BERGEY’S MANUAL� OF Systematic Bacteriology Second Edition Volume Two The Proteobacteria Part B The Gammaproteobacteria BERGEY’S MANUAL� OF Systematic Bacteriology Second Edition Volume Two The Proteobacteria Part B The Gammaproteobacteria Don J. Brenner Noel R. Krieg James T. Staley EDITORS, VOLUME TWO George M. Garrity EDITOR-IN-CHIEF EDITORIAL BOARD James 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 339 COLLEAGUES George M. Garrity, Sc.D. Bergey’s Manual Trust Department of Microbiology and Molecular Genetics Michigan State University East Lansing, MI 48824-4320 USA Library of Congress Cataloging-in-Publication Data TO COME This volume is dedicated to our colleagues, David R. Boone, Don J. Brenner, Richard W. Castenholz, and Noel R. Krieg, who retired from the Board of Trustees of Bergey’s Manual Trust as this edition was in preparation. We deeply appreciate their efforts as editors and authors; they have devoted their time and many years in helping the Trust meet its objectives. EDITORIAL BOARD AND TRUSTEES OF BERGEY’S MANUAL TRUST James T. Staley, Chairman David R. Boone, Vice Chairman George M. Garrity Paul De Vos Michael Goodfellow Fred A. Rainey Karl-Heinz Schleifer Don J. Brenner, Emeritus Richard W. Castenholz, Emeritus John G. Holt, Emeritus Noel R. Krieg, Emeritus John Liston, Emeritus James W. Moulder, Emeritus R.G.E. Murray, Emeritus Charles F. Niven, Jr., Emeritus Norbert Pfennig, Emeritus Peter H.A. Sneath, Emeritus Joseph G. Tully, Emeritus Stanley T. Williams, Emeritus ix Preface to Volume Two of the Second Edition of Bergey’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 that the new edition is a departure from the earlier ones. As this volume goes to press, however, we recognize a need to deviate from this practice, by offering a separate preface to each volume within this edition. In part, this departure is necessary because the size and complexity of this edition far exceeded our expec- tations, as has the amount of time that has elapsed between publication 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 methodological advances that will ultimately lead to a more perfect and useful classification scheme. Clearly, the pace has been accelerating as evidenced in the super-linear rate at which new taxa are being described. Much of the increase can be attributed to rapid ad- vances in sequencing technology, which has brought about a major shift in how we view the relationships among Bacteria and Archaea. While the possibility of a universally applicable natural classification was evident as the First Edition was in preparation, it is only recently that the sequence databases became large enough, and the taxonomic coverage broad enough to make such an arrangement feasible. We have relied heavily upon these data in organizing the contents of this edition of Bergey’s Manual of Systematic Bacteriology, which will follow a phylogenetic frame- work based on analysis of the nucleotide sequence of the small ribosomal subunit RNA, rather than a phenotypic structure. This departs from the First Edition, as well as the Eighth and Ninth Editions of the Determinative Manual. While the rationale for pre- senting the content of this edition in such a manner should be evident to most readers, they should bear in mind that this edi- tion, as in all preceding ones represents a progress report, rather than a final classification of procaryotes. The Editors remind the readers that the Systematics Manual is a peer-reviewed collection of chapters, contributed by authors who were invited by the Trust to share their knowledge and expertise of specific taxa. Citation should refer to the author, the chapter title, and inclusive pages rather than to the Editors. The Trust is indebted to all of the contributors and reviewers, without whom this work would not be possible. The Editors are grateful for the time and effort that each expended on behalf of the entire scientific community. We also thank the authors for their good grace in accepting comments, criticisms, and editing of their manuscripts. We would also like to thank Drs. Hans Tru¨per, Brian Tindall, and Jean Euze´by for their assistance on matters of nomenclature and etymology. We would like to express our thanks to the Department of Microbiology and Molecular Genetics at Michigan State Univer- sity for housing our headquarters and editorial office and for providing a congenial and supportive environment for microbial systematics. We would also like to thank Connie Williams not only for her expert secretarial assistance, but also for unflagging dedication to the mission of Bergey’s Manual Trust and Drs. Julia Bell and Denise Searles for their expert editorial assistance and diligence in verifying countless pieces of critical information and to Dr. Timothy G. Lilburn for constructing many of the phylogenetic trees used in this volume. We also extend our thanks to 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 the strong and continued support of a dedicated publisher, and we have been most fortunate in this regard. We would also like to express our gratitude to Springer-Verlag for supporting our ef- forts and for the development of the Bergey’s Document Type Definition (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 in designing and developing our DTD, and Jeri Lambert and Leslie Grossberg of Impressions Book and Journal Services for their efforts during the pre-production and production phases. We would 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 my personal thanks to my fellow trustees for providing me with the opportunity to participate in this effort, to Drs. Don Brenner, Noel Krieg, and James Staley for their enormous efforts as volume editors 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 Microbiology and Molecular Genetics, 6162 Biomedical and Physical Sciences Building, Michigan State University, East Lansing, MI, USA 48824-4320. Email: