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Fodinicurvata halophila sp. nov., a moderately halophilic bacterium from a marine saltern

Abstract

A Gram-stain-negative, rod-shaped, facultatively anaerobic, moderately halophilic bacterium, designated strain BA45ALT, was isolated from water of a saltern located in Santa Pola, Alicante, Spain. Cells were motile, and catalase-and oxidase-positive. Strain BA45ALT grew at temperatures in the range 14-45 °C (optimally at 37 °C), at pH 5.0-9.0 (optimally at pH 7.5), and in media containing 5-20% (w/v) salts [optimally in media containing 10% (w/v) salts]. Phylogenetic analysis based on the comparison of 16S rRNA gene sequences revealed that strain BA45ALT is a member of the genus Fodinicurvata. The closest relatives to the novel strain were Fodinicurvata fenggangensis YIM D812T and Fodinicurvata sediminis YIM D82T with sequence similarities of 98.2% and 97.4%, respectively. DNA-DNA hybridization between the novel isolate and these phylogenetically related species revealed relatedness values of 30% and 15%, respectively, with respect to the aforementioned species. The major cellular fatty acids of strain BA45ALT were C18: 1ɷ7c, C16: 0 and iso-C15: 0. The G+C content of the genomic DNA of strain BA45ALT was 58.0 mol%, and the polar lipid pattern consisted of diphosphatidylglycerol, phosphatidylmethylethanolamine and a number of unknown phospholipids and lipids. Based on phenotypic, chemotaxonomic and phylogenetic data presented in this study, strain BA45ALT constituted a novel species of the genus Fodinicurvata, for which the name Fodinicurvata halophila sp. nov. is suggested. The type strain is BA45ALT (=CCM 8504T=CECT 8472T=JCM 19075T=LMG 27945T).

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Fodinicurvata halophila sp. nov., a moderately halophilic bacterium from a marine saltern

Author: Infante Domínguez, Carmen; Lawson, Paul A.; Johnson, Crystal N.; Sánchez-Porro Álvarez, Cristina; Ventosa Ucero, Antonio
Publisher: Microbiology Society
Year: 2015
DOI: 10.1099/ijs.0.000010
Source: https://idus.us.es/bitstreams/7ca9e596-a900-4d1e-94b0-0806687bc56f/download
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“This is an Accep ed Manusc ip o an a icle published in In e na ional
Jou nal o Sys ema ic and E olu iona y Mic obiology on 01 Ma ch 2015,
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Fodinicu a a halophila sp. no ., a mode a ely halophilic bac e ium om a ma ine
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sal e n
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Ca men In an e-Dominguez1, Paul A. Lawson2, C ys al N. Johnson2, C is ina Sánchez-
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Po o1 and An onio Ven osa1
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1Depa men o Mic obiology and Pa asi ology, Facul y o Pha macy, Uni e si y o
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Se illa, 41012 Se illa, Spain.
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2Depa men o Mic obiology and Plan Biology, Uni e si y o Oklahoma, No man,
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Oklahoma, 73019 USA.
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Running i le: Fodinicu a a halophila sp. no .
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Subjec ca ego y: New axa, P o eobac e ia
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Au ho o co espondence: An onio Ven osa, Depa men o Mic obiology and
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Pa asi ology, Facul y o Pha macy, Uni e si y o Se illa, 41012 Se illa, Spain. E.-mail:
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en [email p o ec ed]s
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The GenBank/EMBL/DDBJ accession numbe o he 16S RNA gene sequence o
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s ain BA45ALT is HG764424.
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Two supplemen a y igu es and a supplemen a y able a e a ailable wi h he online
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e sion o his pape .
