BERGEY'S MANUAL® OF Systematic Bacteriology Second Edition

Volume One The Archaea and the Deeply Branching and Phototrophic

David R. Boone Richard W. Castenholz EDITORS, VOLUME ONE

George M. Garrity EDITOR-IN-CHIEF

EDITORIAL BOARD James T. Staley, Chairman, David R. Boone, Vice Chairman, Don J. Brenner, Richard W. Castenholz, George M. Garrity, Michael Goodfellow, Noel R. Krieg, Fred A. Rainey, Karl-Heinz Schleifer

WITH CONTRIBUTIONS FROM 105 COLLEAGUES

Springer Contents

Preface to the Second Edition of Bergey's Manual® of Systematic Bacteriology vjj Preface to the First Edition of Bergey's Manual® of Systematic Bacteriology jX Preface to the First Edition of Bergey's Manual® of Determinative Bacteriology xi Contributors xjx The History of Bergey's Manual 1 On Using the Manual -15 Procaryotic Domains 21 Classification of Procaryotic Organisms and the Concept of Bacterial Speciation 27 Identification of Procaryotes 33 Numerical 39 Polyphasic Taxonomy 43 Overview: A Phylogenetic Backbone and Taxonomic Framework for Procaryotic Systematics 49 Nucleic Acid Probes and Their Application in Environmental Microbiology 67 Bacterial Nomenclature 83 Etymology in Nomenclature of Procaryotes 89 Microbial Ecology—New Directions, New Importance 101 Culture Collections: An Essential Resource for Microbiology 111 Intellectual Property of Procaryotes 115 The Road Map to the Manual 119

DOMAIN ARCHAEA

PHYLUM Al Crenarchaeota 169 Class I. Thermoprotei 169 Order I. Thermoproteales 170 Family I. Thermoproteaceae 170 I. Thermoproteus 171 Genus II. Caldivirga 173 Genus III. Pyrobaculum 174 Genus IV. Thermocladium 177 Family II. Thermofilaceae 178 Genus I. Thermofilum 178 Order II. Desulfurococcales 179 Family I. Desulfurococcaceae 180 Genus I. Desulfurococcus 181 Genus II. Aeropyrum 183 Genus III. Ignicoccus 184 Genus IV. Staphylothermus 186 Genus V. Stetteria 187 Genus VI. Sulfophobococcus 188

xiii xiv CONTENTS

Genus VII. Thermodiscus 189 Genus VIII. Thermosphaera 190 Family II. Pyrodictiaceae 191 Genus I. Pyrodictium 192 Genus II. Hyperthermus 195 Genus III. Pyrolobus 196 Order III. Sulfolobales 198 Family I. Sulfolobaceae 198 Genus I. Sulfolobus 198 Genus II. Acidianus 202 Genus III. Metallosphaera 204 Genus IV. Stygiolobus 207 Genus V. Sulfurisphaera 208 Genus VI. Sulfurococcus 209

PHYLUM AH Euryarchaeota 211 Taxonomy of Methanogenic Archaea 211 Class I. Methanobacteria 213 Order Methanobacteriales 214 Family I. Methanobacteriaceae 214 Genus I. Methanobacterium 215 Genus II. Methanobrevibacter 218 Genus III. Methanosphaera 226 Genus IV. Methanothermobacter 230 Family II. Methanothermaceae 233 Genus I. Methanothermus 233 Class II. Methanococci 235 Order I. Methanococcales 236 Family I. Methanococcaceae 236 Genus I. Methanococcus 236 Genus II. Methanothermococcus 241 Family II. Methanocaldococcaceae 242 Genus I. Methanocaldococcus 243 Genus II. Methariotorris 245 Order II. Methanomicrobiales 246 Family I. Methanomicrobiaceae 247 Genus I. Methanomicrobium 247 Genus II. Methanoculleus 251 Genus III. Methanofollis 253 Genus IV. Methanogenium 256 Genus V. Methanolacinia 258 Genus VI. Methanoplanus 259 Family II. Methanocorpusculaceae 262 Genus I. Methanocorpusculum 262 Family III. Methanospirillaceae 264 Genus I. Methanospirillum 264 Genus Incertae Sedis I. Methanocalculus 267 Order III. Methanosarcinales 268 Family I. Methanosarcinaceae 268 Genus I. Methanosarcina 269 Genus II. Methanococcoides 276 Genus III. Methanohalobium 279 Genus IV. Methanohalophilus 281 Genus V. Methanolobus 283 Genus VI. Methanosalsum 287 Family II. Methanosaetaceae 289 CONTENTS xv

Genus I. Methanosaeta 289 Class III. Halobacteria 294 Order I. Halobacteriales 294 Family I. Halobacteriaceae 299 Genus I. Halobacterium 301 Genus II. Haloarcula 305 Genus III. Halobaculum 309 Genus IV. Halococcus 311 Genus V. Haloferax 315 Genus VI. Halogeometricum 318 Genus VII. Halorubrum 320 Genus VIII. Haloterrigeria 324 Genus IX. Natrialba 325 Genus X. Natrinema 327 Genus XI. Natronobacterium 329 Genus XII. Natronococcus 330 Genus XIII. Natronomonas 332 Genus XIV. Natronorubrum 333 Class IV. Thermoplasmata 335 Order I. Thermoplasmata/es 335 Family f. Thermoplasmataceae 335 Genus I. Thermoplasma 335 Family El. Picrophilaceae 339 Genus I. Picrophilus 339 Class IV. Thermococci 341 Order I. Thermococcales 341 Family I. Thermococcaceae 341 Genus I. Thermococcus 342 Genus II. Pyrococcus 346 Class VI. Archaeoglobi 349 Order I. Archaeoglobales 349 Family I. Archaeoglobaceae 349 Genus I. Archaeoglobus 349 Genus II. Ferroglobus 352 Class VII. Methanopyri 353 Order I. Methanopyrales 353 Family I. Methanopyraceae 353 Genus I. Methanopyrus 354

DOMAIN BACTERIA

PHYLUM BS 359 Class I. Aquificae 359 Order I. Aquificales 359 Family I. Aquificaceae 360 Genus I. Aquifex 360 Genus II. Calderobacterium 362 Genus III. Hydrogenobacter 363 Genus IV. Thermocrinis 364 Genus Incertae Sedis I. Desulfurobacterium 366

PHYLUM BII 369 Class I. Thermotogae 369 Order I. Thermotogales 369 Family I. Thermotogaceae 370 Genus I. Thermotoga 370 xvi CONTENTS

Genus II. Fervidobacterium 375 Genus III. Geotoga 377 Genus IV. Petrotoga 382 Genus V. Thermosipho 385

PHYLUM Bill Thermodesulfobacteria 389 Class I. Thermodesulfobacteria 389 Order I. Thermodesulfobacteriales 389 Family I. Thermodesulfobacteriaceae 390 Genus I. Thermodesulfobacterium 390

PHYLUM BIV "Deinococcus-Thermus" 395 Class I. Deinococci 395 Order I. Deinococcales 395 Family I. Deinococcaceae 395 Genus I. Deinococcus 396 Order II. Thermales 403 Family I. Thermaceae 403 Genus I. Thermus 404 Genus II. Meiothermus 414

PHYLUM BV Chrysiogenetes 421 Class I. Chrysiogenetes 421 Order I. Chrysiogenales 421 Family I. 421 Genus I. Chrysiogenes 422

PHYLUM BVI 427 Class I. "Chloroflexi" 427 Order I. "Chloroflexales" 427 Family I. "Chloroflexaceae" 427 Filamentous Anoxygenic Phototrophic Bacteria 427 Genus I. Chloroflexus 429 Genus II. Chloronema 437 Genus III. Heliothrix 438 Genus IV. Oscillochloris 440 Order II. "Herpetosiphonales" 444 Family I. "Herpetosiphonaceae" 445 Genus I. Herpetosiphon 445

PHYLUM BVSl Thermomicrobia 447 Class I. Thermomicrobia 447 Order I. Thermomicrobiales 447 Family I. Thermomicrobiaceae 447 Genus I. Thermomicrobium 448

PHYLUM BVISI 451 Class I. "Nitrospira" 451 Order I. "Nitrospirales" 451 Family I. "Nitrospiraceae" 451 Genus I. Nitrospira 451 Genus II. Leptospirillum 453 Genus III. "Candidates Magnetobacterium" 457 Genus IV. Thermodesulfovibrio 460 CONTENTS xvii

PHYLUM BIX Deferribacteres 465 Class I. Deferribacteres 465 Order I. Deferribacterales 465 Family I. Deferribacteraceae 465 Genus I. Deferribacter 466 Genus II. Flexistipes 468 Genus III. Geovibrio 468 Genus Incertae Sedis I. Synergistes 470

PHYLUM BX 473 Oxygenic Photosynthetic Bacteria 473 Oxygenic Photosynthetic Bacteria 474 General Characteristics of the Cyanobacteria 474 Phylogenetic Relationships Among the Cyanobacteria Based on 16S rRNA Sequences 487 Subsection 1 493 Form-genus I. Chamaesiphon 495 Form-genus II. Chroococcus 496 Form-genus 111. Cyanobacterium 497 Form-genus IV. Cyanobium 498 Form-genus V. Cyanothece 499 Form-genus VI. Dactylococcopsis 501 Form-genus VII. Gloeobacter 502 Form-genus VIII. Gloeocapsa 503 Form-genus IX. Gloeothece 504 Form-genus X. Microcystis 505 Form-genus XI. Prochlorococcus 506 Form-genus XII. Prochloron 507 Form-genus XIII. Synechococcus 508 Form-genus XIV. Synechocystis 512 Subsection II 514 Genera Reproducing by Multiple Fissions Only, or in Combination with Limited (1-3) Binary Fissions 520 Form-genus I. Cyanocystis 520 Form-genus II. Dermocarpella 520 Form-genus III. Stanieria 523 Form-genus IV. Xenococcus 524 Genera in Which Extensive Vegetative Binary Fission Precedes Multiple Fission 528 Form-genus I. Chroococcidiopsis 528 Form-genus II. Myxosarciria 531 Pleurocapsa-group 533 Subsection III 539 Form-genus I. Arthrospira 542 Form-genus II. Borzia 543 Form-genus III. Crinalium 543 Form-genus IV. Geitlerinemia 544 Form-genus V. Leptolyngbya 544 Form-genus VI. Limnothrix 546 Form-genus VII. Lyngbya 547 Form-genus VIII. Microcoleus 548 Form-genus IX. Oscillatoria 550 Form-genus X. Planktothrix 553 Form-genus XI. Prochlorothrix 554 Form-genus XII. Pseudanabaena 554 xviii CONTENTS

