International Journal of Systematic Bacteriology (1998), 48, 11 29-1 143

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Phylogenetic relationships among the Chromatiaceae, their taxonomic reclassification and description of the new genera Allochromatium, Halochromatium, Isochromatium,Marichromatiurn , Thiococcus, Thiohalocapsa and Thermochromatium
Johannes F. Imhoff, Jorg W i n g and Ralf Petri
Author for correspondence: Johannes F. Imhoff. Tel: +49 431 697 3850. Fax: +49 431 565876. e-mail : jimhoff(n ifm.uni-kiel.de
lnstitut fur Meereskunde an der Universitat Kiel, Abteilung Marine Mikrobiologie, Dusternbrooker Weg 20, D-24105 Kiel, Germany

Sequences of the 16s rDNA from all available type strains o f Chromatium species have been determined and were compared to those of other Chromatiaceae, a few selected Ectothiorhodospiraceae and Escherichia coli. The clear separation o f Ectothiorhodospiraceae and Chromatiaceae is confirmed. Most significantly the sequence comparison revealed a genetic divergence between Chromatium species originated f r o m freshwater sources and those o f truly marine and halophilic nature. Major phylogenetic branches marine and halophilic species, (ii) freshwater o f the Chromatiaceae contain (i) Chromatium species together with Thiocystis species and (iii) species o f the genera Thiocapsa and Amoebobacter as recently reclassified [Guyoneaud, R. & 6 other authors (1998). lnt J Syst Bacteriol48, 957-9641, namely Thiocapsa roseopersicina, Thiocapsa pendens (formerly Amoebobacter pendens), Thiocapsa rosea (formerly Amoebobacter roseus), Amoebobacter purpureus and Thiolamprovum pedioforme (formerly Amoebobacter pedioformis). The genetic relationships between the species and groups are n o t in congruence with the current classification o f the Chromatiaceae and a reclassification is proposed on the basis o f 16s rDNA sequence similarity supported by selected phenotypic properties. The proposed changes include the transfers of Chromatium minus and Chromatium violascens to Thiocystisminor comb. nov. and Thiocystis violascens comb. nov., o f Chromatium vinosum, Chromatium minutissimum and Chromatium warmingii t o the new genus Allochromatium as Allochromatium vinosum comb. nov., Allochromatium minutissimum comb. nov., and Allochromatium warmingii comb. nov., o f Chromatium tepidum t o the new genus Thermochromatiumas Thermochromatium tepidum comb. nov., of Chromatium salexigens and Chromatium glycolicum to the new genus Halochromatium as Halochromatium salexigens comb. nov. and Halochromatium glycolicum comb. nov., o f Chromatium gracile and Chromatium purpuratum to the new genus as Marichromatium gracile comb. nov. and Marichromatium purpuratum comb. nov., of Thiocapsa pfennigii t o Thiococcus pfennigii gen. nom. rev., of Thiocapsa halophila t o the new genus Thiohalocapsaas Thiohalocapsahalophila comb. nov., and o f Chromatium buderi t o the new genus lsochromatium as lsochromatium buderi comb. nov.

~

Keywords : phylogeny, Chromatiaceae, taxonomic reclassification

The EMBL accession numbers for the 16s rDNA sequences determined in this study are given in Table 1.

1129

.. 1996). The first branch consists of Chromatium species which have been isolated from marine and hypersaline environments and which specifically and characterInternational Journal of Systematic Bacteriology 48 _________~ ~ 1130 . All strains of purple sulfur bacteria used for this study are listed in Table 1. the DSM numbers when available and the EMBL accession numbers for their 16s rDNA sequences. 1987). The distance matrix was calculated on the basis of the algorithm of Jukes & Cantor (1969) with the DNADIST program within the PHYLIP package (Felsenstein. Halorhodospira halophila strain DSM 244’ (M26630). 1984). 1991..J. DNA was extracted and purified by using the QIAGEN genomic DNA buffer set. 1996). Caumette et al. 1974).. 16s rDNA sequences of only a few species of the Chromatiaceae were available until recently (DeWeerd et al. First surveys on the relationships of purple sulfur bacteria have been made by comparison of oligonucleotide catalogues (Fowler et al. Pure cultures were grown in 100 ml screw-capped bottles completely filled with synthetic medium and incubated in the light. Stackebrandt et al. 1984). (1998). 1994). in part separated from and in part intermixed with non-phototrophic bacteria. For marine and halophilic strains.. Guyoneaud et al. Cell material for 16s rDNA sequencing was taken from 1-2 ml of well-grown liquid cultures. METHODS Source and culture of bacterial strains. Suling and R. In the present work 16s rDNA sequences of all available type strains of Chromatium species are compared to related purple sulfur bacteria and used to characterize their genetic relationships. while another one includes freshwater Chromatiurn species together with known Thiocystis species. 1980) and oligonucleotides obtained from 16s rRNA digestion with specific endonucleases (Gibson et al. according to the Escherichia coli 16s rRNA numbering of the International Union of Biochemistry).. Among the phototrophic purple bacteria. The PCR products were purified by using the QIAquick PCR purification kit. The traditional taxonomic system of these bacteria is not phylogenetically oriented. Petri INTRODUCTION Strains were cultivated in synthetic medium according to Pfennig & Triiper (1992).. F. J. Nucleotide sequence accession numbers. Ectothiorhodospira shaposhnikovii strain DSM 239 (X93480). as required. 1979. The final pH was 7. sequence analyses and comparisons lead to the separation of the Ectothiorhodospiraceae into two genera that can also be recognized by their salt responses and other phenotypic properties (Imhoff & Suling. It is quite apparent from their genetic relationships that a major deeply branching cluster with marine and halophilic Chromatiaceae species is clearly separated from another branch with predominantly freshwater species. Wahlund et al. based on the data from this distance matrix (Fig.. 1979. 1). 1990. phylogenetic information became available only with the establishment of sequencing techniques for proteins and nucleic acids on the basis of sequences of cytochrome c (Ambler et al.. 1987). 1987) which was started with the following primers : 5’-GTTTGATCCTGGCTCAG-3’ and 5’-TACCTTGTTACGACTTCA-3’ (positions 1 1-27 and 1489-1 506. Sequences were aligned using the CLUSTAL w program (Thompson et a/. Fowler et al. Fowler et al. Within this latter branch. 1979. the original strain designations.. 1984. Cultures of all strains are maintained in our laboratory (see Table 1). The FITCH program in the PHYLIP package fitted a tree to the evolutionary distances. Recombinant Taq polymerase was used for PCR (Mullis & Faloona. The 16s rDNA sequences determined in this study have been deposited in the EMBL database under accession numbers given in Table 1. and inconsistencies between the taxonomic treatment and genetic relationships of the purple sulfur bacteria have been discussed previously (Gibson et al. RESULTS PCR amplification and 16s rDNA sequencing and analysis. It was obvious already from the early investigations on the genetic relationships that phototrophic bacteria are found in several major branches of a sequence-based phylogenetic bacterial system (see Woese. Dickerson. 1984.. Truper. has been based on easily recognizable phenotypic properties (see Pfennig & Triiper. 1997. It is primarily of practical relevance. Indeed. Imhoff. Stackebrandt et al. 1977) using an automated laser fluorescence sequencer (Pharmacia). and Halorhodospira halochloris DSM 1059T (M59152) (see Imhoff & Suling. Historically. Ectothiorhodospira haloalkaliphila strain ATCC 5 1935’ (X93479). 1-10% NaCl was added to the medium. later by sequences of the 16s rRNA gene. 1989). Sequences were obtained by cycle sequencing with the SequiTherm sequencing kit (Biozym) and the chain-termination reaction (Sanger et a/. one cluster contains Thiocapsa and Amoebobacter species according to Guyoneaud et al.. 1998). On this basis and with the additional support of phenotypic properties a reclassification of these bacteria is proposed. Further sequences (accession nos in parentheses) from Ectothiorhodospiraceae and Escherichia coli (K02555) were available from the EMBL database: Ectothiorhodospira mobilis strain DSM 237T (X9348 1)... which in addition shows the previously used and the newly proposed names. Accession numbers of additional 16s rDNA sequences of Chromatiaceae that were published previously are also contained in Table 1. separate phylogenetic lines are represented by the purple sulfur bacteria (Chromatiaceae and Ectothiorhodospiraceae) in the y-Proteobacteria and by the purple non-sulfur bacteria in the a-Proteobacteria and the P-Proteobacteria. A positive correlation between differences in several phenotypic properties to similarities of 16s rDNA sequences is discussed and in particular these properties are considered to be of importance for the classification of these bacteria.. ~~~~ The relationships of the investigated type strains of Chromatium species and related purple sulfur bacteria based on almost complete sequences of the 16s rDNA is shown in a distance matrix with similarity values and Knuc values (Table 2) and a phylogenetic tree. 1984. the taxonomy of phototrophic purple sulfur bacteria. 1997.2. 1984. Stackebrandt et al. respectively. The alignment length was from positions 29 to 1381 according to the Escherichia coli numbering (including gaps 1400 positions). More recently.

