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Laboratorium voor Microbiologie, Faculteit Wetenschappen, Universiteit Gent, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium
Correspondence
Peter Vandamme
peter.vandamme{at}ugent.be
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, the type species of the genus Wautersia, is a later synonym of Cupriavidus necator Makkar and Casida 1987, the type species of the genus Cupriavidus. In conformity with Rules 15, 17, 23a and 37a(1) of the International Code of Nomenclature of Bacteria, the genus name Cupriavidus has priority over the genus name Wautersia, and all other members of the genus Wautersia are reclassified into Cupriavidus as Cupriavidus basilensis comb. nov. (type strain LMG 18990T=DSM 11853T), Cupriavidus campinensis comb. nov. (type strain LMG 19282T=CCUG 44526T), Cupriavidus gilardii comb. nov. (type strain LMG 5886T=CCUG 38401T), Cupriavidus metallidurans comb. nov. (type strain LMG 1195T=DSM 2839T), Cupriavidus oxalaticus comb. nov. (type strain LMG 2235T=CCUG 2086T=DSM 1105T), Cupriavidus pauculus comb. nov. (type strain LMG 3244T=CCUG 12507T), Cupriavidus respiraculi comb. nov. (type strain LMG 21510T=CCUG 46809T) and Cupriavidus taiwanensis comb. nov. (type strain LMG 19424T=CCUG 44338T).
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to accommodate a non-obligate bacterial predator of various Gram-negative and Gram-positive soil bacteria and fungi (Byrd et al., 1985; Sillman & Casida, 1986; Zeph & Casida, 1986). The single known isolate, strain N-1T (=LMG 8453T), was obtained from soil in the vicinity of University Park, PA, USA. When confronted with Agromyces ramosus mycelia during the so-called ‘attack–counter-attack’ predation process, this strain produces several chemical signals, one of which chelates copper. C. necator is highly resistant to copper and its growth initiation is strongly stimulated by copper (Makkar & Casida, 1987).
Makkar & Casida (1987) reported the DNA base ratio and a wide range of morphological, biochemical and nutritional properties of this organism but did not examine its phylogenetic position through 16S rRNA studies as is currently standard procedure in prokaryotic taxonomy. They noticed several characteristics their organism shared with members of the genus Alcaligenes, which, at that time, comprised multiple species, including Alcaligenes faecalis (the type species), Alcaligenes xylosoxidans and allied species (now all classified in the genus Achromobacter; Yabuuchi et al., 1998) and Alcaligenes eutrophus [first reclassified in the genus Ralstonia (Yabuuchi et al., 1995) and recently transferred again, to the novel genus Wautersia (Vaneechoutte et al., 2004)]. However, a few unique biochemical characteristics and the spectacular predatory activity convinced Makkar & Casida (1987) to classify their strain into a novel genus and species.
In the course of a long-term study of the biodiversity of various Burkholderia cepacia-like bacteria, we discovered a nearly complete 16S rRNA gene sequence that was deposited for C. necator in the public database under the accession number AF191737. This sequence was very similar to that of Wautersia eutropha isolates. Fig. 1 shows the result of the comparison of the 16S rRNA gene sequence of C. necator LMG 8453T with those of strains representing Wautersia, Ralstonia and other 
-Proteobacteria. The phylogenetic analysis, including a bootstrap analysis with 1000 replicates, was performed using the BioNumerics 3.5 software package (Applied Maths) and was based on the neighbour-joining method using nearly complete sequences and excluding undetermined base positions from the calculations. The similarity level between the 16S rRNA gene sequences of strains C. necator LMG 8453T and W. eutropha LMG 1199T was 99·7 %.
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), after growth of the isolates for 48 h at 37 °C on trypticase soy agar (BBL). Densitometric analysis, normalization and interpolation of the protein profiles were performed using the GelCompar 4.2 software package (Applied Maths). The whole-cell protein profiles of C. necator LMG 8453T and of W. eutropha LMG 1199T (=TF93T) and LMG 1201 (=H16), two established W. eutropha reference strains (Jenni et al., 1988), were very similar (Fig. 2).
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, using an HTS7000 Bio Assay Reader for the fluorescence measurements. The hybridization temperature was 50 °C. DNA was prepared as described by Pitcher et al. (1989). The DNA–DNA binding values obtained were 100 % between W. eutropha LMG 1199T and LMG 1201 (which is in perfect agreement with a previously published value; Jenni et al., 1988), 79 % between W. eutropha LMG 1199T and C. necator LMG 8453T and 92 % between W. eutropha LMG 1201 and C. necator LMG 8453T. These data indicate unambiguously that the three isolates represent the same genospecies and confirm that, in the genera Ralstonia and Wautersia, high protein electrophoretic similarity correlates with a high level of DNA–DNA hybridization (Coenye et al., 1999, 2003a; Vandamme et al., 1999; Goris et al., 2001).
