Siderocapsa type genus of the family; -aceae suffix to
denote a family; M.L. fem.pl.n. Siderocapsaceae the Siderocapsa family.
Cells spherical, ellipsoidal or bacilliform. Frequently embedded in a thick, mucilaginous
capsule in which iron or manganese compounds may be deposited. Motile stages, where
known, are polar flagellate. Free-living in surface films or attached to the surface of sub-
merged objects. Form deposits of iron and manganese compounds. Autotrophic, faculta-
tively autotrophic and heterotrophic species are included in the family. Found in fresh wa-
ter.
The morphology of the bacteria of this family is best determined after dissolving the
iron or manganese compounds with weak acids and staining with Schiff's reagent.
The type genus is Siderocapsa Molisch.
Key to the genera of family Siderocapsaceae.
I. Cells surrounded by capsular matter with iron compounds deposited either on the sur-
face or throughout the capsular material.
A. Cells coccoid.
1. Cells in masses in a common capsule.
Genus I. Siderocapsa, p*. 218.
2. Cells always in pairs in a gelatinous capsule.
Genus II. Siderosphaera, p. 220.
B. Cells ellipsoidal to bacilliform.
1. Cells heavily encapsulated but do not possess a torus. f
a. Cells in chains in a gelatinous capsule.
b. Chains of ellipsoidal cells embedded in a gelatinous capsule, the outlines
of which follow the form of the cells.
Genus III. Sideronema, p. 220.
bb. Rods in pairs or chains in surface films.
* Manuscript prepared by Prof. Robert S. Breed, Cornell Universitj^, Geneva, New York,
December, 1953; further revision with the introduction of additional genera and species
by Prof. Dr. Herbert Beger, Institutfur Wasser-, Boden- und Lufthygiene, Berlin-Dahlem,
Germanj% March, 1954.
t The so-called torus is a marginal thickening of a thin capsule. The torus is heavily im-
pregnated with iron compounds so that the torus of an individual cell looks like the link
of a chain or, if incomplete, like a horseshoe.
218 ORDER I. PSEUDOMONADALES
Genus IV. Ferribacterium, p. 221.
aa. Coccoid to rod-shaped cells in masses in a gelatinous capsule. Usually show
an irregular arrangement of cells.
Genus V. Sideromonas , p. 222.
2. Cells with a thin capsule with a torus,
a. Torus completely surrounds the cells.
Genus VI. Naumanniella, p. 223.
aa. Torus open at one pole giving the wall the appearance of a horseshoe.
Genus VII. Ochrobium, p. 225.
II. Non-encapsulated cells which form deposits of iron compounds in the cell wall, on
the surface of the cells or in the surrounding medium.
A. Cells coccoid.
Genus VIII. Siderococcus , p. 225.
B. Cells rod-shaped.
1. Found in neutral or alkaline waters.
Genus IX. Siderobacter , p. 226.
2. Found in acid mine wastes.
Genus X. Ferrobacillus, p. 227.
Genus I. Siderocapsa Molisch, 1909.
(Ann. Jard. Bot. Buitenzorg, 2 S^r., Supp. 3, 1909, 29; also see Die Eisenbakterien,
Jena, 1910, 11.)
Si.de.ro.cap'sa. Gr. noun siderus iron; L. noun capsa a box; M.L. fem.n. Siderocapsa
iron box.
One to many spherical to ellipsoidal, small cells embedded without definite arrangement
in a primary capsule. The primary capsules may be surrounded by a large secondary cap-
sule, and these may then be united into larger colonies. Iron compounds are predominantly
stored on the surface of the primary capsule, and when a secondary capsule is present, it
is also completely covered.
The type species is Siderocapsa treubii Molisch.
Key to the species of genus Siderocapsa.
I. Several cells in each capsule.
A. Attached forms.
1. Cells small; up to 8 in number in each capsule.
1. Siderocapsa treubii,
2. Cells larger; up to 100 and more in each capsule.
2. Siderocapsa major.
B. Plankton forms.
1. Primary capsules 3.5 to 9 microns in diameter, each containing 2 to 8 cells.
3. Siderocapsa coronata.
2. Primary capsules 10 to 20 microns in diameter, each containing up to 60 or more
cells.
4. Siderocapsa eusphaera.
II. Only one cell in each capsule. Cells always small.
A. Capsules attached.
5. Siderocapsa monoeca.
B. Capsules form unattached iron floes.
6. Siderocapsa botryoides.
FAMILY VI. SIDEROCAPSACEAE
219
1. Siderocapsa treiibii Molisch, 1909.
{Siderocapsa Treuhii (sic) Molisch, Ann.
Jard. Bot. Buitenzorg, 2 Ser., Supp. 3, 1909,
29; also see Die Eisenbakterien, Jena, 1910,
11.)
treu'bi.i. M.L. gen. noun treuhii of Treub;
named for Prof. Treub, director of the
Tropical Garden at Buitenzorg, Java.
Cocci, 0.4 to 0.6 micron in diameter. As
many as 8 cells may be embedded in zoo-
gloeal masses surrounded by ferric hy-
droxide and other iron or manganese com-
pounds; these masses are 1.8 to 3.6 microns
in diameter.
Deposits ferric hydro.xide on the surfaces
of water plants.
Regarded by Hardman and Henrici (Jour.
Bact., 37, 1939, 97) as a heterotrophic organ-
ism that utilizes the organic radicle of
organic iron compounds, depositing the iron
as a waste product on the capsules of the
colonies.
Source: Found attached to the roots, root
hairs and leaves of water plants {Elodea,
Nymphaea, Sagittaria, Salvinia, etc.) in Java.
Habitat: Widely distributed in fresh
water. Epiphytic on submerged plants or
on other objects. Abundant in alkaline,
hard-water lakes of the drainage type in
Minnesota and Wisconsin according to
Hardman and Henrici {ibid., 103). Absent
in neutral or acid soft-water lakes of the
seepage type.
2. Siderocapsa major Molisch, 1909.
(Ann. Jard. Bot. Buitenzorg, 2 S^r., Supp.
3, 1909, 29; also see Die Eisenbakterien,
Jena, 1910, 13.)
ma'jor. L. comp.adj. major larger.
Cells colorless, coccus-like, short rods,
0.7 by 1.8 microns. A colony may consist of
100 or more cells in the same mucilaginous
capsule.
Similar to Siderocapsa treuhii except that
the cells are larger and the gelatinous
capsule is less sharply defined. May be free-
floating in surface films or may be attached
to submerged objects.
Forms intermediate between Siderocapsa
major and Siderocapsa treuhii have been
observed by Hardman and Henrici (Jour.
Bact., 37, 1939, 97).
Source: Found on the surface of a Spiro-
gyra sp. near Prague.
Habitat: Widely distributed in fresh
water.
3. Siderocap.sa coronata Redinger, 1931.
(Arch. f. Hydrobiol., ££, 1931, 410.)
co.ro.na'ta. L. part. adj. coronatus
crowned.
Coccoid cells, about 1.0 micron in diam-
eter, occurring in the primary capsule in
groups of 2 to 8. These groups are sur-
rounded by secondary gelatinous capsules
which may unite into foamy, irregular
masses 5 to 10 or more cm in diameter. The
capsular material contains deposits of iron
and manganese. Free-floating. Yellowish to
dark brown in color.
Source: Found in water from Upper
Lake, Lunz, Austria. Foamy masses are
formed in the winter time. Ruttner (Arch,
f. Hydrobiol., 33, 1937, 167) reports that the
distribution of this organism in Alpine lakes
is related to the o.xygen stratification
therein : it was found most frequently at
depths of from 17.5 to 27.5 meters, where the
oxygen range was 0.12 to 0.30 mg per liter.
4.66 mg per liter was the highest oxygen
tension at which it was found.
Habitat: Presumably widely distributed
in water.
4. Siderocapsa eusphaera Skuja, 1948.
(Symbolae Bot. Upsal., 9 (3), 1948, 12.)
eu.sphae'ra. Gr. prep, eu true; Gr. noun
sphaera ball, sphere; M.L. noun eusphaera
a true sphere.
Cells coccus-shaped, 1 to 2 microns in
diameter, 2 to 60 and more in a primary
capsule. The latter are 10 to 20 microns in
diameter and are surrounded by a large
secondary capsule up to 50 microns in diam-
eter. The secondary capsules are united into
large colonies with a common mucilaginous
layer. The secondary capsule stores com-
pounds of iron and manganese.
Source: Found in lakes in Sweden; found
in the plankton at levels where the oxygen
tension is low.
Habitat: Presumably widely distributed
in fresh-water lakes.
220
ORDER I. PSEUDOMONADALES
5. Siderocapsa nionoeca Nauniann,
1922. (Siderocapsa monoica (sic) Naumann,
Kgl. Svensk. Vetensk. Akad. Handl., I, 62,
1922, 49.)
mo.noe'ca. Gr. adj. monus alone, solitary;
Gr. noun oecus house, dwelling; M.L. adj.
monoecus solitary dwelling.
Cells single, coccus-shaped or ellipsoidal,
0.5 to 0.7 micron in diameter, surrounded
by a more or less thick layer of iron and
manganese compounds in which, at least
when young, a rounded space is kept free.
The cell may be seen in this clear space.
Although the cells are found in great num-
bers in close proximity to each other, they
are distinctly isolated. Form iron and man-
ganese deposits on the surface of water
plants and submerged objects.
Source: Isolated from the surface of
Poiamogeton nutans in Sweden.
Habitat: Found in ponds, rivers and
waterworks; presumably widely distributed.
6. Siderocapsa botryoides Beger, 1949.
(Zent. f. Bakt., I Abt., Orig., 15J^, 1949, 65.)
bot.ry.o.i'des or bot.ry.oi'des. Gr. adj.
botryoides like a bunch of grapes.
Cells coccus-shaped, spherical or ellipsoi-
dal, 0.6 to 0.8 micron in diameter. With
the capsule they are 0.8 to 2.0 microns in
diameter, lying singly only when young,
later forming spherical to clustered colonies
up to 0.3 cm long.
The encrusted colonies form iron or
manganese floes.
Source: Found in wells and waterworks
near Berlin.
Habitat: Presumably widely distributed.
Genus II. Siderosphaera Beger, 1944-
(Ber. d. deutsch. Bot. Ges., 62, (1944) 1950, 7.)
Si.de.ro.sphae'ra. Gr. noun siderus iron; L. noun sphaera ball, sphere; M.L. fem.n.
Siderosphaera iron sphere.
Small, coccoid cells, always occurring in pairs and embedded in a primary capsule. After
cell division the daughter pairs, with the primary capsules, are surrounded by a new, com-
mon capsule. This division continues up to the formation of eight pairs and results in a
round, ball-shaped Gloeocapsa-like stage which stores compounds of iron. A number of these
balls unite to form larger floes which may lie on the surface of bottom mud in fresh-water
ditches and swamps.
The type species is Siderosphaera conglomerata Beger.
1. Siderosphaera conglomerata Beger,
1950. (Ber. d. deutsch. Bot. Ges., 62, (1944)
1950, 7.)
con.glo.me.ra'ta. L. part. adj. conglomer-
ates rolled together.
Cocci, 1.0 to 1.2 microns in diameter,
each with a sheath about 2.0 microns in
diameter. These cells divide to form 2, 4 or,
at times, 8 pairs of cells in a clear, spherical,
gelatinous colony 8 to 10 microns in diam-
eter.
Spherical to ellipsoidal floes containing
these colonies may be as much as 500 mi-
crons in diameter. In general appearance
these floes resemble those formed b}^ Sidero-
capsa coronata found in the Upper Lake at
Lunz. When dilute HCl is added, the jelly-
like colonies may be liberated as the iron
salts dissolve. Manganese salts are appar-
ently not present.
Source: Found in a small ditch near Lunz
(Austrian Alps).
Habitat : Found on the surface of mud in
Genus III. Sideronema Beger, 1941-
(Zent. f. Bakt., II Abt., 103, 1941, 321.)
Si.de.ro.ne'ma. Gr. noun siderus iron; Gr. noun yiema thread; M.L. neut.n. Sideronema
iron thread.
Coccoid cells occurring in short chains which are enclosed in a gelatinous sheath. The
FAMILY Vr. SIDEROCAPSACEAE
221
cell membrane contains an abundance of ferric hydroxide whereas the sheath is relatively
devoid of this substance. Non-motile and unattached. Found in iron-bearing waters.
The type species is Sidcronemn qlobuUfervm Beger.
1. Sideroiiema globuliferiim Boger,
1941. (Sideronemn glohulijern (sic) Beger,
Zent. f. Bakt., II Abt., 103, 1941, 321.)
glob.u.li.fe'rum. L. divcv.noun cjlob^dus a,
small sphere, globule; L. v. fero to bear,
carry; M.L. adj. (/lobulijerus globule-bear-
ing.
Cells coccoid, round to egg-shaped, 4.8
to 5.0 by 6.5 microns. Occur in chains (3 to
S cells) which are enclosed in a gelatinous
sheath 1.6 microns thick; the cells in these
chains are non-confluent. Ferric hydroxide
is found in the cell membrane but only
sparingly so in the sheath. Non-motile and
unattached.
Source: Found on gla.ss slides submerged
in spring water near Magdeburg, Germany.
Habitat: Presumably widely distributed
in iron-bearing waters.
Genus IV. Ferribacterium Brussoff, 1916.
(Brussoff, Cent. f. Bakt., II Abt., .(5, 1916, 547; Sideroderma Naumann, Kungl. Svenska
Vetenskapsakad. Handl., 62, 1922, 54.)
Fer.ri.bac.te'ri.um. L. nonn fernnn iron;Gr. dim. noun bacterium a small rod; M.L.
neut.n. Ferribacterium iron rodlet.
Rods, with rounded or square ends, usually occurring in pairs, sometimes appearing singly
or in short chains. Motility occasionally observed; presumably the cells are polar flagellate.
In most cultures the cells are enclosed in a gelatinous capsule which is ordinarily surrounded
hy deposits of iron compounds. Produces a pellicle on the surface of liquid media and wa-
ter. Found in iron- or manganese-bearing water.
The type species is Ferribacterium duplex Brussoff.
1. Ferribacterium duplex Brussoft",
1916. (Brussoff", Cent. f. Bakt., II Abt., ^5,
1916, 5-i7; Sideroderma duplex Naumann, Kgl.
Svenska Vetenskapsakad. Handl., 62, 1922,
55 and 63.)
du'plex. L. adj. duplex two-fold, double.
Rods, with rounded ends, 1.2 by 2.5 to
5.0 microns, occurring usually in pairs,
sometimes singl}' or in short chains. Re-
ported as non-motile. Cells enclosed in a
gelatinous capsule which is ordinarily sur-
rounded by iron compounds. According to
Sauer (Inaug. Diss., Kiel, 1934, 33) the cells
are motile and Gram-negative.
Peat-infusion agar: In old cultures the
gelatinous capsule is surrounded by a dark
sheath, never bj- an iron secretion; the
sheath is generally ellipsoidal. Irregular
forms are also found.
Iron ammonium citrate broth: Pellicle
scarcely visible, appearing yellow under
the microscope.
Iron peptone broth: Produces a barelj-
visible pellicle which appears yellow under
the microscope.
Water: A pellicle is formed which is
weakly iridescent or of a metallic sheen.
Aerobic.
Source: Isolated from an ochre-colored
sediment from two samples of tap-water
from Breslau labelled "Schwentniger" and
"Pirschamer".
Habitat: Found in iron-bearing waters.
2. Ferribacterium rectangulare (Nau-
mann, 1922) Beger, comb. nov. (Sideroderma
rectangulare Naumann, Kungl. Svenska
Vetenskapsakad. Handl., 62, No. 4, 1922, 54;
Sideroderma tenue Naumann, loc. cit.)
rec.tang.u.la're. L. adj. rectus straight;
L. adj. angularis angular; M.L. adj. rectan-
gularis rectangular.
Rods, with square ends, 0.5 by 3.0 mi-
crons. Embedded in capsular material in
pairs. Iron compounds deposited outside
222
ORDER I. PSEUDOMONADALES
the capsules which are found in surface
films. No motility observed.
Aerobic.
Comment: The differences between the
two species placed by Naumann (loc. cit.)
in the genus Sideroderma are not very sig-
nificant. Moreover, as surface-film organ-
isms, these are so much like F errihacterium
duplex Brussoff that they clearly belong in
the same genus.
Source: Found in the Anebodae region of
Sweden.
Habitat : Found in fresh water, in swampy
ditches and in small streams.
Genus V. Sideromonas Cholodny, 1922.
(Cholodny, Ber. d. deutsch. Bot. Ges., Ifi, 1922, 326; also see Die Eisenbakterien, Pflanzen-
forschung. Heft 4, 1926, 55; Siderothece Naumann, Kungl. Svenska Vetenskapsakad.
Handl., 62, No. 4, 1922, 18; Siderocystis Naumann, ibid., 42; also see Dorff, Die
Eisenorganismen, Pflanzenforschung, Heft 16, 1934, 12; and Beger, Ber. d.
deutsch. Bot. Ges., 62, (1944) 1950, 8.)
Si.de.ro.mo'nas. Gr. noun sider^ls iron; Gr. noun monas a unit, monad; M.L. fem.n.
Sideromonas iron monad.
Short, coccoid to rod-shaped cells each embedded in a rather large, sharply outlined
capsule. The number of cells may increase within the capsule, and older capsules may unite
to form larger colonies, the outlines of which are ill-defined. The capsules are impregnated
with iron or manganese compounds or are completely encrusted with them.
The type species is Sideromonas confervarum Cholodny.
1. Sideromonas confervarum Cho-
lodny, 1922. (Cholodnyi (sic), Ber. d.
deutsch. Bot. Ges., 40, 1922, 326; also see
Cholodny, Die Eisenbakterien, Pflanzen-
forschung, Heft 4, Jena, 1926, 55.)
con.fer.va'rum. M.L. fem.n. Conferva a
genus of algae; M.L. gen. pi. noun confer-
varum of confer vae
Coccobacteria, 0.5 to 0.6 by 0.8 to 1.0
micron, occurring in chains embedded in
gelatinous masses 10 to 100 microns in
diameter. Chains become visible when the
gelatinous mass is treated with formalin
followed by dilute HCl, washed in water
and stained with gentian violet or carbol-
fuchsin. No motility observed.
Form deposits of iron salts within the
gelatinous mass surrounding the chains.
Probably facultative autotrophic (Dorff,
Tabulae Biologicae, 16, 1933, 222).
Cause the algal cells that are surrounded
bj^ the zoogloeal masses to become darker
green than normal.
Source: Found on the surface of algae
{Conferva) in water containing iron salts
near Kiew (Ukraine); also found near
Berlin, Central and Southern Sweden,
Hungary and Moravia.
Habitat: Widely distributed on fresh-
water green
2. Sideromonas duplex (Naumann,
1922) Beger, comb. nov. {Siderocystis duplex
Naumann, Kungl. Svenska Vetenskap-
sakad. Handl., 62, No. 4, 1922, 43.)
du'plex. L. adj. duplex two-fold, double.
Slender rods with rounded ends, 0.5 by
1.5 to 3.0 microns, occurring singly or in
pairs. Embedded in capsules that may fuse
to form zoogloeal masses. Iron compounds
impregnate the capsular material. No
motility observed.
Comment: The differences between this
species and Ferribacterium duplex Brussoff
are slight; cultural studies may show these
two to be identical.
Source: Found in the Aneboda region of
Sweden.
Habitat: Found on submerged objects in
swampy ditches and small streams.
3. Sideromonas vulgaris (Naumann,
1922) Beger, comb. nov. {Siderocystis vulgaris
Naumann, Kungl. Svenska Vetenskapsa-
kad. Handl., 62, No. 4, 1922, 42.)
vul.ga'ris. L. adj. vulgaris common.
FAMILY VI. SIDEROCAPSACEAE
223
Slender rods measuring 0.5 by 2.5 microns.
Rods, several to many placed irregularly in
a gelatinous envelope, form, when old,
zoogloea-like masses as much as 7.5 microns
in diameter. The rods are surrounded by
primary capsules which are impregnated
with iron compounds and which later fuse.
Reported by Dorff (Tabulae Biologicae,
16, 1938, 221) to be autotrophic.
Comment: The characters of the genus
Siderocystis, as described by Naumann
{loc. cit.), do not seem adequate to distin-
guish it from the genus Sideromonas, estab-
lished earlier by Cholodny.
Source: Found in the Aneboda region in
Sweden.
Habitat: Forms deposits on submerged
objects in ditch and river waters.
4. Sideromonas major (Naumann, 1922)
Beger, comb. nov. {Siderothece major Nau-
mann, Kungl. Svenska Vetenskapsakad.
Handl., 62, No. 4, 1922, 17; Siderothece
minor Naumann, loc. cit.; Siderocystis
minor Naumann, ibid., 43.)
ma'jor. L. comp.adj. major larger.
Rods broader than those of Sideromonas
vulgaris, 0.5 to nearly 1.5 by 1.0 to 1.5 mi-
crons. Each rod is surrounded by a large
primary capsule; the capsules later fuse
and form a gelatinous envelope in which
the cells are irregularly arranged. Form
zoogloea-like masses up to 10 microns in
diameter. Iron compounds deposited within
the capsular substance.
Possibly autotrophic (Dorff, Talnilae
Biologicae, 16, 1938, 221).
Aerobic.
Comments: While there are some differ-
ences in the sizes of the organisms placed
by Naumann in the three different species
named above, such differences may, in
reality, not be significant: these differences
may be due to variations in the nutritive
value of the water in which each of the
organisms was growing.
Source: Found in the Aneboda region in
Sweden.
Habitat: Develop concretionary deposits
(microscopic particles) on submerged ob-
jects in swampy ditch and river waters;
also found in wells and pipes in waterworks.
Genus VI. Naumanniella Dorff, 1934.
(Die Eisenorganismen, Pflanzenforschung, Heft 16, 1934, 19.)
Nau.man.ni.el'la. M.L. dim. ending -ella; M.L. fem.n. Naumanniella named for Einar
Naumann, a Swedish limnologist.
Cells ellipsoidal or rod-shaped with rounded ends, occurring singly or in short chains;
the rods may be straight or curved and frequently are constricted in the middle. Each cell
is surrounded by a small capsule and a marginal thickening (torus) heavily impregnated
with iron and manganese compounds. Gelatinous capsules of the type found in Siderocapsa
are absent. Cell division occurs simultaneously with constriction and separation of the
torus. The species in this genus have not been cultured. Found at the surface and in or on
the bottom mud of iron-bearing water.
The type species is Naumanniella neuslonica Dorff.
Key to the species of genus Naumanniella.
I. Cells rod-shaped.
A. Cells occur singly.
1. Cell diameter greater than 1.2 microns with the torus.
a. Cells with the torus 1.8 to 3.3 by 4.0 to 10.0 microns.
1. Naumanniella neustonica.
aa. Cells with the torus 1.2 to 1.5 by 3.1 to 3.6 microns.
2. Naumanniella minor.
2. Cells 1 by 2 microns with the torus.
224
ORDER I. PSEUDOMONADALES
B. Cells occur in chains.
II. Cells ellipsoidal.
3. N aunianniella pygtnaea.
4. Naumanniella catenata.
6. Naumanniella ellipiica.
1. Naumanniella nenstonica Dorff,
1934. (Die Eisenorganismen, Pflanzen-
forschung, Heft 16, 1934, 21.)
neus.to'ni.ca. Gr. adj. neustus swimming,
floating; M.L. adj. neusfonicus of the neus-
ton (surface film).
Cells, including the torus, 1.8 to 3.3 by
4.9 to 10 microns; never curved but may be
slightly constricted. Without the torus the
cells measure 2.5 by 6.0 microns. Occur
singly in the surface film of water, rarely
on submerged plants.
Source: Found on the surface of iron-
bearing water from wells near Freienwalde
(1931) and Stolzenhagen (1932) in Mark
Brandenburg; also isolated at Brisbane,
Australia.
Habitat: Widely distributed in swamp
water.
2. Naumanniella minor Dorff, 1934.
(Die Eisenorganismen, Pflanzenforschung,
Heft 16, 1934,21.)
mi'nor. L. comp.adj. minor smaller.
Cells, including the torus, 1.2 to 1.5 by
3.1 to 3.6 microns; occur singly in the form
of rods which frequently are curved or
spiral-shaped. The cells are 0.9 by 3.0 mi-
crons irrespective of the torus. Usually
found in or on the bottom mud of fresh-
water ponds and swampy areas.
Source: Found at Wurms (Rhein) in the
bottom of a well which contained iron-bear-
ing water.
Habitat: Widely distributed in swamp
water; also found on ore or on the submerged
leaves of water plants.
3. Naumanniella pygmaea Beger, 1949.
(Zent. f. Bakt., I Abt., Orig., 154, 1949, 65.)
pyg.mae'a. Gr. adj. pygmaeus dwarfish.
Small, straight rods, with rounded ends,
1 by 2 microns with the torus. Occur singly.
Source: Isolated from pipes and deep
wells of waterworks near Berlin. Found on
the surface of the gelatinous mass formed
by Zoogloea filipendula Beger.
Habitat: Presumably widely distributed.
4. Naumanniella catenata Beger, 1941.
(Zent. f. Bakt., II Abt., 103, 1941, 32.)
ca.te.na'ta. L. part. adj. catenatus in
chains.
Cells 0.4 to 0.5 by 4.6 to 5.2 microns; with
the torus, 1.0 to 1.2 by 4.9 to 5.5 microns.
Cells elongated or slightly curved with
thick walls impregnated with iron. After
division the cells remain connected in
chains of several to many (3 to 12). These
cells are joined together in such a manner
that, because of the iron-impregnated,
marginal thickenings and the relatively
clear cells inside, they give the appearance
of a chain with elongated links. Non-motile
and unattached.
Source: Found on glass slides submerged
in spring water near Magdeburg, Germany.
Habitat: Presumably widely distributed
in or on the bottom mud of iron-bearing
waters.
5. Naumanniella elliptica Beger, 1949.
(Zent. f. Bakt., I Abt., Orig., 154, 1949, 63
and 65.)
el.lip'ti.ca. Gr. adj. ellipticus defective,
elliptical.
Cells ellipsoidal, 2.0 by 2.5 to 3.0 microns,
with a pronounced torus.
Source: Found in pipes and deep wells
of waterworks near Berlin. Found on masses
of Crenothrix polyspora threads lying on the
bottom mud.
Habitat: Presumably widelj' distributed
in or on the bottom mud of iron-bearing
waters.
FAMILY VI. SIDEROCAPSACEAE
225
Genus VII. Ochrobium Perfiliev, 1921.
(Perfiliev, in Wislouch, Bull. Institut Hydrobiol., Russia, 1921; Sideroderma in part,
Naumann, Kungl. Svenska Vetenskapsakad. Handl., 62, Part 4, (1921) March 20,
1922, 32; also see Naumann, Zent. f. Bakt., II Abt., 78, 1929, 514.)
O.chro'bi.um. Gr. noun ochra yellow ochre, iron oxide; Gr. noun bins life, dwelling;
M.L. neut.n. Ochrobium ochre-dweller.
Ellipsoidal to rod-shaped cells that are partially surrounded by a marginal thickening
(torus) that is heavily impregnated with iron. This torus remains open at one end so that
it resembles a horseshoe. The cells are surrounded b}' a delicate, transparent capsule that
contains a very small amount of iron. Polar flagellate. Widely distributed in fresh water.
The type species is Ochrobium tectum Perfiliev.
1. Ochrobium tectum Perfiliev, 1921.
(Perfiliev, in Wislouch, Bull. Institut
Hydrobiol., Russia, 1921; also see Nach-
richten des Sapropelkommittees, Leningrad,
1922, 1; and Verhandl. Intern. Verein. f.
theor. und angew. Limnologie (1925)3, T. 3,
1927; Sideroderma limneticum Naumann,
Kungl. Svenska Vetenskapsakad. Handl.,
62, 1922, 32.)
tec'tum. L. v. tego to cover; L. past part.
tectus covered.
Cells small, ellipsoidal to rod-shaped, 0.5
to 3.0 hj 1.5 to 5.0 microns. Each cell is
surrounded by a heavily iron-impregnated
torus which is open at one pole. Pairs of
cells appear like a pair of horseshoes with
the open ends together. The cells are cov-
ered with a delicate outer capsule, and they
may be united in small colonies. When
motile, they bear two unequal polar flagella.
Comment: The cells are much like those
found in the algal genus Pteromonas, only
smaller.
Source: Originally found in the region
about Leningrad; then found independently
by Naumann (Zent. f. Bakt., II Abt., 78,
1929, 514) in Sweden and later by Beger
(Zent. f. Bakt., I Abt., Orig., 154, 1949, 65)
in wells of waterworks near Berlin.
Habitat: Widely distributed iniron-bear-
ing waters.
Genus VIII. Siderococcus Dorff, 1934.
(Die Eisenorganismen, Pflanzenforschung, Jena, Heft 16, 1934 9.)
Si.de.ro.coc'cus. Gr. noun siderus iron; Gr. noun coccus a berry, sphere; M.L. mas.n.
Siderococcus iron coccus.
Cells cocciform and of small size. Lack a gelatinous capsule. Not encrusted with iron
compounds; these are deposited entirely outside of the cells.
The type species is Siderococcus limoniticus Dorff.
1. Siderococcus limoniticus Dorff,
1934. (Die Eisenorganismen, Pflanzenfor-
schung, Jena, Heft 16, 1934, 9.)
li.mo.ni'ti.cus. Gr. noun limon meadow,
bog; M.L. noun limonitum limonite, a min-
eral, ferrous iron o.xide; ALL. adj. limon-
iticus of limonite.
Cocci 0.2 to 0.5 micron in diameter. No
evident capsule. Utilize inorganic iron
compounds and deposit them outside of
the cells.
In liquid cultures, the cells produce, on
a glass slide, a sharply marked zone beneath
the surface in which iron compounds are
deposited on the slide. When the iron com-
pounds are dissolved with dilute HCl, very
tiny cocci are left on the slide.
Source: Isolated from limonite deposits
in a bay of Teufelsee near Freienwalde,
Austria. Also found in Russian and Swedish
iron ore deposits as well as in Java, Sumatra
and Borneo.
Habitat : Widely distributed in swamps
and lakes where limonite deposits are
forming.
