Nitrosopumilus maritimus genome reveals unique - de la Torre ...

Nitrosopumilus maritimus genome reveals unique 

mechanisms for nitrification and autotrophy in 

globally distributed marine crenarchaea 

C. B. Walker a,b , J. R. de la Torre a , M. G. Klotz c , H. Urakawa a , N. Pinel a , D. J. Arp d , C. Brochier-Armanet e , P. S. G. Chain f,g,h , 

P. P. Chan i , A. Gollabgir j , J. Hemp k , M. Hügler l,m , E. A. Karr n , M. Könneke o , M. Shin f,g , T. J. Lawton p , T. Lowe i , W. Martens- 

Habbena a , L. A. Sayavedra-Soto d , D. Lang f,g , S. M. Sievert q , A. C. Rosenzweig p , G. Manning j , and D. A. Stahl a,1 

a Department of Civil and Environmental Engineering, University of Washington, Seattle, WA 98195; b Geosyntec Consultants, Seattle, WA 98101; 

c Department of Biology, University of Louisville, Louisville, KY 40292; d Department of Botany and Plant Pathology, Oregon State University, Corvalis, OR 

97331; e Université de Provence Aix-Marseille I, Laboratoire de Chimie Bactérienne, Centre National de la Recherche Scientifique Unité Propre de Recherche, 

Marseille, 13402 France; f Biosciences Division, Lawrence Livermore National Laboratory, Livermore, CA 94550; g Microbial Program, Joint Genome Institute, 

Walnut Creek, CA 94598; h Center for Microbial Ecology, Michigan State University, East Lansing, MI 48824; i Department of Biomolecular Engineering, 

University of California, Santa Cruz, CA 95064; j Razavi Newman Center for Bioinformatics, Salk Institute for Biological Studies, La Jolla, CA 92037; k School of 

Chemical Sciences, University of Illinois, Urbana, IL 61801; l Leibniz-Institut für Meereswissenschaften, Kiel, 24105 Germany; m Water Technology Center, 

Karlsruhe, 76139 Germany; n Department of Botany and Microbiology, University of Oklahoma, Norman, OK 73019; o Institut für Chemie und Biologie des 

Meeres, Universität Oldenburg, Oldenburg, 26129 Germany; p Departments of Biochemistry, Molecular Biology and Cell Biology, and Chemistry, Northwestern 

University, Evanston, IL 60208; and q Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA 02543 

Edited by David Karl, University of Hawaii, Honolulu, HI, and approved April 2, 2010 (received for review December 6, 2009) 

Ammonia-oxidizing archaea are ubiquitous in marine and terrestrial 

environments and now thought to be significant contributors 

to carbon and nitrogen cycling. The isolation of Candidatus “Nitrosopumilus 

maritimus” strain SCM1 provided the opportunity for 

linking its chemolithotrophic physiology with a genomic inventory 

of the globally distributed archaea. Here we report the 1,645,259bp 

closed genome of strain SCM1, revealing highly copper-dependent 

systems for ammonia oxidation and electron transport that 

are distinctly different from known ammonia-oxidizing bacteria. 

Consistent with in situ isotopic studies of marine archaea, the 

genome sequence indicates N. maritimus grows autotrophically 

using a variant of the 3-hydroxypropionate/4-hydroxybutryrate 

pathway for carbon assimilation, while maintaining limited capacity 

for assimilation of organic carbon. This unique instance of archaeal 

biosynthesis of the osmoprotectant ectoine and an 

unprecedented enrichment of multicopper oxidases, thioredoxinlike 

proteins, and transcriptional regulators points to an organism 

responsive to environmental cues and adapted to handling reactive 

copper and nitrogen species that likely derive from its distinctive 

biochemistry. The conservation of N. maritimus gene content 

and organization within marine metagenomes indicates that the 

unique physiology of these specialized oligophiles may play a significant 

role in the biogeochemical cycles of carbon and nitrogen. 

ammonia oxidation | marine microbiology | archaea | nitroxyl 

Marine Group I archaea are among the most abundant 

microorganisms in the global oceans (1–3). Originally discovered 

through ribosomal RNA gene sequencing (3, 4), recent 

metagenomic, biogeochemical, and microbiological studies 

established the capacity of these organisms to oxidize ammonia, 

thus linking this abundant microbial clade to one of the key steps of 

the global nitrogen cycle (5–9). For a century following the discovery 

of autotrophic ammonia oxidizers, only Bacteria were 

thought to catalyze this generally rate-limiting transformation in 

the two-step process of nitrification (10). Despite recent enrichment 

of mesophilic as well as thermophilic ammonia-oxidizing 

archaea (AOA) (6, 11, 12), only a single Group I-related strain, 

isolated from a gravel inoculum from a tropical marine aquarium, 

has thus far been successfully obtained in pure culture (7). 

The isolation of Nitrosopumilus maritimus strain SCM1 ultimately 

confirmed an archaeal capacity for chemoautotrophic 

growth on ammonia. More detailed characterization of this strain 

revealed cytological and physiological adaptations critical for life 

in an oligotrophic open ocean environment, most notably one of 

the highest substrate affinities yet observed (13). Among characterized 

ammonia oxidizers, only N. maritimus is capable growing at 

the extremely low concentrations of ammonia generally found in 

the open ocean (7, 13). This strain therefore provided an excellent 

opportunity to investigate the core genetic inventory for ammoniabased 

chemoautotrophy by Group I crenarchaea. 

The gene content and gene order of N. maritimus is highly 

similar to environmental populations represented in marine bacterioplankton 

metagenomes, confirming on a genomic level its 

close relationship to many oceanic crenarchaea. Thus, an evaluation 

of the genomic inventory of N. maritimus should offer 

a framework to identify features shared among ammonia-oxidizing 

Group I crenarchaea, resolve physiological diversity among AOA, 

and refine understanding of their ecology in relationship to the 

larger assemblage of marine archaea—not all of which are ammonia 

oxidizers. In support of this expectation, the physiological 

and genomic profiles together show that many of the “nonextreme” 

archaea identified in metagenomic studies, and currently 

assigned to the Crenarchaeota kingdom, are AOA that contribute 

to global carbon and nitrogen cycling, possibly determining rates 

of nitrification in a variety of environments (6, 8, 9, 13). 

Results and Discussion 

Primary Sequence Characteristics. N. maritimus strain SCM1 contains 

a single chromosome of 1,645,259 bp encoding 1,997 predicted 

genes and no extrachromosomal elements or complete 

prophage sequences (Table 1). No unambiguous origin of replication 

could be determined on the basis of local gene content 

or GC skew, as commonly observed for other archaeal genomes 

(14). Approximately 61% of the N. maritimus open-reading 

Author contributions: C.B.W., J.R.d.l.T., P.S.G.C., and D.A.S. designed research; C.B.W., 

J.R.d.l.T., M.G.K., H.U., N.P., C.B-A., P.P.C., A.G., M.H., E.A.K., M.K., M.S., T.L., W.M-H., 

M.S., D.L., S.M.S., A.C.R., G.M., and D.A.S. performed research; C.B.W. and J.R.d.l.T. contributed 

new reagents/analytic tools; C.B.W., J.R.d.l.T., M.G.K., H.U., N.P., D.J.A., C.B.-A., 

P.P.C., A.G., J.H., M.H., E.A.K., M.K., T.J.L., T.L., W.M.-H., L.A.S.-S., S.M.S., A.C.R., G.M., and 

D.A.S. analyzed data; and C.B.W., J.R.d.l.T., M.G.K., H.U., N.P., C.B.-A., J.H., M.H., E.A.K., 

M.K., T.L., S.M.S., A.C.R., G.M., and D.A.S. wrote the paper. 

The authors declare no conflict of interest. 

This article is a PNAS Direct Submission. 

Data deposition: The sequence reported in this paper has been deposited in the NCBI 

database (accession no. NC_010085). 

1 To whom correspondence should be addressed. E-mail: dastahl@u.washington.edu. 

This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. 

1073/pnas.0913533107/-/DCSupplemental. 

www.pnas.org/cgi/doi/10.1073/pnas.0913533107 PNAS Early Edition | 1of6 

MICROBIOLOGY

Table 1. Genome features of N. maritimus SCM1, C. symbiosum, sequenced AOB, and crenarchaeal genome fragments 

Nitrosopumilus Cenarchaeum 

maritimus SCM1 symbiosum 

Nitrosococcus 

oceani 

ATCC 19707 

frames (ORFs) could be assigned to clusters of orthologous 

groups of proteins (COGs), a lower percentage than for genomes 

of ammonia-oxidizing bacteria (AOB) (Table S1) but similar to 

Cenarchaeum symbiosum (15). The genome possesses a relatively 

high coding density (91.9%), with a larger fraction dedicated to 

energy production/conservation, coenzyme transport/metabolism, 

and translation genes than other characterized Crenarchaeota, but 

similar to two common species of photoautotrophic marine 

Bacteria, Prochlorococcus, and Synechococcus. 

Energy Metabolism. The stoichiometry of ammonia oxidation to 

nitrite is similar to that of characterized aerobic, obligate chemolithoautotrophic 

AOB (13), yet the contributing biochemistry is 

distinctly unique. All AOB share a common pathway where hydroxylamine, 

produced by an ammonia monooxygenase (AMO), is 

oxidized to nitrite by a heme-rich hydroxylamine oxidoreductase 

(HAO) complex; the oxidation of hydroxylamine supplies electrons 

to both the AMO and a typical electron transport chain 

composed of cytochrome c proteins. N. maritimus lacks genes 

encoding a recognizable AOB-like HAO complex and pertinent 

cytochrome c proteins, indicating an alternative archaeal pathway. 

The numerous copper-containing proteins, including multicopper 

oxidases and small blue copper-containing proteins (similar to 

plasto-, halo-, and sulfocyanins), suggest an alternative electron 

transfer mechanism (Table S2). These predicted periplasmic 

proteins likely serve functionally similar roles to soluble cytochrome 

c proteins in other Archaea, including Natronomonas 

pharaonis and thermoacidophiles such as Sulfolobus (16). This 

apparent reliance on copper for redox reactions is a major divergence 

from the iron-based electron transfer system of AOB. 

The N. maritimus genome contains genes coding for six soluble 

periplasmic multicopper oxidase (MCO) proteins: two nearly 

identical NO-forming nitrite reductase proteins (NirK; 

Nmar_1259 and -1667), each with three cupredoxin domains; 

two NcgA-like (nirK cluster gene A) MCOs (Nmar_1131 and 

-1663) with two cupredoxin domains; one MCO (Nmar_1136) 

with three cupredoxin domains; and one MCO (Nmar_1354) 

with two domains fused to a blue copper-containing protein 

(Table S2). Two (Nmar_1131 and Nmar_1663) of the three 

genes that are classified as belonging to the emerging class of 

two-domain MCOs (2dMCOs) resemble the general architecture 

of the 2dMCO NcgA present in Nitrosomonas europaea. Although 

the overall sequence identity is low, clustering of NcgA 

with a nitrite reductase suggests it may play a supporting role in 

nitrite reduction (17). Genes Nmar_1131 and Nmar_1663 are also 

colocated with a member of the DtxR family of metal regulators and 

a member of the ZIP metal transport family, suggesting a role in 

metal homeostasis (see below). The third 2dMCO (Nmar_1354), 

possessing a fused blue copper-containing domain, has not been 

found in AOB and appears to be unique to AOA. Redox inter- 

Nitrosomonas 

europaea 

ATCC 19718 

Nitrosomonas 

eutropha C91 

Nitrosospira 

multiformis 

ATCC 25196 

Fosmid 

4B7 

Cosmid 

DeepAnt-EC39 

Size (bp) 1,645,259 2,045,086 3,481,691 2,812,094 2,661,057 3,184,243 39,297 33,347 43,902 

Percent coding 91.90% 91.20% 86.80% 88.40% 85.60% 85.60% 89.10% 86.10% 84.00% 

GC content 34.20% 57.70% 50.30% 50.70% 48.50% 53.90% 34.40% 34.10% 32.60% 

ORFs 1,997 2,066 3,186 2,628 2,578 2,827 41 41 51 

ORF density (ORF/kb) 1.19 0.986 0.889 0.876 0.952 0.86 0.992 1.17 1.12 

Avg. ORF length (bp) 757 924 964 1009 890 980 898 737 753 

Standard tRNAs 44 45 45 41 41 43 0 0 0 

rRNAs 1 1 2 1 1 1 1 1 1 

Plasmids 0 0 1 0 2 3 NA NA NA 

NA, not analyzed. 

Fosmid 

74A4 

actions with the MCOs (and other predicted redox proteins) are 

likely mediated by eight soluble and nine membrane-anchored 

copper-binding proteins containing plastocyanin-like domains 

(Table S2). The corresponding genes appear to be the result of 

a series of duplications within the N. maritimus lineage (Fig. S1). 

A second family of predicted redox active periplasmic proteins, 

composed of 11 thiol-disulfide oxidoreductases from the 

thioredoxin family (Nmar_0639, _0655, _0829, _0881, _1140, 

_1143, _1148, _1150, _1181, _1658, and Nmar_1670), show low 

(but recognizable) identity with the better characterized disulfide 

bond oxidases/isomerases found in Bacillus subtilis (BdbD) and 

Escherichia coli (DsbA, DsbC, and DsbG). The mean percentage 

of sequence identity between the N. maritimus proteins and 

BdbD is 21 ± 3%. The significantly lower mean percentage of 

sequence identity to DsbA, DsbC, and DsbG (9.2, 10.7, and 

10.4%, respectively) is comparable to that shared between the 

E. coli proteins (10–11%). Although functional equivalency 

cannot be established, all but Nmar_0881 preserve the conserved 

thioredoxin-like active site FX4CXXC sequence (18–20). In 

E. coli, both DsbA and DsbC rectify nonspecific disulfide bonds 

catalyzed by copper (21), whereas up-regulation of dsbA by the 

Cpx regulon occurs during copper stress (22, 23). Eukaryotic 

protein disulfide isomerase (PDI) homologs sequester and/or 

reduce oxidized Cu(II), perhaps serving as copper acceptors/ 

donors for copper-containing proteins (24). Another described 

function of PDIs is the capture and transport of nitric oxide (25, 

26), a possible intermediate or by-product of ammonia oxidation. 

The related protein family in N. maritimus may function in part 

to alleviate copper and nitric oxide toxicity. 

Pathways for Ammonia Oxidation and Electron Transfer. The three 

genes (Nmar_1500, _1503, and _1502) annotated as amoA, 

amoB, and amoC and coding for a putative ammonia monooxygenase 

complex are the only recognizable genetic hallmarks 

of ammonia oxidation in the genome sequence. However, the 

N. maritimus sequences are no more similar (in either content or 

organizational structure) to bacterial amo genes than they are to 

the genes encoding bacterial particulate methane monooxygenases 

(pMMO), suggestive of functional differences between 

the archaeal and bacterial versions of AMO (7, 27). 

Notably, mapping the sequence encoded by amoB onto the 

pmoB crystal structure of Methylococcus capsulatus (Bath) (28) 

reveals the conservation of the ligands to the pMMO metal 

centers and the complete absence of both a transmembrane helix 

and a C-terminal cupredoxin domain predicted to be present in 

bacterial AMO (Fig. S2). 

The structural differences in the archaeal AMO, the lack of 

genes encoding the hydroxylamine–ubiquinone redox module 

(29), and a periplasm enriched in redox active proteins together 

suggest significant divergence from the bacterial pathway of 

2of6 | www.pnas.org/cgi/doi/10.1073/pnas.0913533107 Walker et al.

ammonia oxidation. There are two hypothetical mechanistic 

alternatives (Fig. 1, Table S2): either a unique biochemistry 

exists for the oxidation of hydroxylamine or the divergent AMO 

does not actually produce hydroxylamine. If the former is true, 

hydroxylamine oxidation may occur via one of the periplasmic 

MCOs (CuHAO). Given the lack of cytochrome c proteins, the 

four electrons would then be transferred to a quinone reductase 

(QRED) via small blue copper-containing plastocyanin-like 

electron carriers. The protein encoded by Nmar_1226, which 

contains four transmembrane-spanning regions and two plastocyanin-like 

domains, may serve as an analog of the membranebound 

cytochrome cM552 quinone reductase present in AOB 

(29) and is a good candidate for the QRED. 

In an alternative scenario, the archaeal AMO produces not 

hydroxylamine, but the reactive intermediate nitroxyl (nitroxyl 

hydride, HNO). Nitroxyl is a highly toxic and reactive compound 

recently recognized as having biological significance in a number 

of systems (30, 31). During archaeal ammonia oxidation, nitroxyl 

might be formed by a unique monooxygenase function of archaeal 

AMO. Alternatively, the archaeal AMO may act as 

a dioxygenase and insert two oxygen atoms into ammonia, 

producing nitroxyl from the spontaneous decay of HNOHOH. 

Both reaction sequences eliminate the requirement for reductant 

recycling during the initial oxygenase reaction, a simplification 

offering significant ecological advantage (when compared with 

AOB) in nutrient poor environments. In this pathway, one of the 

MCO-like proteins may act as a nitroxyl oxidoreductase (NXOR) 

and facilitate the oxidation of nitroxyl to nitrite with the extraction 

of two protons and two electrons in the presence of water. 

The proposed NXOR would relay the two extracted electrons 

into the quinone pool via the QRED pathway described above. 

In this proposed model, the electrons extracted by either 

a CuHAO or a NXOR (and transferred into the quinone pool) 

would generate a proton motive force (PMF) through complexes 

III (plastocyanin-like subunit, Nmar_1542; Rieske-type subunit, 

Nmar_1544; transmembrane subunit, Nmar_1543) and IV 

(Nmar_0182-5), driving the generation of ATP by an F0F1-type 

ATP synthase (Nmar_1688–1693). The production of reductant 

(i.e., NADH) would require the reverse operation of complex I 

(NuoABCDHIJKMLN, Nmar_0276–286) as a quinol oxidase 

driven by a PMF. The proposed biochemistry involving nitroxyl 

produces the same net gain as bacterial ammonia oxidation, 

providing two electrons for reduction of the quinone pool and 

subsequent linear electron flow and the generation of a PMF. 

The presence of a copper-containing (versus heme) complex III 

and the unique evolutionary placement of terminal oxidase 

(complex IV) between two of the heme–copper oxygen reductase 

families further distinguish this proposed ammonia oxidation 

pathway from that in AOB. 