Form-genus XIII. Spirulina 557 Form-genus XIV. Starria 559 Form-genus XV. Symploca 559 Form-genus XVI. Trichodesmium 560 Form-genus XVII. Tychonema 561 Subsection IV. 562 Subsection I V.I 565 Form-genus I. Anabaena 566 Form-genus II. Anabaenopsis 568 Form-genus III. Aphanizomenon 569 Form-genus IV. Cyanospira 570 Form-genus V. Cylindrospermopsis 571 Form-genus VI. Cylindrospermum 572 Form-genus VII. Nodularia 574 Form-genus VIII. Nostoc 575 Form-genus IX. Scytonema 580 Subsection I V.I I 582 Form-genus I. Calothrix 582

Form-genus II. Rivularia , 586 Form-genus III. Tolypothrix 587 Subsection V. 589 Form-genus I. Chlorogloeopsis 591 Form-genus II. Fischerella 593 Form-genus III. Geitleria 595 Form-genus IV. lyengariella 598 Form-genus V. Nostochopsis 598 Form-genus VI. Stigonema 599 PHYLUM BXi Chlorobi 601 Class I. "Chlorobia" 601 Subclass 1 601 Order I. "Chlorobiales" 601 Family I. "Chlorobiaceae" 601 Green Sulfur Bacteria 601 Genus I. Chlorobium 605 Genus II. Ancalochloris 610 Genus III. Chloroherpeton 612 Genus IV. Pelodictyon 614 Genus V. Prosthecochloris 617 Addendum to the Green Sulfur Bacteria: Phototrophic Green Sulfur Bacteria Living in Consortia with Other Microorganisms 620 PHYLUM BXEIB 625 Class I. "Clostridia" 625 Order I. "Clostridiales" 625 Family VI. "Heliobacteriaceae" 625 Genus I. Heliobacterium 626 Genus II. Heliobacillus 629 Genus 111. Heliophilum 629 Genus IV. Heliorestis 630 The Anoxygenic Phototrophic Purple Bacteria 631 Bibliography 639 Index of Scientific Names of Archaea and Bacteria 703 BERGEY'S MANUAL® OF Systematic Bacteriology Second Edition

Volume Two The

Part A Introductory Essays

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 Schlelfer

WITH CONTRIBUTIONS FROM 339 COLLEAGUES

^ Springer Contents

Preface to Volume Two of the Second Edition of Bergey's Manual® of Systematic Bacteriology sx Preface to the First Edition of Bergey's Manual® of Systematic Bacteriology xi Preface to the First Edition of Bergey's Manual® of Determinative Bacteriology xisi Contributors xvii The History of Bergey's Manual 1

On Using the Manual . 15 Procaryotic Domains 21 Classification of Procaryotic Organisms and the Concept of Bacterial Speciation 27 Identification of Procaryotes 33 Numerical Taxonomy 39 Polyphasic Taxonomy 43 Overview: A Phylogenetic Backbone and Taxonomic Framework for Procaryotic Systematics 49 Nucleic Acid Probes and Their Application in Environmental Microbiology 67 Bacterial Nomenclature 83 Etymology of Nomenclature of Procaryotes 89 Microbial Ecology—New Directions, New Importance 101 Culture Collections: An Essential Resource for Microbiology 111 Intellectual Property of Procaryotes 115 Anoxygenic Phototrophic Purple Bacteria 119 Aerobic Bacteria Containing Bacteriochlorophyll and Belonging to the Alphaproteobacteria 133 Nitrifying Bacteria 137 The Lithoautotrophic Ammonia-Oxidizing Bacteria 141 The Lithoautotrophic Nitrite-Oxidizing Bacteria 149 Bacteria that Respire Oxyanions of Chlorine 155 The Revised Road Map to the Manual 159 Bibliography 221 Index 281

XV 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

Springer Contents

Preface to Volume Two of the Second Edition of Bergey's Manual® of Systematic Bacteriology ix Preface to the First Edition of Bergey's Manual® of Systematic Bacteriology xi Preface to the First Edition of Bergey's Manual® of Determinative Bacteriology xiii Contributors xix

PHYLUM XIV. Proteobacteria 1 Class III. Gammaproteobacteria 1 Order I. Chromatiales 1 Family I. Chromatiaceae 3 Genus I. Chromatium 10 Genus II. Allochromatium 12 Genus III. Halochromatium 14 Genus IV. Isochromatium 15 Genus V. Lamprobacter 16 Genus VI. Lamprocystis 18 Genus VII. Marichromatium 20 Genus VIII. Nitrosococcus 21 Genus IX. Pfennigia 22 Genus X. Rhabdochromatium 23 Genus XI. Thermochromatium 24 Genus XII. Thioalkalicoccus 25 Genus XIII. Thiocapsa 26 Genus XIV. Thiococcus 28 Genus XV. Thiocystis 29 Genus XVI. Thiodictyon 31 Genus XVII. Thioflavicoccus 33 Genus XVIII. Thiohalocapsa 34 Genus XIX. Thiolamprovum 35 Genus XX. Thiopedia 36 Genus XXI. Thiorhodococcus 37 Genus XXII. Thiorhodovibrio 38 Genus XXIII. Thiospirillum 39 Family II. Ectothiorhodospiraceae 41 Genus I. Ectothiorhodospira 43 Genus II. Arhodomonas 48 Genus III. Halorhodospira 49 Genus IV. Nitrococcus 52 Genus V. Thioalkalivibrio 56 Genus VI. Thiorhodospira 57 Family III. Halothiobacillaceae 58 Genus I. Halothiobacillus 58 xvi CONTENTS

Order II. Acidithiobacillales 60 Family I. Acidithiobacillaceae 60 Genus I. Acidithiobacillus 60 Family II. Thermithiobacillaceae 62 Genus I. Thermithiobacillus 62 Order III. Xanthomonadales 63 Family I. Xanthomonadaceae 63 Genus I. Xanthomonas 63 Genus II. Frateuria 91 Genus III. Luteimonas 93 Genus IV. Lysobacter 95 Genus V. Nevskia 101 Genus VI. Pseudoxanthomonas 105 Genus VII. Rhodanobacter 106 Genus VIII. Schineria 106 Genus IX. Stenotrophomonas 107 Genus X. Thermomonas 116 Genus XI. Xylella 119 Order IV. Cardiobacteriales 123 Family I. Cardiobacteriaceae 123 Genus I. Cardiobacterium 123 Genus II. Dichelobacter 126 Genus III. Suttonella 130 Order V. Thiotrichales 131 Family I. Thiotrichaceae 131 Genus I. Thiothrix 131 Genus II. Achromatium 142 Genus III. Beggiatoa 148 Genus IV. Leucothrix 162 Genus V. Thiobacterium 169 Genus VI. Thiomargarita 169 Genus VII. Thioploca 171 Genus VIII. Thiospira 178 Family II. Piscirickettsiaceae 180 Genus I. Piscirickettsia 180 Genus II. Cycloclasticus 184 Genus III. Hydrogenovibrio 188 Genus IV. Methylophaga 190 Genus V. Thioalkalimicrobium 193 Genus VI. Thiomicrospira 193 Family III. Francisellaceae 199 Genus I. Francisella 200 Order VI. Legionellales 210 Family I. Legionellaceae 210 Genus I. Legionella 212 Family II. Coxiellaceae 237 Genus I. Coxiella 237 Genus II. Rickettsiella 241 Order VII. Methylococcales 248 Family I. Methylococcaceae 252 Genus I. Methylococcus 254 Genus II. Methylobacter 258 Genus Hi. Methylocaldum 261 Genus IV. Methylomicrobium 262 Genus V. Methylomonas 265 Genus VI. Methylosarcina 268 Genus VII. Methylosphaera 269 CONTENTS xvii

Order VIII. Oceanospirillales 270 Family I. Oceanospirillaceae 271 Genus I. Oceanospirillum 271 Genus II. Baineatrix 282 Genus III. Marinomonas 284 Genus IV. Marinospirillum 290 Genus V. Neptunomonas 292 Family II. Alcanivoraceae 295 Genus I. Alcanivorax 295 Family III. Hahellaceae 299 Genus I. Hahella 300 Family IV. Halomonadaceae 300 Genus I. Halomonas 300 Genus II. Carnimonas 313 Genus III. Chromohalobacter 316 Genus IV. Zymobacter 319 Order IX. Pseudomonadales 323 Family I. Pseudomonadaceae 323 Genus I. Pseudomonas 323 Genus II. Azomonas 379 Genus III. Azotobacter 384 Genus IV. Cellvibrio 402 Genus V. Mesophilobacter 403 Genus VI. Rhizobacter 404 Genus VII. Rugamorias 407 408 Genus VIII. Serpens 411 Fam i ly 11. Moraxellacea 4^7 Genus I. Moraxella 425 Genus II. Acinetobacter 437 Genus III. Psychrobacter 441 Family III. Incertae Sedis 441 Genus I. Enhydrobacter 443 Order X. Alteromonadales 443 Family I. Alteromonadaceae 444 Genus I. Alteromonas 447 Genus II. Alishewanella 447 Genus III. Colwellia 454 Genus IV. Ferrimonas 456 Genus V. Glaciecola 458 Genus VI. Idiomarina 459 Genus VII. Marinobacter 464 Genus VIII. Marinobacterium 464 Genus IX. Microbulbifer 465 Genus X. Moritella 467 Genus XI. Pseudoalteromonas 478 Genus XII. Psychromonas 480 Genus XIII. Shewanella 491 Order XI. "Vibrionales" 491 Family I. Vibrionaceae 494 Genus I. Vibrio 546 Genus II. Photobacterium 552 Genus III. Salinivibrio 556 Order XII. Aeromonadales 556 Family I. Aeromonadaceae 557 Genus I. Aeromonas 578 Genus II. Oceanimonas 579 Genus Incertae Sedis III. Tolumonas xviii CONTENTS