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Manusc ip Including Re e ences (Wo d documen )
Click he e o download Manusc ip Including Re e ences (Wo d documen ): Fodinicu a a halophila- e .doc
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A G am-s ain-nega i e, od-shaped, acul a i ely anae obic, mode a ely halophilic
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bac e ium, designa ed s ain BA45ALT, was isola ed om wa e o a sal e n o
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San a Pola, Alican e, Spain. Cells we e mo ile, ca alase and oxidase posi i e. S ain
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BA45ALT g ew a empe a u es in he ange 14-45 ºC (op imally a 37 ºC) and a
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pH 5.0-9.0 (op imally a pH 7.5). I g ew in media con aining 5-20 % (w/ ) sal s
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and op imally in media con aining 10 % (w/ ) sal s. Phylogene ic analysis based
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on he compa ison o 16S RNA gene sequences e ealed ha s ain BA45ALT is a
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membe o he genus Fodinicu a a. The closes ela i es o his s ain we e
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Fodinicu a a enggangensis YIM D812T and Fodinicu a a sediminis YIM D82T
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wi h sequence simila i ies o 98.2 % and 97.4 %, espec i ely. DNA-DNA
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hyb idiza ion be ween he no el isola e and hese phylogene ically ela ed species
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e ealed ela edness o 15 % and 30 %, espec i ely, wi h espec o he
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a o emen ioned species. The majo cellula a y acids o s ain BA45ALT we e
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C18:1ω7c, C16:0 and iso-C15:0. The G+C con en o he genomic DNA o his s ain
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was 58.0 mol%. I s pola lipid pa e n consis ed o diphospha idylglyce ol,
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phospha idylme hyle hanolamine and a numbe o unknown phospholipids and
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lipids. Based on he pheno ypic, chemo axonomic and phylogene ic da a p esen ed
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in his s udy s ain BA45ALT cons i u ed a no el species o he genus
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Fodinicu a a, o which he name Fodinicu a a halophila sp. no . is sugges ed.
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The ype s ain o Fodinicu a a halophila is s ain BA45ALT (= CCM 8504T =
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CECT 8472T = JCM 19075T = LMG 27945T).
45
46
47
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The genus Fodinicu a a belongs o he amily Rhodospi illaceae, wi hin he o de
48
Rhodospi illales (P ening & T üpe , 1971) o he class Alphap o eobac e ia. This genus
49
was o iginally p oposed by Wang e al. (2009) and o da e i comp ises wo species,
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Fodinicu a a sediminis (Wang e al., 2009) and Fodinicu a a enggangensis (Wang e
51
al., 2009). Bo h species we e isola ed om a sedimen sample collec ed om a sal
52
mine in Yunnan, sou h-wes China. The species o he genus Fodinicu a a s ain G am-
53
nega i e, a e acul a i ely anae obic, non-mo ile, ib ioid o od-shaped. Ca alase and
54
oxidase a e p oduced and polyhyd oxybu y a e (PHB) g anules a e obse ed. The
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p edominan pola lipids consis o diphospha idylglyce ol,
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phosphai idylme hyle hanolamine and phospha idylcholine. Phospa idylinosi ol is
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a iable be ween species. Thei DNA G+C con en anges om 61.5 – 62.3 mol%
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(Wang e al., 2009).
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Recen me agenomic s udies ha e p o ided new insigh s in o he mic obial di e si y
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along he salini y g adien o sal e ns loca ed in San a Pola and Isla C is ina in Spain
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(Ghai e al., 2011; Fe nández e al., 2014 a,b; León e al., 2014; Ven osa e al., 2014).
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Du ing he cou se o a s udy o sea ch o he mos abundan mic oo ganisms in hese
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sal e ns, a no el, mode a ely halophilic, yellow o c eam pigmen ed, G am-s ain-
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nega i e bac e ium, s ain BA45ALT, was isola ed om a wa e sample. The pH o he
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wa e was 7.4 and i s salini y was 12 % (w/ ) sal s. A he ime o sampling, he
66
empe a u e o he wa e was 28 ºC. In his pape we desc ibe he axonomic ea u es o
67
his no el bac e ium and p opose i as a new species o he genus Fodinicu a a, as
68
Fodinicu a a halophila sp. no .
