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volume3-tables.csv
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#####
TABLE 1. Phenotypic characteristics of genera belonging to the family Bacillaceae a
1. Bacillus 2. Alkalibacllus 3. Amphibacillus 4. Anoxybacillus 5. Cerasibacillus 6. Filobacillus 7. Geobacillus 8. Gracilibacillus 9. Halobacillus 10. Halolactibacillus 11. Lentibacillus 12. Marinococcus 13. Oceanobacillus 14. Paraliobacillus 15. Pontibacillus 16. Saccharococcus 17. Tenuibacillus 18. Thalassobacillus 19. Virgibacillus
Number of species in genus 141 4 3 10 1 1 17 4 4 2 4 3 3 1 2 1 1 1 9
Gram reaction +/v/− + + + + − +/− + + + v + + + + + + + +
Predominant cell shape:
Rods + + + + + + + + + + + + + + + + +
Cocci + + +
Mean cell width:
<0.5 mm + + + + + +
0.5–1.0 mm + + + + + + + + + + + + +
1.0 or >1.0 mm + + + + + + + +
Motility +/− + + +/− +/− + +/− +/− + + + + − + + +
Spore formation +/− + + + + + + + + − + − + + + − + + +
Spore shape:
Ellipsoidal + + + + + + + + + + +
Cylindrical + + +
Spherical + + + + + + + + + + + +
Oxygen requirements:
Aerobes + + + + + + + + + + + + + + + + + + +
Facultative anaerobes + − + + − + + − + − − + + − − − +
Strict anaerobes +
Growth in media with added NaCl:
0% + − + + − + v − + − v + + − + − − v
5% + + + + + + + + + + + + + + + + + +
10% + + − − + + + + + + + + + + + + +
20% + + + + + + + + + + − + + +
Growth at:
10 °C + + + + + + +
20 °C + + + + + + + + + + + +
30 °C + + + + + + + + + + + + + + + + +
40 °C + + + + + + + + + + + + + + +
50 °C + + + + + + + + +
60 °C + − + + + +
Growth at pH:
5 + − + + +
6 + − + + + + + + + + + + + +
7 + + + + + + + + + + + + + + + + + +
8 + + + + + + + + + + + + + + + +
9 + + + + + + + + + + + + + +
10 + + + + + + + + + +
Catalase +/− + − +/− + + +/− + + − + + + + + + + + +
Oxidase +/− − − +/− − v + − v − v +/− + + − +
footnote a: Symbols: +, at least one species within the genus gives a positive reaction; +/−, some species are positive, some are negative; v, varies within the genus; −, negative; w, weak.
#####
TABLE 2. Murein cross-linkage types found in Bacillus species and in former Bacillus species that have been transferred to other genera
Murein cross-linkagea Reference
Bacillus
B. subtilis meso-DAP direct Schleifer and Kandler (1972)
B. anthracis (meso-DAP direct) Schleifer and Kandler (1972)
B. aquimaris meso-DAPb Yoon et al. (2003a)
B. barbaricus DAPb Taubel et al. (2003)
B. badius meso-DAP direct Schleifer and Kandler (1972)
B. cereus meso-DAP direct Schleifer and Kandler (1972)
B. coagulans meso-DAP direct Schleifer and Kandler (1972)
B. fastidiosus meso-DAP direct Claus and Berkeley (1986)
B. firmus (meso-DAP direct) Schleifer and Kandler (1972)
B. funiculus DAPb Ajithkumar et al. (2002)
B. halophilus meso-DAP direct Ventosa et al. (1989)
B. hwajinpoensis meso-DAPb Yoon et al. (2004b)
B. horti meso-DAPb Yumoto et al. (1998)
B. indicusc l-Orn-d-Asp Suresh et al. (2004)
B. jeotgali meso-DAP direct Yoon et al. (2001a)
B. lentus (meso-DAP direct) Schleifer and Kandler (1972)
B. licheniformis meso-DAP direct Schleifer and Kandler (1972)
B. marisflavi meso-DAPb Yoon et al. (2003a)
B. megaterium (meso-DAP direct) Schleifer and Kandler (1972)
B. methanolicus meso-DAP direct Arfman et al. (1992)
B. mycoides meso-DAP direct Claus and Berkeley (1986)
B. oleronius meso-DAP direct Kuhnigk et al. (1995)
B. pumilus meso-DAP direct Schleifer and Kandler (1972)
B. schlegelii meso-DAP direct Krüger and Meyer (1984)
B. smithii DAPb Nakamura et al. (1988)
B. thermocloacae meso-DAP direct Demharter and Hensel (1989b)
B. thuringiensis meso-DAP direct Schleifer and Kandler (1972)
B. vietnamensis meso-DAPb Noguchi et al. (2004)
Alkaliphilic and alkalitolerant Bacillus species
B. cohnii l-Orn-d-Asp Spanka and Fritze (1993)
B. halmapalus No DAP Nielsen et al. (1994)
Alkaliphilic species in 6th 16S rRNA group
of Nielsen et al. (1994)
B. horikoshii No DAP Nielsen et al. (1994)
Spherical-spored Bacillus species
B. fusiformisd l-Lys-d-Asp Ahmed et al. (2007c)
B. insolitus Orn-d-Glu Stackebrandt et al. (1987)
B. neidei l-Lys-d-Glu Nakamura et al. (2002)
B. psychrodurans Orn-d-Glu Abd El-Rahman et al. (2002)
B. psychrotolerans Orn-d-Glu Abd El-Rahman et al. (2002)
B. pycnus l-Lys-d-Glu Nakamura et al. (2002)
B. silvestris l-Lys-d-Glu Rheims et al. (1999)
B. sphaericusd l-Lys-d-Asp Schleifer and Kandler (1972)
Alkalibacillus
A. haloalkaliphilus meso-DAP direct Fritze (1996b)
Brevibacillus
Br. Brevis meso-DAP direct Schleifer and Kandler (1972)
Br. laterosporus meso-DAP direct Schleifer and Kandler (1972)
Geobacillus
G. stearothermophilus (meso-DAP direct) Schleifer and Kandler (1972)
G. thermoleovorans DAPb Zarilla and Perry (1987)
G. pallidus meso-DAP direct Scholz et al. (1987)
Gracilibacillus
Gr. dipsosauri meso-DAP direct Lawson et al. (1996)
Marinibacillus
M. marinus l-Lys-direct Yoon et al. (2001b)
Paenibacillus
P. polymyxa (meso-DAP direct) Schleifer and Kandler (1972)
P. alvei meso-DAP direct Schleifer and Kandler (1972)
P. amylolyticuse (meso-DAP direct) Schleifer and Kandler (1972)
P. lentimorbus meso-DAP direct Schleifer and Kandler (1972)
P. macerans meso-DAP direct Schleifer and Kandler (1972)
Sporolactobacillus
S. laevolacticus meso-DAP direct Andersch et al. (1994)
Sporosarcina
S. ureae l-Lys-Gly-d-Glu Stackebrandt et al. (1987)
S. globisporus l-Lys-d-Glu Stackebrandt et al. (1987)
S. psychrophilus l-Lys-d-Glu Stackebrandt et al. (1987)
S. pasteurii l-Lys-d-Asp Ranftl and Kandler (1973)
Ureibacillus
U. thermosphaericus l-Lys-d-Asp Andersson et al. (1995)
Virgibacillus
V. pantothenticus meso-DAP direct Schleifer and Kandler (1972)
V. halodenitrificans meso-DAP direct Denariaz et al. (1989)
V. marismortui meso-DAPb Arahal et al. (1999)
V. salexigens meso-DAPb Garabito et al. (1997)
footnote a: Data in parentheses were not obtained from the type strain of the species.
footnote b: Configuration not determined.
footnote c: This neutrophilic species is closely related to the alkaliphilic species Bacillus cohnii and Bacillus halmapalus.
footnote d: Ahmed et al. (2007c) proposed the transfer of these species to the new genus Lysinibacillus.
footnote e: The strain analyzed by Schleifer and Kandler (1972) as Bacillus circulans (ATCC 9966) has been reallocated to Paenibacillus amylolyticus.
