BERGEY’S MANUAL� OF

SystematicBacteriology

Second Edition

Volume TwoThe Proteobacteria

Part CThe Alpha-, Beta-, Delta-, and

Epsilonproteobacteria

BERGEY’S MANUAL� OF

SystematicBacteriology

Second Edition

Volume TwoThe Proteobacteria

Part CThe Alpha-, Beta-, Delta-, and

Epsilonproteobacteria

Don J. BrennerNoel R. Krieg

James T. StaleyEDITORS, VOLUME TWO

George M. GarrityEDITOR-IN-CHIEF

EDITORIAL BOARDJames T. Staley, Chairman, David R. Boone, Vice Chairman,

Don J. Brenner, Paul De Vos, George M. Garrity, Michael Goodfellow,Noel R. Krieg, Fred A. Rainey, Karl-Heinz Schleifer

WITH CONTRIBUTIONS FROM 222 COLLEAGUES

George M. Garrity, Sc.D.Bergey’s Manual TrustDepartment of Microbiology and Molecular GeneticsMichigan State UniversityEast Lansing, MI 48824-4320USA

Library of Congress Control Number: 2005926296 © 2005, 1984–1989 Bergey’s Manual Trust This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright Clearance Center. Violations are liable to prosecution under the respective Copyright Law. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. While the advice and information in this book are believed to be true and accurate at the date of publication, neither the authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that may be made. The publisher makes no warranty, express or implied, with respect to the material contained herein. Printed on acid-free paper Printed in the United States of America. (APEX/MVY) Springer is part of Springer Science+Business Media (www.springer.com)

ISBN ISBN-13: 978-0-387-24145-6

-10: 0-387-24145-0

This volume is dedicated to our colleagues,David R. Boone, Don J. Brenner,

Richard W. Castenholz, and Noel R. Krieg, whoretired from the Board of Trustees of Bergey’s ManualTrust as this edition was in preparation. We deeplyappreciate their efforts as editors and authors; theyhave devoted their time and many years in helping

the Trust meet its objectives.

EDITORIAL BOARD AND TRUSTEESOF BERGEY’S MANUAL TRUST

James T. Staley, ChairmanDavid R. Boone, Vice ChairmanGeorge M. GarrityPaul De VosMichael GoodfellowFred A. RaineyKarl-Heinz SchleiferDon J. Brenner, EmeritusRichard W. Castenholz, EmeritusJohn G. Holt, EmeritusNoel R. Krieg, EmeritusJohn Liston, EmeritusJames W. Moulder, EmeritusR.G.E. Murray, EmeritusCharles F. Niven, Jr., EmeritusNorbert Pfennig, EmeritusPeter H.A. Sneath, EmeritusJoseph G. Tully, EmeritusStanley T. Williams, Emeritus

ix

Preface to Volume Two of the Second Edition ofBergey’s Manual� of Systematic Bacteriology

There is a long-standing tradition for the Editors of Bergey’s Man-ual to open their respective editions with the observation thatthe new edition is a departure from the earlier ones. As thisvolume goes to press, however, we recognize a need to deviatefrom this practice, by offering a separate preface to each volumewithin this edition. In part, this departure is necessary becausethe size and complexity of this edition far exceeded our expec-tations, as has the amount of time that has elapsed betweenpublication of the first volume of this edition and this volume.

Earlier, we noted that systematic procaryotic biology is a dy-namic field, driven by constant theoretical and methodologicaladvances that will ultimately lead to a more perfect and usefulclassification scheme. Clearly, the pace has been accelerating asevidenced in the super-linear rate at which new taxa are beingdescribed. Much of the increase can be attributed to rapid ad-vances in sequencing technology, which has brought about amajor shift in how we view the relationships among Bacteria andArchaea. While the possibility of a universally applicable naturalclassification was evident as the First Edition was in preparation,it is only recently that the sequence databases became largeenough, and the taxonomic coverage broad enough to makesuch an arrangement feasible. We have relied heavily upon thesedata in organizing the contents of this edition of Bergey’s Manualof Systematic Bacteriology, which will follow a phylogenetic frame-work based on analysis of the nucleotide sequence of the smallribosomal subunit RNA, rather than a phenotypic structure. Thisdeparts from the First Edition, as well as the Eighth and NinthEditions of the Determinative Manual. While the rationale for pre-senting the content of this edition in such a manner should beevident to most readers, they should bear in mind that this edi-tion, as in all preceding ones represents a progress report, ratherthan a final classification of procaryotes.

The Editors remind the readers that the Systematics Manual isa peer-reviewed collection of chapters, contributed by authorswho were invited by the Trust to share their knowledge andexpertise of specific taxa. Citation should refer to the author, thechapter title, and inclusive pages rather than to the Editors. TheTrust is indebted to all of the contributors and reviewers, withoutwhom this work would not be possible. The Editors are gratefulfor the time and effort that each expended on behalf of theentire scientific community. We also thank the authors for theirgood grace in accepting comments, criticisms, and editing of

their manuscripts. We would also like to thank Drs. Hans Truper,Brian Tindall, and Jean Euzeby for their assistance on mattersof nomenclature and etymology.

We would like to express our thanks to the Department ofMicrobiology and Molecular Genetics at Michigan State Univer-sity for housing our headquarters and editorial office and forproviding a congenial and supportive environment for microbialsystematics. We would also like to thank Connie Williams notonly for her expert secretarial assistance, but also for unflaggingdedication to the mission of Bergey’ s Manual Trust and Drs.Julia Bell and Denise Searles for their expert editorial assistanceand diligence in verifying countless pieces of critical informationand to Dr. Timothy G. Lilburn for constructing many of thephylogenetic trees used in this volume. We also extend our thanksto Alissa Wesche, Matt Chval and Kristen Johnson for their as-sistance in compilation of the bibliography.

A project such as the Systematics Manual also requires thestrong and continued support of a dedicated publisher, and wehave been most fortunate in this regard. We would also like toexpress our gratitude to Springer-Verlag for supporting our ef-forts and for the development of the Bergey’ s Document TypeDefinition (DTD). We would especially like to thank our Exec-utive Editor, Dr. William Curtis for his courage, patience, un-derstanding, and support; Catherine Lyons for her expertise indesigning and developing our DTD, and Jeri Lambert and LeslieGrossberg of Impressions Book and Journal Services for theirefforts during the pre-production and production phases. Wewould also like to acknowledge the support of ArborText, Inc.,for providing us with state-of-the-art SGML development and ed-iting tools at reduced cost. Lastly, I would like to express mypersonal thanks to my fellow trustees for providing me with theopportunity to participate in this effort, to Drs. Don Brenner,Noel Krieg, and James Staley for their enormous efforts as volumeeditors and to my wife, Nancy, and daughter, Jane, for their con-tinued patience, tolerance and support.

Comments on this edition are welcomed and should be di-rected to Bergey’s Manual Trust, Department of Microbiologyand Molecular Genetics, 6162 Biomedical and Physical SciencesBuilding, Michigan State University, East Lansing, MI, USA48824-4320. Email: garrity@msu.edu

George M. Garrity

xi

Preface to the First Edition of Bergey’s Manual� ofSystematic Bacteriology

Many microbiologists advised the Trust that a new edition of theManual was urgently needed. Of great concern to us was thesteadily increasing time interval between editions; this intervalreached a maximum of 17 years between the seventh and eightheditions. To be useful the Manual must reflect relatively recentinformation; a new edition is soon dated or obsolete in partsbecause of the nearly exponential rate at which new informationaccumulates. A new approach to publication was needed, andfrom this conviction came our plan to publish the Manual as asequence of four subvolumes concerned with systematic bacte-riology as it applies to taxonomy. The four subvolumes are di-vided roughly as follows: (a) the Gram-negatives of general, med-ical or industrial importance; (b) the Gram-positives other thanactinomycetes; (c) the archaeobacteria, cyanobacteria and re-maining Gram-negatives; and (d) the actinomycetes. The Trustbelieved that more attention and care could be given to prep-aration of the various descriptions within each subvolume, andalso that each subvolume could be prepared, published, andrevised as the area demanded, more rapidly than could be thecase if the Manual were to remain as a single, comprehensivevolume as in the past. Moreover, microbiologists would have theoption of purchasing only that particular subvolume containingthe organisms in which they were interested.