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Aliquo s o he wa e sample we e dilu ed wi h s e ile sal solu ion, whe e 100
70
mic oli e s was pla ed on saline media and incuba ed ae obically a 37º C. The isola ion
71
4
medium used was p epa ed wi h a 15% (g l-1) sal s mix u e: NaCl, 117; MgCl2.6H2O,
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19.5; MgSO4.7H2O, 30.5; CaCl2.2H2O, 0.3; KCl, 3; NaHCO3, 0.06; NaB , 0.23 (Subow,
73
1931), supplemen ed wi h 0.25 % (w/ ) yeas ex ac (Di co) and 1. 8 % (w/ ) aga
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(Di co). The pH o he medium was adjus ed o 7.5 wi h 1 M KOH. A pu e cul u e o
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s ain BA45ALT was ob ained a e se e al ans e s on he same medium. Fo ou ine
76
g ow h, he s ain was cul i a ed in he same medium bu p epa ed wi h 10 % (w/ )
77
o al sal s. The cul u e was main ained a -80 ºC in he ou ine medium con aining 50 %
78
( / ) glyce ol. Fodinicu a a enggangensis DSM 21160T and Fodinicu a a sediminis
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DSM 21159T ob ained om he DSMZ (Ge man Collec ion o Mic oo ganisms and Cell
80
Cul u es) we e used as e e ence s ains and we e g own ollowing he
81
ecommenda ions o he cul u e collec ion.
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Cell mo phology and mo ili y we e examined using an Olympus CX41 mic oscope
83
equipped wi h phase-con as op ics. Cells we e mo ile, bo h od- and ib ioid-shaped
84
(Supplemen a y Fig. S1, a ailable a IJSEM online). Colony mo phology was obse ed
85
on ou ine medium unde op imal g ow h condi ions a e incuba ion a 37º C o 5
86
days. Colonies o s ain BA45ALT ha o med on aga pla es we e ci cula , smoo h,
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en i e, opaque, yellow-c eam pigmen ed, wi h a size o 0.3-0.5 mm. The s ain was able
88
o accumula e PHB, as was obse ed by using an Olympus CX41 mic oscope equipped
89
wi h phase-con as op ics.
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Genomic DNA om s ain BA45ALT was ob ained using he me hod desc ibed by
91
Ma mu (1961). The 16S RNA gene o he isola e we e ampli ied by PCR using
92
o wa d p ime 16F27 and e e se p ime 16R1488 (Mellado e al., 1995). An almos
93
comple e 16S RNA gene sequence (1392 n ) o s ain BA45ALT was ob ained. The
94
iden i ica ion o he closes phylogene ic neighbou s and calcula ion o pai wise 16S
95

5
RNA gene sequence simila i y we e achie ed using he EzTaxon-e ool (Kim e al.,
96
2012). The 16S RNA gene sequence simila i ies be ween BA45ALT and he mos
97
closely ela ed Fodinicu a a ype s ains, Fodinicu a a enggangensis YIM D812T
98
and Fodinicu a a sediminis YIM D82T we e 98.2 and 97.4 %, espec i ely. The 16S
99
RNA gene sequence was aligned and checked agains bo h p ima y and seconda y
100
s uc u es o he 16S RNA molecule using he alignmen ool o he ARB so wa e
101
package. Phylogene ic ees we e cons uc ed using h ee di e en me hods, namely he
102
maximum-likelihood (Felsens ein, 1981), maximum-pa simony (Fi ch, 1971) and
103
neighbou -joining (Sai ou & Nei, 1987) algo i hms in eg a ed in he ARB so wa e. The
104
maximum-likelihood phylogene ic ee is p esen ed in Fig. 1. Simila ee opologies
105
we e ob ained using o he eeing me hods: neighbou -joining and maximum-pa simony
106
(Fig. S2)
107
Al hough no p ecise co ela ion exis s be ween he pe cen age o 16S RNA sequence
108
di e gence and species delinea ion, i is gene ally ecognized ha di e gence alues o
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1.3% o mo e a e signi ican (S ackeb and & Goebel, 1994). Howe e , due o he high
110
sequence simila i y alues o i s close ela i es, DNA-DNA hyb idiza ion s udies we e
111
pe o med. These s udies we e ca ied ou using he compe i ion p ocedu e o he
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memb ane me hod (Johnson, 1994) as desc ibed by Ven osa e al. (2004). The le els o
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DNA-DNA ela edness o s ain BA45ALT wi h espec o Fodinicu a a
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enggangensis DSM 21160T and Fodinicu a a sediminis DSM 21159T we e 30 and 15
115
%, espec i ely. These da a con i med ha s ain BA45ALT ep esen s a no el species
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o he genus Fodinicu a a, ha ing DNA-DNA hyb idiza ion alues <70 % wi h espec
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o he ype s ains o he mos closely ela ed species o he genus Fodinicu a a (Waine
118
e al., 1987; S ackeb and & Goebel, 1994).
