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: nj ; r^ = ru id a D i r^ a CD s r^ = CD # 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, O...

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BERGEY'S MANUAL OF

DETERMINATIVE BACTERIOLOGY

BERGEY'S MANUAL

DETERMINATIVE

BACTERIOLOGY BY

ROBERT Ixite

S.

BREED

Professor Emerilus, Cnrnell Universily, Geneva,

New

York

MURRAY

E. G. D.

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

THE WILLIAMS

BALTIMORE & WILKINS 1957

COMPANY

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 &

Rhizobium

285

Andrewes, C. H.

Virales

Barker, H. Albert

Methanococcus Butyrihacterium Zymobacterium

985 473 577

Beger, H.

Caulobacteraceae

Allen, 0. N., Mr.

Mrs.

577 212 217

Siderocapsaceae Chlamydobacteriales Bier, Otto

Borman, E. K. Branham, Sara E. Breed, R. S.f

262 418

Calymmatobacterium Paracolobactrum

346

480

Neisseria

Introduction Considerations influencing the classification

74 78 193 200 204 212 217 229

Methanomonadaceae Thiobaderiaceae Pholobacterium Protaminobacter

Mycoplana Caulobacteraceae

Siderocapsaceae Vibrio

250 250 252 253 262 292 297 300 309 322 332 335

Methanobacterium Cellvibrio

Cellfalcicula

Spirillum Chlamydobacteriales

Chromobacterium Alcaligenes

Achromobacter Flavobacterium Agarbacterium Enter obacteriaceae Escherichia Aerobacter

341

Klebsiella

344

Serratia

359

Pasteurella

Bacteroidaceae

395 423

Sphaerophorus

441

Micrococcaceae

454

Micrococcus Sarcina

455 467 490

Brevibacteriaceae

578 830 985

Cory neb acteriaceae Caryophanales Virales

Numerous contributions Broom, t

J. C.

1

4

to various other taxa

907

Leptospira

Deceased.

V

LIST OF CONTRIBUTORS

VI

How

Buchanan, R. E.

Bacteria are

Named and

Paraspirilluni

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

t

Deceased.

Identified

15

257

Beggiatoales

837

Beggiatoaceae

837

Leucotrichaceae

850

Pseudomonas Xanthomonas

152

89

Agrobacterium

288

Erwinia

349

Corynebacterium Beneckea Cellulomonas

579 328 601

Pasteurella

Eubacterium

395 552

Catenabacterium

560

Ramibacterium

563

Cillobacterium

566 288

Agrobacterium Alcaligenes Actinomycetales

297

Actinoplanaceae Spirochaetales

825 892

694

Propionibacteriaceae

569

Microbacterium Hyphomicrobiales Peptococcus

600 276

Shigella

384 207

Halubacterium Chromobaclerium

474

Staphylococcus

292 464

Streptobacillus

451

Mycoplasmatales

914

Bacillus

613

Mycobacterium Mycobacterium

695

Erysipelothrix Actinobacillus

Pseudomonas Dialister

695 599 414

89 440

Azotomonas

198

Azotobacteraceae

283 436 441

Fusobacterium Sphaerophorus Virales

Parasites of protozoa

Micrococcus Gaffkya Leuconostoc Brucella Thiobacteriaceae Thiobacterium

985 927 455 466 531

404 78 79

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.

Macromonas

80

Thiovulum

81

Thiospira

82

Salmonella

368 421 424 927

Noguchia Bacteroides Parasites of protozoa

Zymomonas

199

Erysipelothrix

599 227

Ferrohacillus

Photobacterium Selenomonas Myconostoc Pasteurella

Euhacterium Catenabacterium Ramibacterium Vilreoscillaceae

Lochhead, A. G.

Arthrobacter

McClung, L.

Clostridium

S.

McCoy, Elizabeth

Clostridium

Merchant, I. A. Morse, E. V.

Pasteurella

Murray, E. G. D.

Vibrio

Corynebacterium Enter obacteriaceae Moraxella Bacteroidaceae Neisseria Lactobacillaceae

Diplococcus Corynebacterium Listeria

Neitz,

W, O.

Anaplasma

Nellis, Lois

Mycococcus

Niven, C. F.

Lactobacillaceae

Streptococcus

0rskov, J. Parker, Pederson, Carl S.

CD.

Microcyclus Thiobacillus Lactobacillaceae

Pediococcus Leuconostoc Lactobacillus

Pelczar,

M.

J.

M. A.

Philip, Cornelius B.

Pittman, Margaret

t

Deceased.

552

560 563 844

605 634 634 395 229 579

332 419 423 480 505 507 579 597 981 707

505 508 253 83

505 529 531 542

Caryophanales Microtatobiotes

931 and 933

Rickettsiales

934

Anaplasmataceae

980 402 406 844

Bordetella

Haemophilus Pringsheim, E. G.