Family II. Succinivibrionaceae 581 Genus I. Succinivibrio 581 Genus II. Anaerobiospirillum 582 Genus III. Ruminobacter 584 Genus IV. Succinimonas 586 Order XIII. "Enterobacteriales" 587 Family I. Enterobacteriaceae 587 Genus I. Escherichia 607 Genus II. Alterococcus 625 Genus III. Arsenophonus 626 Genus IV. Brenneria 628 Genus V. Buchnera 633 Genus VI. Budvicia 639 Genus VII. Buttiauxella 641 Genus VIII. Calymmatobacterium 645 Genus IX. Cedecea 648 Genus X. Citrobacter 651 Genus XI. Edwardsiella 657 Genus XII. Enterobacter 661 Genus XIII. Erwinia 670 Genus XIV. Ewingella 679 Genus XV. Hafnia 681 Genus XVI. Klebsiella 685 Genus XVII. Kluyvera 694 Genus XVIII. Leclercia 698 Genus XIX. Leminorella 702 Genus XX. Moellerella 705 Genus XXI. Morganella 707 Genus XXII. Obesumbacterium 710 Genus XXIII. Pantoea 713 Genus XXIV. Pectobacterium 721 Genus XXV. "Candidatus Phlomobacter" 730 Genus XXVI. Photorhabdus 732 Genus XXVII. Plesiomonas 740 Genus XXVIII. Pragia 744 Genus XXIX. Proteus 745 Genus XXX. Providencia 753 Genus XXXI. Rahnella 759 Genus XXXII. Saccharobacter 762 Genus XXXIII. Salmonella 764 Genus XXXIV. Serratia 799 Genus XXXV. Shigella 811 Genus XXXVI. Sodalis 823 Genus XXXVII. Tatumella 825 Genus XXXVIII. Trabulsiella 827 Genus XXXIX. Wigglesworthia 828 Genus XL. Xenorhabdus 831 Genus XLI. Yersinia 838 Genus XLI I. Yokenella 848 Order XIV. Pasteurellales 850 Family I. Pasteurellaceae 851 Genus I. Pasteurella 857 Genus II. Actinobacillus 866 Genus III. Haemophilus 883 Genus IV. Lonepinella 904 Genus V. Mannheimia 907 Genus VI. Phocoenobacter 912 Bibliography 913 Index "I085 BERGEY'S MANUAL® OF Systematic Bacteriology Second Edition

Volume Two The Proteobacteria

Part C The Alpha-, Beta-, Delta-, and Epsilonproteobacteria

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 222 COLLEAGUES

<£) Springer Contents

Preface to Volume Two of the Second Edition of Bergey's Manual of Systematic Bacteriology sx Preface to the First Edition of Bergey's Manual of Systematic Bacteriology xi Preface to the First Edition of Bergey's Manual® of Determinative Bacteriology xiii Contributors xxiii Class I. Alphaproteobacteria 1 Order I. Rhodospirillales 1 Family I. Rhodospirillaceae 1 Genus I. Rhodospirillum 1 Genus II. Azospirillum 7 Genus III. Levispirillum 27 Genus IV. Magnetospirillum 28 Genus V. Phaeospirillum 32 Genus VI. Rhodocista 33 Genus VII. Rhodospira 35 Genus VIII. Rhodovibrio 36 Genus IX. Roseospira 37 Genus X. Roseospirillum 39 Genus XI. Skermanella 39 Genus Incertae Sedis XII. "Sporospirillum" 40 Family II. Acetobacteraceae 41 Genus I. Acetobacter 51 Genus II. Acidiphilium 54 Genus III. Acidisphaera 62 Genus IV. Acidocella 65 Genus V. Acidomonas 68 Genus VI. Asaia 69 Genus VII. Craurococcus 70 Genus VIII. Gluconacetobacter 72 Genus IX. Gluconobacter 77 Genus X. Paracraurococcus 81 Genus XI. Rhodopila 83 Genus XII. Roseococcus 85 Genus XIII. Roseomonas 88 Genus XIV. Stella 93 Genus XV. Zavarzinia 95 Order II. Rickettsiales 96 Family I. Rickettsiaceae 96 Genus I. Rickettsia 96 Genus II. Orientia 114 Family II. Ariaplasmataceae 117 Genus I. Anaplasma 117 Genus II. Ehrlichia 125 xvi CONTENTS

Genus III. Neorickettsia 132 Genus IV. Wolbachia 138 Genus Incertae Sedis V. Aegyptianella 143 Family III. Holosporaceae 146 Genus I. Holospora 149 Genus Incertae Sedis II. Caedibacter 152 Genus Incertae Sedis III. Lyticum 156 Genus Incertae Sedis IV. Candidatus Odyssella 157 Genus Incertae Sedis V. Candidatus Paracaedibacter 157 Genus Incertae Sedis VI. Pseudocaedibacter 158 Genus Incertae Sedis VII. "Pseudolyticum" 160 Genus Incertae Sedis VIII. Tectibacter 160 Order III. Rhodobacterales 161 Family I. Rhodobacteraceae 161 Genus I. Rhodobacter 161 Genus II. Ahrensia 167 Genus III. Amaricoccus 168 Genus IV. Antarctobacter 172 Genus V. Gemmobacter 174 Genus VI. Hirschia 176 Genus VII. Hyphomonas 179 Genus VIII. Ketogulonicigenium 187 Genus IX. Maricaulis 188 Genus X. Methylarcula 192 Genus XI. Octadecabacter 194 Genus XII. Paracoccus 197 Genus XIII. Rhodobaca 204 Genus XIV. Rhodovulum 205 Genus XV. Roseibium 209 Genus XVI. Roseinatronobacter 209 Genus XVII. Roseivivax 210 Genus XVIII. Roseobacter 212 Genus XIX. Roseovarius 215 Genus XX. Rubrimonas 217 Genus XXI. Ruegeria 218 Genus XXII. Sagittula 219 Genus XXIII. Staleya 221 Genus XXIV. Stappia 223 Genus XXV. Sulfitobacter 224 Genus Incertae Sedis XXVI. Rhodothalassium 228 Order IV. Sphingomonadales 230 Family I. Sphingomonadaceae 233 Genus I. Sphingomonas 234 Genus II. Blastomonas 258 Genus III. "Citromicrobium" 263 Genus IV. Erythrobacter 267 Genus V. Erythromicrobium 268 Genus VI. Erythromonas 274 Genus VII. Porphyrobacter 275 Genus VIII. Sandaracinobacter 279 Genus IX. Zymomonas 282 Order V. Caulobacterales 287 Family I. Caulobacteraceae 287 Genus I. Caulobacter 287 Genus II. Asticcacaulis 303 Genus III. Brevundimonas 308 Genus IV. Phenylobacterium 316 CONTENTS xvii

Order VI. Rhizobiales 324 Family I. Rhizobiaceae 324 Genus I. Rhizobium 325 Genus II. Agrobacterium 340 Genus III. Allorhizobium 345 Genus IV. Carbophilus 346 Genus V. Chelatobacter 347 Genus VI. Ensifer 354 Genus VII. Sinorhizobium 358 Family II. Bartonellaceae 362 Genus I. Bartonella 362 Family III. Brucellaceae 370 Genus I. Brucella 370 Genus II. Mycoplana 386 Genus III. Ochrobactrum 389 Family IV. Phyllobacteriaceae 393 Genus I. Phyllobacterium 394 Genus II. Aminobacter 397 Genus III. Aquamicrobium 399 Genus IV. Defluvibacter 400 Genus V. Candidatus Liberibacter 400 Genus VI. Mesorhizobium 403 Genus VII. Pseudaminobacter 409 Family V. Methylocystaceae 411 Genus I. Methylocystis 413 Genus II. Albibacter 416 Genus III. Methylosinus 417 Genus Incertae Sedis IV. Methylopila 420 Family VI. Beijerinckiaceae 422 Genus I. Beijerinckia 423 Genus II. Chelatococcus 433 Genus III. Methylocella 437 Family VII. Bradyrhizobiaceae 438 Genus I. Bradyrhizobium 438 Genus II. Afipia 443 Genus III. Agromonas 448 Genus IV. Blastobacter 452 Genus V. Bosea 459 Genus VI. Nltrobacter 461 Genus VII. Oligotropha 468 Genus VIII. Rhodoblastus 471 Genus IX. Rhodopseudomonas 473 Family VIII. Hyphomicrobiaceae 476 Genus I. Hyphomicrobium 476 Genus II. Ancalomicrobium 494 Genus III. Ancylobacter 497 Genus IV. Arigulomicrobium 501 Genus V. Aquabacter 504 Genus VI. Azorhizobium 505 Genus VII. Blastochloris 506 Genus VIII. Devosia 509 Genus IX. Dichotomicroblum 513 Genus X. Filomicrobium 518 Genus XI. Gemmiger 520 Genus XII. Labrys 523 Genus XIII. Methylorhabdus 525 Genus XIV. Pedomicrobium 527 xviii CONTENTS

