BERGEY’S MANUAL OF Systematic Bacteriology - Springer

1 BERGEY S MANUAL OFSystematicBacteriologySecond EditionVolume TwoTheProteobacteriaPart CTheAlpha-, Beta-, Delta-, andEpsilonproteobacteriaBERGEY S MANUAL OFSystematicBacteriologySecond EditionVolume TwoTheProteobacteriaPart CTheAlpha-, Beta-, Delta-, andEpsilonproteobacteriaDon J. BrennerNoel R. KriegJames T. StaleyEDITORS, VOLUME TWOG eorge M. GarrityEDITOR-IN-CHIEFEDITORIAL BOARDJ ames 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 SchleiferWITH CONTRIBUTIONS FROM 222 COLLEAGUESG eorge M. Garrity, s MANUAL TrustDepartment of Microbiology and Molecular GeneticsMichigan State UniversityEast Lansing, MI 48824-4320 USAL ibrary of Congress Control Number: 2005926296 2005, 1984 1989 Bergey s MANUAL Trust This work is subject to copyright.

2 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.

3 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.

4 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 ( ) ISBN ISBN-13: 978-0-387-24145-6-10: 0-387-24145-0 This volume is dedicated to our colleagues,David R.

5 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 helpingthe Trust meet its BOARD AND TRUSTEESOF BERGEY S MANUAL TRUSTJ ames 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, Murray,EmeritusCharles F.

6 Niven, Jr.,EmeritusNorbert Pfennig,EmeritusPeter Sneath,EmeritusJoseph G. Tully,EmeritusStanley T. Williams,EmeritusixPreface to Volume Two of the Second Edition ofBergey s MANUAL of Systematic BacteriologyThere is a long-standing tradition for the Editors ofBergey s Man-ualto 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 , 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.

7 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 amongBacteriaandArchaea. 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 ofBergey 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.

8 Thisdeparts from the First Edition, as well as the Eighth and NinthEditions of theDeterminative 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 Editors remind the readers that theSystematics Manualisa 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.

9 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 oftheir manuscripts. We would also like to thank Drs. Hans Tru per,Brian Tindall, and Jean Euze by for their assistance on mattersof nomenclature and 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.

10 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 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 project such as theSystematics Manualalso requires thestrong and continued support of a dedicated publisher, and wehave been most fortunate in this regard.