260 395

480 485 830

Neisseria Veillonella

Peshkoff,

193 258

Vilreoscillaceae

LIST OF CONTRIBUTORS

Vlll

Rake, Geoffrey W. Reed, Guilford B.f Robinson, Elliott S. Robinson, George H.f

Diplococcus Spirochaetales

Schatz, Albert

Hydrogenomonas

Seeley, H.

W.

Chlamydiaceae Mycobacterium

957

695 507 892 facilis

Streptococcus

H. Sherman, James W.f Seeliger,

Corynebacterium Lactobacillaceae

Streptococcus

Skerman, V. B. D. Smit, Jan Smith, Louis DeSpain

Artificial

Keys

Sarcina Lactobacillaceae

Peptostreptococcus

Smith, N. R.

76 508 579

505 508 987 467 505

Bacteroidaceae

533 423

Bacillus

613

Snieszko, S. F.

Aeromonas

189

Spray, R. S. Stanier, R. Y. Starkey, R. L. Steinhaus, E. A. Stuart, C. A.

Clostridium

634

Myxobacterales

854

Nitrobacteraceae Virales

68 985

Proteus

364

Thiobacillus

83 202

Temple, Kenneth L. Thj0tta, Th.t Tobie,

van

W.

Alginornonas Alginobacter

Chromobacterium

C.

Niel, C. B.

348 292

Rhodobacteriineae

35

Propionibacterium Achromatiaceae

569 851

Vaughn, Reese Verona, Onorato

Acetobacter

183

Cellvibrio

250

Waksman,

Actinomycetales

694

Mycococcus Actinomycetaceae

707

Streptomycetaceae

744

Zoogloea

206

Spirochaetales

892

Weeks, Owen B.

Flavobacterium

309

Weinman, David Wolff, J. W. Yale, N. W.

Bartonellaceae

Leptospira

968 907

Escherichia

335

Aerobacter

341

Desulfovibrio

248

Wattie,

S.

A.

Elsie

Lackey)

ZoBell, Claude E. t

Deceased.

713

(Mrs.

PREFACE TO SEVENTH EDITION Manual of Determinative Bacteriology differs but Uttle from that of the sixth edition. However, 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 bacteria and the descriptions of the taxa from Class to Species and Subspecies, together 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

general format of the seventh edition of Bergey's

Manual itself contains descriptions of many more more adequate descriptions than have former editions; the transfer much material to the Index Bergeyana has meant a reduction in the number pages and a book of more convenient size and greater usefulness. The Index

The net

result is that the

species with of of

Bergeyana should prove containing, as

it will,

to be

an invaluable tool

for the research microbiologist,

references to the whole field of systematic bacteriology

an index to the names

of described species,

and

both valid and invalid.

Much material of historical value and interest in the sixth and earlier editions Manual has been excluded, not because it is lacking in real value to the

of 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

and the names

United States are

To

Manual all of

is

nomenof taxa

new combinations are to be ascribed to these authors, and of the Manual. Contributors from countries other than the more numerous than in the earlier editions. In other words,

of all

not to the editors the

all

new names

are to be regarded in

clatural matters as strictly the authors of their sections. All

rapidly assuming the character of an international publication.

these contributors the sincere thanks of the Board of Editors,

ticularly of the Editor-in-Chief, are due.

patience, care

and

scientific

acumen

The seventh

edition

is

and par-

a tribute to the

of these individuals. Special note should

taken of the assistance rendered in the

office of

be

the Editor-in-Chief by Mr.

Erwin Lessel, Miss Maude Hogan, Mrs. Eleanore Heist CUse and Mrs. Margaret Edson Breed.

PREFACE TO SEVENTH EDITION

X

The keys

to the several categories of taxa (orders, families, tribes, genera and 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 species)

for use in

making appropriate revisions in the 6th Edition of the Manual. The Code of Nomenclature of the Bacteria and Viruses is about

revised International

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 Manual, was regarded by some eminent virologists as perhaps inadvisable because it was premature. They felt strongly that the problems of morphology, of the

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

agreement

will

The

made with

be reached to

the full expectation that sufficient international

make

possible adequate treatment in the eighth

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

edition.

Editorial

of the first edition of the

"The

assistance of

all

Manual:

bacteriologists

is

earnestly solicited in the correction of possible

errors in the text."

Among of

the tasks of the several editions of the

an increasingly satisfactory

the nomenclature of the past.

Manual

has been the codification

classification of the bacteria

The

and the correction

present volume undoubtedly has

that were not caught notwithstanding a most earnest

effort.

many

of

errors

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

Bergey's

of Trustees of

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 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

and devoted

devotion, energy and in

Dr. Breed over a period of

skill of

many

years, as

shown The

the organization of better bacterial nomenclature and classification.

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

have been corrected before publication. Breed leaves the Editorial Committee without a chairman and editor-in-chief. The Board of Trustees is unanimous in the belief that a successor to Dr. Breed should be found outside its present membership. Dr. Breed was als...