International Journal of Systematic and Evolutionary Microbiology (2015), 65, 1860 – 1865 DOI 10.1099/ijs.0.000186 Chryseobacterium shandongense sp. nov., isolated from soil Fan Yang,† Hong-ming Liu,† Rong Zhang, Ding-bin Chen, Xiang Wang, Shun-peng Li and Qing Hong Correspondence Qing Hong [email protected] Key Laboratory of Agricultural Environmental Microbiology, Ministry of Agriculture, College of Life Sciences, Nanjing Agricultural University, Nanjing, Jiangsu 210095, PR China YF-3T is a Gram-stain-negative, non-motile, non-spore-forming, yellow – orange, rod-shaped bacterium. Optimal growth conditions were at 30 8C, pH 7.0 and 1 % (w/v) NaCl. Phylogenetic analysis, on the basis of the 16S rRNA gene sequence, showed that strain YF-3T was closely related to the strains Chryseobacterium hispalense AG13T and Chryseobacterium taiwanense Soil-3-27T with 98.71 % and 96.93 % sequence similarity, respectively. Strain YF-3T contained MK-6 as the main menaquinone and had a polyamine pattern with sym-homospermidine as the major component. Its major polar lipid was phosphatidylethanolamine. The dominant fatty acids of strain YF-3T were iso-C15 : 0, iso-C17 : 0 3-OH, summed feature 9 (comprising iso-C17 : 1v9c and/or C16 : 0 10-methyl) and summed feature 3 (comprising C16 : 1v7c and/or C16 : 1v6c). The DNA G þ C content of strain YF-3T was 37 mol%. The DNA –DNA relatedness levels between strain YF-3T and the most closely related strains, C. hispalense AG13T and C. taiwanense Soil-3-27T, were 31.7 ^ 2.1 % and 28.4 ^ 5.4 %, respectively. Based on these results, a novel species named Chryseobacterium shandongense sp. nov. is proposed. The type strain is YF-3T ( ¼ CCTCC AB 2014060T ¼ JCM 30154T). The genus Chryseobacterium represents a genera with one of the fastest growing number of species (Herzog et al., 2008), which are found in a wide variety of environments. The genus Chryseobacterium was first described by Vandamme et al. (1994) and at the time of writing it contains 88 species with validly published names (http://www. bacterio.net/). In 1994 – 2000, the genus only comprised six species: Chryseobacterium balustinum, Chryseobacterium gleum, Chryseobacterium indologenes, Chryseobacterium indoltheticum, Chryseobacterium meningosepticum and Chryseobacterium scophthalmum. However, the number of species increased by 45 and 33 between 2001 and 2010, and 2011 and 2014, respectively. These newly discovered members of the genus Chryseobacterium are distributed in a variety of environments, such as roots (Park et al., 2006), a lake (Joung & Joh, 2011), clinical samples (Vaneechoutte et al., 2007), soil (Li & Zhu, 2012), sludge (Pires et al., 2010), raw milk (Hantsis-Zacharov & Halpern, 2007), the midgut of insects (Kampfer et al., 2010a), food products (including raw cow’s milk, fish, poultry and lactic acid beverages) (Hantsis-Zacharov et al., 2008) and †These authors contributed equally to this work. The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene sequence of strain YF-3T is KJ644318. Four supplementary figures and one supplementary table are available with the online Supplementary Material. 1860 human clinical sources (Yassin et al., 2010). In this study, strain YF-3T was isolated from soil from Qingdao, Shandong province, China (358 359 –378 099 N 1198 309– 1218 009 E). During the isolation of strains, 10 g of soil was added to a flask containing 100 ml trypticase soy broth (TSB; 17.0 g pancreatic digest of casein, 3.0 g soy peptone, 2.5 g dextrose, 5.0 g NaCl, 2.5 g K2HPO4; pH 7.0 ^ 0.2) and incubated at 30 8C at 150 r.p.m. for 2 days. Trypticase soy agar (TSA; per litre distilled water: 17.0 g pancreatic digest of casein, 3.0 g soy peptone, 2.5 g dextrose, 5.0 g NaCl, 2.5 g K2HPO4, 15.0 g agar; pH 7.0 ^ 