Carbon Fixation and Mixotrophy. N. maritimus, like all known 

AOB, grows chemolithoautotrophically by using inorganic carbon 

as the sole carbon source (7, 32). However, whereas AOB use the 

Calvin–Bassham–Benson cycle with the CO 2-fixing enzyme ribulose 

NH3 + O2 Out 

In 

H 2O + NH 2OH 

H O + HNO 

2 

AMO 

2 e - 

Q 

QH2 Ammonia Metabolism 

CuHAO 

NXOR 

QRED 

2 H + 

4 e- 

2 e- 

Q 

QH2 H + 

4H+ + HNO2 

2H+ + HNO 

C I 

H + 

NDH I 

Nitrite Reduction 

Q 

QH2 NAD 

Complex I 

+ + + H NADH 

H + 

2 

CuNIR 

NO + H 2 O 

pcy 

pcy 

H + 2 H + 

pcy 

aa 3 

bisphosphate carboxylase/oxygenase (RubisCO) as the key enzyme, 

the absence of genes in N. maritimus coding for RubisCO and other 

enzymes of this cycle points to an alternative pathway for carbon 

fixation. The most likely mechanism supported by the genome sequence 

is the 3-hydroxypropionate/4-hydroxybutyrate pathway elucidated 

in the thermophilic crenarchaote Metallosphaera sedula and 

suggested as a potential pathway of carbon fixation in C. symbiosum 

(33). The pathway has two parts: a sequence including two carboxylation 

reactions transforming acetyl-CoA to succinyl-CoA and 

amultistepsequenceconvertingsuccinyl-CoA into two molecules of 

acetyl-CoA. Genes identified in the N. maritimus genome coding for 

key enzymes of the pathway (Fig. S3) includeabiotin-dependent 

acetyl-CoA/propionyl-CoA carboxylase (Nmar_0272–0274), methylmalonyl-CoA 

epimerase and mutase (Nmar_0953, _0954, and 

_0958), and 4-hydroxybutyrate dehydratase (Nmar_0207). With the 

exception of one gene (Nmar_1608), all of the genes implicated in the 

3-hydroxyproprionate/4-hydrobutyrate pathway for carbon assimilation 

in N. maritimus are present and show highest similarity to the 

genes of C. symbiosum (34, 35). Although N. maritimus and M. sedula 

most likely use the same CO 2-fixation reaction sequences, not all 

individual reactions appear to be catalyzed by identical enzymes. In 

one instance, the stepwise reductive transformation of malonyl-CoA 

to propionyl-CoA involves five enzymes in M. sedula (33, 36, 37). 

Although the N. maritimus genome lacks any close homologs of 

the M. sedula genes, it contains alternative alcohol dehydrogenases, 

aldehyde dehydrogenases, acyl-CoA synthetases, and enoyl-CoA 

hydratases possibly fulfilling the same functions. Similarly, M. sedula 

catalyzes the activation of 3-hydroxypropionate to 3-hydroxypropionyl- 

CoA, using an AMP-forming 3-hydroxypropionyl-CoA synthetase 

(37). The N. maritimus genome lacks an obvious homolog, although 

does code for an ADP-forming acyl-CoA synthetase (Nmar_1309) that 

suggests a more energy efficient alternative. 

In addition to the genes coding for the 3-hydroxypropionate/4hydroxybutyrate 

pathway, the genome of N. maritimus contains 

a number of genes encoding enzymes of the tricarboxylic acid 

(TCA) cycle. No homologs for genes coding for a citrate-cleaving 

enzyme (ATP citrate lyase or citryl-CoA lyase) (38) were identified, 

permitting exclusion of the reductive TCA cycle as a pathway 

for carbon fixation. The lack of these genes suggests that N. maritimus 

utilizes either an incomplete (or horseshoe-type) TCA cycle 

for strictly biosynthetic purposes or possibly a complete oxidative 

TCA cycle. 

N. maritimus grows on a completely inorganic medium, indicating 

the genes coding for essential biosynthetic capacity (SI 

Materials and Methods), yet its genomic inventory also suggests 

some flexibility in the utilization of organic sources of phosphorus 

and carbon. Two systems for phosphorous acquisition are 

suggested: the high-affinity, high-activity phosphate transport 

system (pstSCAB, Nmar_0479, Nmar_0481–0483) and a phosphonate 

transporter (Nmar_0873–0875). However, because the 

genome lacks genes encoding known C-P lyases and hydrolases 

(39), and phosphate limitation is not alleviated by supplementation 

with phosphonates common in the marine environment 

4 H + 

ATPase 

H 

Complex III Complex IV Complex V 

+ 

4 H + 

0.5 O2 H O 2 

4 H + 

ADP + Pi ATP 

Fig. 1. Proposed AOA respiratory pathway. Text indicates 

the described possible hydroxylamine (blue text 

and arrows) and nitroxyl (green) pathways. Red arrows 

indicate electron flow not involved in ammonia oxidation. 

Blue shading denotes blue copper-containing proteins. 

Pink box indicates possible alternative respiratory 

electron sink. Hexagons containing Q and QH2 represent 

the oxidized and reduced quinone pools, respectively. 

Walker et al. PNAS Early Edition | 3of6 

MICROBIOLOGY

(e.g., aminoethylphosphonate), there is as yet no support for 

a functional phosphonate utilization pathway. Numerous organic 

transport functions are also evident, broadly encompassing transporters 

for different amino acids, dipeptides/oligopeptides, sulfonates/taurine, 

and glycerol. Additional physiological characterization 

will likely demonstrate some capacity for mixotrophic 

growth, as suggested by isotopic studies of natural populations 

(40–42). No genomic evidence exists for growth on urea, as 

N. maritimus lacks the homologs of the putative urease and urea 

transporter genes identified in C. symbiosum (35). 

Noncoding RNA Genes. The N. maritimus genome contains a full 

complement of essential noncoding RNA (ncRNA) genes, including 

one copy each of 5S/16S/23S ribosomal RNAs, RNase P, 

SRP RNA, and 44 transfer RNAs (Table S3). In addition to six 

normally placed canonical tRNA introns, noncanonical introns 

were found at different positions in six of the tRNAs (ValCAC, Met, 

Trp, Arg CCT, Leu TAA, and Glu TTC), a phenomenon previously 

observed only in thermophiles and C. symbiosum (34, 43). All 

other sequenced crenarchaea (including C. symbiosum) contain at 

least 46 tRNAs. N. maritimus lacks tRNA sequences coding for 

ProCGG or ArgCCG, perhaps resulting from (or related to) the low 

G + C content of the genome and preference for protein codons 

ending in A/T. Other archaeal genomes with relatively low G + C 

content, such as the euryarchaeon Haloquadratum walsbyi, also 

lack these tRNAs, while possessing the exact complement of 44 

tRNAs found in N. maritimus (43). This occurrence likely reflects 

a difference in posttranscriptional modification of the wobble base 

of tRNAs ProTGG and ArgTCG, allowing more efficient decoding 

of the rare codons CCG and CGG, respectively (44). 

Six candidates for C/D box small RNAs (sRNAs) were identified 

(Nmar_sR1–sR6). Most C/D box sRNAs guide the precise positioning 

of posttranscriptional 2′-O-methyl group addition to rRNAs 

or tRNAs, a process also occurring in eukaryotes, but not Bacteria.In 

N. maritimus,predictedtargetsof2′-O-methylation may include the 

wobble base of the Leu CAA tRNA and two different positions separated 

by 26 nucleotides in the 23S rRNA. Before the N. maritimus 

and C. symbiosum genomes, multiple C/D box sRNAs were found 

almost exclusively in hyperthermophilic archaea (45). Detection of 

these conserved, syntenic guide sRNAs in the two mesophilic crenarchaeal 

genomes supports an RNA stabilization/chaperone function 

not seen in other archaeal mesophiles and possibly more similar 

to their predicted function(s) in eukaryotes (46). 

Regulation of Transcription. The genome contains at least eight 

transcription factor B (TFB) and two TATA-box binding protein 

A B 

(TBP) (Table S3) genes required for starting site-specific transcription 

initiation, making N. maritimus among the densest and 

richest archaeal genomes for these transcription factors. TFB 

and TBP are thought to serve functions similar to the bacterial 

sigma factors (e.g., modulating cellular function in response to 

fluctuating environmental conditions) in Archaea with genomes 

coding for multiple copies, with optimal TFB/TPB partners (47). 

Although many other archaeal genomes contain multiple copies 

of these transcription factors, only the haloarchaea have more 

than five TFB genes (47). The functional significance of this 

exceptionally high density of regulatory factors in an apparently 

metabolically specialized organism will likely be informed by 

future transcriptional analyses of different growth states. Genes 

for two widely distributed types of archaeal chromatin proteins 

are present, an archaeal histone (Nmar_0683) and two Alba 

genes (Nmar_0255 and Nmar_0933). These are thought to 

maintain chromosomal material in a state permitting polymerase 

accessibility, with differential expression possibly providing for 

altered global transcription (48). 

Unique Cell Division Machinery and Previously Uncharacterized Instance 

of Archaeal Biosynthesis of Hydroxyectoine. The N. maritimus genome 

contains elements homologous with two systems of cell division: ftsZ 

(Nmar_1262) and cdvABC (Nmar_0700, _0816, and _1088). The 

cdvABC operon, induced at the onset of genome segregation and cell 

division, codes for machinery related to the eukaryotic endosomal 

sorting complex (49, 50). With the exception of N. maritimus, C. 

symbiosum, andtheThermoproteales(wherethecelldivisionmachinery 

remains uncharacterized), all available archaeal genomes 

have either the FtsZ or the Cdv cell division machinery, but not both. 

The two cell division systems may comprise a hybrid mechanism or 

serve two distinct processes in marine Crenarchaeota. 

The genome of N. maritimus also encodes for synthesis of 

the compatible solute hydroxyectoine: ectoine synthase (ectC,Nmar_ 

1344), a L-2,4-diaminobutyrate transaminase (ectB, Nmar_1345), 

aL-2,4-diaminobutyrateacetyltransferase(ectA,Nmar_1346),andan 

ectoine hydroxylase (ectD,Nmar_1343).Althoughwidelydistributed 

among Bacteria (in particular the genome sequences of marine 

organisms), the presence of these genes in N. maritimus represents 

auniqueindicationofarchaeal biosynthesis. 

Relationship to C. symbiosum. The genome of N. maritimus differs 

significantly in G + C content and size from that of the closely related 

sponge symbiont (∼97% 16S rRNA gene sequence identity). 

Despite the differences in overall genomic G + C content (34.2% 

for N. maritimus versus 57.7% for C. symbiosum), the G + C content 

Fig. 2. Synteny plots comparing the N. maritimus genome with (A) the Cenarchaeum symbiosum Atypegenome,(B) crenarchaealgenomefragments,and 

(C) Sargasso Sea fosmids. Vertical gray bar indicates location of ribosomal RNA operon. 

4of6 | www.pnas.org/cgi/doi/10.1073/pnas.0913533107 Walker et al. 

C

Fig. 3. Comparison of N. maritimus genome to metagenomic sequence reads 

from GOS. (A) Percentage of reads from each GOS site that align to the N. 

maritimus genome by protein sequence similarity. (B) NumberofGOSreads 

homologous to each N. maritimus protein-coding gene. Counts of 40 are mostly 

due to highly conserved proteins that include contaminants from other clades. 

for the rRNAs is largely similar between both organisms (50–52%). 

The higher ORF density (1.19 ORF/kb) relative to C. symbiosum 

(0.986 ORF/kb) results principally from the 0.4 Mbp smaller genome 

of N. maritimus, not from a large disparity in the number of 

predicted ORFs. The two genomes share 1,267 genes in common 

(when compared via reciprocal BLAST with expectation cutoff 

values of 10 −4 ), yet there is little conservation of synteny (Fig. 2C, 

Table S4). Most of the increased size of the C. symbiosum genome 

and much of the divergence in gene content are associated with 

discrete regions (“islands”), although no obvious functionality could 

be assigned to individual islands. Homologs for a majority of genes 

implicated in the archaeal ammonia oxidation pathway (51 of 69 

listed in Table S2) appear present in C. symbiosum. 

Phylogeny and Evolution. The widely distributed Group I archaeal 

lineage with which N. maritimus is affiliated was earlier assigned to 

the hyperthermophilic Crenarchaeota (3). Questions regarding this 

association arose through phylogenetic analysis of C. symbiosum 

ribosomal proteins, indicating possible divergence before the 

Crenarchaeota–Euryarchaeota split and therefore deserving provisional 

assignment to a new archaeal kingdom, the Thaumarchaeota 

(51). The basal position of the Group I archaea previously 

inferred from protein sequences encoded by the C. symbiosum genome 

was reexamined by reanalysis of the combined dataset, using 

patterns of gene distribution (Table S5) and phylogeny inference. 

Maximum-likelihood analyses confirmed the basal branching with 

significant statistical support (bootstrap value = 90%, Fig. S4). 

Bayesian analysis of a selection of species from the same dataset 

produced results linking C. symbiosum and N. maritimus as sister 

taxa of Crenarchaeota, albeit with nonsignificant support (posterior 

probability = 0.88). Although a definitive placement within the 

Archaea still must be confirmed by inclusion of genomes from more 

distantly related lineages, both analyses strongly support a lineage 

distinct from all other cultivated Crenarchaeota. 

High Similarity to the Metagenome of the Globally Distributed Marine 

Group I Archaea. The genome of N. maritimus shares remarkable 

conservation of gene content and gene order with numerous archaeal 

sequences previously recovered in fosmid libraries and recent 

oceanic surveys. The Antarctic genome fragments DeepAnt- 

EC39 (taken from 500 m depth) (52) and cosmid 74A4 (from surface 

waters) (53) both share very high synteny with portions of the N. 

maritimus genome (Fig. 2B and Table S6) despite significant differences 

in rRNA sequence identity (93 and 98%, respectively). 

Sixteen Sargasso Sea contigs (93% 16S rRNA gene sequence 

identity with N. maritimus) also have high synteny (Fig. 2C and 

Table S6). Retrieval of DNA fragments from the Global Ocean 

Sampling (GOS) database using differential protein sequence 

similarity showed that N. maritimus-like sequences constituted an 

average of 1.15% of all sequences across a wide range of temperatures 

(9–29 °C), salinities (freshwater to seawater, 0.1–63 practical 

salinity units), two open oceans, and several coastal environments 

(Fig. 3A). The Block Island, NY, coastal site and the Lake Gatun, 

Panama Canal, site (neither of which share any notable physical/ 

chemical characteristics other than sample depth) both exhibited 

notably large increases in density (>2.5%). The available GOS 

sequences provided almost complete and uniform coverage of the 

N. maritimus genome (Fig. 3B), although at least three significant 

gaps in coverage exist (possibly corresponding with unique 

N. maritimus genomic islands). Whereas some of the coverage may 

result in matches to bacterial sequences, particularly for very highly 

conserved proteins, the majority of recruited proteins had >50% 

sequence identity to N. maritimus proteins. Together, these shared 

genomic features suggest N. maritimus is representative of many of 

the globally abundant marine Group I Crenarchaeaota and that it 

should provide a useful model for developing an understanding of 

the basic physiology of these abundant and cosmopolitan organisms. 

A comparison of the sequence content and genome organization 

hints at functionally more divergent marine population 

types. N. maritimus has limited syntenic similarity to a deepwater 

population represented by the North Pacific fosmid 4B7 

(93% 16S rRNA sequence identity), but shares proteins highly 

similar to most of those coded on this fosmid. Previous comparison 

of marine crenarchaeal genomic fragments reported 

changes in genomic organization with sampling depths, suggestive 

of depth-related habitat types (54). Coupled with recent 

evidence indicating varied physiological lifestyles along depth 

and latitudinal gradients, distinct crenarchaeal ecotypes likely 

exist, analogous to that observed in marine cyanobacteria (2, 55). 

However, no clear correlations currently exist between environmental 

parameters and the similarity of recruited fragments. 

The genome sequence presented here also offers further insight 

into the ecological success of AOA. For example, using the likely more 

energy-efficient 3-hydroxypropionate/4-hydroxybutyrate pathway for 

CO2 fixation rather than the Calvin–Bassham–Benson cycle used by 

AOB could provide a growth advantage. Further ecological advantage 

may be conferred by their potential capacity for mixotrophic growth or 

the use of copper as a major redox-active metal for respiration in 

generally iron-limited oceans. However, a deeper understanding of 

the remarkable success of this archaeal lineage will come only from 

more detailed physiological, biochemical, and genetic characterization 

of N. maritimus and additional environmental isolates. 

Materials and Methods 

Genome sequencing was performed on high-molecular-weight DNA 

extracted from two cultures of N. maritimus. Whole-genomeshotgunsequencing 

of 3-, 8-, and 40-kb DNA libraries by the Joint Genome Institute 

produced at least 8× coverage. Annotation of the closed genome was performed 

using Department of Energy (DOE) computational support at Oak 

Ridge National Laboratory and The Institute for Genomic Research (TIGR) 

Autoannotation Service in conjunction with Manatee visualization software. 

Complete details describing high-molecular-weight DNA purification and 

sequence analysis are found in SI Materials and Methods. 

ACKNOWLEDGMENTS. The authors thank David Bruce and Paul Richardson 

from the Joint Genome Institute for facilitating genome sequencing. This 

Walker et al. PNAS Early Edition | 5of6 

MICROBIOLOGY

work was supported by the Department of Energy Microbial Genome 

Program, by National Science Foundation Microbial Interactions and Processes 

Grant MCB-0604448 (to D.A.S. and J.R.d.l.T.), by National Science Foundation 

Molecular and Cellular Biosciences Grant MCB-0920741 (to D.A.S.), by National 

Science Foundation Biological Oceanography Grants OCE-0623174 (to D.A.S.) 

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6of6 | www.pnas.org/cgi/doi/10.1073/pnas.0913533107 Walker et al.

Supporting Information 

Walker et al. 10.1073/pnas.0913533107 

SI Materials and Methods 

High-Molecular-Weight Genomic DNA Preparation. Two cultures of 

Nitrosopumilus maritimus strain SCM1 were grown as previously 

described in 500 mL of media in 1-L flasks (1). Cells from both 

cultures were harvested in late-exponential phase using Sterivex 

filters for one culture and a 0.1-μm filter for the other. Highmolecular-weight 

DNA was isolated as previously described using 

either agarose plugs (2) or a modified guanadinium thiocynate 

protocol (3). Cells from the Sterivex filter were resuspended in 

1 mL of 2× STE buffer [1 M NaCl, 0.1 M EDTA (pH 8.0), 10 mM 

Tris (pH 8.0)], extracted from the filter, and mixed with 1 vol of 1% 

molten SeaPlaque LMP agarose (FMC). The mixture was cooled 

to 40 °C, immediately drawn into a 1-mL syringe, and placed on ice 

for 10 min. The agarose plug was mixed with 10 mL of lysis buffer, 

incubated at 37 °C for 1 h, and then transferred to 40 mL of ESP 

buffer (1% Sarkosyl–1 mg of Proteinase K per ml in 0.5 M EDTA). 

After incubation at 55 °C for 16 h, the solution was replaced with 

fresh ESP buffer and incubated at 55 °C for another hour. DNA 

was purified using phenol:chloroform:isoamyl alcohol (24:24:1) 

and recovered by precipitation with isopropanol. 

Cells collected on the 0.1-μm filter were resuspended in 100 μLof 

Tris-EDTA (pH 8.0) and 100 mg/mL lysozyme before incubating at 

37 °C for 30 min. Then, 3.0 mL of a solution containing 5 M guanidinium 

thiocyanate, 100 mM EDTA (pH 8.0), and 0.5% (vol/vol) 

sarkosyl was added. The solution was mixed gently for 15 min before 

being cooled on ice for 10 min. After cooling, an equal volume of cold 

7.5 M ammonium acetate was added, and the solution was mixed 

gently and cooled on ice. Purification and precipitation of DNA were 

performed with chloroform:isoamyl alcohol (24:1) and isopropanol. 

Genome Sequencing. Acompletelysequencedandclosedgenome 

of N. maritimus was obtained through collaboration with the 

Joint Genome Institute. Whole-genome shotgun sequencing of 

3-, 8-, and 40-kb DNA libraries produced at least 8× coverage of 

the entire genome. Specifics of clone library generation, sequencing, 

and assembly strategies may be found at the DOE JGI 

website (www.jgi.doe.gov/sequencing/index.html). 

