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1 Institut für Angewandte Mikrobiologie, Justus-Liebig Universität Giessen, IFZ–Heinrich-Buff-Ring 26-32, D-35392 Giessen, Germany
2 Grup d'Oceanografia Interdisciplinari, Institut Mediterrani d'Estudis Avançats (CSIC-UIB), Esporles, Mallorca, Spain
3 CCUG, Culture Collection University of Göteborg, Göteborg, Sweden
4 Institut für Bakteriologie, Mykologie und Hygiene, Veterinärmedizinische Universität, Wien, Austria
5 DSMZ, Mascheroder Weg 1b, D-38240 Braunschweig, Germany
Correspondence
Peter Kämpfer
peter.kaempfer{at}agrar.uni-giessen.de
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ABSTRACT |
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Phylogenetic trees constructed using the neighbour-joining and maximum-parsimony methods are available as supplementary figures in IJSEM Online.
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, 1994) for rRNA group 3 bacilli, mainly on the basis of 16S rRNA gene sequence data and the fact that this group was distinct from other groups which they defined within the bacilli. The genus description was emended by Shida et al. (1997) and, to date, the genus comprises more than 60 species, most of which were listed by Aguilera et al. (2001), Berge et al. (2002), Bosshard et al. (2002), Elo et al. (2001), Genersch et al. (2006), Iida et al. (2005), Kim et al. (2004), Meehan et al. (2001), Montes et al. (2004), Rivas et al. (2005a, b), Rodríguez-Díaz et al. (2005), Saha et al. (2005), Sánchez et al. (2005), 
merda et al. (2005), Takeda et al. (2005), von der Weid et al. (2002) and Yoon et al. (2003, 2005). The taxonomic dissection of the genus Bacillus (initiated by Ash et al., 1991), based largely on a 16S rRNA gene sequence perspective, has been summarized by Stackebrandt & Swiderski (2002).
Strain CCUG 47242T was isolated on blood agar at 37 °C during hygiene control checks at a starch-producing company in Sweden. Subcultivation was done on tryptone soy agar (TS agar; Oxoid) at 30 °C for 24 h. On this agar, strain CCUG 47242T was able to grow at 20–55 °C, but not at 10 or 60 °C. Growth at 30 °C was also observed on nutrient agar and R2A agar, but not on Salmonella–Shigella agar (SS agar; all media from Oxoid). Gram-staining was performed as described by Gerhardt et al. (1994). Cell morphology was observed under a Zeiss light microscope at x1000, using cells that had been grown for 24 h at 30 °C on TS agar. Details of cell morphology are given in the species description. The 16S rRNA gene sequence was analysed as described previously (Kämpfer et al., 2003). Analysis of the sequence data was performed by using MEGA version 2.1 (Kumar et al., 2001), after multiple alignment of sequences by CLUSTAL_X (Thompson et al., 1997). A distance-matrix method (distance options according to the Kimura two-parameter model) that used clustering with the neighbour-joining method (see Supplementary Fig. S1 in IJSEM Online), as well as a discrete character-based maximum-parsimony method (Supplementary Fig. S2 in IJSEM Online), were performed. In each case, bootstrap values were calculated based on 1000 replications. The 16S rRNA gene sequence of strain CCUG 47242T was a continuous stretch of 1486 bp. Sequence similarity calculations indicated that strain CCUG 47242T showed the greatest degree of similarity to ‘Paenibacillus hongkongensis’ HKU3 (GenBank accession no. AF433165; 96·58 %), which was described by Teng et al. (2003) (this name has not been validly published to date). Significantly lower sequence similarities (<94·5 %) were found with all other recognized species of the genus Paenibacillus.
Chemotaxonomic analyses were performed as follows: respiratory quinones (Tindall, 1990), polar lipids (Tindall, 1990) and fatty acids (Kämpfer & Kroppenstedt, 1996). Although respiratory quinones have low resolution within the bacilli, the presence of MK-7 supports the affiliation of strains CCUG 47242T and HKU3 to the genus Paenibacillus, where all species investigated to date have MK-7 as the major quinone. MK-7 represented 98 % of the quinones of strains CCUG 47242T and HKU3; in addition, small amounts of MK-6 (1 %) and MK-8 (1 %) were detected.
Strains CCUG 47242T and HKU3 displayed a complex polar lipid profile, which consisted of diphosphatidylglycerol, phosphatidylglycerol, phosphatidylethanolamine, lysyl-phosphatidylglycerol, two unknown phospholipids and four unknown aminophospholipids (Fig. 1).