Genus XV. Prosthecomicrobium 538 Genus XVI. Rhodomicrobium 543 Genus XVII. Rhodoplanes 545 Genus XVIII. Seliberia 549 Genus XIX. Starkeya 554 Genus XX. Xanthobacter 555 Family IX. Methylobacteriaceae 567 Genus I. Methylobacterium 567 Family X. Rhodobiaceae 571 Genus I. Rhodobium 571 Class II. Betaproteobacteria 575 Order I. Burkholderiales 575 Family I. Burkholderiaceae 575 Genus I. Burkholderia 575 Genus II. Cupriavidus 600 Genus III. Lautropia 604 Genus IV. Pandoraea 605 Genus V. Paucimonas 607 Genus VI. Polynucleobacter 607 Genus VII. Ralstonia 609 Genus VIII. Thermothrix 620 Family II. Oxalobacteraceae 623 Genus l. Oxalobacter 624 Genus II. Duganella 628 Genus III. Herbaspirillum 629 Genus IV. Janthinobacterium 636 Genus V. Massilia 643 Genus VI. Telluria 643 Family III. Alcaligenaceae 647 Genus I. Alcaligenes 653 Genus II. Achromobacter 658 Genus III. Bordetelia 662 Genus IV. Derxia 671 Genus V. Oligella 674 Genus VI. Pelistega 678 Genus VII. Pigmentiphaga 681 Genus VIII. Sutterella 682 Genus IX. Taylorella 684 Family IV. Comamonadaceae 686 Genus I. Comamonas 689 Genus II. Acidovorax 696 Genus III. Brachymonas 704 Genus IV. Delftia 706 Genus V. Hydrogenophaga 710 Genus VI. Lampropedia 716 Genus VII. Macromonas 721 Genus VIII. Polaromonas 724 Genus IX. Rhodoferax 727 Genus X. Variovorax 732 Genus Incertae Sedis XI. Aquabacterium 735 Genus Incertae Sedis XII. Ideonella 738 Genus Incertae Sedis XIII. Leptothrix 740 Genus Incertae Sedis XIV. Roseateles 746 Genus Incertae Sedis XV. Rubrivivax 749 Genus Incertae Sedis XVI. SphaerotUus 750 Genus Incertae Sedis XVII. Tepidimonas 755 Genus Incertae Sedis XVIII. Thiomonas 757 CONTENTS xix

Genus Incertae Sedis XIX. Xylophilus 759 Order II. Hydrogenophilales 763 Family I. Hydrogenophilaceae 763 Genus I. Hydrogenophilus 763 Genus II. Thiobacillus 764 Order III. Methylophilales 770 Family I. Methylophilaceae 770 Genus I. Methylophi/us 770 Genus II. Methylobacillus 771 Genus III. Methylovorus 773 Order IV. Neisseriales 774 Family I. Neisseriaceae 775 Genus I. Neisseria 777 Genus II. Alysiella 798 Genus III. Aquaspirillum 801 Genus IV. Chromobacterium 824 Genus V. Eikenella 828 Genus VI. Formivibrio 832 Genus VII. lodobacter 833 Genus VIII. Kingella 836 Genus IX. Microvirgula 840 Genus X. Prolinoborus 841 Genus XI. SimonsieHa 844 Genus XII. Vitreoscilla 851 Genus XIII. Vogesella 858 Genus Incertae Sedis XIV. Catenococcus 859 Genus Incertae Sedis XV. Morococcus 861 Order V. Nitrosomonadales 863 Family I. Nitrosomonadaceae 864 Genus I. Nitrosomonas 864 Genus II. Nitrosolobus 868 Genus III. Nitrosospira 868 Genus IV. "Nitrosovibrio" 869 Family II. Spirillaceae 870 Genus I. Spirillum 870 Family III. Gallionellaceae 880 Genus I. Gallionella 880 Order VI. Rhodocyclaies 887 Family I. Rhodocyclaceae 887 Genus I. Rhodocycius 887 Genus II. Azoarcus 890 Genus III. Azonexus 901 Genus IV. Azospira 902 Genus V. Azovibrio 902 Genus VI. Dechloromorias 903 Genus VII. Dechlorosoma 904

G e n u s V111. Ferribacterium 904 Genus IX. Propionibacter 905 Genus X. Propionivibrio 906 Genus XI. Thauera 907 Genus XII. Zoogioea 913 Class IV. Deltaproteobacteria 922 Order I. Desulfurellales 922 Family I. Desulfureliaceae 923 Genus I. Desulfurella 923 Genus II. Hippea 925 Order II. Desulfovibrionales 925 XX CONTENTS

Family I. Desulfovibrionaceae 926 Genus I. Desulfovibrio 926 Genus II. Bilophila 938 Genus III. Lawsonia 940 Family II. Desulfomicrobiaceae 944 Genus I. Desulfomicrobium 944 Family III. Desulfohalobiaceae 948 Genus I. Desulfohalobium 949 Genus II. Desulfonatronovibrio 953 Genus III. Desulfothermus 955 Family IV. Desulfonatronumaceae 956 Genus I. Desulfonatronum 956 Order III. Desulfobacterales 959 Family I. Desulfobacteraceae 959 Genus I. Desulfobacter 961 Genus II. Desulfobacterium 965 Genus III. Desulfobacula 968 Genus IV. Desulfobotulus 970 Genus V. Desulfocella 971 Genus VI. Desulfococcus 972 Genus VII. Desulfofaba 975 Genus VIII. Desulfofrigus 976 Genus IX. Desulfonema 977 Genus X. Desulfosarcina 981 Genus XI. Desulfospira 984 Genus XII. Desulfotignum 987 Family II. Desulfobulbaceae 988 Genus I. Desulfobulbus 988 Genus II. Desulfocapsa 992 Genus III. Desulfofustis 994 Genus IV. Desulforhopalus 995 Genus V. Desulfotalea 997 Family III. Nitrospinaceae 999 Genus I. Nitrospina 999 Order IV. Desulfarcales 1003 Family I. Desulfarculaceae 1003 Genus I. Desulfarculus 1004 Order V. Desulfuromonales 1005 Family I. Desulfuromonaceae 1006 Genus I. Desulfuromonas 1007 Genus II. Desulfuromusa 1010 Genus III. Malonomonas 1012 Genus IV. Pelobacter 1013 Family II. Geobacteraceae 1017 Genus I. Geobacter 1017 Genus II. Trichlorobacter 1020 Order VI. Syntrophobacterales 1021 Family I. Syntrophobacteraceae 1021 Genus I. Syntrophobacter 1021 Genus II. Desulfacinum 1027 Genus III. Desulforhabdus 1029 Genus IV. Desulfovirga 1030 Genus V. Thermodesulforhabdus 1031 Family II. Syntrophaceae 1033 Genus I. Syntrophus 1033 Genus II. Desulfobacca 1035 Genus III. Desulfomoriile 1036 CONTENTS xxi

Genus IV. Smithella 1039 Order VII. Bdellovibrionales 1040 Family I. Bdellovibrionaceae 1040 Genus I. Bdellovibrio 1041 Genus II. Bacteriovorax 1053 Genus III. Micavibrio 1057 Genus IV. Vampirovibrio 1058 Order V111. Myxococcales 1059 Family I. Myxococcaceae 1073 Genus I. Myxococcus 1074 Genus II. Corallococcus 1079 Genus III. Pyxicoccus 1083 Family II. Cystobacteraceae 1085 Genus I. Cystobacter 1086 Genus II. Archangium 1097 Genus III. Hyalangium 1099 Genus IV. Melittangium 1101 Genus V. Stigmatella 1104 Family III. Polyangiaceae 1109 Genus I. Polyangium 1110 G en us 11. Byssophaga 1118 Genus III. Chondromyces 1121 Genus IV. Haploangium 1129 Genus V. Jahnia 1130 Genus VI. Sorangium 1132 Family IV. Nannocystaceae 1136 Genus I. Nannocystis 1137 Family V. Kofleriaceae 1143 Genus I. Kofleria 1143 Class V. Epsilonproteobacteria 1145 Order I. Campylobacterales 1145 Family I. Campylobacteraceae 1145 Genus I. Campylobacter 1147 Genus II. Arcobacter 1161 Genus III. Sulfurospirillum 1165 Family II. Helicobacteraceae 1168 Genus I. Helicobacter 1169 Genus II. Thiovulum 1189 Genus III. Woliriella 1191 Bibliography 1195 Index 1363 BERGEY'S MANUAL OF Systematic Bacteriology Second Edition

Volume Three The Firmicutes

Paul De Vos, George M. Garrity, Dorothy Jones, Noel R. Krieg, Wolfgang Ludwig, Fred A. Rainey, Karl-Heinz Schleifer and William B.Whitman EDITORS, VOLUME THREE

William B.Whitman DIRECTOR OF THE EDITORIAL OFFICE

Aidan C. Parte MANAGING EDITOR

EDITORIAL BOARD Michael Goodfellow, Chairman, Peter Kampfer, Vice Chairman, Paul De Vos, Fred A. Rainey, Karl-Heinz Schleifer and William B.Whitman WITH CONTRIBUTIONS FROM 165 COLLEAGUES

Springer Contents

Preface to Volume Three of the Second Edition of Bergey's Manual of Systematic Bacteriology ix

Contributors xvis

On using the Manual xxv Revised road map to the phylum Firmicutes 1 Taxonomic outline of the phylum Firmicutes 15