Genome Sequence Analysis. Autoannotation of the closed genome 

sequence was performed by both the Computational Biology 

group at Oak Ridge National Laboratory (http://genome.ornl. 

gov/microbial/nmar/02jul07/) and the TIGR Autoannotation 

Service (now hosted by JCVI; details available from http://www. 

jcvi.org/cms/research/projects/annotation-service/overview/). The 

genome visualization software Manatee (release 2.4.1; latest 

version available from http://manatee.sourceforge.net/) was used 

for manual curation. Analysis of potential transporter genes was 

performed using the Transporter Automatic Annotation Pipeline 

(TransAAP) through the TransportDB genomic comparison 

tool (membranetransport.org). 

Direct comparisons with the C. symbiosum genome and genome 

fragments were performed using the Artemis Comparison Tool (3) 

1. Könneke M, et al. (2005) Isolation of an autotrophic ammonia-oxidizing marine 

archaeon. Nature 437:543–546. 

2. Stein LY, et al. (2007) Whole-genome analysis of the ammonia-oxidizing bacterium, 

Nitrosomonas eutropha C91: Implications for niche adaptation. Environ Microbiol 9: 

2993–3007. 

3. Carver TJ, et al. (2005) ACT: The Artemis comparison tool. Bioinformatics 21:3422–3423. 

4. Bland C, et al. (2007) CRISPR recognition tool (CRT): A tool for automatic detection of 

clustered regularly interspaced palindromic repeats. BMC Bioinformatics 8:209. 

5. Brochier-Armanet C, Boussau B, Gribaldo S, Forterre P (2008) Mesophilic Crenarchaeota: 

Proposal for a third archaeal phylum, the Thaumarchaeota. Nat Rev Microbiol 6:245–252. 

with a comparison library generated through WebACT (www. 

webact.org/WebACT/home). Orthologous genes shared between 

these two organisms were identified through reciprocal BLAST 

searches, with an expectation cutoff value of 10 −4 and a minimum of 

75% alignable N. maritimus sequence. Comparisons with the 

Global Ocean Sampling (GOS) and Sargasso Sea metagenomic 

datasets were performed using several single-copy universal archaeal 

genes to determine a count of ∼15 archaeal genomes in the 

GOS dataset. An initial set of candidate N. maritimus-like proteins 

was found by BLASTP searches with each N. maritimus proteincoding 

gene, using a cutoff of 100 hits and a maximum expected 

value of e = 10. This set consisted of 125,326 peptides drawn from 

107,223 scaffolds. Neighboring ORFs on any scaffold with two or 

more hits were added to make a total of 319,585 peptides, 

amounting to 5.2% of all ORFs in GOS. These sequences were 

filtered by BLAST alignment to four sequence datasets, containing 

24 complete proteomes from diverse euryarchaeota, crenarchaeota, 

and bacteria. Sequences scoring more highly to N. maritimus and/or 

C. symbiosum proteins than any other entry in these datasets were 

retained, giving a filtered dataset of 21,278 ORFs. Coverage of the 

N. maritimus proteome was measured by bidirectional BLASTP of 

N. maritimus proteins against the filtered dataset to assign putative 

orthologs. The average coverage was ∼11×, although some highly 

conserved genes had a much greater number of hits, probably due to 

recruitment of nonarchaeal homologs: 48 ORFs had >30 hits, of 

which most were highly conserved. Searches for CRISPR regions 

were performed using the Java-based CRISPR recognition tool 

(CRT) with least stringent settings (4). 

Maximum-Likelihood and Bayesian Trees of the Archaeal Domain. 

The trees are based on the concatenation of ribosomal proteins 

used by Brochier-Armanet et al. (5) but including sequences from N. 

maritimus and from “Candidatus Korarchaeum cryptofilum” OPF8. 

Sequences were aligned using MUSCLE (6). Resulting alignments 

were visually inspected and improved with the MUST software (7). 

Regions where homology between sites was doubtful were removed 

from further phylogenetic analyses. A total of 6,142 positions were 

kept for the phylogenetic analyses. The maximum-likelihood tree 

was computed with PHYML, using the WAG model corrected by 

a gamma law to take into account evolutionary rate among site 

variations (8). The parameter alpha of the gamma distribution as the 

proportion of invariable sites was estimated from the dataset. The 

robustness of each branch was estimated by the bootstrap procedure 

implemented in PHYML. A Bayesian tree analysis on a subset of 29 

taxa was performed using MrBayes 3.2 (9) with a mixed model of 

amino acid substitution and a gamma distribution (eight discrete 

categories and an estimated proportion of invariant sites) to take 

into account among-site rate variation. MrBayes was run with four 

chains for 1 million generations and trees were sampled every 100 

generations. To construct the consensus tree, the first 1,500 trees 

were discarded as ‘‘burn-in.’’ The reduction of the taxonomic sampling 

was necessary to reduce the computation time. 

6. Edgar RC (2004) MUSCLE: Multiple sequence alignment with high accuracy and high 

throughput. Nucleic Acids Res 32:1792–1797. 

7. Philippe H (1993) MUST, a computer package of management utilities for sequences 

and trees. Nucleic Acids Res 21:5264–5272. 

8. Guindon S, Gascuel O (2003) A simple, fast, and accurate algorithm to estimate large 

phylogenies by maximum likelihood. Syst Biol 52:696–704. 

9. Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under 

mixed models. Bioinformatics 19:1572–1574. 

Walker et al. www.pnas.org/cgi/content/short/0913533107 1of5

0.5 

A. 

B. 

Thermoplasma volcanium GSS1 [rusticyanin; gi13542283] 

Acidithiobacillus ferrooxidans [rusticyanin; gi3282064] 

Thermobaculum terrenum ATCC BAA-798 [plastocyanin; gi227373287] 

Sphaerobacter thermophilus DSM 20745 [plastocyanin; gi229876615] 

Streptomyces sp. AA4 [hypothetical protein; gi256667890] 

Burkholderia cenocepacia PC184 [plastocyanin; gi254250279] 

95 

Catenulispora acidiphila DSM 44928 [blue (type 1) copper domain protein; gi256391720] 

Mycobacterium intracellulare ATCC 13950 [hypothetical protein; gi254819204] 

83 Mycobacterium avium subsp. avium ATCC 25291 [copper-binding protein; gi254776845] 

99 Mycobacterium avium 104 [copper-binding protein; gi118462975] 

Nmar0747 

96 Nmar0361 

Candidatus Koribacter versatillis [plastocyanin-like protein; gi94968815] 

Nmar0902 

Chromobacterium violaceum ATCC 12472 [hypothetical protein; gi34498952] 

Methylocella silvestris BL2 [blue (type 1) copper domain protein; gi217976580] 

Methylobacterium nodulans ORS [blue (type 1) copper domain protein; gi220920817] 

Methylocella silvestris BL2 [blue (type 1) copper domain protein; gi217976580] 

61 Nitrobacter hamburgensis X14 [blue (type 1) copper domain protein; gi92118183] 

Bradyrhizobium japonicum USDA 110 [putative Amicyanin precursor; gi27378126] 

Paracoccus denitrificans PD1222 [amicyanin; gi119387457] 

Nmar0190 

Methanosarcina mazei Go1 [hypothetical protein; gi21226177] 

64 Methanosarcina acetivorans C2A [hypothetical protein; gi20090537] 

C. symbiosum A [copper-binding protein; gi118575216] 

99 

Nmar0167 

99 Nmar0621 

Methanosarcina mazei Go1 [copper-binding protein; gi21228445] 

Methanosarcina acetivorans C2A [plastocyanin/azurin family copper-binding protein; gi20090218] 

62 

Methanosarcina mazei Go1 [copper-binding protein; gi21228446] 

100 Methanosarcina acetivorans C2A [plastocyanin/azurin family copper-binding protein; gi20090219] 

Jannaschia sp. CCS1 [blue (type 1) copper domain protein; gi89056161] 

Nmar0718 

Thermobaculum terrenum ATCC BAA-798 [plastocyanin; gi227373254] 

Deinococcus deserti VCD115 [copper-binding protein; gi226356528] 

Synechococcus sp. WH 8102 [plastocyanin; gi33866031] 

91 Prochlorococcus marinus MIT 9312 [plastocyanin; gi78778966] 

88 

Anabaena variabilisATCC 29413 [plastocyanin; gi75908957] 

79 

Gloeobacter violaceus PCC 7421 [plastocyanin; gi35212909] 

Methanococcus maripaludis C7 [copper-binding protein; gi150402166] 

82 Methanococcus maripaludis S2 [copper-binding protein; gi45358560] 

Nmar1913 

82 

Nmar1789 

Nmar0300 

Nmar0762 


100 C. symbiosum A [copper-binding protein; gi118576965] 

Nmar0516 

Nmar1087 

C. symbiosum A [hypothetical protein; gi118575848] 

98 C. symbiosum A [hypothetical protein; gi118575831] 


79 

Nmar0121 

uncultured marine Crenarchaeote [putative copper-binding protein; gi167045254] 

100 uncultured marine Crenarchaeote [putative copper-binding protein; gi167044521] 

100 uncultured marine Crenarchaeote [putative copper-binding protein; gi167042919] 


100 Nmar0273 

Nmar0732 

Nmar0121 

Nmar0324 

79 

Nmar0167 

80 


Nmar0190 

Nmar0273 

Nmar0300 

Nmar0324 

Nmar0361 

Nmar0516 

Nmar0621 

Nmar0718 

Nmar0732 

Nmar0747 

Nmar0762 

Nmar1087 

Nmar1789 

Nmar1913 

100 aa 

0.1 

Bacillus subtilis subsp. subtilis str. 168 [BdbD thiol-disulfide oxidoreductase; gi 16080401] 

Bacillus amyloliquefaciens FZB42 [BdbD; gi 154687467] 

Paenibacillus larvae subsp. larvae BRL-230010 [disulfide dehydrogenase D; gi 167461952] 

Deinococcus geothermalis DSM 11300 [DsbA oxidoreductase; gi 94984799] 

Thermobaculum terrenum ATCC BAA-798 [protein-disulfide isomerase; gi 227374421] 

100 

C. 

100 

87 

82 

75 

Gemmatimonas aurantiaca T-27 [putative oxidoreductase; gi 226228008] 

Sphaerobacter thermophilus DSM 20745 [protein-disulfide isomerase; gi 229876989] 

Thermomicrobium roseum DSM 5159 [DsbA oxidoreductase; gi 221635547] 

Thermus aquaticus Y51MC23 [DsbA oxidoreductase; gi 218295130] 

Thermobifida fusca YX [protein-disulfide isomerase; gi 72161925] 

Mycobacterium kansasii ATCC 12478 [DsbA oxidoreductase; gi 240169146] 

Rubrobacter xylanophilus DSM 9941 [DsbA oxidoreductase; gi 108804655] 

Streptomyces coelicolor A3(2) [hypothetical protein SCO5993; gi 21224330] 

100 

Streptomyces ghanaensis ATCC 14672 [hypothetical protein SghaA1_08748; gi 239928300] 

Symbiobacterium thermophilum IAM 14863 [hypothetical protein STH2058; gi 51893196] 

Roseiflexus castenholzii DSM 13941 [DsbA oxidoreductase; gi 156741642] 

100 

99 

100 

Chloroflexus aggregans DSM 9485 [DsbA oxidoreductase; gi 219849651] 


Marinobacter algicola DG893 [DsbA oxidoreductase; gi 149377658] 

Vibrio harveyi HY01 [DsbA oxidoreductase; gi 153832115] 

Vibrio parahaemolyticus RIMD 2210633 [hypothetical protein VPA0994; gi 28900849] 

Moritella sp. PE36 [putative membrane protein; gi 149908466] 

99 

Shewanella benthica KT99 [hypothetical protein KT99_10483; gi 163751466] 

Roseiflexus castenholzii DSM 13941 [protein-disulfide isomerase-like protein; gi 156741356] 

75 

Roseiflexus castenholzii DSM 13941 [DsbA oxidoreductase; gi 156743646] 

Chloroflexus aurantiacus J-10-fl [DsbA oxidoreductase; gi 163848707] 

Herpetosiphon aurantiacus ATCC 23779 [DsbA oxidoreductase; gi 159896786] 

72 

100 

Chloroflexus aurantiacus J-10-fl [DsbA oxidoreductase; gi 163845898] 


Stigmatella aurantiaca DW4/3-1 [conserved hypothetical protein; gi 115374845] 

Solibacter usitatus Ellin6076 [DsbA oxidoreductase; gi 116625220] 

Solibacter usitatus Ellin6076 [DsbA oxidoreductase; gi 116624599] 

82 

Syntrophobacter fumaroxidans MPOB [DsbA oxidoreductase; gi 116751066] 

Anaeromyxobacter dehalogenans 2CP-1 [DsbA oxidoreductase; gi 220919173] 

100 

Myxococcus xanthus DK 1622 [putative lipoprotein; gi 108762810] 

Stigmatella aurantiaca DW4/3-1 [disulfide interchange protein; gi 115379912] 

Archaeoglobus fulgidus DSM 4304 [hypothetical protein AF1354; gi 11498950] 

Cenarchaeum symbiosum A [protein-disulfide isomerase; gi 118575694] 

Nmar0752 

93 

91 

Nmar0179 

Nmar0218 

73 

79 

Nmar0740 

Nmar0175 

Natrialba magadii ATCC 43099 ]DsbA oxidoreductase; gi 224823141] 

Nmar1863 

90 


Nmar1805 

67 

100 


Nmar1606 

72 

63 

88 

Nmar0164 

Nmar0169 

Nmar1590 


Fig. S1. Phylogeny of plastocyanin-like protein sequences. Sequences with significant matches to COG3794 (PetE: Plastocyanin [Energy production and 

conversion]) were used to query the non-redundant protein sequence database from NCBI. Sequences from the top 20–30 non-N. maritimus hits were retrieved 

and their match to the above conserved domain model verified. Sequences from experimentally characterized proteins were obtained from the 

available literature and included, aligned with ClustalW and then curated manually. Distance-based phylogenies were inferred in Phylip using the Neighbor- 

Joining algorithm and 100 bootstrap replicates. Bootstrap support values >60% are displayed. Nodes with

Fig. S2. Archaeal ammonia monooxygenase AmoB sequence mapped onto the crystal structure of the particulate methane monooxygenase (PDB accession 

code 1YEW). The pmoA subunit is shown in dark gray, the pmoC subunit in light gray, and the pmoB subunit in red and pink. The red part represents the 

region of pmoB conserved in the predicted N. maritimus AmoB. The transmembrane helix and C-terminal cupredoxin domain shown in pink are missing in the 

predicted N. maritimus AmoB. Cyan spheres represent copper ions. The grey sphere represents a zinc ion. 


Acetyl-CoA carboxylase 

(Nmar_0272, 0273, 0274) 

Acetoacetyl-CoA -ketothiolase 

(Nmar_0841 or Nmar_1631) 

Malonyl-CoA reductase 

Malonate semialdehyde reductase 

(unknow n) 

3-Hydroxypropionyl-CoA synthetase 

3-Hydroxypropionyl-CoA dehydratase 

Acryloyl-CoA reductase (unknow n) 

Propionyl-CoA carboxylase 

(Nmar_0272, 0273, 0274) 

Methylmalonyl-CoA epimerase 

Methylmalonyl-CoA mutase 

(Nmar_0953, 0954, 0958) 

3-Hydroxybutyryl-CoA dehydrogenase 

(Nmar_1028) 

Crotonyl-CoA hydratase 

(Nmar_1308) 

4-Hydroxybutyryl-CoA dehydratase 

(Nmar_0207) 

4-Hydroxybutyryl-CoA synthetase 

(Nmar_0206) 

Succinate semialdehyde reductase 

(Nmar_1110 or Nmar_0161) 

Succinyl-CoA reductase 

(Nmar_1608) 

Fig. S3. Proposed 3-hydroxypropionate/4-hydroxybutyrate cycle for autotrophic carbon fixation by N. maritimus. 


A. 

0.1 

B. 

100 

90 

52 

Other Supporting Information Files 

Table S1 (DOC) 






96 

100 

97 

78 

100 

100 

38 

100 

55 

95 

100 

61 

88 

64 

70 

100 

100 

100 

100 

100 

100 

100 

97 

51 

86 

100 

61 

100 

100 

100 

100 

90 

100 

100 

Giardia lamblia 

Entamoeba histolytica 

Leishmania major 

Cenarchaeum symbiosum 

Candidatus Nitrosopumilus maritimus 

Candidatus Korarchaeum cryptofilum 

Thermofilum pendens 

Caldivirga maquilingensis 

100 

94 

100 

Pyrobaculum calidifontis 

Pyrobaculum islandicum 

Pyrobaculum aerophilum 

Pyrobaculum arsenaticum 

Ignicoccus hospitalis 

Staphylothermus marinus 

Aeropyrum pernix 

Hyperthermus butylicus 

Metallosphaera sedula 

Sulfolobus solfataricus 

Thermococcus gammatolerans 

Thermococcus kodakarensis 

Pyrococcus furiosus 

Pyrococcus abyssi 

Pyrococcus horikoshii 

Methanopyrus kandleri 

100 

100 

100 

100 

100 

100 

100 

Trypanosoma cruzi 

Trypanosoma brucei 

Cryptosporidium parvum 

Theileria parva 

Plasmodium yoelii 

Plasmodium falciparum 

Arabidopsis thaliana 

Oryza sativa 

Dictyostelium discoideum 

Homo sapiens 

Anopheles gambiae 

Saccharomyces cerevisiae 

Schizosaccharomyces pombe 

Sulfolobus acidocaldarius 

Sulfolobus tokodaii 

Nanoarchaeum equitans 

Methanosphaera stadtmanae 

Methanothermobacter thermautotrophicus 

Methanocaldococcus jannaschii 

Methanococcus aeolicus 

100 

100 

Methanococcus maripaludis 

Methanococcus vannielii 

Picrophilus torridus 

Ferroplasma acidarmanus 

Thermoplasma acidophilum 

Thermoplasma volcanium 

100 

100 

100 

Archaeoglobus fulgidus 

69 

Halobacterium sp 

Natronomonas pharaonis 

71 

Haloarcula marismortui 

100 

Halorubrum lacusprofundi 

100 

Haloquadratum walsbyi 

100 Haloferax volcanii 

Methanocorpusculum labreanum 

100 

Methanoculleus marisnigri 

75 

Candidatus Methanoregula 

82 

Methanospirillum hungatei 

Methanosaeta thermophila 

Methanococcoides burtonii 

100 

Methanosarcina barkeri 

100 Methanosarcina acetivorans 

100 Methanosarcina mazei 

1.00 

Saccharomyces cerevisiae 

Dictyostelium discoideum 

Oryza sativa 

Candidatus Korarchaeum cryptofilum 

Candidatus Nitrosopumilus maritimus 

1.00 

Cenarchaeum symbiosum 

0.88 

Pyrobaculum aerophilum 

1.00 

1.00 

Thermofilum pendens 

1.00 1.00 

Aeropyrum pernix 

Hyperthermus butylicus 

0.88 

1.00 

1.00 

Sulfolobus acidocaldarius 

Metallosphaera sedula 

Nanoarchaeum equitans 

Thermococcus kodakarensis 

1.00 

1.00 Pyrococcus abyssi 

Methanopyrus kandleri 

1.00 

1.00 

Methanothermobacter thermautotrophicus 

1.00 

Methanosphaera stadtmanae 

1.00 

1.00 

Methanocaldococcus jannaschii 

Methanococcus maripaludis 

Thermoplasma volcanium 

1.00 

1.00 

Ferroplasma acidarmanus 

1.00 

Archaeoglobus fulgidus 

Haloarcula marismortui 

1.00 

1.00 

Natronomonas pharaonis 

1.00 1.00 

Methanosarcina mazei 

Methanosaeta thermophila 

0.70 

Methanospirillum hungatei 

0.1 

1.00 

Methanocorpusculum labreanum 

Eucarya 

Thaumarchaeota 

Korarchaeota 

Thermoproteales 

Desulfurococcales 

Sulfolobales 

Nanoarchaeota 

Thermococcales 

Methanopyrales 

Methanobacteriales 

Eucarya 

Methanococcales 

Thermoplasmatales 

Archaeoglobales 

Halobacteriales 

Methanomicrobiales 

Methanosarcinales 

Korarchaeota 

Thaumarchaeota 

Thermoproteales 

Desulfurococcales 

Sulfolobales 

Nanoarchaeota 

Thermococcales 

Methanopyrales 

Methanobacteriales 

Methanococcales 

Thermoplasmatales 

Archaeoglobales 

Halobacteriales 

Methanosarcinales 

Methanomicrobiales 

Fig. S4. (A) Maximum-likelihoodphylogenyofGroup1Archaea. The phylogeny was inferred using an alignment of concatenated R-proteins (66 taxa, 6,142 

positions). WAG+Inv+Gamma (4 classes); 100 replicates. (B) BayesiantreeofmesophilicGroup1Archaea inferred using an alignment of concatenated Rproteins 

(29 taxa, 6,142 positions). Mixed model + Gamma (4 classes); 100 replicates. 