The Trust also believed that the scope of the Manual neededto be expanded to include more information of importance forsystematic bacteriology and bring together information dealingwith ecology, enrichment and isolation, descriptions of speciesand their determinative characters, maintenance and preserva-tion, all focused on the illumination of bacterial taxonomy. Toreflect this change in scope, the title of the Manual was changedand the primary publication becomes Bergey’ s Manual of SystematicBacteriology. This contains not only determinative material suchas diagnostic keys and tables useful for identification, but alsoall of the detailed descriptive information and taxonomic com-ments. Upon completion of each subvolume, the purely deter-minative information will be assembled for eventual incorpora-tion into a much smaller publication which will continue theoriginal name of the Manual, Bergey’ s Manual of DeterminativeBacteriology, which will be a similar but improved version of thepresent Shorter Bergey’ s Manual. So, in the end there will be twopublications, one systematic and one determinative in character.

An important task of the Trust was to decide which generashould be covered in the first and subsequent subvolumes. Wewere assisted in this decision by the recommendations of our

Advisory Committees, composed of prominent taxonomic au-thorities to whom we are most grateful. Authors were chosen onthe basis of constant surveillance of the literature of bacterialsystematics and by recommendations from our Advisory Com-mittees.

The activation of the 1976 Code had introduced some novelproblems. We decided to include not only those genera that hadbeen published in the Approved Lists of Bacterial Names in Jan-uary 1980 or that had been subsequently validly published, butalso certain genera whose names had no current standing innomenclature. We also decided to include descriptions of certainorganisms which had no formal taxonomic nomenclature, suchas the endosymbionts of insects. Our goal was to omit no im-portant group of cultivated bacteria and also to stimulate taxo-nomic research on “neglected” groups and on some groups ofundoubted bacteria that have not yet been cultivated and sub-jected to conventional studies.

The invited authors were provided with instructions and ex-emplary chapters in June 1980 and, although the intended dead-line for receipt of manuscripts was March 1981, all contributionswere assembled in January 1982 for the final preparations. TheManual was forwarded to the publisher in June 1982.

Some readers will note the consistent use of the stem -varinstead of -type in words such as biovar, serovar and pathovar.This is in keeping with the recommendations of the Bacterio-logical Code and was done against the wishes of some of theauthors.

We have deleted much of the synonymy of scientific nameswhich was contained in past editions. The adoption of the newstarting date of January 1, 1980 and publication of the ApprovedLists of Bacterial Names has made mention of past synonymyobsolete. We have included synonyms of a name only if theyhave been published since the new starting date, or if they werealso on the Approved Lists and, in rare cases with certain path-ogens, if the mention of an old name would help readers asso-ciate the organism with a clinical problem. If the reader is in-terested in tracing the history of a name we suggest he or sheconsult past editions of the Manual or the Index Bergeyana andits Supplement. In citations of names we have used the abbrevia-tion AL to denote the inclusion of the name on the ApprovedLists of Bacterial Names and VP to show the name has beenvalidly published.

In the matter of citation of the Manual in the scientific lit-erature we again stress the fact that the Manual is a collection

PREFACE TO THE FIRST EDITIONxii

of authored chapters and the citation should refer to the author,the chapter title and its inclusive pages, not the Editor.

To all contributors, the sincere thanks of the Trust is due; theEditor is especially grateful for the good grace with which theauthors accepted comments, criticisms and editing of their man-uscripts. It is only because of the voluntary and dedicated effortsof these authors that the Manual can continue to serve the sci-ence of bacteriology on an international basis.

A number of institutions and individuals deserve special ac-knowledgment from the Trust for their help in bringing aboutthe publication of this volume. We are grateful to the Departmentof Biology of the Virginia Polytechnic Institute and State Uni-versity for providing space, facilities and, above all, tolerance forthe diverted time taken by the Editor during the preparation ofthe book. The Department of Microbiology at Iowa State Uni-versity of Science and Technology continues to provide a wel-come home for the main editorial offices and archives of theTrust and we acknowledge their continued support. A grant

(LM-03707) from the National Library of Medicine, NationalInstitutes of Health to assist in the preparation of this and thenext volume of the Manual is gratefully acknowledged.

A number of individuals deserve special mention and thanksfor their help. Professor Thomas O. McAdoo of the Departmentof Foreign Languages and Literatures at the Virginia PolytechnicInstitute and State University has given invaluable advice on theetymology and correctness of scientific names. Those assistingthe Editor in the Blacksburg office were R. Martin Roop II, DonD. Lee, Eileen C. Falk and Michael W. Friedman and their helpis sincerely appreciated. In the Ames office we were ably assistedby Gretchen Colletti and Diane Triggs during the early periodof preparation and by Cynthia Pease during the major portionof the editing process. Mrs. Pease has been responsible for theconstruction of the List of References and her willingness tohandle the cumbersome details of text editing on a big computeris gratefully acknowledged.

John G. Holt

xiii

Preface to the First Edition of Bergey’s Manual� ofDeterminative Bacteriology

The elaborate system of classification of the bacteria into families,tribes and genera by a Committee on Characterization and Clas-sification of the Society of American Bacteriologists (1911, 1920)has made it very desirable to be able to place in the hands ofstudents a more detailed key for the identification of species thanany that is available at present. The valuable book on “Deter-minative Bacteriology” by Professor F. D. Chester, published in1901, is now of very little assistance to the student, and all pre-vious classifications are of still less value, especially as earliersystems of classification were based entirely on morphologic char-acters.

It is hoped that this manual will serve to stimulate efforts toperfect the classification of bacteria, especially by emphasizingthe valuable features as well as the weaker points in the newsystem which the Committee of the Society of American Bacte-riologists has promulgated. The Committee does not regard theclassification of species offered here as in any sense final, butmerely a progress report leading to more satisfactory classifica-tion in the future.

The Committee desires to express its appreciation and thanksto those members of the society who gave valuable aid in thecompilation of material and the classification of certainspecies. . . .

The assistance of all bacteriologists is earnestly solicited in thecorrection of possible errors in the text; in the collection ofdescriptions of all bacteria that may have been omitted from thetext; in supplying more detailed descriptions of such organismsas are described incompletely; and in furnishing complete de-scriptions of new organisms that may be discovered, or in di-recting the attention of the Committee to publications of suchnewly described bacteria.

David H. Bergey, ChairmanFrancis C. HarrisonRobert S. BreedBernard W. HammerFrank M. HuntoonCommittee on Manual.August, 1923.

xv

Contents

Preface to Volume Two of the Second Edition of Bergey’s Manual� ofSystematic Bacteriology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

Preface to the First Edition of Bergey’s Manual� of SystematicBacteriology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi

Preface to the First Edition of Bergey’s Manual� of DeterminativeBacteriology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii

Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxiiiClass I. Alphaproteobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Order I. Rhodospirillales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Family I. Rhodospirillaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Genus I. Rhodospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Genus II. Azospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Genus III. Levispirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Genus IV. Magnetospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28Genus V. Phaeospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Genus VI. Rhodocista . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33Genus VII. Rhodospira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35Genus VIII. Rhodovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36Genus IX. Roseospira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Genus X. Roseospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39Genus XI. Skermanella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39Genus Incertae Sedis XII. ‘‘Sporospirillum’’ . . . . . . . . . . . . . . . . . . . . 40

Family II. Acetobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Genus I. Acetobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51Genus II. Acidiphilium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54Genus III. Acidisphaera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62Genus IV. Acidocella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65Genus V. Acidomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68Genus VI. Asaia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69Genus VII. Craurococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Genus VIII. Gluconacetobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72Genus IX. Gluconobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77Genus X. Paracraurococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81Genus XI. Rhodopila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83Genus XII. Roseococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85Genus XIII. Roseomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88Genus XIV. Stella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Genus XV. Zavarzinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

Order II. Rickettsiales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96Family I. Rickettsiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