226
ORDER I. PSEUDOMONADALES
2. Siderococcus communis Dorff, 1934.
(Die Eisenorganismen, Pflanzenforschung,
Jena, Heft 16, 1934, 9.)
com.mu'nis. L. adj. communis common.
Cocci to short rods, 0.4 to 1.0 micron in
diameter, occurring singly or in chains. No
capsules observed. Utilize organic iron
compounds (ferrous ammonium citrate)
and produce precipitates of ferric oxide.
Do not grow in water containing inorganic
iron compounds such as iron carbonate. Do
not grow on glass slides submerged in w^ater
containing organic iron compounds but are
found in the precipitate that is formed.
Source: Found in many European coun-
tries and in North America.
Habitat: Widely distributed in water
containing organic iron compounds.
Genus IX. Siderobacter Nauinann, 1922.
(Kungl. Svenska Vetenskapsakad. Handl., 62, No. 4, 1922, 55.)
Si.de.ro.bac'ter. Gr. noun siderus iron; M.L. noun bacter the masculine form of the Gr.
neut. n. hactrum a small rod; M.L. mas.n. Siderobacter iron rodlet.
Cells bacilliform with rounded ends; occur singly, in pairs or in short chains or are united
to form colonies. Lack a gelatinous capsule. Iron or manganese compounds are deposited
on the surfaces or in the membranes of the cells; the deposit may also be entirely outside
of the cells. Flagellated cells may occur. Found in neutral or alkaline waters.
The type species is Siderobacter linearis Naumann.
Key to species of genus Siderobacter.
I. Cells less than 1.0 micron in diameter. Found on the surface of zoogloeal masses.
A. Cells less than 0.5 micron in diameter.
1. Siderobacter gracilis.
B. Cells 0.8 micron in diameter.
2. Siderobacter hrevis.
II. Cells 1.0 micron or greater than 1.0 micron in diameter.
A. Cells 1.0 micron in diameter.
3. Siderobacter linearis.
B. Cells greater than 1.0 micron in diameter.
1. Cells in pairs and 1.5 microns in diameter.
4. Siderobacter duplex.
2. Cells 2.5 microns in diameter. Participate in the formation of iron and lime
concretions of macroscopic size.
5. Siderobacter latus.
1. Siderobacter gracilis Beger, 1949.
(Zent. f. Bakt., I Abt., Orig., ISJt, 1949, 65.)
gra'ci.lis. L. adj. gracilis slim, slender.
Cells 0.4 by 3.0 microns. Encrusted cells
are 5.0 to 7.0 microns long. Occur singly.
Participate in the formation of deposits of
iron compounds.
Source: Found on the surface of masses of
Zoogloea filipendula. This species formed
thick coatings on the walls of two wells
supplying rapid sand filters near Berlin,
Germany. The filters required frequent
washing because the coatings were easily
detached.
Habitat: Found in the cool waters of deep
wells.
2. Siderobacter brevis Beger, 1949.
(Zent. f. Bakt., I Abt., Grig., 1S4, 1949, 65.)
bre'vis. L. adj. brevis short.
Cells, 0.8 to 1.0 by 3.0 to 4.0 microns,
usually occurring singly. Participate in the
formation of deposits of iron compounds.
Source : Found on the surface of masses of
Zoogloea filipendula. This species formed
thick coatings on the walls of two wells
supphdng rapid sand filters near Berlin,
Germany. The filters required frequent
washing because the coatings were easily
detached.
3. Siderobacter linearis Naumann, 1922.
FAMILY VI. SIDEROCAPSACEAE
227
(Kungl. Svenska Vetenskapsakad. Handl.,
62, 1922, 55.)
li.ne.a'ris. L. adj. linearis linear.
The type species of genus Siderobacter
Naumann.
Cells 1.0 by 5.0 microns after the encrust-
ing iron compounds have been dissolved
away with dilute HCl. Opaque, encrusted
cells, 1.2 b}- 7.0 microns. Always occur
singl}' in contrast to the larger-celled
Siderobacter duplex, where the cells occur in
pairs.
Source: Found in the Aneboda region in
Sweden.
Habitat: Found in surface films and on
submerged objects.
4. Siderobacter duplex Naumann, 1922.
(Kungl. Svenska Vetenskapsakad. Handl.,
62, No. 4, 1922, 55.)
du'plex. L. adj. duplex two-fold, double.
Cells, 1.5 by 3.5 microns after encrusting
iron compounds are removed with dilute
HCl, occurring in pairs.
Source: Found in the Aneboda region in
Sweden.
Habitat: Found in surface films on the
water of swamps and small streams.
5. Siderobacter latus Beger, 1941.
(Zent. f. Bakt., I Abt., Orig., 154, 1949, 63
and 66.)
la'tus. L. adj. latus broad.
Straight or occasionally curved cells, 2.5
b}^ 6.0 to 15.0 microns, usually occurring
singly. Participate in the formation of iron
and lime concretions.
Source: Found on concretions on the
brick walls of two wells suppljang a rapid
sand filter near Berlin, Germany.
Habitat : Found in the cool waters of deep
wells.
Genus X. Ferrobacillus Leathen and Braley, 1954*
(Bact. Proc. 54th General Meeting, Soc. of Amer. Bact., 1954, 44.)
Fer.ro. ba.cil'lus. L. noun /errum iron, here meaning ferrous iron; L. dim. noun bacillus
a small rod; M.L. noun Ferrobacillus ferrous-iron rodlet.
Short, plump, rod-shaped cells occurring singly and in pairs, seldom in chains; the cells
are not united to form colonies. O.xidize ferrous iron to the ferric state in acid environments.
Optimum reaction, pH 3.5.
The type species is Ferrobacillus ferrooxidans Leathen and Braley.
1. Ferrobacillus ferrooxidans Leathen
and Braley, 1954. (Ferrous iron oxidizing
bacterium, Leathen, IMcIntyre and Braley,
Bact. Proc. 52nd General Meeting, Soc. of
Amer. Bact., 1952, 15; also see Leathen,
Braley and Mclntj're, Appl. Microbiol., 1,
1953, 65; Leathen and Braley, Bact. Proc.
54th General Meeting, Soc. of Amer. Bact.,
1954, 44.)
fer.ro.o'xi.dans. L. noun ferruni iron;
Gr. adj. oxys sharp, acid; M.L. v. oxido to
oxidize or make acid; M.L. part. adj. fer-
rooxidans iron-oxidizing.
Rods 0.6 to 1.0 by 1.0 to 1.6 microns.
Motile, presumably polar flagellate. Gram-
negative.
Ferrous iron-silica gel : Colonies are small
and raised with irregular margins. Young
colonies are glistening and tan, but gradu-
ally become granular and brown with
oxidizing iron. A tan to brown area of
oxidized iron is frequently found around the
colony.
Liquid ferrous iron medium (Leathen,
Mclntyre and Braley, Science, 114, 1951,
280): Rapidly oxidized; forms a precipitate
of ferric hydroxide or basic ferric sulfate.
Acid thiosulfate liquid medium: Not
oxidized.
Optimum reaction, pH 3.5. O.xidation
retarded below pH 2.2 and above pH 4.6.
Optimum temperature, between 15°
and 20° C.
Strictly autotrophic. Derives energy by
the o.xidation of ferrous iron to the ferric
state. Utilizes the CO2 of the atmosphere
as a source of carbon.
Aerobic.
*Description of genus and species prepared by Wm. W. Leathen, Mellon Institute, Pitts-
burgh, Pennsylvania.
228 ORDER I. PSEUDOMONADALES
Distinctive characters: By catalytic ac- of even minute traces of ferrous iron medium
tion, this species increases by several fold to an acid thiosulfate medium may cause
the amount of sulfuric acid normally formed decomposition of the thiosulfate, evidenced
by the atmospheric oxidation of pyritic by the development of turbidity due to the
materials found in bituminous coal seams formation of colloidal sulfur; this purely
and associated rock strata. chemical reaction involving thiosulfate
maj^ easily be misinterpreted as a bacterial
oxidation of this same substrate.
Source: Isolated from bituminous coal
Comment: This organism closely resem
bles Thiobacillus ferrooxidans and may, in
fact, be identical with it. However Temple
and Colmer (Jour. Bact., 59, 1950, 317)
report that Thiobacillus ferrooxidans oxi- "^^^« drainages and from waters receiving
dizes thiosulfate while Leathen and Braley ^^^^ discharges.
(op. cit., 1954, 44) report that Ferrobacillus Habitat: Indigenous to bituminous coal
ferrooxidans does not oxidize thiosulfate. regions. Frequently form relatively hard
Thelatter workers (personal communication, granules of ferric iron in which many
May, 1954) further report that the transfer bacteria are entrapped.
FAMILY VII. SPIRILLACEAE MIGULA, 1894.
(Migula, Arb. Bact. Inst. Karlsruhe, 1, 1894, 237; Spirillobacteriaccae Orla-Jensen,
Jour. Bact., 6, 1921, 264.)
Spi.ril.la'ce.ae. M.L. neut.n. Spirillum type genus of the family; -aceac ending to denote
a family; M.L. fem.pl.n. Spirillaceae the Spirillum family.
Cells simple, curved or spirally twisted rods. These frequently remain attached to each
other after transverse division to form chains of spirally twisted cells. Cells are rigid and
usually motile by means of a single flagellum (rarely two) or a tuft of polar flagella. Gram-
negative. Frequently oxidative in their physiology. Aerobic or facultatively anaerobic,
although a few strict anaerobes occur among the vibrios (Desulfovibrio and Vibrio). Largely
water forms, although some are parasitic or pathogenic on higher animals and man.
Key to the genera of family Spirillaceae.
I. Curved, vibrio-like rods that are rarely united into a complete ring.
A. Cells curved; rods never united at the end into a ring-shaped cell. Usually possess
a single, polar flagellum.
1. Curved rods that are not known to attack cellulose.
a. Aerobic to anaerobic, heterotrophic vibrios.
Genus I. Vibrio, p. 229.
aa. Anaerobic, facultatively autotrophic vibrios that produce hydrogen sulfide
or methane.
b. Reduce sulfates to hydrogen sulfide.
Genus II. Desulfovibrio, p. 248.
bb. Reduce carbon dioxide to methane.
Genus III. Methanobacterium, p. 250.
2. Curved rods that attack cellulose.
a. Vibrio-like cells.
Genus IV. Cellvibrio, p. 250.
aa. Pointed, sickle-shaped cells.
Genus V. Cellfalcicula, p. 252.
B. Curved rods that join ends to form a complete ring.
Genus VI. Microcyclus, p. 253.
FAMILY VII. SPIRILLACEAE 229
II. Crescent-shaped to spiral cells that are frequently united into spiral chains of cells.
A. Cells not embedded in zoogloeal masses.
1. Spiral cells with polar flagellation.
a. Possess a tuft of polar flagella.
Genus VII. Spirillum, p. 253.
aa. Possess a single, polar flagellum.
Genus VIII. Paraspirillvm, p. 257.
2. Crescent-shaped cells with a tuft of flagella attached to the middle of the concave
side of the cell.
Genus IX. Selenomonas, p. 258.
B. Crescent- to spiral-shaped cells embedded in a spherical mass of jelly. Found in
fresh water.
Genus X. Myconostoc, p. 260.
Genns I. Vibrio Miiller, 1773 *
(Miiller, Vermium terrestrium et fluviatilum, 1, 1773, 39; Pacinia Trevisan, Atti d. Accad.
Fisio-Medico-Statistica in Milano, Ser. 4, 3, 1885, 83; Microspira Schroeter,
in Cohn, Kryptogamen-Flora von Schlesien, 3, 1, 1886, 168.)
Vib'ri.o. L. v. vibro to move rapidly to and fro, to vibrate; M.L. mas.n. Vibrio that
which vibrates.
Cells short, curved, single or united into spirals. Motile by means of a single, polar flagel-
lum which is usually relativel}^ short; rarely two or three flagella in one tuft. Grow well and
rapidly on the surfaces of standard culture media. Heterotrophic organisms varying greatly
in their nutritional requirements. Aerobic, facultative anaerobic and anaerobic species.
Widely distributed as saprophytic forms in salt- and fresh-water and in soil; also occur as
parasites and as pathogens.
See Genus I, Pseudomonas, of Family IV, Pseudomonadaceae , for a discussion of the border-
line between the genus Vibrio and the genus Pseudomonas.
Few comparative studies have been made on the species in this genus; it is therefore
impossible to prepare a really satisfactory differential key.
The type species is Vibrio comma (Schroeter) Winslow et al.
Kerj to the species of genus Vibrio.
I. Aerobic species.
A. Produce acid but no gas from glucose and usually from other sugars (one lumines-
cent, one halophilic and several agar-digesting species fail to produce acid from
glucose) .
1. Not luminescent, not able to digest agar and do not attack benzene ring com-
pounds or o.\idize oxalates so far as known,
a. Found in fresh water or in the body fluids of animals, including man.
b. Liquefy gelatin.
c. Indole produced.
d. Nitrites produced from nitrates.
6. Milk not coagulated.
f. Cause of cholera.
1. Vibrio comma.
ff. Cholera-like vibrio from fresh water.
2. Vibrio herolinensis.
* Revised by Prof. Robert S. Breed, Cornell University, Geneva, New York, January,
1954; the section covering the microaerophilic and anaerobic animal pathogens was re-
viewed by Dr. E. V. Morse, College of Agriculture, University of Wisconsin, Madison,
Wisconsin, May, 1955.
230 ORDER I. PSEUDOMONADALES
ee. Milk coagulated.
3. Vibrio metschnikovii .
dd. Nitrites not produced from nitrates.
4. Vibrio proteus.
cc. Indole not produced.
d. Found in the human buccal cavity.
5. Vibrio sputigenus.
dd. Cause of abscesses in the African toad.
6. Vibrio xenopus.
bb. Does not liquefy gelatin.
7. Vibrio leonardii.
aa. Require sea-water or heavy brine media for growth on fresh isolation,
b. Isolated from sea water.
c. A diffusible dark brown pigment is usually produced in gelatin
media.
8. Vibrio marinopraesens .
cc. A buff -colored pigment is produced on sea-water agar. No diffusible
pigment produced.
9. Vibrio phytoplanktis .
bb. Found growing in brines.
c. Acid from glucose.
10. Vibrio costicoliis.
cc. No acid from glucose.
11. Vibrio halonitrificans.
2. Luminescent, digest agar, attack benzene ring compounds or oxidize oxalates,
a. Produce luminescence especially on neutral media containing sea water or
the equivalent salt content,
b. Gelatin liquefied.
c. Require alkaline sea water or equivalent media for growth,
d. Optimum growth temperature, between 25° and 28° C.
12. Vibrio himinosus.
dd. Optimum growth temperature, between 30° and 32° C.
13. Vibrio indicus.
cc. Found in fresh water and in intestinal contents.
14. Vibrio albensis.
bb. Does not liquefy gelatin.
15. Vibrio pierantonii.
aa. Not as above.
b. Digest agar either actively or at least soften it.
c. Found in soil and in rotting organic matter,
d. Decomposes both cellulose and agar.
16. Vibrio agarliqvefaciens.
dd. Liquefies agar only.
17. Vibrio andoii.
cc. Found in sea water with rotting algae.
d. Nitrites produced from nitrates.
e. Colonies on agar are white to gray.
18. Vibrio beijerinckii.
ee. Colonies on agar are pale yellow becoming bright yellow
then pale brown.
19. Vibrio fuscus.
FAMILY VII. SPIRILLACEAE 231
dd. Nitrites not produced from nitrates.
20. Vibrio granii.
bb. Not as above.
c. Soil organisms that are known to attack benzene ring compounds,
d. Soil organism that attacks naphthalene.
21. Vibrio neocistes.
dd. Soil organism that attacks phenol and m-cresol.
22. Vibrio cyclosites.
cc. Soil organisms that are known to attack oxalates.
d. Grows well on calcium oxalate agar. White colonies.
23. Vibrio oxaliticus.
dd. Forms film on bottom of liquid oxalate media. Rose-red to
blood-red chromogenesis on oxalate agar.
24. Vibrio extorquens.
B. Do not attack carbohydrates.
1. Soil organism that is known to attack naphthalene.
25. Vibrio cuneatus.
2. Not as above.
a. Do not liquefy gelatin,
b. From fresh water.
26. Vibrio percolans.
bb. Requires sea-water media for growth on fresh isolation.
27. Vibrio adapiaius.
aa. Liquefy gelatin.
b. Causes a disease in fresh-water fishes.
28. Vibrio piscium.
bb. Requires sea-water media for growth on fresh isolation.
29. Vibrio hyphalus.
II. Anaerobic to microaerophilic species (all parasitic, normally pathogenic).
A. Microaerophilic species that are pathogenic to warm-blooded animals.
1. Cause of abortion in cattle and sheep.
30. Vibrio fetus.
2. Not as above.
a. Cause of swine dysentery.
31. Vibrio coli.
aa. Cause of dysentery in cattle and related animals.
32. Vibrio jejuni.
B. Strict anaerobes.
a. Produces gas and bad odors in protein media.
33. Vibrio niger.
aa. Does not produce gas and bad odors in protein media.
34. Vibrio sputorum.
1. Vibrio comma (Schroeter, 1886) wiede, Rogers and Smith, Jour. Bact., 5,
Winslow et al., 1920. (Kommabacillus, 1920 204.)
Koch, Berliner klin. Wochenschr., ^i, 1884, „^/,„„ n- ^ ^„ „^„.^„
o • -77 77 • • r, r. • com'ma. ur. comma a comma.
479; bpinllum cholerae asiaticae Zopf, Die , , , ,
Spaltpilze, 3 Aufl., 1885, 69; Microspira Slightly curved rods, 0.3 to 0.6 by 1.0 to
comma Schroeter, in Cohn, Kryptogamen ^.0 microns, occurring singly and in spiral
Flora V. Schlesien, 3, 1, 1886, 168; Vibrio chains. Cells may be long, thin and delicate
cholerae Neisser, Arch. f. Hyg., 19, 1893, 199; or short and thick. May lose their curved
Winslow, Broadhurst, Buchanan, Krum- form on artificial cultivation. Motile,
232
ORDER I. PSEUDOMONADALES
possessing a single polar flagellum. Gram-
negative.
Gelatin colonies: Small, yellowish white.
Gelatin stab: Rapid, napiform liquefac-
tion.
Agar colonies: Circular, whitish brown,
moist, glistening, translucent, slightlj^
raised, entire.
Agar slant: Brownish gray, moist, glisten-
ing.
McConkey's medium: Good growth,
colonies colorless when young, soon pinkish,
medium becomes darker red.
Broth: Slightly turbid, with fragile,
wrinkled pellicle and flocculent precipitate.
Peptone water: Characteristic rapid
growth, chiefly at surface, where, after 6
to 9 hours, a delicate membrane is formed;
little turbidity, deposit apparently derived
from pellicle (Topley and Wilson, Princip.
Bact. and Immun., 2nd ed., 1936, 388).
Readily isolated from the surface film of
0.1 per cent peptone water.
Litmus milk: Alkaline at the top and
slightly acid at bottom; generally not
coagulated; peptonized; reduced.
Potato: Dirty white to yellowish, moist,
glistening, spreading growth.
Blood serum: Abundant growth, some-
times slow liquefaction.
Blood agar: The blood pigment is di-
gested forming a greenish zone around
colonies; a true soluble hemolysin is not
formed (the El Tor vibrio also digests
blood pigment but in addition produces a
soluble hemolysin; otherwise it is said to
be indistinguishable from the typical
cholera vibrio) .
Indole produced.
Cholera-red reaction, which depends on
production of indole and reduction of
nitrates, is positive.
Hydrogen sulfide produced.
Acid but no gas from glucose, fructose,
galactose, maltose, sucrose and mannitol.
Slowly from glycerol. Does not attack
lactose, inulin or dulcitol.
Group I of Heiberg (Classification of
Vibrio cholerae and Cholera-like Vibrios.
Copenhagen, 1935) ferments mannose and
sucrose but not arabinose.
Hydrolyzes starch actively in alkaline
media.
Nitrites produced from nitrates.
High alkali but low acid tolerance:
optimum pH, between 7.6 and 8.0; for
isolation on Dieudonne's medium, pH 9.0
to 9.6.
Aerobic, grows best in abundant oxygen;
under strict anaerobiosis may fail to grow
altogether.
Optimum temperature, 37° C. Maximum,
42° C. Minimum, 14° C.
Source: Isolated from the intestinal
contents of cholera patients in Egypt and
India.
Habitat: Found in the intestinal contents
of cholera patients and carriers.
The relationships existing among the
cholerigenic and non-pathogenic water
vibrios, although studied intensively, have
not yet been completely defined. As a
working scheme, based on somatic (O) and
flagellar (H) antigen studies, Gardner and
Vankatraman (Jour. Hyg., 35, 1935, 262-
282) suggest the one shown in the graph
on the following page.
Linton (Bact. Rev., 4, 1940, 275) has out-
lined a classification of the vibrios based
upon their protein and polysaccharide struc-
tures. Using chemical methods, it was found
that one polysaccharide and one protein
was commonly obtained from each strain
of vibrio; when exceptions occurred, it was
invariably noted that the strain was under-
going dissociation. Given a single protein
and polysaccharide in each vibrio, it was
possible to divide the strains into six groups,
which were numbered in the order of their
discovery as shown in the table.
A chemical grouping of the cholerigenic
and water vibrios.
Group
Protein Type
Polysaccharide
Type
I
I
I
II
I
II
III
II
II
IV
II
I
V
II
III
VI
I
III
The strains of Groups I and II possess
the same protein and different polysaccha-
FAMILY VII. SPIRILLACEAE
233
Cholera group of vibrios.
(Biochemically similar. Common H antigen.)
0-sub-group I,
Non-hemolytic
(goat cells).
Cholera vibrios.
Types — original ,
variant and mid-
dle.
Hemolytic (goat
cells) .
El Tor vibrios.
Types — original
and variant (?mid-
dle).
I
O sub-groups II, III, IV, V, VI and indi-
vidual races (mostly hemolytic). Para-
cholera, cholera-like, and some El Tor
vibrios.
(Types within sub-groups underlined.)
rides. These are derived from cases of chol-
era and have the serological and biochemical
characteristics of 0-Group I, Vibrio chol-
era. Group I strains are more common than
those of Group II, which have, however,
been isolated from epidemics with a high
mortality. The phospholipid fraction is
common to both tj^pes when isolated in the
early part of an epidemic but is not found
in strains of other groups. The harmless
water vibrios, which are so heterogeneous
serologically (Taylor and Ahuja, Indian
Jour. Med. Res., 26, 1938, 8-32), form a
single chemical group with a homogeneous
structure. They fall into Group III, which
differs in its protein structure from the
authentic cholera vibrios and which re-
sembles Group II in its polysaccharide. The
vibrios of Group IV, which came from El
Tor and from chronic vibrio carriers, are
believed, on epidemiological grounds, to
be harmless, although serological methods
have failed to distinguish them from chol-
erigenic vibrios. Group V, which, like III
and IV, contains protein II, consists, like
Group IV, of strains from chronic vibrio
carriers. Group VI strains are only rarely
isolated in nature, and representatives of
this group are generally found among col-
lections of old laboratory strains. They
appear to be the result of polysaccharide
variation from Group I after long-con-
tinued growth on artificial media.
2. Vibrio berolinensis Xeisser, 1893.
(Arch. f. Hyg., 19, 1893, 200.)
be.ro.li.nen'sis. M.L. Berolinum place
name, Berlin; M.L. adj. berolinensis of
Berlin.
Curved rods, somewhat smaller than those
of Vibrio comma, frequently occurring in
pairs. Pleomorphic. Motile by means of a
single, polar flagellum. Gram-negative.
Gelatin colonies: Small, grayish, slightly
granular, fragmented; very slow liquefac-
tion.
Gelatin stab: Slow, napiform liquefaction.
Agar slant: Grayish yellow, moist, glis-
tening growth.
Broth: Turbid, with gray pellicle.
Litmus milk: No coagulation, no acid.
Potato: Brownish streak.
Indole produced.
Nitrites produced from nitrates.
Not pathogenic for mice, pigeons or
guinea pigs.
Aerobic, facultative.
Optimum temperature, 37° C. Minimum,
above 10° C. Maximum, less than 60° C.
Source: Isolated from filtered Spree river
water.
Habitat: Presumably widely distributed
in polluted water.
3. Vibrio metschnikovii GamaMia, 1888.
{Vibrio metschnikovi (sic) Gamaleia, Ann.
Inst. Past., 2, 1888, 482.)
metsch.ni.ko'vi.i. Named for Metsch-
nikoff, a Russian bacteriologist; M.L. mas.
gen.n. metschnikovii of Metschnikoff.
Curved rods, somewhat shorter and
thicker than those of Vibrio comma. Long,
234
ORDER I. PSEUDOMONADALES
slender chains of cells are formed in old
cultures. Motile by means of a single, polar
flagellum. In the animal body the cells are
nearly coccoid. Gram-negative.
Gelatin colonies: Like those of Vibrio
comma.
Gelatin stab: Rapid, napiform liquefac-
tion.
Agar slant: Yellowish, plumose, moist,
glistening growth.
Broth: Turbid, with thin, white pellicle.
Litmus milk: Acid, coagulated (eighth
day) ; not peptonized.
Potato: Delicate, brownish growth.
Indole produced.
Nitrites produced from nitrates.
Aerobic, facultative.
Optimum temperature, 37° C. Maximum,
less than 45° C.
Pathogenic for pigeons, fowls and guinea
pigs.
Source: Isolated from fowl dead of a chol-
era-like disease.
Habitat: Found in the intestinal contents
of chickens, pigeons and other animals suf-
fering from a cholera-like disease.
4. Vibrio proteus Buchner, 1885. (Kom-
mabacillus der cholera nostras, Finkler and
Prior, Deutsche med. Wochenschr., 1884,
632; Buchner, Sitzungsber. d. Gesel. f.
Morph. u. Physiol., Miinchen, Heft 1, 1885,
10.)
pro'te .us. Gr. noun Proteus a sea-god who
could change his form; M.L. mas.n. Proteus
a generic name.
Description supplemented by material
taken from Lehmann and Neumann (Bakt.
Diag., 7 Aufl., 2, 1927, 541).
Curved rods, 0.4 to 0.6 by 2.4 microns,
often pointed at both ends. Motile by means
of a single, polar flagellum. Gram-negative.
Gelatin colonies: Small, gray, circular,
granular, entire; rapid liquefaction with
the formation of large craters.
Gelatin stab: Rapid, saccate liquefaction.
Agar slant: Dirty grayish, plumose
growth.
Broth: Turbid, with fetid odor.
Litmus milk: Slightly acid; coagulated;
peptonized.
Potato: Grayish, slimy layer.
Indole not produced; indole reaction weak
(Lehmann and Neumann).
Hydrogen sulfide production very slight.
Gas not produced from glucose.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 30° C.
Source: Isolated from feces of patients
suffering from cholera nostras (gastroen-
teritis) .
Habitat: Found in the intestinal contents
in cholera nostras and cholera infantum.
5. Vibrio sputigenus (Migula, 1900)
Bergey et al., 1923. (Vibrio aus Sputum,
Brix, Hyg. Rundschau, 4, 1894, 913; Micro-
spira sputigena Migula, Syst. d. Bakt., 2,
1900, 981; Bergey et al.. Manual, 1st ed.,
1923, 80.)
spu.ti'ge.nus. L. noun sputum spit, spu-
tum; Gr. v. gennao to bear; M.L. adj. sputi-
genus sputum-borne.
Slightly curved rods, about the same
size and form as those of Vibrio comma,
occurring singly, occasionally three or four
in a chain. Motile by means of a single, polar
flagellum. Gram-negative.
Gelatin colonies: Small, circular, slightly
granular, yellowish, becoining brownish.
Gelatin: Crateriform liquefaction.
Agar slant: Grayish white, moist.
Broth: Turbid, no pellicle formed.
Litmus milk: Acid; coagulated.
Potato: Thin, gray layer, spreading.
Indole not produced.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 37° C.
Source: Isolated from sputum.
6. Vibrio xenopus Schrire and Green-
field, 1930. (Trans. Roy. Soc. So. Africa,
17, 1930, 309.)
xe'no.pus. Gr. adj. xenus alien, strange;
Gr. noun pus, podis a foot; M.L. mas.n.
Xenopus strange foot, a genus of toads.
Spiral forms, occurring singly and in
pairs. Non-motile. Gram-negative.
Gelatin stab: Slow, crateriform liquefac-
tion.
Agar colonies: Small, white, glistening,
slimy, entire.
FAMILY VII. SPIRILLACEAE
235
Agar slant: Grayish white, slimy, entire
growth.
Broth: Turbid with flocculent sediment.
Litmus milk: Unchanged.
Potato: Not reported.
Indole not produced.
Blood serum peptonized.
Starch not hydrolyzed.
Acid from glucose, fructose, maltose,
glycerol and sorbitol.
Nitrites produced slowly from nitrates.
Aerobic, facultative.
Optimum temperature, 37° C.
Source: Isolated from an abscess of the
pectoral muscle of an African toad.
7. Vibrio leonardii Mdtalnikov and
Chorine, 1928. (Ann. Inst. Past., Ji2, 1928,
1647.)
le.o.nar'di.i. M.L. gen. noun leonardii of
Leonard; named for A. G. Leonard.
Curved rods with rounded ends, 0.5 to
1.0 by 2.0 to 3.0 microns. Motile by means
of 1 to 3 polar flagella. Gram-negative.
Gelatin stab: No liquefaction.
Agar colonies: Small, transparent, circu-
lar, having a characteristic odor.
Broth: Turbid, with thin pellicle.
Litmus milk: No coagulation, acid, with
reduction of litmus.
Potato: Slight, colorless growth.
Coagulated blood serum not liquefied.
Indole not produced.
Hydrogen sulfide produced.
Acid and gas from glucose, fructose, ga-
lactose, lactose, sucrose and mannitol. No
acid or gas from maltose or glycerol.
Nitrites produced from nitrates.
Aerobic, facultative.
Optimum temperature, 30° C.
Habitat: Highly pathogenic for insects
such as Galleria mellonella L. (wax moth)
and Pyrmista nubialis Hiibn. (European
corn borer).