Walker et al. www.pnas.org/cgi/content/short/0913533107 5of5 

Crenarchaeota Euryarchaeota 

Crenarchaeota Euryarchaeota

Supplemental Table 1: Number and genomic density of genes grouped as clusters of orthologous genes (COGs). 

Nitrosopumilus 

maritimus SCM1 

Nitrosococcus oceani 

ATCC 19707 

Nitrosomonas 

europaea ATCC 19718 

Nitrosomonas 

eutropha C71 

Nitrosospira 

multiformis ATCC 

25196 

Pelagibacter ubique 

HTCC1062 

Procholoroccus 

species 

Synechococcus 

species 

Archaeoglobus 

fulgidus DSM 4304 

Halobacterium sp. 

NRC-1 

Methanococcus 

maripaludis S2 

Thermoplasma 

acidophilum DSM 1728 

Genome size (Mb) 1.65 3.52 2.81 2.78 3.23 1.31 1.64-2.68 2.22-3.05 2.18 2.57 1.66 1.56 1.74-1.91 2.23-2.99 

ORFs 1,997 3,186 2,628 2,578 2,827 1,.389 1,901 - 3,152 2,580 - 3,401 2,471 2,674 1,772 1,548 1,879 - 2,229 2,305 - 3,031 

COG Genes 1,131 2,290 1,995 1,952 2,102 1,131 1,157-1,540 1,459-1,968 1,918 1,812 1,417 1,201 1,421-1,575 1,563-2,105 

# #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb # #/Mb 

Amino acid transport & 

141- 

114- 

150metabolism 

109 66 180 51 139 49 141 51 154 48 182 139 121-152 57-76 178 57-65 149 68 151 59 125 75 130 83 159 66-83 202 63-68 

Carbohydrate transport & 

88metabolism 

46 28 105 30 73 26 81 29 94 29 64 49 62-94 34-41 86-108 29-46 54 25 60 23 50 30 92 59 80-93 45-49 128 37-43 

Cell cycle control, cell division 11 7 31 9 39 14 32 12 31 10 13 10 15-18 6-10 18-33 7-11 23 11 31 12 16 10 8 5 18-21 9-12 8-15 4-5 

Cell motility 6 4 75 21 82 29 75 27 64 20 12 9 1-20 1-7 4-28 2-10 21 10 45 18 27 16 14 9 13-27 7-16 10-14 4-5 

Cell wall/membrane/envelope 

113biogenesis 

49 30 186 53 167 59 152 55 194 60 99 76 86-136 47-60 137 38-60 60 28 59 23 38 23 41 26 46-61 26-32 51-79 19-26 

Coenzyme transport & 

122- 

102metabolism 

95 58 114 32 102 36 99 36 108 33 77 59 106-122 45-66 143 47-56 114 52 109 42 126 76 86 55 70-89 40-47 110 37-49 

Defense mechanisms 13 8 59 17 49 17 41 15 31 10 11 8 14-33 8-14 23-37 9-14 18 8 22 9 11 7 12 8 16-29 9-15 17-22 7-8 

Energy production & 

110- 

104- 

144conservation 

100 61 182 52 122 43 142 51 160 49 99 76 82-106 40-52 139 43-51 227 104 130 51 151 91 103 66 128 60-68 176 59-65 

140- 

189- 99- 147- 

Function unknown 86 52 235 67 172 61 149 54 197 61 78 60 85-145 48-56 179 55-67 259 119 187 73 219 132 116 74 197 113 187 59-69 

171- 

262- 144- 272- 116- 

General function prediction only 

Inorganic ion tranport & 

152 92 282 80 221 79 196 70 228 70 112 86 141-189 70-91 270 77-98 331 152 276 107 207 125 186 119 287 162 346 122 

metabolism 

Intracellular trafficking & 

63 38 135 38 164 58 115 41 105 32 40 31 55-77 27-38 72-148 32-49 91 42 128 50 77 46 51 33 78-99 45-56 63-85 25-28 

secretion 11 7 91 26 91 32 107 38 96 30 38 29 20-52 12-21 27-47 12-17 25 11 23 9 21 13 20 13 20-21 10-12 14-21 5-7 

Lipid transport & metabolism 

Nucleotide transport & 

30 18 72 20 65 23 63 23 78 24 51 39 37-49 18-26 35-51 12-20 109 50 56 22 16 10 56 36 19-25 11-13 70-89 26-40 

metabolism 45 27 56 16 50 18 54 19 54 17 46 35 48-52 18-31 52-63 18-24 51 23 63 25 51 31 50 32 47-52 27-28 62-66 21-28 

Posttranslational modifications 74 45 130 37 103 37 127 46 129 40 61 47 70-95 35-48 90-119 36-42 69 32 105 41 53 32 49 31 51-55 29-30 67-88 27-33 

Replication, recombination & 

67- 

84- 38repair 

Secondary metabolites 

56 34 202 57 170 60 225 81 182 56 54 41 62-84 31-40 78-207 31-70 98 45 169 66 58 35 67 43 109 38-57 326 109 

biosynthesis & transport 15 9 44 12 36 13 49 18 59 18 37 28 19-46 11-17 27-47 9-18 33 15 26 10 5 3 22 14 10-12 6-7 35-53 13-24 

Singal transduction mechanisms 47 29 121 34 126 45 86 31 124 38 34 26 23-49 14-20 42-116 19-42 71 33 76 30 35 21 9 6 19-33 10-19 23-28 

95- 

8-10 

Transcription 76 46 96 27 113 40 92 33 99 31 42 32 36-63 21-26 60-93 24-34 97 45 122 47 75 45 71 45 80-85 42-49 107 34-43 

Translation, ribosomal structure 

137- 

160- 

165- 

& biogenesis 136 83 149 42 154 55 152 55 151 47 113 86 128-146 54-80 144 45-64 160 73 148 58 155 93 136 87 166 87-92 166 55-74 

Pyrococcus 

species 

Sulfolobus 

species

Supplemental Table 2: Predicted coordinates for genes implicated in novel archaeal biochemistry for ammonia oxidation and respiratory chain. 

NCBI locus 

tag 

Protein # Strand Left bp Right bp Annotation Best Hit Analog in Ammonia oxidizing Bacteria 

Nmar_0588 ABX12484 F 523,890 525,197 ammonium transporter AmtB 

Nmar_1698 ABX13594 R 1,559,816 1,558,251 ammonium transporter AmtB 

Nmar_1500 ABX13396 R 1,367,890 1,367,240 putative ammonia monooxygenase, subunit A AMO (AmoA) 

Nmar_1501 ABX13397 R 1,368,387 1,368,025 hypothetical protein 

Nmar_1502 ABX13398 R 1,369,079 1,368,507 putative ammonia monooxygenase, subunit C AMO (AmoC) 

Nmar_1503 ABX13399 F 1,369,326 1,369,895 putative ammonia monooxygenase, subunit B AMO (AmoB) 

Nmar_0815 ABX12711 R 718,761 720,380 

soluble periplasmic BCP (complete upstream CxxxxxC domain, complete 

CxxHxxM domain [C]) 

PetE, plastocyanin cytochrome c552 

Nmar_1102 ABX12998 F 


1,004,439 1,005,395 



Nmar_1443 ABX13339 R 


1,314,030 1,313,572 



Nmar_1637 ABX13533 R 


1,494,547 1,492,676 



Nmar_0004 ABX11904 F 3,463 4,359 

soluble periplasmic BCP (modified upstream CxxxxxC domain, modified 


Halocyanin, P39442 cytochrome c552 

Nmar_1307 ABX13203 F 


1,195,212 1,195,736 


Halocyanin, P39442 cytochrome c552 

Nmar_0918 ABX12814 R 802,019 802,861 

periplasmic membrane BCP (1 TMS [C], modified upstream CxxxxxC domain, 

complete CxxHxxM domain [M]) 

plastocyanin anchored cytochrome c552 

Nmar_1678 ABX13574 R 

periplasmic membrane BCP (1 TMS [C], modified upstream CxxxxxC domain, 

1,538,377 1,537,367 

complete CxxHxxM domain [N]) 


Nmar_1273 ABX13169 R 

periplasmic membrane BCP (1 TMS [C], complete upstream CxxxxxC domain, 

1,169,255 1,168,392 

modified CxxHxxM domain [C]) 


Nmar_1161 ABX13057 R 1063718 1062348 

periplasmic membrane BCP (1 TMS [N], modified upstream CxxxxxC domain, 

modified CxxHxxM domain [M]) 


Nmar_1226 ABX13122 F 

periplasmic membrane BCP (4 TMS [N], modified upstream CxxxxxC domain, 2 

1,126,185 1,127,261 

complete CxxHxxM domains [C]) 

plastocyanin cM552 Nmar_1142 ABX13038 F 

cytoplasmic membrane BCP (1 TMS [N], modified upstream CxxxxxC domain, 

1,045,689 1,046,150 

complete CxxHxxM domain [C]) 

Nmar_1542 "cytoplasmic BCP" 

Nmar_0276 ABX12172 F 243,850 244,182 CI, NADH-ubiquinone/plastoquinone oxidoreductase chain 3 

Nmar_0277 ABX12173 F 244,223 244,747 CI, NADH-quinone oxidoreductase, B subunit 

Nmar_0278 ABX12174 F 244,747 245,349 CI, NADH dehydrogenase (ubiquinone) 30 kDa subunit 

Nmar_0279 ABX12175 F 245,352 246,488 CI, NADH dehydrogenase (quinone) 

Nmar_0280 ABX12176 F 246,489 247,787 CI, NADH dehydrogenase (quinone) 

Nmar_0281 ABX12177 F 247,787 248,284 CI, [4Fe-4S] ferredoxin iron-sulfur binding domain protein Complex I 

Nmar_0282 ABX12178 F 248,277 248,789 CI, NADH-ubiquinone/plastoquinone oxidoreductase chain 6 

Nmar_0283 ABX12179 F 248,770 249,075 CI, NADH-ubiquinone oxidoreductase chain 4L 

Nmar_0284 ABX12180 F 249,075 250,628 CI, proton-translocating NADH-quinone oxidoreductase, chain M 

Nmar_0285 ABX12181 F 250,630 252,708 CI, proton-translocating NADH-quinone oxidoreductase, chain L 

Nmar_0286 ABX12182 F 252,721 254,205 CI, proton-translocating NADH-quinone oxidoreductase, chain N 

Nmar_1542 ABX13438 R 

CIII, cytoplasmic membrane BCP, 1 TMS (N), modified upstream CxxxxxC 

1,404,537 1,404,073 

domain, complete CxxHxxM domain (C) 

Nmar_1142 membrane-bound cytochrome c1 Nmar_1543 ABX13439 R 1,406,185 1,404,590 CIII, Cytochrome b/b6 domain Complex III cyt. B 

Nmar_1544 ABX13440 R 1,406,774 1,406,169 CIII, Rieske [2Fe-2S] domain protein Complex III [Fe-S] 

Nmar_0182 ABX12078 F 166,066 166,326 CIV, hypothetical protein HCO subunit IV 

Nmar_0183 ABX12079 F 166,329 166,760 CIV, heme-copper oxidase subunit II HCO subunit II 

Nmar_0184 ABX12080 F 166,798 168,324 CIV, heme-copper oxidase subunit I HCO subunit I 

Nmar_0185 ABX12081 F 168,337 169,284 

CIV, periplasmic membrane BCP, 1 TMS (N), complete upstream CxxxxxC 

domain, complete CxxHxxM domain (C) 

Nmar_1354 cytochrome c subunit III 

Nmar_1259 ABX13155 R 1,155,995 1,154,583 NirK, soluble periplasmic MCO (Cu-oxidase_1; Cu-oxidase_2; Cu-oxidase_3), 

Nwi_2648; 

Nham_3281; NE0924; 

Neut_1403; Noc_0089 

3dMCO, NirK 

Nmar_1661 ABX13557 R 1,519,832 1,519,440 Transcriptional regulator, MarR family (winged helix-turn-helix HxlR type) HTH -regulator MarR 

Nmar_1662 ABX13558 R 1,521,322 1,520,144 Metal cation transporter (ZIP Zinc transporter; cl00437) Zip 

Nmar_1663 ABX13559 R 

soluble periplasmic 2dMCO (Cu-oxidase_2; Cu-oxidase_3), similar to NcgA 

1,522,373 1,521,312 

(BCO) 

Nwi_2651; 

Nham_3284; NE0927; 

Neut_1406 

2dMCO, NcgA (BCO) 

Nmar_1664 ABX13560 R 1,522,974 1,522,483 metal (Mn, Fe) dependent repressor protein, DtxR family Nmar_1132 HTH -regulator DtxR 

Nmar_1665 ABX13561 R 


1,523,638 1,523,135 


Halocyanin, P39442 monoheme cytochrome c 

Nmar_1666 ABX13562 R 

beta-propeller structure oxidase (Kelch repeat-containing protein 

1,524,788 1,523,754 

[CDD:121499]) 

Nmar_1133 beta propeller structure 

Nmar_1667 ABX13563 R 1,526,542 1,525,130 

Nmar_1136 ABX13032 R 1,041,807 1,040,200 

NirK, soluble periplasmic 3d MCO (Cu-oxidase_1; Cu-oxidase_2; Cuoxidase_3), 

NsrR motif in upstream region. 

soluble periplasmic 3dMCO (Cu-oxidase_1; Cu-oxidase_2; Cu-oxidase_3), 

similar to NcgA 

Nwi_2648; 

Nham_3281; NE0924; 

Neut_1403; Noc_0089 

Nwi_2651; 

Nham_3284; NE0927; 

Neut_1406 

3dMCO, Nirk 

3dMCO- modified NcgA (BCO) 

Nmar_1135 ABX13031 R 1,040,071 1,038,395 putative dehydrogenase with Rossmann-fold NAD(P) + -binding domain [cl09931] Rossmann-fold - DH 


Nmar_1133 ABX13029 R 

beta-propeller structure oxidase (Kelch repeat-containing protein 

1,037,891 1,036,866 

[CDD:121499]) 

Nmar_1666 beta propeller structure 

Nmar_1132 ABX13028 R 1,036,419 1,035,937 metal (Mn, Fe) dependent repressor protein, DtxR family Nmar_1664 HTH -regulator DtxR 

Nmar_1131 ABX13027 R 

soluble periplasmic 2d MCO (Cu-oxidase_2; Cu-oxidase_3), similar to NcgA 

1,035,788 1,034,733 

(BCO) 

Nwi_2651; 

Nham_3284; NE0927; 

Neut_1406 

2dMCO NcgA (BCO) 

Nmar_1130 ABX13026 R 1,034,743 1,033,565 Metal cation transporter (ZIP Zinc transporter; cl00437) Zip 

Nmar_1129 ABX13025 R 

cytoplasmic membrane BCP (3 TMS [1N, 2C], complete upstream CxxxxxC 

1,031,846 1,033,210 

domain, complete CxxHxxM domain [N) 

"cytoplasmic BCP" 

Nmar_1128 ABX13024 R 

periplasmic binding protein (transport of ferric siderophores and metal ions such 

1,031,477 1,030,587 

as Mn2+, Fe3+, Cu2+ and/or Zn2+) 

cation BP 

Nmar_1352 ABX13248 R 1,238,660 1,237,860 Transcriptional regulator, ArsR family (winged helix-turn-helix type, COG4742) HTH -regulator 

Nmar_1353 ABX13249 R 1,239,270 1,238,728 Uncharacterized conserved protein [COG3945; Function unknown] 

Nmar_1354 ABX13250 R 

Fusion Protein: soluble periplasmic 2d MCO (Cu-oxidase_2; Cu-oxidase_3) - 

1,240,615 1,239,281 

BCP (modified upstream CxxxxxC domain & CxxHxxM domain [C]) 

Nmar_1131; 

Nmar_0185 

2dMCO - BCP bi-functional MCO 

Nmar_1355 ABX13251 R 1,242,507 1,241,821 hypothetical protein

Nmar_1356 ABX13252 R 1,243,134 1,242,610 IsiB, flavodoxin/nitric oxide synthase [cl00438] flavodoxin/nitric oxide synthase 

Nmar_1357 ABX13253 F 1,243,214 1,243,858 nitroreductase [Nitro_FMN_reductase] Nitro_FMN_reductase 

Nmar_0650 ABX12546 R 588,034 587,297 Nitrilase/cyanide hydratase and apolipoprotein N-acyltransferase 

Nmar_1650 ABX13546 R 

periplasmic membrane BCP (1 TMS [C], complete upstream CxxxxxC domain, 

1,508,803 1,509,615 

complete CxxHxxM domain [M]) 

plastocyanin monoheme cytochrome c 


Nmar_1652 ABX13548 F 

periplasmic Cu resistance membrane hybrid protein: CopC - [8 TMS] - CopD - [1 

1,510,361 1,513,144 

TMS] 

CENSYa_1798 copC-copD 3' of amoCAB(E)D 

Nmar_1653 ABX13549 R 1,513,952 1,513,134 ABC-3 protein; ABC-ATPase subunit interface [CDD:119348] 

Nmar_1654 ABX13550 R 1,514,677 1,513,955 ABC-type Mn/Zn transport systems, ATPase component [COG1121] 

Nmar_1655 ABX13551 R 

periplasmic solute binding protein [ABC transport of ferric siderophores and metal 

1,515,591 1,514,671 cation BP 

ions such as Mn 2+ , Fe 3+ , Cu 2+ and/or Zn 2+ ] 

NikR; transcriptional regulator with C-terminal nickel binding domain 

Nmar_1656 ABX13552 R 1,516,066 1,515,683 

[pfam08753] 

Nmar_1657 ABX13553 F 1,516,198 1,516,479 hypothetical protein 

protease inhibitor, cytoplasmic membrane BCP (1 TMS [N], complete upstream 

Nmar_1250 ABX13146 R 1,148,436 1,147,717 cytoplasmic protease inhibitor 

CxxxxxC domain, complete CxxHxxM domain [C]) 

Red labelling indicates presence in C. symbiosum genome. 

Green shading indicates gene cluster encoding a 2dMCO (Nmar_1663) and a 3dMCO-NirK (Nmar_1667) preceeded by a NsrR binding site. Both genes flank a gene encoding a soluble periplasmic blue copper redox 

protein. This resembles AOB and nitrite-oxidizing bacteria in which the MCO-encoded genes flank genes encoding cytochrome c proteins. 