.. ..... . .. . Chronm t ium salexigens and Chromatium glycolicwn form one line.. . The second cluster of this branch showed some distant relationships to the freshwater Chromatium species... Thiorhodococcus minor and Chromatium buderi form well-separated lines within this cluster (Fig..... Thiocy st is gela tinosa and Chromat ium violascens.. as could be delineated from the comparison of its oligonucleotide catalogue (Fowler et al... including Thiolamprovum pediofornw (formerly Amoebobacter pedioformis)..... . Also. . ..... . also has a number of significant positions in its 16s rDNA sequence in common specifically with Chromatium okenii.... In this cluster are Thiolamprovum pedioforme (formerly Amoebobacter pedioformis).. .st I s violacea Thiocjystis gelutinosa Rhabdoc‘liroma tium inarinurti Thiorhodococcus minus Amoehohucter purpureus Tliio caps^ I roseope rs icin a Amoehohucter pendens Unchanged Allochromatiuni vinosum Allochromatiuni ininutissiinum Allochromatium warmingii Thermochromatiuni tepidum Thiocystis minor Thiocyst is violascens Marichromatiuni purpuratum Marichromatiuin gracile Isochromutiuni huderi Hulochrortiatiuni salexigens Halochromatiurn glycolicum Tliiococcus p fennigii Thiohalocapsa halophila Unchanged Unchanged Unchanged Unchanged Thiorhodococcus minor” Unchanged Unchanged Thiocapsa pendens? 5710 5410 5500 5210 5610 63 10 6410 4250 4270 4910 4340 4310 6510 6680 4450 4210 4410 * Corrected epithet.. A pure culture of Chromatiurn weissei is at present not available and therefore the comparison of its complete sequence was not possible.... Chromat ium okenii.. Matsunaga). ... two major clusters can be recognized... ........ 1994 .... .... . 1998). A second subcluster is represented by Chroma t ium niinus.. 16s rDNA sequence of this strain as personal communication of T..... 1 .. In the first cluster Chromatiurn warmingii... Y 12376 M26629 Y 12369 Y 12365 M59 150 Y 12372 A5224438 A5224439 x93473 A5224430 X98597 X93472 Y 12373 A5002796 Y 12368 Y 11315 Y11317 X843 16 Y11316 Y 12366 Y 12364 A5002797 Chrotnutiurn okenii Chrorncitiurn vinosum Clir omci t iunz n inu t issimum z Chroniutiurn warmingii Chr onwt i urn tep idum Chr o n i t~uin ininus i Chr oina t I um violascens Chromit I uin purpura t urn Chrotnutiurn gracile Chrotnatiuin huderi Chrotnutiurn salexigens Clirotnut iuin glycolicum ThiocqmI pfennig ii Thiocup.. . is only distantly related to other bacteria of this cluster... Chromatium weissei. Thiocapsa rosea (formerly Amoebobacter roseus) and Thiocapsa roseopersicina (Fig. . a number of 1131 -~ ~ - lnterna tiona I Jo urnaI of Systematic Bacteriology 48 .. ... Type strains are denoted by T. Chromatium gracile and Chromatium purpuratum form a second line together with an isolate that has been first described as ‘ Rhodohurter marinus’ (Burgess et al........ 1984) with the sequence of Chromatium okenii. .. not available in this collection... .... ..... NN. Thiorhodovibrio winogradskyi.. the type species of the genus Chromatium. I-See Guyoneaud et al.. ... Amoehohacter purpureus. .. .... the other with Thiocapsa and Amoehohacter species.. Thiocapsapfennigii and Thiocapstr halophila belong to this cluster and therefore certainly can no longer be classified as Thiocapsa species (Guyoneaud et al.. .. ..... Species and strains of Chromatiaceae compared in this study .. 1).. istically require various concentrations of salt for growth . .. .. In addition.... which is phenotypically quite similar to Chromatium okenii...... The second branch mostly contains species that originate from freshwater habitats and is significantly distinct from the branch of truly marine and halophilic species. .. .. 6010 51 10 5310 5810 NN EMBL no... . ..s1i halophila Thiorh or10 vibr io winogradskji Thiocj3.... Rhabdochromatium marinum. Chromatium vinosum and Chromatium minutissimum form one ~ subcluster to which Chromatium tepidum is distantly related.. ... 169T 1SOT 1376’ 173T 3771T 17ST 16ST 1591T 203T 176’ 4395* 11080’ 226 62 1OT 6702T 207’ 215’ 526 1 1 151ST 4197’ 217’ 236’ Other designation(s) 1111 D MSU 6512 MC 1211 6111 984 861 1 8111 SG320 1 SL320 1 8013 SG3202 0651 1 271 1 261 1 8315 CE2203 ThSchl2 1711 1314 Authors’ no.... Although most of these species characteristically are found in freshwater habitats..... 1998)..... ... . Previous name ~~ ~~ New name DSM no. (1998)...... one with Chromatium and Thiocj*stis species... Thiocapsa pendens (formerly Amoebobacter pendens).....Phylogenetic taxonomy of Chrornatiaceae Table 1. On the basis of the sequence data. Thiocy st is vioIacea .. Guyoneaud et al.. .