8. Vibrio niarinopraesens ZoBell and
Upham, 1944. (Bull. Scripps Inst, of Ocean-
ography, Univ. Calif., 5, 1944, 256.)
ma.ri .no.prae'sens. L. adj. marinus of the
sea; L. part. adj. praesens present; M.L.
adj. marinopraesens present in the sea.
Curved rods, 0.4 to 0.5 by 1.2 to 2.4 mi-
crons, occurring singly and in spiral chains.
Polar staining. Motile by means of one or
two polar flagella. Gram-negative.
Note: All differential media except the
fresh-water broth, litmus milk and potato
were prepared with sea water.
Gelatin colonies: Circular, 1 mm in di-
ameter, dense center, brown discoloration
of gelatin.
Gelatin stab: Stratiform above, infundi-
buliform below; complete liquefaction in
5 days; brown discoloration of gelatin.
Agar colonies: Convex, circular, 0.5 mm
in diameter, entire, translucent.
Agar slant: Abundant, filiform, glisten-
ing, butyrous growth with no pigment.
Sea-water broth: Heavy turbidity; slight
viscid sediment; surface ring.
Fresh-water broth: No visible growth.
Litmus milk: Completely decolorized.
Potato: No visible growth.
Indole not produced.
Hydrogen sulfide rapidly produced.
Acid but no gas from glucose and maltose.
Glycerol, xylose, lactose, sucrose, mannitol
and salicin not fermented.
Starch is hydrolyzed.
Non-lipolytic.
Nitrites not produced from nitrates.
Ammonia produced from peptone but not
from urea.
Casein not digested.
Aerobic, facultative.
Optimum temperature, between 20° and
25° C.
Source: Isolated from sea water.
Habitat: Common; probably widely dis-
tributed.
9. Vibrio phytoplanktis ZoBell and
Upham, 1944. (Bull. Scripps Inst, of Ocean-
ography, Univ. Calif., 5, 1944, 261.)
phy.to.plank'tis. Gr. neut.n. phylum
plant; Gr. adj. plancius wandering; M.L.
neut.n. plankton (plancium) plankton; M.L.
neut.n. phytoplankton plant plankton; M.L.
adj. ^phytoplanktis of the phytoplankton.
Curved rods, 0.5 to 0.6 by 2.0 to 5.4 mi-
crons, occurring mostly singly with some
short spiral chains. Bipolar staining. Motile
by means of a single polar flagcllum. Gram-
negative.
Note: All differential media except the
236
ORDER I. PSEUDOMONADALES
fresh-water broth, litmus milk and potato
were prepared with sea water.
Gelatin colonies: Diffuse, irregular; waxy
appearance, slightly depressed; rapid lique-
faction.
Gelatin stab: Slow, crateriform, lique-
faction becoming stratiform. Buff pigment.
Agar colonies: 1 to 2 mm in diameter,
translucent, smooth, convex, circular.
Agar slant: Luxuriant, echinulate; water.y
appearance; slightly mucoid, glistening
growth with buff or cream pigment.
Sea-water broth: Heavy turbidity; abun-
dant, flocculent sediment; surface ring.
Fresh-water broth: No visible growth.
Litmus milk: No visible change.
Potato: No visible growth.
Indole not produced.
Hydrogen sulfide is produced.
Acid but no gas from glucose, maltose
and sucrose. Glycerol, xylose, lactose, man-
nitol and salicin not fermented.
Starch not hydrolj^zed.
Non-lipolytic.
Nitrites not produced from nitrates.
Ammonia produced from peptone but
not from urea.
Casein is digested.
Aerobic, facultative (good anaerobic
growth).
Optimum temperature, between 20° and
25° C.
Source: Lsolated from sea water and ma-
rine phytoplankton.
Habitat: Presumably widely distributed.
10. Vibrio costicolus Smith, 1938. (Roy.
Soc. Queensland, Proc. for 1937, 49, 1938,
29.)
cos.ti'co.lus. L. noun, cos/o rib; L. v. colo
to dwell; M.L. adj. costicolus rib dwelling
(from bacon).
Curved rods, 0.5 by 2.0 to 4.0 microns;
old cells coccoid. Actively motile by means
of a single, polar flagellum. Young cultures
show pronounced beaded staining. Gram-
negative.
No growth on media which does not con-
tain salt. Limit for growth, 2 to 23 per cent
NaCl; optimum, 6 to 12 per cent.
Gelatin stab: No liquefaction. However,
some strains liquefy within 2 days at 32° C;
these may represent a distinct variety or a
separate species.
Agar colonies: Circular, entire, convex,
glistening, non-viscid.
Agar slant: Abundant, filiform, trans-
parent or translucent growth.
Broth : Pellicle formation varies from
absent to pronounced, whitish and non-
coherent.
Litmus milk: Not coagulated.
Potato: Sparse, moist, brownish growth.
Indole not produced.
Hj^drogen sulfide produced.
Acid from glucose, fructose, sucrose, man-
nose, mannitol and glycerol. No acid from
galactose, lactose, maltose, rhamnose, raf-
finose, arabinose, xylose, sorbitol, dextrin,
starch or salicin.
Acetjdmethylcarbinol not produced.
Non-lipolytic.
Nitrites produced from nitrates.
Catalase-positive.
Aerobic, facultative.
Optimum temperature, between 30° and
35° C; temperature range, 2° to 42° C.
Related organisms: Robinson (A Possible
Explanation of Microbial Halophilism, The-
sis, McGill University, 1950, 92 pp.) isolated
a similar organism from bacon-curing brines
in Canada. At concentrations of 11.7 and
17.5 per cent NaCl, cells are spirillum-
shaped and sluggishly motile. Pellicle formed
on broth. Gelatin liquefied. Acetjdmeth-
ylcarbinol produced. Catalase and urease
absent. Acid from raffinose and inulin. No
acid from mannose, dulcitol, cellobiose,
adonitol or ethyl alcohol. Organism will
note grow in absence of salts, but NaCl may
be replaced by KCl, NaBr, NaoS.Oa , LiCl
or MgCU (also see Flannery, Doetsch and
Hansen, Jour. Bact., 64, 1952, 713-17).
Source: Isolated from the tainted ribs
of bacon and tank brines in bacon factories
in Australia.
Habitat: Found in cured meats and meat-
curing brines.
11. Vibrio halonitrificans* Smith, 1938.
* This organism is, in realitj-, a denitrifier,
inappropriate.
not a nitrifier, and therefore this name
FAMILY VII. SPIRILLACEAE
237
(Roy. Soc. Queensland, Proc. for 1937, 49,
1938, 29.)
ha.lo.ni.tri'fi.cans. Gr. noun hals , hairs
the sea salt; M.L. part. adj. nitrificans ni-
trifying; M.L. adj. halonitrificans nitrifying
salt.
Curved rods, usually 0.3 by 1.2 to 2.5 mi-
crons, occurring singly. Motile by means of
a single, polar flagellum. No marked varia-
tion in form in media of varied salinity.
Stain somewhat faintly with the usual
stains. Gram-negative.
No growth on media which does not con-
tain salt. Limit for growth, 1 to 23 per cent
salt; optimum growth in 4.0 to 6.0 per cent
salt.
Gelatin stab: Liquefied within 7 days at
35° C.; at 20° C. shallow, superficial lique-
faction was evident in 20 days.
Agar colonies: Light amber, glistening,
convex, transparent, non-viscid, slightly
spreading.
Agar slants: Growth slow at 4° C., col-
onies appearing in 14 days.
Nutrient and nitrate broths: Growth.
No growth, however, when covered with a
paraffin oil seal.
Litmus milk: Not coagulated; growth
slight or absent.
Potato: Growth moist, fairly abundant,
whitish.
Indole not produced.
Hydrogen sulfide not produced.
Glucose, fructose, sucrose, mannose,
rhamnose, galactose, lactose, maltose, raf-
finose, sorbitol and glycerol not fermented.
Acetylmethylcarbinol not produced.
Non-lipolytic.
Nitrites produced from nitrates.
Catalase-negative.
Aerobic.
Temperature relations: Optimum, be-
tween 30° and 35° C. Slow growth at 4° C.
Killed in 10 minutes in 6 per cent saline
broth at 55° C.
Limiting reactions for growth, pH 5.4
and pH 9.2.
Not pathogenic for guinea pigs or mice.
Source: Five strains were isolated from
tank brines from bacon-curing factories
in Australia. The strains showed little var-
iation in characters. Except for its ability
to liquefy gelatin, this species resembles
the organisms isolated by Sturges and
Heideman (Absts. Bact., 7, 1923, 11; ibid.,
8, 1924, 14; ibid., 9, 1925, 2) in the U. S. A.
Habitat: Known to be found in meat-
curing brines but probably more widely
distributed.
12. Vibrio luminosus Beijerinck, 1888.
(Vibrio luminosus (nomen nudtim) Beije-
rinck, Botan. Zeitg., 46, 1888, 763; Photo-
bacterium luminosum Beijerinck, Arch.
N^erl. d. Sci. Exact, et Natur., 23, 1889, 401 ;
Microspira luminosa Migula, Syst. d. Bakt.,
2, 1900, 1015.)
lu.mi.no'sus. L. adj. luminosus Inminons.
Small rods having the size and form of the
cholera vibrio when grown in media con-
taining little nitrogen and carbohydrates.
Sometimes form chains of vibrios which
resemble spirilla. In richer media the cells
become much larger. Motile. Gram-nega-
tive (Chester, 1897).
Gelatin: Liquefied. In presence of 0.5 per
cent asparagine and 0.5 per cent peptone,
offensive odors not produced. Putrefaction
of the gelatin occurs when the nitrogen
source is insufficient.
Peptonized meat bouillon gelatin: No
growth. Good growth and luminescence
with the addition of 3.0 to 3.5 per cent of
sea salt, potassium chloride or magnesium
chloride.
Agar: Growth rapid, shines feebly.
Sea-water broth: Produces forms which
resemble the bacteroids of legume bacteria.
Blood serum: No growth. Addition of 3.0
to 3.5 per cent of sea salt, potassium chloride
or magnesium chloride allows good growth
and luminescence.
Nitrates not reduced.
Indigo-blue not readily reduced.
Salt tolerance: In order to assure phos-
phorescence and good growth, the osmotic
tension of inorganic salt solutions used for
cultivation should be equivalent to that
produced in a 3 per cent sodium chloride
solution.
Optimum temperature for growth and
luminescence, between 25° and 28° C.
Aerobic.
Quality of luminescence: Bluish green,
persisting for 1 to 2 weeks.
Distinctive characters: Develops only
238
ORDER I. PSEUDOMONADALES
on neutral or f eebl}'^ alkaline media : a slight
quantity of acid completely prevents growth
and the production of luminescence. Lu-
minescence occurs on organic matter only
when a sufficient proportion of inorganic
salt is present.
Source: Isolated in Holland from coastal
sea water, dead sea fish and Crustacea.
Habitat: Found in coastal sea water, on
dead fish, Crustacea and other salt-water
animals, on meat and even on soldiers'
wounds where they produce no known harm-
ful effects. No food poisoning has ever been
traced to meat on which these organisms
have grown (Niven, Circular No. 2, Amer-
ican Meat Inst. Foundation, 1951, 1-11).
13. Vibrio indicus (Beijerinck, 1889)
Lehmann and Neumann, 1896. (Bacillus
phosphorescens Fischer, Ztschr. f. Hyg., S,
1887, 58; also see Anonymous, Sitzber. d.
Gesell. naturf. Freunde zu Berlin, 1886,
162; Photobacterium indicum Beijerinck,
Arch. Need. d. Sci. Exact, et Natur., 23,
1889, 401; not Photobacterium phosphores-
cens Beijerinck, loc. cit.; Lehmann and
Neumann, Bakt. Diag., 1 Aufl., 2, 1896,
341; Pseudomonas phosphorescens Bergej'
et al.. Manual, 3rd ed., 1930, 177.)
in'di.cus. L. adj. indicus of India.
Description taken from Fischer (op. cit.,
1887, 58) and Beijerinck (op. cit., 1889, 401).
Small, thick rods 2 to 3 times as long as
wide with rounded ends; occasionally spiral
and short, irregularly-curved filamentous
forms are found. Motile. Stain lightly with
aniline dyes. Gram-negative (Chester, 1897).
Gelatin colonies: After 36 hours, small,
circular, grayish white, punctiform. Lique-
faction, forming a slightly concave surface.
Blood serum: Grayish white, slimy
growth.
Potato: Thin, white laj'er in 2 to 3 days.
Cooked fish: Abundant growth. Entire
surface covered with a grayish white, slimy
growth. Bluish white phosphorescence.
Alkaline broth: Slight turbidity in 24
hours. Pellicle in 3 days.
Acid broth: No turbidity. No phospho-
rescence.
Milk: No growth.
No gas produced.
Nitrates not reduced.
Indigo-blue not readily reduced.
Not pathogenic for laboratory animals.
Salt tolerance: To assure phosphorescence
and good growth, the osmotic tension of
inorganic salt solutions used for cultivation
should be equivalent to that produced in a
3 per cent sodium chloride solution.
Optimum temperature for growth and
luminescence, between 30° and 32° C. Min-
imum, 15° C.
Aerobic.
Quality of luminescence: Bluish to green,
persisting for 1 to 2 weeks.
Distinctive character: Luminescence on
organic matter occurs only when a sufficient
proportion of inorganic salt is present.
Source: Isolated from sea water of the
West Indies.
Habitat: Found in coastal sea water and
on dead fish, Crustacea and other salt-water
animals; they are also found on meat and
even on soldiers' wounds where they pro-
duce no known harmful effects. No food
poisoning has been traced to meat on which
these organisms have grown (Niven, Cir-
cular No. 2, American Meat Inst. Founda-
tion, 1951, 1-11).
14. Vibrio albensis Lehmann and Neu-
mann, 1896. (Elbe vibrio, Dunbar, Deutsche
med. Wochnschr., 19, 1893, 799; Lehmann
and Neumann, Bakt. Diag., 1 Aufl., 2, 1896,
340; Microspira dunbari Migula, Syst. d.
Bakt., 2, 1900, 1013.)
al.ben'sis. M.L. adj. aZftensj's pertaining
to the (river) Elbe.
Early descriptions merel.y report this
organism as morphologically and culturally
(except for phosphorescence and patho-
genicity) indistinguishable from Vibrio
comma. Some of the early workers even
failed to observe phosphorescence. Descrip-
tion taken from Gorham (in Dahlgren, Jour.
Franklin Inst., 180, 1915, table following
714) and Warren (Jour. Bact., 49, 1945,
549); also see Sonnenschein (Cent. f. Bakt.,
I Abt., Orig., 123, 1931, 92).
Curved rods, 1.2 by 2.1 microns, occurring
singly and in pairs. Motile b}^ means of a
single, polar flagellum. Not encapsulated.
Gram-negative.
FAMILY VII. SPIRILLACEAE
239
Gelatin colonies: Small, j^ellowish white.
Gelatin stab: Liquefaction. Growth at
the surface and along the stab.
Agar: Abundant growth.
Agar slant: Growth dull and wrinkled.
Blood agar: Good growth and lumines-
cence; beta hemolysis.
Broth: Pellicle formed.
Koser's citrate medium: Growth and lu-
minescence.
Milk: Growth.
Potato: Luxuriant growth.
Indole produced.
Hydrogen sulfide not produced.
Acid but no gas from glucose and sucrose.
No acid or gas from lactose.
Starch hydrolyzed.
Gives a cholera-red reaction, i.e., pro-
duces both indole and nitrites.
Nitrites produced from nitrates.
Optimum salt concentration, 0.9 per cent.
Temperature relations: Optimum, 22° C.;
growth at 37.5° C.
Aerobic, facultative.
Distinctive characters: Morphologically
and culturally like Vibrio comma. Lumi-
nescent. Pathogenic to guinea pigs and
pigeons.
Source: Originally isolated from the Elbe
River. If Vibrio phosphorescens Jermoljewa
(Cent. f. Bakt., I Abt., Orig., 100, 1926,
170) is accepted as identical with this spe-
cies, then it has also been found in the in-
testinal contents of three cholera patients,
one gastroenteritis and one typhoid pa-
tient; Jermoljewa (ibid., 171) also isolated
his organism from the bile of a cadaver.
Sonnenschein (op. cit., 12S, 1931, 92) reiso-
lated this species from a fish taken from
the Elbe River and found that it main-
tained its ability to luminesce when grown
in o.x bile.
Habitat: Found in fresh water, in human
feces and in bile. Probably widely distrib-
uted.
15. Vibrio pierantonii (Zirpolo, 1918)
Meissner, 1926. (Bacillus pierantonii Zir-
polo. Boll. Soc. Nat. Napoli, 30, (1917)
1918, 206; Meissner, Cent. f. Bakt., II Abt.,
67, 1926, 200.)
pier.an.to'ni.i. M.L. gen. noun pierantonii
of Pierantoni; named for Prof. U. Pieran-
toni, an Italian scientist.
Rods, 0.5 by 1.5 microns, with rounded
ends. Rods curved and vibrio-shaped ac-
cording to Meissner (ibid., 201). Motile by
means of one to three polar flagella. Gram-
negative.
Gelatin colonies: Circular and irregularly
lobulate.
Gelatin stab: No liquefaction.
Agar colonies: Circular, light green,
smooth, entire.
Glycerol agar slant: Slightly luminous
streak.
Broth: Turljid, with pellicle.
Indole not produced.
Acid from glucose and maltose. Some
strains also attack lactose, sucrose and
mannitol.
Best growth in alkaline media.
Aerobic, facultative.
Optimum temperature, 37° C.
Source: Isolated from the photogenic
organ of the cephalopod Sepiola intermedia
Naef.
16. Vibrio agarliqiiefaciens (Gray and
Chalmers, 1924) Bergey et al., 1934. (Mi-
crospira agar-liquefaciens (sic) Gray and
Chalmers, Ann. Appl. Biol., 11, 1924, 325;
Bergey et al., Manual, 4th ed., 1934, 119.)
a.gar.li.que.fac'i.ens. Malay agar, a jelly
from seaweeds; L. v. liquefacio to liquefy;
M.L. part. adj. agarliquefaciens liquefying
agar.
Short, curved rods, usually c-shaped,
with occasional s-shaped and coccoid forms.
Cells 2.0 microns long by 0.5 to 0.7 micron
broad; 3.0 to 5.0 microns long in division
stages. Coccoid forms stained, 0.5 to 0.7
micron long. Motile by means of a single,
polar flagellum. Gram stain not reported.
Gelatin stab: Very slight surface growth
after one month; the streak then shows a
beaded line. No liquefaction.
Agar colonies: Surface colonies appear
as a whitish growth in a depression, sur-
rounded by a white ring. The colony is later
surrounded by a ring of liquid agar. Deep
colonies show a clear area and may be irreg-
ular, oval or angular.
Agar slant: A deep groove is cut along
240
ORDER I. PSEUDOMONADALES
the inoculation streak, whitish growth along
sides. The gel is later much weakened.
Broth: Slightly turbid. No pellicle.
Acid from glucose, lactose and maltose.
No acid from sucrose or glycerol.
Starch hydrolyzed.
Decomposes cellulose and agar. The pres-
ence of one per cent glucose prevents the
liquefaction of agar.
Nitrites produced from nitrates.
Utilizes ammonium salts as a source of
nitrogen.
Aerobic.
Temperature relations: Optimum, 25° C;
will grow at 16° but not at 34° C.
Habitat: Soil.
17. Vibrio andoii Aoi and Orikura, 1928.
(Eine neue Agarzersetzende Bodenbak-
terienart, Aoi, Cent. f. Bakt., II Abt., 63,
1924, 30; Aoi and Orikura, Cent. f. Bakt.,
II Abt., 74, 1928, 331.)
an.do'i.i. M.L. gen .no\in andoii of Andoi;
named for Andoi, a Japanese scientist.
Curved rods with more or less tapering
ends, c- or s-shaped, 0.5 to 0.8 by 1.5 to 2.5
microns. Motile by means of a single, polar
flagellum. Gram-negative.
Gelatin: No growth.
Peptone agar media: No growth.
Peptone broth: No growth.
Litmus milk: No growth.
Potato: No growth.
Ammonium sulfate agar colonies: Puncti-
form, circular, concave, surrounded with
a clear zone.
Ammonium sulfate agar slant: Growth
grayish, becoming straw-yellow, sinking
into the medium as the agar liquefies.
Glucose, fructose, galactose, mannose,
xylose and "konjac" assimilated. Konjac,
a Japanese food in tablets and strips, re-
sembles gelatinized agar; it is prepared from
the tuber of the konjac plant, Amor-pho-
phallus rivieri.
Starch hydrolyzed.
Cellulose and lignin not attacked.
Xylan decomposed.
Cellobiose decomposed.
Aerobic, facultative.
Temperature relations: Optimum, be-
tween 25° and 28° C. Minimum, 8° C. Max-
imum, 37° C.
Optimum pH, between 6.8 and 7.5.
Distinctive characters : When grown sym-
biotically with a second, unnamed species
found in rotted manure, the latter species
is able to hydrolyze cellulose in straw, prob-
ably because the first species (Vibrio andoii)
decomposes the xylan that protects the
cellulose from the action of the second spe-
cies.
Source: Isolated from rotted stable ma-
nure.
Habitat: Presumably decomposing or-
ganic matter.
18. Vibrio beijerinckii Stanier, 1941.
(Tyrosine vibrio of the sea, Beijerinck, Proc.
Sect. Sci., Kon. Akad. Vetenschappen,
Amsterdam, 13, 1911, 1072; Stanier, Jour.
Bact., 42, 1941, 539.)
beij .er.inck'i.i.M.L. gen. nounbeijerinckii
of Beijerinck; named for Prof. M. W. Beije-
rinck, the Dutch biologist who first dis-
covered this species.
Small, curved rods, 0.4 to 1.0 by 2.0 to
6.0 microns, usually single, sometimes oc-
curring in short chains; in older cultures,
occur mostly as straight rods. Actively
motile by means of polar flagella. Encap-
sulated. Gram-negative.
Sea-water peptone agar colonies : Round,
smooth, glistening, mucoid, entire. White
to gray in color. After 24 hrs, 3 to 4 mm in
diameter. The agar softens and clears for
a distance of 3 to 5 mm from the edge of the
colony, the outer edge of the gelase field
being sharply defined. The colonies eventu-
ally grow to as much as 10 mm in diameter
with a gelase margin of 2 to 3 cm.
Sea-water nitrate agar: Growth is slower
than with peptone, but pigment production
is much more marked. After 48 hours, col-
onies are 1 mm in diameter with a dark
brown to black center and a colorless mar-
gin. Pigmented granules may be seen lying
among the cells.
Sea-water peptone agar slant: Abundant
growth after 24 hours, spreading, slightly
raised, smooth, glistening, mucoid, dirty-
white to dark gray in color. Agar digestion
is evidenced onlj' by a general softening of
the slant. After several days, a pale brown,
diffusible pigment is produced by some
strains.
FAMILY VII. SPIRILLACEAE
241
Sea-water nutrient gelatin slant: Good,
filiform, gray growth after 24 hours, with
considerable liquefaction. Slant completely
liquefied after one week.
Sea-water nutrient gelatin stab: Fair,
filiform growth after 24 hours, best at sur-
face. Napiform liquefaction, complete after
7 to 10 days.
Sea-water peptone broth: Heavily turbid
after 24 hours. Gray pellicle and flocculent;
gray sediment. Later a light brown, soluble
pigment is formed.
Indole not produced.
Hydrogen sulfide not produced.
Very slight or no acid from glucose, ga-
lactose, maltose, lactose and cellobiose.
Arabinose, xylose and sucrose not fer-
mented. Agar is extensively softened but
not liquefied. Cellulose, chitin and alginic
acid not attacked.
Starch is rapidly hydrolyzed.
Nitrites produced from nitrates.
Ammonia and nitrates utilized as sole
sources of nitrogen.
Urease-negative.
Catalase-positive.
Aerobic.
Optimum temperature, 23° C.; grows
between 5° and 30° C.
Salt range: 0.25 to 6.0 per cent. Optimum,
between 2.0 and 4.0 per cent.
Source: Found in sea-water and, in the
winter months, in the plankton; also found
in fresh water and in sewage. Isolated both
in Holland and in California. Along the
coast of California it appears to be the most
common species of marine agar-digester.
Habitat: Widely distributed in sea water
and also in fresh water.
19. Vibrio fuscus Stanier, 1941. (Jour.
Bact., 4^, 1941,540.)
fus'cus. L. adj. fiiscus dark or tawny.
Small, slightly curved rods, 0.7 by 1.5 to
5.0 microns, usually occurring singly, some-
times in short chains. Very actively motile
by means of a single, polar flagellum. Not
encapsulated. Gram-negative.
Sea-water peptone agar colonies: 1 mm
in diameter after 48 hours; round, smooth,
glistening, translucent, entire, pale ^yellow
and slightly sunken in the agar. Colonies
several mm in diameter after 7 days, be-
coming bright yellow, then pale brown.
They are sharply sunken into the agar and
are surrounded b}' a narrow, sharply de-
fined gelase field. Liquefaction does not
occur except on heavily seeded plates.
Sea-water peptone agar slant : Fair growth
after 48 hours, filiform, smooth, glistening,
translucent, pale yellow, slightly sunken
in the agar. Later a pale yellow, diffusible
pigment may be produced, and the streak
tends to become light brown in color. On
old slants the agar is slightly liquefied.
Sea-water nutrient gelatin slant: Fili-
form, smooth, pale j-ellow growth after 48
hours with slight liquefaction; liquefaction
almost complete after 7 days.
Sea-water gelatin stab: Filiform growth
after 48 hours with slight liquefaction; col-
orless; growth best at surface. Later the
liquefaction becomes stratiform and almost
complete.
Sea-water peptone broth: Good growth
after 48 hours; turbid with a granular sedi-
ment and yellow pellicle.
Indole not produced.
Hydrogen sulfide not produced.
Glucose, galactose, sucrose, maltose,
lactose, xylose and cellobiose attacked.
Arabinose not utilized. Cellulose is attacked
to a slight extent, and agar is softened and
sometimes liquefied. Chitin and alginic acid
are not attacked.
Starch not hydrolyzed.
Nitrites produced from nitrates.
Urease-negative.
Catalase-positive.
Aerobic.
Optimum temperature, between 20° and
25° C. Grows between 5° and 30° C.
Salt range, 1.0 to 5.0 per cent. Optimum,
between 2.0 and 4.0 per cent.
Source: Found only once in a marine cel-
lulose-enrichment culture in California.
Habitat: Presumably salt water.
20. Vibrio granii (Lundestad, 1928)
Stanier, 1941. (Bacterium granii Lundestad,
Cent. f. Bakt., II Abt., 75, 1928, 330; Stan-
ier, Jour. Bact., 42, 1941, 538.)
gra'ni.i. M.L. gen. noun granii of Gran;
named for Prof. H. H. Gran, who first de-
tected agar-liquef3'ing bacteria.
Rods, 0.6 to 0.8 by 1.4 to 2.4 microns, with
242
ORDER I. PSEUDOMONADALES
rounded ends, occurring singly, in pairs,
and at times in short chains. Motile. Polar
flagellate (Stanier, loc. cit.). Gram-negative.
Fish-gelatin colonies: Punctiform, black,
glistening.
Fish-gelatin stab: Slow, crateriform lique-
faction.
Sea-weed agar colonies: Circular, flat,
opaque, glistening, white, slimy, entire.
Agar is dissolved.
Fish-agar slant: Flat, white, elevated,
glistening, undulate growth. Liquefaction.
Broth: Turbid with grayish white, slimy
sediment.
Indole not produced.
No action on sugars.
Starch usuallj^ hydrolyzed.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, between 20° and
25° C. Minimum, between 0° and 5° C. Max-
imum, between 30° and 32° C.
Source: Isolated from sea-water of the
Norwegian Coast.
Habitat: Presumably found in sea water
and on sea weeds.
22. Vibrio cyclosites Gray and Thorn-
ton, 1928. (Cent. f. Bakt., II Abt., 73, 1928,
92.)
cyc.lo.si'tes. Gr. nonn ajclus a ring: Gr.
V. sited to eat; M.L. adj. cyclosites feeding
on rings, i.e., on ring compounds.
Curved rods 0.5 to 1.0 by 1.5 to 4.0 mi-
crons. Motile by means of a single, polar
flagellum. Gram-negative.
Gelatin colonies: Circular, buff to brown,
flat, smooth, glistening, entire.
Gelatin stab: No liquefaction.
Agar colonies: Circular to irregular, pale
buff (later greenish), smooth, entire.
Agar stab: Filiform, greenish buff, raised,
smooth, undulate.
Broth: Turbid.
Indole not reported.
Acid from glucose.
Starch not hydrolj'zed.
Nitrites not produced from nitrates.
Attacks phenol and ?«-cresol.
Aerobic, facultative.
Optimum temperature, between 30° and
35° C.
Habitat: Soil.
21. Vibrio neocistes Gray and Thorn-
ton, 1928. (Cent. f. Bakt., II Abt., 73, 1928,
92.)
ne.o.cis'tes. Gr. adj. nensnev;; Gr. noun
ciste box; M.L. fern. gen. n. neocistes of New-
ark, a city.
Curved rods 0.5 to 1.0 by 1.0 to 4.0 mi-
crons. Motile by means of one to three polar
flagella. Gram stain not recorded.
Gelatin colonies: Liquefied.
Gelatin stab: Liquefied. Medium red-
dened.
Agar colonies: Circular or amoeboid, buff
to brownish, convex, smooth, glistening,
entire.
Agar slant: Filiform, fluorescent, raised,
smooth, glistening, undulate.
Broth: Turbid.
Acid from glucose.
Starch not hydrolyzed.
Nitrites not produced from nitrates.
Attacks naphthalene.
Aerobic, facultative.
(Optimum temperature, between 30° and
35° C.
Habitat: Soil.
23. Vibrio oxaliticus Bhat and Barker,
1948. (Jour. Bact., 55, 1948, 359.)
ox.a.li'ti .cus. Gr. noun oxalis sorrel, a
sour plant; Gr. adj. lyticus dissolving; M.L.
adj. oxaliticus intended to mean decom-
posing oxalate.
Curved rods 0.4 by 1.3 microns. Actively
motile by means of a single, polar flagellum.
Not encapsulated. Gram-negative.