Given the reported difference in DNA base ratio for the two taxa [67 mol% G+C for W. eutropha (Goris et al., 2001) versus 57 mol% for C. necator (Makkar & Casida, 1987)], we determined the DNA base ratio of C. necator LMG 8453T by two approaches. The DNA was enzymically degraded into nucleosides as described by Mesbah et al. (1989). The obtained nucleoside mixture was then separated by HPLC using a Waters SymmetryShield C8 column thermostatted at 37 °C. The solvent was 0·02 M NH4H2PO4 (pH 4·0) with 1·5 % acetonitrile. Non-methylated lambda phage DNA (Sigma) was used as the calibration reference. In addition, the DNA base ratio was also determined by thermal denaturation and calculated as described by De Ley (1970). The former method yielded a G+C content of 65 mol%, the latter 66 mol%. These values are similar to values previously determined for W. eutropha (Goris et al., 2001; Jenni et al., 1988) and clearly different from the value for C. necator determined by Makkar & Casida (1987). We believe that this difference is due to experimental error in the original study.
The results of the extensive biochemical characterization of C. necator LMG 8453T generally correlate well with those provided by Yabuuchi et al. (1995) and De Baere et al. (2001) for W. eutropha. Both organisms are reported as Gram-negative, peritrichously flagellated bacteria with an oxidative metabolism. They produce catalase and oxidase and reduce nitrate to nitrite but exhibit no DNase activity. They hydrolyse Tween 80, but not urea, gelatin or aesculin. The remarkable resistance to (and growth stimulation by) copper was one of the key arguments for excluding strain LMG 8453T from the genus Alcaligenes (Makkar & Casida, 1987). However, resistance to copper and a range of other metals is well-documented for species now classified in Wautersia and is often plasmid-borne (Mergeay et al., 2003).
The results of the present study indicate that, in the 1980s, the isolate described by Makkar & Casida (1987) should have been classified as Alcaligenes eutrophus Davis 1969. Alcaligenes eutrophus was reclassified in the novel genus Ralstonia, together with two former Burkholderia species, Burkholderia solanacearum and Burkholderia pickettii (Yabuuchi et al., 1995). Subsequently, the genus Ralstonia was divided into Ralstonia sensu stricto and the novel genus Wautersia, with W. eutropha as the type species. As outlined above, the name C. necator was validly published in 1987 and the names Ralstonia and Wautersia were only published much later. Rule 23a of the International Code of Nomenclature of Bacteria (Lapage et al., 1992) specifies that each taxon above a species can bear only one correct name, that is, the earliest that is in accordance with the Rules of the Code.
In addition, the nomenclatural type of a taxon is that element of the taxon with which it is permanently associated. Rule 42 specifies that, in the case of subspecies, species, subgenera and genera, if two or more taxa of the same rank are united, the oldest legitimate name or epithet is retained. Therefore, the genus name Wautersia is a later synonym of the genus Cupriavidus, for which the type species is C. necator (Rule 15 of the Code). Furthermore, the Code stipulates that the type determines the application of the name of a taxon if the taxon is subsequently divided or united with another taxon (Rule 17). While renaming and subsequent further renaming of bacterial species causes confusion and, not the least, irritation in the wider microbiological community, adhering to the rules of nomenclature is essential for establishing a truly systematic taxonomy. Therefore, while it may be inconvenient to deal with two name changes for a bacterium within one year (i.e. Ralstonia to Wautersia to Cupriavidus), in the long run, such reorganizations of the taxonomy of organisms are warranted as new data come to light.
Conforming to Rule 37a(1), that the name of a taxon must be changed if the nomenclatural type is excluded, we propose that the name Wautersia be replaced by Cupriavidus and that all species of the genus Wautersia be considered species of the genus Cupriavidus. We consequently propose to reclassify the remaining Wautersia species, i.e. Wautersia basilensis (Steinle et al. 1999) Vaneechoutte et al. 2004, Wautersia campinensis (Goris et al. 2001) Vaneechoutte et al. 2004, Wautersia gilardii (Coenye et al. 1999) Vaneechoutte et al. 2004, Wautersia metallidurans (Goris et al. 2001) Vaneechoutte et al. 2004, Wautersia oxalatica (Sahin et al. 2000) Vaneechoutte et al. 2004, Wautersia paucula (Vandamme et al. 1999) Vaneechoutte et al. 2004, Wautersia respiraculi (Coenye et al. 2003b) Vaneechoutte et al. 2004 and Wautersia taiwanensis (Chen et al. 2001) Vaneechoutte et al. 2004, into the genus Cupriavidus.
In contrast with the etymology presented by Makkar & Casida (1987), the gender of the genus name Cupriavidus is masculine [Rule 65(2)]. We therefore propose to reclassify the former Wautersia species as follows.
Emended description of the genus Cupriavidus
The description of the emended genus Cupriavidus is based on that presented by Makkar & Casida (1987) with some modifications. Cells are Gram-negative, peritrichously flagellated rods. Chemoheterotrophic or chemolithotrophic. The metabolism is oxidative. Several amino acids are used as sole carbon and nitrogen sources. Catalase and oxidase activity is produced. Resistance to various metals is widespread. The respiratory quinone Q8 has been reported in W. eutropha (Yabuuchi et al., 1995). The DNA G+C content is between 63 and 69 mol%. Species occur in soil and human clinical specimens, particularly in samples from debilitated patients. The type species is Cupriavidus necator.