34 89.14 87.19 88.063 0.28 88.099 0.13 8 0. 5 .106 0. 3.75 93.20 88.58 0.082 90.69 88. 6.092 0.32 88.70 84.112 0.80 90.20 91. Ectothiorhodospira marismortui DSM 4 180T.104 0.50 92.91 90.50 92.16 84. 33.11 85. Ectothiorhodospira vacuolata DSM 2 11lT.36 88.26 89.77 90.121 0.101 0.27 87.074 91.03 87. 87.070 0. Thermochromatium tepidum DSM 3771T.26 22 92.31 93.110 0.092 0.48 86.44 89. Thiocystis violascens DSM 168T.101 0.87 91.51 93. 1 2 3 4 5 6 89. 88.69 89.69 91.091 0-102 0.60 93.093 0.073 0.28 88.34 91. 88.091 4 0.02 92. 10.43 89.37 90.66 90.28 88.78 91.113 2 0. Thiocaps 9.06 89.53 92.38 25 83.18 89.03 88.69 90.69 88.76 88.65 90.97 86.42 94.05 90.59 87.55 84.68 88.40 88.54 91.52 91.90 89. Thiocystis violacea DSM 207T.16 87. 88.102 1 0.21.103 0.87 18 19 89.98 93. 89.28 89.05 88.25 90.41 91. 89.107 0-118 0. Thiohalocapsa halophila DSM 62 loT.099 0.131 0. Ectothiorhodospira haloalkaliphila ATCC 5 193 DSM 237T.047 0.107 9 0.027 97.99 90.28 89.62 90.065 0.17 91.24 90.28 0.26 90.85 91. Marichromatium gracile DSM 203T.95 88.63 89.30 91.098 5 0.39 90.093 0.30 90. Thiocystis gelatinosa DSM 2 15 14. Rhabdochromatium marinum DSM 5261T. Halorhodospira halophila DSM 244T.097 0.52 23 24 91.104 0.77 90. bacteria and Escherichia coli as reference species The values on the upper right are the uncorrected sequence similarities (%).40 87. 86.77 17 90.95 88.87 88.102 0.02 90.102 7 0.27 91.095 0. Organisms: 1.22 90.23 91.083 0.41 91.33 98.66 89.102 0.97 85. 0. 16.18 94.30 87.31 88.44 90.100 0.119 0.69 91.058 0. 85-67 89.095 0.52 87.093 0.109 4 0.104 0.82 92.096 0.13 91.02 88. 88..08 93.39 86.070 5 0.36 91.126 0.89 89.57 88.98 92. Allochromatium vinosum DSM 180T.138 0.10 87.113 0.114 0.054 0.44 92.109 0.70 89.03 88.13 91.096 0.089 0.31 91.01 91.98 88.82 90.89 86.12 90.28 90.04 89. 17.03 93.105 0. 88. 11.35 21 87.54 88.90 90.03 88. Halochromatium glycolicum DS salexigens DSM 4395T.121 0.69 90.72 94.93 93.71 87. 83.78 89.068 0.122 0.126 0.077 0.43 92. 28.115 0.55 83.83 89.38 91. 82-62 85.09 92.44 89.38 95.04 93.106 0.133 0.08 90.093 0.78 92. Thiorhodococcus minor DSM 11518T.93 92.69 95.97 90.28 90. Halorhodospira halochloris DSM 1059T.51 87.37 90.90 89.44 94.17 89.69 91.29 90.44 90.115 0.61 85.34 88.40 90.01 92.28 92.79 84.89 91.093 0.108 0.84 93.34 89.098 3 0.095 .68 90.119 0.095 0.120 0. Ectothiorhodospira s 27.22 9254 92.22 DSM 6702T.123 0 0.124 0.34 87.104 0.29 95.44 90.18 89.094 0.10 92.040 0.32 90.08 92.41 83.15.88 88. and the values on the lower left are K.46 90. Chromatium okenii DSM 169T.45 90.130 0.91 91.08 91.45 91.19 91.42 86.79 89.40 88.118 0.38 92.88 87.94 89.46 89. Halorhodospira abdelmalekii DSM 21 loT.99 91.129 0.49 94.099 0.086 0.07 90.28 94.097 0.093 0. Marichromatium purpuratum DSM 1591T.86 92.53 87.86 91.112 0.32 83.100 0. 88.39 93. 25.08 91.50 7 8 9 10 11 12 13 14 15 16 90.85 93.19.05 90.42 96. Thiolamprovumpedioforme DSM 3802T. Amoebobacterpurpureus DSM 4197T.01 91.097 0.71 92.105 6 0.102 0.79 94-77 94. Isochromatium buderi DSM 176T. 7. 26.20.43 92.46 91.30 90. values corrected for multiple base & Cantor (1969).070 0.047 0.104 0.68 85.102 0.10 89. Escherichia coli.099 0.99 88.20 84.08 90. 29.37 92.69 90.04 92.109 0.74 84-24 86.138 89.35 90.14 91.13 91.32 92.78 89.36 85.03 91.55 89.35 90.87 91.059 0.76 92.21 91.94 91.88 92.076 0.53 88.102 0.90 90.104 0.064 0.050 0.082 0.89 90. 8.24 93.94 88.83 88.1 I5 6 0.106 0.27 90.75 87. Thiocapsa rosea DSM 235T.12 92.48 90.07 89. 86.97 88.51 20 87..55 93.61 92.086 0.24 85.24 2 0.062 3 0. Thiococcuspfennigii DSM 226.20 91.16 91. Allochromatium minutissimum D warmingii DSM 173T.096 0.104 0.35 88.49 90.23 90.24.85 88.Table 2 Levels of 165 rDNA sequence similarity and evolutionary distances of presently recognized Chromatiurn species with oth .35 88.2.16 89.39 89.63 90.13 92.53 90.89 85.090 0.18 91.056 0.53 90. 12.10 88.04 92.05 92.46 90.61 90.112 0.112 0.094 0.053 0.61 90.32.85 87.23. 1 94. 3 1.15 26 27 87.93 89.39 87.106 0. Thiocapsa pendens DSM 236T.124 0.08 90.062 0.75 92.58 89.65 93.

118 0.07 87.155 0.149 0.163 0.088 0.159 0.138 0.182 0.100 0.153 0.075 0.22 84.097 0.155 0.198 0.155 0.199 0.155 0.104 0.134 0.145 0.128 0.143 0.122 0.080 0.166 0.118 0.180 0.110 0.159 16 17 0.150 0.57 0.155 0.136 0.111 0.88 90.160 0.32 19 91.32 89.147 0.079 0.129 0.148 0.157 0.142 0.170 0.116 0.092 0.75 17 0.126 0.78 85.160 0.08 85.101 0.132 0.112 0.110 0.150 .133 0.152 0.115 0.134 19 0.148 0.184 0.38 88.175 0.174 0.070 0.97 95.177 0.144 0.130 93.132 0.52 89.120 0.131 0.065 0.163 0.04 87.095 0.143 0.153 0.104 18 0.129 0.112 0.158 0.137 0.119 0.177 0.088 5 6 7 8 9 18 93.081 0.081 0.118 0.125 0.127 0.40 88.207 0.179 0.115 0.153 0.146 0.104 0.149 0.66 91.105 0.154 0.139 0.178 0.65 86.165 0.153 0.169 0.139 0.162 0.171 0.167 0.130 0.127 0.127 0.162 0.201 0.124 27 0.106 0.044 0.126 0.120 0.163 0.134 0.151 0.53 92.179 0.164 0.54 0.132 0.177 0.148 0.143 0.087 0.102 0.133 0.58 8 1.152 0.01 0.074 0.62 90.128 0.153 0.086 0.164 0.129 0.096 0.093 0.095 0.162 0.45 89.132 0.42 0.117 0.190 0.152 0.133 0.136 21 0.183 0.084 0.119 0.77 92.165 0.0 13 0.165 0.128 0.126 0.103 0.157 32 0.143 0.016 0.159 0.20 86.114 0.082 0.057 0.136 0.133 0.164 0.135 0.194 0.076 0.32 8 1.183 0.148 0.091 0.138 31 0.126 0.120 22 0.084 0.18 89.181 0.145 0.22 84.02 85.91 89.079 0.136 0.136 0.142 0.065 0.094 12 13 0.103 0.148 0.160 0.56 90.066 0.150 0.198 0.122 0.097 0.094 0.127 0.143 0.151 0.24 87.163 0.135 0.47 87.118 0.153 0.163 0.114 0.084 0.Table 2 (cont.191 0.045 0.07 83.141 0.101 0.122 0.151 0.084 0.106 0.150 0.156 0.042 0.169 0.105 0.073 0.07 1 0.61 21 89.120 0.145 0.130 0.12 90.120 0.141 0.131 0.125 0.35 89.164 0.39 90.194 0.094 0.130 0.135 0.088 0.118 0.083 0.084 0.108 0.164 0.185 0.044 0.145 0.094 0.074 0.160 0.155 0.184 0.146 0.104 0.63 82.181 0.152 0.080 0.145 20 88.096 0.078 0.081 0.114 0.160 0.155 0.137 0.107 0.) 1 4 10 14 ~ 2 11 15 0.148 0.090 0.209 0.142 0.186 0.127 0.122 0.129 30 0.131 0.133 29 0.125 0.159 0.156 0.96 94.126 20 0.122 0.084 0.105 0.157 0.069 0.129 0.031 0.096 0.148 0.160 0.131 0.08 1 0.087 0.103 0.65 87.104 0.098 0.070 0.08 1 0.97 98.128 0.154 0.185 0.098 0.161 0.132 0.04 84.101 0.099 0.130 0.166 0.140 0.105 0.133 0.168 0.136 0.179 0.161 0.8 1 89.1 14 0.157 33 0.123 0.124 0.125 0.119 0.101 22 90.144 0.170 0.208 0.186 0.069 0. I28 0.162 0.115 0.103 0.111 0.186 0.1 15 0.142 0.162 0.81 87.148 0.155 0.062 0.110 0.095 0.060 0.130 0.75 90.156 0.083 0.144 0.087 0.120 0.075 23 0.115 0.086 0.130 0.08 88.212 0.137 0.78 89.92 27 88.112 0.137 0.112 0.143 0.138 0.37 89.080 0.152 0.115 0.122 0.121 0.180 0.118 0.173 0.140 0.178 0.148 0.134 28 0.1 10 23 24 25 26 90.116 0.124 0.137 26 0.142 0.132 0.151 0.082 0.158 0.145 0.042 0.07 90.136 0.119 0.143 0.108 0.25 82.082 24 0.035 0.134 0.127 0.079 25 0.150 0.086 0.129 0.105 3 0.090 0.161 0.132 0.129 0.168 0.123 0.88 86.82 85.151 0.125 0.111 0.161 0.084 0.102 0.1 11 0.46 88.