#####
TABLE 3. Differential characteristics of the species of the genus Bacillusa,b
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; v, variation within strains; w, weak reaction; −/w, negative or weak reaction; d/w, different strains give different reactions and reactions are weak when positive; ng, no growth in the test medium; no entry indicates that no data are available.
footnote b: Compiled from Larkin and Stokes (1967); Nakayama and Yanoshi (1967); Gordon et al. (1973), Pichinoty et al. (1976, 1983, 1984); Aragno (1978); Schenk and Aragno (1979); Pichinoty (1983); Bonjour and Aragno (1984); Logan and Berke- ley (1984), Claus and Berkeley (1986), Priest et al. (1987, 1988); Nakamura et al. (1988, 1999, 2002); Demharter and Hensel (1989b); Denariaz et al. (1989); Nakamura (1989, 1998); Ventosa et al. (1989); Tomimura et al. (1990); Nagel and Andreesen (1991); Arfman et al. (1992); Spanka and Fritze (1993); Andersch et al. (1994); Roberts et al. (1994, 1996); Agnew et al. (1995); Boone et al. (1995); Combet-Blanc et al. (1995); Nielsen et al. (1995a); Fritze (1996a); Fujita et al. (1996); Kuhnigk et al. (1995); Kuroshima et al. (1996); Pettersson et al. (2000, 1996); Shelobolina et al. (1997); Lechner et al. (1998); Switzer Blum et al. (1998); Yumoto et al. (2004c, 2003, 1998); Rheims et al. (1999); Logan et al. (2002a, b, 2000, 2004b); Palmisano et al. (2001); Yoon et al. (2001a, 2003a); Abd El-Rahman et al. (2002); Ajithkumar et al. (2002); Kanso et al. (2002); Li et al. (2002); Reva et al. (2002); Venkateswaran et al. (2003); Gugliandolo et al. (2003a); Heyrman et al. (2003a, 2005a, 2004); Taubel et al. (2003); De Clerck et al. (2004b, 2004c); Heyndrickx et al. (2004); La Duc et al. (2004); Ivanova et al. (2004a); Noguchi et al. (2004); Santini et al. (2004); Scheldeman et al. (2004); Suresh et al. (2004).
footnote c: See Table 5 for a comparison of the subspecies of Bacillus subtilis and the closely related species Bacillus subtilis, Bacillus atrophaeus, Bacillus mojavensis and Bacillus vallismortis, and the species Bacillus amyloliquefaciens, Bacillus licheniformis, Bacillus pumilus, and Bacillus sonorensis.
footnote d: See Table 6 for a comparison of alkaliphilic species: Bacillus agaradhaerens, Bacillus alcalophilus, Bacillus algicola, Bacillus arseniciselenatis, Bacillus clarkii, Bacillus cohnii, Bacillus halodurans, Bacillus horti, Bacillus krulwichiae, Bacillus okuhidensis, Bacillus pseudoalcaliphilus, Bacillus pseudofirmus, Bacillus selenitireducens, Bacillus thermocloacae, Bacillus vedderi.
footnote e: See Table 7 for comparison of the closely related species Bacillus anthracis, Bacillus cereus, Bacillus mycoides, Bacillus pseudomy- coides, Bacillus thuringiensis, and Bacillus weihenstephanensis.
footnote f: See Table 8 for comparison of the thermophilic species (optimum growth at 50 °C or above): Bacillus aeolius, Bacillus fumarioli, Bacillus infernus, Bacillus methanolicus, Bacillus schlegelii, Bacillus thermoamylovorans, Bacillus thermocloacae, and Bacillus tusciae.
footnote g: See Table 9 for comparison of the neutrophilic, non-thermophilic species that form spherical spores: Bacillus fusiformis, Bacillus insolitus, Bacillus neidei, Bacillus psychrodurans, Bacillus psychrotolerans, Bacillus pycnus, and Bacillus sphaericus.
footnote h: Pigmentation: Bacillus subtilis may form pigments, varying from cream through yellow, orange, pink and red, to brown or black, on potato or agar media containing glucose, and strains forming brown or black pigment were often formerly called "Bacillus subtilis var. aterrimus"; Bacillus algicola produces semitransparent, creamy, slightly yellowish colonies; Bacil- lus aquimaris colonies are pale orange-yellow; Bacillus arseniciselenatis and Bacillus selenitireducens will produce red colonies, owing to elemental selenium precipitation, on selenium oxide media; Bacillus atrophaeus forms a dark brownish-black soluble pigment in 2–6 d on media containing tyrosine or other organic nitrogen source; Bacillus carboniphilus produces grayish yellow pigment on nutrient agar and brownish red pigment on trypto-soya agar; some strains of Bacillus cereus may produce a yellowish-green fluorescent pigment on various media, some strains may produce a pinkish brown diffusible pigment on nutrient agar, and on starch-containing media containing sufficient iron some strains produce the red pigment pulcher- rimin; Bacillus clarkii colonies may be cream-white to pale yellow in color, and one of the three strains described produces dark yellow colonies with age; Bacillus endophyticus colonies may be white or pink-red, even on the same plate, and media containing ampicillin and lysozyme commonly yield red colonies; colonies of Bacillus fastidiosus on uric acid medium may become yellowish; Bacillus firmus colonies are creamy-yellow to pale orangey-brown after 3 d on TSA at 30 °C; Bacillus gibsonii colonies are yellow; Bacillus indicus colonies are yellowish-orange; Bacillus hwaijinpoensis colonies are light yellow; Bacillus jeot- gali colonies are cream-yellow to light orange-yellow; many strains of Bacillus licheniformis can produce red pigment (assumed to be pulcherrimin) on carbohydrate media containing sufficient iron, and colonies on glycerol/glutamate medium are reddish-brown; Bacillus marisflavi colonies are pale yellow; Bacillus megaterium colonies may become yellow and then brown or black on long incubation; Bacillus pseudofirmus colonies are yellow; Bacillus sonorensis colonies are yellowish-cream on routine media, and bright yellow on pH 5.6 agar; Bacillus subterraneus colonies are dark yellow to orange on tryptic soy agar.
footnote i: Citrate test results may vary according to the test method used; Gordon et al. (1973) found citrate utilization to be a variable property among 23 strains of Bacillus anthracis, while Logan and Berkeley (1984) and Logan et al. (1985) obtained negative results for 37 strains using the API 20E test method. For Bacillus badius, Gordon et al. (1973) obtained negative results with two strains, while Logan and Berkeley (1984) obtained positive results for two strains using the API 20E test method.
footnote j: Strains of Bacillus cereus of serovars 1, 3, 5, and 8, which are particularly associated with outbreaks of emetic-type food poison- ing, do not produce acid from salicin and starch, whereas strains of Bacillus cereus of other serotypes are usually positive for these reactions. See Table 7.
footnote k: For Bacillus fumarioli, acid production from carbohydrates is tested at pH 6 – see Logan et al. (2000) and Testing for special characters.
footnote l: Gordon et al. (1973) found the Voges–Proskauer reaction to be negative for 60 strains of Bacillus megaterium, while Logan and Berkeley (1984) obtained positive results for all but one of 33 strains using the API 20E test method.
footnote m: Acid production from carbohydrates by Bacillus naganoensis is slow, and shows only after extended (>14 d) incubation.
footnote n: The published description of Bacillus pseudomycoides (Nakamura, 1998) records that 7% salt is tolerated, but the differentia- tion table in that publication indicates the opposite result.
footnote o: Spores of Bacillus psychrodurans and Bacillus psychrotolerans are rarely formed; on casein-peptone soymeal-peptone agar spores are predominantly spherical, but on marine agar they are predominantly ellipsoidal.
footnote p: Mosquitocidal strains of Bacillus sphaericus produce parasporal toxin crystals which are smaller than those produced by Bacil- lus thuringiensis, but which are nonetheless visible by phase-contrast microscopy.
footnote q: Growth occurs within the range pH 7.2–9.5 in media adjusted with NaOH and HCl, but not in media where the pH is adjusted using buffered systems as described by Nielsen et al. (1995a).
footnote r: Negative when incubated at 30 °C, but may become positive slowly when incubated at 40 °C.
footnote s: Growth is poor in the absence of NaCl.
#####
TABLE 4. Additional data for differentiation of Bacillus speciesa,b
footnote a: Symbols: +, >85% positive; d, variable (16–84% positive); −, 0–15% positive; w, weak reaction; d/w, variable and weak when positive; no entry indicates that no data are available.
footnote b: Compiled from Larkin and Stokes (1967); Nakayama and Yanoshi (1967); Gordon et al. (1973), Pichinoty et al. (1984, 1976, 1983); Aragno (1978); Schenk and Aragno (1979); Pichinoty (1983); Bonjour and Aragno (1984); Logan and Berkeley (1984); Claus and Berkeley (1986); Priest et al. (1987, 1988); Nakamura et al. (1988, 1999, 2002) Demharter and Hensel (1989b)<qu ref=81>; Denariaz et al. (1989); Nakamura (1989, 1998); Ventosa et al. (1989); Tomimura et al. (1990); Nagel and Andreesen (1991); Arfman et al. (1992); Spanka and Fritze (1993); Andersch et al. (1994); Roberts et al. (1994, 1996); Agnew et al. (1995); Boone et al. (1995); Combet-Blanc et al. (1995); Nielsen et al. (1995a); Fritze (1996a); Fujita et al. (1996); Kuhnigk et al. (1995); Kuroshima et al. (1996); Pettersson et al. (2000, 1996); Shelobolina et al. (1997); Lechner et al. (1998); Switzer Blum et al. (1998); Yumoto et al. (1998, 2003, 2004c); Rheims et al. (1999); Logan et al. (2002a, b, 2000, 2004b); Palmisano et al. (2001); Yoon et al. (2001a); Abd El-Rahman et al. (2002); Ajithkumar et al. (2002); Kanso et al. (2002); Li et al. (2002); Reva et al. (2002); Venkateswaran et al. (2003); Gugliandolo et al. (2003a); Heyrman et al. (2003a, 2005a, 2004); Taubel et al. (2003); Yoon et al. (2003a); De Clerck et al. (2004b, 2004c); Heyndrickx et al. (2004); La Duc et al. (2004); Ivanova et al. (2004a); Noguchi et al. (2004); Santini et al. (2004); Scheldeman et al. (2004); Suresh et al. (2004).
footnote c: Reactions differ between strains of the emetic biotype of Bacillus cereus for these substrates.
footnote d: Results obtained when inocula are supplemented with 7% NaCl.
footnote e: Results obtained when grown at pH 10.
footnote f: Assimilation data for Bacillus simplex are for the type strain only.
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TABLE 5. Differentiation of Bacillus subtilis from closely related Bacillus speciesa,b
1. B. subtilis subspp. subtilis and spizizeniic 6. B. amyloliquefaciens 11. B. atrophaeus 51. B. licheniformis 57. B. mojavensis 73. B. pumilus 83. B. sonorensis 91. B. vallismortis
Characteristic
Pigmented colonies
Yellow-pink-red −d − − − − − we −
Dark brown/black −d − +f − − − − −
Anaerobic growth − − − + − − + −
Acid from:
Glycogen + + nd + nd − nd nd
Methyl α-d-mannoside − − nd − nd + nd nd
Starch + + nd + nd − nd nd
Hydrolysis of starch + + + + + − + +
Utilization of propionate − − − + − − + −
Nitrate reduction + + + + + − + +
Growth in NaCl:
5% + + + + + + − +
7% + nd + + + + − +
10% nd d nd nd + + − +
Growth at:
5 °C − − d − d d − d
10 °C d − + − + d − +
50 °C d + + + + d + +
55 °C − − d d d − + −
65 °C − − − − − − − −
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% posi- tive; w, weak reaction; nd, no data are available.
footnote b: Compiled from Claus and Berkeley (1986), Priest et al. (1987), Nakamura (1989), Roberts et al. (1994, 1996), Nakamura et al. (1999), and Palmisano et al. (2001).
footnote c: The subspecies of Bacillus subtilis are not distinguishable by routine phenotypic tests.
footnote e: Colonies are yellowish-cream on routine media, and bright yellow on pH 5.6 agar.
footnote f: This species accommodates strains forming brownish-black pigment on tyrosine (and so often evident on the crude media available to earlier workers), and often formerly called "Bacillus subtilis var. niger".
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TABLE 6. Differentiation of alkaliphilic Bacillus speciesa,b
3. B. agaradhaerens 4. B. alcalophilus 5. B. algicola 9. B. arseniciselenatis 20. B. clarkii 23. B. cohnii 40. B. halodurans 43. B. horti 49. B. krulwichiae 65. B. okuhidensis 67. B. pseudalcalophilus 68. B. pseudofirmus 76. B. selenitireducens 88. B. thermocloacae 92. B. vedderi
Characteristic
Colonies pigmented − − +c −c +c − − − − − − +c −c − −
Motility nd nd + − nd + nd + + + nd nd − − +
Spores:
Ellipsoidal + + + nd + + − + + + + + nd + v
Spherical − − − nd − − + − − − − − nd − v
Borne terminally − − + nd − + + − − − − − nd + +
Swollen sporangia + − − nd d + + + − v + − nd + +
Catalase nd + w + nd + nd + + + nd nd + + +
Anaerobic growth nd − − +d nd nd nd − + nd nd nd + − +
Hydrolysis of:
Casein + + − nd + + d/w + d + + + nd − −
Gelatin + + + nd + + + + d + + + nd − w
Starch + + + nd − + w + + + + + nd − −
Nitrate reduction + − w nd + + − + + + − − nd − nd
Growth at pH:
6 − − − − − − − − − + − − − − −
7 − − + − − − + + − + − d − − −
8 + + + w nd nd + + + + + + − + −
9 + + + + + + + + + + + + + + +
10 + d + + + + + + + + + nd + − +
Optimum pH 10 9–10 9 8.5–10 10 9.7 9–10 8–10 8–10 10.5 10 9 8.5–10 8–9 10
Growth in NaCl:
2% + + + − nd + + d + + nd nd + + +
5% + + − + + + + + + + + + + w +
7% + + nd + + nd + + + + + + + − +
10% + − nd + + − + + + + + + + − −
NaCl required for growth + − − + + − − d − − − − + − −
Growth at:
10 °C + + + nd − + − − − − + + nd − nd
20 °C + + + + + + + + nd − + + + − nd
30 °C + + + nd + + + + nd + + + nd −
40 °C + + + nd + + + + + + + + nd + +
50 °C − − − nd − − − − − + − − nd + d
55 °C − − − nd − − − − − + − − nd + −
Deamination of phenylalanine − − nd nd + − − − nd − − + nd nd nd
Respiratory growth with As(V) nd nd nd + nd nd nd nd nd nd nd nd + nd nd
Respiratory growth with Se(IV) nd nd nd + nd nd nd nd nd nd nd nd + nd nd
or Se(VI)
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; v, variation within strains; w, weak reaction; d/w, d, different strains give different reactions, but positive reactions are weak; nd, no data are available.
footnote b: Compiled from Claus and Berkeley (1986), Demharter and Hensel (1989b); Spanka and Fritze (1993); Agnew et al. (1995); Nielsen et al. (1995a); Fritze (1996a); Yumoto et al. (1998, 2003); Switzer Blum et al. (2001); Li et al. (2002); Ivanova et al. (2004a).
footnote c: Bacillus algicola produces semitransparent, creamy, slightly yellowish colonies; Bacillus arseniciselenatis and Bacillus selenitireducens will produce red colonies, owing to elemental selenium precipitation, on selenium oxide media; Bacillus clarkii colonies may be cream-white to pale yellow in color, and one of the three strains described produces dark yellow colonies with age; Bacillus pseudofirmus colonies are yellow.
footnote d: Bacillus arseniciselenatis does not grow aerobically; Bacillus selenitireducens grows weakly in microaerobic conditions.
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TABLE 7. Differentiation of Bacillus cereus from closely related Bacillus speciesa,b.
7. B. anthracis 18. B. cereus 60. B. cereus (emetic biovar)c 58. B. mycoides 69. B. pseudomycoides 89. B. thuringiensis 95. B. weihenstephanensis
Characteristic
Motility − + + − − + +
Rhizoid colonies − − − + + − −
Cell diameter >1.0 μm + + + + v + +
Parasporal crystals − − − − − + −
Acid from:
Glycerol − + d + nd + nd
Glycogen + + − + nd + +
Salicin − d − d nd d d
Starch + + − + nd + +
Arginine dihydrolase − d d d nd + nd
Utilization of citrate dd + + d d + +
Nitrate reduction + d + d + + d
Growth at:
5 °C − − nd − − − +
10 °C − d nd d − d +
40 °C + + nd d + + −
Degradation of tyrosine − + nd d + + +
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; v, variation within strains; nd, no data are available.
footnote b: Compiled from Gordon et al. (1973); Logan and Berkeley (1984); Logan et al. (1985); Claus and Berkeley (1986); Lechner et al. (1998); Nakamura (1998).
footnote c: Strains of Bacillus cereus of serovars 1, 3, 5 and 8, which are particularly associated with outbreaks of emetic-type food poisoning.
footnote d: Citrate test results may vary according to the test method used; Gordon et al. (1973) found citrate utilization to be a variable property among 23 strains of Bacillus anthracis, while Logan and Berkeley (1984) and Logan et al. (1985) obtained negative results for 37 strains using the API 20E test method.
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TABLE 8. Differentiation of thermophilic Bacillus speciesa,b
2. B. aeolius 33. B. fumarioli 46. B. infernus 56. B. methanolicus 75. B. schlegelii 87. B. thermoamylovorans 88. B. thermocloacae 90. B. tusciae
Characteristic
Motility + + − − + + − +
Spore formation: nd + − + + − d +
Ellipsoidal + + nd + − nd + +
Cylindrical + + nd − − nd − −
Spherical − − nd − + nd − −
Borne terminally + − nd − + nd + −
Swollen sporangia nd − nd + + nd + +
Catalase − + − + + + + w
Aerobic growth + + − + + + + +
Anaerobic growth − − + − − + − −
Voges–Proskauer + + nd nd − nd − nd
Acid from:
l-Arabinose + −c − nd − + − −
d-Glucose + +c + nd − + − −
Glycogen − −c nd − nd + nd −
d-Mannitol + +c nd + − − nd −
d-Mannose + +c − nd nd + − −
Salicin + −c nd − nd + nd −
Starch + −c nd − nd + nd −
d-Xylose + −c − nd − + − −
Hydrolysis of:
Casein + − − − w nd − nd
Gelatin + + − nd − nd − nd
Starch + nd − d − + − −
Nitrate reduction − − + − + − − +
Growth at pH:
5 − + − nd nd − − +
6 − + − − + + − w
7 + − + + + + − −
8 + − + nd nd + + −
9 + − − nd nd − + −
10 − − − nd nd − − −
Growth in NaCl:
2% + nd + d + nd + −
5% + nd + − − nd w −
7% − nd + nd − nd − −
10% − nd + nd − nd − −
Growth at:
30 °C − + − − − nd − −
40 °C + + − + − nd + −
50 °C + + + + + + + +
55 °C + + + + + + + +
60 °C + − + + + − + nd
65 °C + − − − + − + −
70 °C − − − − + nd − −
Optimum growth 55 50 61 55 70 50 55–60 55
temperature (°C)
Autotrophic with − − − − + − − +
H2 + CO2 or CO
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; nd, no data are available.
footnote b: Compiled from Schenk and Aragno (1979); Bonjour and Aragno (1984); Demharter and Hensel (1989b); Arfman et al. (1992); Boone et al. (1995); Combet-Blanc et al. (1995); Logan et al. (2000); Gugliandolo et al. (2003a).
footnote c: For Bacillus fumarioli, acid production from carbohydrates is tested at pH 6 – see Logan et al. (2000) and Testing for special characters.
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TABLE 9. Differentiation of spherical-spored Bacillus speciesa,b
35. B. fusiformis 47. B. insolitus 61. B. neidei 64. B. odysseyi 70. B. psychrodurans 72. B. psychrotolerans 74. B. pycnus 75. B. schlegelii 78. B. silvestris 84. B. sphaericus
Characteristic
Cell diameter >1.0 mm − v − − − − + − − −
Spores:
Ellipsoidal − + − − +c +c − − − −
Spherical + v + + +c +c + + + +
Borne terminally d + + + + + + + + +
Sporangia swollen + − + + + + + + + +
Parasporal crystals − − − − − − − − − −d
Hydrolysis of:
Casein + ng − − nd − − w − d
Gelatin + − nd − d d nd − − d
Starch − − − − + + − − − −
Utilization of citrate + − − nd − − − − − d
Nitrate reduction − − − − + − − + − −
Growth at pH:
5 − nd nd − − − nd nd nd −
6 + nd + + nd nd + + nd d
7 + + + + + + + + + +
8 + nd nd + nd nd nd nd nd +
9 + nd nd + nd nd nd nd nd +
10 − nd nd + nd nd nd nd nd −
Growth in NaCl:
2% + d + + + + nd + + +
5% + − + + d d − − + +
7% + − nd − − − − − − −
NaCl required for growth nd − − nd nd nd nd nd nd −
Growth at:
5 °C − + d − + + d − − −
10 °C nd + + − + + + − + d
20 °C + + + − + + + − + +
30 °C + − + + + + + − + +
40 °C nd − + + − d + − + d
50 °C nd − − nd nd nd nd + − nd
Deamination of d d nd nd − − nd nd − +
phenylalanine
Autotrophic with nd nd nd nd nd nd nd + nd nd
H2 + CO2 or CO
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; d/w, different strains give different reactions, but positive reactions are weak; v, variation within strains; w, weak reaction; ng, no growth in test medium; nd, no data are available.
footnote b: Compiled from Larkin and Stokes (1967); Schenk and Aragno (1979); Logan and Berkeley (1984); Claus and Berkeley (1986); Priest et al. (1988); Fritze (1996a); Rheims et al. (1999); Abd El-Rahman et al. (2002); Nakamura et al. (2002); Priest (2002); La Duc et al. (2004).
footnote c: Spores of Bacillus psychrodurans and Bacillus psychrotolerans are rarely formed; on casein-peptone soymeal-peptone agar spores are predominantly spherical, but on marine agar they are predominantly ellipsoidal.
footnote d: Mosquitocidal strains of Bacillus sphaericus produce parasporal toxin crystals which are smaller than those produced by Bacillus thuringiensis, but which are none- theless visible by phase-contrast microscopy.