Genus I. Rickettsia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96Genus II. Orientia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114

Family II. Anaplasmataceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Genus I. Anaplasma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Genus II. Ehrlichia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

CONTENTSxvi

Genus III. Neorickettsia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132Genus IV. Wolbachia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138Genus Incertae Sedis V. Aegyptianella . . . . . . . . . . . . . . . . . . . . . . . 143

Family III. Holosporaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146Genus I. Holospora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Genus Incertae Sedis II. Caedibacter . . . . . . . . . . . . . . . . . . . . . . . . 152Genus Incertae Sedis III. Lyticum . . . . . . . . . . . . . . . . . . . . . . . . . . . 156Genus Incertae Sedis IV. Candidatus Odyssella . . . . . . . . . . . . . . . . 157Genus Incertae Sedis V. Candidatus Paracaedibacter . . . . . . . . . . . . 157Genus Incertae Sedis VI. Pseudocaedibacter . . . . . . . . . . . . . . . . . . 158Genus Incertae Sedis VII. ‘‘Pseudolyticum’’ . . . . . . . . . . . . . . . . . . . . 160Genus Incertae Sedis VIII. Tectibacter . . . . . . . . . . . . . . . . . . . . . . . 160

Order III. Rhodobacterales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161Family I. Rhodobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Genus I. Rhodobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161Genus II. Ahrensia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167Genus III. Amaricoccus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168Genus IV. Antarctobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172Genus V. Gemmobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174Genus VI. Hirschia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176Genus VII. Hyphomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179Genus VIII. Ketogulonicigenium . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187Genus IX. Maricaulis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188Genus X. Methylarcula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192Genus XI. Octadecabacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194Genus XII. Paracoccus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197Genus XIII. Rhodobaca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204Genus XIV. Rhodovulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205Genus XV. Roseibium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209Genus XVI. Roseinatronobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209Genus XVII. Roseivivax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210Genus XVIII. Roseobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212Genus XIX. Roseovarius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215Genus XX. Rubrimonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217Genus XXI. Ruegeria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218Genus XXII. Sagittula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219Genus XXIII. Staleya . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221Genus XXIV. Stappia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223Genus XXV. Sulfitobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224Genus Incertae Sedis XXVI. Rhodothalassium . . . . . . . . . . . . . . . . . 228

Order IV. Sphingomonadales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230Family I. Sphingomonadaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233

Genus I. Sphingomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234Genus II. Blastomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258Genus III. ‘‘Citromicrobium’’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263Genus IV. Erythrobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267Genus V. Erythromicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268Genus VI. Erythromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274Genus VII. Porphyrobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275Genus VIII. Sandaracinobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279Genus IX. Zymomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282

Order V. Caulobacterales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Family I. Caulobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287

Genus I. Caulobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Genus II. Asticcacaulis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303Genus III. Brevundimonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308Genus IV. Phenylobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316

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Order VI. Rhizobiales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324Family I. Rhizobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324

Genus I. Rhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325Genus II. Agrobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 340Genus III. Allorhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345Genus IV. Carbophilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 346Genus V. Chelatobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347Genus VI. Ensifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354Genus VII. Sinorhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 358

Family II. Bartonellaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362Genus I. Bartonella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362

Family III. Brucellaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370Genus I. Brucella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370Genus II. Mycoplana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386Genus III. Ochrobactrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389

Family IV. Phyllobacteriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393Genus I. Phyllobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394Genus II. Aminobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397Genus III. Aquamicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399Genus IV. Defluvibacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400Genus V. Candidatus Liberibacter . . . . . . . . . . . . . . . . . . . . . . . . . . . 400Genus VI. Mesorhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403Genus VII. Pseudaminobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409

Family V. Methylocystaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411Genus I. Methylocystis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413Genus II. Albibacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416Genus III. Methylosinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417Genus Incertae Sedis IV. Methylopila . . . . . . . . . . . . . . . . . . . . . . . . 420

Family VI. Beijerinckiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 422Genus I. Beijerinckia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423Genus II. Chelatococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433Genus III. Methylocella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437

Family VII. Bradyrhizobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438Genus I. Bradyrhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438Genus II. Afipia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 443Genus III. Agromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 448Genus IV. Blastobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452Genus V. Bosea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 459Genus VI. Nitrobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 461Genus VII. Oligotropha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 468Genus VIII. Rhodoblastus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471Genus IX. Rhodopseudomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473

Family VIII. Hyphomicrobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476Genus I. Hyphomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476Genus II. Ancalomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494Genus III. Ancylobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497Genus IV. Angulomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501Genus V. Aquabacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 504Genus VI. Azorhizobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505Genus VII. Blastochloris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 506Genus VIII. Devosia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 509Genus IX. Dichotomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513Genus X. Filomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 518Genus XI. Gemmiger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520Genus XII. Labrys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 523Genus XIII. Methylorhabdus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525Genus XIV. Pedomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 527

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Genus XV. Prosthecomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . 538Genus XVI. Rhodomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 543Genus XVII. Rhodoplanes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545Genus XVIII. Seliberia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 549Genus XIX. Starkeya . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 554Genus XX. Xanthobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 555

Family IX. Methylobacteriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 567Genus I. Methylobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 567

Family X. Rhodobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571Genus I. Rhodobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571

Class II. Betaproteobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575Order I. Burkholderiales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575

Family I. Burkholderiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575Genus I. Burkholderia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575Genus II. Cupriavidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 600Genus III. Lautropia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 604Genus IV. Pandoraea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 605Genus V. Paucimonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607Genus VI. Polynucleobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607Genus VII. Ralstonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 609Genus VIII. Thermothrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 620

Family II. Oxalobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623Genus I. Oxalobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 624Genus II. Duganella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 628Genus III. Herbaspirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629Genus IV. Janthinobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 636Genus V. Massilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643Genus VI. Telluria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643

Family III. Alcaligenaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647Genus I. Alcaligenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653Genus II. Achromobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 658Genus III. Bordetella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 662Genus IV. Derxia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 671Genus V. Oligella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 674Genus VI. Pelistega . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 678Genus VII. Pigmentiphaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 681Genus VIII. Sutterella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 682Genus IX. Taylorella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 684

Family IV. Comamonadaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 686Genus I. Comamonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 689Genus II. Acidovorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696Genus III. Brachymonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 704Genus IV. Delftia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 706Genus V. Hydrogenophaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 710Genus VI. Lampropedia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 716Genus VII. Macromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 721Genus VIII. Polaromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 724Genus IX. Rhodoferax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 727Genus X. Variovorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 732Genus Incertae Sedis XI. Aquabacterium . . . . . . . . . . . . . . . . . . . . . 735Genus Incertae Sedis XII. Ideonella . . . . . . . . . . . . . . . . . . . . . . . . . 738Genus Incertae Sedis XIII. Leptothrix . . . . . . . . . . . . . . . . . . . . . . . . 740Genus Incertae Sedis XIV. Roseateles . . . . . . . . . . . . . . . . . . . . . . . 746Genus Incertae Sedis XV. Rubrivivax . . . . . . . . . . . . . . . . . . . . . . . . 749Genus Incertae Sedis XVI. Sphaerotilus . . . . . . . . . . . . . . . . . . . . . . 750Genus Incertae Sedis XVII. Tepidimonas . . . . . . . . . . . . . . . . . . . . . 755Genus Incertae Sedis XVIII. Thiomonas . . . . . . . . . . . . . . . . . . . . . . 757

CONTENTS xix

Genus Incertae Sedis XIX. Xylophilus . . . . . . . . . . . . . . . . . . . . . . . . 759Order II. Hydrogenophilales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763

Family I. Hydrogenophilaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763Genus I. Hydrogenophilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763Genus II. Thiobacillus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 764

Order III. Methylophilales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 770Family I. Methylophilaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 770

Genus I. Methylophilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 770Genus II. Methylobacillus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 771Genus III. Methylovorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 773

Order IV. Neisseriales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 774Family I. Neisseriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 775