so far much less considered properties. Caumette et al.. 1990. upon which the current taxonomy is based.J. some of the spherical cells containing gas vesicles (former Amoebobacter species) are now also recognized as members of this genus. 1997. The differential characteristics of the genera and species of the Chromatiaceae are summarized in Table 3. Nonetheless. Non-motile spherical purple sulfur bacteria without gas vesicles. 1998. DISCUSSION Morphological properties The available 16s rDNA sequence information of Ectothiorhodospiraceae (Imhoff & Suling. 1134 The principal classification schemes of Chromatiaceae genera use cell form.. Derived from the distance matrix (Table 2) and calculated as indicated in Methods. for example. and this work) support the separation of these two families (Imhoff. Guyoneaud & P. indicating that the presence of gas vesicles should no longer be a genus determinative property either. 1997. genetic relationships indicate that two of the three known species (Thiocapsa halophila and Thiocapsa pfennigii) are not at all related to the type species of this genus. one of the major phylogenetic clusters of Chromatiaceae is constituted out of spherical cells without flagellar movement and with or without gas vesicles (Fig. Caumette. 1974. Imhoff. 1998). supposedly are of higher significance in this regard. Wahlund et al. 1996) and Chromatiaceae species (DeWeerd et al. 1. Ectothiorhodospira haloalkaliphila ATCC SI 935' Ectothiorhodospira marismortui DSM 41 80' Ectothiorhodospira vacuolata DSM2III' Ectothiorhodospira shaposhnikovii DSM243' Escherichia coli I Fig... On the other hand. 1984a) and at the same time demonstrate the inconsistency of the present taxonomic classification of the Chromatiaceae with their genetic relationships. Suling and R. have to be regarded as being of minor importance in support of a genetically oriented taxonomic classification. motility by flagella and presence of gas vesicles as key properties (Imhoff & Truper. 1991. However. Though it would be an exaggeration to state that these properties have no taxonomic meaning. 1.. and are discussed below. 1982. Most of the morphological and physiological properties.. Guyoneaud et al. F.Pfennig & Truper. Guyoneaud et al.. International Journal of Systematic Bacteriology 48 .05 K. which is Thiocapsa roseopersicina. Phylogenetic tree showing the relationships of the investigated type strains of Chromatiurn species and related purple sulfur bacteria. it is clear that they are insufficient taxonomic characteristics and not sufficient to decide on the assignment of bacteria to a particular genus. while others. Petri Thiohnlocapsa halophila DSM 62Id Thiococcuspfmnig'i DSM226 Halochmmatium salexigens DSM 4395' Halochmmatium glycolicum DSM 110 d 8 Isochivmatium buderi DSM I7# Rhabdochromatiummarinum DSM526Ir Thiorhodovibrio winograhkyi DSM 670P I ' - - Mnichromatium gracile DSM203' Thermochromatium tepidum DSM 3 771' Allochromatium warrningii DSM 173' Allochivmatium minutimimum DSM 1376' Chivmatium okenii DSM I 6 f Thiocystis violacea DSM 207' Thiocystis minor DSM I781 I I I 0. J. ~ isolates from coastal environments do grow in the presence of low concentrations of salt but do not require salt for growth (R. 1989). personal communication). had been united within the genus Thiocapsa.

6 66-3 65.5-600 3.0 2.0 Rod 2.2 61.0 48.2 62'8-67.4 68.5 Rod 1.54.c.5 Bl2 ~.2-1.9-66-6 694-69.5-2.3 64.0 Rod 1.1-60.0 Sphere 1.5 Sphere 1-0-2.3 Rod I '5-2.5 62'5-63.0-1.0 Sphere 1.2-1.5-6.skyi A llochromatium A Ilo chso n icr t iuin vinosum A llochr on ici t ium minu t issimum Allochroniatium warmingii Thermochromatium T h e mociirom a t ium tepidum Chromatium Ciiromcr t I iiin okenii Cliromcrtium rceissei Thiocysti.5 Spiral 2.0 1.0 63-8 Chemoautotrophy Yes No No Yes Yes Yes Yes Yes Yes No No No No No No No No No Yes Yes Yes No No No No No No No No Yes Yes Yes No Yes Yes Yes Yes Yes No Yes Rod 2.3 66.5 Sphere 1*0-2*0 Rod 1'0.0-2.7 Rod 3.8-64'3 6 3.5 48.3 65-5 63. International Journal of Systematic Bacteriology 48 1135 .8 66.0 Sphere 1.0-3.5-2'5 Sphere 1.7 55.+ Thiocj3.5-2.5-4.5-3.9-2. Genus and new species name Halochroma tium Haloc*iironiatium salexigens Halochromatiurn glycolicum Thiohalocupsa Thiohaloc upsu haloph ila Thiococcus TlZiocwcwiJpjennigii Rhubdochromatium Riiabck.0-50.3 62.\ violascens Thiocapsa Thioccrpbtr roseopevsicina Tiiioctrpw rosea Thiocapstr pendens Thiolamprovum Thiolampsovumpediofovme Amoebobacter Amoebohricter purpureus Thiodicty on Tliiodictjvn elegans Tiiiodic.0-50.9 60.0 2.5ti.iiromatium marinum Isochromatium Isociirom~rtiuni buderi Thiorhodococcus Thioriiotlococcus minor Marichromatium Mariciirotnatium gracile Mariciiromatiurn purpuratum Thiorhodovibrio Thioriioclovibrio \iinograd.4 Rod 2.5 2.0-3.5 65.ijv)n bacillosum Lamprobucter Lamprohrrcter modestohalophilus Lamprocystis Lamproc~!~sris roseopersicina Thiospirillum Thiospiriiluin jenense Thiopediu Tliiopecr'icr sosea Motile Gas vesicles Cell form and size (rtm) Optimum salinity (%) $8) 20-30 3&35 20-30 20-3 5 30 25-30 30-35 30-35 25-30 33 30-35 30-3 5 25-30 48-50 25-30 25-30 30 30 25-35 30-35 20-3 5 20-35 20-35 37 23-25 20-25 20-30 Vitamins required G +C content (mol %) 64.~ .9 61.9-70.5 Sphere 3.54.9 6 1*O-62.8-1.0 None BE Bl2 Yes Yes Yes No No No Yes Yes 0 5-7 1-2 1'5-5 1-3 2 2-3 2-7 2-3 0 None None Bl2 None None 0 None None None Bl2 Yes Yes Yes No No No No Yes Yes Yes Yes Yes Yes No Yes Yes No No Yes No No No None None None None None None None 0 None Bl2 Bl2 0 None None None None None 4 2 0 None None None None None None Bl2 (I 3 1 2 1 0 None None B12 1-2 None None None 25-27 20-30 20-25 20 None Bl2 45.0 1.4 62'2-62.0-2.5 Sphere 2.0 Rod 4.\ riolacea Tiiiocj9sti.3 63.5 0.3 64. _ Phylogenetic taxonomy of Chrongatiaceae Table 3 Differential characteristics of genera and species of t h e Chromatiaceae family .5-1.1 3 1.2 3.3-66.3 65.7 Spiral 0.0 1.0-2.0 61.54-0 Sphere/rod 3-0 2.2-3.9 68.8-1.4 6 1*3-66.0 * 10 2.