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TABLE 10. Differential characteristics of Alkalibacillus speciesa
1. A. haloalkaliphilus 2. A. filiformis 3. A. salilacus 4. A. silvisoli
Characteristic
Motility + − + +
Gram stain − + + +
NaCl range for growth (%, w/v) >0–25.0 0–18.0 5.0–20.0 5.0–25
pH optimum 9.7 9.0 8.0 9.0–9.5
Growth temperature range (°C) >50 15–45 15–40 20–50
Catalase + w + +
Oxidase + − − −
Acid production from:
d-Fructose − − + −
d-Galactose − − − +
Maltose − − − +
Trehalose − − − +
d-Mannitol − − − +
Hydrolysis of:
Starch w − − −
Casein −/w − − +
Gelatin + + − +
Hippurate + − − −
Esculin + − + −
Nitrate reduction − − − +
footnote a: Data are based on Table 1 in Usami et al. (2007). w, Weak.
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TABLE 11. Differential characteristics of Amphibacillus species and members of closely related genera in the phylogenetic treea
Amphibacillus xylanusb Amphibacillus fermentumc Amphibacillus tropicusc Halolactibacillus halophilusd Halolactibacillus miurensisd Paraliobacillus ryukyuensise Gracilibacillus halotoleransf Gracilibacillus dipsosaurif
Characteristic
Spore formation + + + − − + + +
Anaerobic growth + + + + + + − −
Catalase − − − − − + + +
Cytochromes − − − − − + + +
Quinones − − − − − + + +
Products formed during anaerobic growth:
Acectate + + + + + + − −
Formate + + + + + + − −
Ethanol + + + + + + − −
Lactate − + w + + + − −
Pyruvate w −g −g − − − − −
Mol% G + C 36–38 42 39 40 39 36 38 39
footnote a: Symbols: +, positive; −, negative; w, weakly positive. All species grow aerobically.
footnote b: Niimura et al. (1989).
footnote c: Zhilina et al. (2001a).
footnote d: Ishikawa et al. (2005).
footnote e: Ishikawa et al. (2002).
footnote f: Wainø et al. (1999).
footnote g: Pyruvate is produced under aerobic conditions.
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TABLE 12. Distinctive characteristics of the genus Anoxybacillus and other endospore-forming generaa
Anoxybacillusd Alkaliphiluse Amphibacillusf Anaerovirgulag Bacillush Clostridiumi Desulfotomaculumj Sporolactobacillusk Tindallial,m
Feature
Motility +/− + + + + + +/− + +/−
Gram reaction + + + + + +/− +/− + +
Relation to O2 An/Aer oblig An f Aer Oblig An An/f Aer An (atl) oblig An An/f Aer oblig An
Reduction of:
SO42− to H2S − −b − − − − + − −
NO32− to NO2− +/− − − − +/− +/− − − −
Activity of:
Catalase +/− ND − − + − − − −
Oxidase +/− ND − − +/− − − − −
NaCl (3–12%) − − − − +/− +/− +/− − +
requirement
CO32− require- +/− −c − − − − +/− − −
ment
Lactate as − ND − − − − − + −
sole end
product
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; ND, not determined. Abbreviations: An, anaerobe; An (atl), aerotolerant anaerobe; f Aer, facultative aerobe; oblig An, obligative anaerobe.
footnote b: Reduction of sulfur, thiosulfate, and fumarate.
footnote c: Takai and Fredrickson, personal communication.
footnote d: Data from Pikuta et al. (2000a).
footnote e: Data from Takai et al. (2001).
footnote f: Data from Pikuta et al. (2000a).
footnote g: Data from Pikuta et al. (2006).
footnote h: Data from Pikuta et al. (2000a).
footnote i: Data from Claus and Berkeley (1986).
footnote j: Data from Pikuta et al. (2000a, 2000b).
footnote k: Data from Pikuta et al. (2000a).
footnote l: Data from Pikuta et al. (2003b).
footnote m: Data from Kevbrin et al. (1998).
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TABLE 13. Diagnostic characteristics for species of genus Anoxybacillusa
1. A. pushchinoensisb 2. A. amylolyticusc 3. A. ayderensisd 4. A. contaminanse 5. A. flavithermusf 6. A. gonensisg 7. A. kamchatkensish 8. A. kestanbolensisi 9. A. rupiensisj 10. A. voinovskiensisk
Characteristic
Yellow colonies − − − − + − − − − −
Motility − + + + + + + + + −
Gram reaction + + + Variable + + + + + +
CO32− requirement + − − − − − ND − − −
Catalase − + + + + + − + + +
NO3− reduction + + + + + − ND + − +
Growth on:
Peptone − ND + ND + + + + + +
Xylose − − + + ND + − − + +
Gelatin hydrolysis − − + + − + − + − −
Casein hydrolysis − − − − + − − − + −
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; ND, not deter- mined.
footnote b: Data from Pikuta et al. (2000a).
footnote c: Data from Poli et al. (2006).
footnote d: Data from Dulger et al. (2004c).
footnote e: Data from De Clerck et al. (2004c).
footnote f: Data from Heinen et al. (1982).
footnote g: Data from Belduz et al. (2003).
footnote h: Data from Kevbrin et al. (2005).
footnote i: Data from Dulger et al. (2004).
footnote j: Data from Derekova et al. (2007).
footnote k: Data from Yumoto et al. (2004a).
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TABLE 14. Descriptive table of Anoxybacillus speciesa
1. A. pushchinoensisd 5. A. flavithermuse 6. A. gonensisf 4. A. contaminansg 10. A. voinoskiensish 3. A. ayderensisi 8. A. kestanbolensisj 2. A. amylolyticusk 9. A. rupiensisl 7. A. kamchattkensism
Characteristic
Cell diameter, length 0.5–0.6, 3.0–5.0 0.85, 2.3–7.1 0.75, 5.0 0.7–1.0, 4.0–10.0 0.4–0.6, 0.55, 4.6 0.65, 4.75 0.5, 2.0–2.5 0.7–1.5, 1.0, 2.5–8.8
(μm) 1.5–5.0 3.3–7.0
Gram reaction + + + Variable + + + + + +
Motility − + + + − + + + + +
Spore location Terminal Terminal Terminal Terminal/ −b Terminal Terminal Terminal Terminal Terminal
subterminal
Temperature range 37–65 (62) 30–72 (60) 40–70 (55–60) 40–60 (50) 30–64 (54) 30–70 (50) 40–70 (50–55) 45–65 (61) 35–67 (55) 38–67 (60)
(optimum) (°C)
pH range (optimum) 8.0–10.5 (9.5–9.7) 6–9 6–10.0 4.5–10.0 (7.0) 7–8 6.0–11.0 6–10.5 (7.5–8.5) 5.6 (ND) 5.5–8.5 5.7–9.9
(7.5–8.0) (7.5–8.5) (6.0–6.5) (6.8–8.5)
NaCl range (optimum) 0–3.0 (0.5–1.0) 0–2.5 0–4.0 (2.0) 0–5 (0.5) 0–3 (ND) 0–2.5 (1.5) 0–4 (2.5) 0.6 (ND) ND (ND) ND (ND)
(%, w/v)
Relation to O2 Aerotolerant Facultative Facultative Facultative Facultative Facultative Facultative Facultative Strict Facultative
anaerobe anaerobe anaerobe anaerobe anaerobe anaerobe anaerobe anaerobe aerobe aerobe
Catalase − + + + + + + + + −
Oxidase ND + + − + + + − ND −
NO3− reduction + + − + + + + − − ND
Substrates:
d-Glucose + + + + + + + + + +
d-Fructose + ND + + + + + ND + +
Starch + + + + − + − + + −
Peptone − + + − +c + + ND ND +
Gelatin − − + + − + + − − −
Casein − + − − − − − − + −
Antibiotics:
Sensitive to B ND A, S, T, C ND ND A, S, K, T, G A, S, K, T, G A,P,C,K,F,G,S, T,G,S,E,C ND
T,N,L
Resistant to P, A, V, S, C ND ND ND ND ND ND ND A,O,P ND
G + C mol% 42.2 41.6 57.0 44.4 43.9 54.0 50.0 43.5 41.7 42.3
Source Manure, Russia Hot spring, Hot spring, Gelatin batches, Hot spring, Hot spring, Hot spring, Geothermal soil, Hot spring, Hot spring,
New Zealand Turkey Belgium Russia Turkey Turkey Antarctica Bulgaria Russia
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; ND, not determined. Antibiotics: A, ampicillin; B, bacitracin; C, chloramphenicol; E, erytromycin; F, fusidic acid; G, gentamicin; P, penicillin; O, oxacillin; S, streptomycin; V, vancomycin; K, kanamycin; T, tetracycline; N, novobiocin; L, linkomycin.