Blue shading indicates gene cluster that may have arisen by incomplete gene duplication of the cluster shaded in green (loss of BCP gene), followed by modification of the 3dMCO gene (loss of nirK-specific domains) 

and acquisition of genes Nmar_1134 and Nmar_1135). 

Orange shading indicates gene cluster containing a unique combination of gene homologues implicated in nitrogen oxide processing. 

Gene cluster Nmar_1650-7 may encode inventory responsible for copper uptake and homeostatis. Nmar_1652 encodes a fusion protein of the CopC and CopD proteins, whose encoding genes are found in tandem 

immediately downstream of amo genes.

Supplemental Table 3: Gene prediction coordinates for stable RNAs and TFBs/TBPs 

Gene Name Start End Product Note 

STABLE RNAs 

Nmar_rR16S 896,240 897,711 

Nmar_rR23S 893,105 896,109 

Nmar_rR5S 262,500 262,619 

Nmar_SRP 95,541 95,248 

Nmar_RNaseP 569,667 569,388 

Nmar_sR1 100,802 100,856 

Nmar_sR2 537,407 537,466 

Nmar_sR3 256,057 256,113 

Nmar_sR4 1,131,193 1,131,136 

Nmar_sR5 471,299 471,354 

Nmar_sR6 1,376,278 1,376,344 

16S 

ribosomal 

RNA 

23S 

ribosomal 

RNA 

5S 

ribosomal 

RNA 

SRP 

RNA 

RNase P 

RNA 

C/D box 

guide 

sRNA 

C/D box 


sRNA 

C/D box 


sRNA 

C/D box 


sRNA 

C/D box 


sRNA 

C/D box 


sRNA 

Nmar_tR01 88,221 88,305 LeuCAG 

Nmar_tR02 171,008 171,081 LysCTT 

Nmar_tR03 177,670 177,744 ValTAC 

Small subunit ribosomal RNA 

Large subunit ribosomal RNA 

5S ribosomal RNA; detected 

by Rfam model 

Signal recognition particle 

RNA; detected by Rfam model 

Detected by Rfam model 

Predicted to modify tRNA 

tR12-LeuCAA at Cm36 

(anticodon) by snoscan; 

conserved region in C. 

symbiosum 

Predicted to modify 23S rRNA 

at Cm2014 by snoscan 


tR06-AlaGGC at Am60 by 

snoscan; conserved sRNA in 

C. symbiosum 


tR35-TyrGTA at Am83 by 

snoscan; conserved sRNA in 

C. symbiosum 

No predicted target, but 

conserved with sRNA in C. 

symbiosum 

No predicted target, but 

conserved with sRNA in C. 

symbiosum 

No introns; tRNAscan-SE 

v1.23 Score 53.62 


v1.23 Score 71.92 


v1.23 Score 74.7 

Nmar_tR04 188,725 188,796 AlaCGC 


v1.23 Score 70.62 

Nmar_tR05 210,730 210,813 SerTGA No introns; tRNAscan-SE

Nmar_tR06 224,909 224,982 AlaGGC 

Nmar_tR07 232,880 232,957 AspGTC 

Nmar_tR08 255,938 256,011 MetCAT 

Nmar_tR09 270,209 270,280 HisGTG 

Nmar_tR10 281,501 281,585 LeuGAG 

Nmar_tR11 366,393 366,493 SerGGA 

Nmar_tR12 454,316 454,416 LeuCAA 

Nmar_tR13 504,317 504,391 ArgGCG 

Nmar_tR14 537,291 537,367 ProTGG 

Nmar_tR15 647,814 647,895 SerCGA 

Nmar_tR16 1,029,515 1,029,611 ValCAC 

Nmar_tR17 1,079,755 1,079,829 ThrTGT 

Nmar_tR18 1,212,832 1,212,945 MetCAT 

Nmar_tR19 1,313,500 1,313,571 CysGCA 

Nmar_tR20 1,349,774 1,349,927 TrpCCA 

Nmar_tR21 1,364,392 1,364,466 IleGAT 

Nmar_tR22 1,366,079 1,366,152 ThrGGT 

Nmar_tR23 1,403,611 1,403,685 ArgTCT 

Nmar_tR24 1,430,303 1,430,377 iMetCAT 

Nmar_tR25 1,592,406 1,592,498 ArgCCT 

v1.23 Score 65.85 


v1.23 Score 62.77 


v1.23 Score 64.16 


v1.23 Score 60.27 


v1.23 Score 44.15 


v1.23 Score 59.05 

Intron at 366431-366444; 

tRNAscan-SE v1.23 Score 

59.06 

Intron at 454356-454371; 


50.5 


v1.23 Score 62.07 


v1.23 Score 74.43 


v1.23 Score 55.73 

Contains a non-canonical 

intron; tRNAscan-SE v1.23 

Score 70.29 


v1.23 Score 68.33 



Score 80.94 


v1.23 Score 25.47 

Two introns: one at 1349839- 

1349891 and other is noncanonical; 

tRNAscan-SE v1.23 

Score 67.18 


v1.23 Score 72.57 


v1.23 Score 69.19 


v1.23 Score 65.64 


v1.23 Score 66.66 



Score 55.09

Nmar_tR26 1,630,610 1,630,537 ProGGG 

Nmar_tR27 1,591,588 1,591,515 ValGAC 

Nmar_tR28 1,479,706 1,479,633 GlyTCC 

Nmar_tR29 1,365,987 1,365,864 LeuTAA 

Nmar_tR30 1,363,723 1,363,650 AlaTGC 

Nmar_tR31 1,317,848 1,317,775 ThrCGT 

Nmar_tR32 1,263,222 1,263,136 SerGCT 

Nmar_tR33 754,037 753,964 PheGAA 

Nmar_tR34 707,158 707,082 AsnGTT 

Nmar_tR35 470,128 470,002 TyrGTA 

Nmar_tR36 418,681 418,604 GluCTC 

Nmar_tR37 397,959 397,883 GlyGCC 

Nmar_tR38 321,890 321,800 GlnCTG 

Nmar_tR39 232,783 232,710 LysTTT 

Nmar_tR40 198,740 198,653 LeuTAG 

Nmar_tR41 162,816 162,742 ArgTCG 

Nmar_tR42 81,511 81,439 GlnTTG 

Nmar_tR43 59,676 59,603 GlyCCC 

Nmar_tR44 255,184 255,090 GluTTC 

TRANSCRIPTION FACTOR B 

Nmar_0013 12,800 13,720 TFB TFB5 

Nmar_0020 16,889 17,794 TFB TFB6 

Nmar_0519 464,679 464,978 TFB TFB7 


v1.23 Score 63.49 


v1.23 Score 60.87 


v1.23 Score 70.32 

Two introns: one at 1365923- 

1365909 and other is noncanonical; 

tRNAscan-SE v1.23 

Score 56.34 


v1.23 Score 73.24 


v1.23 Score 76.73 


v1.23 Score 70.89 


v1.23 Score 73.7 


v1.23 Score 64.81 

Intron at 470090-470041; 


57.47 


v1.23 Score 70.32 


v1.23 Score 70.33 

Intron at 321852-321835; 


52.48 


v1.23 Score 70.32 


v1.23 Score 70.33 


v1.23 Score 70.34 


v1.23 Score 70.35 


v1.23 Score 70.36 



Score 73.75

Nmar_0624 564,972 565,880 TFB TFB8 

Nmar_0979 856,416 857,327 TFB TFB1 

Nmar_0987 863,158 864,105 TFB TFB2 

Nmar_1340 1,227,903 1,228,823 TFB TFB3 

Nmar_1341 1,229,190 1,230,119 TFB TFB4 

TRANSCIPTION BINDING PROTEINS 

Nmar_0598 534,324 534,884 TBP TBP2 

Nmar_1519 1,382,425 1,382,988 TBP TBP1

Supplemental Table 4: Gene prediction coordinates shared between the N. maritimus and C. symbiosum genomes. 

N. maritimus 

ORF 

C. symbiosum 

ORF 

aa % 

indentity Start 

N. maritimus SCM1 

Length 

End 

(aa) 

Strand Start 

C. symbiosum A 

Length 

End 

(aa) 

N. maritimus Annotation of Top BLAST Hit 

Strand 

146 1,348 400 f 1,814,836 1,816,068 410 f cell division control protein 6 family protein 

Nmar_0002 CENSYa_1840 58 1,325 2,770 481 r 1,816,083 1,817,513 476 r DNA-directed DNA polymerase 

Nmar_0003 CENSYa_1841 53 2,881 3,387 168 f 1,817,646 1,818,176 176 f hypothetical protein 

Nmar_0005 CENSYa_1842 65 4,356 5,072 238 r 1,818,183 1,818,902 239 r precorrin-6y C5,15-methyltransferase (decarboxylating), CbiE subunit 

Nmar_0006 CENSYa_1843 58 5,075 5,566 163 r 1,818,899 1,819,387 162 r hypothetical protein 

Nmar_0007 CENSYa_1844 42 5,639 7,168 509 f 1,819,536 1,821,173 545 f peptidylprolyl isomerase 

Nmar_0008 CENSYa_1845 70 7,169 8,524 451 r 1,821,115 1,822,368 417 r FAD-dependent pyridine nucleotide-disulphide oxidoreductase 


Nmar_0010 CENSYa_1847 63 9,370 9,861 163 f 1,823,259 1,823,753 164 f heat shock protein HSP20 

Nmar_0011 CENSYa_1848 72 9,889 12,030 713 f 1,823,778 1,825,907 709 f AAA family ATPase, CDC48 subfamily 

Nmar_0013 CENSYa_1825 64 12,800 13,720 306 r 1,803,378 1,804,292 304 f transcription factor TFIIB cyclin-related 

Nmar_0018 CENSYa_1820 41 15,531 15,830 99 f 1,800,611 1,800,907 98 r hypothetical protein 


Nmar_0020 CENSYa_1876 88 16,889 17,794 301 f 1,849,320 1,850,231 303 f transcription factor TFIIB cyclin-related 

Nmar_0021 CENSYa_1877 63 17,795 19,249 484 r 1,850,234 1,851,682 482 r sodium/hydrogen exchanger 

Nmar_0022 CENSYa_1879 52 19,365 19,907 180 f 1,852,830 1,853,375 181 f conserved hypothetical protein 

Nmar_0023 CENSYa_1880 80 19,920 20,459 179 f 1,853,445 1,853,939 164 f adenylylsulfate kinase 

Nmar_0024 CENSYa_1881 49 20,451 21,266 271 r 1,853,945 1,854,727 260 r inositol monophosphatase 






Nmar_0037 CENSYa_1892 58 28,946 29,659 237 f 1,861,823 1,862,539 238 f radical SAM domain protein 

Nmar_0038 CENSYa_1893 75 29,723 30,280 185 f 1,862,589 1,863,158 189 f GTP cyclohydrolase I 


Nmar_0040 CENSYa_1895 62 30,578 31,252 224 r 1,863,524 1,864,195 223 r ExsB family protein 

Nmar_0041 CENSYa_1896 59 31,256 32,179 307 r 1,864,192 1,864,965 257 r homoserine kinase 

Nmar_0042 CENSYa_1897 64 32,179 32,937 252 r 1,865,115 1,865,855 246 r protein of unknown function ATP binding 


Nmar_0044 CENSYa_1899 61 34,156 34,950 264 f 1,866,962 1,867,729 255 f short-chain dehydrogenase/reductase SDR 



Nmar_0048 CENSYa_1902 41 37,114 38,277 387 r 1,869,343 1,870,470 375 r thiamine biosynthesis ATP pyrophosphatase-like protein 

Nmar_0049 CENSYa_1903 68 38,305 39,030 241 f 1,870,540 1,871,325 261 f adenylylsulfate reductase, thioredoxin dependent 

Nmar_0050 CENSYa_1904 68 39,032 40,174 380 f 1,871,322 1,872,461 379 f sulfate adenylyltransferase 


Nmar_0052 CENSYa_1906 71 40,855 41,511 218 r 1,873,004 1,873,837 277 r DNA-(apurinic or apyrimidinic site) lyase 

Nmar_0053 CENSYa_1907 59 41,561 42,715 384 f 1,873,844 1,875,040 398 f protein of unknown function DUF521 


Nmar_0055 CENSYa_1920 49 43,073 45,031 652 r 1,885,975 1,887,870 631 f protein of unknown function DUF814 


Nmar_0057 CENSYa_1918 61 45,627 46,217 196 f 1,884,678 1,885,274 198 r precorrin-6Y C5,15-methyltransferase (decarboxylating), CbiT subunit 

Nmar_0058 CENSYa_1917 73 46,253 46,975 240 f 1,883,935 1,884,660 241 r precorrin-2 C20-methyltransferase 

Nmar_0059 CENSYa_1916 68 46,968 47,738 256 f 1,883,031 1,883,933 300 r precorrin-4 C11-methyltransferase 

Nmar_0061 CENSYa_1915 71 48,580 49,230 216 f 1,882,322 1,882,966 214 r conserved hypothetical protein 

Nmar_0062 CENSYa_1914 62 49,278 50,438 386 r 1,881,108 1,882,265 385 f glycosyl transferase family 2 

Nmar_0063 CENSYa_1924 72 50,600 52,057 485 r 1,889,509 1,890,846 445 r glycyl-tRNA synthetase 

Nmar_0064 CENSYa_1925 64 52,088 52,927 279 r 1,890,979 1,891,821 280 r apurinic endonuclease Apn1 

Nmar_0065 CENSYa_1926 59 53,013 54,038 341 f 1,891,958 1,892,971 337 f DNA primase, large subunit 

Nmar_0066 CENSYa_1927 53 54,038 55,159 373 f 1,892,964 1,894,082 372 f DNA primase small subunit 


Nmar_0068 CENSYa_1929 41 58,063 58,605 180 f 1,897,039 1,897,578 179 f glutamine amidotransferase class-I 

Nmar_0069 CENSYa_1930 50 58,640 59,602 320 f 1,897,613 1,898,572 319 f tetratricopeptide TPR_2 repeat protein 


Nmar_0071 CENSYa_1934 60 60,101 60,514 137 f 1,899,102 1,899,503 133 f ribosomal protein S6e 

Nmar_0072 CENSYa_1935 74 60,546 61,808 420 f 1,899,621 1,900,799 392 f protein synthesis factor GTP-binding 


Nmar_0074 CENSYa_1937 53 62,205 62,654 149 f 1,901,192 1,901,644 150 f PEBP family protein 

Nmar_0075 CENSYa_1939 47 62,723 64,861 712 f 1,902,465 1,904,675 736 f dolichyl-diphosphooligosaccharide--protein glycotransferase 

Nmar_0077 CENSYa_1948 67 65,021 65,818 265 r 1,922,910 1,923,689 259 f RNA binding S1 domain protein 

Nmar_0078 CENSYa_1947 62 65,859 66,389 176 r 1,922,333 1,922,863 176 f cob(I)alamin adenosyltransferase 

Nmar_0079 CENSYa_1944 52 66,477 67,829 450 f 1,908,224 1,909,495 423 r cobyrinic acid a,c-diamide synthase 

Nmar_0080 CENSYa_1943 61 67,818 68,459 213 r 1,907,632 1,908,222 196 f precorrin-8X methylmutase CbiC/CobH 

Nmar_0081 CENSYa_1942 60 68,449 69,201 250 r 1,906,845 1,907,600 251 f cobalamin (vitamin B12) biosynthesis CbiX protein 

Nmar_0082 CENSYa_1941 58 69,198 70,250 350 r 1,905,805 1,906,848 347 f cobalamin (vitamin B12) biosynthesis CbiG protein 

Nmar_0083 CENSYa_1940 61 70,282 71,367 361 r 1,904,838 1,905,773 311 f cobalamin biosynthesis protein CbiD 

Nmar_0084 CENSYa_1949 35 71,458 72,327 289 f 1,923,956 1,924,705 249 f NAD-dependent epimerase/dehydratase 

Nmar_0085 CENSYa_1950 70 72,304 73,470 388 r 1,924,679 1,925,845 388 r methionine adenosyltransferase 

Nmar_0086 CENSYa_1951 80 73,486 73,731 81 r 1,925,860 1,926,180 106 r like-Sm ribonucleoprotein core 

Nmar_0087 CENSYa_1952 37 73,840 74,493 217 f 1,926,247 1,926,903 218 f RecA/RadA recombinase-like protein 

Nmar_0088 CENSYa_1953 66 74,592 77,333 913 f 1,927,046 1,929,784 912 f DEAD/DEAH box helicase domain protein 

Nmar_0089 CENSYa_1954 54 77,330 78,730 466 r 1,929,781 1,931,187 468 r nucleic acid binding OB-fold tRNA/helicase-type 

Nmar_0090 CENSYa_1955 66 78,944 79,882 312 f 1,931,295 1,932,236 313 f ABC transporter related 

Nmar_0091 CENSYa_1956 73 79,866 80,627 253 f 1,932,292 1,932,981 229 f ABC-2 type transporter 

Nmar_0092 CENSYa_0589 40 80,633 81,373 246 r 539,411 540,283 290 r UDP-glucose/GDP-mannose dehydrogenase 

Nmar_0093 CENSYa_1958 44 81,548 82,600 350 r 1,933,158 1,934,201 347 r protein of unknown function DUF354 


Nmar_0095 CENSYa_1960 46 83,839 84,402 187 f 1,935,423 1,935,986 187 f GrpE protein 

Nmar_0096 CENSYa_1961 79 84,405 86,315 636 f 1,935,989 1,937,959 656 f chaperone protein DnaK 

Nmar_0097 CENSYa_1962 65 86,365 87,450 361 f 1,937,993 1,939,048 351 f chaperone protein DnaJ 

Nmar_0099 CENSYa_1967 49 88,308 90,212 634 r 1,941,188 1,943,074 628 r putative aspartyl-tRNA(Asn) amidotransferase, B subunit 

Nmar_0100 CENSYa_1968 57 90,221 91,510 429 r 1,943,097 1,944,275 392 r glutamyl-tRNA(Gln) amidotransferase, subunit D

Nmar_0101 CENSYa_1969 89 91,543 93,711 722 r 1,944,442 1,946,628 728 r AAA family ATPase, CDC48 subfamily 

Nmar_0102 CENSYa_1970 79 93,816 94,550 244 f 1,946,846 1,947,577 243 f ribosomal protein L2 

Nmar_0105 CENSYa_0057 62 95,588 96,502 304 r 42,784 43,593 269 f PP-loop domain protein 

Nmar_0107 CENSYa_0056 72 97,656 98,852 398 r 41,429 42,655 408 f peptidase M50 

Nmar_0108 CENSYa_1015 52 98,896 99,207 103 f 1,074,559 1,074,888 109 f CutA1 divalent ion tolerance protein 

Nmar_0109 CENSYa_1016 50 99,204 100,253 349 r 1,074,878 1,075,909 343 r eRF1 domain 1 protein 


Nmar_0111 CENSYa_1018 71 100,838 101,290 150 f 1,076,501 1,076,920 139 f protein of unknown function UPF0047 


Nmar_0113 CENSYa_1014 75 101,499 101,951 150 r 1,074,077 1,074,523 148 f iron (metal) dependent repressor, DtxR family 

Nmar_0114 CENSYa_1013 71 101,953 102,720 255 r 1,073,307 1,074,074 255 f transcriptional regulator, TrmB 

Nmar_0115 CENSYa_1012 91 102,822 104,501 559 f 1,071,473 1,073,242 589 r radical SAM domain protein 

Nmar_0116 CENSYa_1002 26 104,553 105,578 341 f 1,057,826 1,058,809 327 f glycosyl transferase group 1 