Genus I. Neisseria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777Genus II. Alysiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 798Genus III. Aquaspirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 801Genus IV. Chromobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824Genus V. Eikenella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 828Genus VI. Formivibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 832Genus VII. Iodobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 833Genus VIII. Kingella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 836Genus IX. Microvirgula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 840Genus X. Prolinoborus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 841Genus XI. Simonsiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 844Genus XII. Vitreoscilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 851Genus XIII. Vogesella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 858Genus Incertae Sedis XIV. Catenococcus . . . . . . . . . . . . . . . . . . . . . 859Genus Incertae Sedis XV. Morococcus . . . . . . . . . . . . . . . . . . . . . . . 861

Order V. Nitrosomonadales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 863Family I. Nitrosomonadaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 864

Genus I. Nitrosomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 864Genus II. Nitrosolobus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 868Genus III. Nitrosospira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 868Genus IV. ‘‘Nitrosovibrio’’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 869

Family II. Spirillaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 870Genus I. Spirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 870

Family III. Gallionellaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 880Genus I. Gallionella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 880

Order VI. Rhodocyclales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 887Family I. Rhodocyclaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 887

Genus I. Rhodocyclus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 887Genus II. Azoarcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 890Genus III. Azonexus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 901Genus IV. Azospira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 902Genus V. Azovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 902Genus VI. Dechloromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903Genus VII. Dechlorosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 904Genus VIII. Ferribacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 904Genus IX. Propionibacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 905Genus X. Propionivibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 906Genus XI. Thauera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 907Genus XII. Zoogloea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 913

Class IV. Deltaproteobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 922Order I. Desulfurellales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 922

Family I. Desulfurellaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 923Genus I. Desulfurella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 923Genus II. Hippea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 925

Order II. Desulfovibrionales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 925

CONTENTSxx

Family I. Desulfovibrionaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 926Genus I. Desulfovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 926Genus II. Bilophila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 938Genus III. Lawsonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 940

Family II. Desulfomicrobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 944Genus I. Desulfomicrobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 944

Family III. Desulfohalobiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 948Genus I. Desulfohalobium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 949Genus II. Desulfonatronovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 953Genus III. Desulfothermus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 955

Family IV. Desulfonatronumaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 956Genus I. Desulfonatronum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 956

Order III. Desulfobacterales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 959Family I. Desulfobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 959

Genus I. Desulfobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 961Genus II. Desulfobacterium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 965Genus III. Desulfobacula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 968Genus IV. Desulfobotulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 970Genus V. Desulfocella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 971Genus VI. Desulfococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 972Genus VII. Desulfofaba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 975Genus VIII. Desulfofrigus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 976Genus IX. Desulfonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 977Genus X. Desulfosarcina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 981Genus XI. Desulfospira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 984Genus XII. Desulfotignum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 987

Family II. Desulfobulbaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 988Genus I. Desulfobulbus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 988Genus II. Desulfocapsa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 992Genus III. Desulfofustis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 994Genus IV. Desulforhopalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 995Genus V. Desulfotalea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 997

Family III. Nitrospinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 999Genus I. Nitrospina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 999

Order IV. Desulfarcales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1003Family I. Desulfarculaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1003

Genus I. Desulfarculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1004Order V. Desulfuromonales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1005

Family I. Desulfuromonaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1006Genus I. Desulfuromonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1007Genus II. Desulfuromusa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1010Genus III. Malonomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1012Genus IV. Pelobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1013

Family II. Geobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1017Genus I. Geobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1017Genus II. Trichlorobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1020

Order VI. Syntrophobacterales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1021Family I. Syntrophobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1021

Genus I. Syntrophobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1021Genus II. Desulfacinum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1027Genus III. Desulforhabdus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1029Genus IV. Desulfovirga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1030Genus V. Thermodesulforhabdus . . . . . . . . . . . . . . . . . . . . . . . . . . . 1031

Family II. Syntrophaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1033Genus I. Syntrophus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1033Genus II. Desulfobacca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1035Genus III. Desulfomonile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1036

CONTENTS xxi

Genus IV. Smithella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1039Order VII. Bdellovibrionales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1040

Family I. Bdellovibrionaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1040Genus I. Bdellovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041Genus II. Bacteriovorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1053Genus III. Micavibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1057Genus IV. Vampirovibrio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1058

Order VIII. Myxococcales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1059Family I. Myxococcaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1073

Genus I. Myxococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1074Genus II. Corallococcus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1079Genus III. Pyxicoccus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1083

Family II. Cystobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1085Genus I. Cystobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1086Genus II. Archangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1097Genus III. Hyalangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1099Genus IV. Melittangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1101Genus V. Stigmatella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1104

Family III. Polyangiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1109Genus I. Polyangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1110Genus II. Byssophaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1118Genus III. Chondromyces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1121Genus IV. Haploangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1129Genus V. Jahnia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1130Genus VI. Sorangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1132

Family IV. Nannocystaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1136Genus I. Nannocystis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1137

Family V. Kofleriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1143Genus I. Kofleria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1143

Class V. Epsilonproteobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1145Order I. Campylobacterales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1145

Family I. Campylobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1145Genus I. Campylobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1147Genus II. Arcobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1161Genus III. Sulfurospirillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1165

Family II. Helicobacteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1168Genus I. Helicobacter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1169Genus II. Thiovulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1189Genus III. Wolinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1191

Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1195Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1363

xxiii

Contributors

Wolf-Rainer AbrahamChemical Microbiology Group, GBF-National Research Centrefor Biotechnology, Mascheroder Weg 1, D-38124 Braunschweig,GermanyPaula AguiarPortland State University, Portland, OR 97207-0751, USAMilton J. AllisonDepartment of Microbiology, Iowa State University, Ames, IA50011-3211, USARudolf AmannNachwuchsgruppe Molekulare Okologie, Max Planck-Institutefur Marine Mikrobiologie, Celsiusstrasse 1, D-28359 Bremen, Ger-manyGeorg AulingInstitute fur Mikrobiologie, Universitat Hannover, Schneider-berg 50, D-30167 Hannover, GermanyMarcie L. BaerBiology Department, Shippensburg University, Shippensburg, PA17257, USASimon C. BakerBirkbeck College, Malet Street, Bloomsbury, London WC1E7HX, United KingdomJose Ivo BaldaniCentro Nacional de Pesquisa de Agrobiologia, Empresa Brasileirade Pesquisa Agropecuaria, Room 247-23851-970 Seropedica,Caixa Postal 74.505, Rio de Janeiro 465, BrazilVera Lucia Divan BaldaniCentro Nacional de Pesquisa de Agrobiologia, Empresa Brasileirade Pesquisa Agropecuaria, Room 247-23851-970 Seropedica,Caixa Postal 74.505, Rio de Janeiro 465, BrazilDavid L. BalkwillDepartment of Biological Science, Florida State University, Tal-lahassee, FL 32306-4470, USAMenachem BanaiMinistry of Agriculture, Veterinary Services & Animal Health,Kimron Veterinary Institute, P.O. Box 12, Bet Dagan 50 250, Is-raelClaudio BandiDipartimento di Patologia Animale, Igiene e Sanita Pubblica Ve-terinaria, Sezione di Patologia Generale e Parassitologia, Univ-ersita degli Studi di Milano, Via Celoria 10 20133 Milano, ItalyEllen Jo BaronClinical Microbiology/Virology Laboratory, Stanford UniversityMedical Center, Stanford, CA 94305-5250, USAJaniche BeederSection for Biotechnology, Novsk Hydro ASA Research Centre,P. O. Box 2560, N-3901 Porsgruun, NorwayJulia A. BellDepartment of Microbiology and Molecular Genetics, MichiganState University, East Lansing, MI 48824-4320, USA