Nutrient agar colonies: Small, moist,
raised, entire; no chromogenesis. Pin-point
in size in 48 hours, growing slowly until
they reach a diameter of 1.5 mm in 6 days.
Nutrient broth: Moderate growth after
24 hours, appearing at first as a thin film
while a slight, general turbidity develops
in another 24 to 48 hours.
Calcium oxalate agar: Growth rapid and
colonies small; medium becomes alkaline.
Oxalate broth: Becomes turbid following
the formation of a slight surface film.
Oxalates and pyruvates support growth
within 3 to 4 days when added to a mineral
medium as the sole carbon source; formates
support growth only when the incubation
period is extended. The following do not
FAMILY VII. SPIRILLACEAE
243
support growth under any of the above
conditions when added to the mineral me-
dium: butyrates, citrates, lactates, malates,
malonates, succinates, tartrates or glucose.
Indole not produced.
Hydrogen sulfide not produced.
Nitrites not produced from nitrates.
Aerobic.
Source: Five strains were isolated from
Boston, Mass., and Berkeley, California,
soils by inoculation of soil into a medium
containing potassium oxalate and other
minerals in distilled water. All soil samples
tested showed the presence of this species.
Ayers, Rupp and Johnson (U. S. Dept. Agr.
Bull. No. 782, 1919, 38 pp.) and den Dooren
de Jong (Dissertation, Delft, 1926, Table
XVIII) tested over 125 strains of bacteria
without finding any that decomposed ox-
alate. Bassilik (Jahrb. wiss. Bot., 63, 1913,
255) found only three strains out of 90 tested
which decomposed oxalate, two slowly;
the third was the species described by him
(Vibrio extorquens).
Habitat: Widely distributed in soil.
24. Vibrio extorquens (Bassalik, 1913)
Bhat and Barker, 1948. (Bacilhis extorquens
Bassalik, Jahrb. f. wiss. Bot., 53, 1913, 255;
Bhat and Barker, Jour. Bact., 55, 1948, 367;
Pseudomonas extorquens Janota, Med. Dos-
wiadczalna i Mikrobiol., 2, 1950, 131; see
Biol. Abstracts, .25,1951, Abs. no. 34148.)
ex.tor'quens. L. puTt.ad] . extorquens twist-
ing out.
Slightly curved rods, 1.5 by 3.0 microns.
Motile by means of a single, polar flagel-
lum. Gram-negative.
Gelatin media: Poor growth. Colonies
small (less than 1 mm in diameter in 7 days),
round, entire, butyrous. Surface colonies
dirty yellow to yellowish red, eventually
becoming a beautiful red color. No lique-
faction.
Oxalate and similar mineral media:
Growth rapid and abundant.
Peptone-agar colonies: Growth slower
than on gelatin.
Liquid oxalate media. Grows rapidly as
a rose-colored film on the bottom and walls
of the flask, leaving the liquid clear.
Potato: Slow growth with darkening of
potato.
Litmus milk: Not coagulated. Reaction
becomes alkaline but growth is poor.
Aerobic, facultative.
Optimum temperature, between 25° and
30° C. Poor growth at 37° C.
Optimum pH : Prefers media with an alka-
line reaction.
Distinctive characters: In old cultures in
liquid calcium oxalate media and especially
in media made with plant materials con-
taining oxalate crystals, the cells become
encrusted with a surface deposit. This ap-
pears to be calcium carbonate and is easily
dissolved with dilute acid, especially dilute
HCl.
Source: Originally isolated by adding the
e.xcreta of earthworms that had ingested
plant materials containing oxalate crystals
to a liquid medium containing ammonium
oxalate. Pure cultures were isolated with
difficulty by using a silica gel medium con-
taining ammonium oxalate. Later these
organisms were found to be generally pres-
ent in forest and garden soils in Switzerland.
Habitat: Presumably widely distributed
in soil.
25. Vibrio cuneatus Gray and Thornton,
1928. (Cent. f. Bakt., II Abt., 73, 1928, 92.)
cu.ne.a'tus. L. part. adj. amea/z/s wedge-
shaped.
Curved rods, 1.0 by 1.0 to 3.0 microns,
the cells tapering at one extremity. Motile
by means of one to five polar flagella. Gram-
negative.
Gelatin colonies: Liquefied.
Gelatin stab: Liquefied.
Agar colonies: Circular to amoeboid,
white to buff, flat to convex, smooth, trans-
lucent, border entire.
Agar slant: Filiform, whitish, smooth,
glistening.
Indole production not recorded.
No acid from carbohydrate media.
Starch not hydrolyzed.
Nitrites not produced from nitrates.
Attacks naphthalene.
Aerobic, facultative.
Optimum temperature, between 30° and
35° C.
Source: One strain was isolated from
soil from Rothamsted, England.
Habitat: Soil.
244
ORDER I. PSEUDOMONADALES
26. Vibrio percolans Mudd and Warren,
1923. (Jour. Bact., 8, 1923, 447.)
per'co.lans. L. v. percolo to filter through;
L. part. adj. percolans filtering through.
Curved rods, 0.3 to 0.4 by 1.5 to 1.8 mi-
crons, occurring singly or in short chains.
Pleomorphic. Actively motile by means of
1 to 3 polar flagella. Gram-negative.
Gelatin stab: No liquefaction.
Agar colonies: Circular, slightly convex,
amorphous, entire.
Agar slant: Bluish white, glistening
streak.
Broth: Turbid. Pellicle, sediment.
Litmus milk: Unchanged.
Potato: White, slimy streak.
Coagulated blood serum not liquefied.
Indole not produced.
No action on carbohydrates.
Starch not hydrolyzed.
Nitrites not produced from nitrates.
Passes through bacterial filters (Berkefeld
V candles).
Aerobic, facultative.
Optimum temperature, 30° C.
Non-pathogenic.
Relationships to other species: Except
for polar flagellation, this species has char-
acters much like those of Alcaligenes faecalis
Castellani and Chalmers. The two species
are frequently confused. For example Leh-
mann and Neumann renamed Alcaligenes
faecalis as Vibrio alcaligenes in their Bakt.
Diag., 7 Aufl., 2, 1927, 548, in the mistaken
idea that the former organism is polar flagel-
late. Leifson and Hugh (personal com-
munication, 1954), who recognize the spe-
cies Vibrio alcaligenes, report that Vibrio
alcaligenes produces nitrites from nitrates
and that it does not hydrolyze urea (with
possible rare exceptions). They report also
that Vibrio alcaligenes occurs more fre-
quently in the intestine than does Alca-
ligenes faecalis Petruschky.
Source: Isolated from a hay infusion.
Habitat: Water.
27. Vibrio adaptatiis ZoBell and Up-
ham, 1944. (Bull. Scripps Inst, of Ocean-
ography, Univ. Calif., 6, 1944, 258.)
a.dap.ta'tus. L. part. adj. adaptatus fitted,
adapted.
Curved rods, 0.4 to 0.5 by 1.6 to 2.3 mi-
crons, only slightly curved, occurring singly
and sometimes in pairs. Motile by means
of a single, polar flagellum. Gram-negative.
Note: All differential media except the
fresh-water broth, litmus milk and potato
were prepared with sea water.
Gelatin colonies: Pin-point, yellow.
Gelatin stab: No liquefaction. Yellow,
filiform growth along stab.
Agar colonies: Punctiform, yellow,
opaque, pulvinate, smooth.
Agar slant: Luxuriant, filiform, shiny
growth with waxy yellow pigment.
Sea-water broth: Moderate turbidity;
thick, yellow pellicle; slight, flocculent
sediment.
Fresh-water broth: Moderate growth.
Litmus milk: No visible change.
Potato : No visible growth.
Indole not produced.
Hydrogen sulfide not produced.
Glucose, sucrose, maltose, lactose, xylose,
glycerol, mannitol and salicin not fer-
mented.
Starch not hydrolyzed.
Non-lipolytic.
Nitrites not produced from nitrates.
Ammonia produced from peptone but
not from urea.
Casein not digested.
Aerobic, facultative (poor anaerobic
growth) .
Optimum temperature, between 20° and
25° C.
Source: Isolated from sea water and from
marine sediments.
Habitat: Common; probably widely dis-
tributed.
28. Vibrio piscium David, 1927. (Cent,
f. Bakt., I Abt., Orig., 102, 1927, 46.)
pis'ci.um. L. noun piscis a fish; L. gen.pl.
piscium of fishes.
Curved rods 0.3 to 0.5 by 2.0 microns.
Motile by means of a single, polar flagellum.
Gram-negative.
Gelatin colonies: Circular, granular,
opaque.
Gelatin stab: Napiform liquefaction.
Agar colonies: Yellowish, circular,
smooth, entire, iridescent.
Agar slant: Light yellow, transparent
streak.
FAMILY VII. SPIRILLACEAE
245
Broth: Slightly turbid; thin pellicle.
Litmus milk: Soft coagulum. Peptonized,
alkaline.
Potato : Brownish red streak.
Indole produced.
Hj^drogen sulfide produced.
No action in sugar media.
Nitrites not produced from nitrates.
Pathogenic for frogs.
Aerobic, facultative.
Optimum temperature, between 18° and
20° C.
Habitat: Causes epidemic infection in
fish.
29. Vibrio hyphalus ZoBell and Up-
ham, 1944. (Bull. Scripps Inst, of Ocean-
ography, Univ. Calif., 5, 1944, 277.)
hy.pha'lus. Gr. adj. hyphalus under the
sea, submarine.
Curved rods, 0.6 by 1.6 to 4.0 microns,
with rounded ends, occurring singly. Motile
by means of one or occasionally two polar
fiagella. Granular staining. Gram-negative.
Note: All differential media e.xcept the
fresh-water broth, litmus milk and potato
were prepared with sea water.
Gelatin colonies: Circular or irregular
with liquefaction; yellowish gray.
Gelatin stab: Napiform liquefaction.
Filiform growth along line of stab.
Agar colonies: 2 to 3 mm in diameter,
circular, undulate, convex, radial folds,
smooth.
Agar slant: Abundant, echinulate, glis-
tening, gummy growth with pale pink pig-
ment.
Sea-water broth: Scant pellicle; moder-
ate turbidity; moderate, flocculent sedi-
ment.
Fresh-water broth: No visible growth.
Litmus milk: No visible change.
Potato: No visible growth.
Indole not produced.
Hydrogen sulfide is produced.
No acid or gas from glucose, sucrose, lac-
tose, glycerol, xylose, mannitol or salicin.
Starch not hydrolyzed.
Non-lipolytic.
Nitrites produced from nitrates.
Ammonia produced from peptone but
not from urea.
Casein is digested.
Aerobic, facultative.
Optimum temperature, between 20° and
25° C.
Source: Isolated from marine bottom
deposits.
Habitat: Probably widely distributed.
30. Vibrio fetus Smith and Taylor, 1919.
(Spirillum causing abortion in sheep, Mac-
Fadyean and Stockman, Rept. Dept. Comm.
Ministry Agric. on Epizootic Abortion,
London, 1909, 156; also see MacFadyean
and Stockman, ibid., 1913, 111; Spirillum
associated with infectious abortion. Smith,
Jour. Exp. Med., 28, 1918, 701; Smith and
Taylor, ibid., 30, 1919, 299.)
fe'tus. L. noun fetus a fetus; L. mas. gen. n.
fetus of a fetus.
Description taken primarily from Plast-
ridge, Williams, Easterbrooks, Walker and
Beccia (Storrs Agr. Exp. Sta., Bull. 281,
1951, 11) and from Rhoades (Bact. Proc,
53rd Gen. Meeting, Soc. Amer. Bact., San
Francisco, 1953, 34).
Curved rods that are minute, comma-
and S-shaped forms on initial isolation.
On transfer, very long, filamentous forms
may appear. 0.2 to 0.5 by 1.5 to 5.0 microns.
Motile, the comma forms possessing a single,
polar fiagellum, and the S forms usually
possessing a single fiagellum at each pole.
Prolonged incubation and transfer to dry
slants or semisolid media produces coccoid
forms with one or more fiagella. Occasion-
ally encapsulated. Granules present in older
cultures. Gram-negative.
Gelatin: No or poor growth on ordinary
gelatin; with the addition of proper nutri-
ments, good to excellent growth may occur
in 3 to 5 days. No liquefaction.
Agar plates: No growth. Reich, Morse and
Wilson (Amer. Jour. Vet. Res., 17, 1956,
140), however, report growth when cultures
are incubated in an atmosphere of either
helium or nitrogen.
Agar slant: No surface growth by freshly
isolated strains; laboratory strains produce
a scant, grayish white, glistening surface
growth. Good growth is obtained when
cultures are incubated in an atmosphere of
helium or nitrogen (Reich et al., loc. cit.).
Sub-surface agar colonies: Small, yellow,
opaque.
246
ORDER I. PSEUDOMONADALES
Blood agar plates (in 10 per cent CO2
atmosphere) : Growth.
Thiol agar (prepared by adding 35.0 gm
of granular agar and 0.05 gm of glutathione
to 1.0 liter of thiol medium (supplied in
dehydrated form by Difco Laboratories)
and adjusting the pH to 8.9): Moderate
growth. Colonies vary from small (1 mm in
diameter), transparent and convex to trans-
lucent or opaque, light tan colonies up to
3 mm in diameter. Masses of growth are
translucent and light gray or light tan.
Broth: A viscid ring pellicle may appear;
faint clouding of the medium occurs; a
filmy, stringy deposit may settle out.
Litmus milk: No growth.
Potato: No growth.
Indole not produced.
Hydrogen sulfide not produced.
Nitrites produced from nitrates (Bryner
and Frank, Amer. Jour. Vet. Res., 16, 1955,
76).
Blood serum slant: Feeble growth. No
liquefaction.
No gas from carbohydrates. No change
or slightl.y acid from glucose, lactose and
sucrose. No acid from the following carbo-
hydrates when each was added to a medium
of beef infusion with peptone, agar and
Andrade's indicator: glucose, fructose,
galactose, arabinose, raffinose, trehalose,
sucrose, maltose, lactose, dextrin, inulin,
salicin, dulcitol, mannitol and sorbitol.
Temperature relations: Optimum, 37° C.
Minimum, 15° C. Maximum, 40.5° C. With-
stands 55° C. for 5 minutes.
Strains isolated from cases of abortion
are catalase-positive (Bryner and Frank,
loc. cit.).
Salt tolerance: Tolerates 1.5 to 2.0 per
cent NaCl in a semisolid medium.
Bile tolerance: Most strains grow in a
semisolid medium containing 10 per cent
fresh ox bile; all strains grow in 5 per cent
ox bile media (Schneider and Morse, Cor-
nell Vet., J!^5, 1955, 84).
Aerobic to microaerophilic.
Pathogenicity: Infection with Vibrio
fetus (vibriosis) causes abortion in cattle
and sheep. Pathogenic for guinea pigs, ham-
sters and embryonated chicken eggs (see
Webster and Thorp, Amer. Jour. Vet. Res.,
14, 1953, 118; Ristic and Morse, ibid., 399;
and Ristic, Morse, Wipf and McNutt, ibid.,
15, 1954, 309). Non-pathogenic to rabbits,
rats and mice when injected intraperitone-
ally.
Source: Twenty-two strains were isolated
from the placentas or fetuses of cows having
abortion.
Habitat: Causes abortion in cattle and
sheep.
31. Vibrio coli Doyle, 1948. (Comma-
shaped microorganisms. Whiting, Doyle
and Spray, Purdue Univ. Agr. Exp. Sta.
Bull. 257, 1921, 12; Vibrio of swine dysen-
tery, Doyle, Amer. Jour. Vet. Res., 5, 1944,
3; Doyle, ibid., 9, 1948, 50.)
co'li. Gr. noun colum or colon the large
intestine, colon; M.L. gen. noun coli of the
colon.
Description taken from Doyle {loc. cit.)
and Hauduroy et al. (Diet. d. Bact. Path.,
2nd ed., 1953, 649).
Curved rods, comma- and sometimes
spiral-shaped, 0.2 to 0.5 by 1.5 to 5.0 mi-
crons. Motile by means of a single, polar
flagellum. Gram-negative.
Agar colonies: Transparent and color-
less. Good growth only when the medium
contains 10 per cent of defibrinated blood
and when the atmosphere contains 15 per
cent CO2 ; abundant growth in the moisture
of condensation.
Gelatin: Not liquefied.
Litmus milk: No growth; not coagulated.
Indole not produced.
Glucose, sucrose, lactose, maltose and
mannitol not utilized.
Nitrites not produced from nitrates.
Coagulated blood serum not hemolyzed.
Pathogenicity: Injection causes no dis-
ease in calves, rabbits, rats, mice, guinea
pigs or chickens. Injection causes dj-sentery
in swine.
Source: Isolated from the mucosa of the
colon of a swine which had died of dys-
entery.
Habitat: Causes dysentery in swine.
32. Vibrio jejuni Jones et al., 1931.
(Jones, Orcutt and Little, Jour. Exp. Med.,
53, 1931, 853.)
FAMILY VII. SPIRILLACEAE
247
je.ju'ni. L. adj. jejunus insignificant,
meagre; ALL. noun jejunuin the jejunum.
Pleomorphic, occurring in three different
form.s in the same culture: the first forms
are short, slightly convoluted and activel}^
motile with either a single polar flagellum
or a single flagellum at each pole; the sec-
ond are less active and have two or more
complete coils; the remaining forms are
extremely long and rarely motile. In older
cultures clumps occur, and these usually
degenerate into fragments and granules.
Gram-negative.
Blood agar: Within 4 or 5 days the con-
densation fluid becomes slightly turbid;
delicate lines then appear at the border of
the agar. After several transfers these lines
Ijecome well defined, and a delicate film
spreads over the nether portion of the slant.
Gelatin: Not liquefied.
Coagulated blood serum not lifiuefied.
Carbohydrates not utilized.
Aerobic.
Optimum pH, 7.6. No growth in either
slightly acid or definitely alkaline media.
Temperature relations : Optimum, 37.5° C.
(Merchant, Vet. Bact. and Virology, 4th
cd., 1950, 343). Killed in 5 minutes at 55° C.
Pathogenicity: Non-pathogenic to lab-
oratory animals under the usual conditions.
Some strains produce multiple necrotic foci
of the liver when injected intraperitoneally
into white mice. Febrile reactions are pro-
duced in rabbits when injected intrave-
nously with certain strains. The enteritis
produced experimentally in calves is less
severe than that occurring spontaneously.
Source: Isolated from the small intestine
of calves suffering from diarrhoea.
Habitat: Causes diarrhoea in cows and
calves where it is found in the small in-
testine and feces.
33. Vibrio niger (Rist, 1898) Prevot,
1948. (Spirillum nigrum Rist, These med.,
Paris, 1898; also see Cent. f. Bakt., I Abt.,
30, 1901, 299; Prevot, Man. d. Classif. Bact.
Anaer., 2nd ed., 1948, 124.)
ni'ger. L. adj. niger black.
Description taken from Rist (op. cil.,
1901, 299) and Hauduroy et al. (Diet. d.
Bact. Path., 2nd ed., 1953, 658).
Long, slender, comma- or S-shaped cells
rounded at the ends; 1.0 to 2.0 by 3.0 mi-
crons. Motile. Possess a black granule which
swells the cell and which may be terminal.
Gram-negative.
Glucose agar colonies: Lenticular, dark
black, opaque, 2 to 3 mm in diameter.
Deep agar colonies: Lenticular, black,
cloudy; gas is produced.
Deep blood serum agar colonies: Small,
thin, delicate, non-hemolytic.
Brain medium: Blackened; hydrogen
sulfide, ethanol and butyric and lactic acids
are produced.
Gelatin colonies: Black, opaque; putrid
odor. No liquefaction.
Glucose broth: Dark gray turbidity;
putrid odor; gas and hydrogen sulfide are
produced.
Peptone broth: Poor growth.
Milk: Coagulated slowly then digested.
Coagulated ascitic fluid: Not liquefied.
Indole not produced.
Hydrogen sulfide produced.
Neutral red reduced.
Obligate anaerobe.
Temperature relations: Growth range,
21° to 37° C. Death occurs at 55° C. Can
withstand freezing.
Pathogenicity: Fatal for guinea pigs in
two weeks; macroscopic lesions not demon-
strable.
Source: Isolated from purulent otitis,
mastoiditis and pulmonary gangrene; also
isolated from cases of meningitis and appen-
dicitis.
Habitat: Found rather frequently in man
under pathological conditions.
34. Vibrio sputoruin Prevot, 1940. (An
anaerobic vibrio from bronchitis, Tunni-
cliff. Jour. Inf. Dis., 15, 1914, 350; A small
anaerobic vibrio from Vincent's angina.
Smith, ibid., Jfi, 1930, 307; Prevot, Man. de
Classif. des Bact. Anaer., Paris, 1940, 85.)
spu.to'rum. L. noun sputum spit, spu-
tum; L. gen.pl. noun sputorum of sputa.
Description taken from Prevot {loc. cit.)
and from Macdonald (Motile, Non-sporu-
lating. Anaerobic Rods of the Oral Cavity,
Toronto, 1953, 53).
Straight or slightly curved rods, 0.5 to
248
ORDER I. PSEUDOMONADALES
0.8 by 2.0 to 8.0 microns, occurring singly,
in pairs or in short chains. Active, darting
motility by means of 1 to 3 polar flagella.
Gram-negative.
Grows only in media to which body fluids
(blood, ascites, etc.) have been added or in
other enriched media.
Ascitic fluid broth: Almost imperceptible
turbidity.
Thioglycollate broth: Light, floccular
turbidity.
Blood agar colonies : After 5 days, smooth,
convex, grayish yellow, dull and translucent
with a finely fimbriate margin. Less than
0.5 mm in diameter. Frequently surrounded
by a narrow zone of green.
Coagulated blood serum: Cloth-like
growth. No odor.
Genus II. Desulfovibrio Kluyver and van Niel, 1936.*
(Kluyver and van Niel, Zent. f. Bakt., II Abt., H, 1936, 369; Sporovibrio
Starkey, Arch. f. Mikrobiol., 9, 1938, 300.)
De.sul.fo.vib'ri.o. L. pref. de from; L. sulfur sulfur; L. v. vibro to vibrate; M.L. mas.n.
Vibrio that which vibrates, a generic name; M.L. mas.n. Desulfovibrio a vibrio that reduces
sulfur compounds.
Slightly curved rods of variable length, usually occurring singly but sometimes in short
chains which have the appearance of spirilla. Swollen pleomorphic forms are common.
Actively motile by means of a single polar flagellum. Strict anaerobes which reduce sulfates
to hydrogen sulfide. Found in sea water, marine mud, fresh water and soil.
The type species is Desulfovibrio desulfuricans (Beijerinck) Klu3^ver and van Niel.
Nitrites produced from nitrates by some
strains.
Indole not produced.
Hydrogen sulfide produced.
Carbohydrates not utilized.
Optimum pH, 7.2. Feeble growth between
pH 6.0 and 9.7.
Serology: Cross-reacting, somatic anti-
gens have been demonstrated.
Anaerobic.
Optimum temperature, 37° C. Growth
feeble at 27° and 45° C. No growth at 20° C.
Source: Isolated by Pr^vot from a case of
bronchitis.
Habitat: Found in the human oral cavity
and in fusospirochetal diseases of the mouth.
1. Desulfovibrio desulfuricans (Beijer-
inck, 1895) Kluyver and van Niel, 1936.
(Bacterium hydrosulfureum ponticum Zelin-
sky, Proc. Russ. Phys. and Chem. Soc, 25,
1893, 298; Spirillum desulfuricans Beijer-
inck, Cent. f. Bakt., II Abt., 1, 1895, 1;
Kluyver and van Niel, Zent. f. Bakt., II
Abt., 94, 1936, 369; Sporovibrio desulfuricans
Starkey, Koninkl. Nederland. Akad. v.
Wetenschappen, Proc, 41, 1938, 426; also in
Arch. f. Mikrobiol., 9, 1938, 268.)
de.sul.fur'i.cans. L. pref. de from; L.
noun sulfur sulfur; M.L. part. adj. desul-
furicans reducing sulfur compounds.
Slightly curved rods, 0.5 to 1.0 by 1 to 5
microns, usuall}- occurring singly but some-
times in pairs and short chains which cause
them to look like spirilla. Swollen pleomor-
phic forms are common. Older cells appear
black due to precipitated ferric sulfide.
Actively motile, possessing a polar flagel-
lum. Stains readily with carbol fuchsin.
Gram-negative.
Gelatin: No liquefaction.
Grows best in fresh-water media. Fails to
develop in sea water upon initial isolation.
Produces opalescent turbidity in absence
of oxygen in mineral media enriched with
sulfate and peptone.
Media containing iron salts and sulfur
compounds blackened. Bacteria found asso-
ciated with precipitated ferrous sulfide.
Peptone-glucose agar colonies (in absence
of air) : Small, circular, slightly raised, dull,
entire, soft in consistency.
Peptone, asparagine, glycine, alanine,
* Prepared by Dr. Claude E. ZoBell, Scripps Institution of Oceanography, La Jolla, Cali-
fornia, January, 1943; revised January, 1953.
FAMILY VII. SPIRILLACEAE
249
aspartic acid, ethanol, propanol, butanol,
glycerol, glucose, lactate, succinate and
malate known to be utilized as hydrogen
donors. Some varieties oxidize H2 .
Produces up to 3100 mg HjS per liter.
Nitrites not produced from nitrates.
Reduces sulfate to hydrogen sulfide;
also reduces sulfites, thiosulfates and
hyposulfites.
Temperature relations: Optimum, be-
tween 25° and 30° C. Maximum, between
35° and 40° C.
Chemical tolerance: Optimum pH, be-
tween 6 and 7.5. Limits for growth, between
pH 5 and 9.
Cytochrome is produced.
Anaerobic.
Habitat : Soil, sewage and water.
2. Desulfovibrio aestuarii (van Delden,
1904; ZoBell, 1948. (Microspira aestuarii van
Delden, Cent. f. Bakt., II Abt., 11, 1904,81;
ZoBell, in Manual, 6th ed., 1948, 208.)
aes.tu.a'ri.i. L. noun uestuarium an
estuary, inlet; L. gen. noun aestuarii of an
estuary.
Morphologically indistinguishable from
Desulfovibrio desulfuricans described above,
although it has a greater tendency to pleo-
morphism and is slightly larger. Motile,
possessing a polar flagellum. Gram-nega-
tive.
Gelatin: No liquefaction.
Grows preferentially in media prepared
with sea water or 3 per cent mineral salt
solution enriched with sulfate and peptone.
According to Baars (Over Sulfaatreductie
door Bakterien, Diss. Delft, 1930, 164 pp.)
the marine species can be acclimatized to
tolerate hj^potonic salt solutions, but Rit-
tenberg (Studies on Marine Sulfate-
Reducing Bacteria, Thesis, Univ. of Calif.,
1941, 115 pp.) was unable to confirm this
observation. Likewise Rittenberg was
unable to acclimatize D. aestuarii to tolerate
temperatures exceeding 45° C. or to produce
endospores.
Produces faint turbidity in absence of
oxygen in sea water enriched with sulfate
and peptone. Organisms most abundant in
sediment.
Agar colonies: Small, circular, slightly
raised, darker centers, entire, soft con-
sistency.
Peptone, asparagine, glycine, alanine,
glucose, fructose, ethanol, butanol, gh^cerol,
acetate, lactate and malate known to be
utilized in presence of sulfate. Some strains
utilize molecular hydrogen as the sole source
of energy.
Reduces sulfate to hj'drogen sulfide. Also
reduces sulfites, thiosulfates and hy-
posulfites.
Produces up to 950 mg of hydrogen sulfide
per liter.
Nitrites not produced from nitrates.
Temperature relations: Optimum, be-
tween 25° and 30° C. Maximum, between
35° and 40° C.
Chemical tolerance: Optimum pH, be-
tween 6 and 8. Limits for growth, between
pH 5.5 and 8.5.
Anaerobic.
Habitat: Sea water, marine mud, brine
and oil wells.
3. Desulfovibrio rubentschikii (Baars,
1930) ZoBell, 1948. {Vibrio riibentschickii
(sic) Baars, Over Sulfaatreductie door Bak-
terien, Diss. Delft, 1930, 89; ZoBell, in Man-
ual, 6th ed., 1948, 208.)
ru.ben.tschi'ki.i. M.L. gen. noun ruben-
tschikii of Rubentschik; named for Dr. L.
Rubentschik.
Slightly curved rods, 0.5 to 1.0 by 1 to 5
microns, usually occurring singly, some-
times in pairs and short chains. Actively
motile, possessing a polar flagellum. Gram-
negative. Morphologically indistinguishable
from Desulfovibrio desulfuricans.
Reduces sulfate to hydrogen sulfide; also
reduces sulfites, thiosulfates and hypo-
sulfites.
Culturally and phj'siologically like D.
desulfuricans except that D. rubentschikii
utilizes propionic, butyric, valeric, palmitic
and stearic acids and galactose, sucrose,
lactose and maltose.
Anaerobic.
Habitat: Soil and ditch water.
250 ORDER I. PSEUDOMONADALES
Genus III. Methanobacterium Kluyver and van Niel, 1936*
(Zent. f. Bakt., II Abt., 94, 1936, 399.)
Me.tha.no.bac.te'ri.um. Gr. noun iiiethy wine; M.L. noun methanum methane; Gr.
neut.n. bacterium a small rod; M.L. noun Methanobacterium the methane (-producing)
rodlet.
Straight or slightly curved rods, sometimes united in bundles or long chains. Reported to
be non-motile. Anaerobic. Chemo-heterotrophic or chemo-autotrophic, oxidizing various
organic or inorganic compounds and reducing carbon dioxide to methane. Gram-negative.
The anaerobic genus Methanobacterium was proposed by Kluyver and van Niel in 1936 with
an indication that they regarded Sohngen's methane bacterium as the type species of the
genus. Later, Barker (1936) found organisms that he regarded as identical with those pre-
viously isolated by Sohngen, and he proposed the name Methanobacterium sohngenii for this
species. While the organisms belonging to this genus are reported to be non-motile, the
curved form of their cells and their physiology places them near the species placed in De-
suFovibr-io.
The type species is Methanobacterium soehngenii Barker.