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, showed a similar fatty acid content to strain CCUG 47242T with respect to C16 : 0 and C17 : 0, but not for the other fatty acids. Kämpfer (2002) compiled fatty acid data for paenibacilli and found that the major fatty acids of the genus Paenibacillus (with ranges as percentages of total given in parentheses) are anteiso-C15 : 0 (36–80 %), iso-C16 : 0 (0·5–6 %), iso-C15 : 0 (1–12 %) and anteiso-C17 : 0 (6·7 %).
Although some progress has been made in the reclassification of the bacilli, there is still some question as to what constitutes the genus Bacillus and whether other genera created over the years should be split further or combined with other genera. Redefining the genus Bacillus or the genus Paenibacillus would need extensive comparative work that would at present hinder the creation of novel taxa that may be closely associated with the genera Bacillus or Paenibacillus. This problem is clearly evident in a recent publication by Heyrman et al. (2005), where novel species within Bacillus group 2 are, by default, also placed in the genus Bacillus, although this group also contains members of the genera Kurthia, Caryophanon and Sporosarcina. The alternative is to define clearly the properties of the type species of the genera Bacillus and Paenibacillus and to create novel genera where clear differences can be found without having first to wait for extensive taxonomic rearrangements. This approach can be most effectively undertaken using a combination of 16S rRNA gene sequence data and chemical composition analysis. Such an approach leads to a rather different perspective of the potential taxonomic groupings than that based on 16S rRNA gene sequence data alone. It should be noted that features such as physiology, biochemistry and morphology may also be correlated with these data and all such data should be used to evaluate the taxonomic significance once groupings have been established. A clear distinction has to be made between groupings and genera. In the future, this may be expanded to include more detailed information from total genome analysis, although it is essential that such information should be coupled to other aspects such as biochemical pathways and structural proteins etc. (i.e. the epigenetic level).
It is well documented that the chemical composition of members of the bacilli is heterogeneous (Minnikin & Goodfellow, 1981; O'Leary & Wilkinson, 1988). However, in recent years, much emphasis has been put on the use of fatty acids. While some differentiation is possible, they appear to be of limited value in further differentiating within the bacilli. In the case of respiratory lipoquinones, menaquinones dominate, with isoprenoid chain lengths from 6 to 9. The most commonly encountered menaquinone is MK-7. Closer examination of polar lipids indicates a greater degree of variation, although this method has not been widely applied in recent years. In the case of members of Bacillus group 1 (Ash et al., 1991), there is chemical heterogeneity within the group (Minnikin & Goodfellow, 1981; O'Leary & Wilkinson, 1988). However, Bacillus subtilis, the type species of the genus Bacillus, produces diphosphatidylglycerol, phosphatidylglycerol, diphosphatidylethanolamine, an aminoacylphosphatidylglycerol and a glycolipid, which has been identified as a 
-gentiobiosyldiacylglycerol (Minnikin & Goodfellow, 1981). Species sharing this polar lipid pattern, together with MK-7 and the fatty acid pattern listed in Table 1, should, in the future, constitute the ‘core’ of the genus Bacillus. The polar lipid composition of the type species of the genus Paenibacillus, Paenibacillus polymyxa, is different. The single glycolipid present does not have the same Rf value as -gentiobiosyldiacylglycerol. While the major phospholipids present are diphosphatidylglycerol, phosphatidylglycerol and diphosphatidylethanolamine, they only serve (together with the presence of menaquinones and iso/anteiso fatty acids) to confirm that this species belongs to the bacilli. The presence of a number of unidentified phospholipids serves to differentiate this species. Preliminary work on the polar lipid composition of the genus Paenibacillus indicates that this group is also heterogeneous (S. Kirschner and B. J. Tindall, unpublished; Minnikin & Goodfellow, 1981; O'Leary & Wilkinson, 1988); only those species sharing a similar chemical composition to the type species should be retained in the genus Paenibacillus in the future.