Nmar_0120 CENSYa_0995 11 109,495 110,676 393 r 1,049,888 1,051,114 408 r glycosyl transferase group 1 

Nmar_0161 CENSYa_0549 45 149,412 150,347 935 f 480,454 481,398 315 r alcohol dehydrogenase, class IV 

Nmar_0167 CENSYa_0590 44 154,844 155,815 323 r 540,324 541,418 364 r transcriptional regulator, RpiR family 

Nmar_0168 CENSYa_1003 32 155,950 156,876 308 f 1,058,746 1,059,645 299 r NAD-dependent epimerase/dehydratase 

Nmar_0169 CENSYa_0591 66 156,890 157,927 345 f 541,417 542,457 346 f putative translation initiation factor, aIF-2BI family 


Nmar_0173 CENSYa_1972 60 160,619 161,266 215 r 1,948,340 1,949,260 306 r short-chain dehydrogenase/reductase SDR 

Nmar_0177 CENSYa_1974 66 162,861 164,036 391 f 1,949,468 1,950,619 383 f aminotransferase class I and II 

Nmar_0178 CENSYa_1975 58 164,030 164,431 133 r 1,950,616 1,950,939 107 r pyridoxamine 5'-phosphate oxidase-related FMN-binding 


Nmar_0180 CENSYa_1977 60 164,664 165,335 223 r 1,951,240 1,951,860 206 r NADP oxidoreductase coenzyme F420-dependent 

Nmar_0181 CENSYa_1978 59 165,387 166,010 207 f 1,951,880 1,952,566 228 f phosphoglycerate mutase 


Nmar_0183 CENSYa_1980 82 166,329 166,760 143 f 1,952,975 1,953,406 143 f cytochrome c oxidase subunit II 

Nmar_0184 CENSYa_1981 85 166,798 168,324 508 f 1,953,456 1,954,982 508 f cytochrome c oxidase subunit I 

Nmar_0185 CENSYa_1982 64 168,337 169,284 315 f 1,954,992 1,955,579 195 f blue (type 1) copper domain protein 




Nmar_0190 CENSYa_1377 46 172,976 173,530 184 r 1,405,238 1,405,786 182 f methyltransferase type 11 

Nmar_0191 CENSYa_1376 60 173,569 173,730 53 r 1,405,056 1,405,232 58 f hypothetical protein 

Nmar_0192 CENSYa_1375 64 173,772 174,686 304 r 1,404,088 1,405,023 311 f branched-chain amino acid aminotransferase 

Nmar_0193 CENSYa_1371 59 175,136 176,551 471 r 1,400,934 1,402,349 471 f aspartate/glutamate/uridylate kinase 

Nmar_0194 CENSYa_1370 43 176,568 177,590 340 r 1,399,888 1,400,904 338 f glycosyltransferase 28 domain 


Nmar_0196 CENSYa_0389 50 178,141 178,488 115 r 338,160 338,510 116 r hypothetical protein 

Nmar_0197 CENSYa_0461 36 178,619 178,984 365 f 385,999 386,391 131 f hypothetical protein 

Nmar_0198 CENSYa_0009 68 178,988 179,500 170 r 7,457 7,954 165 f hypothetical protein 


Nmar_0201 CENSYa_0018 71 180,299 181,180 293 r 12,522 13,436 304 r radical SAM domain protein 

Nmar_0205 CENSYa_0020 61 182,361 182,717 118 f 13,798 14,151 117 f ribonuclease H 

Nmar_0206 CENSYa_0021 74 182,783 184,879 698 f 14,200 16,305 701 f CoA-binding domain protein 

Nmar_0207 CENSYa_0022 80 184,953 186,479 508 f 16,378 17,901 507 f vinylacetyl-CoA Delta-isomerase 

Nmar_0208 CENSYa_0023 51 186,634 187,344 236 r 17,915 18,889 324 r peptidase S26B, signal peptidase 


Nmar_0210 CENSYa_0025 82 188,271 188,597 108 f 19,871 20,203 110 f thioredoxin 

Nmar_0211 CENSYa_0027 57 188,840 189,199 119 f 21,002 21,361 119 r hypothetical protein 


Nmar_0214 CENSYa_0031 60 190,720 192,114 464 f 21,858 23,306 482 f dihydropyrimidinase 


Nmar_0216 CENSYa_1923 50 192,965 193,582 205 r 1,888,900 1,889,499 199 f AMMECR1 domain protein 

Nmar_0217 CENSYa_0284 68 193,616 194,686 356 f 247,243 248,373 376 f radical SAM domain protein 

Nmar_0218 CENSYa_0034 53 194,923 195,597 224 r 25,253 25,930 225 r ribose 5-phosphate isomerase 


Nmar_0220 CENSYa_0036 85 196,558 196,725 55 r 26,520 26,687 55 r ribosomal protein L37e 

Nmar_0221 CENSYa_0037 83 196,736 196,972 78 r 26,696 26,932 78 r like-Sm ribonucleoprotein core 

Nmar_0222 CENSYa_0038 42 197,030 197,752 240 r 26,994 27,704 236 r creatininase 

Nmar_0223 CENSYa_0039 65 197,758 198,627 289 r 27,701 28,543 280 r formyl transferase domain protein 

Nmar_0224 CENSYa_0292 54 198,788 200,137 449 f 252,466 253,737 423 f phosphoglucosamine mutase 

Nmar_0225 CENSYa_0293 85 200,199 200,585 128 f 253,819 254,199 126 f ribosomal protein L7Ae/L30e/S12e/Gadd45 

Nmar_0226 CENSYa_0294 82 200,582 200,794 70 f 254,196 254,408 70 f ribosomal protein S28e 

Nmar_0227 CENSYa_0295 84 200,804 201,004 66 f 254,420 254,617 65 f ribosomal protein L24E 

Nmar_0228 CENSYa_0296 73 201,007 201,408 133 f 254,730 255,020 96 f nucleoside-diphosphate kinase 

Nmar_0229 CENSYa_0297 71 201,415 203,196 593 f 255,027 256,808 593 f translation initiation factor aIF-2 

Nmar_0230 CENSYa_1987 52 203,199 203,600 133 r 1,957,626 1,958,033 135 r thioredoxin 

Nmar_0231 CENSYa_1734 45 203,816 205,486 556 r 1,723,506 1,725,080 524 r Ig family protein 


Nmar_0234 CENSYa_0852 56 206,483 207,019 178 f 878,130 878,741 203 f SNARE associated Golgi protein 

Nmar_0236 CENSYa_1730 50 207,468 208,751 427 f 1,719,258 1,720,511 417 r histidyl-tRNA synthetase 

Nmar_0237 CENSYa_1729 57 208,732 209,397 221 r 1,718,596 1,719,261 221 f putative translation initiation factor eIF-6 

Nmar_0238 CENSYa_1728 74 209,508 209,999 163 f 1,717,976 1,718,470 164 r 4Fe-4S ferredoxin iron-sulfur binding domain protein 


Nmar_0240 CENSYa_1725 66 210,817 211,767 316 r 1,716,277 1,717,212 311 f Replication factor C 


Nmar_0242 CENSYa_1723 71 212,475 214,562 695 f 1,713,470 1,715,563 697 r MCM family protein 

Nmar_0243 CENSYa_1722 58 214,559 216,688 709 f 1,711,350 1,713,473 707 r DEAD/DEAH box helicase domain protein 

Nmar_0244 CENSYa_1721 64 216,685 217,677 330 f 1,710,360 1,711,349 329 r glycosyl transferase family 4 

Nmar_0245 CENSYa_1720 56 217,901 218,377 158 r 1,709,666 1,710,142 158 f acetyltransferase 


Nmar_0247 CENSYa_1718 62 218,647 219,582 311 r 1,708,460 1,709,395 311 f oxidoreductase domain protein 

Nmar_0248 CENSYa_1717 58 219,583 220,500 305 r 1,707,546 1,708,463 305 f branched-chain amino acid aminotransferase 

Nmar_0249 CENSYa_1630 15 220,541 221,920 459 r 1,633,373 1,635,529 718 r histidine kinase 


Nmar_0253 CENSYa_1714 74 223,939 224,784 281 r 1,705,909 1,706,721 270 f oxidoreductase FAD/NAD(P)-binding domain protein 

Nmar_0255 CENSYa_1711 84 225,638 225,925 95 f 1,705,085 1,705,375 96 r alba, DNA/RNA-binding protein 

Nmar_0256 CENSYa_1710 70 225,951 226,817 288 r 1,704,218 1,705,060 280 f putative methylisocitrate lyase 

Nmar_0257 CENSYa_1708 65 227,020 227,529 169 r 1,703,109 1,703,915 268 f transcriptional regulator, PadR-like family

Nmar_0258 CENSYa_1705 82 227,624 229,336 570 f 1,700,274 1,701,332 352 r succinate dehydrogenase or fumarate reductase, flavoprotein subunit 



Nmar_0261 CENSYa_1702 69 230,117 230,866 249 f 1,698,720 1,699,496 258 r succinate dehydrogenase and fumarate reductase iron-sulfur protein 


Nmar_0263 CENSYa_1700 53 231,236 232,693 485 f 1,696,813 1,698,324 503 r argininosuccinate lyase 


Nmar_0265 CENSYa_1657 49 233,806 234,882 358 f 1,661,549 1,662,619 356 f glutamyl-tRNA reductase 


Nmar_0267 CENSYa_1659 70 235,140 235,982 280 r 1,662,970 1,663,809 279 r oxidoreductase FAD/NAD(P)-binding domain protein 

Nmar_0268 CENSYa_1652 58 236,068 236,868 266 r 1,657,288 1,658,103 271 r Inositol-phosphate phosphatase 


Nmar_0270 CENSYa_1654 62 238,081 239,238 385 f 1,659,318 1,660,478 386 f AAA ATPase central domain protein 


Nmar_0272 CENSYa_1660 79 239,653 241,200 515 f 1,663,911 1,665,458 515 f carboxyl transferase 

Nmar_0273 CENSYa_1661 77 241,206 242,693 495 f 1,665,512 1,666,942 476 f carbamoyl-phosphate synthase L chain ATP-binding 

Nmar_0274 CENSYa_1662 56 242,700 243,212 170 f 1,666,944 1,667,453 169 f biotin/lipoyl attachment domain-containing protein 

Nmar_0275 CENSYa_1663 81 243,229 243,693 154 r 1,667,459 1,667,929 156 r alkyl hydroperoxide reductase/ Thiol specific antioxidant/ Mal allergen 

Nmar_0276 CENSYa_1664 77 243,850 244,182 110 f 1,668,048 1,668,419 123 f NADH-ubiquinone/plastoquinone oxidoreductase chain 3 

Nmar_0277 CENSYa_1665 94 244,223 244,747 174 f 1,668,474 1,668,956 160 f NADH-quinone oxidoreductase, B subunit 

Nmar_0278 CENSYa_1666 64 244,747 245,349 200 f 1,668,953 1,669,765 270 f NADH dehydrogenase (ubiquinone) 30 kDa subunit 

Nmar_0279 CENSYa_1667 81 245,352 246,488 378 f 1,669,768 1,670,907 379 f NADH dehydrogenase (quinone) 

Nmar_0280 CENSYa_1668 75 246,489 247,787 432 f 1,670,907 1,672,205 432 f NADH dehydrogenase (quinone) 

Nmar_0281 CENSYa_1669 87 247,787 248,284 165 f 1,672,202 1,672,702 166 f 4Fe-4S ferredoxin iron-sulfur binding domain protein 

Nmar_0282 CENSYa_1670 77 248,277 248,789 170 f 1,672,728 1,673,207 159 f NADH-ubiquinone/plastoquinone oxidoreductase chain 6 

Nmar_0283 CENSYa_1671 91 248,770 249,075 101 f 1,673,218 1,673,493 91 f NADH-ubiquinone oxidoreductase chain 4L 

Nmar_0284 CENSYa_1672 77 249,075 250,628 517 f 1,673,493 1,675,055 520 f proton-translocating NADH-quinone oxidoreductase, chain M 

Nmar_0285 CENSYa_1673 83 250,630 252,708 692 f 1,675,057 1,677,132 691 f proton-translocating NADH-quinone oxidoreductase, chain L 

Nmar_0286 CENSYa_1674 72 252,721 254,205 494 f 1,677,146 1,678,633 495 f proton-translocating NADH-quinone oxidoreductase, chain N 

Nmar_0287 CENSYa_1675 69 254,195 255,046 283 r 1,678,623 1,679,441 272 r Polyprenyl synthetase 

Nmar_0288 CENSYa_1684 60 255,156 255,896 246 f 1,684,842 1,685,531 229 f TENA/THI-4 domain protein 

Nmar_0289 CENSYa_1686 46 256,122 256,727 201 r 1,685,789 1,686,391 200 r orotate phosphoribosyltransferase 


Nmar_0291 CENSYa_1688 64 257,852 258,958 368 r 1,687,508 1,688,602 364 r peptidase M50 

Nmar_0292 CENSYa_1690 76 259,014 259,529 171 f 1,689,758 1,690,264 168 f ribosomal-protein-alanine acetyltransferase 

Nmar_0293 CENSYa_1691 70 259,567 259,785 72 f 1,690,295 1,690,513 72 f transcription regulator containing HTH domain-like protein 

Nmar_0294 CENSYa_1692 56 259,787 260,614 275 r 1,690,514 1,691,341 275 r protein of unknown function Met10 

Nmar_0295 CENSYa_1694 73 260,620 262,407 595 r 1,691,905 1,693,689 594 r ABC transporter related 

Nmar_0296 CENSYa_0283 56 262,794 264,260 488 r 245,772 247,238 488 r leucyl aminopeptidase 

Nmar_0299 CENSYa_0086 67 265,235 266,902 555 f 61,147 62,661 504 f ribulose-phosphate 3-epimerase 

Nmar_0300 CENSYa_0087 71 266,899 267,873 324 f 62,658 63,614 318 f transketolase central region 

Nmar_0301 CENSYa_0088 71 267,870 268,538 222 f 63,665 64,276 203 f putative transaldolase 

Nmar_0303 CENSYa_1633 63 268,906 269,928 340 r 1,637,318 1,638,322 334 f Mg transporter protein CorA family protein 




Nmar_0307 CENSYa_1620 67 271,956 272,225 89 r 1,622,173 1,622,520 115 f protein of unknown function UPF0147 

Nmar_0309 CENSYa_0195 20 272,612 273,388 258 f 177,352 178,161 269 f GCN5-related N-acetyltransferase 

Nmar_0310 CENSYa_1619 59 273,380 274,249 289 r 1,621,263 1,622,129 288 f 5-carboxymethyl-2-hydroxymuconate Delta-isomerase 

Nmar_0311 CENSYa_1618 57 274,255 275,964 569 r 1,619,568 1,621,256 562 f glutamyl-tRNA synthetase 

Nmar_0312 CENSYa_1617 61 275,986 276,966 326 r 1,618,606 1,619,571 321 f polyprenyl synthetase 

Nmar_0313 CENSYa_1616 57 276,953 277,603 216 r 1,617,977 1,618,609 210 f isopentenyl-diphosphate delta-isomerase, type 1 

Nmar_0314 CENSYa_1615 54 277,603 278,346 247 r 1,617,237 1,617,977 246 f aspartate/glutamate/uridylate kinase 

Nmar_0315 CENSYa_1614 44 278,371 279,312 313 r 1,616,279 1,617,208 309 f mevalonate kinase 

Nmar_0316 CENSYa_1612 58 279,341 279,964 207 r 1,614,826 1,615,452 208 f ribosomal protein S2 

Nmar_0317 CENSYa_1611 62 279,972 281,210 412 r 1,613,584 1,614,822 412 f phosphopyruvate hydratase 

Nmar_0318 CENSYa_1610 71 281,212 281,484 90 r 1,613,346 1,613,582 78 f RNA polymerase, N/8 Kd subunit 

Nmar_0319 CENSYa_1608 53 281,601 282,215 204 r 1,612,595 1,613,230 211 f translin 

Nmar_0320 CENSYa_1603 67 282,289 282,936 215 f 1,607,129 1,607,770 213 r carbohydrate kinase, YjeF related protein 

Nmar_0321 CENSYa_1602 62 282,933 283,541 202 f 1,606,521 1,607,132 203 r beta-lactamase domain protein 

Nmar_0322 CENSYa_1601 57 283,546 284,607 353 f 1,605,464 1,606,504 346 r nicotinate-nucleotide-dimethylbenzimidazole phosphoribosyltransferase 

Nmar_0323 CENSYa_1600 61 284,600 286,360 586 r 1,603,720 1,605,477 585 f glucosamine--fructose-6-phosphate aminotransferase, isomerizing 








Nmar_0336 CENSYa_1577 85 298,821 300,401 526 r 1,586,170 1,587,705 511 f radical SAM domain protein 

Nmar_0337 CENSYa_1576 54 300,474 301,250 258 r 1,584,905 1,585,708 267 f 1-(5-phosphoribosyl)-5-amino-4-imidazole- carboxylate (AIR) carboxylase 

Nmar_0338 CENSYa_1575 77 301,284 302,198 304 r 1,583,961 1,584,869 302 f lactate/malate dehydrogenase 

Nmar_0339 CENSYa_1574 46 302,289 303,509 406 f 1,582,613 1,583,830 405 r protein of unknown function DUF111 

Nmar_0340 CENSYa_1573 60 303,510 304,307 265 f 1,581,822 1,582,616 264 r ExsB family protein 

Nmar_0341 CENSYa_1572 56 304,304 305,470 388 f 1,580,668 1,581,792 374 r cysteine desulfurase 





Nmar_0346 CENSYa_1568 81 309,252 309,503 83 f 1,577,515 1,577,766 83 r RNA polymerase Rpb5 

Nmar_0347 CENSYa_1567 88 309,504 312,851 1115 f 1,574,167 1,577,514 1115 r RNA polymerase Rpb2 domain 6 

Nmar_0348 CENSYa_1566 82 312,851 316,642 1263 f 1,570,385 1,574,167 1260 r DNA-directed RNA polymerase subunit A' 


Nmar_0352 CENSYa_0447 47 319,896 320,219 107 f 374,252 374,557 101 r ribosomal protein L7Ae/L30e/S12e/Gadd45 

Nmar_0353 CENSYa_0446 74 320,271 320,735 154 f 373,714 374,178 154 r NusA family KH domain protein 

Nmar_0354 CENSYa_0445 88 320,739 321,176 145 f 373,274 373,711 145 r ribosomal protein S23 (S12) 

Nmar_0355 CENSYa_0444 82 321,179 321,778 199 f 372,672 373,271 199 r ribosomal protein S7 

Nmar_0357 CENSYa_0442 61 322,289 322,993 234 f 371,462 372,409 315 r protein of unknown function UPF0153 

Nmar_0359 CENSYa_0276 83 323,446 324,444 332 r 239,994 241,007 337 r dehydrogenase (flavoprotein)-like protein 

Nmar_0366 CENSYa_1913 58 330,106 330,759 217 r 1,880,399 1,881,082 227 f uracil-DNA glycosylase superfamily

Nmar_0367 CENSYa_1256 55 330,772 333,639 955 r 1,278,565 1,280,301 578 f leucyl-tRNA synthetase 

Nmar_0368 CENSYa_1254 62 333,646 336,315 889 r 1,274,761 1,277,445 894 f alanyl-tRNA synthetase 



Nmar_0374 CENSYa_1242 78 341,230 341,547 105 f 1,268,526 1,268,822 98 r ribosomal protein 60S 