Nancy M.C. Bleumink-PluymDept. of Bacteriology, Inst. of Infectious Diseases & Immunology,Vet. Medicine, Universitat Utrecht, Yalelaan 1, 3584 CL Utrecht,The NetherlandsEberhard BockInst. fur Allgemeine Botanik und Botanischer Garten, UniversitatHamburg, Ohnhorststrasse 18, D-22609 Hamburg, GermanyDavid R. BooneDepartment of Environmental Biology, Portland State University,Portland, OR 97207-0751, USAEdward J. BottoneDepartment of Infectious Diseases, The Mount Sinai Hospital,New York, NY 10029-6574, USAJohn P. BowmanSchool of Agricultural Science, University of Tasmania, AntarticCRC, Private Bag 54, Hobart 7001, Tasmania, AustraliaKristian K. BrandtSection of Genetics and Microbiology, Department of Ecology,Royal Veterinary and Agricultural University, DK-1871 Frederiks-berg, DenmarkDon J. BrennerMeningitis & Special Pathogens Branch Laboratory Section, Cen-ters for Disease Control & Prevention, Atlanta, GA 30333, USAHans-Jurgen BusseInstitut fur Bakteriologie, Mykologie und Hygiene, Veterinar-medizinische Universitat Wien, Veterinarplatz 1, A-1210 Wien,AustriaDouglas E. CaldwellDept. of Applied Microbiology and Food Science, University ofSaskatchewan, Saskatoon, 51 Campus Drive, Saskatchewan S7N5A8 SK, CanadaWen Xin ChenDepartment of Microbiology, Biology College, Beijing Agricul-tural University, Beijing, P.R. ChinaJohn D. CoatesPlant and Microbial Biology, University of California, Berkeley,Berkeley, CA 94720-3102, USAJohn D. CoatesPlant and Microbial Biology, University of California, Berkeley,Berkeley, CA 94720-3102. USAMichael J. CorbelNational Institute for Biol. Standards & Control, Blanche Lane,South Mimms, Potters Bar, Hertfordshire EN6 3QG, United King-domMilton S. da CostaDepartamento de Zoologia, Centro de Neurociencias, Universi-dade de Coimbra, Apartado 3126, P-3004-517 Coimbra, PortugalSubrata K. DasInstitute of Life Sciences, Nalco square, Bhubaneswar 751 023,India

CONTRIBUTORSxxiv

Gregory A. DaschDivision of Viral and Rickettsial Diseases, Viral and RickettsialZoonoses Branch, National Center for Infectious Diseases, Cen-ters for Disease Control and Prevention, Atlanta, GA 30333, USAFrank B. DazzoDepartment of Microbiology and Molecular Genetics, MichiganState University. East Lansing, MI 48824-4320, USAKim A. DeWeerdDepartment of Chemistry, State University of New York, Univer-sity at Albany, Albany, NY 12222, USAEwald B.M. DennerAbteilung Mikrobiologie und Biotechnologie, Institut fur Mik-robiologie und Genetik, Dr. Bohr-Gasse 9, A-1030 Wein, AustriaRichard DevereuxNHEERL, Gulf Ecology Division, U.S.E.P.A., Gulf Breeze, FL32561, USAFloyd E. DewhirstDepartment of Molecular Genetics, The Forsyth Institute, 140The Fenway, Boston, MA 02115-3799, USAJohanna Dobereiner (Deceased)Centro Nacional de Pesquisa de Agrobiologia, Empresa Brasilierade Pesquisa Agropecuaria, Room 247, 23851-970 Seropedica,Caixa Postal 74.505, Rio de Janeiro 465, BrazilNina V. DoroninaInst. of Biochemistry & Physiology of Micro-organisms RAS, Lab-oratory of Methylotrophy, Russian Academy of Sciences, Push-chino-on-the-Oka, Moscow Region 142290, RussiaGalina A. DubininaInstitute of Microbiology, Russia Academy of Sciences, Prospect6—let. Oktyabrya 7/2, Moscow, RussiaJ. Stephen DumlerDivision of Microbiology, Department of Pathology, The JohnsHopkins Hospital, Univ. School of Medicine, Baltimore, MD21287-7093, USAJurgen EberspacherInstitut fur Mikrobiologie (250), Universitat Hohenheim, Gar-benstrasse 30, D-70599 Stuttgart, GermanyThomas W. EgliDepartment of Microbiology, EAWAG, Uberlandstrasse 133, CH8600 Duebendorf, SwitzerlandStefanie J.W.H. Oude ElferinkID TNO Animal Nutrition, P.O. Box 65, 8200 AB Lelystad, TheNetherlandsTakayuki T. EzakiDepartment of Microbiology and Bioinformatics, Regenerationand Advanced Medical Science, Gifu University School of Medi-cine, 40 Tsukasa-machi, Gifu 500 8705, JapanMark FeganCoop. Research Centre for Tropical Plant Protection, Dept. ofMicro. & Parasitology, The University of Queensland, St. Lucia,Brisbane, Queensland 4072, AustraliaAndreas FesefeldtGeibelallee 12a, 24116 Kiel, GermanyKai W. FinsterDepartment of Microbial Ecology, Institute of Biological Sci-ences, University of Aarhus, Building 540, Ny, Munkegade, DK-8000 Aarhus C, DenmarkJames G. FoxDepartment of Comparative Medicine, Massachusetts Institute ofTechnology, Cambridge, MA 02139, USAMichael FriedrichAbteilung Biogeochemie, Max Planck-Institut fur TerrestrischeMikrobiologie, Karl-von-Frisch-Strasse, D-35043 Marburg, Ger-many

Georg FuchsMikrobiologie, Institut fur Biologie II, Albert-Ludwigs-UniversitatFreiburg, D-79104 Freiburg, GermanyJohn A. FuerstCenter for Bacterial Diversity and Identification, Department ofMicrobiology, University of Queensland, Brisbane, Queensland4072, AustraliaJean-Louis GarciaLaboratoire de Microbiologie, ORSTOM-ESIL-Case 925, Univ-ersite de Provence, 163, Avenue de Luminy, 13288 Marseille,Cedex 9, FranceMonique Garnier (Deceased)Institut National de la Recherche Agronomique et Universite Vic-tor Segalen, Laboratoire de Biologie Cellulaire et Moleculaire,Bordeaux 2, 33883, BP 81, Villenave d’Ormon Cedex, FranceGeorge M. GarrityDept. of Microbiology and Molecular Genetics, Michigan StateUniversity, East Lansing, MI 48824, USARainer GebersDepenweg 12, D-24217 Schonberg/Holstein, GermanyConnie J. GebhartDivision of Comparative Medicine, University of MinnesotaHealth Center, Minneapolis, MN 55455, USABarbara R. Sharak GenthnerCenter for Environmental Diagnostics and Bioremediation, Uni-versity of West Florida, Pensacola, FL 32514, USAPeter Gerner-SmidtDepartment of Gastrointestinal Infections, Statens Serum Insti-tut, Artillerivej 5, DK-2300 Copenhagen S, DenmarkMonique GillisLaboratorium voor Microbiologie Vakgroep WE 10V, UniversiteitGent, K.L. Ledeganckstraat 35, B-9000 Gent, BelgiumChristian GliescheInstitut fur Okologie, Ernst-Moritz-Arndt-Universitat, GreifswaldSchwedenhagen 6, D-18565 Kloster/Hiddensee, GermanyJose M. GonzalezDepartamento de Microbiologia y Biologia Celular, Facultad deFarmacia, Universidad de La Laguna, 38071 La Laguna. Tenerife,SpainYvonne E. GoodmanDepartment of Medical Bacteriology, University of Alberta, Medi-cal Services Building, Edmonton, Alberta, CanadaVladimir M. GorlenkoInstitute of Microbiology, Russian Academy of Sciences, Prospect60-letiya, Oktyabrya 7, korpus 2, Moscow 117312, RussiaHans-Dieter GortzDepartment of Zoology, Biologisches Institut, Universitat Stutt-gart, Pfaffenwaldring 57, D-70550 Stuttgart, GermanyJohn J. GosinkAmgen, Inc., Seattle, WA 98101, USAJennifer Gossling8401 University Drive, St. Louis, MO 63105-3641, USAPeter N. GreenNational Collection of Industrial & Marine Bacteria, 23 St. Ma-char Drive, Aberdeen AB24 3RY, United KingdomLotta E-L. HallbeckDepartment of Cell and Molecular Biology, Goteborg University,Medicinaregatan 9 C, Box 462, S-405 30 Goteborg, SwedenTheo A. HansenDepartment of Microbial Physiology, Groningen BiomolecularSci. & Biotech. Inst., University of Groningen, P. O. Box 14, 9750AA Haren, The Netherlands