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In particular, the lipid labelled LPG had the same Rf value as authentic lysyl-phosphatidylglycerol, which has been identified in Staphylococcus (Nahaie et al., 1984) and Listeria (Fischer & Leopold, 1999) species. The biosynthesis of this lipid is unusual in that lysine tRNA serves as the source of lysine (Gould & Lennarz, 1967, 1970; Lennarz et al., 1966). A specific enzyme has been identified that is responsible for the incorporation of lysine into the parent phospholipids (Oku et al., 2004). This lipid, and other amino acid derivatives of phospholipids, have also been found in other bacilli (Minnikin & Goodfellow, 1981; O'Leary & Wilkinson, 1988), as well as in members of the genera Enterococcus, Lactobacillus, Listeria, Vagococcus and Staphylococcus (Fischer & Leopold, 1999; Fischer & Arneth-Seifert, 1998; Nahaie et al., 1984; O'Leary & Wilkinson, 1988). The ability to synthesize such lipids (including other amino acid derivatives) must also be seen in a taxonomic and evolutionary context, being particularly prevalent in this branch of the Gram-positive bacteria. This study, and previous work, clearly indicates the value of chemotaxonomy within the bacilli and further supports the need for the inclusion of such analyses in all future taxonomic work on this group. This is also consistent with the often forgotten remarks of the ad hoc committee (Wayne et al., 1987), which emphasized the need to carry out more chemotaxonomic work.
The results of physiological characterization, using methods that have been described previously (Kämpfer et al., 1991), are given in the species descriptions. Strain CCUG 47242T was able to utilize many carbohydrates, but organic acids and amino acids were not utilized. In the case of HKU3, no positive results for carbon source utilization tests were reported (Teng et al., 2003). On the basis of the results presented, we propose that strains CCUG 47242T and HKU3 represent separate novel species of a new genus, Cohnella gen. nov.
Description of Cohnella gen. nov.
Cohnella (Coh.nel'la. N.L. fem. dim. n. Cohnella named after Ferdinand Cohn, the German microbiologist who first described the bacterial genus Bacillus in 1872).
Cells are Gram-positive, spore-forming, aerobic, non-motile, rod-shaped and thermotolerant. Good growth occurs after 24 h incubation on TS and nutrient agars at 25–30 °C; good growth also occurs at 55 °C. Main menaquinone is MK-7. Predominant polar lipids are diphosphatidylglycerol, phosphatidyglycerol, phosphatidylethanolamine and lysyl-phosphatidylglycerol. In addition, two unknown phospholipids and four unknown aminophospholipids are present. The major fatty acids are iso C16 : 0, anteiso C15 : 0 and C16 : 0. The type species is Cohnella thermotolerans.
Description of Cohnella thermotolerans sp. nov.
Cohnella thermotolerans (ther.mo.tol'er.ans. Gr. n. therme heat; L. pres. part. tolerans tolerating; N.L. part. adj. thermotolerans able to tolerate high temperatures).
Displays the following properties in addition to those given in the genus description. Oxidase-positive and shows an oxidative metabolism. The fatty acid profile of the type strain comprises iso C16 : 0 (45·5 %), anteiso C15 : 0 (28·4 %), anteiso C17 : 0 (6·7 %), C16 : 0 (6·6 %), C18 : 1
7c (4·0 %), iso C15 : 0 (3·2 %), iso C14 : 0 (2·1 %), C15 : 0 (1·4 %), iso C17 : 0 (1·1 %), C14 : 0 (1·0 %) and C17 : 16c (1·0 %). Aesculin and p-nitrophenol (pNP) -D-glucopyranoside are hydrolysed. Arbutin, L-arabinose (weakly), D-cellobiose, D-fructose, D-galactose, gluconate, D-glucose, D-maltose, D-mannose, 
-D-melibiose, L-rhamnose (weakly) and D-ribose are utilized as sole sources of carbon. No acid production from glucose, lactose, sucrose, D-mannitol, dulcitol, salicin, adonitol, inositol, sorbitol, L-arabinose, raffinose, rhamnose, maltose, D-xylose, trehalose, cellobiose, methyl D-glucoside, erythritol, melibiose, D-arabitol or D-mannose. No hydrolysis of o-NP -D-galactopyranoside, pNP -D-glucuronide, pNP -D-glucopyranoside, pNP -D-xylopyranoside, bis-pNP phosphate, pNP phenylphosphonate, pNP phosphorylcholine, 2-deoxythymidine-5'-pNP phosphate, L-alanine p-nitroanilide (pNA), L-glutamate-
-3-carboxy pna or L-proline pNA. The following carbon sources are not utilized: N-acetyl-D-galactosamine, N-acetyl-D-glucosamine, sucrose, salicin, D-trehalose, D-xylose, adonitol, i-inositol, maltitol, D-mannitol, D-sorbitol, putrescine, acetate, propionate, cis-aconitate, trans-aconitate, adipate, 4-aminobutyrate, azelate, citrate, fumarate, glutarate, DL-3-hydroxybutyrate, itaconate, DL-lactate, L-malate, mesaconate, oxoglutarate, pyruvate, suberate, L-alanine, -alanine, L-aspartate, L-histidine, L-leucine, L-ornithine, L-phenylalanine, L-proline, L-serine, L-tryptophan, 3-hydroxybenzoate, 4-hydroxybenzoate and phenylacetate. The DNA G+C content is 59 mol%.