Although some specialized species. Thiocapsa pendens (formerly Amoebobacter pendens) and Lamprobacter modestohalop h ilus. Sulfide is required. thiosulfate and hydrogen are not used as electron donors. Salt relationships Though salt dependence has been used already in the past to distinguish between species and even genera. Chromatiurn weissei. Thiospirillum jenense and Thiocapsu pfennigii. the other primarily freshwater isolates. None of these species assimilates sulfate as a sulfur source. The salt requirement apparently is of general taxonomic importance in phototrophic bacteria and already has been considered in the taxonomy of purple non-sulfur bacteria and Ectothiorhodospiraceae. brackish water and marine habitats. On the one hand. So far. others like Chromatiurn warmingii are moderately related to the versatile Chromatium vinosum. Some species even grow as chemoautotrophs. Among these species are several small-celled Chromatium species. F. and from the freshwater species of Chromatium vinosurpi and relatives.. Furthermore. sequence relationships indicate that a number of strains isolated from the marine environment and assigned to this species could be distinguished at the species level (Guyoneaud et al. These bacteria do not grow with organic electron donors and chemotrophic growth is not possible. Therefore. All species are able to grow photoautotrophically under anoxic conditions in the light using sulfide or elemental sulfur as an electron donor. In particular the so far known genera Chromatium and Thiocapsa have representatives in both of these phylogenetic branches. Petri Physiological properties Physiological properties always have been important characteristics in taxonomic considerations. Species such as Chomatium vinosum and Thiocapsa roseopersicina have been reported to be found in freshwater. Among these species are Chromatiurn okerzii. Thus. J. It has been found to be highly variable in some of the known Chromatiaceae genera (e. both the genetic relationships and the salt responses enable differentiation of the halophilic Chromatiurn species C'lzromatiunz salexigem and Chromatium glycolicum from the marine species Chromatiurn gracile and Chromatiunz purpuratum. Most of them are able to grow with sulfate and in the absence of reduced sulfur sources with organic substrates as electron donors for photosynthesis. Suling and R. Chromatiurn buderi and Thiocapsa pfennigii form well-separated genetic lines. expressed as the G + C content. being halotolerant but not halophilic. at present the versatility versus specialization of the species appears not to be a good taxonomic criterion. Further studies will be needed to determine whether differences in the G + C content are correlated with the occurrence of these bacteria in marine or freshwater environments. 1. In regard to Thiocapsa roseopersicina.. Major phylogenetic branches within the Chromatiaceae can be distinguished on the basis of their salt requirement (Fig. in a few species the congruence between their phylogenetic position and their salt response requires further attention. the former Rhodospirillum species (Imhoff et al. DNA base ratio The DNA base ratio. Only acetate and pyruvate (or propionate) are photoassimilated in the presence of sulfide and CO. These bacteria behave indifferently to low salt concentrations and may be poorly adapted to the marine environment. 1980. although their taxonomic significance is ambiguous. On the other hand. 1974. Chronzatiurn wwrmingii. 1994) and also genera of the spiral-shaped purple non-sulfur aProteobacteria. 1998). versatile and specialized species.0 to International Journal of Systematic Bacteriology 48 . we distinguish two major physiological groups. Genetic relationships of the Chromatiaceae (Fig. Large differences in the G + C content of the DNA of strains assigned to this species (61-3-66-3 mol YO) may indicate genetic heterogeneity. 1. which always has been regarded as a taxonomically important parameter. Thiocystis violacea. the salt relation should be considered as property for characterization of ' genetically defined ' genera. Thiocapsa rosea (formerly Amoebobacter roseus). Although salt relationships obviously are important properties for the taxonomic characterization of Clzromatiaceae. is gaining new attention in view of a genetically oriented taxonomic system. phylogenetic relationships of the Chlorobiaceae indicate that salt relationships may be a useful criterion also for these bacteria (Overmann & Tuschak. Thiocapsa roseopersicina. As a consequence thereof. Kampf & Pfennig. one exclusively containing marine and halophilic species.. 1976. such as Chromatiurn okenii. 1998).J. values from 48. Kondratieva et al. physiological properties of the Chromatiaceae are quite conservative in regard to the use of sulfur sources and the restricted use of organic carbon sources. Chroulzat iurn buderi. the phylogenetic and taxonomic importance of this property has so far not been fully recognized. 1986).. Table 2). and a few species can grow chemoheterotrophically in addition (Gorlenko. The specialized species depend on strictly anoxic conditions and are obligately phototrophic. 1996). Rhodovulum and Rhodobacter (Hiraishi & Ueda. This implies principally that a separate evolutionary development has occurred in the marine and in the freshwater environments and that therefore the salt response should be considered as an important 1136 taxonomic criterion. justifying their inclusion into one and the same genus. Table 2) revealed the obvious divergence of freshwater and saltwater bacteria. In regard to Chromatiurn vinosum the situation is unclear at present. 1997).g. The versatile species photoassimilate a greater number of organic substrates. Imhoff. It proved to be of relevance to distinguish the genera Halorlzodospira and Ectothiorhodospira (Imhoff & Suling. clear proof of identity of various isolates from marine habitats assigned to Chromatium vinosum on the species level is lacking.