footnote b: Spores were never observed.
footnote c: Personal communication.
footnote d: Data from Pikuta et al. (2000a).
footnote e: Data from Heinen et al. (1982).
footnote f: Data from Belduz et al. (2003).
footnote g: Data from De Clerck et al. (2004c).
footnote h: Data from Yumoto et al. (2004a).
footnote i: Data from Dulger et al. (2004c).
footnote j: Data from Dulger et al. (2004c).
footnote k: Data from Poli et al. (2006).
footnote l: Data from Derekova et al. (2007).
footnote m: Data from Kevbrin et al. (2005).
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TABLE 15. Characteristics differentiating Filobacillus from other physiologically or morphologically similar taxaa
Filobacillus milensisb Bacillus agaradhaerensc Bacillus haloalkaliphilusc Bacillus halophilusd Bacillus marismortuie Bacillus pseudofirmusc Gracilibacillus dipsosaurif Gracilibacillus halotoleransg Halobacillus halophilush Halobacillus litoralisi Marinococcus albusi Salibacillus salexigensj Virgibacillus pantothenticusk
Characteristic
Spore shape S E S E E E S E S E/S − E E/S
Sporangium position T ST T C T/ST ST T T C/T C/ST NA C/ST T
Gram reaction − + −a + + + + + + + + + +
Growth in the presence of 20% NaCl + − + + + − − + + + + + −
Growth at 50 °C − − − + + − + + − − ND − +
Murein type Orn-d-Glu ND m-Dpm m-Dpm m-Dpm ND m-Dpm m-Dpm Orn-d-Asp Orn-d-Asp m-Dpm m-Dpm m-Dpm
Acid produced from:
Glucose − + − + + + + + − + − + +
Trehalose − + − + − + + + − + − − +
Xylose − − − + − − + + − + − − −
Casein − + − − + + − − + − − + +
Hydrolysis of:
Gelatin − + + − + + + + + + − + +
Starch − + − − − + + + + − − − +
Nitrate reduction + + + − + − + + − − + + +
G + C content (mol%) 35 39.3−39.5 37–38 51.5 40.7 39–40.8 39.4 38 40.1–40.9 42 44.9 39.5 36.9
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; ND, not determined. Abbreviations: E, ellipsoidal; S, spherical; C, central; ST, subterminal; T, terminal; NA, not applicable; ND, no data.
footnote b: Data from Schlesner et al. (2001).
footnote c: Data from Fritze (1996a).
footnote d: Data from Ventosa et al. (1989).
footnote e: Data from Arahal et al. (1999).
footnote f: Data from Lawson et al. (1996).
footnote g: Data from Wainø et al. (1999).
footnote h: Data from Spring et al. (1996).
footnote i: Data from Hao et al. (1984).
footnote j: Data from Garabito et al. (1997).
footnote k: Data from Heyndrickx et al. (1998).
#####
TABLE 16. Characteristics of Filobacillus milensisa
Characteristic Result
Acid from carbohydrates:
Glucose −
Galactose −
Fructose −
Maltose −
Mannitol −
Sucrose −
Trehalose −
Xylose −
Hydrolysis of:
Casein −
DNA +
Esculin −
Gelatin −
Hippurate +
Pullulan −
Starch −
Tributyrin +
Sensitive to:
Ampicillin +
Chloramphenicol +
Kanamycin −
Streptomycin +
Tetracycline +
Vancomycin +
Voges–Proskauer reaction −
Production of:
Catalase +
Cytochromoxidase −
l-Alanine aminopeptidase −
Phosphatase −
footnote a: Symbols: +, >85% positive; −, 0–15% positive.
#####
TABLE 17. Differentiation of Geobacillus speciesa,b
1. G. stearothermophilus 2. G. caldoxylosilyticus 3. G. debilis 4. G. gargensis 5. G. jurassicus 6. G. kaustophilus 7. G. lituanicus 8. G. pallidus 9. G. subterraneus 10. G. tepidamans 11. G. thermocatenulatus 12. G. thermodenitrificans 13. G. thermoglucosidasius 14. G. thermoleovorans 15. G. toebii 16. G. uzenensis 17. G. vulcani
Characteristic
Motility + + + + + +d + + + + + +d + − + + +
Spore shape:
Ellipsoidal + + +d + + + + + + + +d + + + + + +
Cylindrical d dd +d −d −d + −d + −d − + −d dd −d −d −d
Spore position:
Subterminal + −d − −d +d + + + + −d +d + +d +d + +d dd
Terminal d + + + + + +d + + + + + + + + + +
Sporangia swollen d + −e +f + +g +f + − + +f − + +f + d dd
Aerobic growth + + + + + + + + + + + + + + + + +
Anaerobic growth − d/w −e −e wd +d + − +d −e + d −e −e wd +d −e
Acid from:
l-Arabinose d + d/w − + d +f − − +d − + −e − − +f −
Cellobiose − + −e + + d + − + +d + + + +f − + +
Galactose − + −e + + d + − + + + +f − + −e + +
Glycerol + wd − + + d +d +d + +d + +f + de − + +
Glycogen + +d −d −d wd −d −d −d +d −d −d −d − +d −d +d +
myo-Inositol − −d −d − − d +d +d − +d − − − −e + −e −e
Lactose − + − − − −g −d − − w − +f − − − − +
d-Mannitol d −d −d + + + + −d +d −d + wd + + −e + −e
Mannose + +d − + + + + −e + + + + + + +d + +
l-Rhamnose − d −d − − − −d −h − − − d + − − − −
Sorbitol − −d d/w − − − − +d − − +f − + − − − −
Trehalose + + + + + + +d d + + + + + + +d + +
d-Xylose d + − − −e + + − −e + − + + de − −e +
Utilization of:
n-Alkanes + d/w + + + + + + + + + +
Acetate + − + + + + + − + − + −
Citrate − d − − − d − − − + d +/−c + − − +
Formate − −/w − − + − − − −
Lactate − + + d + + − − + −
Hydrolysis of:
Casein d/w + d +f −e + + − − − w −/w +f + +f − −
Esculin d +d +d + + d +d de + +d + +d + +d − + +
Gelatin + +d d −e + d +d − −e − −e wd −e − −e + +
Starch + + − + + + + w + w − d +g + −e +f +
Urea − − − − − − −d − − − − − +f − − − −g
Nitrate reduction d + −d + −e +g +d − + +f +f + + + + + −e
(continued)
#####
TABLE 17. (continued)
1. G. stearothermophilus 2. G. caldoxylosilyticus 3. G. debilis 4. G. gargensis 5. G. jurassicus 6. G. kaustophilus 7. G. lituanicus 8. G. pallidus 9. G. subterraneus 10. G. tepidamans 11. G. thermocatenulatus 12. G. thermodenitrificans 13. G. thermoglucosidasius 14. G. thermoleovorans 15. G. toebii 16. G. uzenensis 17. G. vulcani
Characteristic
Growth at pH:
6 + + +d + − + +d − + + − + − − + −e +
8 + + +d + − +d +d + −e + + + + − + −e +
8.5 −e d +d + − +d + −e + + d + − + −e +
Growth in NaCl:
1% + −d wd + + +d − + + + + + + + + + +
2% +f −d −d − + d −d + + d + +f − − + + +f
3% +f − −d − + −d −d + + − + +f −d − + + +f
4% +f −d −d − + −d −d + d −d + − −d − + + −
5% df −d −d − + − −d + d −d +d − −d − −e − −
Growth at:
35 °C d − −d − − − − − − − + − − − − − −
40 °C + − −d −e − + − + − + + − − − − −e +
45 °C + + − + + + − + + + + d + + + + +
70 °C +f + + + − d + d d − + d + + + −e +
75 °C df − − − − − − − − −d + − − − − − −
footnote a: Symbols: +, >85% positive; d, variable (16–84% positive); −, 0–15% positive; w, weak reaction; +/w, positive or weakly positive; d/w, variable, but weak when positive; −/w, occasional strains are weakly positive; no entry indicates that no data are available.