The type strain, CCUG 47242T (=DSM 17683T=CIP 108492T), was isolated from a sample of industrial starch production in Sweden.
Description of Cohnella hongkongensis sp. nov.
Cohnella hongkongensis (hong.kong.en'sis. N.L. fem. adj. hongkongensis pertaining to Hong Kong).
The name ‘Paenibacillus hongkongensis’ has been used for this organism, but it has not been validly published and cannot be formally cited as a synonym.
In addition to the description given by Teng et al. (2003), the type strain utilizes several sole carbon sources after 7 days of incubation: N-acetyl-D-galactosamine, N-acetyl-D-glucosamine, arbutin, L-arabinose (weakly), D-cellobiose, D-fructose, D-galactose, gluconate, D-glucose, D-maltose, D-mannose, -D-melibiose, L-rhamnose and D-ribose. The following carbon sources are not utilized: sucrose, salicin, D-trehalose, D-xylose, adonitol, i-inositol, maltitol, D-mannitol, D-sorbitol, putrescine, acetate, propionate, cis-aconitate, trans-aconitate, adipate, 4-aminobutyrate, azelate, citrate, fumarate, glutarate, DL-3-hydroxybutyrate, itaconate, DL-lactate, L-malate, mesaconate, oxoglutarate, pyruvate, suberate, L-alanine, -alanine, L-aspartate, L-histidine, L-leucine, L-ornithine, L-phenylalanine, L-proline, L-serine, L-tryptophan, 3-hydroxybenzoate, 4-hydroxybenzoate and phenylacetate. The fatty acid profile of the type strain contains iso C16 : 0 (11·9 %), anteiso C15 : 0 (31·2 %), anteiso C17 : 0 (2·6 %), C16 : 0 (25·3 %), iso C15 : 0 (8·1 %), anteiso C13 : 0 (0·8 %), iso C14 : 0 (2·3 %), C15 : 0 (8·0 %), C16 : 111c (0·9 %), iso C17 : 0 (1·9 %), C14 : 0 (5·0 %) and C17 : 0 (1·2 %). The DNA G+C content is 60·9 mol% (method according to Peña et al., 2005).
The type strain is HKU3T (=DSM 17642T=CIP 107898T=CCUG 49571T).
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R. A. Albert, J. Archambault, M. Lempa, B. Hurst, C. Richardson, S. Gruenloh, M. Duran, H. L. Worliczek, B. E. Huber, R. Rossello-Mora, et al. Proposal of Viridibacillus gen. nov. and reclassification of Bacillus arvi, Bacillus arenosi and Bacillus neidei as Viridibacillus arvi gen. nov., comb. nov., Viridibacillus arenosi comb. nov. and Viridibacillus neidei comb. nov. Int J Syst Evol Microbiol, December 1, 2007; 57(12): 2729 - 2737. [Abstract] [Full Text] [PDF]
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E.-A. Cho, J.-S. Lee, K. C. Lee, H.-C. Jung, J.-G. Pan, and Y.-R. Pyun Cohnella laeviribosi sp. nov., isolated from a volcanic pond Int J Syst Evol Microbiol, December 1, 2007; 57(12): 2902 - 2907. [Abstract] [Full Text] [PDF]
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I. Ahmed, A. Yokota, A. Yamazoe, and T. Fujiwara Proposal of Lysinibacillus boronitolerans gen. nov. sp. nov., and transfer of Bacillus fusiformis to Lysinibacillus fusiformis comb. nov. and Bacillus sphaericus to Lysinibacillus sphaericus comb. nov. Int J Syst Evol Microbiol, May 1, 2007; 57(5): 1117 - 1125. [Abstract] [Full Text] [PDF]
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