0.2) plates were spread with 0.1 ml diluted soil suspension and incubated at 30 8C for 2 days. A yellow 2 orange colony was selected, purified and then the isolate was cultivated at 30 8C on the same medium and preserved in 20 % (v/v) glycerol at 2 80 8C. Strain YF-3T was cultivated on TSA plates at 30 8C for 2 days. The presence of flexirubin-type pigments was investigated by noting whether a colour shift occurred when the colony was flooded with 20 %(w/v) KOH (Fautz & Reichenbach, 1980). Gram staining was performed by the modified method of Gerhardt et al. (1994). Cell motility was determined according the procedure described by Smibert & Krieg (1994). A colony of strain YF-3T was picked from the last quadrant streak after it was grown on TSA at 30 8C for 24 h. Then the cell morphology and dimensions were determined by Downloaded from www.microbiologyresearch.org by 000186 G 2015 IUMS IP: 88.99.165.207 On: Mon, 19 Jun 2017 05:16:41 Printed in Great Britain Chryseobacterium shandongense sp. nov. transmission electron microscopy (H-7650; Hitachi) (Fig. S1, available in the online Supplementary Material). Growth at different temperatures (4, 20, 25, 30, 37 and 42 8C) and pH 4 –10 (at 1 pH unit intervals) was assessed in TSB, the pH range was buffered with citrate/phosphate buffer or Tris/HCl buffer (Breznak & Costilow, 1994). Salt tolerance was investigated on TSB supplemented with 0 – 9 % (w/v) NaCl (at 1 % intervals). The OD600 value was determined after 3 days of incubation in order to evaluate the growth of strain YF-3T. In addition, growth on Luria – Bertani (LB; 10.0 g peptone, 5.0 g yeast extract, 10.0 g NaCl, 15.0 g agar; pH 7.0 ^ 0.2) agar, R2A (0.5 g proteose peptone, 0.5 g yeast extrct, 0.5 g casamino acids, 0.5 g glucose, 0.5 g soluble starch, 0.3 g sodium pyruvate, 0.3 g K2HPO4, 0.05 g MgSO4.7H2O, 15.0 g agar; pH 7.0 ^ 0.2) agar, nutrient agar (NA; 5.0 g peptone, 3.0 g beef extract, 5.0 g NaCl, 15.0 g agar; pH 6.8 ^ 0.2), cetrimide agar (CA; 20.0 g pancreatic digest of gelatin, 1.4 g MgCl, 10.0 g K2SO4, 0.3 g cetrimide, 15.0 g agar; pH 7.2 ^ 0.2), Simmons’ citrate (SC; 5.0 g NaCl, 2.0 g sodium citrate, 1.0 g NH4H2PO4, 1.0 g K2HPO4, 0.2 g MgSO4, 0.08 g bromothymol blue, 15.0 g agar; pH 6.9 ^ 0.2) agar and MacConkey agar (17.0 g peptone, 3.0 g proteose tryptone, 10.0 g lactose, 1.5 g bile salts, 5.0 g NaCl, 0.001 g crystal violet, 0.03 g neutral red, 13.5 g agar; pH 7.1 ^ 0.2) was also evaluated. Catalase activity was determined by bubble production with 3 % (v/v) H2O2. Oxidase activity was assayed using filter-paper discs (grade 388; Sartorius) impregnated with a 1 % (w/v) solution of N,N,N9,N9-tetramethyl-p-phenylenediamine (Sigma-Aldrich). A positive result was indicated by the development of a blue – purple colour after applying biomass to the filter paper. Antibiotic susceptibility tests were performed with the disc (Hangzhou Tianhe Microbial Reagent) diffusion method on TSA plates incubated at 30 8C for 2 days. Strains were considered sensitive when the diameter of the inhibition zone was $ 10 mm (Jorgensen & Ferraro, 2009). The hydrolysis of DNA, cellulose, starch and casein were investigated as described by Smibert & Krieg (1994). Basic biochemical, enzyme activity and carbon source tests were performed using the API 20NE, API 20E and API ZYM systems (bioMerieux), and GN2 MicroPlates (Biolog), according to the manufacturers’ instructions. For the analysis of whole-cell fatty acids, strain YF-3T and the reference strains, Chryseobacterium hispalense AG13T and Chryseobacterium taiwanense Soil-3-27T, were grown on TSA at 30 8C for 24 h. The biomass, which was always harvested from the same sector (the last quadrant streak) was freeze-dried, then the fatty acid methyl esters were extracted according to