Phylum XIII Firmicutes 19 Class I 19 Order I Bacillales 20 Family I Bacillaceae 20 Genus i Bacillus 21 Genus II Alkalibacillus 128 Genus III Amphibacillus 130 Genus IV Anoxybacillus 134 Genus V Cetasibacillus 141 Genus VI Filobacillus 142 Genus VII Geobacillus 144 Genus VIII Gracihbacillus 160 Genus IX Halobacillus 164 Genus X Halolactibacillus 168 Genus XI Lentibacillus 175 Genus XII Marinococcus 178 Genus XIII Oceanobacillus 181 Genus XIV Paraliobacillus 185 Genus XV Pontibacillus 189 Genus XVI Saccharococcus 190 Genus XVII Tenuibacillus 191 Genus XVIII Thalassobacillus 193 Genus XIX Virgibacillus 193 Family II Alicyclobacillaceae 229 Genus I Ahcyclobacillus 229 Family III Listeriaceae 244 Genus I Listeria 244 Genus II Brochothrix 257 Family IV 'Paenibacillaceae' 269 Genus I Paembacillus 269 Genus II Ammoniphilus 296 Genus III Aneurinibacillus 298 Genus IV Brevibacillus 305 Genus V Cohnella 316 Genus VI Oxalophagus 320 Genus VII Thermobacillus 321

XI XII CONTENTS

Family V Pasteunaceae 328 Genus 1 Pasteuria 328 Family VI Planococcaceae 348 Genus 1 Planococcus 348 Genus II Caryophanon 354 Genus III Filibacter 359 Genus IV Jeotgalibacillus 364 Genus V Kurthia 364 Genus VI Marinibacillus 370 Genus VII Planomicrobium 373 Genus VIII Sporosarcina 377 Genus IX Ureibacillus 381 Family VII "Sporolactobacillaceae" 386 Genus 1 Sporolactobacillus 386

' Family VIII Staphylococcaceae" 392 Genus 1 Staphylococcus 392 Genus II Jeotigalicoccus 421 Genus III Macrococcus 422 Genus IV Sahmcoccus 426 Family IX "Thermoactiriomycetaceae 434 Genus 1 Thermoactinomyces 443 Genus II Laceyella 444 Genus III Mechercharimyces 445 Genus IV Planifilum 446 Genus V Seinonella 447 Genus VI Shimazuella 448 Genus VII Thermoflavimicrobium 449 Family X Incertae Sedis 454 Genus 1 Thermicanus 454 Family XI Incertae Sedis 455 Genus 1 Gemella 455 Family XII Incertae Sedis 460 Genus 1 Exiguobacterium 460 Order II "Lactobacillales" 464 Family 1 Lactobacillaceae 465 Genus 1 Lactobacillus 465 Genus II Paralactobacillus 511 Genus III Pediococcus 513 Family II "Aerococcaceae" 533 Genus 1 Aerococcus 533 Genus II Abiotrophia 536 Genus III Dolosicoccus 538 Genus IV Eremococcus 540 Genus V Facklamia 541 Genus VI Globicatella 544 Genus VII Ignavigranum 546 Family III "Carnobacteriaceae" 549 Genus 1 Carnobacterium 549 Genus II Alkahbacterium 557 Genus III Allofustis 559 Genus IV Alloiococcus 562 Genus V Atopobacter 563 Genus VI Atopococcus 565 Genus VII Atopostipes 566 Genus VIII Desemzia 568 CONTENTS xill

Genus IX. Dolosigrariulum 572 Genus X. Granulicatella 574 Genus XI. Isobaculum 579 Genus XII. Marirtilactibacillus 530 Genus XIII. Trichococcus 535 IV. Family "Enterococcaceae" 594 Genus I. Enterococcus 594 Genus II. Melissococcus 607 Genus III. Tetragenococcus 611 Genus IV. Vagococcus 61 g V. Family "Leuconostocaceae" 624 Genus I. Leuconostoc 624 Genus II. Oenococcus 635 Genus III. Weissella 643 VI. Family 655 Genus I. Streptococcus 655 Genus II. Lactococcus 71-j Genus III. Lactovum 722 Class II. "Clostridia" 736 Order I. Clostridiales 730 I. Family Clostridiaceae 735 Genus I. Clostridium 738 II. Genus Alkaliphilus 828 Genus III. Anaerobacter 830 Genus IV. Anoxynatronum 831 Genus V. Caloramator 834 Genus VI. Caloranaerobacter 839 Genus VII. Caminicella 840 GenusVIII. Natronirico/a 841 Genus IX. Oxobacter 842 Genus X. Sarcina 843 Genus XI. Thermobrachium 848 Genus XII. Thermohalobacter 848 Genus XIII. Tindallia 851 Family II. "Eubacteriaceae" 865 Genus I. Eubacterium 865 Genus II. Acetobacterium 891 Genus III. Alkalibacter 896 Genus IV. Anaerofustis 897 Genus V. Garciella 900 Genus VI. Pseudoramibacter. 902 Family III. "Gracilibacteraceae" 910 Genus I. Gracilibacter 910 Family IV. "Heliobacteriaceae" 913 Genus I. Heliobacterium 916 Genus II. Heliobacillus 918 Genus III. Heliophilum 918 Genus IV. Heliorestis 919 Family V. "Lachnospiraceae" 921 Genus I. Lachnospira 921 Genus II. Acetitomaculum 923 Genus III. Anaerostipes 924 Genus IV. Bryantella 925 Genus V. Butyrivibrio 927 Genus VI. Catonella 937 XIV CONTENTS

Genus VII. Coprococcus 940 Genus VIII. Dorea 941 Genus IX. Hespellia 944 Genus X. Johnsonella 945 Genus XI. Lachnobacterium 946 Genus XII. Moryella 948 Genus XIII. Oribacterium 950 Genus XIV. Parasporobacterium 952 Genus XV. Pseudobutyrivibrio 952 Genus XVI. Roseburia 954 Genus XVII. Shuttleworthia 957 Genus XVIII. Sporobacterium 959 Genus XIX. Syntrophococcus 961 Family VI. Peptococcaceae 969 Genus I. Peptococcus 971 Genus II. Cryptanaerobacter 971 Genus ill. Dehalobacter 974 Genus IV. Desulfitobacterium 975 Genus V. Desulfonispora 982 Genus VI. Desulfosporosinus 983 Genus VII. Desulfotomaculum 989 Genus VIII. Pelotomaculum 996 Genus IX. Sporotomaculum 999 Genus X. Syritrophobotulus 1000 Genus XI. Thermincola 1001 Family VII. "Peptostreptococcaceae" 1008 Genus I. Peptostreptococcus 1008 Genus II. Filifactor 1009 Genus III. Tepidibacter 1013 Family VIII. "Ruminococcaceae" 1016 Genus I. Ruminococcus 1016 Genus II. Acetanaerobacterium 1019 Genus III. Acetivibrio 1020 Genus IV. Anaerofilum 1022 Genus V. Anaerotruncus 1023 Genus VI. Faecalibacterium 1026 Genus VII. Fastidiosipila 1028 Genus VIII. Oscillospira 1031 Genus IX. Papillibacter 1033 Genus X. Sporobacter 1034 Genus XI. Subdoligranulum 1037 Family IX. Syntrophomonadaceae 1044 Genus I. Syrttrophomonas 1045 Genus II. Pelospora 1051 Genus III. Syntrophospora 1052 Genus IV. Syntrophothermus 1053 Genus V. Thermosyntropha 1055 Family X. Veillonellaceae 1059 Genus I. Veillonella 1059 Genus II. Acetonema 1065 Genus III. Acidaminococcus 1067 Genus IV. Allisonella 1068 Genus V. Anaeroarcus 1069 Genus VI. Anaeroglobus 1071 Genus VII. Anaeromusa 1073 CONTENTS XV

GenusVIII Anaerosinus 1074 Genus IX Anaerovibrio 1075 Genus X Centipeda 1077 Genus XI Dendrosporobacter 1079 Genus XII Dialister 1080 Genus XIII Megasphaera 1082 Genus XIV Mitsuokella 1090 Genus XV Pectinatus 1094 Genus XVI Phascolarctobacterium 1100 Genus XVII Propionispira 1102 Genus XVIII Propionispora 1103 Genus XIX Quinella 1104 Genus XX Schwartzia 1105 Genus XXI Selenomonas 1106 Genus XXII Sporomusa 1112 Genus XXIII Succiniclasticum 1116 Genus XXIV Succinispira 1117 Genus XXV Thermosinus 1118 Genus XXVI Zymophilus 1119 Family XI Incertae Sedis 1130 Genus I Anaerococcus 1130 Genus II Finegoldia 1131 Genus III Galhcola 1132 Genus IV Helcococcus 1132 Genus V Parvimonas 1135 Genus VI Peptoniphilus 1136 Genus VII Sedimentibacter 1137 Genus VIII Soehngema 1141 Genus IX Sporanaerobacter 1143 Genus X Tissierella 1146 Family XII Incertae Sedis 1150 Genus I Acidaminobacter 1150 Genus II Fusibacter 1151 Genus III Guggenheimella 1154 Family XIII Incertae Sedis 1156 Genus I Anaerovorax 1156 Genus II Mogibactetium 1157 Family XIV Incertae Sedis 1161 Genus I Anaerobranca 1161 Family XV Incertae Sedis 1165 Genus I Ammobacterium 1165 Genus II Aminomonas 1167 Genus III Anaerobaculum 1170 Genus IV Dethiosulfovibrio 1174 Genus V Thermanaerovibrio 1176 Family XVI Incertae Sedis 1180 Genus I Carboxydocella 1180 Family XVII Incertae Sedis 1181 Genus I Sulfobacillus 1181 Genus II Thermaerobacter 1184 Family XVIII Incertae Sedis 1188 Genus I Symbiobacterium 1188 Family XIX Incertae Sedis 1190 Genus I Acetoanaerobium 1190 Order II Halanaerobiales 1191 XVI CONTENTS