footnote b: Compiled from: Golovacheva et al. (1965, 1975); Suzuki et al. (1983); Logan and Berkeley (1984); Claus and Berkeley (1986); Zarilla and Perry (1987); Priest et al. (1988); Scholz et al. (1988); White et al. (1993); Sunna et al. (1997b); Ahmad et al. (2000b); Caccamo et al. (2000); Manachini et al. (2000); Fortina et al. (2001a); Nazina et al. (2004, 2005b, 2001); Sung et al. (2002); Banat et al. (2004); Kuisiene et al. (2004); Schaffer et al. (2004).
footnote c: Tests as Christensen’s citrate-positive and Simmons’ citrate-negative (Suzuki et al., 1983).
footnote d: Data from Dinsdale and Logan (unpublished); all species were negative for lysine decarboxylase, ornithine decarboxylase, H2S production, and acid from d-arabinose, l-arabitol, dulcitol, erythritol, d-fucose, gluconate, 2-keto- d-gluconate, d-lyxose, and methyl-xyloside.
footnote e: Positive according to Dinsdale and Logan (unpublished).
footnote f: Negative according to Dinsdale and Logan (unpublished).
footnote g: Variable according to Dinsdale and Logan (unpublished).
footnote h: Weak according to Dinsdale and Logan (unpublished).
#####
TABLE 18. Additional characters of the species of the genus Geobacillusa,b
1. G. stearothermophilus 2. G. caldoxylosilyticus 3. G. debilis 4. G. gargensis 5. G. jurassicus 6. G. kaustophilus 7. G. lituanicus 8. G. pallidus 9. G. subterraneus 10. G. tepidamans 11. G. thermocatenulatus 12. G. thermodenitrificans 13. G. thermoglucosidasius 14. G. thermoleovorans 15. G. toebii 16. G. uzenensis 17. G. vulcani
Characteristic
Voges–Proskauer de −e − −e −e − wd − −e −e −e −g −g −e + −e +
ONPGd − − + − − d + − − + − − − − − − −
Arginine dihydrolased − − + − − − − − − − − − − − − − −
Methyl red d − − + + − w − − −
Utilization of:
Benzoate + + +
Butyrate + + + + +
Fumarate + + + +
Malate + − + +
Phenylacetate − + + +
Propionate − − d +
Pyruvate + + + + +
Succinate + + + + +
Butanol − − + +
Ethanol − + + +
Phenol − + +
Acid from:
N-Acetylglucosamine − −d +d −d −d −d wd −d −d +d +/wd dd +d wd −d −d +d
Adonitold − − − − − − − + − − − − − − − − −
Amygdalind − + + − − − − + − + + − + − − − d
d-Arabitold − − − − − − − + − − − − − − − − −
Arbutind − + w + − − − + − + + − + − − w −
Fructose + + −e + + + + + + + + + + + +d + +
Gentiobiosed − + + − − − − − d + − + − − − −
d-Glucose + + −e + + + + + + + + + + + + + +
Inulind + − − − − − − − − − − − − − − − −
Maltose + + − + + + + d + + + + + + +d + +
d-Melezitose + −d −d +d +d − wd −d +d +d +d dd −d +/wd −d −d +d
d-Melibiose + +d −d +d −d − +d −d −d −d +d +d dd +d +d −d +d
Methyl d-glucoside + wd −d −d +d −d +d +d −d +d wd −d +d −/wd −d +d dd
Methyl d-mannosided d − − − − − − − − − − − − − − − −
Raffinose + +d +f −e − − −d −d − +d wd −/wd d +d wd − +d
Ribose − + d/w + + d + −e +f wf + + −e + −d −d +
Salicin − +d +d +d −d d −d +d +d +d +d dd + wd −d +d +d
l-Sorbosed − − − − − − − d − − − − d − w − −
Starch + +d +d +d wd dd +d +d +d +d +d +d + +d −d +d +d
Sucrose + + − + + d + + + + + +g + + −d + +
d-Turanose d +d wd −d +d −d −d wd −d +d wd dd +d wd −d −d −d
Xylitold − − − − − − − + − − − − − − − − −
Phenylalanine deamination − − −
Gas from nitrate − d − − − − − + − + + − − + − −
footnote a: See Table 17 for explanation of footnotes.
#####
TABLE 19. Differential characteristics for Gracilibacillus speciesa,b
Characteristic 1. G. halotolerans 2. G. boraciitolerans 3. G. dipsosauri 4. G. orientalis
Growth pigment Creamy white Light pink to red White Cream
Anaerobic growth − ND + −
Optimal growth temperature,°C 47 25–28 45 37
Spore shape E S/E S S
NaCl growth range (% w/v) 0–20 0–11 0–15 1–20
Boron tolerance (mM) 0–50 0–450 0–150 ND
Oxidase test + + + −
Gelatin hydrolysis + − + +
Urea hydrolysis + − − −
H2S production + − − −
Nitrate reduction to nitrite + − + −
Voges–Proskauer test − + − ND
DNA G + C content (mol%) 38 35.8 39.4 37.1
footnote a: Taken from Wainø et al. (1999), Ahmed et al. (2007b), Lawson et al. (1996), and Carrasco et al. (2006).
footnote b: Symbols: +, positive, −; negative; E, ellipsoidal; S, spherical; ND, no data.
#####
TABLE 20. Characteristics helpful in differentiating Gracilibacillus from closely related genera based on type strain reactionsa,b
Paraliobacillus Amphibacillus Halolactibacillus
Characteristic Gracilibacillus halotolerans ryukyuensis xylanus halophilus
Spore formation + + + −
Anaerobic growth −c +(F) +(F) +(F)
Glucose required for aerobic growth − + + +
Major isoprenoid quinones MK-7 MK-7 None None
footnote a: Taken from Isikawa et al. (2005).
footnote b: Symbols: +, positive, −; negative; F, fermentation.
footnote c: Gracilibacillus dipsosauri exhibits anaerobic respiration (see species description).