the standard procedure of the Microbial Identification System (MIDI) (Sasser, 1990). Extracts were analysed using a Hewlett Packard model 6890 gas chromatograph equipped with a flame-ionization detector (Kampfer & Kroppenstedt, 1996) and a 5 % phenyl-methyl-silicone capillary column. The extraction of the respiratory quinones was carried out from freezedried cell material according to the method of http://ijs.sgmjournals.org Collins et al. (1977) and determined by HPLC (Tamaoka et al., 1983). Polar lipids were extracted from 100 mg of freeze-dried cell material using a chloroform/methanol/ 0.3 % (w/v) aqueous NaCl mixture 1 : 2 : 0.8 (by vol.) (Tindall et al., 2007). Then polar lipids were recovered into the chloroform phase by adjusting the chloroform/methanol/ 0.3 % aqueous NaCl (w/v) mixture to a ratio of 1 : 1 : 0.9 (by vol.). Polar lipids were separated by two-dimensional silica gel TLC. Total lipid material was detected using molybdophosphoric acid and specific functional groups were detected using spray reagents specific for defined functional groups (Tindall et al., 2007) (DSMZ service) (Fig. S2). Cells of strain YF-3T used for polyamine analysis were grown on TSB, harvested at the late exponential growth phase and lyophilized. Extraction of polyamines was performed as described by Busse & Auling (1988) and analysis was conducted using the HPLC equipment described by Stolz et al. (2007). The genomic DNA of strain YF-3T was extracted and purified according to the method described by Sambrook & Russell, (2001) and the DNA G þ C content was determined by reversed-phase HPLC (Tamaoka & Komagata, 1984) using Escherichia coli K-12 as a standard. DNA – DNA hybridizations were performed at 50 8C, with photobiotin-labelled probes in microplate wells, as described by Ezaki et al. (1989). A bioassay plate reader (HTS 7000; Perkin Elmer) was used to measure the fluorescence, and reciprocal experiments were performed on each pair of strains investigated. The 16S rRNA gene of strain YF-3T was amplified with the bacterial universal primers 27F and 1492R (Lane, 1991). The PCR products were purified using an AxyPrep PCR Purification kit (AxyGen) and were cloned into a pMD 19-T Vector. The purified plasmid DNA was sequenced with an automated sequencer (model 3730; Applied Biosystems). Pairwise sequence similarity was calculated with the known sequences using the EzTaxon-e server (http://eztaxon-e.ezbiocloud.net/; Kim et al., 2012). Phylogenetic analysis was performed by using MEGA version 5.0 (Tamura et al., 2011) after multiple alignments of data using CLUSTAL_X (Thompson et al., 1997). Distances were determined through distance options based on Kimura’s two-parameter system (Kimura, 1980). Unrooted trees were reconstructed via neighbour-joining (Saitou & Nei, 1987) (Fig. 1), maximum-parsimony (Fitch, 1971) (Fig. S3) and maximum-likelihood (Felsenstein, 1981) (Fig. S4) methods. Confidence values for the branches of phylogenetic trees were calculated according to bootstrap analyses (based on 1000 resamplings) (Felsenstein, 1985). YF-3T was Gram-stain-negative, rod-shaped (0.5 –0.7 mm in width, 2.1 – 2.3 mm in length), non-spore-forming and non-motile. Colonies of strain YF-3T on TSA plates were yellow –orange, circular, convex, smooth, translucent and shiny, and the colonies were not visible as single entities after prolonged incubations. The optimal growth conditions for YF-3T were 30 8C, 1 % (w/v) NaCl and pH 7.0 in TSB. Good growth also occurred on LB agar and R2A Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Mon, 19 Jun 2017 05:16:41 1861 F. Yang and others C. gwangjuense THG-A18T (JN196134) 85* C. geocarposphaerae 91A-561T (HG738132) * 0.01 C. defluvii B2T (AJ309324) 99* * ‘C. yeoncheonense’ DCY67 (JX141782) C. aahli