Description of Cupriavidus basilensis comb. nov.
Cupriavidus basilensis [N.L. masc. adj. basilensis from Basilea (Basel), where the type strain was isolated].
Basonym: Ralstonia basilensis Steinle et al. 1999.
The description is identical to that given for Wautersia basilensis by Vaneechoutte et al. (2004). The type strain is LMG 18990T (=DSM 11853T).
Description of Cupriavidus campinensis comb. nov.
Cupriavidus campinensis (N.L. masc. adj. campinensis from the Kempen or Campine, the geographical region of north-east Belgium where this bacterium was initially isolated).
Basonym: Ralstonia campinensis Goris et al. 2001.
The description is identical to that given for Wautersia campinensis by Vaneechoutte et al. (2004). The type strain is LMG 19282T (=CCUG 44526T).
Description of Cupriavidus gilardii comb. nov.
Cupriavidus gilardii (N.L. gen. n. gilardii of Gilardi, named in honour of G. L. Gilardi, an American microbiologist).
Basonym: Ralstonia gilardii Coenye et al. 1999.
The description is identical to that given for Wautersia gilardii by Vaneechoutte et al. (2004). The type strain is LMG 5886T (=CCUG 38401T).
Description of Cupriavidus metallidurans comb. nov.
Cupriavidus metallidurans (N.L. masc. part. adj. metallidurans enduring metal, to indicate that these bacteria are able to survive high heavy-metal concentrations).
Basonym: Ralstonia metallidurans Goris et al. 2001.
The description is identical to that given for Wautersia metallidurans by Vaneechoutte et al. (2004). The type strain is LMG 1195T (=DSM 2839T).
Description of Cupriavidus oxalaticus comb. nov.
Cupriavidus oxalaticus (N.L. masc. adj. oxalaticus pertaining to oxalate).
Basonym: Ralstonia oxalatica (ex Khambata and Bhat 1953) Sahin et al. 2000.
The description is identical to that given for Wautersia oxalatica by Vaneechoutte et al. (2004). The type strain is LMG 2235T (=CCUG 2086T=DSM 1105T).
Description of Cupriavidus pauculus comb. nov.
Cupriavidus pauculus (L. masc. adj. pauculus rare, few, to indicate that these strains only sporadically cause human infections).
Basonym: Ralstonia paucula Vandamme et al. 1999.
The description is identical to that given for Wautersia paucula by Vaneechoutte et al. (2004). The type strain is LMG 3244T (=CCUG 12507T).
Description of Cupriavidus respiraculi comb. nov.
Cupriavidus respiraculi (L. gen. n. respiraculi of the respiratory system).
Basonym: Ralstonia respiraculi Coenye et al. 2003.
The description is identical to that given for Wautersia respiraculi by Vaneechoutte et al. (2004). The type strain is LMG 21510T (=CCUG 46809T).
Description of Cupriavidus taiwanensis comb. nov.
Cupriavidus taiwanensis (N.L. masc. adj. taiwanensis from Taiwan, where root-nodulating strains were isolated).
Basonym: Ralstonia taiwanensis Chen et al. 2001.
The description is identical to that given for Wautersia taiwanensis by Vaneechoutte et al. (2004). The type strain is LMG 19424T (=CCUG 44338T).
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ACKNOWLEDGEMENTS |
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REFERENCES |
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TOPABSTRACTMAIN TEXTREFERENCES |
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Chen, W.-M., Laevens, S., Lee, T.-M., Coenye, T., De Vos, P., Mergeay, M. & Vandamme, P. (2001). Ralstonia taiwanensis sp. nov., isolated from root nodules of Mimosa species and sputum of a cystic fibrosis patient. Int J Syst Evol Microbiol 51, 1729–1735.[Abstract]
Coenye, T., Falsen, E., Vancanneyt, M., Hoste, B., Govan, J. R. W., Kersters, K. & Vandamme, P. (1999). Classification of some Alcaligenes faecalis-like isolates from the environment and human clinical samples as Ralstonia gilardii sp. nov. Int J Syst Bacteriol 49, 405–413.[CrossRef][Medline]
Coenye, T., Goris, J., De Vos, P., Vandamme, P. & LiPuma, J. J. (2003a). Classification of Ralstonia pickettii-like isolates from the environment and various clinical samples as Ralstonia insidiosa sp. nov. Int J Syst Evol Microbiol 53, 1075–1080.
Coenye, T., Vandamme, P. & LiPuma, J. J. (2003b). Ralstonia respiraculi sp. nov., isolated from the respiratory tract of cystic fibrosis patients. Int J Syst Evol Microbiol 53, 1339–1342.
De Baere, T., Steyaert, S., Wauters, G., De Vos, P., Goris, J., Coenye, T., Suyama, T., Verschraegen, G. & Vaneechoutte, M. (2001). Classification of Ralstonia pickettii biovar 3/‘thomasii’ strains (Pickett 1994) and of new isolates related to nosocomial recurrent meningitis as Ralstonia mannitolytica sp. nov. Int J Syst Evol Microbiol 51, 547–558.[Abstract]
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