119
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Op imal condi ions o g ow h and ange we e de e mined by g owing he s ain in he
120
ou ine medium p epa ed a he ollowing salini y alues: 0.5, 3, 5, 7, 7.5, 8, 10, 12, 15,
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20 and 25% (w/ ) o al sal s and a empe a u es om 5 o 45 ºC (in inc emen s o 5 ºC)
122
and om 35 o 40 ºC in inc emen s o one uni . The pH ange o isola e was
123
de e mined in he ou ine medium a op imal salini y adjus ed o pH alues anging
124
om 4 o 10 (in inc emen s o one pH uni ). The g ow h was de e mined by moni o ing
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he op ical densi y using a spec opho ome e a 600 nm.
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S ain BA45ALT g ew in media con aining 5-20 % (w/ ) sal s and op imally in media
127
con aining 10 % (w/ ) sal s. No g ow h was obse ed in he absence o NaCl. I g ew a
128
empe a u es in he ange 14-45 ºC (op imally a 37 ºC) and a pH 5.0-9.0 (op imally a
129
pH 7.5). The s ain was able o g ow anae obically, de e mined a e incuba ion in an
130
anae obic chambe (GasPak Anae obic sys em, BBL).
131
All pheno ypic ea u es we e ca ied ou g owing he s ain in he ou ine medium
132
p epa ed a 10% (w/ ) o al sal s, pH 7.5 and a 37 ºC. The p oduc ion o acid om
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di e en ca bohyd a es was es ed in a medium wi h 0.5 % (w/ ) yeas ex ac and
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supplemen ed wi h 1% (w/ ) o he ca bohyd a e es ed (s e ilized sepa a ely) (Ven osa
135
e al., 1982). To de e mine he u iliza ion o di e en o ganic subs a es as ca bon and
136
ene gy sou ces, a medium con aining 0.05 % (w/ ) yeas ex ac and supplemen ed wi h
137
1 % (w/ ) o he es ed subs a e, was used (Ven osa e al., 1982). The esul s o he
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u iliza ion o di e en subs a es a e included in he species desc ip ion.
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Ca alase ac i i y was de e mined by adding a 1% (w/ ) H2O2 solu ion o colonies on
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solid medium. The oxidase es was pe o med using a D ySlide assay (Di co). S ain
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BA45ALT was ca alase and oxidase posi i e. The ollowing es s we e conduc ed as
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desc ibed by Cowan and S eel (1977): hyd olysis o aesculin, casein, gela in, Tween 80,
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7
and s a ch; he u ease, me hyl ed, and indole p oduc ion, Vogues-P oskaue es ; ni a e
144
and ni i e educ ion, and Simmons ci a e. The ollowing es s we e nega i e: me hyl
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ed, Voges-P oskaue , indole p oduc ion om yp one and Simmons’s ci a e, casein
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and s a ch hyd olysis. The s ain was posi i e o ni a e educ ion bu ni i e was no
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educed. Bo h gela in and aesculin we e hyd olyzed and he u ease es was posi i e.
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O he pheno ypic cha ac e is ics o s ain BA45ALT a e included in he species
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desc ip ion.
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An imic obial compounds sensi i i y es s we e pe o med by sp eading he cul u e
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suspension on solid medium pla es and applying discs imp egna ed wi h an imic obial
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compounds ollowing he echnique o Baue -Ki by (Baue e al., 1966). Suscep ibili y
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o an imic obial compounds was de e mined on aga pla es using an imic obial agen
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discs wi h he ollowing concen a ions: ampicillin (10 µg), chlo amphenicol (30 µg),
155
e y h omycin (15 µg), gen amicin (10 µg), nalidixic acid (30 µg), neomycin (10 µg),
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no obiocin (30 µg), penicillin G (10 U) and i ampicin (30 µg). S ain BA45ALT was
157
esis an o ampicillin, e y h omycin, gen amicin, nalidixic acid, neomycin, no obiocin,
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penicillin G, and i ampicin bu suscep ible o chlo amphenicol.