Family I. Halanaerobiaceae 1195 Genus I. Halanaerobium 1196 Genus II. Halocella 1201 Genus III. Halothermothrix 1202 Family II. Halobacteroidaceae 1207 Genus I. Halobacteroides 1208 Genus II. Acetohalobium 1209 Genus III. Halanaerobacter 1212 Genus IV. Halonatronum 1215 Genus V. Natrortiella 1216 Genus VI. Orenia 1217 Genus VII. Selenihalanaerobacter 1221 Genus VIII. Sporohalobacter 1222 Order III. Thermoanaerobacterales 1224 Family I. Thermoanaerobacteraceae 1225 Genus I. Thermoanaerobacter 1225 Genus II. Ammonifex 1240 Genus III. Caldanaerobacter 1241 Genus IV. Carboxydothermus 1244 Genus V. Gelria 1246 Genus VI. Moorella 1247 Genus VII. Thermacetogenium 1253 Genus VIII. Thermanaeromonas 1256 Family II. Thermodesulfobiaceae 1268 Genus I. Thermodesulfobium 1268 Genus II. Coprothermobacter 1271 Family III. Incertae Sedis 1275 Genus I. Caldicellulosiruptor 1275 Genus II. Thermoanaerobacterium 1279 Genus III. Thermosediminibacter 1287 Genus IV. Thermovenabulum 1290 Family IV. Incertae Sedis 1296 Genus I. Mahella 1296 Class III. Erysipelotrichia 1298 Order I. Erysipelotrichales 1298 Family I. Erysipelotrichaceae 1299 Genus I. Erysipelothrix 1299 Genus II. Allobaculum 1306 Genus III. Bulleidia 1307 Genus IV. Catenibacterium 1309 Genus V. Coprobacillus 1310 Genus VI. Holdemania 1310 Genus VII. Solobacterium 1312 Genus VIII. Turicibacter 1314

Appendix 1. Validly published names, conserved and rejected names, and taxonomic opinions cited in the International Journal of Systematic and Evolutionary Microbiology since publication of Volume 2 of the Second Edition of the Systematics 1319 BERGEY'S MANUAL OF Systematic Bacteriology Second Edition

Volume Four The , , Tenericutes (Mollicutes), , , , Dictyoglomi, , Lentisphaerae, , , and

Noel R. Krieg, James T. Staley, Daniel R. Brown, Brian P. Hedlund, Bruce J. Paster, Naomi L.Ward, Wolfgang Ludwig and William B.Whitman EDITORS, VOLUME FOUR

William B.Whitman DIRECTOR OF THE EDITORIAL OFFICE

Aidan C. Parte MANAGING EDITOR

EDITORIAL BOARD Michael Goodfellow, Chairman, Peter Kampfer, Vice Chairman, Jongsik Chun, Paul De Vos, Fred A. Rainey and William B.Whitman WITH CONTRIBUTIONS FROM 129 COLLEAGUES

^ Springer Contents

Preface to volume 4 ix Contributors xix On using the Manual xxv Road map of volume 4 phyla 1 Taxonomic outline of volume 4 21

Phylum XIV. Bacteroidetes phyl. nov 25 Class I. Bacteroidia class, nov 25

Order I. Bacteroidales ord. nov 25 Family I. Bacteroidaceae 25 Genus I. Bacteroides 27 Genus II. Acetofilamentum 41 Genus III. Acetomicrobium 42 Genus IV. Acetothermus 44 Genus V. Anaerorhabdus 45 Family 11. Marinilabiliaceae fam. nov 49 Genus I. Marinilabilia 49 Genus II. Anaerophaga 51 Genus III. Alkaliflexus 53 Family 111. Rikeriellaceae fam. nov 54 Genus I. Rikenella 55 Genus II. Alistipes 56 Family IV. Porphyromonadaceae fam. nov 61 Genus I. Porphyromonas 62 Genus II. Barnesiella 70 Genus III. Dysgonomonas 71 Genus IV. Paludibacter 76 Genus V. Petrimonas 77 Genus VI. Proteiniphilum 77 Genus VII. Tannerella 78 Family V. Prevotellaceae fam. nov 85 Genus I. Prevotella 86 Genus II. Xylanibacter 102 Class II. Flavobacteriia class, nov 105 Order I. Flavobacteriales ord. nov 105 Family I. Flavobacteriaceae 106 G e n u s I. Fla vobacterium 112 Genus II. Aequorivita 155 Genus III. Algibacter 157 Genus IV. Aquimarina 158 Genus V. Arenibacter 161 Genus VI. Bergeyella 165

XI xii CONTENTS

Genus VII. Bizionia 166 Genus VIII. Capnocytophaga 168 Genus IX. Cellulophaga 176 Genus X. Chryseobacterium 180 Genus XI. Cloacibacterium 197 Genus XII. Coenonia 198 Genus XIII. Costertonia 199 Genus XIV. Croceibacter 199 Genus XV. Dokdonia 201 Genus XVI. Donghaeana 201 Gen us XV11. Elizabethkingia 202 Genus XVIII. Empedobacter 210 Gen us XIX. Epilithonimonas 212 G e n u s XX. Fla viramulus 213 Genus XXI. Formosa 214 Genus XXII. Gaetbulibacter 218 Genus XXIII. Gelidibacter 219 Genus XXIV. GiUisia 221 Genus XXV. Gramella 226 Genus XXVI. Kaistella 227 Genus XXVII. Kordia 228 Genus XXVIII. Krokinobacter 230 Genus XXIX. Lacinutrix 231 Genus XXX. Leeuwenhoekiella 232 Genus XXXI. Lutibacter 234 Genus XXXII. Maribacter 235 Genus XXXI11. Maririiflexile 238 Genus XXXIV. Mesonia 239 Genus XXXV. Muricauda 240 Genus XXXVI. Myroides 245 Genus XXXVII. Nonlabens 248 Genus XXXVIII. Olleya 249 Genus XXXIX. Ornithobacterium 250 Genus XL. Persicivirga 254 Genus XLI. Polaribacter 255 GenusXLII. Psychroflexus 258 Genus XLIII. Psychroserpens 261 Genus XLIV. Riemerella 262 Genus XLV. Robiginitalea 264 Genus XLVI. Salegentibacter 266 Genus XLVI I. Sandarakiriotalea 269 Genus XLVIII. Sediminicola 270 Genus XLIX. Sejongia 271 Genus L. Stenothermobacter 275 Genus LI. Subsaxibacter 275 Genus LI I. Subsaximicrobium 277 Genus LIII. Tenacibaculum 279 Genus LIV. Ulvibacter 283 Genus LV. Vitellibacter 284 Genus LVI. Wautersiella 285 Genus LVI I. Weeksella 286 Genus LVIII. WirtogradskyeUa 288 Genus LIX. Yeosuana 291 292 Genus LX. Zhouia - Genus LXl. Zobellia 292 CONTENTS xiii

Family II. Blattabacteriaceae fam. nov 315 Genus i. Blattabacterium 315 Family III. Cryomorphaceae 322 Genus I. Cryomorpha 323 Genus II. Brumimicrobium 323 Genus III. Crocinitomix 326 Genus IV. Fluviicola 327 Genus V. Lishizhenia 327 Genus VI. Owenweeksia 328 Class III. Sphingobacteriia class, nov 330 Order I. Sphingobacteriales ord. nov 330 Family I. Sphingobacteriaceae 331 Genus I. Sphingobacterium 331 Genus II. Pedobacter 339 Family II. Chitinophagaceaelam. nov 351 Genus I. Chitinophaga 351 Genus II. Terrimonas 356 Family III. Saprosp/Vaceae fam. nov. 358 Genus I. Saprospira 359 Genus II. Aureispira 361 Genus III. Haliscomenobacter 363 Genus IV. Lewinella 366 Class IV. Cytophagia class, nov 370 Order I. Cytophagales 370 Family I. Cytophagaceae 371 Genus I. Cytophaga 371 Genus II. Adhaeribacter 375 Genus III. Arcicella 377 Genus IV. Dyadobacter. 380 Genus V. Effluviibacter 387 Genus VI. Emticicia 388 Genus VII. Flectobacillus 389 Genus VI11. Flexibacter 392 Genus IX. Hymenobacter 397 Genus X. Larkinella 404 Genus XI. Leadbetterella 405 Genus XII. Meniscus 406 Genus XIII. Microscilla 408 Genus XIV. Pontibacter. 410 Genus XV. Runella 412 Genus XVI. Spirosoma 415 Genus XVII. Sporocytophaga 418 Family II. Cyclobacteriaceae fam. nov 423 Genus I. Cyclobacterium 423 Genus II. Algoriphagus 426 Genus III. Aquiflexum 433 Genus IV. Belliella 434 Genus V. Echinicola 437 Genus VI. Rhodonellum 440 Family III. Flammeovirgaceae fam. nov 442 Genus I. Flammeovirga 442 Genus II. Fabibacter 447 Genus III. Flexithrix 448 Genus IV. Persicobacter 450 Genus V. Reichenbachiella 452 xiv CONTENTS

Genus VI. Roseivirga 453 Order II. Incertae sedis 457 Family I. Rhodothermaceae farm, nov 457 Genus I. Rhodothermus 458 Genus II. Salinibacter 460 Order III. Incertae sedis 465 Genus I. Thermonema 465 Order IV. Incertae sedis 467 Genus I. Toxothrix 467

Phylum XV. Spirochaetes phyl. nov 471 Class I. Spirochaetia class, nov 471 Order I. Spirochaetales 471 Family I. Spirochaetaceae 473 Genus I. Spirochaeta 473 Genus II. Borrelia 484 Genus III. Cristispira 498 Genus IV. Treponema 501 Family II. Brachyspiraceae 531 Genus I. Brachyspira 531 Family III. Brevinemataceaefam. nov. 545 Genus I. Brevinema 545 Family IV. Leptospiraceae 546 Genus I. Leptospira 546 Genus II. Leptonema 556 Genus III. Turneriella 558 Hindgut spirochetes of termites and Cryptocercus punctulatus 563