1. Methanobacterium soehngenii Acetate and n-butyrate but not pro-
Barker, 1936. (Methane bacterium, Sohn- pionate are fermented with the production
gen, Dissertation, Delft, 1906; Barker, Arch. of methane and carbon dioxide,
f. Mikrobiol., 7, 1936, 433.) Ethyl and n-butyl alcohols not fermented.
soehn.ge'ni.i. M.L. gen.noun soehngenii Obligate anaerobe,
of Sohngen; named for Prof. N. L. Sohngen, g^^^.^^ . Enrichment cultures containing
the bacteriologist who first studied this acetate or butyrate as the only organic com-
?,,■., ,. , , , , pound. Four strains were isolated from ace-
Rods straight to slightly curved, moder- "■' . . ^ ,^ rr,, ,.,
^ , , ,, ,., ^, ,. tate enrichment cultures. The cultures were
ately long. ISon-motile. Gram-negative. .^ , , . ,
In liquid cultures, cells are character- highly purified but not strictly pure,
istically joined into long chains which often Habitat: Canal mud, sewage. Probably
lie parallel to one another so as to form occurs widely in fresh-water sediments
bundles. where anaerobic conditions prevail.
Genus IV. Cellvibrio Winogradsky, 1929.]
(Ann. Inst. Pasteur, J^S, 1929, 577.)
Cell.vib'ri.o. L. noun cella a room, cell; L. v. vibro to vibrate; M.L. mas.n. Vibrio that
which vibrates, a generic name; M.L. mas.n. Cellvibrio cell vibrio, but here the cell is an
abbreviation of cellulose, hence, cellulose vibrio.
Long slender rods, slightly curved, with rounded ends, showing deeply staining granules
which appear to be concerned in reproduction. Monotrichous. Most species produce a yellow
or brown pigment with cellulose. Oxidize cellulose, forming oxycellulose. Growth on ordi-
nary culture media is feeble. Found in soil.
The type species is Cellvibrio ochraceus Winogradsky.
Key to the species of genus Cellvibrio.
I. No growth on glucose or starch agar.
A. Ochre-yellow pigment produced on filter paper.
1. Cellvibrio ochraceus.
* Revised by Prof. Robert S. Breed, Cornell University, Geneva, New York, May, 1955.
t Revised by Prof. Robert S. Breed, Cornell University, Geneva, New York, with the
assistance of Prof. Onorato Verona, University of Pisa, Pisa, Italy, September, 1953.
FAMILY VII. SPIRILLACEAE
251
II. Growth on glucose and starch agar.
A. Poor growth on starch agar.
1. Cream-colored pigment which becomes brown with age is produced on filter paper.
2. Cellvibrio flavescens.
B. Abundant growth on starch agar.
1. Scant growth on glucose agar.
a. Intense yellow pigment produced on filter paper.
3. Cellvibrio fulvus.
2. Abundant growth on glucose agar.
a. No pigment produced on filter paper.
4. Cellvibrio vulgaris.
1. Cellvibrio ochraceus Winogradsky,
1929. (Ann. Inst. Pasteur, 43, 1929, 549, 601.)
och.ra'ce.us. Gr. noun ochra ochre; M.L.
adj. ochraceus like ochre, rust-colored.
Plump, curved rods with rounded ends,
2.0 to 4.0 microns long, rarely occurring as
spirals. Chromatic granule frequently found
in center. Motile by means of a single flagel-
lum. Gram-negative.
Produces diffuse, light ochre-colored,
mucilaginous colonies on cellulose silica gel
medium.
No action or growth on plain agar. No
growth on peptone, glucose, starch or
tragacanth gum agar.
Grows well on hydrocellulose agar without
producing clearings.
Cellulose is oxidized to acid oxycellulose
without the production of reducing sub-
stances or volatile by-products; a soluble,
non-reducing product may be formed.
Filter paper streaks : Entire paper colored
ochre-yellow in 48 hrs.
Aerobic, facultative.
Optimum temperature, 20° C.
Distinctive character: Rapid, ochre-
colored growth.
Habitat: Soil. Disintegrates vegetable
fibers.
2. Cellvibrio flavescens Winogradsky,
1929. (Ann. Inst. Pasteur, 43, 1929, 608.)
fla.ves'cens. L. v. flavesco to become
golden yellow; L. part. adj. flavescens be-
coming yellow.
Plump, curved rods, flexuous, with
rounded ends, 0.5 by 2.5 to 5.0 microns.
Show metachromatic granules. Motile by
means of a single flagellum. Gram-negative.
Produces diffuse, cream-colored growth
becoming brownish; mucilaginous colonies
on cellulose silica gel medium.
Good growth on peptone agar. Colonies
I mm in 4 days. Grows poorly on glucose,
starch and gum agars.
Filter paper streaks: Almost as rapid in
growth as Cellvibrio ochraceus and colors
entire paper in 2 to .3 days.
Aerobic, facultative.
Optimum temperature, 20° C.
Distinctive characters: Smaller, less
curved rods which grow on a greater variety
of media than Cellvibrio ochraceus but which
do not attack cellulose as readily.
Source: Isolated from a pile of old damp
sawdust. A variety of this organism has been
isolated from sea water by Kadota (Bull.
Japan. Soc. Sci. Fish., 16, 1951, 63-70).
Habitat: Soil. Disintegrates vegetable
fibers.
3. Cellvibrio fulvus Stapp and Bortels,
1934. (Culture Y, Dubos, Jour. Bact., 15,
1928, 230; Stapp and Bortels, Zent. f. Bakt.,
II Abt.,50, 1934,42.)
ful'vus. L. ad}, fulvus deep yellow.
Slightly curved rods, 0.3 to 0.4 by 1.5 to
3.0 microns. Show involution forms. Motile
by means of a single, polar flagellum. Gram-
negative.
Cellulose is decomposed. Grows on filter
paper with an intense egg-yellow color
which in older cultures may deepen to rust
brown.
Glucose agar: Very scant growth.
Sucrose agar: Very slight growth.
Maltose agar: Abundant yellow growth.
Lactose agar: Fairly abundant yellow
growth.
252
ORDER I. PSEUDOMONADALES
Starch agar: Very abundant, In-ight
vellow growth which later turns brown.
Nutrient broth: No growth.
Temperature relations: Optimum, be-
tween 25° and 30° C. Minimum, 5° C. Maxi-
mum, between 32° and 35° C. No growth at
37° C. Thermal death point, between 39°
and 40° C.
Aerobic.
Source: Isolated from forest soil in Ger-
many and from soil in the United States.
Habitat: Widely distributed in soils.
4. Cellvibrio vulgaris Stapp and Bor-
tels, 1934. (Culture Co, Dubos, Jour. Bact.,
15, 1928, 230; Stapp and Bortels, Zent. f.
Bakt., II Abt., 50, 1934,44.)
vul.ga'ris. L. adj. vulgaris common.
Curved rods, 0.3 by 2.9 to 4.0 microns.
Show involution forms. Motile by means of
a single polar flagellum. Gram-negative.
Cellulose is decomposed. Grows on filter
paper without the formation of pigment.
Glucose agar: Abundant growth. No pig-
ment.
Sucrose agar: Abundant, slightly yellow
growth.
Maltose agar: Abundant, .yellowish
growth.
Lactose agar: Very heavy growth.
Starch agar: Very abundant, yellowish
growth.
Nutrient broth: No growth.
Temperature relations: Optimum, be-
tween 25° and 30° C. Minimum, 5° C. Maxi-
mum, between 32° and 35° C. No growth at
37° C. Thermal death point, between 44°
and 45° C.
Aerobic.
Source: Isolated from forest soil in Ger-
many and from soils in the United States.
Habitat: Widelv distributed in soils.
Genus V. Cellfalcicula Winogradsky, 1929.*
(Ann. Inst. Pasteur, 43, 1929, 616.)
Cell.fal.ci'cu.la. L. noun cella a room, cell; M.L. noun ceUulosum cellulose; L. noun
falcicula a sickle; M.L. fem.n. Cellfalcicula cell sickle, but here the cell is an abbreviation of
cellulose, hence, cellulose sickle.
Short rods or spindles, not exceeding 2 microns in length, with pointed ends, containing
metachromatic granules. Old cultures show coccoid forms. Monotrichous. Oxidize cellulose,
forming oxycellulose. Growth on ordinary culture media is feeble. Soil bacteria.
The type species is Cellfalcicula viridis Winogradsky.
1. Cellfalcicula viridis Winogradsky,
1929. (Ann. Inst. Pasteur, 43, 1929, 616.)
vi'ri.dis. L. adj. viridis green.
Plump, small spindles, 0.7 by 2.0 microns,
with rounded ends. Motile by means of a
single flagellum. Gram-negative.
Produces diffuse green, mucilaginous
colonies on cellulose silica gel medium.
Filter paper streaks: Rapid spreading
growth colored green in 3 days at 30° C.
Hydrocellulose agar: Growth rapid,
green; minute, yellowish green, mucous
colonies on streaking.
No growth on peptone, glucose, starch
or gum agar.
Aerobic, facultative.
Optimum temperature, 20° C.
Habitat: Soil.
2. Cellfalcicula mucosa Winogradsky,
1929. (Ann. Inst. Pasteur, 43, 1929, 621.)
mu.co'sa. L. adj. mucosus slimy.
Plump, curved spindles, with slightly
pointed ends. Motile by means of a single
polar flagellum. Contain a single chromatic
granule. Gram-negative.
Produces diffuse, cream-colored, muci-
laginous colonies on cellulose silica gel
medium.
Hydrocellulose agar: Abundant grayish
growth.
No growth on peptone, glucose, starch or
gum agar.
Aerobic, facultative.
Optimum temperature, 20° C.
Habitat: Soil.
* Revised by Prof. Robert S. Breed, Cornell LTniversity, Geneva, New York, September,
1937.
FAMILY VII. SPIRILLACEAE
253
3. Cellfalcicula fusca Winogradsky,
1929. (Ann. Inst. Pasteur, A3, 1929, 622.)
fus'ca. L. adj. fuscus dark, tawny.
Plump, curved spindles, 0.5 by 1.2 to 2.5
microns, with slightly pointed ends and a
central chromatic granule. Motile by means
of a single, polar flagellum. Gram-negative.
Produces diffuse, brownish, slightly
marbled or veined colonies on cellulose silica
gel medium.
Filter paper streak: Paper becomes a
partially transparent, dry, non-mucilagi-
nous pellicle adherent to gel.
Aerobic, facultative.
Optimum temperature, 20° C.
Source: Isolated from a pile of old, damp
sawdust.
Habitat: Probably rotting wood.
Genus VI. Microcyclus 0rskov, 1928.*
(Cent. f. Bakt., I Abt., Orig., 107, 1928, 180; also see Riassunti d. Communicazioni,
VI Cong. Internaz. d. Microbiol., Roma, 1, 1953, 24.)
Micro. cyc'lus. Gr. adj. micrus small, little; Gr. cyclus round, circle; M.L. mas.n. Mic-
rocyclus small circle.
Small, slightly curved, non-motile rods which form a closed ring during growth. These
rings grow into bodies which subdivide again into rod-shaped elements as at the beginning.
Encapsulated. Attack few sugars and then only slowly. From fresh-water ponds and from
soil.
The type species is Microcyclus aquaticus ^rskov.
1. Microcyclus aquaticus 0rskov, 1928.
(Cent. f. Bakt., I Abt., Orig., 107, 1928, 180;
also see Riassunti d. Communicazioni, VI
Cong. Internaz. d. Microbiol., Roma, 1,
1953, 24.)
a.qua'ti.cus. L. adj. aquaticus living in
water.
Very small, slightly curved rods about 1
micron in length. During growth, the rods
form closed rings 2 to 3 microns in diameter.
The next stage is a body consisting of horse-
shoe-shaped halves that are fastened to-
gether without visible divisional lines.
These halves further subdivide into separate
rods ; the rods then form rings and start the
cycle of growth all over again. Form and
capsule are seen most distinctly with direct
agar microscopy and direct India ink agar
microscopy. Encapsulated. Non-motile.
Gram-negative.
Gelatin: No liquefaction in one month.
butyrous. This species is not fastidious in its
growth requirements, although colonies are
small. It grows well on tap-water agar plus
0.5 per cent peptone.
No acid from glucose, sucrose, lactose,
maltose, adonitol, dulcitol, sorbitol, inosi-
tol, rhamnose and salicin. After six weeks,
slight acid in arabinose and xylose.
Indole not produced.
Non-hemolytic.
Grows at temperatures between 5° and
30° C. No growth at 37° C.
Source: Originally found in the waters of
a woodland lake. Later isolated from fresh-
water ponds and occasionally from soil.
Sturges (Absts. of Bact., 7, 1923, 11) briefly
reports the presence of organisms with the
same unusual morphology in ham-curing
brines.
Habitat: Presumably widely distributed
in fresh water and in soil.
Agar colonies: Round, smooth edges.
Genus VII. Spirillum Ehrenberg, 1832.^
(Physik. Abhandl. k. Akad. Wissensch. Berlin, 1832, 38.)
Spi.ril'lum. Gr. noun spira a spiral; M.L. dim.neut.n. Spirillum a small spiral, generic
name.
* Prepared by Dr. J. 0rskov, Director, Statens Seruminstitut, Copenhagen, Denmark,
November, 1953.
t Revised by Prof . Robert S. Breed, Cornell University, Geneva, New York, April, 1954,
based on a Monograph by Giesberger, Inaug. Diss., Delft, Nov. 30, 1936.
254 ORDER I. PSEUDOMONADALES
Cells form either long screws or portions of a turn. Volutin granules are usually present.
Usually motile by means of a tuft of polar flagella (5 to 20) which may occur at one or both
ends of the cells. Aerobic, growing well on ordinary culture media except for one saprophyte
and the pathogenic species; these have not yet been cultivated. Usually found in fresh and
salt water containing organic matter.
The type species is Spirillum undula (Miiller) Ehrenberg.
Key to the species of genus Spirillum.
I. One micron or less in diameter.
A. Volutin granules present.
1. Slow to rapid liquefaction of gelatin.
a. Grajash to brown growth on potato.
1. Spirilluni undula.
aa. Light yellowi.sh orange growth on potato.
2. Spirillum serpens.
2. No liquefaction of gelatin. Of small size (0.5 micron in diameter).
a. Colonies on agar white becoming brownish black and slightly wrinkled.
3. Spirillum itersonii.
aa. Colonies on agar white and smooth.
4. Spirillum tenue.
B. No volutin granules observed.
1. Single fiagellum. From sea water.
5. Spirillum virginianum.
2. Tuft of flagella. From blood of rats and mice.
6. Spirillum minus.
II. Over 1 micron in diameter.
A. Grow poorly or not at all on peptone agar media.
1. Grows poorly on peptone agar and potato.
7. Spirillum kutscheri.
2. Has not been cultivated on artificial media. Very evident volutin granules.
8. Spirillum volutans.
B. Grows abundantly on peptone media. Cells may be deformed with fat droplets.
9. Spirillum lipoferum.
1. Spirillum undula (Miiller, 1786) Agar colonies: Grayish white, smooth.
Ehrenberg, 1832. {Vibrio undula Miiller, Broth: Turbid.
Animalcula infusoria et marina, 1786; Potato: Grayish brown growth.
Ehrenberg, Physik. Abhandlungen d. k. Indole not produced.
Berl. Akad., 1832, 38.) Catala.se-positive.
un'du.la. L. noun unda a wave; M.L. Nitrites not produced from nitrates,
dim.fem.n. undula a small wave. Aerobic, facultative.
Stout threads, 0.9 micron in diameter, Optimum temperature, 25° C.
with one-half to three turns. The wave Cohn (Beitrage z. Biol. d. Pflanzen, 1,
lengths are 6 microns. Width of spiral, 3.0 Heft 2, 1875, 132) reports that he could not
microns. Tufts of three to nine flagella at distinguish this organism from Vibrio
each pole. Volutin granules present. Gram- prolifer Ehrenberg.
negative. Habitat: Putrid and stagnant water.
Gelatin colonies: The surface colonies are
circular, granular, greenish yellow, entire. 2. Spirillum serpens (Miiller, 1786)
Gelatin stab: Thick, white, rugose sur- Winter, 1884. {Vibrio serpens Miiller, Ani-
face growth. Very slow liquefaction. malcula infusoria et marina, 1786, 43;
FAMILY VII. SPIRILLACEAE
255
Winter, in Rabenhorst's Kryptogamen-
Flora, ;, Die Pilze, 1884, 63.)
ser'pens. L. v. serpo to crawl or creep; L.
part. adj. serpens creeping.
Long, curved rods with two (o three wave-
like undulation.s, 0.8 to 1.0 micron in di-
ameter; wave length, 8 to 9 micron.s. Width
of spiral, 1.5 to 1.8 microns. Volutin granules
in cytoplasm. Motile, possessing tufts of
fiagella at both poles. Gram-negative.
Gelatin colonies: Yellowish to brownish,
granular, entire.
Gelatin stab: Yellowish surface growth.
Slow liquefaction.
Agar colonies: Heavy cream-colored
growth.
Agar slant: Grayish, with yellowish
center, granular, entire.
Broth: Turbid.
Litmus milk: Unchanged.
Potato: Clear orange-yellow growth.
Indole not produced.
Catalase-positive.
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 35° C.
Habitat: Stagnant water.
3. Spirillum itersonii Giesberger, 1936.
(Inaug. Diss., Utrecht, 1936, 46 and 57.)
i.ter.so'ni.i. M.L. gen. noun itersonii of
Iterson; named for G. van Iterson, a Dutch
bacteriologist.
The smallest of the spirilla isolated from
water. First observed by van Iterson (Proc.
Kon. Akad. v. Wetensch. Amsterdam, 5,
1902, 685).
Small spirals, 0.5 micron in diameter.
Wave length, 3 to 3.5 microns. Spiral width,
1 to 1.5 microns. Motile by means of bipolar
tufts of fiagella. Gram-negative.
Gelatin stab: No liquefaction.
Grows readily on peptone agar. White
colonies becoming brownish black and
slightly wrinkled.
Potato: Brownish orange growth.
Acid from glucose, fructose, ethyl alcohol,
n -propyl alcohol, n-butyl alcohol and
glj'cerol. Utilizes acetic, propionic, n-bu-
tyric, tartaric, fumaric, lactic, citric and
succinic acids.
Grows well in pe])tone broth. Also utilizes
ammonium compounds.
Catalase-positive.
Anaerobic growth in the presence of
nitrates when organic or ammonia nitrogen
is also available.
Optimum temperature, 30° C.
Source: Isolated from water.
Habitat: Water.
4. Spirillum tenue Khrenberg, 1838.
(Die Infusionsthierchen als vollkommende
Organismen. Leipzig, 1838; also see Bon-
hoff. Arch. f. Hyg., 26, 1896, 162.)
te'nu.e. L. adj. tenuis thin.
Slender spirals 0.7 micron in diameter.
Wave lengths, 4.5 to 5.0 microns; width of
wave, 1.5 to 1.8 microns. Activel}' motile in
peptone water by means of tufts of fiagella
at each pole. Volutin granules present.
Gram-negative.
Gelatin stab: No liquefaction.
Agar colonies: White, smooth.
Peptone agar slant: Heavy growth.
Potato: Light brown growth.
Acid from glucose and fructose. Slight
acid from several other sugars and glycerols.
Utilizes salts of acetic, propionic, n-butyric,
tartaric, lactic, citric, malic and succinic
acids.
Ammonia compounds are used as a source
of nitrogen.
Catalase-positive.
Optimum temperature, 30° C.
Source: Isolated from putrefying vege-
table matter.
Habitat: Putrefying materials.
5. Spirillum virginianum Dimitroff,
1926. (Jour. Bact., 12, 1926, 19.)
vir.gi.ni.a'num. M.L. adj. virginianus
Virginian; named for the State of Virginia.
Spirals consisting of )^ to 3 complete
turns in young cultures, older cultures show-
ing 7 turns; 0.6 to 0.9 by 3 to 11 microns. No
volutin granules observed (Giesberger,
Inaug. Diss., Delft, 1936, 60). Motile by
means of a single, polar flagellum on one or
both ends. Gram-negative.
Gelatin colonies: Entire, conve.x, circular,
moist, colorless.
256
ORDER I. PSEUDOMONADALES
Gelatin stab: Growth along entire stab.
No liquefaction (Dimitroff, op. cit., 12,
1926, 31). Active liquefaction (Giesberger,
op. cit., 1936, 65).
Agar colonies: Dew drop, convex, entire,
moist, colorless.
Agar slant: Dew drop, isolated colonies.
Broth: Cloudy, no flocculation.
Uschinsky's protein-free medium: Abun-
dant growth.
Litmus milk: No growth.
Loeffler's blood serum: Convex, isolated
dew drop colonies. No liquefaction.
Potato: No growth.
Methyl red negative; acetylmethylcar-
binol not produced.
Indole not produced.
Hydrogen sulfide not produced.
No acid or gas from carbohydrates.
Lactates and citrates utilized (Giesberger,
loc. cit.).
Nitrites not produced from nitrates.
Aerobic, facultative.
Optimum temperature, 35° C.
Source: Isolated from mud on an oyster
shell.
Habitat: Probably the muddy bottom of
brackish water.
6. Spirillum minus Carter, 1888. (Car-
ter, Sci. Mem. Med. Officers Army India, 3,
1888, 45; Spirochaeta muris Wenyon, Jour.
Hyg., 6, 1906, 580.)
mi'nus. L. comp.adj. minus less, smaller.
Description taken from Adachi (Jour.
Exp. Med., S3, 1921, 647) and Giesberger
(Inaug. Diss., Delft, 1936, 67).
Short thick cells, 0.5 by 3.0 microns,
having 2 or 3 windings which are thick,
regular and spiral. Actively motile by means
of bipolar tufts of flagella. Gram-negative.
Has not been cultivated on artificial
media.
Aerobic, facultative.
Pathogenic for man, monkeys, rats, mice
and guinea pigs.
This species is regarded by some as a
spirochaete. Because of its habitat and
wide distribution it has been described
under many different names. It is possible
that some of these names indicate varieties
or even separate species. See Beeson (Jour.
Amer. Med. Assoc, 123, 1943, 332) for im-
portant literature.
Source: Found in the blood of rats and
mice.
Habitat: A cause of rat-bite fever. Widely
distributed.
7. Spirillum kutscheri Migula, 1900.
{Spirillum undula majus Kutscher, Cent. f.
Bakt., I Abt., 18, 1895, 614; Migula, Syst. d.
Bakt., 2, 1900, 1024.)
ku'tsche.ri. M.L. gen. noun kutscheri of
Kutscher; named for K. H. Kutscher, the
German bacteriologist who first isolated
this organism.
Stout, spiral -shaped threads 1.5 microns
in diameter. Wave lengths, 10.5 to 12.5 mi-
crons; width, 3.0 to 4.5 microns. The spiral
form may be lost on continued cultivation.
Volutin granules present. Motile by means
of tufts of flagella at the poles. Gram-nega-
tive.
Gelatin plate: Surface colonies are trans-
parent and round; deep colonies are dark
brown.
Gelatin stab: Slow liquefaction.
Agar colonies grow poorly; granular.
Deep colonies yellowish green to dark
brown.
Agar slant: Delicate, transparent growth.
Potato: Limited growth.
Utilizes malic and succinic acids.
Grows well on peptone broth. Also
utilizes ammonia compounds.
Catalase-positive.
Optimum temperature, between 22° and
27° C.
Source: Isolated from putrid materials
and liquid manure.
Habitat: Putrefying liquids.
8. Spirillum volutans Ehrenberg, 1832.
(Prototype, Vibrio spirillum Miiller, Ani-
malcula infusoria et marina, 1786; Ehren-
berg, Physik. Abhandlungen d. k. Akad.
Berlin, 1832, 38.)
vo'lu.tans. L. v. voluto to tumble about;
L. part. adj. volutans tumbling about.
The largest of the spirilla; probably first
seen by Miiller.
Spiral-shaped cells 1.5 microns in di-
ameter. Wave length, 13 to 14 microns;
FAMILY VII. SPIRILLACEAE
257
width 4 to 5 microns. Slightly attenuated
ends. Dark granules of volutin are present
in the cytoplasm. Motile, possessing a tuft
of ten to fifteen fiagella at each pole. Gram-
negative.
Migula (Syst. d. Bakt., 2, 1900, 1025)
reports that this species has not been
cultivated on artificial media and that the
cultures described by Kutscher (Ztschr. f.
Hyg., 20, 1895, 58) as Spirillum volutans
are of a different species. Vahle (Cent. f.
Hakt., II Abt., 25, 1910, 237) later describes
the cultural characters of an organism which
he regards as identical with Kutscher's
organism. Giesberger (Inaug. Diss., Delft,
1936, 65) saw what he felt was the true
Spirillum volutans but could not cultivate
it.
Optimum temperature, 35° C.
Habitat: Stagnant water.
9. Spirillum lipoferum Beijerinck, 1925.
(Azotobacier spirillum Beijerinck, Kon.
Akad. Wetensch. Amsterdam, 30, 1923, 431,
([uoted from Giesberger, Inaug. Diss.,
Delft., 1936, 24; Beijerinck, Cent. f. Bakt.,
II Abt., 63, 1925, 353.)
li.po'fe.rum. Gr. noun lipus fat; L. v.fero
to carry; M.L. adj. lipoferus fat-bearing.
Curved cells with one-half to one spiral
turn. Contain minute fat droplets which
ma}^ deform the cells. Motile by means of a
tuft of polar fiagella. Gram-negative.
Calcium malate agar colonies: Circular,
small, transparent, dry. The malate is oxi-
dized to calcium carbonate. Cells contain
fat droplets.
Peptone agar colonies: More abundant
development. Cells lack fat droplets and
are typically spirillum in form.
Glucose peptone broth: Cells actively
motile with large fat droplets.
Fixes atmospheric nitrogen in partially
pure cultures, i.e., free from Azotobacter
and Clostridium (Beijerinck, loc. cit.).
Schroder (Cent. f. Bakt., II Abt., 85, 1932,
17) failed to find fixation of nitrogen when
she used cultures derived from a single cell.
Aerobic.
Optimum temperature, 22° C.
Beijerinck regards this as a transitional
form between Spirillum and Azotobacter.
Giesberger (op. cit., 1936, 64) thinks it a
Vibrio.
Habitat: Garden soil.
Genus VIII. Paraspirillum Dobell, 1912.*
(Arch. f. Protistenk., 2^, 1912, 97.)
Pa.ra.spi.ril'lum. Gr. pref. para beside; M.L. neut.n. Spirillum a genus of bacteria;
M.L. neut.n. Paraspirillum Spirillum-like (organisms).
Cells spiral or S-shaped, tapering toward the ends, wdth a well marked thickening toward
the middle of the body; resemble much elongated and spirally twisted spindles. Motile by
means of a single, polar flagellum. Found in fresh water.
Dobell {loc. cit.) believes that this organism belongs to the Spirillaceae rather than to the
Spirochaetaceae .
The type species is Paraspirillum vejdovskii Dobell.
1. Paraspirillum vejdovskii Dobell,
1912. (Arch. f. Protistenk., 24, 1912, 97.)
vej .dov'ski.i. M.L. gen. noun vejdovskii of
Vejdovsky; named for Prof. F. Vejdovsky.
Spiral or S-shaped rods, tapering toward
the ends, 8 to 25 microns in length, averag-
ing 15 microns. Width, in the middle, 1.5 to
to 2.0 microns. A definite spherical to ellip-
soidal nucleus is present. The cytoplasm
immediately about the nucleus is hyaline
or very finely granular. Volutin granules
are numerous between the hyaline area and
the ends of the cell. Locomotion is screw-
like, resembling that characteristic of spe-
cies of Spirillum. The motion is reversible,
and cells may swim in either direction. In
motion the cell seems to be rigid, but it may
increase or decrease the amount of bending.
Sometimes the cell is much-curved, at other
Prepared by Prof. R. E. Buchanan, Iowa State College, Ames, Iowa, July, 1952.
258 ORDER I. PSEUDOMONADALES
times it is almost straight. A single, polar This organism has not been cultivated,
flagellum is demonstrable; such a flagellum Source: Encountered only once in a cul-
may occur at each end. Division of the cell ture of Oscillatoriae in water from the River
is transverse and is preceded by a division Granta near Cambridge, England,
of the nucleus. Habitat: Fresh water.
Genus IX. Selenonionas von Prowazek, 1913*
(Von Prowazek, Cent. f. Bakt., I Abt., Orig., 70, 1913 (July), 36; Selenomastix
Woodcock and Lapage, Quart. Jour. Micro. Sci., S9 (N.S.), 1913 (November), 433.)
Se.le.no. mo'nas. Gr. noun selene the moon; Gr. noun monas a unit, monad; IM.L. fem.n.
moon monad.
Cells kidney- to crescent-shaped with blunt ends. Motile by means of a tuft of flagella
attached to the middle of the concave side. The flagella are thicker at the base than at the
free end and are usually about one and a half times as long as the cell. Gram-negative.
Anaerobic. Parasites found in the alimentary tracts of mammals, including man.
Three species are described, and it is possible that when comparative studies are made
the three will be found to belong to but a single species. On the other hand it is equally pos-
sible that not only these three but also additional species will be recognized (Lessel and
Breed, Bact. Rev., 18, 1954, 167).
The type species is Selenomonas palpitans Simons.
Key to the species of genus Selenonionas.
I. Found in the coeca of guinea pigs.
1. Selenonionas palpitans.
II. Found in the human mouth cavity.
2. Selenonionas sputigena.
III. Found in the rumen juices of ruminants.
3. Selenomonas ruminantiuni .
1. Selenomonas palpitans Simons, 1922. a true nucleus, dividing as the cell divides.
(Guinea pig selenomonad, da Cunha, Brasil Boskamp {ibid., 65) was unable to deter-
Medico, 29, 1915, 33; Selenomonas palpitans mine whether this division was mitotic or
Simons {nomen dubium), Cent. f. Bakt., I amitotic. With Giemsa's stain the cyto-
Abt., Orig., 57, 1921, 50; Simons, inBoskamp, plasm is blue whereas the chromatin ma-
ibid., 88, 1922, 58.) terial, the cell membrane and the flagella
pal 'pi. tans. L. part. adj. palpitans trem- stain red. Gram-negative,
bling. Anaerobic (?) as presumed by Boskamp
Description taken from Boskamp {loc. (^^^^^ ^^^^^ fron^ the fact that these
f^'^i-)- organisms, in feces, died quickly when
Kidney- to crescent-shaped cells with ^^^^^^^ ^^ ^j^. ^.j^^^ likewise did not grow
blunt ends, 1.8 to 2.3 by 6.8 to 9.1 microns. ,. „ „ j-v.^^ , ^^^;„
' . / ,^ , a ^^ ^u aerobically on ordmary media.