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Se e al pheno ypic di e ences we e obse ed be ween s ain BA45ALT and he wo
160
closely ela ed Fodinicu a a species, including hose conce ning mo ili y,
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physiological and cul u al ea u es, ni a e educ ion, hyd olysis o gela in and aesculin,
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H2S p oduc ion, and u iliza ion o se e al ca bohyd a es (Table 1).
163
The G+C con en o he genomic DNA o s ain BA45ALT was de e mined om he
164
mid-poin alue (Tm) o he he mal dena u a ion p o ile (Ma mu & Do y, 1962),
165
ob ained wi h a Pe kin-Elme UV-Vis Lambda 20 spec opho ome e a 260 nm. The
166
8
DNA G+C con en o s ain BA45ALT was 58.0 mol%, a alue close o hose epo ed
167
o species o he genus Fodinicu a a (Wang e al., 2009).
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Cell biomass o a y acid, pola lipid analyses and quinone composi ion o s ain
169
BA45ALT, Fodinicu a a sediminis DSM 21159T and Fodinicu a a enggangensis
170
DSM 21160T was ob ained by cul i a ion in nu ien aga (NA) supplemen ed wi h 5 %
171
NaCl, incuba ed a 28 ºC o 5 days. The whole-cell a y acid composi ion o s ain
172
BA45ALT and he e e ence s ains was de e mined on cul u es eaching he la e-
173
exponen ial s age o g ow h acco ding o he ou quad an s s eak me hod (Sasse ,
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1990). Fa y acid me hyl es e s we e p epa ed and analysed acco ding o he s anda d
175
p o ocol o he Mic obial Iden i ica ion Sys em (MIDI, Ve sion 6.1; Iden i ica ion
176
Lib a y TSBA40 4.1; Mic obial ID). Ex ac s we e analyzed using a Hewle Packa d
177
model HP6890A gas ch oma og aph equipped wi h a lame-ioniza ion de ec o as
178
desc ibed by Kämp e & K oppens ed (1996). The a y acid p o iles o s ain
179
BA45ALT included C18:1 ω7c (35.6 %), iso-C16:0 (16.3 %) and iso-C15:0 (10.1 %) as he
180
majo a y acids, ollowed by iso-C17:1 ω9c (5.8 %), C17:0 (5.5 %), C16:0 (3.3 %), C18:1
181
2OH (3 %). The a y acid p o ile o s ain BA45ALT was clea ly di e en om hose o
182
he ela ed axa. Al hough all h ee species p oduce C18:1 ω7c as a majo p oduc , he
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no el o ganism also p oduces majo amoun s o iso-C16:0 and iso-C15:0 which a e lacking
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in Fodinicu a a sediminis DSM 21159T and Fodinicu a a enggangensis DSM
185
21160T (Table S1).
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The pola lipids o s ain BA45ALT and he e e ence s ains we e analyzed as desc ibed
187
by G o h e al. (1996). The pola lipids de ec ed in s ain BA45ALT we e
188
diphospha idylglyce ol, phospha idylme hyle hanolamine and a numbe o unknown
189
phospholipids and lipids (Supplemen a y Figu e S2). I is impo an o no e ha in ou
190
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Thompson, J. D., Gibson, T. J., Plewniak, F., Jeanmougin, F. & Higgins, D. J.
312
(1997). The clus al X windows in e ace: Flexible s a egies o mul iple sequence
313
alignmen aided by quali y analysis ools. Nucleic Acids Res 24, 4876–4882.
314
Tindall, B. J., Rosselló-Mó a, R., Busse, H.-J., Ludwig, W. & Kämp e , P. (2010).
315
No es on he cha ac e iza ion o p oka yo e s ains o axonomic pu poses. In J Sys
316
E ol Mic obiol 60, 249–266.