T68T (JX287893) * ‘C. massiliae’ 90B (AF531766) * C. gambrini 5-1St1aT (AM232810) C. daecheongense CPW 406T (AJ457206) * C. wanjuense R2A10-2T (DQ256729) * * C. taiwanense BCRC 17412T (DQ318789) C. gregarium P 461/12T (AM773820) * C. hagamense RHA2-9T (DQ673672) C. camelliae THG C4-1T (JX843771) 71* Chryseobacterium shandongense YF-3T (KJ644318) 98* C. hispalense AG13T (EU336941) C. taeanense PHA3-4T (AY883416) 96* * C. taichungense CC-TWGS1-8T (AJ843132) C. vietnamense GIMN1.005T (HM212415) 100 C. gleum ATCC 35910T (ACKQ01000057) C. arthrosphaerae CC-VM-7T (FN398101) 86* C. indologenes LMG 8337T (AM232813) C. nakagawai NCTC 13529T (JX100822) 98* C. lactis NCTC 11390T (JX100821) C. viscerum 687B-08T (FR871426) Ornithobacterium rhinotracheale LMG 9086T (L19156) Fig. 1. Neighbour-joining phylogenetic tree according to 16S rRNA gene sequences indicating the relationship of strain YF3T to closely related species of the genus Chryseobacterium. Asterisks indicate branches that were also recovered using the maximum-parsimony and maximum-likelihood algorithms. Bootstrap values of more than 70 % (based on 1000 replications) are shown at branching points. The 16S rRNA gene sequence of Ornithobacterium rhinotracheale LMG 9086T was used as an outgroup. Bar, 0.01 substitutions per nucleotide position. agar; weak growth occurred on NA; no growth occurred on CA, SC agar or MacConkey agar. YF-3T was resistant to tenebrimycin, streptomycin, amikacin, oxacillin, kanamycin, aztreonam and gentamicin, but sensitive to chloromycetin, cefotaxime, spectinomycin, minocycline, levofloxacin, cefuroxime, cefoperazone, cefoxitin, norfloxacin, furadantin, ciprofloxacin, midecamycin, polymyxin B, vancomycin, ofloxacin, erythromycin, clindamycin, tetracycline, benzylpenicillin, cefazolin, cefepime, ampicillin, ceftriaxone, trimethoprim-sulfamethoxazole, ceftazidime, cefalotin and piperacillin. The morphological, cultural, physiological and biochemical characteristics of strain YF-3T are listed in the species description. The differences between strain YF-3T and the reference strains (C. hispalense AG13T and C. taiwanense Soil-3-27T) are presented in Table 1. The predominant fatty acids of strain YF-3T ($ 5 %) were iso-C15 : 0, iso-C17 : 0 3-OH, summed feature 9 (comprising iso-C17 : 1v9c and/or C16 : 0 10-methyl) and summed feature 3 (comprising C16 : 1v7c and/or C16 : 1v6c), which was 1862 consistent with those of its closest phylogenetic neighbours grown under the same conditions. Smaller amounts of anteiso-C15 : 0 and iso-C15 : 0 3-OH were also present. The detailed fatty acid composition of strain YF-3T is shown in Table 2 where it is compared with that of the reference strains C. hispalense AG13T and C. taiwanense Soil-3-27T. The polar lipid profile of strain YF-3T consisted of the predominant compounds phosphatidylethanolamine, five unknown lipids and two unknown aminolipids. The main respiratory quinone was menaquinone MK-6. Polyamine analysis indicated that sym-homospermidine [43.2 mmol (g dry weight)21] was the major component and that minor amounts of spermidine [3.1 mmol (g dry weight)21] and spermine [2.8 mmol (g dry weight)21] and traces of 1,3-diaminopropane, cadaverine and putrescine [, 0.1 mmol (g dry weight)21] were also present, which is consistent with the characteristics of other members of the genus Chryseobacterium (Hamana & Matsuzaki, 1990; Kampfer et al., 2003). The DNA G þ C content of strain YF-3T was 37 mol%. Downloaded from www.microbiologyresearch.org by International Journal of Systematic and Evolutionary Microbiology 65 IP: 88.99.165.207 On: Mon, 19 Jun 2017 05:16:41 Chryseobacterium shandongense sp. nov. Table 1. Differential biochemical characteristics of strain YF-3T and type strains of closely related