Motile by means of a tuft of flagella on the ., t^ j • xi_ c
•' . . . ^, ,1 rru a 11 Source: Found in the cecum oi a guinea
concave side of the cell. The flagella are
thicker at the base than at the free end and P*^
are
usually about one to one and a half times
Habitat : Found in the ceca of guinea pigs.
long as the cell. A highly refractive Not found in the buccal cavity, the stomach
granule is found on the concave side at the or the small intestine. Decrease rapidly in
base of the tuft of flagella; this granule number in the upper large intestine and dis-
stains with nuclear stains and appears to be appear entirely in the lower part.
* Prepared by Mr. Erwin F. Lessel, Jr., Cornell University, Geneva, New York, January,
1954.
FAMILY VII. SPIRILLACEAE
259
2. Selenonionas sputigena (Fliigge,
1886, emend. Muhlens, 1909) Dobell, 1932.
(Spirillum sputigenum Fliigge, Die Mikro-
organismen, 2 Aufl., 1886, 387; Muhlens,
Cent. f. Bakt., I Abt., Orig., 48, 1909, 524;
Selenomonas sputigena Boskamp (nomen
provisorium) , Cent. f. Bakt., I Abt., Orig.,
88, 1922, 70; Dobell, Antony van Leeuwen-
hoek and His "Little Animals". New York,
1932, 239, plate XXIV, and 245, footnote 2;
Vibrio sputigenus Pr^vot, Man. de Classif.
des Bact. Anaer., Paris, 1« ed., 1940, 85; not
Vibrio sputigenus Bergey et al., Manual,
1st ed., 1923, 80.)
spu.ti'ge.na. L. noun sputum spit, spu-
tum; L. V. gigno to produce; M.L. adj.
sputigenus sputum-produced.
Description taken from Muhlens (op. cit.,
1909, 524). Also see Hoffmann and von Pro-
wazek (Cent. f. Bakt., I Abt., Orig., 4i ,
1906, 820), von Prowazek (ibid., 70, 1913, 36)
and Macdonald (Thesis, Univ. of Toronto,
1953,95 pp.).
Comma- and crescent-shaped cells,
thicker and longer than the cholera vibrio,
occasionally occurring in pairs in the form
of an S. Motility is vibratory, rotating,
whirling and boring in nature. Dobell (op.
cit., 1932, 245) feels that the type of motility
depicted in Leeuwenhoek's drawing (ibid.,
239, plate XXIV, Fig. B, with motion shown
in C to D) is so characteristic of Spirillum
sputigenum that the organism labelled Fig.
B by Leeuwenhoek is, in all probability.
Spirillum sputigenum Miller. Hoffmann and
von Prowazek {op. cit., 1906, 820) describe
this organism as a crescent-shaped rod with
a thick flagellum that appears to be attached
on the concave side. Muhlens {op. cit., 1909,
525) reports 1 to 3 flagella, the majority of
the organisms having a single thick flagel-
lum (a tuft of flagella) on the concave side.
Von Prowazek {op. cit., 1913, 36) later shows
excellent figures of these thick flagella
treated with Giemsa's stain. These figures
show that the thick flagellum is really a tuft
of flagella which may separate like the
bristles of a paint brush. Stains pale red
with Giemsa's stain.
Horse-serum agar stab: Fine, hazy
colonies develop in the low portions. Growth
begins in 1 to 3 days as fine, cloudy colonies
with somewhat thicker, yellowish centers
and increases to a thick streak, opaque in
the center and cloudy-transparent at the
edges.
Kutscher's placenta agar: Good growth.
Anaerobic growth in the lower two-thirds
of stab and shake cultures. No visible gas
produced.
Serum broth: No growth.
The following characters are from Mac-
donald {op. cit. 1953) :
Blood agar: Growth occurred only when
plates were reduced immediately after
streaking. Colonies were smooth, convex,
grayish yellow, and less than 0.5 mm in
diameter.
Difco thioglycollate broth: Growth heavy
and granular in 48 hrs. The best fluid me-
dium in which to maintain cultures.
Difco thioglj'coUate agar shake cultures:
Irregularly shaped, yellow colonies.
Litmus milk: Acid and coagulated.
Acid from glucose and sucrose. Slight
acid from mannitol.
Indole not produced.
Hydrogen sulfide not produced.
Nitrites produced from nitrates.
Optimum temperature, 37°C. Growth
range, from 20° to 45° C. The pH range for
growth is 4.5 to 8.6 with the best growth
between 5.5 and 8.6.
Not pathogenic for guinea pigs injected
subcutaneously or intracardially, nor for
mice injected intraperitoneally.
Distinctive characters: Early investiga-
tors described an organism of this type but
were unable to culture it, e.g. Lewis (Lancet,
Sept. 20, 1884, who regarded the organism
he found as identical with the cholera
vibrio) and Miller (The Microorganisms of
the Human Mouth, Philadelphia, 1890, 75) ;
Miller also gives an excellent discussion of
the early work. Muhlens {op. cit., 1909, 526)
described a variety of this species with
smaller cells.
While Macdonald describes this species
as peritrichous, he reports (personal com-
munication) that others have felt that his
electron micrographs could be interpreted
as showing a cluster of flagella attached at
the middle of the concave side of the cres-
cent-shaped cells, and that in dark field
examination he observed, as had earlier
workers, a single heavy flagellum attached
260
ORDER I. PSEUDOMONADALES
at the middle of the concavity. The so-called
nuclear body shows plainly in the electron
micrographs.
Source: Isolated from the buccal cavity of
man.
Habitat: Found in the buccal cavity.
3. Selenoinonas runiinantium (Certes,
1889) Wenyon, 1926. {Ancyromonas runiinan-
tium Certes, Bull. Soc. Zool., France, H,
1889, 70; Selenomastix ruminantium Wood-
cock and Lapage, Quart. Jour. Micro. Sci.,
59 (N.S.), 1913-1914, 433; Wenyon, Proto-
zoology. 1, 1926,311.)
ru.mi.nan'ti.um. L. part. adj. ruminans,
ruminantis ruminating; M.L. neut.pl.n.
ruminantia ruminants; M.L. pi. gen. noun
ruminantium of ruminants.
Rigid, crescent-shaped cells which meas-
ure 2 to 3 by 9.5 to 11 microns. Woodcock and
Lapage {op. cit., 434) state that the cells are
only slightly crescentic and never assume
the S shape as reported by Certes (op. cit.,
439); furthermore, they report that the
curve lies in but one plane. A tuft of flagella
which attains a length of 8.0 to 9.5 microns
springs from the center of the concavity.
The protoplasm stains homogeneously ex-
cept at the base of the flagella where a
deepl}^ staining mass is easily demonstrable.
Reproduction is by binary fission transverse
to the long axis of the cell and through the
flagellar region. Each half of the flagella
passes to one of the two pear-shaped daugh-
ter cells where it is attached near the blunt
end; later the flagella undergo an apparent
shift in position to the center of the con-
cavity.
Probably anaerobic but does not grow on
ordinary media either aerobically or an-
aerobically.
Woodcock and Lapage (op. cit., 445 ff.)
found ellipsoidal, non-motile organisms
mixed abundantly with the motile crescents
and felt that these might represent a stage in
the life history of the crescents although
they could not demonstrate this. Wenyon
{op. cit., 311) also thinks that a rounded
flagellate organism may be a stage of the
crescent-shaped organism, but he presents
no proof to support this conclusion.
Source: Found by Certes {op. cit., 70) by
microscopical examination of rumen juice
of cattle, sheep and deer. Later found by
Woodcock and Lapage {op. cit., 433) very
abundantly in the rumen juice of goats.
Habitat : Found as a predominant organ-
ism on microscopical examination of rumen
juices from herbivorous mammals.
Genus X. Myconostoc Cohn, 1875.*
(Beitrage z. Biol. d. Pflanzen, 1, Heft 3, 1875, 183.)
My.co.nos'toc. Gr. noun myces fungus; M.L. neut.n. Nostoc a genus of algae; i\LL. neut.n.
Myconostoc fungus nostoc.
Curved, colorless cells occurring singly or in curved or spiral chains. Embedded in small,
spherical, gelatinous masses. Found in fresh- or sulfur-water containing decomoosine
organic matter.
The type species is Myconostoc gregarium Cohn.
1. Myconostoc gregarium Cohn, 1875.
(Cohn, Beitrage z. Biol. d. Pflanzen, 1,
Heft 3, 1875,_ 183; Spirosoma gregarium
Migula, Syst. d. Bakt., ^, 1900, 960.)
gre.gar'i.um. L. adj. gregarius of or be-
longing to a flock or group.
Cells curved to comma-shaped, 1 by 5 to
10 microns, often joined together as spiral
chains which may resemble horse-shoes or
which may twist around each other to form
coiled, non-septate, non-motile, colorless
filaments. The filaments are usually en-
closed in a spherical, solid, microscopic,
gelatinous mass which measures 10 to 17
microns in diameter; these masses may
clump together and form a cluster, usually
on the surface of the water, which is visible
to the naked eye. Excellent illustrations de-
picting the nature of this species are shown
in Zopf (Die Spaltpilze, 3 Aufl., 1885, 23).
* Prepared by Mr. Erwin F. Lessel, Jr., Cornell University, Ithaca, New York. Septem-
ber, 1953.
FAMILY VII. SPIRILLACEAE
261
When the gelatinous mass disintegrates,
swarm cells are formed which are pre-
sumably polar flagellate. The individual
cells are granular and stain rather poorly.
During reproduction, the filaments be-
come somewhat elongated and expand the
gelatinous mass to an ellipsoidal shape. As
the gelatinous sphere undergoes transverse
fission, there is a concomitant division of the
filament, the daughter cells finally separat-
ing from each other. Lankester (Quart.
Jour. Micros. Sci., 13, (N.S.), 1873, 408)
erroneously believed this gelatinous form
to be a stage in the life cycle of a Spii-ilhim,
probably Spirillum undula Ehrenberg.
These organisms have not been cultivated
on artificial media.
Source : Found in the surface scum of sul-
fur-water in a jar with decomposing algae,
especially Spirogyra sp. (Cohn, op. cit.,
1875, 183). Also found by Migula {op. cit.,
1900, 960) in peat bogs between Weingarten
and Karlsruhe. Hansgirg (Osterr. Vot.
Ztschr., 38, 1888, 265) frequently found this
organism among his algal cultures in Bo-
hemia.
Habitat: Fresh-water ponds.
ORDER II. CHLAMYDOBACTERIALES BUCHANAN, 1917.*
(Buchanan, Jour. Bact., 2, 1917, 162.)
Chla.my.do.bac.te.ri.a'les. M.L. fern. pi. n. Chlamdobacteriaceae type family of the order
Chlamydobacteriales ; -ales ending to denote an order ; M.L. fern. pi. n. Chlamydobacteriales the
Chlam ydobacteriaceae order .
Colorless, alga-like bacteria which occur in trichomes. May or may not be ensheathed.
They may be unbranched or may show false branching. False branching arises from a lateral
displacement of the cells of the trichome within the sheath; this gives rise to a new trichome
so that the sheath is branched while the trichomes are separate. The sheaths may be com-
posed of an organic matrix impregnated with iron or manganese oxides, or they may be
composed of an organic matrix free from these oxides. Gram-negative. Reproduction may be
by flagellate swarm spores or by non-motile conidia. Endospores of the type found in Bacil-
lus are never developed. Fresh-water and marine forms.
Key to the families of order Chlamydobacteriales.
I. Conidia, when formed, are motile by means of a sub-polar tuft of flagella.
A. Possesses trichomes in which false branching may occur. Motile swarm cells may be
formed.
Family I. Chlamydobacteriaceae . p. 262.
B. Possesses ensheathed, unbranching trichomes which may be very long (0.5 cm).
Found in fresh water.
Family II. Peloplocaceae, p. 270.
II. Non-motile conidia are produced.
Family III. Crenotrichaceae, p. 272.
FAMILY I. CHLAMYDOBACTERIACEAE MIGULA, 1894.
(Arb. bakt. Inst. Hochschule, Karlsruhe, 1, 1894, 237.)
Chla.my.do.bac.te.ri.a'ce.ae. Gr. noun chlamys, chlamydis a cloak; Gr. neut.n. bac-
terium a small rod; -aceae ending to denote a family; M.L. fem.pl.n. CJdaui ydobacteriaceae
the family of the sheathed bacteria.
Bacteria which occur in trichomes and which frequently show false branching. Sheaths,
when present, may or may not be impregnated with ferric and/or manganese oxides. Cells
divide transversely. Swarm cells, if developed, are usually motile by means of a tuft of
flagella. Usually found in fresh water.
Key to the genera of family Chlamydobacteriaceae.
I. Trichomes surrounded by sheaths which are usually not impregnated with iron or
manganese oxides and which do not dissolve in hydrochloric acid. Large forms, mostl}^
sessile.
* Rearranged and revised by Prof. Robert S. Breed, Cornell University, Geneva, New
York, November, 1953; further revision, with the introduction of an additional family and
genera and species, by Prof. Dr. Herbert Beger, Institut fiir Wasser-, Boden- und Luft-
hygiene, Berlin-Dahlem, Germany, January, 1955.
262
FAMILY I. CHLAMYDOBACTERIACEAE 263
Genus I. Sphaerotilus, p. 263.
II. Trichomes surrounded by sheaths impregnated with oxides of iron or manganese which
dissolve in strong h.ydrochloric acid. Free-living or sessile.
A. Individual trichomes, each with a sheath.
Genus II. Leptothrix, p. 264.
B. Sheaths contain more than one trichome; the trichomes are sometimes in a fan-like
arrangement.
Genus III. Toxothrix, p. 269.
Genus I. Sphaerotilus Kiitzing, 1833.
(Kiitzing, Linnaea, 8, 1833, 385; Cladothrix Cohn, Beitr. z. Biol. d.
Pflanz., 1, Hefts, 1875, 185.)
Sphae.ro'ti.lus. Gr. noun sphaera a sphere; Gr. noun tilus anything shredded, flock,
down; M.L. mas.n. Sphaerotilus sphere down.
Attached or free-floating, colorless trichomes showing false branching, though this may
be rare in some species. When e.xamined under the electron microscope, the sheath shows a
homogeneous structure. Sheath may become j'ellowish or brown with the deposition of iron
oxide. The deposition of iron is dependent on environmental factors, not on the ph3'siologi-
cal ability to store iron. Trichomes consist of rod-shaped or ellipsoidal cells surrounded by
a firm sheath. Multiplication occurs both by non-motile conidia and by motile swarm cells,
the latter with a subpolar tuft of flagella. Gram-negative so far as known. Found in fresh
water.
The systematic positions of the species placed in Sphaerotilus, Leptothrix and related ge-
nera are uncertain. Pringsheim (Phil. Trans. Roy. Soc. London, Series B, 233, 1949, 605,
and Biol. Reviews, 24, 1949, 200) would combine some of the species now placed in Lepto-
thrix with Sphaerotilus nutans and broaden the definition of Sphaerotilus to include other
species here placed in Leptothrix and Clonothrix. However, Beger and Bringmann (Zent. f.
Bakt., II Abt., 107, 1953, 318) indicate differences in the structures of the sheaths of Sphaero-
tilus and Leptothrix and give other reasons why it may be better to keep the earlier groupings
as they have been.
The type species is Sphaerotilus natui^s Kiitzing.
1. Sphaerotilus natans Kiitzing, 1833. nitrogen, does not grow in the ordinary pep-
(Kiitzing, Linnaea, 8, 1833, 385; not (Sp/jae?o- tone solution, grows best with low concen-
tilus natans Sack, Cent. f. Bakt., II Abt., trations of meat extract (Zikes, Cent. f.
65, 1925, 116.) Bakt., II Abt., 4S, 1915, 529). See Stokes
na'tans. L. part. adj. wafa/is swimming. (Jour. Bact., 67, 1954, 278) for a recent study
Colorless, slimy trichomes which attain of the cultural and physiological charac-
a length of several millimeters. The tri- teristics of this species,
chomes are ensheathed, show false branching Distinctive characters: This species
and are either free-floating or attached at thrives in great tassels on solid substrata
one end by means of a small disc. The indi- covered by dirty running water. These tas-
vidual cells are cylindrical, 1 by 2 to 6 mi- sels are composed of trichomes of bacterial
crons, and vacuolated (Lackey and Wattie, cells held together by slim}^, tubular
U. S. Pub. Health Ser., Pub. Health Repts., sheaths. The latter maj' become softened
55,1940,975). and dissolved, releasing Pseudomonas-hke
]\Iultiplication occurs through the forma- swarm cells. The same organism grows in
tion of conidia within the sheath of the a quite different state in quiet waters
vegetative cells, from which thej' swarm out with only a little organic matter, forming
at one end, swim about for a time, then branched structures occurring in trichomes,
attach themselves to objects and develop the sheaths of which are not slimy. A third
into delicate trichomes. form is produced when ferrous compounds
Gelatin rapidly liquefied, requires organic and very little organic substance are pres-
264
ORDER II. CHLAMYDOBACTERIALES
ent. The sheaths become brittle and glass-
like in appearance by deposition of ferric
hj^droxide in a hard colloidal form. Prings-
heim (Endeavour, 11, 1952, 209) states that
under these conditions it is identical with
Leptothrix ochracea, which looks ochre-like
in bulk but never brown under the micro-
scope.
Source: Originally found in polluted
waters. May become a real nuisance in
sewage purification plants of the activated
sludge type (Lackey and Wattie, op. cit.,
1940, 975) and in streams polluted with sul-
fite liquor from pulp and paper mills
(Lackey, Mimeographed Rept., U. S.
Pub. Health Ser., 1941).
Habitat: Stagnant and running water,
especially sewage-polluted streams. Widely
distributed throughout the world in fresh
water.
2. Sphaerotilus dichotonius (Cohn,
1875) Migula, 1900. (Cladothrix dichotoma
Cohn, Beitr. z. Biol. d. Pflanz., 1, Heft 3,
1875, 185; Migula, Syst. d. Bakt., 2, 1900,
1033.)
di.cho'to.mus. Gr. adj. dichotomus di-
vided, forked.
The identity of this species as distinct
from Sphaerotilus natans has been ques-
tioned. In his text, Cohn reports the di-
ameter of the trichomes to be 0.3 micron.
This clearly is an error as his figures at
600X show the diameter of the trichomes to
be greater than the diameter of Bacillus
anthracis spores shown at the same mag-
nification. Such spores are 1.3 to 1.5 microns
in diameter.
Zikes (Cent. f. Bakt., II Abt., 43, 1915,
529) gives the following differential charac-
ters : Cells smaller than those of Sphaerotilus
natans, 1.5 to 2.5 microns; false branching
constant; grows best in high concentrations
of meat extract; will grow in ordinary pep-
tone solutions; can utilize inorganic nitro-
gen; liquefies gelatin slowly.
Source: Isolated by Cohn from water con-
taining Myconostoc.
Habitat: Comparatively unpolluted fresh
water capable of sustaining algae.
3. Sphaerotilus flviitans (Migula, 1895)
Schikora, 1899. {Streptothrixfluitans Migula,
in Engler and Prantl, Die natiirl. Pflanzen-
fam., 1, la, 1895, 38; Schikora, Ztschr. f.
Fischerei, 7, 1899, 1-28.)
flu'i.tans. L. part. adj. fluitans floating.
Verj thin, attached trichomes as much as
1 cm in length. The trichomes are sur-
rounded by a soft sheath from which almost
spherical conidia issue, usually attaching
themselves to the exterior of the sheath
where they multiply.
Source: Found attached to pieces of wood
and stems of plants in running water.
Habitat: Fresh water.
Genus II. Leptothrix Kiitzing, 1843.
(Kiitzing, Phycologia Generalis, 1843, 198; not Leptotrichia Trevisan, Reale
1st. Lombardo di Sci. e Lettere, Ser. 2, 12, 1879, 138.)
Lep'to.thrix. Gr. adj. leptus fine, small; Gr. noun thrix hair; M.L. fem.n. Leptothrix fine
hair.
Trichomes of cylindrical, colorless cells with a sheath at first thin and colorless, later
thicker, yellow or brown, encrusted with iron or manganese oxide. The oxides may be dis-
solved by dilute acid, whereupon the inner cells show up well. If the sheath contains man-
ganese oxide, it does not dissolve completel}^ in weak acids. When examined under the elec-
tron microscope, the sheath shows an alveolar structure. Multiplication is by cell division
with individual cells occasionally slipping out of the sheath as reproductive cells. These
are sometimes motile with a tuft of flagella. False branching may occur. Gram-negative and
not acid-fast so far as known. Usually found in fresh water.
The type species is Leptothrix ochracea Kiitzing.
Key to the species of genus Leptothrix.
I. Trichomes straight, not spirally twisted.
A. Trichomes free-floating and unbranched. Sheath thin.
FAMILY I. CHLAMYDOBACTERIACEAE 265
1. Trichomes 1 to 3 microns thick.
1. Lcplothrix ochrncea.
2. Trichomes 0.4 to 0.5 micron thick.
2. Leptothrix thermnlis.
B. Trichomes attached.
1. Attachment is by means of a hoklfast.
a. Trichomes arise singly, each from its own holdfast.
b. Sheath very thin, encrusted only at the base of the trichome.
3. Leptothrix sideropons.
bb. Sheaths thick.
c. Trichomes showing no or only a few false branches. Cells up to 1
micron thick.
4. Leptothrix discophora .
cc. Trichomes always with numerous false branches. Cells 1.4 microns
thick.
5. Leptothrix major.
aa. Numerous trichomes arising from a common holdfast.
b. Sheaths not tapering to the tip. Trichomes form sessile, hemispherical
clusters.
6. Leptothrix lopholea.
bb. Sheaths tapering to the tip. Trichomes usually form free-living colonies
in which the trichomes radiate like the spokes of a wheel.
7. Leptothrix echinata.
2. Attached by gelatinous masses.
8. Leptothrix epiphytica.
II. Trichomes spirally twisted (except in a variety of Leptothrix pseudovacuolata).
A. Epiphytic, growing twisted around thread-like algae.
9. Leptothrix voluhilis.
B. Free-living in water or on the surface of mud.
1. Trichomes very thin; sheaths tapering slowly to the tip, ending in a sharp point.
a. Cells 0.3 micron in diameter.
10. Leptothrix skujae.
aa. Cells 0.9 micron in diameter.
11. Leptothrix winogradskii.
2. Trichomes thick, sometimes not twisted; sheaths rounded at the tip.
12. Leptothrix pseudovamwlata.
1. Leptothrix ochracea (Roth, 1797) rounded by a delicate sheath which later
Kiitzing, 1843. {Conferva ochracea Roth, becomes yellow to brown in color. Sheath
Catal. bot. I, 1797, Table V, Fig. 2; also see alveolar, completely dissolving in dilute
Dillwyn, Syn. Conf., 1802, Table 62; Kiit- hydrochloric acid. When the sheath becomes
zing, Phycologia generalis, 1843, 198.) very thick, the trichomes slip out of the
o.chra'ce.a. Gr. noun ochra j-ellow ochre; sheath and secrete a new one so that many
M.L. adj. ochraceus like ochre. empty sheaths are found. Presumably polar
Description taken from Kiitzing (loc. cit.) flagellate swarm cells have been observed,
and Cataldi (Estudio Fisiologico y Siste- Not acid-fast. Gram-negative,
mdtico de Algunas Chlamydobacteriales. Gelatin: No liquefaction.
Thesis, University of Buenos Aires, 1939, Iron citrate and ammonium agar colonies:
58 and 66). Filamentous and spreading, with wavy
Cells rod-like, colorless, 0.8 to 1.0 micron. edges.
Motile. Trichomes long, free-floating, never Manganese acetate agar colonies: Fila-
attached to a substrate, never branching, 1 mentous, not very large,
micron in thickness. Young trichomes sur- Iron citrate and ammonium agar slant :
266
ORDER II. CHLAMYDOBACTERIALES
Growth very abundant, spreading over the
entire surface; iridescent.
Peptone and manganese acetate broth:
Abundant growth in the form of loose fiakes.
Indole not produced.
Hydrogen sulfide not produced.
Acetylmethylcarbinol not produced.
Nitrites produced from nitrates.
Optimum temperature, 28° C.
Optimum pH, 8.0.
Aerobic, but growth favored by the pres-
ence of CO2 .
Habitat: Found in iron-bearing, fresh
waters; widely distributed.
2. Leptothrix thermalis (Molisch, 1925)
Dorff, 1934. (Chlamydothrix thermalis Mo-
lisch, Sc. Rept. Tohoku Imp. Univ., 4 ser.
Biol., Sendae, Japan, 1, 1923, 135 (or pos-
sibly 1, 1925, 146); Dorff, Die Eisenorganis-
men, Pflanzenforschung, Heft 16, 1934, 38.)
ther.ma'lis. Gr. noun therme heat; M.L.
adj. thermalis pertaining to heat.
Unbranched trichomes, 0.4 to 0.5 micron
thick, united in bundles. Surrounded by a
sheath which may store iron and turn
brown.
Source : From warm and hot spring waters
in Japan.
Habitat: Found in warm and hot (37° to
74° C.) spring waters.
3. Leptothrix sideropous (Molisch,
1910) Cholodny, 1926. {Chlamydothrix sider-
opous Molisch, Die Eisenbakterien, 1910,
14; Cholodny, Die Eisenbakterien, Pflanzen-
forschung, Heft 4, 1926, 25.)
si.de'ro.pous. Gr. adj. sideropus or
sideropous iron-footed.
Description taken from Molisch {op. cit.
1910, 14) and Cataldi (Estudio Fisiologico
y Sistemdtico de Algunas Chlamydobac
teriales. Thesis, University of Buenos Aires
1939, 62 and 66).
Cells rod-shaped, 0.5 to 0.8 micron
Motile. Trichomes short and unbranched
Sheath very thin and colorless, giving an
iron reaction only at the base of the tri
chome. Attached by a broad holdfast which
gives a marked iron reaction. Not acid-fast.
Gram-negative.
Gelatin: No liquefaction.
Iron citrate and ammonium agar colonies :
Very filamentous. Colonies and filaments
encompassed by a spattering of rust-colored
spots.
Manganese acetate agar colonies: Large
and filamentous, the filaments being
strongly colored.
Iron citrate and ammonium agar slant:
Growth in the form of isolated colonies;
strongly colored.
Manganese acetate agar slant: Abundant
growth which adheres to the medium except
in those places covered with water of con-
densation.
Peptone and manganese acetate broth:
Firm pellicle with a metallic sheen.
Indole not produced.
Hydrogen sulfide not produced.
Acetylmethylcarbinol not produced.
Nitrites produced from nitrates.
Optimum temperature, between 25° and
28° C.
Optimum pH, 8.0.
Aerobic; growth not favored by the pres-
ence of CO2 .
Habitat: Found growing on the surfaces
of objects submerged in water; widely dis-
tributed.
4. Leptothrix discophora (Schwers,
1912) Dorff, 1934. (Megalothrix discophora
Schwers, Cent. f. Bakt., II Abt., 33, 1912,
273; Leptothrix crassa Cholodny, Cent. f.
Bakt., II Abt., 61, 1924, 292; Dorff, Die
Eisenorganismen, Pflanzenforschung, Heft
16, 1934, 31.)
dis.co'pho.ra. Gr. noun discus a disc; Gr.
adj. phorus bearing; M.L. adj. discophorus
disc-bearing.
Description taken from Schwers (op. cit.,
1912, 273) and Cataldi (Estudio Fisiologico
y Sistemdtico de Algunas Chlamydobac-
teriales. Thesis, University of Buenos
Aires, 1939, 60 and 66).
Cells 0.5 by 0.8 micron. Motile. Tri-
chomes long, slender, articulated, composed
of elements of varjdng length showing
false branching (Cholodny, Cent. f. Bakt.,
II Abt., 61, 1924, 297). Usually attached to
a submerged substrate, but may be free-
floating. A sheath, thick (10 to 15 microns)
at the base, tapering toward the free tip
FAMILY I. CHLAMYDOBACTERIACEAE
267
and heavily impregnated with iron oxide,
surrounds the trichomes. Reproduction by
motile swarm cells liberated from the tip
and also by the emergence of the trichome
from the sheath, with subsequent breaking
up into individual, non-motile cells. Not
acid-fast. Gram-negative.
Gelatin not liquefied.
Iron citrate and ammonium agar colonies:
More or less rounded, with oily inclusions,
filamentous border.
Manganese acetate agar colonies: Fila-
mentous growth, the filaments being rather
large and showing false branching.
Iron citrate and ammonium agar slant:
Growth only in the water of condensation,
rarely on the slant.
Peptone and manganese acetate broth:
Abundant growth in the form of loose flakes.
Indole not produced.
Hydrogen sulfide not produced.
Acetylmethylcarbinol not produced.
Nitrites produced from nitrates.
Optimum temperature, between 25° and
28° C.
Optimum pH, 8.5.
Aerobic; growth not favored by the pres-
ence of CO2 .
Habitat : Found in fresh water; widely dis-
tributed.
5. Leptothrix major Dorff, 1934. (Dorff
Die Eisenorganismen, Pflanzenforschung,
Heft 16, 1934, 35; also see Beger and Bring-
mann,Zent. f.Bakt., IIAbt., 107, 1953,323.)
ma'jor. L. comp.adj. major larger.
Trichomes, up to 1 and more cm in length,
attached by a holdfast, richly branched,
forming tufts. Trichomes composed of rod-
like cells, 1.4 by 5 to 10 microns, which con-
tain small false vacuoles. Giant cells up to
75 microns in length. Two trichomes may be
found in the same sheath. The sheath may
be as much as 12 microns in thickness, taper-
ing to the tip, storing manganese and iron.
Resembles the sheath of Leptothrix disco-
phora but is firmer in texture. Light to dark
l)rown in color.
Source: From the Spree River near Berlin.
Habitat: Found in fresh- water streams.