CONTRIBUTORS xxv

Anton HartmannInstitute of Soil Ecology, Rhizosphere Biology Division, GSF Re-search Center, PO Box 1129, D-85764 Neuherberg, Munchen,GermanyFawzy M. HashemSustainable Agriculture Laboratory, Animal and Natural Re-sources Institute, Beltsville Agricultural Research Institute,USDA-ARS, Beltsville, MD 20705, USABrian P. HedlundDepartment of Biological Sciences, University of Nevada, Las Ve-gas, Las Vegas, NV 89154-4004, USAJohann HeiderMikrobiologie, Institut fur Biologie II, Universitat Freiburg,Schanzlestrasse 1, D-79104 Freiburg, GermanyKarl-Heinz HinzKlinik fur Geflugel der Tierarztlichen Hochschule, Bunteweg 17,D-30559 Hannover, GermanyAkira HiraishiDepartment of Ecological Engineering, Toyohashi University ofTechnology, Tempaku-cho, Toyohashi 441-8580, JapanPeter HirschInstitut fur Allgemeine Mikrobiologie der Biozentrum, Univer-sitat Kiel, Am Botanischen Garten 1-9, D-24118 Kiel, GermanyBecky HollenDepartment of Biological Sciences, Louisiana State University,Baton Rouge, LA 70803, USABarry HolmesPublic Health Laboratory Service, Central Public Health Labo-ratory, National Collection of Type Cultures, 61 Colindale Ave-nue, London NW9 5HT, United KingdomJohn HoltDepartment of Microbiology and Molecular Genetics, MichiganState University, East Lansing, MI 48824-1101, USAPhilip HugenholtzEcosystem Sciences Division, Department of Environmental Sci-ence, Policy, and Management, University of California, Berkeley,Berkeley, CA 94720-3110, USAThomas HurekArbeitsgruppe Symbioseforschung, Planck-Institut fur Terrestris-che Mikrobiologie, Karl-von-Frisch-Strasse, D-35043 Marburg,GermanyJohannes F. ImhoffInstitut fur Meereskunde, Abt. Marine Mikrobiologie, UniversitatKiel, Dusternbrooker Weg 20, D-24105 Kiel, GermanyKjeld IngvorsenDepartment of Microbial Ecology, Institute of Biological Sci-ences, University of Aarhus, Building 540, Ny Munkegade, DK-8000 Aarhus C, DenmarkFrancis L. JacksonMedical Microbiology and Immunology, University of Alberta, 1-41-Medical Sciences Building, Edmonton, Alberta AB T6G 2H7,CanadaCheryl JenkinsDepartment of Microbiology, University of Washington, Seattle,WA 98195-0001, USA

Sibylle KalmbachStudienstiftung des Deutschen Volkes, Mirbachstrasse 7, D-53173Bonn, Germany

Peter KampferInstitut fur Angewandte Mikrobiologie, Justus-Liebig-UniversitatGiessen, Heinrich-Buff-Ring 26-32, IFZ, D-35392 Giessen, Ger-many

Yoshiaki KawamuraDepartment of Microbiology, Gifu University School of Medicine,40 Tsukasa-machi, Gifu 500 8705, JapanDonovan P. KellyDepartment of Biological Sciences, University of Warwick, Cov-entry CV4 7AL, United KingdomSuzanne V. KellyProfessor of Biology, Scottsdale Community College, Scottsdale,AZ 85250, USAChristina KennedyDepartment of Plant Pathology, College of Agriculture, The Uni-versity of Arizona, Tucson, AZ 85721-0036, USAChristina KennedyDivision of Plant Pathology and Microbiology, Department ofPlant Pathology, The University of Arizona, Tucson, AZ 85721-0036, USAAllen KerrWaite Agricultural Research Institute, The University of Adelaide,Glen Osmond 5064, South AustraliaKarel KerstersLab. voor Microbiologie, Vakgroep Biochemie, Fysiologie en Mi-crobiologie, Rijksuniversiteit Gent, K.L. Ledeganckstraat 35, B-9000 Gent, BelgiumHans-Peter KlenkVP Genomics, Epidauros Biotechnology Inc., Am Neuland 1, D-82347 Bernried, GermanyOliver KlimmekBiozentrum Niederursel, Institut fur Mikrobiologie der JohannWolfgang Goethe-Universitat, Marie-Curie-Strasse 9, D-60439Frankfurt am Main, GermanyAllan E. KonopkaDepartment of Biological Science, Purdue University, West La-fayette, IN 47907-2054, USAHans-Peter KoopsAbteilung Mikrobiologie, Inst. fur Allgemeine Botanik und Bo-tanischer Garten, Universitat Hamburg, Ohnhorststrasse 18, D-22609 Hamburg, GermanyYoshimasa KosakoThe Institute of Physical and Chemical Research, Japan Collec-tion of Microorganisms, RIKEN, Wako-shi, Saitama 351-0198, Ja-panJulius P. KreierDepartment of Microbiology, The Ohio State University, Colum-bus, OH 43201, USANoel R. KriegDepartment of Biology, Virginia Polytechnic Institute & StateUniversity, Blacksburg, VA 24061-0406, USAAchim Kroger (Deceased)Biozentrum Niederursel, Institut fur Mikrobiologie der JohannWolfgang Goethe-Universitat, Marie-Curie-Strasse 9, D-60439Frankfurt am Main, GermanyJ. Gijs KuenenFaculty of Chemical Tech. & Materials Science, Kluyver Labora-tory for Biotechnology, Delft University of Technology, 2628 BCDelft, The NetherlandsJan KueverDepartment of Microbiology, Institute for Material Testing,Foundation Institute for Materials Science, D-28199 Bremen,GermanyL. David KuykendallMolecular Plant Pathology Laboratory, Plant Sciences Institute,United States Department of Agriculture, Beltsville, MD 20705-2350, USA

CONTRIBUTORSxxvi

David P. LabedaNatl. Ctr. for Agricultural Utilization Research, Microbial Prop-erties Research, U.S. Department of Agriculture, Peoria, IL61604-3999, USAMatthias LabrenzInstitut fur Allgemeine Mikrobiologie, Biologiezentrum, Univer-sity of Kiel, Am Botanischen Garten 1-9, 24118 Kiel, GermanyAdrian LeeSchool of Microbiology and Immunology, University of NewSouth Wales, Kensington, Sydney, AustraliaWerner LiesackMax Planck-Institut fur Terrestrische Mikrobiologie, Karl-von-Frisch-Strasse, D-35043 Marburg, GermanyTimothy LilburnATCC Bioinformatics, Manassas, VA 20110-2209, USANiall A. LoganSchool of Biological and Biomedical Sciences, Glasgow Caledon-ian University, Cowcaddens Road, Glasgow G4 0BA, United King-domDerek R. LovleyDepartment of Microbiology, University of Massachusetts, Physi-ology & Ecology of Anaerobic Micro., Amherst, MA 01003, USAWolfgang LudwigLehrstuhl fur Mikrobiologie, Technische Universitat Munchen,Am Hochanger 4, D-85350 Freising, GermanyBarbara J. MacGregorMax Planck-Institute for Marine Microbiology, Celsiusstrasse 1,D-28359 Bremen, GermanyMichael T. MadiganDepartment of Microbiology, Life Science II, Southern IllinoisUniversity, Carbondale, IL 62901-6508, USAAsa MalmqvistANOX AB, Klosterangsvagen 11A, S-226 47 Lund, SwedenWerner ManzSection G3, Ecotoxicology and Biochemistry, German Federal In-stitute of Hydrology, Kaiserin-Augusta-Anlagen 15-17, P. O. Box20 02 53, D-56002 Koblenz, GermanyEsperanza Martınez-RomeroCentro de Investigacion sobre Fijacion de Nitrogeno, UNAM, ApPostal 565–A, Cuernavaca, Morelos, MexicoAbdul M. MaszenanEnvironmental Engineering Research Centre, School of Civil andStructural Engineering, Nanyang Technological University,Block N1, #1a-29, 50 Nanyang Avenue, Singapore 639798Michael J. McInerneyDepartment of Botany and Microbiology, The University of Okla-homa, Norman, OK 73019-6131, USASteven McOristDepartment of Biomedical Sciences, Tufts University College ofVeterinary Medicine, North Grafton, MA 01536, USARoy D. Meredith (Deceased)Joris MergaertLaboratorium voor Microbiologie Vakgroep Biochemie, Fysiolo-gie en Microbiol., Universiteit Gent, K.L. Ledeganckstraat 35, B-9000 Gent, BelgiumOrtwin D. MeyerLehrstuhl fur Mikrobiologie, Universitat Bayreuth, Universitatss-trasse 30, D-95440 Bayreuth, GermanyEdward R.B. MooreProgramme of Soil Quality and Protection, The Macaulay Re-search Institute, Macaulay Dr., Craigiebuckler, AB15 8QH Aber-deen, United Kingdom