317
Tindall, B. J., Siko ski, J., Smibe , R. A. & K ieg, N. R. (2007). Pheno ypic
318
cha ac e iza ion and he p inciples o compa a i e sys ema ics. In Me hods o Gene al
319
and Molecula Mic obiology, pp. 330-393. Edi ed by C. A. Reddy, T. J. Be e idge, J.
320
A. B eznak, G. A. Ma zlu , T. M. Schmid and R. L. Snyde . Washing on, D. C.: ASM
321
P ess.
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To eblanca, M., Rod iguez-Vale a, F., Juez, G., Ven osa, A.,Kameku a, M. &
323
Ka es, M. (1986). Classi ica ion o non-alkaliphilic halobac e ia based on nume ical
324
axonomy and pola lipid composi ion, and desc ip ion o Haloa cula gen. no . and
325
Halo e ax gen. no . Sys Appl Mic obiol 8, 89–99.
326
Ven osa, A., Quesada, E., Rod íguez-Vale a, F., Ruiz-Be aque o, F. & Ramos-
327
Co menzana, A. (1982). Nume ical axonomy o mode a ely halophilic G am-nega i e
328
ods. J Gen Mic obiol 128, 1959–1968.
329
Ven osa, A., Gu ié ez, M. C., Kameku a, M., Z yagin se a, I. S. & O en, A.
330
(2004). Taxonomic s udy o Halo ub um dis ibu um and p oposal o Halo ub um
331
e es e sp. no . In J Sys E ol Mic obiol 54, 389–392.
332

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Ven osa, A., Fe nández, A. B., León, M. J., Sánchez-Po o, C. & Rod iguez-
333
Vale a, F. (2014). The San a Pola sal e n as a model o s udying he mic obio a o
334
hype saline en i onmen s. Ex emophiles 18, 811-824.
335
Wang, Y. X., Liu, J.-H., Zhang, X. X., Chen, Y.-G., Wang, Z. G., Chen, Y., Li, Q.
336
Y., Peng, Q. & Cui, X. L. (2009). Fodinicu a a sediminis gen. no ., sp. no . and
337
Fodinicu a a enggangensis sp. no ., poly-ß-hyd oxybu y a e-p oducing bac e ia in
338
he amily Rhodospi illaceae. In J Sys E ol Mic obiol 59, 2575–2581.
339
Wayne, L. G., B enne , D. J., Colwell, R. R., G imon , P. A. D., Kandle , O.,
340
K iche sky, M. I., Moo e, L. H., Moo e, W. E. C., Mu ay, R. G. E., S ackeb and ,
341
E., S a , M. P. & T üpe , H. G. (1987). Repo o he ad hoc commi ee on
342
econcilia ion o app oaches o bac e ial sys ema ics. In J Sys Bac e iol 37, 463-464.
343
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Table 1. Cha ac e is ics ha di e en ia e s ain BA45ALT om ela ed species o he
344
genus Fodinicu a a.
345
Taxa: 1, s ain BA45ALT; 2, Fodinicu a a enggangensis DSM 21160T; 3,
346
Fodinicu a a sediminis DSM 21159T. All da a a e om he p esen s udy, excep
347
whe e indica ed. +, Posi i e eac ion; -, nega i e eac ion.
348
Cha ac e is ic
1
2
3
Colony pigmen a ion
Yellow-c eam
C eam-whi e
C eam-whi e
Cell size (wid h x leng h; µm)
0.05-0.1 x 0.2-1.5
0.2-04 x 0.5-1.3
0.3-0.5 x 0.7-1.5
Mo ili y
+
-
-
Tempe a u e ange (ºC)
14-45
15-42
15-42
NaCl ange (%, w/ )
5-20
1.5-20
1.5-20
pH ange
5-9
6.5-8.5
6.5-8.5
Ni a e educ ion
+
-
-
Hyd olysis o :
Aesculin
Gela in
+
+
-
-
+
-
U iliza ion o ca bon sou ces:
F uc ose
Lac ose
Fuma a e
Hippu a e
L-o ni hine
+
+
+
+
+
-
-
-
-
-
-
-
-
-
-
Simmons ci a e
-
+
+
H2S p oduc ion
+
-
-
Majo a y acids
C18:1 ω7c, iso-C16:0 and iso-
C15:0
C18:1 ω7c, 2-OH-C18:1 and
C16:0
C18:1 ω7c, 2-OH-C18:1 and
C16:0
Pola lipids
Diphospha idylglyce ol,
phospha idylme hyle hanola
mine and unknown
phospholipids and lipids
Diphospha idylglyce ol,
phospha idylme hyle hanola
mine and ou unknown
phospholipids and wo
unknown lipids
Diphospha idylglyce ol,
phospha idylme hyle hanol
a mine and ou unknown
phospholipids and wo
unknown lipids
DNA G+C con en (mol%)
58.0
62.3
61.5
Sou ce
Sal wa e
Deposi o sal mine*
Deposi o sal mine*
*Da a om Wang e al. (2009).