1-60.2 mol YO). The differentiation of the Chromatiaceae species on the basis of some diagnostic properties is shown in Table 3. ('the other Chromatium') for the new genus.8-67. Thiemann & Imhoff. the lipid composition of Thiocapsa pfennigii is different from all other so far analysed phototrophic purple bacteria (J. nov. it appears that these components. is congruent with the sequence-based similarity of these bacteria and may well turn out to be a significant property to distinguish genetically related groups of Clzromatinceae in the future. Imhoff. Further studies are needed to clarify whether analytical problems have caused these variations or misclassification of some strains is indicated. contain the same lipids. Therefore.4 mol (YOG C in the genus Chronzatiunz). respectively. Chromatium vinosum (61-3-66-3 mol YO). Chromatium violascens and the two known Tlziocystis species). (2) Chromatium vinosum. specific adaptation mechanisms are involved. Clzromatium vinosum and Chromatium warmingii. Molecular parameters Though only scattered information is available on fatty acid and quinone composition of Chromatiaceae. and Thiocj7. The only species of the Chromatiaceae so far known. 1996). two species of the proposed new genus Allochromatium. 1995). nov. despite surprisingly similar patterns in genetically related species.However. we propose to transfer these species to the genus Tlziocystiswith the new names Thiocystis minor comb. which exhibits a very high optimum temperature response is Chronzntium tepidum (optimum at 48-50 "C). Variation at the intraspecies level of more than 5 and up to 10 mol YOG + C content of the DNA is by far beyond an acceptable level of natural variation within bacteria as closely related as to justify their classification into one and the same species.Phylogenetic taxonomy of Chromatiaceae 70. The 16s rDNA sequence data justify this conclusion. which is approximately 63-66 mol YO the Thiocapsa/Amoebobacter cluster. the proposed members of the genus Thiocystis (Chromatium minus. and Thiocystis violascens comb. as additional species of the genus. the search for more of such properties must be continued and those properties that are of relevance must be analysed in all known species. the following taxonomic changes are proposed : (1) Chromatium minus and Chromatium violascens are most closely related to Thiocystis gelatinosa. as far as known. Their lipid composition is different from a second group of species. Therefore. because of their genetic distance from Chromatium okenii. In the true Chromcitium species it is as low as 48-50 mol YO. respectively. Imhoff. Thus. it is very well indicated to taxonomically consider the successful adaptation to the extremes of temperature. 1991. unpublished results). the pattern of polar lipid composition. as far as the temperature extremes are concerned.9 mol YO) have surprisingly high intraspecies variation of their G + C content. However. genetically closely related species of the Chroniatiaceae in general have a quite narrow range of their G C content.s violacea (62. Despite these first attempts to define phenotypic properties that are congruent with the genetic relationships of the purple sulfur bacteria. nov. shows significant variation between International Journal of Systematic Bacteriology 48 On the basis of their genetic relationships and in consideration of the preceding discussion on the relevance of various phenotypic properties. Chromutiurn warmingii (55. which also show uniformity in this property. Imhoff & Bias-Tmhoff. unpublished results). Chromatium tepidum is the only thermophilic isolate of the purple sulfur bacteria described so 1137 . as the type species and Allochromatium minutissimum comb. which also is genetically distinct from other species within the cluster of freshwater species.sti. For example. F. Because Chromatium vinosum is among the best-studied purple sulfur bacteria and the name Chromatium is very much associated with this species. (3) Because Chromatium tepidum quite obviously can not be retained within the genus Chromatium and it is neither specifically related to Tlziocystis nor to Allochromatium. + + groups of species (J. Chromatium minutissimum and Chromatium tvarnzingii are closely related species that should be placed into a new genus. However. which are wellaccepted tools for identification and taxonomic differentiation of phototrophic bacteria (Imhoff. in between 6 1-64 mol YO the proposed Thiocystis group in (with the exception of a few strains of Thiocystis violaceu). show minor variation and are of low significance for taxonomic differentiation within the Clzromcrtiliceae (Imhoff & Bias-Imhoff. specific environmental niches are inhabited and a different evolutionary background can be expected. 1984b. with Allochromatium vinosum comb. Indeed. the polar lipid composition. we propose the name Allochromatium gen. More analytical data are required to support this concept. nov. nov. nov. at 69-70 mol YO in the proposed Marichromatium species and at 64-66 mol% in the proposed Halochromatium species. Proposed taxonomic changes Temperature response The temperature response of many species is not wellresolved and in most cases probably of minor significance for taxonomic purposes. A more natural system would be expected to group bacteria together that have much less variation in their G + C content. few A species. and Allochromatium warmingii comb. 1995. but different from this bacterium at the species level. it is assumed that to a large extent the intrageneric and also the intraspecies variability represents the nonphylogenetic nature of the current taxonomic system and misclassification of strains.. it should also be placed in a new genus. that has been analysed so far only in several freshwater species. F. In contrast.

as the type species and Halochromatiuni glycolicum comb. J. Cells are rod-shaped or ovoid to spherical. M. Chromatium the one which is coloured). (4) Chromatium salexigens and Chromatium glycolicum shall be transferred to the new genus Halochromatium gen. Suling and R. Multiply by binary fission. is required. Thiocystis sulfur bag).J. 60AL Thiocystis (Thi.cys'tis.L. Multiply by binary fission. F..0-50-0 mol% (Bd). Contains internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll a and carotenoids are located. is proposed as the new genus name and Thiocapsa halophila is transferred to the new genus as Tliiohalocapsa halophilu comb. motile by means of flagella. globules of elemental sulfur are transiently stored inside the cells. was not validly published and is illegitimate and therefore was not included into the Approved Lists of Bacterial Names. with Halochromatium salexigens comb. Cells are straight or slightly curved rods. According to Rule 33c of the International Code of Nomenclature of Bacteria (Lapage et al. Gr. 174AL Chromatiurn (Chro. 1970). Certainly. n. 174AL). Gram-negative. because it is significantly different from this genus on the basis of the available genetic sequence information and belongs to the cluster of the halophilic Chromatiaceae species. Vitamin B. Assimilatory sulfate reduction is lacking. Mesophilic bacteria International Journal of Systematic Bacteriology 48 . we propose to revive the genus name Thiococcus Eimhjellen et al. (8) Chromatium buderi is a marine species of the Clzromatiaceae. Habitat: ditches. In the presence of sulfide and bicarbonate. thios sulfur. cells are capable of photolithoautotrophic growth with sulfide or sulfur as electron donor. chroma colour . Elemental sulfur is formed as an intermediate oxidation product and stored in the form of highly refractile globules inside the cells. Its phenotypic traits together with the results on its genetic relationships to other Chromutiaceae led us to consider this species as a separate genus. Sulfate is the ultimate oxidation product. neut.. we regard the thermophilic nature and high temperature optimum of this species as a valid criterion for genus differentiation and propose Thermochromatium gen. Gram-negative. M. The name Thiococcus. ponds and lakes with stagnant freshwater containing hydrogen sulfide and exposed to light. which is the highest within the Chromatiaceae. are proposed for these two species. nov. Petri far. Because of significant genotypic and phenotypic properties to these bacteria.. Most likely also Chromatiurn weissei is a species of this genus. n. the bacterium described as Thiocapsa Iialophila is not related to the genus Thiocapsa and is most similar on the basis of its 16s rDNA sequence to Thiocupsa pfennigii. (6) By its intracytoplasmic membrane system and its pigment composition. freshwater bacteria without a distinct requirement for salt. rev. and Marichromatiuin purpuratum comb. n. The G + C content of the DNA is 48. cystis the bladder. 1992). as the new genus name and Thermochromatium tepidum comb. nov. Emended description of the genus Thiocystis Winogradsky 1888. nov. Marichromatium gracile is the proposed type species. nov. more than 3 pm wide. nov. fem. Chromatiurn salexigens and Chromatium glycolicum (Fig. n. (7) Phylogenetically. strictly anaerobic. Contain internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll a and carotenoids are located. 1967 and to transfer Thiocapsa pfennigii to Thiococcus pjennigii gen. originally used by Eimhjellen et al. Type species is Chromatiurn okenii (Ehrenberg 1838. ( 5 ) Chromatium gracile and Chromatium purpuratum are transferred to the new genus Marichromatium. because of their extended salt dependence and salt tolerance. before cell division often diplococcus-shaped. as the new name for this species. Emended description of the genus Chromatiurn Perty 1852. Eimhjellen. n. require sulfide-reduced media. Although it is not well-established whether and to what extent the temperature response is of phylogenetic and taxonomic significance. several simple organic substrates are photoassimilated. may grow in irregular aggregates surrounded by slime. Thiocapsa pfennigii is unique among the family Chromatiaceae (Eimhjellen et al.. Tmhoff. 1). comb. nom. The new names Marichrornatium gracile comb. The name is proposed according to the marine origin and the moderate salt dependence that is characteristically found in typical marine bacteria. it can no longer be regarded as a species of the genus Thiocapsa. The final oxidation product is sulfate. nov. Gr. However. During sulfide oxidation. Gr. Perty 1852. Under anoxic conditions. Cells occur singly or in pairs.L. Mesophilic with optimal growth temperatures of 20-30 "C. Chromatium buderi is transferred to this new genus as Isochromatium huderi comb. Cells occur singly or in pairs. as an additional species. nov. it should be recognized as a separate and new genus. able to photoassimilate only a few simple organic compounds such as acetate and pyruvate. (1967) to describe this bacterium. Obligately phototrophic. Facultatively chemoautotrophic or mixotrophic under micro-oxic to oxic conditions.um.o. thiosulfate may also be used.ma'ti. Under anoxic conditions capable of photolithoautotrophic growth with sulfide or elemental sulfur as electron donor. motile by a polar tuft of flagella. nov. nov. In the presence of sulfide and bicarbonate. according to its 16s rDNA sequence it is dissimilar to any of the other described 1138 species of this branch to a degree that suggests its recognition as a separate genus for which the name Isochromatium is proposed. Thiohalocapsa gen. nov. a bag. nov. 1967.