R.G.E. MurrayDepartment of Microbiology and Immunology, The University ofWestern Ontario, London, Ontario N6A 5C1, Canada

Yasuyoshi NakagawaBiological Resource Center (NBRC), Department of Biotechnol-ogy, National Institute of Technology and Evaluation, 2-5-8, Ka-zusakamatari, Kisarazu, Chiba 292-0818, Japan

M. Fernanda NobreDepartmento de Zoologia, Universidade de Coimbra, Apartado3126, P-3000 Coimbra, Portugal

Jani L. O’RourkeSchool of Microbiology and Immunology, University of NewSouth Wales, Kensington, Sydney, Australia

Bernard OllivierLaboratoirede Microbiologie—LMI, ORSTOM, Case 925, Univ-ersite de Provence, ESIL, 163 Avenue de Luminy, Marseille 13288Cedex 09, France

Stephen L.W. OnDanish Veterinary Institute, Bulowsvej 27, DK-1790, CopenhagenV, Denmark

Ronald S. OremlandWater Research Division, U.S. Geological Survey, Menlo Park, CA94025-3591, USA

Norberto J. PalleroniRutgers, North Caldwell, NJ 07006-4146, USA

Bruce J. PasterDepartment of Molecular Genetics, The Forsyth Institute, 140The Fenway, Boston, MA 02115-3799, USA

Bharat K.C. PatelMicrobial Discovery Research Unit, School of Biomolecular Sci-ences, Griffith University, Nathan Campus, Kessels Road, Bris-bane, Queensland 4111, Australia

Dominique PatureauLaboratoire de Biotechnologie de l’Environnement, INRA Nar-bonne, avenue des etangs, 11 100 Narbonne, France

Karsten PedersenDepartment of Cell and Molecular Biology, Goteborg University,Medicinaregatan 9 C, Box 462, S-405 30 Goteborg, Sweden

Jeanne S. PoindexterDepartment of Biological Sciences, Barnard College, ColumbiaUniversity, New York, NY 10027-6598, USA

Andreas Pommerening-RoserAbteilung Mikrobiologie, Inst. fur Allgemeine, Botanik und Bo-tanischer Garten, Universitat Hamburg, Ohnhorststrasse 18, D-22609 Hamburg, Germany

Bruno PotScience Department, Yakult Belgium, Joseph Wybranlaan 40, B-1070 Brussels, Belgium

Fred A. RaineyDepartment of Biological Sciences, Louisiana State University,Baton Rouge, LA 70803, USA

Didier RaoultFaculte de Medecine, CNRS, Unite des Rickettsies, 27 BoulevardJean Moulin, 13385 Marseille Cedex 05, France

Christopher RathgeberDepartment of Microbiology, The University of Manitoba, Win-nipeg, Manitoba R3T 2N2, Canada

Gavin N. ReesMurray-Darling Freshwater Research Centre, CRC FreshwaterEcology, Ellis Street, Thurgoona, PO Box 921, Albury NSW 2640,Australia

CONTRIBUTORS xxvii

Hans ReichenbachArbeitsgruppe Mikrobielle Sekundarstoffe, Gesellschaft fur Bio-technologische Forschung mbH, Mascheroder Weg 1, D-38124Braunschweig, GermanyBarbara Reinhold-HurekUniversitat Bremen, Fachbereich 2, Allgemeine Mikrobiologie,P. O. Box 330440, D-28334 Bremen, GermanyAnna-Louise ReysenbachDepartment of Environmental Biology, Portland State University,Portland, OR 97207, USAYasuko RikihisaDepartment of Veterinary Biosciences, The Ohio State Univer-sity, 1925 Coffey Road, Columbus, OH 43210-1093, USALesley A. RobertsonKluyver Laboratory for Biotechnology, Delft University of Tech-nology, Julianalaan 67, P. O. Box 5057, 2628BC Delft, The Neth-erlandsTakeshi SakaneInstitute for Fermentation, Osaka, Yodogawa-ku, Osaka 532-8686,JapanAbigail A. SalyersDepartment of Microbiology, University of Illinois-Urbana,Champaign, Urbana, IL 61801-3704, USAGary N. SandenEpidemic Investigations Laboratory, Meningitis and Special Path-ogens Branch, Division of Bacterial and Mycotic Diseases, Centersfor Disease Control and Prevention, Atlanta, GA 30333, USAHiroyuki SawadaNational Institute of Agro-Environmental Sciences, 3-1-1 Kannon-dai, Tsukuba, Ibaraki 305-8604, JapanBernhard H. SchinkFakultat fur Biologie, Lehrstuhl fur Mikrobielle Okologie, Univ-ersitat Konstanz, Postfach 55 60, D-78457 Konstanz, GermanyKarl-Heinz SchleiferLehrstuhl fur Mikrobiologie, Technische Universitat Munchen,Am Hochanger 4, Freising D-85350, GermanyKarl-Heinz SchleiferLehrstuhl fur Mikrobiologie, Technische, Universitat Munchen,Am Hochanger 4, Freising D-85350, GermanyHeinz SchlesnerInstitut fur Allgemeine Mikrobiologie, Universitat Kiel, Am Bo-tanischen Garten 1-9, Biologiezentrum, D-24118 Kiel, GermanyHelmut J. SchmidtBiological Faculty, University of Kaiserslautern, Building 14, Pf3049, D-67653 Kaiserslautern, GermanyJean M. SchmidtDepartment of Microbiology, Arizona State University, Tempe,AZ 85287-2701, USADirk SchulerMax Planck-Institute for Marine Microbiology, Celsiusstrasse 1,D-28359 Bremen, GermanyBernard La ScolaCNRS UMR6020, Unite des Rickettsies, 27 Boulevard Jean Mou-lin, 13385 Marseille Cedex 05, FrancePaul SegersLab. voor Microbiologie Vakgroep WE 10V, Universiteit Gent,K.L. Ledeganckstraat 35, B-9000 Gent, BelgiumRobert J. SeviourBiotechnology Research Centre, La Trobe University, P.O. Box199, Bendigo VIC 3550, AustraliaRichard SharpSchool of Applied Sciences, South Bank University, 103 BoroughRoad, London SE1 0AA, United Kingdom

Tsuneo ShibaShimonoseki University of Fisheries, Dept. of Food Science andTechnology, Yoshimi-Nagatahoncho Shimonose, Yamaguchi 759-65, Japan

Martin SieversUniversity of Applied Sciences, Department of Biotechnology,Molecular Biology, P. O. Box 335, CH 8820 Wadenswil, Switzer-land

Lindsay I. SlyCentre for Bacterial Diversity and Identification, Department ofMicrobiology and Parasitology, University of Queensland, St. Lu-cia, Brisbane, Queensland 4072, Australia

Peter H.A. SneathDepartment of Microbiology and Immunology, School of Medi-cine, University of Leicester, P.O. Box 138, Leicester LE1 9HN,United Kingdom

Martin SobierajDepartment of Environmental Biology, Portland State University,P. O. Box 751, Portland, OR 97207-0751, USA

Dimitry Y. SorokinInstitute of Microbiology, Russian Academy of Sciences, Prospect60-let. Oktyabrya 7/2, Moscow 117811, Russia