species of the genus Chryseobacterium T Strains: 1, Chryseobacterium shandongense sp. nov. YF-3 ; 2, C. hispalense AG13T; 3, C. taiwanense BCRC 17412T. All data are from this study. þ , Positive; 2, negative; W , weakly positive. Characteristic API 20 NE strips Reduction of nitrate L -Malic acid b-Glucosidase Utilization of (GN 2 plate): L -Serine Histidine Dextrin D -Fructose L -Fucose D -Galactose Gentiobiose a-D -Glucose Maltose D -Mannose L -Rhamnose D -Sorbitol D -Glucuronic Acid Inosine L -Proline Glycyl-L -aspartic acid Turanose Enzymic activities (API ZYM strip) Esterase (C4) Esterase lipase (C8) Lipase (C14) Trypsin a-Chymotrypsin N-Acetyl-b-glucosaminidase Cystine aminopeptidase 1 2 Strains: 1, Chryseobacterium shandongense sp. nov. YF-3T; 2, C. hispalense AG13T; 3, C. taiwanense BCRC 17412T. All data are from this study. Values are percentages of total fatty acids. Fatty acids amounting to less than 1 % of the total fatty acids are not shown. TR , Trace (less than 1 %); 2 , not detected/reported. 3 Fatty acid 2 2 þ þ þ 2 2 2 2 þ þ þ þ þ 2 þ þ þ 2 2 þ 2 þ þ þ þ 2 2 þ þ þ þ þ þ þ þ þ 2 þ 2 2 þ þ þ 2 2 2 2 2 2 2 2 2 2 2 2 þ 2 2 2 þ þ þ 2 2 2 2 2 þ 2 2 þ 2 2 2 2 2 W W Þ W A nearly full-length 16S rRNA gene sequence (1478 bp) of strain YF-3T was determined. Similarity analysis of 16S rRNA gene sequences showed that strain YF-3T was most closely related to C. hispalense AG13T (98.71 %) and C. taiwanense Soil-3-27T (96.93 %). Phylogenetic analysis based on sequence similarities of the 16S rRNA gene indicated there was a relationship between strain YF-3T and members of the genus Chryseobacterium (Fig. 1). DNA – DNA hybridization with its closest relatives was carried out to determine further the taxonomic status of YF-3T. In the present study, the levels of DNA –DNA relatedness between strain YF-3T and C. hispalense AG13T and C. taiwanense Soil3-27T were found to be 31.7 ^ 2.1 % and 28.4 ^ 5.4 %, respectively (Table S1), far below the value of 70 % that is commonly accepted to define a novel species (Wayne et al., 1987). http://ijs.sgmjournals.org Table 2. Fatty acid compositions of strain YF-3T and closely related species of the genus Chryseobacterium C16 : 0 iso-C15 : 0 iso-C17 : 0 anteiso-C15 : 0 iso-C15 : 0 3-OH iso-C17 : 0 3-OH C16 : 0 3-OH C18 : 1v9c C20 : 1v9c Summed features: 3* 9† 1 2 3 1.44 47.06 1.31 42.48 1.96 44.17 1.17 TR TR 2.85 2.74 15.06 1.33 1.04 1.71 1.93 2.68 13.59 1.02 2 2 2 5.57 13.76 10.54 19.25 9.47 16.16 TR TR 4.31 17.92 TR *Summed feature 3 contains C16 : 1v7c and/or C16 : 1v6c. †Summed feature 9 contains iso-C17 : 1v9c and/or C16 : 0 10-methyl. Therefore, on the basis of these phylogenetic, phenotypic and chemotaxonomic data, strain YF-3T represents a novel species of the genus Chryseobacterium, for which the name Chryseobacterium shandongense sp. nov. is proposed. Description of Chryseobacerium shandongense sp. nov. Chryseobacterium shandongense (shan.dong.en9se. N.L. neut. adj. shandongense pertaining to Shandong province, the location of the soil sample from which the type strain was isolated). Cells are Gram-stain-negative, non-motile, non-sporeforming, rod-shaped, and approximately 0.5 – 0.7 mm in width and 2.1 – 2.3 mm in length. Best growth occurs on TSA; good growth occurs on LB agar and R2A agar; weak growth occurs on NA; no growth occurs on CA, SC agar or MacConkey agar. Colonies grown for 24 h on TSA are about 3 mm in diameter, circular with a shiny surface and entire edges, yellow – orange (flexirubin-type, nondiffusible), translucent and mucoid. Growth conditions are 25 – 37 8C (optimum 30 8C), at pH 5.0 –8.0 (optimum 7.0), with 0 –5 % (w/v) NaCl (optimum 1 %). Positive for the hydrolysis of DNA, starch and casein and for oxidase and catalase activities. Negative