Nmar_0381 CENSYa_1250 70 345,558 346,424 288 r 1,272,219 1,273,043 274 r ribosomal protein L10 


Nmar_0383 CENSYa_0345 79 347,271 347,708 145 f 290,622 291,101 159 r transcriptional regulator, AsnC family 

Nmar_0385 CENSYa_0344 79 348,548 349,027 159 r 290,143 290,625 160 f ribosomal protein L11 

Nmar_0386 CENSYa_0343 74 349,062 349,520 152 r 289,715 290,110 131 f NusG antitermination factor 

Nmar_0387 CENSYa_0341 58 349,736 350,512 258 f 288,664 289,440 258 r protein of unknown function DUF516 

Nmar_0388 CENSYa_0340 59 350,509 350,808 99 f 288,353 288,667 104 r methylated-DNA--protein-cysteine methyltransferase 

Nmar_0389 CENSYa_1261 68 350,810 351,205 131 r 1,283,501 1,283,893 130 f glyoxalase/bleomycin resistance protein/dioxygenase 

Nmar_0390 CENSYa_0339 73 351,250 351,705 151 r 287,904 288,356 150 f ribosomal protein L19e 

Nmar_0391 CENSYa_0338 68 351,689 352,093 134 r 287,516 287,920 134 f ribosomal protein L32e 

Nmar_0392 CENSYa_0337 70 352,349 353,848 499 f 285,725 287,344 539 r phosphoenolpyruvate carboxykinase (ATP) 

Nmar_0394 CENSYa_0336 85 355,951 356,571 206 f 285,044 285,664 206 r superoxide dismutase 

Nmar_0395 CENSYa_0332 51 356,654 357,091 145 f 282,430 282,951 173 r prefoldin, alpha subunit 

Nmar_0396 CENSYa_0331 54 357,095 358,636 513 f 281,085 282,353 422 r signal recognition particle-docking protein FtsY 

Nmar_0397 CENSYa_0330 60 358,633 359,547 304 f 280,183 281,088 301 r ornithine carbamoyltransferase 

Nmar_0398 CENSYa_0329 68 359,584 360,066 160 f 279,669 280,148 159 r ribosomal protein L18P/L5E 

Nmar_0399 CENSYa_0328 79 360,068 360,784 238 f 279,010 279,669 219 r ribosomal protein S5 

Nmar_0400 CENSYa_0327 60 360,791 361,258 155 f 278,538 279,005 155 r ribosomal protein L30 

Nmar_0401 CENSYa_0326 71 361,260 361,724 154 f 278,121 278,537 138 r ribosomal protein L15 

Nmar_0402 CENSYa_0325 85 361,717 363,147 476 f 276,684 278,120 478 r preprotein translocase, SecY subunit 

Nmar_0403 CENSYa_0324 59 363,134 363,712 192 f 276,119 276,655 178 r adenylate kinase 

Nmar_0404 CENSYa_0323 74 363,716 364,339 207 f 275,487 276,113 208 r membrane protein-like protein 

Nmar_0405 CENSYa_0322 64 364,336 364,896 186 f 274,930 275,487 185 r cytidylate kinase 

Nmar_0406 CENSYa_0321 69 364,893 365,891 332 f 273,932 274,933 333 r pseudouridylate synthase TruB domain protein 


Nmar_0408 CENSYa_0388 38 366,847 367,329 160 f 337,276 338,094 272 f secreted protein 

Nmar_0409 CENSYa_0316 60 367,332 368,645 437 r 269,701 271,068 455 f FAD dependent oxidoreductase 



Nmar_0412 CENSYa_0313 55 369,607 370,575 322 r 267,895 268,941 348 f glyoxylate reductase 

Nmar_0413 CENSYa_0311 64 370,822 372,732 636 f 265,794 267,722 642 r pyruvate flavodoxin/ferredoxin oxidoreductase domain protein 

Nmar_0414 CENSYa_0310 79 372,722 373,684 320 f 264,842 265,771 309 r pyruvate ferredoxin/flavodoxin oxidoreductase, beta subunit 




Nmar_0419 CENSYa_0306 56 375,731 376,510 259 r 262,391 263,173 260 f uncharacterised conserved protein UCP005026 

Nmar_0420 CENSYa_1484 77 376,584 377,852 422 f 1,485,551 1,486,840 429 f 3-isopropylmalate dehydratase 

Nmar_0421 CENSYa_1485 74 377,852 378,334 160 f 1,486,837 1,487,322 161 f 3-isopropylmalate dehydratase, small subunit 

Nmar_0422 CENSYa_1486 71 378,343 379,758 471 f 1,487,324 1,488,742 472 f 3-isopropylmalate dehydratase, large subunit 

Nmar_0423 CENSYa_1487 56 379,760 380,344 194 f 1,488,783 1,489,307 174 f 3-isopropylmalate dehydratase, small subunit 

Nmar_0424 CENSYa_1488 55 380,322 381,041 239 r 1,489,285 1,490,004 239 r nucleotidyl transferase 

Nmar_0425 CENSYa_1489 78 381,045 381,494 149 r 1,490,008 1,490,457 149 r ribosomal protein S9 



Nmar_0428 CENSYa_1544 67 382,339 383,004 221 r 1,553,362 1,554,000 212 f RNA polymerase insert 


Nmar_0430 CENSYa_1538 74 383,542 384,228 228 f 1,549,535 1,550,221 228 r Shwachman-Bodian-Diamond syndrome protein 

Nmar_0431 CENSYa_1537 70 384,228 384,908 226 f 1,548,858 1,549,535 225 r KH type 1 domain protein 

Nmar_0432 CENSYa_1536 76 384,908 385,642 244 f 1,548,127 1,548,858 243 r exosome complex exonuclease 1 

Nmar_0433 CENSYa_1535 72 385,645 386,463 272 f 1,547,315 1,548,127 270 r 3' exoribonuclease 


Nmar_0436 CENSYa_0007 53 386,930 387,319 129 f 4,974 5,492 172 r prefoldin, beta subunit 

Nmar_0437 CENSYa_0010 71 387,372 388,058 228 f 7,982 8,677 231 f ERCC4 domain protein 

Nmar_0439 CENSYa_0011 50 388,545 389,024 159 r 8,668 9,255 195 r nucleotide binding protein, PINc 

Nmar_0440 CENSYa_1369 70 389,014 389,850 278 r 1,398,983 1,399,816 277 r ATP-NAD/AcoX kinase 

Nmar_0441 CENSYa_1378 83 389,913 390,758 281 f 1,405,811 1,406,680 289 f rhodanese domain protein 



Nmar_0444 CENSYa_1125 73 392,111 394,177 688 f 1,155,560 1,156,894 444 r glycosyl transferase family 2 


Nmar_0446 CENSYa_1129 73 396,135 397,460 441 f 1,158,729 1,160,099 456 f sodium/hydrogen exchanger 



Nmar_0450 CENSYa_1136 58 398,850 399,449 199 f 1,164,797 1,165,360 187 f tRNA intron endonuclease 

Nmar_0451 CENSYa_1137 57 399,439 400,095 218 r 1,165,401 1,166,060 219 r DNA repair helicase 





Nmar_0462 CENSYa_1142 35 415,626 416,150 525 r 1,167,952 1,168,620 222 f HEAT repeat 






Nmar_0477 CENSYa_0565 35 429,251 430,828 525 f 494,502 499,202 1566 f NHL repeat containing protein 

Nmar_0479 CENSYa_0570 60 432,960 434,213 417 f 505,584 507,107 507 f phosphate ABC transporter, periplasmic phosphate-binding protein 

Nmar_0481 CENSYa_0576 69 434,638 435,615 325 f 512,356 513,321 321 f phosphate ABC transporter, inner membrane subunit PstC 

Nmar_0482 CENSYa_0577 63 435,612 436,544 310 f 513,354 514,232 292 f phosphate ABC transporter, inner membrane subunit PstA 

Nmar_0483 CENSYa_0578 70 436,541 437,374 277 f 514,235 515,056 273 f phosphate ABC transporter, ATPase subunit 

Nmar_0484 CENSYa_1367 66 437,374 438,033 219 f 1,397,483 1,398,139 218 r phosphate uptake regulator, PhoU 

Nmar_0489 CENSYa_1359 42 440,199 440,882 227 f 1,385,705 1,386,367 220 r major intrinsic protein 

Nmar_0490 CENSYa_1357 58 440,908 442,182 424 f 1,383,004 1,384,374 456 r glutamate-1-semialdehyde-2,1-aminomutase 

Nmar_0491 CENSYa_1356 60 442,179 443,114 311 f 1,382,072 1,383,007 311 r porphobilinogen deaminase 

Nmar_0492 CENSYa_1355 68 443,111 443,851 246 f 1,381,335 1,382,075 246 r uroporphyrin-III C-methyltransferase

Nmar_0493 CENSYa_1354 56 443,851 444,654 267 f 1,380,559 1,381,338 259 r uroporphyrinogen III synthase HEM4 

Nmar_0495 CENSYa_1353 89 444,970 446,370 466 f 1,379,081 1,380,493 470 r FeS assembly protein SufB 

Nmar_0496 CENSYa_1352 67 446,390 447,790 466 f 1,377,672 1,379,057 461 r FeS assembly protein SufD 

Nmar_0497 CENSYa_1350 71 447,804 449,048 414 f 1,376,114 1,377,358 414 r cysteine desulfurase, SufS subfamily 

Nmar_0498 CENSYa_1349 67 449,045 449,488 147 f 1,375,687 1,376,094 135 r SUF system FeS assembly protein, NifU family 


Nmar_0500 CENSYa_1347 70 449,827 451,053 408 r 1,374,119 1,375,339 406 f phosphoglycerate kinase 

Nmar_0501 CENSYa_1392 52 451,172 451,840 222 f 1,417,127 1,417,753 208 f phosphoesterase PA-phosphatase related 




Nmar_0506 CENSYa_1400 59 454,758 455,378 206 f 1,422,629 1,423,249 206 f SNARE associated Golgi protein 


Nmar_0509 CENSYa_1402 64 456,304 457,284 326 r 1,423,535 1,424,524 329 r porphobilinogen synthase 

Nmar_0510 CENSYa_1403 63 457,316 458,581 421 r 1,424,556 1,425,809 417 r glutamyl-tRNA reductase 

Nmar_0511 CENSYa_1404 38 458,578 459,231 217 r 1,425,806 1,426,459 217 r siroheme synthase 

Nmar_0512 CENSYa_1301 69 459,323 460,348 341 f 1,310,966 1,311,985 339 f putative transcriptional regulator, AsnC family 

Nmar_0514 CENSYa_1302 69 460,691 461,689 332 r 1,311,986 1,312,966 326 r aldo/keto reductase 

Nmar_0515 CENSYa_1416 65 461,765 462,442 225 r 1,436,202 1,436,873 223 f chlorite dismutase 

Nmar_0516 CENSYa_1414 82 462,514 463,284 256 r 1,434,079 1,434,846 255 f FeS assembly ATPase SufC 

Nmar_0517 CENSYa_1413 64 463,382 464,338 318 r 1,433,062 1,434,027 321 f zinc finger TFIIB-type domain protein 


Nmar_0519 CENSYa_1411 62 464,679 464,978 99 r 1,432,434 1,432,739 101 f signal recognition particle, subunit SRP19 (SRP19) 

Nmar_0520 CENSYa_1410 73 464,980 465,363 127 r 1,432,040 1,432,429 129 f ribosomal protein S8e 

Nmar_0521 CENSYa_1409 60 465,432 466,010 192 r 1,431,384 1,431,977 197 f chromosome segregation and condensation protein ScpB 

Nmar_0522 CENSYa_1408 58 466,065 466,742 225 r 1,430,712 1,431,380 222 f chromosome segregation and condensation protein ScpA 

Nmar_0523 CENSYa_1407 66 466,837 467,889 350 f 1,429,504 1,430,562 352 r alcohol dehydrogenase GroES domain protein 

Nmar_0526 CENSYa_0779 73 469,360 469,671 103 f 731,412 731,969 185 f translation initiation factor eIF-1A 

Nmar_0527 CENSYa_0497 77 469,717 469,908 63 f 419,650 419,841 63 f cold-shock DNA-binding domain protein 


Nmar_0529 CENSYa_0419 78 470,881 471,291 136 f 355,433 355,906 157 f translation initiation factor eIF-5A 

Nmar_0530 CENSYa_0420 51 471,402 472,091 229 f 355,998 356,804 268 f putative ATP binding protein 

Nmar_0531 CENSYa_0421 73 472,084 473,412 442 f 356,806 357,999 397 f GTP-binding signal recognition particle SRP54 G- domain 

Nmar_0532 CENSYa_0422 33 473,435 474,580 381 f 358,039 359,247 402 f pseudouridylate synthase-like protein 


Nmar_0534 CENSYa_0425 45 475,477 475,650 57 r 360,216 360,383 55 r ribosomal protein S27a 


Nmar_0536 CENSYa_0430 58 476,150 476,710 186 f 362,275 362,814 179 f protein of unknown function DUF309 

Nmar_0537 CENSYa_0431 61 476,694 477,980 428 r 362,804 364,063 419 r phosphoglycerate mutase 

Nmar_0538 CENSYa_0432 73 478,209 479,225 338 f 364,127 365,182 351 f galactose-1-phosphate uridylyltransferase 


Nmar_0540 CENSYa_0434 58 479,711 480,148 145 r 365,670 366,089 139 r GCN5-related N-acetyltransferase 

Nmar_0541 CENSYa_0435 59 480,145 480,861 238 r 366,104 366,751 215 r nucleotidyl transferase 

Nmar_0542 CENSYa_0221 40 480,908 482,206 432 r 199,804 201,120 438 f phosphomethylpyrimidine kinase 

Nmar_0543 CENSYa_0216 80 482,942 484,270 442 f 195,975 197,330 451 r thiamine biosynthesis protein ThiC 

Nmar_0544 CENSYa_0215 44 484,253 484,657 134 r 195,726 196,031 101 f NUDIX hydrolase 

Nmar_0545 CENSYa_0214 69 484,663 485,514 283 r 194,795 195,640 281 f prephenate dehydrogenase 

Nmar_0546 CENSYa_0213 52 485,511 486,881 456 r 193,434 194,783 449 f aminotransferase class I and II 

Nmar_0547 CENSYa_0212 69 486,886 487,983 365 r 192,343 193,434 363 f chorismate synthase 

Nmar_0548 CENSYa_0205 51 488,310 489,578 422 r 185,867 187,111 414 f 3-phosphoshikimate 1-carboxyvinyltransferase 

Nmar_0549 CENSYa_0204 58 489,568 490,422 284 r 185,020 185,865 281 f shikimate kinase 

Nmar_0550 CENSYa_0203 55 490,422 491,243 273 r 184,217 185,023 268 f shikimate 5-dehydrogenase 

Nmar_0551 CENSYa_0202 49 491,292 491,963 223 r 183,509 184,162 217 f 3-dehydroquinate dehydratase, type I 

Nmar_0552 CENSYa_0201 73 491,965 493,026 353 r 182,496 183,512 338 f 3-dehydroquinate synthase 

Nmar_0553 CENSYa_0200 72 493,019 493,801 260 r 181,742 182,473 243 f predicted phospho-2-dehydro-3-deoxyheptonate aldolase 

Nmar_0554 CENSYa_0199 73 493,936 495,054 372 f 180,425 181,531 368 r radical SAM domain protein 

Nmar_0555 CENSYa_0198 73 495,049 496,491 480 r 178,951 180,393 480 f UbiD family decarboxylase 

Nmar_0556 CENSYa_1358 30 496,619 499,099 826 f 1,384,458 1,385,690 410 f histidine kinase 




Nmar_0560 CENSYa_0191 67 501,721 502,185 154 r 170,706 171,167 153 f alkyl hydroperoxide reductase/ Thiol specific antioxidant/ Mal allergen 

Nmar_0561 CENSYa_0190 57 502,286 502,999 237 f 169,899 170,645 248 r major intrinsic protein 



Nmar_0572 CENSYa_1806 54 508,749 510,014 421 f 1,786,901 1,788,181 426 f conserved hypothetical protein, membrane 

Nmar_0573 CENSYa_1826 57 510,081 510,596 171 f 1,804,394 1,804,939 181 r ferritin Dps family protein 


Nmar_0576 CENSYa_1160 49 511,861 512,307 148 f 1,194,717 1,195,163 148 r pyridoxamine 5'-phosphate oxidase-related FMN-binding 


Nmar_0578 CENSYa_1157 64 513,299 514,888 529 r 1,192,113 1,193,690 525 f lysyl-tRNA synthetase 

Nmar_0579 CENSYa_1156 60 514,892 516,475 527 r 1,190,545 1,192,113 522 f histone acetyltransferase, ELP3 family 

Nmar_0581 CENSYa_1152 76 516,887 518,137 416 r 1,175,420 1,176,811 463 f sodium/hydrogen exchanger 

Nmar_0582 CENSYa_1151 63 518,212 519,234 340 r 1,174,372 1,175,376 334 f phosphoribosylformylglycinamidine cyclo-ligase 

Nmar_0584 CENSYa_0161 33 520,425 522,080 1,656 f 138,954 144,167 1,738 r hypothetical protein 


Nmar_0590 CENSYa_0472 59 525,903 526,793 296 f 395,221 396,105 294 r Na+/Ca+ antiporter, CaCA family 



Nmar_0594 CENSYa_1238 62 530,634 531,773 379 r 1,263,583 1,264,740 385 f 2-alkenal reductase 

Nmar_0595 CENSYa_1236 61 531,812 533,020 402 r 1,261,060 1,262,217 385 f glycosyl transferase family 2 

Nmar_0596 CENSYa_1235 51 533,024 533,440 138 r 1,260,497 1,261,018 173 f cyclase/dehydrase 

Nmar_0598 CENSYa_1234 98 534,324 534,884 186 r 1,259,842 1,260,402 186 f 2-alkenal reductase 



Nmar_0601 CENSYa_1231 71 536,325 537,218 297 r 1,257,517 1,258,437 306 f methionine aminopeptidase, type II 




Nmar_0618 CENSYa_1313 53 558,117 560,579 820 r 1,321,685 1,324,150 821 f solute binding protein-like protein


Nmar_0620 CENSYa_1311 54 561,296 563,041 581 r 1,319,093 1,320,829 578 f membrane protein-like protein 

Nmar_0621 CENSYa_1209 55 563,248 563,577 109 r 1,234,676 1,235,170 164 r Sjogrens syndrome scleroderma autoantigen 1 

Nmar_0622 CENSYa_1212 92 563,679 563,987 102 f 1,237,032 1,237,340 102 f translation initiation factor SUI1 

Nmar_0623 CENSYa_1213 36 563,994 564,983 329 f 1,237,342 1,238,274 310 f abortive infection protein 

Nmar_0624 CENSYa_1876 69 564,972 565,880 909 r 1,849,320 1,850,231 304 f transcription initiation factor, TFIIB 


Nmar_0626 CENSYa_1217 79 566,639 567,598 319 r 1,240,091 1,241,017 308 r LPPG domain containing protein 


Nmar_0629 CENSYa_1314 49 569,705 570,328 207 r 1,324,125 1,324,871 248 r methyltransferase type 11 


Nmar_0631 CENSYa_1317 53 573,682 574,890 402 r 1,328,925 1,329,713 262 r geranylgeranyl reductase 


Nmar_0634 CENSYa_1323 63 575,917 576,678 253 r 1,334,887 1,335,645 252 r short-chain dehydrogenase/reductase SDR 

Nmar_0635 CENSYa_0230 52 576,732 577,610 292 r 206,615 207,463 282 f 2-hydroxy-3-oxopropionate reductase 