Rob J.M. van SpanningDepartment of Molecular Cell Physiology/Molecular MicrobialEcology, Vrije Universiteit, De Boelelaan 1087, NL-1081 HV Am-sterdam, The Netherlands

Eva SpieckInst. fur Allgemeine Botanik und Botanischer Garten, UniversitatHamburg, Ohnhorststrasse 18, D-22609 Hamburg, Germany

Georg A. SprengerForschungszentrum Julich GmbH, Institut fur Biotechnologie 1,P. O. Box 1913, D-52425 Julich, Germany

Stefan SpringDSMZ-Deutsche Sammlung von Mikrooorganismen und Zellk-ulturen, GmbH, D-38124 Braunschweig, Germany

David A. StahlCivil and Environmental Engineering, University of Washington,Seattle, WA 98195-2700, USA

James T. StaleyDepartment of Microbiology, University of Washington, Seattle,WA 98195-0001, USA

Alfons J.M. StamsDepartment of Microbiology, Wageningen Agricultural Univer-sity, Hesselink Van Suchtelenweg 4, NL-6703 CT Wageningen,The Netherlands

Patricia M. StanleyMinntech Corporation, North, Minneapolis, MN 55447-4822,USA

John F. StolzDepartment of Biological Sciences, Duquesne University, Pitts-burgh, PA 15282-2504, USA

Adriaan H. StouthamerDept. of Molecular Cell Physiology/Molecular Microbial Ecology,Vrije Universiteit, De Boelelaan 1087, NL-1081 HV Amsterdam,The Netherlands

William R. StrohlMerck & Company, Rahway, NJ 07065-0900, USA

Joseph M. SuflitaEnvironmental and General Applied Microbiology, Departmentof Botany & Micro., The University of Oklahoma, Norman, OK73019-0245, USA

CONTRIBUTORSxxviii

Jorg SulingInstitut fur Meereskunde, Abt Marine Mikro-biologie, UniversitatKiel, Dusternbrooker Weg 20, D-24105 Kiel, GermanyJean SwingsLaboratorium voor Microbiologie, Vakgroep WE10V, Fysiologieen Microbiologie, Universiteit of Gent, K.L. Ledeganckstraat 35,B-9000 Gent, BelgiumUlrich SzewzykDepartment of Microbial Ecology, Technical University Berlin,Franklinstrasse 29, Secr. OE 5, D-10587 Berlin, GermanyZhiyuan TanDepartment of Microbiology and Molecular Genetics, College ofAgronomy, South China Agricultural University, 510642, ChinaAnders TernstromANOX AB, Klosterangsvagen 11A, S-226 47 Lund, SwedenTone TønjumInstitute of Microbiology, Section of Molecular Microbiology A3,Rikshospitalet (National Hospital), Pilestredet 32, N-0027 Olso,NorwayG. Todd TownsendUniversity of Oklahoma, Norman, OK 73072, USAYuri A. TrotsenkoInstitute of Biochemistry and Physiology of Microorganisms RAS,Laboratory of Methylotrophy, Prospekt Nauki, 5, Moscow Region142290, RussiaHans G. TruperInstitut fur Mikrobiologie und Biotechnologie, Universitat Bonn,Mechenheimer Allee 168, W-53115 Bonn, GermanyRichard F. UnzDepartment of Civil Engineering, The Pennsylvania State Uni-versity, University Park, PA 16802-1408, USATeizi UrakamiBiochemicals Development Div., Mitsubishi Building, MitsubishiGas Chemical Company, 5-2, Marunouchi 2-chome, Chiyoda-ku,Tokyo 100-8324, JapanMarc VancanneytLaboratorium voor Microbiologie, Universiteit Gent, K.L. Lede-ganckstraat 35, B-9000 Gent, BelgiumPeter VandammeLab. voor Microbiologie en Microbiele Genetica, Univeristeit ofGent, Faculteit Wetenschappen, K.L. Ledeganckstraat 35, B-9000Gent, BelgiumLeana V. VasilyevaInstitute of Microbiology RAN, 117811, Russian Academy of Sci-ences, 60-let. Oktyabrya 7 build. 2, Moscow, RussiaHenk W. van VerseveldDept. of Molecular Cell Physiology, Molecular and MicrobialEcology, Vrije Universiteit, De Boelelaan 1087, NL-1081 HV Am-sterdam, The NetherlandsPaul De VosDept. of Biochem., Physiology and Microbiology (WE 10V), Uni-versity of Gent, K.L. Ledeganckstraat 35, B-9000 Gent, BelgiumDavid H. WalkerDepartment of Pathology, University of Texas Medical Branch,30l University Boulevard, Galveston, TX 77555-0609, USAEn Tao WangDepartamento de Microbiologia, Escuela Nacional de CienciasBiologicas, Instituto Politecnico Nacional, Carpio y Plan de AyalaS/N, Mexico D.F. 11340, MexicoNaomi L. WardThe Institute for Genomic Research, Rockville, MD 20850, USARichard I. WebbDepartment of Microbiology, University of Queensland, Bris-bane, Queensland 4072, Australia

Ronald M. WeinerCell Biology Cluster, Division of Molecular and Cellular Biosci-ences, National Science Foundation, Arlington, VA 22230, USADavid F. WelchLaboratory Corporation of America, Dallas, Texas 75230, USAAimin WenFood Science and Technology Program, Pacific Agri-Food Re-search Centre, Summerland BC V0H 1Z0, CanadaHannah M. WexlerDepartment of Veterans Affairs, West Los Angeles Medical Ctr.,UCLA School of Medicine, 11301 Wilshire Boulevard, Los An-geles, CA 90073, USARobbin S. WeyantMeningitis & Special Pathogens Branch, Centers for Disease Con-trol and Prevention, Atlanta, GA 30333, USAAnne M. WhitneyMeningitis & Special Pathogens Branch Lab. Section, MS D-11,Centers for Disease Control & Prevention, Atlanta, GA 30303,USAFriedrich W. WiddelAbteilung Mikrobiologie, Max Planck-Institut fur Marine Mikro-biologie, Celsiusstrasse 1, D-28359 Bremen, GermanyJurgen K.W. WiegelDepartment of Microbiology, University of Georgia, Athens, GA30602-2605, USAAnne WillemsLaboratorium voor Microbiologie, Universiteit Gent, K.L. Lede-ganckstraat 35, B-9000 Gent, BelgiumHenry N. WilliamsDepartment of OCBS, Dental School, University of Maryland atBaltimore, Baltimore, MD 21201-1510, USAAnn P. WoodMicrobiology Research Group, King’s College, London, Div. ofLife Sciences, Franklin-Wilkins Building, 150 Stamford Street,London SE1 8WA, United KingdomEiko YabuuchiAichi Medical University, Omiya 4-19-18, Asahi-ku, Osaka 535-0002, JapanAkira YokotaInstitute of Molecular and Cellular Biosciences, The Universityof Tokyo, Yayoi 1-1-1, Bunkyo-ku, Tokyo 113-0032, JapanJohn M. YoungMt. Albert Research Centre, Landcare Research New ZealandLtd., Private Bage 92 170, Auckland, New ZealandXue-jie YuDepartment of Pathology, University of Texas Medical Branch,30l University Boulevard, Galveston, TX 77555-0609Vladimir V. YurkovDepartment of Microbiology, The University of Manitoba, Win-nipeg, Manitoba R3T 2N2, CanadaGeorge A. ZavarzinInstitute of Microbiology, Russian Academy of Sciences, Building2, Prospect 60-letja Oktyabrya 7a, Moscow 117312, RussiaBernard A.M. van der ZeijstNational Institute of Public Health and Environ., Antonie vanLeeuwenhoeklaan 9, P. O. Box 1, P. O. Box 80.165, 3720 BABilthoven, The NetherlandsTatjana N. ZhilinaInstitute of Microbiology, Russian Academy of Sciences, Prospect60-letja Oktyabrya 7a, Moscow 117312, RussiaStephen H. ZinderDepartment of Microbiology, Cornell University, Ithaca, NY14853-0001, USA