for the reduction of nitrate and cellulose hydrolysis. In API 20E and 20NE kits, positive for citrate utilization, acetoin production, indole production, aesculin hydrolysis and gelatin hydrolysis, but negative for arginine dihydrolase, urease, lysine Downloaded from www.microbiologyresearch.org by IP: 88.99.165.207 On: Mon, 19 Jun 2017 05:16:41 1863 F. Yang and others decarboxylase, ornithine decarboxylase, H2S production and tryptophan deaminase. Acid is produced from D -mannose, but not from mannitol, inositol, sorbitol, melibiose, amygdalin, arabinose, N-acetylglucosamine or potassium gluconate. In GN2 Microplates, utilization of dextrin, glycogen, Tween 40, Tween 80, adonitol, i-erythritol, D -fructose, L -fucose, gentiobiose, a-D -glucose, a-lactose, lactulose, maltose, D -psicose, raffinose, L -rhamnose, sucrose, turanose, xylitol, monomethyl succinate, acetic acid, cis-aconitic acid, D -galactonic acid, lactone, D -galacturonic acid, D -glucosaminic acid, a-hydroxybutyric acid, b-hydroxybutyric acid, p-hydroxyphenylacetic acid, aketobutyric acid, DL -lactic acid, malonic acid, propionic acid, quinic acid, succinic acid, L -alaninamide, L -alanine, L -asparagine, L -aspartic acid, L -glutamic acid, glycyl-L aspartic acid, glycyl-L -glutamic acid, L -proline, L -pyroglutamic acid, inosine, thymidine, 2,3-butanediol, DL -a-glycerol, glucose 1-phosphate, glucose 6-phosphate and N-acetyl-D glucosamine is positive, but not utilization of a-cyclodextrin, N-acetyl-D -galactosamine, L -arabinose, D -arabitol, cellobiose, D -galactose, myo-inositol, methyl b-D -glucoside, trehalose, methyl pyruvate, citric acid, formic acid, D gluconic acid, D -glucuronic acid, c-hydroxybutyric acid, itaconic acid, a-ketoglutaric acid, a-ketovaleric acid, D saccharic acid, sebacic acid, succinamic acid, glucuronamide, D -alanine, L -alanyl-glycine, L -phenylalanine, D serine, L -threonine, DL -carnitine, c-aminobutyric acid, phenyethylamine, putrescine or 2-aminoethanol. In API ZYM tests, alkaline phosphatase, esterase (C4), esterase lipase (C8), lipase (C14), leucine aminopeptidase, valine aminopeptidase, acid phosphatase, naphthol-AS-BI-phosphoamidase, a-glucosidase, N-acetyl-b-glucosaminidase, cystine aminopeptidase (weak), trypsin (weak) and a-chymotrypsin (weak) activities are present, but a-galactosidase, b-galactosidase, b-glucuronidase, a-mannosidase and a-fucosidase activities are absent. Menaquinone-6 is the main respiratory quinone. The predominant fatty acids ($ 5 %) are iso-C15 : 0, iso-C17 : 0 3-OH, summed feature 9 (comprising iso-C17 : 1v9c and/or C16 : 0 10-methyl) and summed feature 3 (comprising C16 : 1v7c and/or C16 : 1v6c). Polar lipids consist of phosphatidylethanolamine, five unidentified lipids and two unidentified aminolipids. sym-Homospermidine is the predominant polyamine, but minor amounts of spermidine and spermine are also present. The type strain is YF-3 ( ¼ CCTCC AB 2014060 ¼ JCM 30154T), which was isolated from farmland soil collected from Qingdao city, Shandong province, PR China. The DNA G þ C content of the type strain is 37 mol%. T T References Breznak, J. A. & Costilow, R. N. (1994). Physicochemical factors in growth. In Methods for General and Molecular Bacteriology, pp. 137 –154. Edited by P. Gerhardt, R. G. E. Murray, W. A. Wood & N. R. Krieg. Washington, DC: American Society for Microbiology. Busse, H. J. & Auling, G. (1988). Polyamine pattern as a chemo- taxonomic marker within the Proteobacteria. Syst Appl Microbiol 11, 1– 8. Collins, M. D., Pirouz, T., Goodfellow, M. & Minnikin, D. E. (1977). Distribution of menaquinones in actinomycetes and corynebacteria. J Gen Microbiol 100, 221 –230. Ezaki, T., Hashimoto, Y. & Yabuuchi, E. (1989). 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