neut. Elemental sulfur globules are transiently stored inside the cells. hals salt. named after Norbert Pfennig. halos of the salt. n.966. cell aggregates may be formed. multiply by binary fission. Thiocystis minor and Thiocyst is violascens. Individual cell surrounded by a strong slime capsule . 1 7424L) (vi. rev. non-motile . lnterna tional Journal of Systematic Bacteriology 48 Thiohalocapsa halophila (Thiocapsa halophila Caumette et al. L. n. Description of Thiohalocapsa gen. 1888. The description is the same as that for Chromatium minus (Winogradsky. n. masc. Elemental sulfur is formed as an intermediate oxidation product and stored in the form of highly refractile globules inside the cells. M. Gram-negative. Habitat: brackish and marine waters and sediments containing hydrogen sulfide and exposed to the light. Gr. globules of elemental sulfur are transiently stored inside the cells. M. 1852. Good growth at 20-35 O C . Thiohalocapsa the sulfur capsule of the salt).o. Cells are spherical with diplococcus-shaped division stages. 1139 . The description is the same as that z for Chromatium violascens (Perty. n. Gr. L. minor less. 1980). pfennigii of Pfennig. 65AL.nig’i.ha. Contain internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll a and carotenoids are located. 1989).lo’phi. Gr. Capable of photolithoautotrophic growth with sulfide and sulfur as electron donor. philos loving. nov. Gr. Salt is required for optimum growth. n.5. final oxidation product is sulfate. Thiococcus pfennigii (Thiocapsa pfennigii Eimhjellen 1970. gen. The genus name Thiococcus was proposed for this bacterium by Eimhjellen et (11.las’cens. Thiococcus (Thi. several of which may be found also in brackish and marine environments.um. The name Thiococcus is revived according to Rule 33c of the International Code of Nomenclature of Bacteria (Lapage et al. final oxidation product is sulfate. violascens becoming violet). multiply by binary fission. masc. 1989).chro.9 mol% (Bd). Thiohalocapsa (Thi. halophila saltloving). nov. The G + C content of the DNA is 69.lo. The description is the same as that for Thiocapsa pfennigii (Eimhjellen.la.cap’sa. 193AL)(pfen. Cells are spherical. Description of Thiohalocapsa halophila comb. Thiocys t is minor (Chroma t ium minus Wino gradsky 1888.Phylogenetic taxonomy of Chrornatiaceae with good growth at 25-35 “C. adj.Approved Lists of Bacterial Names. elemental sulfur and possibly other electron donors. nov.o. However. Cells occur singly or in pairs. Principally freshwater bacteria.5-7.6 mol % (Bd). Internal photosynthetic membrane system consisting of bundles of ribbonlike branched tubes continuous with the cytoplasmic membrane. Description of Thiococcus gen.o. Obligately phototrophic and strictly anaerobic. gen. n. Habitat : sulfide-containing waters and sediments of saltern and other coastal habitats with elevated salt concentrations and exposed to light. Description of Halochromatium gen. Gram-negative. pH 6. Halochrowlatium (Ha. nov. Habitat: anoxic sulfide-containing water and mud of freshwater ponds and lakes as well as brackish water and marine environments exposed to light. fem. a German microbiologist).L. Type species is Thiocystis violacea Winogradsky 1888. 199I).i.5. fem. 1989). halos of the salt.. Gram-negative. Multiply by binary fission.. capsa capsule.lo. coccus sphere. M. The G + C content of the DNA is 61-3-67. Halochromatium the Chromatium of the salt). 99AL) (mi’nor. Assimilatory sulfate reduction lacking. Under anoxic conditions in the light capable of photolithoautotrophic growth with sulfide or elemental sulfur as electron donor. nom. fem. Vitamin B. During oxidation of sulfide.L. Mesophilic bacteria with good growth at 20-30 OC. Photosynthetic pigments are bacteriochlorophyll b and carotenoids. Thiocyst is violascens (Chroma t ium violascens Pert y 1852. colour of cell suspensions yellowish to orange-brown.L. Pfennig & Truper. Type species is Thiococcus pfennigii. In the presence of sulfide and bicarbonate organic substrates are photoassimilated. Pfennig 8 Truper. L. 1970. Chromatium a genus name. L.L. nov. n. n. nov. Gr. Internal photosynthetic membrane system of vesicular type containing the photosynthetic pigments bacteriochlorophyll a and carotenoids. pH 6. n. Description of Thiococcus pfennigii comb.Other species are Thiocystis gelatinosa.5-7. this name has never been validly published and the organism was validly named Thiocapsa pfennigii (Eimhjellen. gen. Individual cells colourless. Description of Thiocystis minor comb. adj.469. The G + C content of the DNA is 65. Chemoautotrophic or chemoorganotrophic growth possible under micro-oxic to oxic conditions in the dark. fem. Description of Thiocystis violascens cornb.. comp. is required. motile by polar flagella. (1967). smaller).coc’cus. 1991) (ha. Type species is Thiohalocapsa ha loph ila . M. M. thios sulfur.L. part. Cells are straight to slightly curved rods. Pfennig & Truper. Gr. The description is the same as that for Thiocapsa halophila (Caumette et al. 1992).6. Under anoxic conditions capable of photoautotrophic growth with sulfide.9 mol YO (Bd).ma’ti. adj. pH 6. n. n. thios sulfur. 1970. Elevated concentrations of sodium chloride are required for growth.5-7. Thiococcus sphere with sulfur).

Thermochroma tium tepidum (Chromatium tepiduni Madigan 1986) (te’pi.J. Gramnegative. L. Imhoff.cigens Caumette et al. HPLC). tepidunz lukewarm). nov. photolithoautotrophic growth with sulfide or elemental sulfur as electron donor under anoxic conditions in the light.le. 1913. which are exposed to light. Pfennig & Truper. Description of Halochromatium salexigens com b. motile by polar flagella. Optimum growth at salt concentrations typical of marine bacteria. Thermochromatium (Ther. gracile thin. and coastal lagoons with elevated salt concentrations. Description of Marichromatium purpura tum comb. Type species o is Thermochromatium tepidum. In the presence of sulfide and bicarbonate. adj. 1988) (sal.a’tum. occur singly or in pairs or may stick together and form clumps. L. maris of the sea. part. Gram-negative.ri . microbial mats in hypersaline environments. 1980a). (Chromatium M a r ichr oma t ium purpurat um purpuratum Imhoff and Truper 1980b.chro. Wing and R. organic substrates can be photoassimilated. The description is the same as that for Chromatium gracile (Strzeszewski. Contain internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll a and carotenoids are located. Chemolithotrophic and chemo-organotrophic growth may be possible under micro-oxic conditions in the dark. Chromntium a genus name.ex’i. Description of Allochromatium gen. purpuratum dressed in purple). thermos hot. L . Hydrogen may be used as electron donor. Description of Marichromatium gen. Habitat : sulfide-containing hot springs of neutral to alkaline pH. Freshwater bacteria without a requirement for significant levels of salt. M.L. The G + C content of the DNA is 68-9-70. An additional species is Halochromatiurn glycolicum. to demand.mo. M. In the presence of sulfide and bicarbonate. motile most likely by polar flagella. In the presence of sulfide and bicarbonate organic substrates can be photoassimilated. Elevated salt concentrations are required for growth. Type species is Marichromatium gracile. nov. The description is the same as that for Chromatium tepidum (Madigan.. Chromatium a genus name. marine sponges and other marine invertebrates. Capable of photolithoautotrophic growth with sulfide or elemental sulfur as electron donor under anoxic conditions in the light. M. neut. Gr. Elemental sulfur is formed as an intermediate oxidation product and stored inside the cells.ma’ti. allos the other. Habitat : anoxic marine sediments and stagnant waters con1140 Cells are straight to slightly curved rods. Elemental sulfur is formed as an intermediate oxidation product and stored in the form of highly refractile globules inside the cells.chro. Description of Marichromatium gracile comb. Hulochromatium salexigens (Chromatium sale. Thermochroma tium the hot Chromatium). 1986). multiply by binary fission. taining hydrogen sulfide and exposed to light. Sulfate is the ultimate oxidation product. Mar ichroma t ium (Ma.pur.666. Contain internal photosynthetic membrane systems of vesicular type in which the pho t 0synt hetic pigments bac teri ochlo ro p h yll a and carotenoids are located. Description of Thermochromatium tepidum cornb. neut. Chemolithotrophic and chemoorganotrophic under micro-oxic conditions in the dark. 1988). Habitat: reduced sediments in salinas.L. L. neut. Sulfate is the ultimate oxidation product. exigo demanding. pH 6.lo. salexigens salt-demanding). The G + C content of the DNA is 64. The description is the same as that for Chromatium glycolicum (Caumette et al. n. Petri Sulfate is the ultimate oxidation product. Allochromatium the other Chromatium). n. Marichromatium gracile (Chromatium gracile Strzeszewski 1913.cum. M. slender). Cells are straight to slightly curved rods. The G + C content of the DNA is 60-63 mol Y (Bd). L.3 mol% (Bd.. Allochromatium (Al.um.L. adj.co’li. neut.6. Type species is Halochromatium salexigens. An additional species is Marichromatium purpuratum. Mnrichromatium the Chromatium of the sea. 1989). Multiply by binary fission.2-7.chr 0. M. nov. neut. nov.4 mol YO (Bd). the truly marine Chromatium).ma’ti. F. The description is the same as that for Chromatiurn purpuratum (Imhoff & Truper.6. nov. gen. nov. International Journal of Systematic Bacteriology 48 .gens. adj. Gr. 1997). adj. um . L. n. Mesophilic with optimal growth temperatures of 20-35 “C. Not capable of assimilatory sulfate reduction. organic substrates can be photoassimilated.ma’ti. Halochroma tium glycolicum (Chromatium glycolicum Caumette et al. ’ Description of Thermochromatium gen. v. nov. adj. J. optimum pH 7. glycolicum related to glycolate as a substrate). adj. nov. 1997) (gly.0. sal salt. Mesophilic bacteria with optimal growth temperatures of 25-35 OC. n. Obligately phototrophic. Chromatium a genus name.L. 321”IJ) (gra’ci.5-7. Molecular hydrogen may be used as electron donor. 60 1 ”) (pur.. adj. Description of Halochromatium glycolicum comb. neut.dum. The description is the same as that for Chromatiuni salexigens (Caumette et ul. Thermophilic bacteria with optimal growth temperatures at 48-50 “C. n. pH optimum is 7.um. neut.L.