#####
TABLE 21. Differential characteristics of Halobacillus species and phylogenetically closely related taxaa
halophilusb Halobacillus karajensisb Halobacillus litoralisb Halobacillus trueperib halophilusc Bacillus Marinococcus albusd
Characteristic
Morphology Coccoid Rod Rod Rod Rod Coccus
Flagella Peritrichous None Peritrichous Peritrichous Peritrichous 1 or 2
Pigmentation Orange Colorless Orange Orange Colorless Colorless
Spores Spherical Ellipsoidal (spheri- Ellipsoidal (spheri- Ellipsoidal (spheri- Ellipsoidal None
cal) cal) cal)
NaCl range (%) 2–20 1–24 0.5–25 0.5–30 3–30 5–20
Temperature range 15–40 10–49 10–43 10–44 15–50 ND
(°C)
Nitrate reduction − − − − − +
Acid from:
d-Galactose − − − + ND −
Glucose − + + + + −
Maltose − + + + − −
d-Xylose − − + − + −
Hydrolysis of:
Casein + + − − − −
Gelatin + + + + − −
Esculin − + − − + −
Starch + + − − − −
Urea − − − − + +
Cell-wall type Orn-d-Asp Orn-d-Asp Orn-d-Asp Orn-d-Asp m-Dpm m-Dpm
G+C content 40.1–40.9 41.3 42 43 51.5 44.9
(mol%)
Source of isolation Salt marsh soil Hypersaline soil Hypersaline Hypersaline Seashore drift wood Solar saltern
sediment sediment
footnote a: Symbols: +, >85% positive; d, different strains give different reactions (16–84% positive); −, 0–15% positive; w, weak reaction; ND, not determined.
footnote b: Data from Amoozegar et al. (2003).
footnote c: Data from Ventosa et al. (1989).
footnote d: Data from Hao et al. (1984).
#####
TABLE 22. Fatty acid composition of type strains of Halobacillus species after growth on MB agar (DIFCO 2216) at 28 °C for 48 h prior to analysisa
Halobacillus halophilus Halobacillus karajensis Halobacillus litoralis Halobacillus trueperi
Fatty acidb
Equivalent chain-length (DSM 2266T) (DSM 14948T) (DSM 10405T) (DSM 10404T)
13.618 C14:0 iso 10.6 2 23.2
14.623 C15:0 iso 7.4 11.3 16.3 6.6
14.715 C15:0 anteiso 42.1 42.4 45.6 19.2
15 C15:0 0.8 0.3
15.387 C16:1 ω7c OH 8.8 9.7 2.6 12.2
15.627 C16:0 iso 14.2 6.9 1.2 28
15.756 C16:1 ω11c 0.6 0.9 1.3 0.7
15.998 C16:0 1 1.1 0.9 1
16.388 C17:1 ω10c iso 1.1 2.5
16.478 Summed feature 4c 0.9 3.3 6.8 1.2
16.631 C17:0 iso 1.7 5 7.8 2.8
16.724 C17:0 anteiso 11.6 16 15 5.1
footnote a: Values are percentages of total fatty acids.
footnote b: The position of the double bond in unsaturated fatty acids is located by counting from the methyl (ω) end of the carbon chain; cis and trans isomers are indicated by the suffixes c and t, respectively.
footnote c: Summed feature 4 contained one or more of the following fatty acids: C17:1 iso I and/or C17:1 anteiso.
#####
TABLE 23. Characteristics differentiating Halolactibacillus speciesa,b
Characteristic H. halophilus H. miurensis
NaCl optima (%) 2.0–3.0 2.5–3.0
NaCl range (%) 0–23.5 to 24.0 0–25.5
pH optima 8.0–9.0 9.5
pH range 6.5–9.5 6.0–6.5 to 10.0
Temperature optimum (°C) 30–37 37–40
Temperature range (°C) 5–10 to 40 5–45
Casein hydrolysis − −
Gelatin hydrolysis − −
Starch hydrolysis w w
Nitrate reduction − −
NH3 from arginine − −
Dextran from sucrose − −
DNase − −
Fermentation of:
d-Glucose, d-fructose, d-mannose, d-galactose, maltose, sucrose, + +
d-cellobiose, lactose, melibiose, d-trehalose, d-raffinose, d-mannitol,
starch, α-methyl-d-glucoside, d-salicin, gluconate
Glycerol + w
d-Ribose (+) +
d-Arabinose, d-rhamnose − (−)
Adonitol, myo-inositol, dulcitol, d-sorbitol − −
l-Arabinose, d-xylose, d-melezitose, inulin − +
Gas from gluconate − −
Yields of lactate from glucose (%) 50–60 50–60
Major fatty acid composition (% of total):c
C12:0 2.5 2.1
C13:0 iso 6.5 5.7
C13:0 ante 19.1 18.8
C14:0 iso/ante − 0.5
C14:0 4.1 4.0
C15:0 iso 3.5 3.7
C15:0 ante 6.2 7.6
C15:0 1.4 1.6
C16:0 iso 0.9 0.8
C16:0 43.1 37.2
C16:1 − 1.3
C16:1 ω7 − 0.7
C17:0 iso 1.5 3.5
C17:0 ante − 2.5
C17:0 − 0.9
C18:0 4.6 5.5
C18:1 ω9 (oleic acid) 2.7 2.2
C18:2 1.1 1.2
footnote a: Symbols: +, all strains positive; (+), most strains positive; w, weakly positive; (−), most strains negative; −, all strains negative.
footnote b: Data from Ishikawa et al. (2005).
footnote c: Fatty acid compositions are of strain IAM 15242T (Halolactibacillus halophilus) and of strain IAM 15247T (Halolactibacil- lus miurensis).
#####
TABLE 24. Effect of initial pH of culture medium on the product balance of glucose fermentation in Halolactibacillus speciesa
H. halophilus IAM 15242T H. miurensis IAM 15247T
Initial pH of culture medium 7 8 9 7 8 9
End products [mol/(mol glucose)]:
Acetate 0.27 0.37 0.74 0.28 0.45 0.51
Ethanol 0.16 0.46 0.47 0.18 0.53 0.32
Formate 0.73 0.81 1.84 0.76 0.81 1.28
Lactate 1.50 1.13 0.45 1.30 1.13 0.73
Lactate yield from consumed 75 57 22 65 57 37
glucose (%)
Carbon recovery (%) 101 98 93 93 103 86
footnote a: Data from Ishikawa et al. (2005).
#####
TABLE 25. Products from glucose under aerobic and anaerobic cultivation conditions for Halolactibacillus speciesa,b
H. halophilus IAM 15242T H. miurensis IAM 15247T
Cultivation Aerobic Anaerobic Aerobic Anaerobic
Glucose consumed (mM) 25.0 24.8 21.0 27.9
Products (mM):
Acetate 13.4 9.8 12.9 9.7
Ethanol ND 12.1 ND 12.8
Formate ND 22.3 ND 24.9
Lactate 19.0 27.6 13.0 29.4
Pyruvate 15.9 ND 13.8 ND
Carbon recovery (%) 88 100 84 94
footnote a: ND, not detected.
footnote b: Data from Ishikawa et al. (2005).
#####
TABLE 26. Characteristics differentiating Halolactibacillus from other related members of the halophilic/halotolerant/alkaliphilic and/or
Halolactibacillusb Alkalib-acteriumc,d,e,f Amphibacillus xylanusg
Characteristic
Spore formation − − +
Anaerobic growth + (F) + (F) + (F)
Catalase − − −
Glucose requirement in aerobic cultivation + ND +
NaCl (range, %) 0–25.5 0–17 3, +; 6, −
NaCl (optimum, %) 2–3 2–13 ND
pH (range) 6–10 8.5–12 8–10