Phylum XVI. Tenericutes 567 Class I. Mollicutes 568 Order I. Mycoplasmatales 574 Family i. Mycoplasmataceae 575 Genus I. Mycoplasma 575 Genus II. Ureaplasma 613 Family II. Incertae sedis 639 Genus I. Eperythrozoon 640 Genus II. Haemobartonella 642 Order II. Entomoplasmatales 644 Family I. Entomoplasmataceae 645 Genus I. Entomoplasma 646 Genus II. Mesoplasma 649 Family II. Spiroplasmataceae 654 Genus I. Spiroplasma 654 Order 111. Acholeplasmatales 687 Family I. Acholeplasmataceae 687 Genus I. Acholeplasma 688 Family II. Incertae sedis 696 Genus I. "Candidatus Phytoplasma" gen. nov. 696 Order IV. Anaeroplasmatales 719 Family I. Anaeroplasmataceae 720 Genus I. Anaeroplasma 720 Gen us 11. Asteroleplasma 722 CONTENTS xv

Phylum XVII. Acidobacteria phyl. nov 725 Class I. Acidobacteriia 727 Order I. Acidobacteriales 727 Family I. Acidobacteriaceae fam. nov 728 Genus I. Acidobacterium 728 Genus II. Edaphobacter 729 Genus 111, Terriglobus 730 Class II. Holophagae 731 Order I. Holophagales 731 Family I. Holophagaceae 732 Genus I. Holophaga 732 Genus II. Geothrix 732 Order 11. Acanthopleuribacterales 734 Family I. Acanthopleuribacteraceae 734 Genus I. Acanthopleuribacter 734

Phylum XVIII. "Fibrobactereg' 737

Class I. Fibrobacteria class, nov. 739

Order I. Fibrobacterales ord. nov 739 Family I. Fibrobacteraceae fam. nov. 739 Genus I. Fibrobacter 740

Phylum XIX."Fusobacteria" 747 Class I. Fusobacteriia class, nov 747

Order I. Fusobacteriales ord. nov 747 Family I. Fusobacteriaceae fam. nov 748 Genus I. Fusobacterium 748 Genus II. Cetobacterium 758 Genus III. Ilyobacter 759 Genus IV. Propionigenium 761 Family \\. Leptotrichiaceae fam. nov. 766 Genus I. Leptotrichia 766 Genus II. Sebaldella 769 Genus III. Sneathia 770 Genus IV. Streptobacillus 771

Phylum XX. Dictyoglomi phyl. nov 775 Class I. Dictyoglomia class, nov 776 Order I. Dictyoglomales ord. nov 776 Family I. Dictyoglomaceae fam. nov 776 Genus I. Dictyoglomus 776

Phylum XXI. Gemmatimonadetes 781 Class I. Gemmatimonadetes 781 Order I. Gemmatimoriadales 781 Family I. Gemmatimonadaceae 782 Genus I. Gemmatimonas 782

Phylum XXII. Lentisphaerae 785 Class I. Leritisphaeria class, nov 787 Order I. Lentisphaerales 788 Family I. Lentisphaeraceae fam. nov 788 Genus I. Lentisphaera 788 CONTENTS

Order II. Victivallales 791 Family I. Victivallaceae fam. nov 791 Genus I. Victivallis 791

Phylum XXIII. Verrucomicrobia phyl. nov 795 Class I. Verrucomicrobiae 799 Order I. Verrucomicrobiales 802 Family I. Verrucomicrobiaceae 803 Genus I. Verrucomicrobium 803 Genus II. Prosthecobacter 805 Family II. Akkermansiaceae fam. nov 809 Genus I. Akkermansia 809 Family 111. Rubritaleaceae fam. nov 812 Genus I. Rubritalea 812 Class II. Opitutae 817 Order I. Opitutales 820 Family I. Opitutaceae 820 Genus I. Opitutus 820 Genus II. Alterococcus 821 Order II. Puniceicoccaies 823 Family I. Puniceicoccaceae 824 Genus I. Puniceicoccus 824 Genus II. Cerasicoccus 825 Genus III. Coraliomargarita 827 Genus IV. Pelagicoccus 829 834 Class 111. Spartobacteria class, nov 836 Order I. Chthoniobacterales ord. nov 837 Family I. Chthoniobacteraceae fam. nov Genus I. Chthoriiobacter 837 Genus II. "Candidatus Xiphinematobacter" 838

843 Phylum XXIV. "Chlamydiae" 844 Class I. Chlamydiia class, nov 844 Order I. Chlamydiales Family I. Chlamydiaceae 845 Genus I. Chlamydia 846 865 Family 11."Candidatus Clavichlamydiaceae" Genus I."Candidatus Clavichlamydia" 865 867 Family III. Criblamydiaceae Genus I. Criblamydia 867 867 Family IV. Parachlamydiaceae Genus I. Parachlamydia 868 Genus II. Neochlarnydia 869 870 Genus III. Protochlamydia gen. nov 372 Family V. "Candidatus Piscichlamydiaceae" Genus I. "Candidatus Piscichlamydia" 872 873 Family VI. Rhabdochlamydiaceae fam. nov S73 Genus I. Rhabdochlamydia gen. nov 874 Family VII. Simkaniaceae Genus I. Simkania 875 Genus II."Candidatus Fritschea" 875 87® Family VIII. Waddliaceae Genus I. Waddiia 877 CONTENTS xvii

Phylum XXV."Planctomycete^ 879 879 Class I. Planctomycetia class, nov Order I. Planctomycetales 879 Family I. Planctomycetaceae 880 Genus I. Planctomyces 881 Genus II. Blastopirellula 895 Genus III. Gemmata 897 Genus IV. Isosphaera 900 Genus V. Pirellula 903 906 G e n u s VI. Rhodopirellula Genus VII. Schlesneria 910 Genus VIII. Singulisphaera 913 Order II. "Candidatus Brocadiales" ord. nov 918 Family I. "Candidatus Brocadiaceae" fam. nov 918

Author index 927

Index of scientific names of Archaea and Bacteria 929 BERGEY'S MANUAL OF Systematic Bacteriology Second Edition

Volume Five The

Michael Goodfellow, Peter Kampfer, Hans-Jurgen Busse, Martha E.Trujillo, Ken-ichiro Suzuki, Wolfgang Ludwig and William B.Whitman EDITORS, VOLUME FIVE

William B.Whitman DIRECTOR OF THE EDITORIAL OFFICE

Aidan C. Parte MANAGING EDITOR

EDITORIAL BOARD Fred A. Rainey, Chairman, Peter Kampfer, Vice Chairman, Paul De Vos, Jongsik Chun, Martha E.Trujillo and William B.Whitman WITH CONTRIBUTIONS FROM 116 COLLEAGUES

^ Springer Contents

Preface to volume 5 ix Contributors xvii On using the Manual xxiii

Part A

Road map of the phylum Actinobacteria 1 Taxonomic outline of the phylum Actinobacteria 29 Phylum XXVI. Actinobacteria phyl. nov 33 Class I. Actinobacteria 34 Order I. 35 Family I. Actinomycetaceae 36 Genus I. Actinomyces 42 Genus II. Actinobaculum 109 Genus III. Arcanobacterium 114 Genus IV. Mobiluncus 126 Genus V. Varibaculum 139 Order II. Actinopolysporales ord. nov 162 Family I. Actinopolysporaceae 163 Genus I. Actinopolyspora 163 Order III. Bifidobacteriales 170 Family I. Bifidobacteriaceae 171 Genus I. Bifidobacterium 171 Genus II. Aeriscardovia 206 Genus III. Ailoscardovia 207 Genus IV. Gardnerelia 208 Genus V. Metascardovia 211 Genus VI. Parascardovia 213 Genus VII. Scardovia 214 Order IV. Catenulisporales ord. nov 225 Family I. Catenuiisporaceae 226 Genus I. Catenulispora 226 Family II. Actinospicaceae 232 Genus I. Actinospica 232 Order V. Corynebacteriales ord. nov 235 Family I. Corynebacteriaceae 244 Genus I. Corynebacterium 245 Genus II. Turicella 289 Family II. Dietziaceae 301 Genus I. Dietzia 301 Family III. Mycobacteriaceae 312 Genus I. Mycobacterium 312

xi xii CONTENTS

Family IV. Nocardiaceae 376 Genus I. Nocardia 376 Genus II. Gordonia 419 G en us 111. Millisia 435 Genus IV. Rhodococcus 437 Genus V. Skermania 464 Genus VI. Smaragdicoccus 467 G e n u s V11. Williamsia 470 Family V. Segniliparaceae 497 Genus I. Segniliparus 497 Family VI. Tsukamurellaceae 500 Genus I. Tsukamurella 500 Order VI. Frankiales ord. nov 509 Family I. Frankiaceae 512 Genus I. Frankia 512 Family II. Acidothermaceae 520 Genus I. Acidothermus 520 Fam ily 111. Cryptosporangiaceae 522 Genus I. Cryptosporangium 522 Genus Incertae sedis I. Fodinicola 525 Family IV. Geodermatophilaceae 528 Genus I. Geodermatophilus 528 Genus II. Blastococcus 531 Genus III. Modestobacter 536 Family V. Nakamurellaceae 539 Genus I. Nakamurella 540 Genus II. Humicoccus 542 Family VI. Sporichthyaceae 544 Genus I. Sporichthya 544 Order VII. Glycomycetales ord. nov. 546 Family I. Glycomycetaceae 547 Genus I. Glycomyces 547 Genus II. Stackebrandtia 553 Order VIII. Jiangellales ord. nov 555 Family I. Jiangellaceae 557 Genus I. Jiangella 557 Genus II. Haloactinopolyspora 560 Order IX. Kineosporiales ord. nov 561 Family I. Kineosporiaceae 562 Genus I. Kineosporia 562 Genus II. Kineococcus 565 Genus III. Quadrisphaera 567 Order X. 569 Family I. Micrococcaceae 571 Genus I. Micrococcus 571 Genus II. Acaricomes 577 Genus III. Arthrobacter 578 Genus IV. Citricoccus 624 Genus V. Kocuria 626 Genus VI. Nestererikonia 636 Genus VII. Renibacterium 643 Genus VIII. Rothia 646 Genus IX. Yaniella 650 Genus X. Zhihengliuella 653 CONTENTS xiii