349
350
351
18
352
Legend o igu e
353
354
Fig. 1. Maximum-likelihood phylogene ic ee based on 16S RNA gene sequences,
355
showing he ela ionships be ween s ain BA45ALT and ep esen a i es o he amily
356
Rhodospi illaceae. Boo s ap pe cen ages (based on 1000 eplica ions) ≥ 70 % a e
357
shown a b anch poin s. Ba ep esen s 0.1 subs i u ions pe nucleo ide posi ion.
358
Azospi illum melinis TMCY 0552T (DQ022958)
Azospi illum la gimobile ACM 2041T (X90759)
Azospi illum o yzae COC8T (AB185396)
Azospi illum zeae N7T (DQ682470)
Azospi illum lipo e um ATCC 29707T (M59061)
Azospi illum hiophilum BV-sT ( EU678791)
Azospi illum canadense DS2T (DQ393891)
Azospi illum ugosum IMMIB AFH-6T (AM419042)
Azospi illum b asilense NCIMB 11860T (Z29617)
Azospi illum halop ae e ens Au4T (X79731)
Rhodocis a cen ena ia ATCC 43720T (D12701)
Rhodocis a pekingensis 3-pT (AF523824)
Ske manella pa ooensis ACM 2042T (X90760)
Inquilinus ginsengisoli Gsoil 080T (AB245352)
Inquilinus limosus AU476T (AY043374)
Thalassobaculum li o eum CL-GR58T (EF203900)
Phaeospi illum molischianum ATCC 14031T (M59067)
Phaeospi illum ul um NCIMB 11762T (D14433)
Phaeospi illum chand amohanii JA145T (AM779061)
Magne ospi illum g yphiswaldense MSR-1T (Y10109)
Magne ospi illum magne o ac icum DSM 3856T (Y10110)
Telma ospi illum sibe iense 26-4b1T (AF524863)
De lu iicoccus anus Ben 114T (AF179678)
Tis ella mobilis IAM 14872T (AB071665)
Fodinicu a a sediminis YIM D82T (FJ357426)
Fodinicu a a enggangensis YIM D812T (FJ357427)
Rhodo ib io salina um NCIMB 2243T (D14432)
Rhodo ib io sodomensis DS1T (M59072)
Caenispi illum bisanense K92T (EF100694)
Roseospi illum pa um 930IT (AJ011919)
Roseospi a mediosalina BN 280T (AJ000989)
Roseospi a ma ina CE2105T (AJ298879)
Roseospi a goensis JA135T (AM283537)
Roseospi a isakhapa namensis JA131T (AM282560)
Rhodospi a uepe i ATCC 70022T (AJ001276)
Rhodospi illum pho ome icum DSM 122T (AJ222662)
Rhodospi illum ub um ATCC 11170T (CP000230)
Thalassospi a lucen ensis QMT2T (AF358664)
Esche ichia coli ATCC 11775T (X80725)
0.10
Fodinicu a a halophila BA45ALT (HG764424)
97
100
97
100
100
100
99
96
98
100
99
100
93
100
96
100
100
77
78
88
72
81
78
Figu e 1
Click he e o download Figu e: Fig. 1.pp
Supplemen a y Ma e ial File
Click he e o download Supplemen a y Ma e ial Files: Suppl Ma e ial Fodinicu a a halophila.pd