6. Leptothrix lopholea Dorff, 1934. (Die
Eisenorganismen, Pflanzenforschung, Heft
16, 1934, 33.)
lo.pho'le.a. Gr. noun lophus a crest;
M.L. dim. adj. lopholeus somewhat crested
or tufted.
Short, slender unbranched trichomes,
uniform in diameter, attached to a sub-
strate, 5 to 13 trichomes arising from a
common holdfast. Trichomes 20 to 33 mi-
crons long, cells 0.5 by 1.0 to 1.3 microns.
Sheaths composed of iron oxide; dissolve
completely in dilute hydrochloric acid.
Trichomes slip out of the sheath as in
Leptothrix: ochracea.
Habitat: Water.
7. Leptothrix echinata Beger, 1935.
(Zent. f. Bakt., II Abt., 92, 1935, 401.)
e.chi.na'ta. Gr. noun echinus the hedge-
hog; M.L. adj. echinatus like the hedgehog,
bristly.
Similar to the preceding species but oc-
curring in larger colonies, 20 to 50 trichomes
arising from a common holdfast. Trichomes
are shorter (9 to 10 microns).
Sheath is thicker at the base and tapers
toward the free tip of the trichome, which is
slightly spiral. The sheath contains an or-
ganic matrix visible after treatment in di-
lute hydrochloric acid.
Habitat: Found in water, especially in
manganese-bearing waters.
8. Leptothrix epiphylica (Migula, 1895)
Schoenichen and Kalberlah, 1900. (Strepto-
thrix epiphytica Migula, in Engler and
Prantl, Die natiirl. Pflanzenfam., /, la,
1895, 36 and 38; Chlamydothrix epiphytica
Migula, Syst. d. Bakt., 2, 1900, 1033; Schoe-
nichen and Kalberlah, Eyferth's Einfachste
Lebensformen, 3rd ed., 1900, 46.)
e.pi.phy'ti.ca. Gr. prep, epi' upon; Gr.
noun phytum plant; M.L. adj. epiphyticns
epiphytic, growing on plants.
Chains of cells enclosed in short, colorless
trichomes which are surrounded by thick,
gelatinous masses; the gelatinous masses are
attached to algae, but never in groups or
clusters.
Habitat: Widely distributed in fresh
water containing algae.
268
ORDER II. CHLAMYDOBACTERIALES
9. Leptothrix volubilis Cholodny, 1924.
{Lynghya epiphytica Hieronymus, in Kirch-
ner, in Engler and Prantl, Die naturl.
Pflanzenfam., 1, la, 1898, 67; Cholodny,
Zent. f. Bakt., II Abt., 61, 1924, 292;
Chlamydothrix epiphytica Naumann,
Ber. d. deutsch. bot. Gesellsch., 46, 1928,
141; not Chlamydothrix epiphytica Migula,
Syst. d. Bakt., 3, 1900, 1033; Leptothrix
epiphytica Dorff, Die Eisenorganismen,
Pflanzenforschung, Heft 16, 1934, 32; not
Leptothrix epiphytica Schoenichen and Kal-
berlah, Ej'ferth's Einfachste Lebensfor-
men, 3rd ed., 1900, 46.)
vo.lu'bi.lis. L. adj. volubilis twisting
spirally around a support, twining.
Cells rod-shaped and colorless, measuring
1 by 2 microns. The cells are enclosed in
long, cylindrical, unbranched trichomes
which grow in a spiral fashion around
threads of Oe(?o^om?/m, Tolypothrix, etc. The
bacterial trichomes are, in turn, surrounded
by cjdindrical, ochre-yellow sheaths, about
3 microns in diameter, which are encrusted
with iron. The cells m&y leave the sheaths as
in Leptothrix ochracea.
Habitat : Found in fresh water containing
algae.
10. Leptothrix skujae Beger, 1953.
{Leptothrix tenuissima Skuja, Symbolae
Botanicae Upsaliensis, 9, 1948, 33; not
Leptothrix tenuissima Naegeli, in Kiitzing,
Species Algarum, 1849, 265; Beger, in Beger
and Bringmann, Zent. f. Bakt., II Abt.,
107, 1953,331.)
sku'jae. M.L. gen. noun skujae of Skuja;
named for H. Skuja, the Swedish algologist
who first described this species.
Unattached trichomes, generally without
false branching, spirally wound together,
0.3 to 0.4 micron in diameter. The sur-
rounding sheath is as much as 18 microns in
diameter and tapers toward the tip. Cells
rod-shaped and colorless, with a few gran-
ules lying in chains.
Resembles Leptothrix discophora Dorff.
Source: From Store Halsjon, Prov. Upp-
land, Sweden. Found between other water
plants and in the plankton.
Habitat: Found near the shore in lakes.
11. Leptothrix winogradskii Cataldi,
1939. (Estudio Fisiologico y Sistemdtico de
Algunas Chlamydobacteriales. Thesis, Uni-
versity of Buenos Aires, 1939, 58.)
wi.no.grad'ski.i. M.L. gen. noun wino-
gradskii of Winogradsky; named for S.
Winogradsky, a Russian bacteriologist.
Cells 0.9 micron in diameter. Motile,
presumably polar flagellate. Trichomes very
long, never attached. Sheath 1.5 microns
thick. Not acid-fast. Gram-negative.
Gelatin not liquefied.
Iron citrate and ammonium agar colonies:
Very filamentous, terminate in spirals,
lusterless red.
Manganese acetate agar colonies: Very
filamentous, red to bright chestnut in color.
Iron citrate and ammonium agar slant:
Colonies quite large and distinct, pale
white.
Manganese acetate agar slant: Filaments
long, red to chestnut-colored and inter-
twined much as are cotton fibers.
Peptone and manganese acetate broth:
Filaments quite long and intertwined like
cotton fibers.
Indole not produced.
Hydrogen sulfide not produced.
Acetylmethylcarbinol not produced.
Nitrites produced from nitrates.
Optimum temperature, 37° C.
Optimum pH, between 5.0 and 9.8.
Aerobic; growth favored by the presence
of COo .
Source: Isolated from fresh water in the
neighborhood of Buenos Aires.
Habitat: Presumably widely distributed.
12. Leptothrix pseudovacuolata (Per-
filiev, 1925) Dorff, 1934. (Spirothrix pseudo-
vacuolata Perfiliev, Verh. d. Int. Verein. f.
theor. u. angew. Limnologie, 1925, Stutt-
gart, 1927; Dorff, Die Eisenorganismen,
Pflanzenforschung, Heft 16, 1934, 36.)
pseu.do. va.cu.o.la'ta. Gr. adj. pseudes
false; L. adj. vacuus empty; M.L. noun
vacuola a vacuole; M.L. adj. pseudovacuola-
tus having false vacuoles.
FAMILY I. CHLAMYDOBACTERIACEAE 269
Trichomes, 85 to 250 microns in length, Cells rounded at the ends, thin-walled,
unbranched, spirally wound, occasionally granular, 1.7 to 2.8 by 3.5 to 30 microns,
straight. Strongly encrusted with ferric Apparently heterotrophic,
hydroxide. Spirals 20 to 24 microns from Habitat: Found in bottom muds of deep
crest to crest. lakes with verj' low oxygen content.
Genus III. Toxothrix MoUsch, 1925.
(Molisch, Sci. Rept. Tahoku Imp. Univ., 4 Ser., Biol., 1925, 144; Cryptothrix Perfiliev, Zur
Mikroflora des Sapropels, Nachrichten des Sapropelkomitees Leningrad, 1, 1922.)
Tox'o.thrix. Gr. noun toxum a bow; Gr. noun thrix, trichis a thread; M.L. fem.n. Toxo-
thrix bent thread.
Trichomes composed of cylindrical, colorless cells with a thin primary sheath; the latter
soon becomes impregnated with iron oxide. The trichomes lie loosely, longitudinally to-
gether, in slightly spirally twisted rolls. The continued repetition of this process leads to the
development of a thick, secondary sheath from which parallel bundles may separate. False
branching maj^ occur. The sheaths do not completely dissolve in weak acids. Cells may slip
out of the sheath and may become motile swarm spores.
The type species is Toxothrix irichogenes (Molisch) Beger.
Key to the species of genus Toxothrix.
I. Long, unattached trichomes not in a gelatinous layer.
1. Toxothrix trichogenes.
II. Short trichomes lying in a gelatinous layer.
2. Toxothrix gelatinosa.
1. Toxothrix trichogenes (Cholodny, the sheaths. The number and diameters of
1924) Beger, 1953. (Leptothrix trichogenes the longitudinally placed trichomes are
Cholodny, Cent. f. Bakt., II Abt., 61, 1924, variable. Giant cells are frequently present.
296; T'oxo^Arz.T/e/TMgrmea Molisch, Sci. Rept. The sheaths, when empty, decay rapidly.
Tahoku Imp. Univ., 4 Ser., Biol., 1925, 13; The trichomes may slip out of their sheaths.
Chlamydothrix trichogenes Naumann, Zent. This species has been cultivated by Teich-
f. Bakt., II Abt., 78, 1929, 513; Sphaerotilus mann (Vergleichende Untersuchungen iiber
trichogenes Pringsheim, Biol. Reviews, die Kultur und Morphologie einiger Eisen-
Cambridge, 24, 1949, 234; Beger, in Beger organismen, Inaug. Diss., Prague, 1935).
and Bringmann, Zent. f. Bakt., II Abt., Beger and Bringmann (op. cit., 1953, 332)
107, 1953, 332.) report a form of this species in which the
tri.cho'ge.nes. Gr. noun thrix, trichis sheaths do not split; the cells are 0.5 by 2.0
hair; Gr. v. gennao to bear; M.L. adj. tri- to 4.0 microns.
chogenes hair-producing. Source: This species has been described
Found in trichomes up to 400 microns in from springs, wells, small rivers, water
length; composed of rod-shaped cells works and rice fields.
which are 0.5 by 1.0 to 2.0 microns. Sur- Habitat: Found in cool, fresh, iron-bear-
rounded by a tubular sheath which splits ing waters,
later so that arched, fan-shaped groups of
threads or irregular groups are formed as 2. Toxothrix gelatinosa Beger, 1953.
the trichomes grow in length. Do not lie in (Beger, in Beger and Bringmann, Zent. f.
a gelatinous layer. The tubular sheath is Bakt., II Abt., 107, 1953, 333.)
longitudinally and somewhat spirally stri- ge.la.ti.no'sa. L. part. adj. gelatus con-
ated with lines about 0.2 micron apart. No gealed; M.L. adj. gelatinosus gelatinous,
false branching. Iron oxide is deposited in The trichomes are up to 22 microns in
270
OEDER 11. CHLAMYDOBACTERIALES
length; including the sheath, they measure
1.5 to 1.7 microns in diameter. Several tri-
chomes arising from the same point and
each trichome developing a few false
branches produces a fan-shaped appearance.
All trichomes end at approximately the
same distance from the starting point. The
trichomes are bent so that the entire fan-
shaped group arches somewhat. The indi-
vidual cells are rod-shaped, 0.5 by 3.0 mi-
crons. Iron oxide is deposited in the sheath.
Around this, a gelatinous mass is formed,
as much as 22 microns in length and ovoid in
form. Iron is not deposited in this gelatinous
mass.
Source: Found on slides submerged in an
aquarium in Berlin in which Cahomba was
growing.
Habitat : Found in fresh water.
F.\]VIILY II. PELOPLOCACEAE BEGER, Fam. Nov.
Pe.lo.plo.ca'ce.ae. M.L. fem.n. Peloploca tj-pe genus of the family; -aceae ending to de-
note a family; M.L. fem.pl.n. Peloplocaceae the Peloploca family.
Long, unbranched trichomes usually enclosed in a thin, delicate sheath. Cells within the
trichomes, when in the living state, contain false vacuoles which are easily discerned by a
reddish gleam of light which they emit; the cytoplasm of the cell appears bluish white. Gen-
erally non-motile, but motile species may occur. Reproduction is by transverse fission of the
cells. Unattached forms found in fresh-water ponds with decomposing algae.
Key to the genera of family Peloplocaceae,
I. Trichomes lie parallel to each other in bundles or bands.
Genus I. Peloploca, p. 270.
II. Trichomes occur singh\
Genus II. Pelonema, p. 271.
Genus I. Peloploca Lauterborn, 1913.
(Allgem. bot. Ztschr., 19, 1913, 99.)
Pe.lo'plo.ca. Gr. adj. pellos or pelos dark-colored; Gr. noun place a twining, a braid or a
twist; M.L. fem.n. Peloploca dark-colored braid or twist.
Trichomes of cylindrical, colorless cells with no evident sheath. Occur as motionless
bundles or bands. Cells contain false vacuoles which emit a reddish gleam of light. Non-
motile. Occur in fresh-water ponds where Char a sp. is undergoing decomposition. Frequently
overlooked because the trichomes resemble plant fibers.
The type species is Peloploca undiilata Lauterborn.
1. Peloploca undulata Lauterborn,
1913. (Allgem. bot. Ztschr., 19, 1913, 99.)
un.du.la'ta. L. adj. undulatus undulated,
with waves.
Cells 6 to 10 microns long. The trichomes
are spirally twisted into wavy bundles that
are tightly wound together. The bundles
reach a length of 60 to 150 microns. Non-
motile.
Source : Found in Germany in ponds wliere
Chara sp. was growing.
Habitat: Presumably widelj' distributed
in fresh-water ponds.
2. Peloploca taeniata Lauterborn, 1913.
(Allgem. bot. Ztschr., 19, 1913, 99.)
tae.ni.a'ta. L. adj. taeniatus band-like.
Cells 3.0 to 4.0 microns long. Trichomes
united into rather broad, frequently twisted
bands. These may have the appearance of a
grating or lattice because of the presence
of pseudo vacuoles in the individual cells.
FAMILY II. PELOPLOCACEAE 271
The bands may reach a length of 700 mi- the surface of bottom mud along with
crons. Beggiatoa, Thiospira, Zoogloea and similar
Source : Found in Germany in ponds where types of bacteria.
C/tara sp. was growing; also found by Beger Habitat: Presumably widelj' distributed
(1954) in decomposing plant materials on in fresh-water ponds.
Genus II. Pelonema Lauterhorn, 1915.
(Verhandl. Naturhist.-med. Verein z. Heidelberg, N.F. 13, 1915, 408.)
Pe.lo.ne'ma. Gr. adj. pellos or pelos dark-colored; Gr. noun neina filament; M.L. neut.n.
Pelonema dark-colored filament.
Long, unbranching trichomes, occurring singly, which are either straight or spirally
twisted. The trichomes are enclosed in a very thin, delicate sheath. Non-motile, but may be-
come motile. Within the trichomes are C3'lindrical, colorless cells which contain one or
several to many false vacuoles which emit a reddish gleam of light. Found on the surfaces
of ponds and lakes w^hich contain decomposing algae and which are poorly aerated.
The type species is Pelonema temie Lauterborn.
Key to the species of genus Pelonema.
I. Cells contain a single false vacuole.
A. Cells 8 to 12 microns long; trichomes are straight and attain a length of up to 300
microns.
1. Pelonema tenue.
B. Cells 4 to 6 microns long; trichomes are straight and are 200 or more microns long.
2. Pelonema hyalinum.
II. Cells contain several to many false vacuoles.
A. Trichomes are straight, measuring up to 500 microns in length.
3. Pelonema pseudovacuolatum.
B. Trichomes are spirally twisted, reaching a length of 40 to 160 microns.
4. Pelonema spirale.
1. Pelonema tenue Lauterborn, 1915. hy. a. li'num. Gr. adj. ^T/ahwi/s of crystal,
(Verhandl. Naturhist.- med. Verein z. glass; M.L. adj. hyalinus hyaline.
Heidelberg, N.F. 13, 1915, 408.) Straight trichomes which measure 200 or
te'nu.e. L. adj. tenuis slender. more microns in length. The cells, 2 by 4 to
Straight trichomes, up to 300 microns 6 microns, contain a single false vacuole
long, which may become motile when the which is quite large and slightly refractive;
water in which they are growing is low in the vacuole is rectangular in shape and has
oxygen content. Cells are 2 by 8 to 12 mi- rounded edges.
crons. Each cell contains a single false Source: From water from Little Ploner
vacuole which nearly fills the cell; the Lake, Schleswig-Holstein, Germany,
vacuole is irregular in shape and emits but Habitat: Found in the upper algae-con-
a small reddish gleam of light. taining layers of deep fresh-water lakes.
Source: From pools in the Rheinebene,
Germany, where Chara was growing. 3. Pelonema pseudovacuolatum Lau-
Habitat: Presumably widely distributed terborn, 1915. (Verhandl. Naturhist.- med.
in fresh-water ponds and lakes which con- Verein z. Heidelberg, N.F. 13, 1915, 408.)
tain decomposing algae. pseu.do. va.cu.o.Ia'tum. Gr. adj. pseudes
false; L. adj. vacuus empty; M. L. noun
2. Pelonema hyalinum Koppe, 1923. vacuola a vacuole; M.L. adj. pseudovacuola-
(Pelonema hyalina (sic) Koppe, Archiv. f. tns having false vacuoles.
Hydrobiologie, H, 1923, 625.) Straight trichomes measuring up to 500
272 ORDER II. CHLAMYDOBACTERIALES
microns in length. The cells are 2 by 4 mi- 1915. (Verhandl. Naturhist.-med. Verein z.
crons and possess several small false vacu- Heidelberg, N.F. 13, 1915, 408.)
oles which are sharply but irregularly out- spi.ra'le. Gr. noun spira a spiral; M.L.
lined. The cytoplasm of the cells emits a adj. spiralis spiral.
marked bluish gleam of light. Spirally twisted trichomes, 1.0 to 1.5 by
Source: From pools and shallow lakes 40 to 160 microns, with a wave length of 8
which contained an abundance of decom- to 14 microns. The cells contain numerous,
posing algae. small but long false vacuoles.
Habitat: Found in fresh water. Source: From a pool in Germany rich in
Chara.
4. Peloneina (?) spirale Lauterborn, Habitat: Found in fresh water.
FAMILY III. CRENOTRICHACEAE HANSGIRG, 1888.
(Oesterr. bot. Ztschr., 36, 1888, 228.)
Cre.no.tri.cha'ce.ae. M.L. fem.n. Crenothrix type genus of the family; -aceae ending to
denote a family; M.L. fem.pl.n. Crenotrichaceae the Crenothrix familJ^
Trichomes attached to a firm substrate and show differentiation of base and tip. Un-
branched or show false branching. Sheaths may be thin, delicate and not encrusted with
oxides of iron or manganese, or they may be plainlj^ visible, thin and colorless at the tip and
thick and encrusted with iron or manganese oxides at the base. Cells disc-shaped to cylindri-
cal, dividing to produce spherical, non-motile conidia. Individual cells may also slip out of
the sheath to grow into new trichomes. Found in fresh and salt waters.
Key to the genera of family Crenotrichaceae.
I. Attached trichomes which are swollen at the free end.
A. Sheath thick, storing iron or manganese o.xides.
Genus I. Crenothrix, p. 272.
B. Sheath very delicate, always colorless.
Genus II. Phragmidiothrix, p. 273.
II. Attached trichomes which are tapered at the free end.
Genus III. Clonothrix, p. 274.
Genus I. Crenothrix Cohn, 1870.
(Beitr. z. Biol. d. Pflanz., 1, Heft 1, 1870, 108.)
Cre'no.thrix. Gr. noun crenus a fountain, spring; Gr. noun thrix, trichis a hair; M.L.
fem.n. Crenothrix fountain hair.
Trichomes attached to a firm substrate and swollen at the free end. Unbranched or show
false branching. The sheaths surrounding the trichomes are plainly visible, thin and color-
less at the tip and encrusted with iron or manganese o.xides at the base. Cells disc-shaped
to cylindrical, dividing to produce spherical, non-motile conidia of two types: micro- and
macroconidia. Individual cells may also slip out of the sheath and form new trichomes.
Found in stagnant and running waters which contain organic matter and iron salts.
The type species is Crenothrix polyspora Cohn.
1. Crenothrix polyspora Cohn, 1870. noun spon/s a seed; M.L. noun spora a spore;
(Beitr. z. Biol. d. Pflanz., 1, Heft 1, 1870, M.L. adj. polysporus many-spored.
108.) Trichomes long (up to 1 cm), articulated,
po.ly'spo.ra. Gr. adj. poly many; Gr. unbranched and sessile. There is consider-
FAMILY III. CRENOTRICHACEAE
273
able variation in the diameter of the indi-
vidual trichomes, the base measuring 1.5
to 5.0 microns and the swollen tip measur-
ing 6.0 to 9.0 microns. Each trichome is sur-
rounded b}' a colorless sheath which later
may become rust-colored and heavily en-
crusted, especially at the base, with deposi-
tions of ferric hydroxide and, to a lesser
e.xtent, manganese oxides. The ensheathed
trichomes may reach a diameter of 12 mi-
srons or more. Cells within the trichomes
are usually about 1.5 times as long as they
are wide and are more or less rectangular in
shape.
During reproduction the cells divide by
longitudinal and transverse fission into non-
motile conidia of two types: microconidia,
which are 1 to 2 microns in diameter, and
macroconidia, which measure about 5
microns in diameter; intermediate forms
may also occur. When the tip of the sheath
ruptures, the conidia are extruded; these
may attach themselves to some object and
grow into trichomes, or they may germinate
upon the exterior of the sheath from which
they were liberated, giving rise to new tri-
chomes attached to the surface of the older
one, thus simulating false branching. The
conidia often form a zoogloeal mass, but
only in the presence of dissolved iron.
In addition to the above-mentioned types
of reproductive cells, Cohn (ibid., 120) ob-
served a third structure which he condi-
tionall}'^ alluded to as a spore. These cells
originate from the swollen terminal cell
which is usually ellipsoidal in shape and
sometimes as much as seven times as long
as it is wide (3.67 by 26.25 microns). The
protoplasm of this terminal cell becomes
finely granular and eventually emerges from
the sheath. From these cells, short, color-
less Oscillaria-like trichomes are produced
which contain no more than eight cylindri-
cal cells measuring 5 to 6 by 10 to 12 microns.
The trichomes have a characteristic, slow,
gliding motion and are surrounded by a fine,
transparent membrane, but no sheath.
Subsequent authors, when describing this
species, have usually failed to mention this
third type of reproductive cell observed by
Cohn.
Cultivation: Has not been grown on
artificial media in pure culture. Grows
readily in water containing organic matter
regardless of the iron content of the water.
Related species: Cholodny believed
Clonothrix fusca to be identical with Creno-
thrix polyspora. However, Clonothrix ftisca
shows genuine false branching and produces
conidia by fission in only one plane so that
the trichomes taper toward the tip instead
of expanding (see Kolk, Amer. Jour. Bot.,
25, 1938, 11, for a differentiation of these
two species).
Comments: Zopf (Entwicklungsgesch.
Unters. (i. Crenoihrix polyspora, die Ursache
der Berliner Wasserkalamitat. Berlin, 1879,
2) regards Leptothrix kuehniana Rabenhorst
as identical with Crenothrix polyspora Cohn,
and there seems to be much evidence in
favor of considering the tw-o species as
identical. If Cohn's organism proves to be
identical with Rabenhorst's, then the spe-
cific epithet kuehniana has priority over
polyspora; however, until the relationship
of the two organisms has been clarified, the
name Crenothrix polyspora is retained here.
Source: This organism is wide-spread in
water pipes, drain pipes and springs w^here
the water contains iron. It frequently fills
pipes under such circumstances and causes
a real nuisance. Found by Cohn in samples
of water from springs in the neighborhood
of Breslau, Germany.
Habitat: Found in stagnant or running
waters containing organic matter and iron
salts. Harmless, but frequently becomes
bothersome in w^ater pipes and city water
supplies; grows as thick, brownish masses.
Genus II. Phragmidiothrix Engler, 1883.
(Vierter Ber. d. Commission z. wissensch. Unters. d. deutsch. Meere in
Kiel fiir 1877 bis 1881, I Abt., 1883, 187.)
Phrag.mi'di.o.thrix. Gr. noun phragma fence; Gr. noun eidus form, shape; Gr. noun
thrix, trichis hair; M.L. fem.n. Phragmidiothrix fence-like hair.
Trichomes are articulated, unbranched and attached, the free ends being swollen. Sur-
274
ORDER II. CHLAMYDOBACTERIALES
rounding the trichomes are very thin, delicate, colorless sheaths which do not store iron or
manganese compounds. The cells are small and disc-shaped and are uniform in size. Conidia
of the same diameter as the cells are produced. Found in salt water.
Hansgirg (Bot. Ztg., 49, 1891, 313) concluded that Phragmidiothrix should be included in
the genus Crenothrix, and that the genus Crenothrix should be divided into two sections,
Eitcrenothrix and Phragmidiothrix.
The type species is Phragmidiothrix viuUise'ptata (Engler) Engler.
1. Phragmidiothrix niultiseptata
(Engler, 1882) Engler, 1883. (Beggiatoa
jnultisepiata Engler, Verhandl. bot. Ver.
Brandenburg, U, 1882, 19; Engler, Vierter
Ber. d. Commission z. wissensch. Unters. d.
deutsch. Meere in Kiel fur 1877 bis 1881, I
Abt., 1883, 187; also see Zopf, Die Spaltpilze,
1883, 104.)
mul.ti.sep.ta'ta. L. mas.n. multus much;
L. adj. septatus fenced; M.L. adj. multisep-
tatus much-fenced, with many septa.
Colorless trichomes, several millimeters
long, which form grayish white tufts. The
trichomes are sessile; when young they are
1.5 microns wide, but when older they meas-
ure 2 to 3 microns at their bases and 5 to 6
microns at their tips. Very thin, delicate
sheaths which are not encrusted with iron
or manganese oxides surround the tri-
chomes. The cells are disc-shaped, their
width being 1.5 to 4.0 microns while their
length is only }4: to l^ this size. Each cell
has a very thin, colorless membrane and
some hyaline granules.
When mature, the cells in the upper por-
tion of the trichomes divide longitudinally
and transversely and form uniformly sized
conidia (1 micron in diameter). These co-
nidia may be extruded, may become free by
decomposition of the sheath, or they may
germinate within the sheath. The extruded
conidia may produce zoogloeal masses be-
fore they germinate.
Source: From the body of a crustacean
(Gammarus locusta) from sea water; also
found on seaweeds in polluted water on the
shores of the northern Adriatic.
Habitat : Found in polluted salt water.
Genus III. Clonothrix Roze, 1896.
(Roze, Jour, de Botanic, 10, 1896, 325; also proposed independently
by Schorler, Cent. f. Bakt., II Abt., 12, 1904, 689.)
Clo'no.thrix. Gr. noun clon, clonis twig, slip; Gr. noun thrix, trichis hair; M.L. fem.n.
Clonothrix twig hair.
Attached trichomes showing false branching as in Sphaerotihis. Sheaths organic, encrusted
with iron or manganese, broader at the base and tapering toward the tip. Cells colorless,
cylindrical. Reproduction by spherical conidia formed in chains by transverse fission of
cells; conidia formation acropetal, limited to short branches of the younger portions of the
trichomes.
The type species is Clonothrix putealis (Kirchner) Beger.
1. Clonothrix putealis (Kirchner, 1878)
Beger, 1953. {Glaucothrix putealis Kirchner,
Kryptogamen-Flora von Schlesien, 2, 1,
1878, 229; Clonothrix fusca Roze, Jour, de
Botanic, 10, 1896, 325; Beger, in Beger and
Bringmann, Zent. f. Bakt., II Abt., 107,
1953, 327.)
pu.te.a'lis. L. adj. putealis belonging to a
well.
Ensheathed trichomes, up to 0.6 mm long,
which show false branching and which taper
towards the tip; the bases of the trichomes
measure 7 microns and the tips measure 2
microns in diameter. The sheaths may be-
come encrusted with oxides of manganese
and/or iron, particularly those of manga-
nese. Sheath encrustations may reach a
thickness of 24 microns when manganese
oxides are prevalent and 10 microns when
iron oxides are abundant. Cells cj'lindrical
with rounded ends, 2 by 10 microns, be-
coming larger toward the base and smaller
toward the tips of the trichomes.
Multiplication by extrusion of single cells
FAMILY III. CRENOTRICHACEAE
275
or by rather uniform, spherical, non-motile
conidia formed into short trichomes in
chains of 2 to 6 or more, their diameters
being about 2 microns.
Historical: This organism was described
by Roze as a blue-green alga, but subse-
quent observers have failed to find pigment.
It was described independently by Schorler
(Cent. f. Bakt., II Abt., 12, 1904, 689) who
also gave it the name Clonothrix fusca.
Cholodny considered it identical with
Crenolhrix polyspora, but Kolk (Amer.
Jour. Bot., 25, 1938, 11) has clearly differ-
entiated these species.
Source : From a well in Proskau, Schlesien.
Habitat: Widely distributed in rivers
and streams with gravelly, manganese-
bearing bottoms; also found in water works
and pipe lines, where it may cause technical
difficulties. May occur in dark brown masses
that are large enough to be seen readily in
tap water.
ORDER III. HYPHOMICROBIALES DOUGLAS, Ordo Nov,
Hy.pho.mi.cro.bi.a'les. M.L. fem.pl. n. Hyphomicrobiaceae type family of the order;
-ales ending to denote an order; M.L. fem.pl.n. Hyphomicrohiales the Hyphomicrobiaceae
order.
Multiplication is by budding or by budding and longitudinal fission. Buds may be sessile
or may be borne at the tip of a slender filament which arises from the pole of a mature cell
or from a filament connecting two cells. Cells may occur singly or in pairs but are found more
commonly in aggregates. In some types the aggregates consist of groups of cells attached
to a surface by stalks which appear to radiate from a common holdfast; in others the ag-
gregates consist of free-floating cell groups in which the cells are attached to one another
by the filament engendered in the budding process. Branching of the filament may result
in groups which contain several hundred cells. Cells are ovoid, ellipsoidal, spherical or
pyriform. If motile, the cells possess a single polar flagellum. Specialized resting stages have
not been found. Gram-negative so far as known. Metabolism may be heterotrophic or
photosynthetic. Found in the mud and water of fresh-water ponds and streams; also para-
sitic on fresh-water Crustacea.
Key to the Families of Order Hyphomicrobiales.
I. Buds borne upon filaments.
Family I. Hyphomicrobiaceae, p. 276.