Hermoso. Triiper (University of Bonn. M. a German plant physiologist). & Matheron. L. neut. nov.1-66-3 mol% (Bd). Pfennig (Uberlingen. Contain internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll CI and carotenoids are located. Description of Allochromatium warmingii comb. 1141 . n. The description is the same as that for Chromatium buderi (Truper & Jannasch.2-62-8 mol YO buderi. Imhoff. named after E. Clzromatium a genus name. Characteritation of Chromatium salexigens sp. M. motile by polar flagella. Internationa I Journal of Systematic Bacteriology 48 Chromutiunzglycolicum sp.nu. nov. J. Meyer.no’sum. R. sequence variation among the recognized species of purple nonsulfur photosynthetic bacteria. 1 1 . (1991). Caumette.5-7. pH range 6-5-7. Multiply by binary fission. Bartsch. T. neut. Photolithoautotrophic growth with sulfide or elemental sulfur as electron donor under anoxic conditions in the light. Type species is Isochromatium 62. Cytochrome c. ponds and lakes with stagnant freshwater containing hydrogen sulfide and exposed to light. multiply by binary fission. Germany) for providing the original cultures and introduction into the handling of these creatures. Caumette. Mesophilic bacteria with optimal growth temperatures of 25-35 O C .. (1997). nov. Gram-negative. The description is the same as that for Chroma t ium minu t issimum (Winogradsky . named after J.L.so. 1O4SAL)(war. warmingii of Warming. adj. 364AL) (bu’deri M. D. minutissimum very small. Allochromrtium minu t issimum (Chromat ium niinutissinium Winogradsky 1888. Description of Allochromatium min utissimum comb. Gramnegative. Syst Appl Microbiol 10. nov.. Description of lsochromatium buderi corn b. M.6. neut. n. nov. Habitat: estuarine salt flats and salt marshes. 1. isos equal. The G C content of the DNA is 55. potentially mixotrophic. Burgess. Gr.L.. gen. Allochrormtium vinosum (Monas vinosa Ehrenberg 1838. The G + C content of the DNA is (Bd). Other species are Allochroniutiurn minutissimum and Allochromatium lvarnzingii. vinosum full of wine). In the presence of sulfide and bicarbonate organic substrates can be photoassimilated. Pfennig & Triiper. 1968). R. 1888. estuaries and salt marshes.Phylogenetic taxonomy of Chromatiaceae ~~ Cells are straight to slightly curved rods. P.tis’si.mum. G.. Yamada.6. P. adj.L. pH range 6. Kawaguchi. Obligately phototrophic and strictly anaerobic. P. Habitat: ditches. Type species is Allochronicrtiuni vinosum. a halophilic Chromatiaceae isolated from Mediterranean Salinas. Nature 278.. E. Baulaigue. REFERENCES Ambler. G . Rhodobacter marinus sp. 659-660. producing photosynthetic bacterium which is sensitive to oxygen and sulphide. & Matheron. Chromatium warmingii Migula 1900.. Description of lsochromatium gen. Baulaigue. except for strains that originate from the marine and brackish water environment and may tolerate or require low concentrations of salt. motile by a polar tuft of flagella. Cells occur singly or in pairs.. 1989). 965-970. Buder. Chromatium vinosum Winogradsky 1888. Arch Microbiol 155. Mesophilic bacteria with optimal growth temperatures of 25-35 “C. & Matsunaga. + Cells are straight to slightly curved rods. also sewage lagoons. (1988). 1900. Elemental sulfur is formed as an intermediate oxidation product and stored in the form of highly refractile globules inside the cells.. M. R. Arch Microbiol 167. Isochromatium buderi (Chromatiurn buderi Triiper and Jannasch 1968. No distinct requirement for salt. Salt is required for growth at concentrations typical for marine bacteria. T. Contain internal photosynthetic membrane systems of vesicular type in which the photosynthetic pigments bacteriochlorophyll a and carotenoids are located. 284-292. Daniel. R. J. Isochrornatium the similar Chromatium. a Danish botanist). Allochrornirtium warmingii (Monas warmingii Cohn 1875. gen.chro. Thiocapsa halophila sp. Pfennig & Truper.min’gi. 1989). J. P. sup. nov. adj. The description is the same as that for Chromatiurii ivnrvlzingii (Migula. R. 167 . Caumette. Elemental sulfur is formed as an intermediate oxidation product and stored in the form of highly refractile globules inside the cells. R. similar..ma’ti. Microbiology 140. nov. In the presence of sulfide and bicarbonate organic substrates can be photoassimilated. R.. 1OOAL) (mi. 170-1 76. nov. 11-18. Warming. The description is the same as that for Chromatium vinosun-1 (Winogradsky. : a new marine-hydrogen Isochromutiuni (1. L. R. Under anoxic conditions in the light capable of photolithoautotrophic growth with sulfide or elemental sulfur as electron donor. 1888. (1979).um. smallest). A.i. & Matheron. 1989). Sulfate is the ultimate oxidation product. Sulfate is the ultimate oxidation product. n.. & Kamen. nov. the bacterium similar to Chromatium).. Molecular hydrogen may be used as electron donor. Germany) for discussion on and proposal of new names and Professor D r N. G. ACKNOWLEDGEMENTS We would like to express our gratitude to Professor D r H. a new halophilic phototrophic purple sulfur bacterium. Description of Allochromatium vinosum cornb. Pfennig & Triiper. (1994).. buderi of Buder. 99“L) (vi. a moderately halophilic purple sulfur bacterium that uses glycolate as substrate. F. Siiling.

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