Family II. Beutenbergiaceae 667 Genus !. Beutenbergia 669 Genus II. Miniimonas 671 Genus III. Salana 673 Genus IV. Serinibacter .. 677 Fam i ly 111. Bogoriellaceae 678 Genus I. Bogoriella 679 Genus II. Georgenia 681 Family IV. Brevibacteriaceae 685 Genus I. Brevibacterium 685 Family V. Cellulomonadaceae 701 Genus I. Cellulomonas 702 Genus II. Incertae sedis Actinotalea 710 Genus III. Incertae sedis Demequina 712 Genus IV. Oerskovia 713 Genus V. Tropheryma 717 Family VI. Dermabacteraceae 727 Genus I. Dermabacter 727 Genus II. Brachybacterium 730 Family VII. Dermacoccaceae 738 Genus I. Dermacoccus 738 Genus II. Demetria 742 Genus III. Kytococcus 744 Family VI11. Derma tophilaceae 748 Genus I. Dermatophilus 749 Genus II. Kineosphaera 752 Family IX. Intrasporangiaceae 754 Genus I. Intrasporangium 759 Genus II. Arsenicicoccus 761 Genus III. Humihabitans 764 Genus IV. Janibacter 765 Genus V. Knoellia 769 Genus VI. Kribbia 773 Genus VII. Lapillicoccus 774 Genus VIII. Ornithinicoccus 775 Genus IX. Ornithinimicrobium 777 Genus X. Oryzihumus 781 Genus XI. Phycicoccus 782 Genus XII. Serinicoccus 786 Genus XIII. Terrabacter 788 Genus XIV. Terracoccus 790 Genus XV. Tetrasphaera 792 Family X. Jonesiaceae 802 Genus I. Jonesia 802 Family XI. Microbacteriaceae 807 Genus I. Microbacterium 814 Genus II. Agreia 852 Genus III. Agrococcus 855 Genus IV. Agromyces 862 Genus V. Clavibacter 877 Genus VI. Cryobacterium 883 Genus VII. Curtobacterium 887 Genus VI11. Frigoribacterium 895 Genus IX. Frondihabitans 898 xiv CONTENTS

Genus X. Gulosibacter 900 Genus XI. Humibacter 904 Genus XII. Labedella 906 Genus XIII. Leifsonia 907 Genus XIV. Leucobacter 923 Genus XV. Microcella 933 Genus XVI. Microterricola 934 Genus XVII. Mycetocola 936 Genus XVIII. Okibacterium 940 Genus XIX. Phycicola 942 Genus XX. Plantibacter 943 Genus XXI. Pseudoclavibacter 949 Genus XXII. Rathayibacter 953 Genus XXIII. Rhodoglobus 964 Genus XXIV. Salinibacterium 969 Genus XXV. Subtercola 973 Genus XXVI. Yorighaparkia 975 Family XII. Promicromonosporaceae 995 Genus I. Promicromonospora 995 Genus II. Cellulosimicrobium 1002 Genus III. Isoptericola 1006 Genus IV. Myceligenerans 1010 Genus V. Xylanibacterium 1012 Genus VI. Xylanimicrobium 1015 Genus VII. Xylanimorias 1016 Family XIII. 1019 Genus I. Rarobacter 1020 Family XIV. Ruaniaceae 1022 Genus I. Ruania 1024 Genus II. Haloactinobacterium 1025 Family XV. Sanguibacteraceae 1027 Genus I. Sanguibacter 1030

Part B

Order XI. Micromonosporales ord. nov 1035 Family I. Micromonosporaceae 1035 Genus I. Micromonospora 1039 Genus II. Actinocatenispora 1057 Genus III. Actinoplanes 1058 Genus IV. Asanoa 1088 Genus V. Catellatospora 1090 Genus VI. Catenuloplanes 1092 Genus VII. Couchioplanes 1094 Genus VIII. Dactylosporangium 1096 Genus IX. Krasilnikovia 1106 Genus X. Lorigispora 1107 Genus XI. Luedemannella 1110 Genus XII. Pilimelia 1111 Genus XIII. Polymorphospora 1117 Ge n u s XIV. Salinispora 1118 G en us XV. Spirilliplanes 1123 Genus XVI. Verrucosispora 1124 Genus XVII. Virgisporangium 1127 Order XII. ord. nov 1137 Family I. 1138 Genus I. Propioriibacterium 1138 CONTENTS xv

Genus 11. Aestuariimicrobium 1155 Genus III. Brooklawnia 1156 Genus IV. Friedmanniella 1159 G e n u s V. Granulicoccus 1165 Genus VI. Luteococcus 1166 Genus VII. Microlunatus 1168 Genus VI11. Micropruina 1173 Genus IX. Propionicicella 1174 G e n u s X. Propionicimonas 1175 Genus XI. Propioniferax 1177 G e n u s X11. Propionimicrobium 1179 Genus XIII. Tessaracoccus 1180 Fami ly 11. Nocardioidaceae 1189 Genus I. Nocardioides 1197 Genus II. Actinopolymorpha 1251 Genus III. Aeromicrobium 1258 Genus IV. Kribbella 1268 Genus V. Marmoricola 1284

Order XIII. Pseudonocardiales ord. nov 1301 Family I. Pseudonocardiaceae 1302 Genus I. Pseudonocardia 1305 Genus 11. Actinoatloteichus 1323 Genus III. Actinokineospora 1325 Genus IV. Actinosynnema 1331 Genus V. Amycolatopsis 1334 Genus VI. Crossiella 1359 Genus VII. Goodfellowiella 1363 Genus VIII. Kibdelosporangium 1366 Genus IX. Kutzneria 1371 Genus X. Lechevalieria 1375 Genus XI. Lentzea 1379 Genus XII. Prauserella 1384 Genus XIII. Saccharomonospora 1390 Genus XIV. Saccharopolyspora 1396 Genus XV. Saccharothrix 1415 Genus XVI. Streptoalloteichus 1419 Genus XVII. Thermocrispum 1423 Genus XVIII. Umezawaea 1427 Order XIV. Streptomycetales ord. nov 1446 Family I. Streptomycetaceae 1446 G e n u s I. Streptomyces 1455 Genus Incertae sedis I. Kitasatospora 1768 Genus Incertae sedis II. Streptacidiphilus 1777 Order XV. Streptosporarigiales ord. nov 1805 Family I. Streptosporangiaceae 1807 G e n u s I. Streptosporangium 1811 G e n u s 11. A crocarpospora 1825 Genus III. Herbidospora 1827 Genus IV. Microbispora 1831 Genus V. Microtetraspora 1838 Genus VI. Nonomuraea 1844 Genus VII. Planobispora 1861 Genus VIII. Planomonospora 1866 Genus IX. Planotetraspora 1873 Genus X. Sphaerisporarigium 1875 Genus XI. Thermopolyspora 1880 xvi CONTENTS

Family II. Nocardiopsaceae 1889 Genus I. Nocardiopsis 1891 Genus II. Haloactinospora 1907 Genus III. Streptomonospora 1908 Genus IV. Thermobifida 1914 Family III. Thermomonosporaceae 1929 Genus I. Thermomonospora 1931 Genus II. Actinocorallia 1935 Genus III. Actinomadura 1940 Genus IV. Spirillospora 1959 Order XVI. Incertae sedis 1967 Genus I. Thermobispora 1967 Class II. Acidimicrobiia class, nov 1968 Order I. Acidimicrobiales 1969 Family I. Acidimicrobiaceae 1970 Genus I. Acidimicrobium 1970 Genus II. Ferrimicrobium 1972 Genus 111. Ferrithrix 1973 Family II. lamiaceae 1974 Genus I. lamia 1975

Class III. Coriobacteriia class, nov 1975 Order I. Coriobacteriales 1976 Family I. Coriobacteriaceae 1977 Genus I. Coriobacterium 1977 Genus II. Atopobium 1978 Genus III. Collinsella 1984 Genus IV. Cryptobacterium 1987 Genus V. Denitrobacterium 1988 Genus VI. Eggerthella 1990 Genus VII. Olsenella 1993 Genus VIIL Slackia 1995 Class IV. Nitriliruptoria class, nov 2000 Order I. Nitriliruptorales 2001 Family I. Nitriliruptoraceae 2001 Genus I. Nitriliruptor 2001 Order II. Euzebyales 2002 Family I. Euzebyaceae 2003 Genus I. Euzebya 2003 Class V. Rubrobacteria class, nov 2004 Order I. Rubrobacterales 2005 Family I. Rubrobacteraceae 2006 Genus I. Rubrobacter 2006 Class VI. Thermoleophilia class, nov 2010 Order I. Thermoleophilales 2010 Family I. Thermoleophilaceae 2011 Genus I. Thermoleophilum 2011 Order II. Solirubrobacterales 2014 Family I. Solirubrobacteraceae 2015 Genus I. Solirubrobacter 2015 Family II. Conexibacteraceae 2018 Genus I. Conexibacter 2019 Family III. Patulibacteraceae 2024 Genus I. Patulibacter 2024

Author index 2029

Index of scientific names of Archaea and Bacteria 2031