II. Buds sessile.
Family II. Pasteuriaceae, p. 278.
FAMILY I. HYPHOMICROBIACEAE BABUDIERI, 1950.
(Rendiconti dell'Istituto Superiore di Sanita, 13, 1950, 589.)
Hy.pho.mi.cro.bi.a'ce.ae. M.L. neut.n. Hyphomicrobium type genus of the family;
-aceae ending to denote a family; M.L. fem.pl.n. Hxjphomicrobiaceae the Hyphomicrobium
family.
These organisms occur mainly as free-floating groups in which the cells are attached to
one another by a slender, sometimes branched, filament. Daughter-cell formation is initi-
ated by the outgrowth of a filament from the pole of a mature cell or from some point on a
filament connecting two mature cells. The daughter cell is formed by enlargement of the tip
of the filament. Gram-negative.
Key to the genera of family Hyphomicrobiaceae.
I. Chemoheterotrophic. Motile.
Genus I. Hyphomicrobium, p. 277.
* New material prepared by and old material rearranged by Prof. H. C. Douglas, Depart-
ment of Microbiology, School of Medicine, University of Washington, Seattle, Washington,
December, 1953.
276
FAMILY I. HYPHOMICROBIACEAE
277
II. Photoheterotrophic. Non-motile.
Genus II. Rhodomicrohium, p. 277.
Genus I. Hyphomicrobium Stutzer and Hartleb, 1898.
(Mitteil. d. landwirtsch. Inst. d. k. Univ. Breslau, 1898; abst. in Cent. f. Bakt., II Abt.,
6, 1899, 678.)
Hy.pho.mi.cro'bi.um. Gr. noun hyphe thread; Gr. adj. micrus small; Gr. noun hius life;
M.L. neut.n. Hyphomicrobium thread (-producing) microbe.
Daughter cells may remain attached to the filaments which connect them to the mother
cells or may tear free of the filament as the result of active movement by means of a single,
polar flagellum. Gram-negative. Non-pigmented. Metabolism is chemoheterotrophic and
oxidative. Aerobic. Found in soil and in fresh water.
The type species is Hyphomicrobium vulgare Stutzer and Hartleb.
1. Hyphomicrobium vulgare Stutzer
and Hartleb, 1898. (Saltpeterpilz, Stutzer
and Hartleb, Cent. f. Bakt., II Abt., 3,
1897, 621; Stutzer and Hartleb, Mitteil. d.
landwirtsch. Inst. d. k. Univ. Breslau, 1898;
abst. in Cent. f. Bakt., II Abt., 5, 1899,
678.)
vul.ga're. L. adj. vxdgaris common.
Description taken from Stutzer and
Hartleb {loc. cit.), Kingma-Boltjes (Arch,
f. Mikrobiol., 7, 1936, 188) and Mevius
(Arch. f. Mikrobiol., 19, 1953, 1).
Mature cells are ovoid, measuring 0.5
by 1.0 nucron; immature cells are spherical.
Motile bj- means of a single, polar flagel-
lum. Daughter cells are borne on filaments
measuring approximately 0.2 micron or
less in diameter and varying in length from
one to several times the length of mature
cells. The predominant growth habit is that
of a dense clump of cells from which fila-
ments radiate outward. Branching of the
filament occurs but is not common.
Daughter cells may tear free of the fila-
ments and e.xist as single, motile cells with
motility sometimes persisting even after
the cell has produced a filament of consid-
erable length. Cells in pairs, connected
by a filament, are common. Gram-nega-
tive.
Gelatin: No growth.
Formate-nitrate agar or silica gel plates:
Colonies are colorless, 0.5 to 1.0 mm in di-
ameter, slightly elevated, entire.
Peptone agar colonies: Much smaller
than those above.
Peptone broth: Poor growth.
Formate-nitrate broth: Growth occurs
as a light, cream-colored, granular sedi-
ment.
Chemoheterotrophic, oxidative. Growth
occurs in mineral media at pH 7.0 to 7.5
with ammonium or nitrate as a nitrogen
source and formate, formaldehyde, meth-
anol, acetate or lactate as a carbon source.
Some growth occurs in mineral media with-
out an added carbon source at the expense
of organic compounds in the air.
Sucrose not attacked.
Asparagine not utilized.
Aerobic.
Temperature range for growth, 20° to
37° C.
Source: Isolated from soil and water.
Commonly found in enrichment cultures
for nitrifying bacteria and in activated
sludge. Babudieri (Rendiconti Istit. Super,
di Sanita, Roma, 13, 1950, 589) has found
this species as a contaminant in Leptospira
canicola cultures.
Habitat: Widely distributed in soil and
in fresh water.
Genus II. Rhodomicrobium Duchow and Douglas, 1949.
(Jour. Bact., 58, 1949, 409.)
Rho.do.mi.cro'bi.um. Gr. noun rhodum the rose; Gr. adj. micrus small; Gr. noun bius
life; M.L. neut.n. Rhodomicrobium red (-producing) microbe.
The daughter cells remain attached to the filaments connecting them to the mother cells.
278
ORDER III. HYPHOMICROBIALES
Non-motile. Gram-negative. Colonies are salmon-pink to orange-red in color. Photohetero-
trophic. Anaerobic. Found in mud and in fresh water.
The type species is Rhodomicrobium vnnnielii Duchow and Douglas.
1. Rhodomicrobium vannielii Duchow
and Douglas, 1949. (Duchow and Douglas,
Jour. Bact., 58, 1949, 409; also see Murray
and Douglas, Jour. Bact., 59, 1950, 157;
and Volk and Pennington, Jour. Bact.,
59, 1950, 169.)
van.niel'i.i. M.L. gen. noun vannielii of
van Niel; named for C. B. van Niel, an
American bacteriologist.
Mature cells are ovoid, measuring 1.2 by
2.8 microns; immature cells are spherical.
Non-motile. The cells are connected by
filaments which are approximately 0.3 mi-
cron in diameter and from one to several
times the length of a mature cell. A mature
cell may produce as many as three daughter
cells: one by formation of a primary fila-
ment from the pole of the cell, and one or
two more by lateral outgrowths of new fila-
ments from the primary filament upon
which the first daughter cell is borne. Be-
cause of the tendency of the cells to remain
attached to the filament, the predominant
growth habit is that of an aggregate con-
taining many cells. Gram-negative.
Agar: In shake tubes, colonies are dark
orange-red, irregular, 2 to 3 mm in diameter
and have a rough, convoluted surface.
Broth: Turbid in young cultures, becom-
ing granular and flocculent; salmon -pink
to deep orange-red, depending on the den-
sity of growth.
Photoheterotrophic. Cells contain bac-
teriochlorophyll and carotenoid pigments.
Growth occurs in the presence of light in a
mineral medium containing an organic hy-
drogen donor and bicarbonate; organic
growth factors are not required. Suitable
hydrogen donors are ethanol, propanol,
butanol, acetate, propionate, butyrate,
valerate, caproate, lactate and malate.
Glucose, mannose, fructose, mannitol,
citrate, tartrate, formate, thiosulfate and
sulfide are not utilized.
Anaerobic.
Optimum temperature, between 25° and
30° C.
Source: Isolated from mud and water
from Washington State.
Habitat: Commonly found in mud, pond,
lake and stream waters.
FAMILY II. PASTEURIACEAE LAURENT, 1890, EMEND.
HENRICI AND JOHNSON, 1935.
(Laurent, Compt. rend. Acad. Sci., Paris, 3, 1890, 754; Henrici and Johnson,
Jour. Bact., 30, 1935, 84.)
Pas.teu.ri.a'ce.ae. M.L. fem.n. Pasteuria type genus of the family; -aceae ending to
denote a family; M.L. fem.pl.n. Pasteuriaceae the Pasteuria family.
Stalked bacteria with spherical or pear-shaped cells; if cells are elongated, the long axis
of the cell coincides with the axis of the stalk. Stalks may be very short or absent, but
when present they are usually very fine and at times arranged in whorls attached to a
common holdfast. Cells multiply by longitudinal fission and/or by budding. Mostly peri-
phytic; one species is parasitic.
The descriptions of the members of this family are largely based upon microscopic exam-
inations of collected materials such as parasitized daphnias (fresh-water Crustacea) or
glass slides submerged at various depths for about two weeks in Lake Alexander, Minne-
sota (Henrici, Jour. Bact., 25, 1932, 277). A few crude cultures were obtained in two liquid
media: one containing a mineral solution with precipitated cellulose and ammonium salts
as a source of nitrogen, the other being a solution of MgS04 and K2HPO4 in tap water to
FAMILY II. PASTEURIACEAE
279
which bits of the exoskeleton of marine crabs were added. No growth took phice on agar
media so that no pure cultures were obtained. Cultures were incubated at room temperature
in the dark. Further information regarding the organisms belonging to the genera of this
family is greatly needed.
Key to the genera of family Pasteuriaceae.
I. Stalks lacking; cells sessile.
Genus I. Pasteuria, p. 279.
II. Stalks long and slender, often in whorls.
Genus II. BlastocauUs, p. 279.
Genus I. Pasteuria Metchnikoff, 1888.
(Ann. Inst. Past., 2, 1888, 166.)
Pas.teu'ri.a. M.L. gen.n. Pasteuria of Pasteur; named for Louis Pasteur, the French
scientist.
Pear-shaped cells attached to each other or to a firm substrate by holdfasts secreted at
the narrow end. Multiplication is by longitudinal fission and by budding of spherical or
ovoid cells at the free end. Non-motile. Non -pigmented. Parasitic on fresh-water Crustacea.
The type species is Pasteuria ramosa Metchnikoff.
1. Pasteuria ramosa Metchnikoff, 1888. found by Henrici and Johnson {ibid., 71
(Ann. Inst. Past., 2, 1888, 166.) and 77) in Lake Alexander, Minnesota;
ra.mo'sa. L. adj. ramosMS much-branched, these appeared frequently on glass slides
Cells 1 to 2 by 4 to 5 microns. Non-motile, submerged in the lake water; they produced
Non-pigmented. Cells grow attached to reproductive bodies apparently by budding
each other in cauliflower-like masses, multi- rather than by an endogenous formation,
plying by longitudinal fission or by intra- Photomicrographs are shown in Henrici
cellular bodies which are extruded as buds, and Johnson {ibid., 93, plate 3, fig. 4) . ZoBell
apparentlj^ reproductive in nature; at times and Allen (Proc. Soc. Exp. Biol, and Med.,
these colonies break up into smaller ones 30, 1933, 1409) and ZoBell and Upham (Bull,
and continue to separate until all of the Scripps Inst. Oceanography, LaJolla, Cal-
individual cells are liberated. Cells and ifornia, 5, 1944, 243) used a submerged-
methods of reproduction resemble those slide technique in sea water and found simi-
of Chamaesiphon, a genus of blue-green lar bacteria.
algae (Henrici and Johnson, Jour. Bact., Source: From the body cavities of Daph-
30, 1935, 71). Gram stain not recorded. nia pulex and D. magna.
Related species: Free-living organisms Habitat: Parasitic on fresh-water crus-
which resembled Pasteuria ramosa were tacea so far as known.
Genus II. Blastocaulis Henrici and Johnson, 1935.
(Jour. Bact., 30, 1935, 84.)
Blas.to.cau'lis. Gr. noun blastus a sprout, shoot, bud; Gr. noun caulis a stalk; M.L.
fem.n. Blastocaulis a bud stalk.
Pear-shaped or globular cells attached to a firm substrate by long, slender stalks with a
holdfast at the base; stalks may occur singly or may arise in clusters from a common hold-
fast. Not cultivable on artificial media. Found on firm substrates in fresh water.
The type species is Blastocaulis sphaerica Henrici and Johnson.
1. Blastocaulis sphaerica Henrici and sphae'ri.ca. Gr. adj. sphaericus spherical.
Johnson, 1935. (Jour. Bact., 30, 1935, 84.) Cells spherical, 1 to 2 microns in diameter.
280
ORDER III. HYPHOMICROBIALES
The cells are attached to long, slender stalks
which radiate from a common center; as
many as 8 stalks may be attached to a com-
mon holdfast; usually thej' are attached
directly to a glass slide, occasionally to
algae or other organisms or to some amor-
phous debris. Multiplication is by budding,
the buds being globular in shape. The
smaller cells stain solidly, but the larger
cells that are budding show a differentiation
of the protoplasm: the free end stains deeply
while that part of the cell which is attached
to the stalk stains more faintly. Young cells
are Gram-positive, but budding individuals
are Gram-negative.
Temperature relations: Found only in
lake water where temperatures do not ex-
ceed 23° C.
Comments: It is believed that the char-
acteristic growth of this organism in whorls
may be best explained by assuming that
when the buds germinate they first undergo
a simple fission, perhaps producing clusters
of cells, and that then, from these clusters,
the individual cells secrete stalks which
thus radiate from a common holdfast.
Related species: Similar stalked bacteria
which reproduce by budding are illustrated
by Henrici and Johnson (ibid., 77 and 91)
but are not named or described in detail.
Source: From glass slides submerged in
Lake Alexander, Minnesota.
Habitat: Presumably widely distributed
in fresh-water ponds. Does not occur closer
to the shore than the 2-meter contour.
Found constantly in the open lake at all
depths up to 13 meters. Occurs more abun-
dantly in the fall months than in the sum-
ORDER IV. EUBACTERIALES BUCHANAN, 1917.
(Jour. Bact., 2, 1917, 102.)
Eu.bac.te.ri.a'les. Gr. pref. eu- well, true; Gr. neut.n. hacterium a small rod; -ales end-
ing to denote an order; M.L. fem.pl.n. Euhacteriales the order of the true bacteria.
Simple, undifferentiated, rigid cells which are either spherical or straight rods. In some
families, for e.xample Corynehacteriaceae, a certain amount of pleomorphism occurs. Only
the simplest forms of branching occur, and these only rarely. There are many non-motile
as well as motile species. The flagella are usually arranged peritrichously, but monotrichous
species do occur in groups where the flagellation is normally peritrichous; such conditions
appear to have been developed from ancestral peritrichous species. Tj'pical endospores
occur in one family (Bacillaceae) . All of the species in certain families are definitely Gram-
negative; in other families and groups, where the majority of species are Gram-positive,
at least in certain stages of growth, species occur which lose their Gram stain so readily
that they are generally classed as Gram-negative. Reproduction is by transverse fission;
occasionally the cells divide in two or three planes perpendicular to each other, thereby
forming tetrads or packets of eight cells. The pigments of chromogenic species are com-
monly non-water-soluble and of a carotenoid nature; other pigments do occur however,
some of which show slight powers of diffusion into agar media. None of these pigments have
the ability to carry out photosynthesis. The order includes saprophytes, parasites and
many pathogenic species; the latter cause diseases of both animals and plants. Found in
salt and fxesh waters, air, soil and in the bodies of animals and plants.
Key to the families of order Euhacteriales.
I. Cells rod-shaped (rarely large, yeast-like cells). Gram-negative.
A. Aerobic or facultatively anaerobic.
1. Large ovoid to rod-shaped cells which may be yeast-like in appearance. Free-
living in soil. Fix free nitrogen.
Family I. Azofobacteraceae, p. 283.
2. Not as above.
a. Heterotrophic rods which may not require organic nitrogen for growth.
Usually motile by means of one to six flagella. Frequently form nodules or
tubercles on roots of plants or show violet chromogenesis. Colonies usually
large and slimy, especially on mannitol agar.
Family II. RMzobiaceae, p. 285.
aa. Not as above.
b. Straight rods which grow readily on ordinary peptone media. May or
may not ferment sugars anaerobically with the production of organic
acids.
c. Glucose usually attacked oxidatively or not at all. Only rarely are
species able to ferment glucose anaerobically. Produce little or no
acid in litmus milk. May or may not reduce nitrates. Many yellow
chromogens. Some species digest agar, others chitin. Primarily
found as saprophytes in foods, in soil and in fresh and salt water.
Family III. Achromobacteraceae, p. 296.
281
282 ORDER IV. EUBACTERIALES
cc. Ferment glucose anaerobically, frequently producing visible gas
(CO2 + H2) from glucose and sometimes lactose. Reduce nitrates
(rare exceptions). Frequently found in the alimentary, respiratory
and urinary tracts of vertebrates, others are free-living, while still
others are plant pathogens.
Famil}^ IV. Enterobacteriaceae, p. 332.
bb. Usuallj^ small, motile or non-motile rods. Obligate animal parasites
which usually require body fluids for growth. Man}' fail to grow on
ordinar}^ media. The majority do not ferment glucose anaerobically.
Family V. Brucellaceae , p. 394.
B. Anaerobic to microaerophilic, rod-shaped organisms which sometimes show branch-
ing.
Family VI. Bacteroidaceae, p. 423.
II. Cells spherical to rod-shaped. Usually Gram-positive, though some cocci and anaerobic
spore-forming rods lose the Gram stain readil3^
A. Cells do not form endospores.
1. Cells spherical, occurring in masses, tetrads or packets of eight cells.
a. Spherical cells. Gram-positive. Aerobic or anaerobic.
Family VII. Micrococcaceae, p. 454.
aa. Cells spherical. Gram-negative. Aerobic or anaerobic. Frequently occur
in pairs.
Family VIII. Neisseriaceae, p. 480.
2. Cells either spherical, occurring in chains, or rod-shaped. Gram-positive, but
cells may lose the Gram stain readily in old cultures.
a. Cells rod-shaped, no pleomorphism or branching of cells. Rarely or never
ferment glucose anaerobically.
Family IX. Brevibacteriaceae, p. 490.
aa. Not as above.
b. Gram-positive cocci and rods which frequentl}' form chains of cells.
Cells ferment sugars anaerobically with the production of lactic, acetic,
propionic, butyric, etc. acids. Microaerophilic to anaerobic,
c. Homo- and hetero-fermentative cocci and rods whose chief product
in fermentation is lactic acid. Do not reduce nitrates.
Family X. LactobaciUaceae, p. 505.
cc. Rod-shaped bacteria whose distinctive product in fermentation is
propionic acid, butyric acid or ethanol. All produce CO2 .
Family XI. Propionibacteriaceae, p. 569.
bb. Cells generally rod-shaped but wedge and club forms are common. The
cells are usually found in angular or picket formations due to snapping
division. Old cells are frequently Gram-negative. Not active in the an-
aerobic fermentation of sugars. May or may not reduce nitrates.
Family XII. Corynebacieriaceae, p. 578.
B. Rod-shaped cells that produce endospores. Aerobic and anaerobic. Some anaerobic
species lose the Gram stain readily.
Family XIII. Bacillaceae, p. 613.
FAMILY I. AZOTOBACTERACEAE
283
FAMILY I. AZOTOBACTERACEAE BERGEY, BREED AND MURRAY, 1938.*
(Azotobacleriaceae (sic) Bergey, Breed and Murray, Preprint, Manual, 5th ed.,
October, 1938, v and 71.)
A.zo.to.bac.te.ra'ce.ae. M.L. mas.n. Azotobacter tj-pe genus of the familj^; -aceae end-
ing to denote a family; M.L. fem.pl.n. Azotobacteraceae the Azotobacter family.
Relatively large rods or even cocci, sometimes almost yeast-like in appearance. Cells
without endospores. The type of flagellation in this genus has been definitely established
as peritrichous. Gram-negative. Obligate aerobes, usually growing in a film on the surface
of the culture medium. Capable of fixing atmospheric nitrogen when provided with carbo-
hydrate or other energy source. Grow best on media deficient in nitrogen. Soil and water
bacteria.
There is a single genus.
Genus I. Azotobacter Beijerinck, 1901.
(Cent. f. Bakt., II Abt., 7, 1901, 567.)
A.zo.to.bac'ter. Gr. adj. azons without life; Fr. noun azote nitrogen; M.L. mas.n, bacter
the masculine equivalent of Gr. neut.n. bactrum a rod or staff; M.L. mas.n. Azotobacter
nitrogen rod.
Description same as for the family.
The type species is Azotobacter chroococcum Beijerinck.
1. Azotobacter chroococcum Beijer-
inck, 1901. (Cent. f. Bakt., II Abt., 7, 1901,
567; also see ibid., 9, 1902, 3.)
chro.o.coc'cum. Gr. noun chroa color,
complexion; Gr. noun coccus a grain; M.L.
neut.n. chroococcum colored coccus.
Rods, 2.0 to 3.0 by 3.0 to 6.0 microns,
occurring in pairs and packets and occasion-
allj^ in chains. The cells show three or four
refractile granules. The organisms are sur-
rounded by a slimy membrane of variable
thickness, usually becoming brownish in
older cultures, due possibly to the con-
version of tyrosine to melanin. The coloring
matter is insoluble in water, alcohol, ether
or chloroform. Motile bj' means of numerous
peritrichous flagella (Hofer, Jour. Bact.,
47, 1944, 415). Gram-negative.
Grows in absence of organic nitrogen.
Gelatin colonies: Very small, circular,
yellow, granular, later becoming yellowish
brown.
Gelatin stab: Onlj' slight growth in the
stab. No liquefaction.
Mannitol agar stab: Gray, may become
brownish.
Nutrient broth: No growth even in the
presence of glucose; peptone utilized with
difficult}'.
Litmus milk: Becoming clearer in 10 to
14 days.
Potato: Glossy, barely visible, slimy to
wrinkled; may become yellowish, brownish
.yellow or chocolate-brown.
The organism fixes atmospheric nitrogen
and gives off CO2, utilizing glucose and
sucrose. Other generally used carbon com-
pounds are fructose, maltose, mannitol,
inulin, dextrin, galactose, arabinose, starch,
glycerol, ethyl alcohol, acetate, butyrate,
citrate, lactate, malate, propionate and
succinate.
Nitrate : Improves growth in amounts
less than 1 gm per liter; greater amounts
are toxic.
Fixes nitrogen moderately actively.
Chemical analysis: Four-day cultures
grown upon mannitol agar, when dried, are
found to contain less than 0.5 per cent of
hemicelluloses, less than 20 per cent of crude
protein, less than 5 per cent of ash, and
more than 30 per cent of lignin-like mate-
rials (Greene, Soil Sci., 39, 1935, 327). The
nitrogen fraction contains less than 1 per
* Revised by Dr. A. W. Hofer, New York State Experiment Station, Cornell University,
Geneva, New York, Jime, 1938; further revision by Dr. A. W. Hofer, May, 1954.
284
ORDER IV. EUBACTERIALES
cent of amide nitrogen, less than 1 per cent
of humin nitrogen and about 1 per cent of
basic nitrogen.
Aerobic.
Optimum temperature, between 25° and
28° C.
Distinctive characters: Inability to grow
in peptone media, even in the presence of
glucose; frequent occurrence of a dark brown
or black pigment.
Source: Isolated from soil.
Habitat : Occurs naturally in the majority
of neutral or alkaline field soils.
2. Azotobacter agilis Beijerinck, 1901.
(Cent. f. Bakt., II Abt., 7, 1901, 577.)
a'gi.lis. L. adj. agilis quick, agile.
Rods, 4 to 6 microns in length, almost
spherical. Actively motile by means of
numerous, peritrichous flagella (Hofer,
Jour. Bact., 47, 1944, 415). Some strains
are reported to be non-motile. Gram-nega-
tive.
Grows in absence of organic nitrogen.
Gelatin: No liquefaction.
Mannitol agar colonies: Circular, gray-
ish white, translucent with whitish center.
Washed agar colonies : Show slight bluish
green fluorescence. The presence of a fluo-
rescent pigment is readily demonstrated
by placing cultures under ultraviolet light,
3600 A. Examination by paper chromotog-
raphy indicates that this pigment is not
fluorescin, the pigment found in fluorescent
pseudomonads (Johnstone, Jour. Bact.,
69, 1955, 481).
Mannitol agar slant: Grayish, translu-
cent, fluorescent.
Plain agar slant : Yellowish white, smooth,
glistening, translucent with opaque center.
Broth: Turbid, with sediment.
Litmus milk: Becoming clear in 10 to 14
days.
Potato: Yellowish white, slimy, becom-
ing yellowish brown.
In the presence of organic acids, a green-
ish or reddish pigment is formed.
The organism fixes atmospheric nitrogen
actively and gives off CO2 .
Aerobic.
Chemical analysis: Four-day cultures
grown upon mannitol agar, when dried, con-
tain more than 4 per cent of hemicelluloses,
more than 45 per cent of crude protein, more
than 7 per cent of ash, and less than 4 per
cent of lignin-Iike materials. The nitrogen
fraction contains more than 1 per cent
amide nitrogen, more than 1 per cent humin
nitrogen, and 2 per cent or more of basic
nitrogen (Greene, Soil Sci., 39, 1935, 327).
Optimum temperature, between 25° and
28° C.
Distinctive characters: Lack of a brown
pigment; occasional fluorescence; growth
in peptone broth containing glucose.
Comment: A non-chromogenic variety
of this species has been recognized by
Kluyver and van den Bout (Arch. f. Mi-
krobiol., 7, 1936,263).
Source: Originally isolated from canal
water at Delft.
Habitat: Occurs in water and soil.
3. Azolobacter indicus Starkey and De,
1939. {Azotobacter iniicum (sic) Starkey and
De, Soil Sci., 47, 1939, 337.)
in'di.cum. L. adj. indicus of India.
Ellipsoidal rods, 0.5 to 1.2 by 1 7 to 2.7
microns when grown on nitrogen-free glu-
cose agar. One of the distinctive character-
istics is the presence of two large, round,
highly refractive bodies in the cells, one
usually at each end. Motile by means of
numerous peritrichous flagella (Hofer, Jour.
Bact., 47, 1944, 415). Gram-negative.
The organism grows slowly but in time
produces large amounts of slime. Has high
acid tolerance, since it grows from pH 3 to 9.
Sucrose or glucose agar plates: Colonies
are colorless, round, very much raised and
uniformly turbid, having much the appear-
ance of heavy starch paste. After two weeks,
a buff to light brown color develops.
Mannitol agar slant: Grows very poorly.
Peptone agar slant with 0.5 per cent glu-
cose: Limited grayish growth.
Nutrient broth: No growth.
Liquid media: Generallj^ turbid with
some sediment.
Fixes atmospheric nitrogen readily with
either glucose or sucrose as source of energy.
Aerobic.
Optimum temperature, 30° C.
FAMILY I. AZOTOBACTERACEAE 285
Distinctive characters: Tolerance of acid- because the organisms in the new genus
ity; wide limits of pH tolerated; abundant differ in morphology and physiology in
slime production; large globules of f;it important respects from the organisms in
within cells. the genus Azotobacier proper. Further
Relationships to other species: Derx comparative studies should be made before
(Kon. Nederl. Akad. v. Wetensch., Amster- this separation is accepted,
dam, Proc. Sect. Sci., 53, 1950, 145; Ann. Source: Isolated from soils of India and
Bogoriensis, 1, 1950, 1) has made this species Java,
the type species of a new genus, Beijerinckia, Habitat : Soils.
FAMILY II. RHIZOBIACEAE CONN, 1938.
(Jour. Bact., 36, 1938, 321.)
Rhi.zo.bi.a'ce.ae. M.L. neut.n. lihizobium type genus of the family; -aceae ending to
denote a family; M.L. fem.pl.n. Rhizohiaceae the Rhizobium family.
Cells without endospores, rod-shaped, sparsely flagellated (one polar or lateral flagellum
or 2 to 4 peritrichous ones); some species are non-motile. Usually Gram-negative. One
genus {Chromobacterium) produces a violet pigment. Grow aerobically on ordinary culture
media containing glucose. Glucose and sometimes other carbohydrates are utilized without
appreciable acid formation. Saprophytes, symbionts and pathogens; the latter are usually
plant pathogens forming abnormal growths on roots and stems.
Key to genera of family Rhizobiaceae.
I. Cells capable of fixing free nitrogen when growing symbiotically on the roots of Legu-
minosae.
Genus I. Rhizobium, p. 285.
II. Either plant pathogens which attack roots or produce hypertrophies on stems or free-
living non-chromogenic soil or water forms. Do not fix nitrogen.
Genus II. Agrobaderiujn, p. 288.
III. Usually free-living soil and water forms which produce a violet chromogenesis.
Genus III. Chromobacterium, p. 292.
Genus I. Rhizobium Frank, 1889.*
(Phytomyxa Schroeter, in Cohn, Kryptogamen-Flora von Schlesien, 3, 1886, 134; Frank,
Ber. d. deutsch. bot. Gesellsch., 7, 1889, 380.)
Rhi.zo'bi.um. Gr. noun rhiza a root; Gr. noun bins life; M.L. neut.n. Rhizobium that
which lives in a root.
Rods which measure 0.5 to 0.9 bj^ 1.2 to 3.0 microns. Motile when young, commonly chang-
ing to bacteroidal forms (a) upon artificial culture media containing alkaloids or glucosides,
or in which aciditj^ is increased, or (b) during symbiosis within the nodule. Gram-negative.
Aerobic. Heterotrophic, growing best with extracts of yeast, malt or other plant materials.
Nitrites may be produced from nitrates. Nitrites are not utilized. Gelatin is not liquefied
or is very slightly liquefied after long incubation. Optimum temperature, 25° C. This group
is capable of producing nodules on the roots of Leguminosae and of fixing free nitrogen during
this symbiosis.
The type species is Rhizobium leguminosarum Frank.
*Original revision by Dr. and Mrs. O. N. Allen, Universit}^ of Wisconsin, Madison, Wis-
consin, January, 1938; further revision by Dr. O. N. Allen, September, 1953.
286 ORDER IV. EUBACTERIALES
Key to the species of genus Rhizobiuni.
I. Litmus milk alkaline.
A. Forms a serum zone in milk. Young cells peritrichous.
1. Causes formation of root nodules on species of the genera Lathyrus, Pisum, Vicia
and Lens. Bacteroids irregular with x-, y-, star- and club-shaped forms.
1. Rhizohium leguminosarum.
2. Causes formation of root nodules on Phaseolus vulgaris, P. muUiflorvs and P.
angustifolius . Bacteroids, vacuolated rods, few branched forms.
2. Rhizohium phaseoli.
3. Causes formation of nodules on species in the genus Tri folium. Bacteroids pear-
shaped, swollen, vacuolated. Pentoses usually not fermented.
3. Rhizohitim trifolii.
B. No serum zone formed in milk. Monotrichous cells usually occur; in 