Nmar_0639 CENSYa_0223 60 579,023 579,685 220 f 201,595 202,233 212 r DSBA oxidoreductase 



Nmar_0642 CENSYa_0386 62 581,336 582,430 364 r 335,323 336,414 363 f DNA-directed DNA polymerase 


Nmar_0644 CENSYa_0384 72 583,285 583,518 77 f 334,474 334,950 158 r DNA-binding protein 

Nmar_0647 CENSYa_0383 49 584,407 585,120 237 r 333,787 334,494 235 f protein of unknown function DUF75 

Nmar_0648 CENSYa_0380 67 585,157 586,098 313 r 326,341 327,279 312 f deoxyhypusine synthase 

Nmar_0649 CENSYa_0379 81 586,202 587,296 364 f 325,073 326,290 405 r myo-inositol-1-phosphate synthase 

Nmar_0651 CENSYa_0378 50 588,210 591,734 1174 f 321,467 324,994 1175 r SMC domain protein 

Nmar_0652 CENSYa_0377 63 591,731 592,699 322 r 320,465 321,421 318 r oligopeptide/dipeptide ABC transporter, ATPase subunit 

Nmar_0653 CENSYa_0372 49 592,793 593,812 339 f 316,933 317,973 346 r conserved hypothetical protein 

Nmar_0655 CENSYa_0223 40 594,507 595,166 660 r 201,595 202,233 213 r protein-disulfide isomerase 


Nmar_0658 CENSYa_0369 68 597,843 599,285 480 r 313,409 314,839 476 f prolyl-tRNA synthetase 

Nmar_0659 CENSYa_0368 74 599,345 599,629 94 r 313,058 313,372 104 f conserved hypothetical protein 

Nmar_0666 CENSYa_0367 72 608,181 608,897 238 f 312,186 312,836 216 r phosphoserine phosphatase SerB 

Nmar_0667 CENSYa_0366 59 608,932 609,657 241 f 311,444 311,998 184 r F420-dependent oxidoreductase, putative 

Nmar_0668 CENSYa_0365 54 609,714 610,628 304 f 310,446 311,363 305 r PHP domain protein 

Nmar_0669 CENSYa_0364 68 610,696 610,932 78 f 310,177 310,398 73 r SirA family protein 

Nmar_0672 CENSYa_0363 69 612,652 613,656 334 r 309,204 310,175 323 f thioredoxin reductase 

Nmar_0674 CENSYa_1325 48 614,254 614,685 143 f 1,336,267 1,336,713 148 r UspA domain protein 


Nmar_0678 CENSYa_0361 79 616,743 617,561 272 r 307,557 308,369 270 f thiazole biosynthesis enzyme 

Nmar_0679 CENSYa_0358 71 617,751 619,559 602 f 302,735 304,540 601 r sulfite reductase (ferredoxin) 

Nmar_0680 CENSYa_0359 63 619,556 620,353 265 f 304,731 305,525 264 r rhodanese domain protein 

Nmar_0681 CENSYa_0357 66 620,354 621,481 375 r 301,484 302,593 369 f tetratricopeptide TPR_2 repeat protein 

Nmar_0682 CENSYa_0352 58 621,596 623,224 542 f 296,630 298,036 468 r collagen triple helix repeat 

Nmar_0684 CENSYa_0349 74 624,050 625,159 369 r 294,723 295,793 356 f small GTP-binding protein 

Nmar_0685 CENSYa_0348 51 625,481 626,176 231 f 293,881 294,684 267 r SPP-like hydrolase 

Nmar_0686 CENSYa_0347 58 626,178 628,064 628 f 292,037 293,860 607 r arginyl-tRNA synthetase 


Nmar_0690 CENSYa_0346 66 631,181 632,062 293 f 291,169 291,993 274 r tRNA methyltransferase complex GCD14 subunit 

Nmar_0691 CENSYa_0257 63 632,107 632,970 287 f 226,468 227,448 326 f carbohydrate kinase, YjeF related protein 

Nmar_0692 CENSYa_0258 69 632,975 634,327 450 f 227,451 228,560 369 f peptidase M20 

Nmar_0693 CENSYa_0260 57 634,324 634,740 138 r 228,794 229,225 143 r DoxX family protein 

Nmar_0694 CENSYa_0261 75 634,836 635,432 198 f 229,403 229,984 193 f proteasome endopeptidase complex 

Nmar_0695 CENSYa_0262 62 635,442 635,918 158 r 229,981 230,454 157 r PUA domain containing protein 

Nmar_0696 CENSYa_0263 69 635,915 636,925 336 r 230,451 231,458 335 r homoserine dehydrogenase 

Nmar_0697 CENSYa_0265 36 637,018 637,425 135 r 232,064 232,441 125 r methionine-R-sulfoxide reductase 

Nmar_0698 CENSYa_0266 61 637,462 639,099 545 r 232,489 234,069 526 r thymidylate synthase complementing protein ThyX 



Nmar_0703 CENSYa_0270 71 642,950 643,972 340 r 236,246 237,268 340 r XPG I domain protein 

Nmar_0710 CENSYa_0075 41 648,858 649,091 234 f 52,786 53,025 80 r transcriptional regulator 



Nmar_0715 CENSYa_1304 61 652,125 652,526 133 f 1,314,397 1,314,801 134 f Mov34/MPN/PAD-1 family protein 



Nmar_0718 CENSYa_1307 63 653,242 654,609 455 r 1,315,480 1,316,775 431 r binding-protein-dependent transport systems inner membrane component 

Nmar_0719 CENSYa_1308 76 654,606 655,655 349 r 1,316,831 1,318,042 403 r binding-protein-dependent transport systems inner membrane component 



Nmar_0726 CENSYa_1802 72 659,913 661,637 574 f 1,782,471 1,784,195 574 f SNF2-related protein 


Nmar_0729 CENSYa_0336 55 662,927 663,550 624 f 285,044 285,664 207 r superoxide dismutase 





Nmar_0734 CENSYa_1809 37 666,166 667,611 481 r 1,793,565 1,794,941 458 r FAD linked oxidase domain protein 



Nmar_0737 CENSYa_1812 34 669,433 669,981 182 r 1,796,748 1,797,263 171 r heat shock protein DnaJ domain protein 

Nmar_0738 CENSYa_1813 67 669,992 670,435 147 r 1,797,267 1,797,725 152 r transcriptional regulator, AsnC family 

Nmar_0739 CENSYa_1814 53 670,559 670,786 75 f 1,797,797 1,798,018 73 f TPR repeat-containing protein 

Nmar_0740 CENSYa_1815 42 670,859 671,335 158 f 1,798,058 1,798,531 157 f TPR repeat-containing protein 


Nmar_0742 CENSYa_1817 77 671,572 672,669 365 f 1,798,830 1,799,927 365 f radical SAM domain protein 



Nmar_0747 CENSYa_0916 40 674,573 675,526 317 f 956,842 957,774 310 r hypothetical protein

Nmar_0748 CENSYa_0915 84 675,576 677,768 730 f 954,593 956,785 730 r translation elongation factor aEF-2 

Nmar_0755 CENSYa_0448 43 681,293 681,910 205 r 374,609 375,172 187 r O-methyltransferase family 3 



Nmar_0764 CENSYa_0909 67 685,965 687,782 605 r 950,069 951,898 609 f CBS domain containing protein 


Nmar_0770 CENSYa_0907 70 690,338 691,429 363 r 948,465 949,556 363 f IMP biosynthesis enzyme PurP domain protein 

Nmar_0771 CENSYa_0902 75 691,728 692,285 185 f 945,138 945,779 213 r 3-octaprenyl-4-hydroxybenzoate carboxy-lyase 


Nmar_0773 CENSYa_0900 80 692,718 692,924 68 r 944,482 944,673 63 f sec-independent translocation protein mttA/Hcf106 

Nmar_0774 CENSYa_0899 65 692,953 693,744 263 r 943,615 944,409 264 f sec-independent protein translocase, TatC subunit 


Nmar_0776 CENSYa_2060 53 688,854 689,246 393 f 2,030,326 2,034,024 1,233 r hypothetical protein 

Nmar_0777 CENSYa_0353 61 695,077 695,337 86 f 298,139 298,399 86 r phosphoribosylformylglycinamidine synthase, purS 

Nmar_0779 CENSYa_1912 66 695,828 696,514 228 f 1,879,631 1,880,305 224 r phosphoribosylformylglycinamidine synthase I 

Nmar_0780 CENSYa_1911 61 696,511 698,676 721 f 1,877,493 1,879,634 713 r phosphoribosylformylglycinamidine synthase II 

Nmar_0781 CENSYa_1910 78 698,669 700,138 489 f 1,876,052 1,877,455 467 r glutamine amidotransferase class-II 

Nmar_0782 CENSYa_0895 70 700,141 700,419 92 f 911,771 912,091 106 r protein of unknown function DUF427 

Nmar_0784 CENSYa_0887 59 700,874 701,698 274 f 906,283 907,104 273 r phosphoribosylaminoimidazolesuccinocarboxamide synthase 

Nmar_0785 CENSYa_0885 58 702,047 702,550 167 f 905,096 905,617 173 r transcriptional regulator, ArsR family 

Nmar_0787 CENSYa_0884 62 702,901 703,305 134 f 904,689 905,087 132 r pyridoxamine 5'-phosphate oxidase-related FMN-binding 




Nmar_0795 CENSYa_0872 79 708,081 708,473 130 r 888,844 889,236 130 f ribosomal protein S8 



Nmar_0798 CENSYa_0869 60 709,185 709,901 238 r 887,432 888,148 238 f ribosomal protein S4E, central domain protein 

Nmar_0799 CENSYa_0868 48 709,901 710,407 168 r 886,929 887,432 167 f KOW domain protein 

Nmar_0800 CENSYa_0867 79 710,412 710,834 140 r 886,493 886,927 144 f ribosomal protein L14b/L23e 


Nmar_0802 CENSYa_0865 31 711,154 711,420 88 r 885,925 886,167 80 f ribonuclease P, Rpp29 





Nmar_0807 CENSYa_0860 73 713,265 713,531 88 r 883,802 884,044 80 f ribosomal protein L25/L23 

Nmar_0808 CENSYa_0859 61 713,528 714,343 271 r 882,972 883,787 271 f ribosomal protein L4/L1e 



Nmar_0811 CENSYa_0991 73 716,385 717,122 245 r 1,045,770 1,046,507 245 f proteasome endopeptidase complex 



Nmar_0815 CENSYa_0033 45 718,761 720,380 1,620 r 24,575 25,207 211 f coppper binding protein, plasocyanin/azurin family 

Nmar_0816 CENSYa_0986 53 720,467 721,117 216 r 1,043,156 1,043,785 209 f SNF7 

Nmar_0817 CENSYa_0985 55 721,209 722,318 369 r 1,041,950 1,043,062 370 f redoxin domain protein 

Nmar_0818 CENSYa_0984 68 722,323 723,057 244 r 1,041,214 1,041,945 243 f cytochrome c biogenesis protein transmembrane region 

Nmar_0819 CENSYa_0982 50 723,259 724,383 374 r 1,039,061 1,040,083 340 f 8-amino-7-oxononanoate synthase 


Nmar_0821 CENSYa_0981 66 725,326 726,306 326 r 1,038,016 1,038,972 318 f biotin synthase 

Nmar_0822 CENSYa_0980 57 726,380 727,660 426 r 1,036,637 1,037,956 439 f aminotransferase class-III 

Nmar_0823 CENSYa_0979 43 728,011 728,697 228 f 1,035,647 1,036,423 258 r dethiobiotin synthase 


Nmar_0828 CENSYa_0977 76 730,969 732,363 464 r 1,033,737 1,035,131 464 f hydroxymethylglutaryl-CoA synthase 

Nmar_0829 CENSYa_0976 56 732,469 733,179 236 r 1,032,975 1,033,655 226 f DSBA oxidoreductase 

Nmar_0831 CENSYa_0972 64 734,100 735,152 350 f 1,030,389 1,031,432 347 r glyceraldehyde-3-phosphate dehydrogenase (NAD(P)(+)) (phosphorylating) 


Nmar_0833 CENSYa_0532 76 736,555 737,892 445 r 464,494 465,816 440 r threonine synthase 



Nmar_0837 CENSYa_0579 53 739,895 740,443 182 r 515,094 515,630 178 f resolvase, Holliday junction-type 



Nmar_0841 CENSYa_0587 52 742,126 743,241 371 r 537,320 538,423 367 r propanoyl-CoA C-acyltransferase 

Nmar_0842 CENSYa_0588 55 743,238 744,203 321 r 538,420 539,382 320 r luciferase family protein 

Nmar_0843 CENSYa_0853 52 744,462 745,130 222 r 878,773 879,432 219 f nucleotidyl transferase 


Nmar_0846 CENSYa_0847 53 745,994 747,136 380 r 847,322 848,458 378 f phosphoribosylaminoimidazole carboxylase, ATPase subunit 

Nmar_0847 CENSYa_1141 61 747,236 747,811 191 f 1,167,335 1,167,880 181 r phosphoribosylaminoimidazole carboxylase, catalytic subunit 

Nmar_0848 CENSYa_0843 72 747,812 749,020 402 r 830,743 831,909 388 f threonine dehydratase 


Nmar_0850 CENSYa_0838 57 750,195 751,031 278 f 815,903 816,799 298 r adenylate/guanylate cyclase 


Nmar_0853 CENSYa_0834 70 752,141 752,551 136 r 814,348 814,761 137 f glyoxalase/bleomycin resistance protein/dioxygenase 

Nmar_0854 CENSYa_0833 68 752,586 752,804 72 r 814,140 814,295 51 f transcriptional regulator, AsnC family 


Nmar_0858 CENSYa_0231 56 754,081 754,617 178 r 207,460 208,083 207 f ATP--cobalamin adenosyltransferase 

Nmar_0859 CENSYa_0234 44 754,698 755,231 177 f 208,426 209,232 268 f metal dependent phosphohydrolase 

Nmar_0860 CENSYa_0234 50 755,231 755,455 225 f 208,426 209,232 269 f HD superfamily hydrolase 


Nmar_0863 CENSYa_0237 79 756,232 757,455 407 f 210,319 210,912 197 f LOR/SDH bifunctional protein conserved domain protein 


Nmar_0868 CENSYa_0238 63 761,773 763,176 467 r 210,958 212,352 464 r glutamyl-tRNA(Gln) amidotransferase, B subunit 

Nmar_0869 CENSYa_0239 62 763,173 764,618 481 r 212,349 213,788 479 r glutamyl-tRNA(Gln) amidotransferase, A subunit 


Nmar_0871 CENSYa_0241 70 764,895 766,205 436 r 214,051 215,619 522 r aspartyl-tRNA synthetase 

Nmar_0872 CENSYa_0242 69 766,788 767,813 341 f 215,698 216,723 341 f IMP biosynthesis enzyme PurP domain protein 

Nmar_0873 CENSYa_1262 75 767,816 768,613 265 r 1,283,890 1,284,627 245 r phosphonate ABC transporter, inner membrane subunit 

Nmar_0874 CENSYa_1263 61 768,591 769,421 276 r 1,284,662 1,285,462 266 r ABC transporter related 

Nmar_0875 CENSYa_1264 61 769,440 770,528 362 r 1,285,472 1,286,515 347 r phosphonate ABC transporter, periplasmic phosphonate-binding protein


Nmar_0877 CENSYa_1266 68 770,972 771,304 110 r 1,286,943 1,287,272 109 r putative transcriptional regulator 

Nmar_0878 CENSYa_1267 43 771,370 772,098 242 r 1,287,406 1,288,119 237 r conserved hypothetical protein 



Nmar_0881 CENSYa_0976 41 773,072 773,716 645 f 1,032,975 1,033,655 227 f protein-disulfide isomerase 

Nmar_0882 CENSYa_1290 73 773,772 775,052 426 f 1,305,181 1,306,416 411 f eRF1 domain 2 protein 

Nmar_0883 CENSYa_1289 61 775,045 775,449 134 r 1,304,689 1,305,114 141 f histidine triad (HIT) protein 

Nmar_0886 CENSYa_1285 79 776,452 778,128 558 f 1,301,101 1,302,717 538 r dihydroxy-acid dehydratase 

Nmar_0888 CENSYa_0208 53 778,828 780,228 466 r 188,290 189,969 559 r helicase c2 

Nmar_0906 CENSYa_0671 30 791,128 791,799 223 r 639,924 640,541 205 r conserved hypothetical protein 

Nmar_0911 CENSYa_1280 63 794,959 796,308 449 f 1,297,926 1,299,197 423 r Anthranilate synthase 

Nmar_0912 CENSYa_1279 66 796,305 796,901 198 f 1,297,339 1,297,929 196 r glutamine amidotransferase of anthranilate synthase 

Nmar_0913 CENSYa_1278 49 796,898 797,944 348 f 1,296,308 1,297,339 343 r anthranilate phosphoribosyltransferase 

Nmar_0914 CENSYa_1277 54 797,937 798,713 258 f 1,295,547 1,296,299 250 r indole-3-glycerol-phosphate synthase 

Nmar_0915 CENSYa_1276 70 798,710 799,897 395 f 1,294,375 1,295,490 371 r tryptophan synthase, beta subunit 

Nmar_0916 CENSYa_1275 50 799,884 800,690 268 f 1,293,558 1,294,337 259 r tryptophan synthase, alpha subunit 

Nmar_0918 CENSYa_1796 48 802,019 802,861 843 r 1,775,021 1,776,016 332 f copper binding protein, plastocyanin/azurin family 

Nmar_0919 CENSYa_1274 71 802,942 804,543 533 f 1,291,934 1,293,526 530 r CTP synthase 

Nmar_0921 CENSYa_1273 65 806,078 807,076 332 r 1,290,936 1,291,937 333 f NAD(+) kinase 

Nmar_0922 CENSYa_1272 46 807,118 807,987 289 r 1,290,026 1,290,889 287 f PfkB domain protein 

Nmar_0923 CENSYa_1382 88 808,250 808,537 95 f 1,408,487 1,408,774 95 r ribosomal protein S26E 

Nmar_0924 CENSYa_1381 59 808,540 809,112 190 r 1,407,918 1,408,490 190 f CDP-alcohol phosphatidyltransferase 

Nmar_0925 CENSYa_1380 65 809,190 810,065 291 r 1,407,021 1,407,881 286 f putative agmatinase 

Nmar_0926 CENSYa_0251 67 810,069 811,934 621 r 222,040 223,887 615 r threonyl-tRNA synthetase 


Nmar_0928 CENSYa_0253 66 812,404 813,468 354 f 224,381 225,442 353 f deoxyhypusine synthase 


Nmar_0930 CENSYa_0255 39 813,793 814,104 103 f 225,650 226,111 153 f small subunit ribosomal protein S25e 

Nmar_0932 CENSYa_0303 58 814,486 815,034 182 f 259,382 259,849 155 r RNA polymerase Rpb6 

Nmar_0933 CENSYa_0302 45 815,086 815,400 104 f 259,055 259,333 92 r alba, DNA/RNA-binding protein 


Nmar_0935 CENSYa_0300 44 815,847 816,866 339 f 257,603 258,754 383 r asparagine synthase 


Nmar_0937 CENSYa_1194 59 817,489 818,589 366 r 1,220,391 1,221,509 372 f aldo/keto reductase 

Nmar_0938 CENSYa_1185 61 818,620 819,510 296 r 1,216,088 1,216,966 292 f UbiA prenyltransferase 

Nmar_0939 CENSYa_1184 63 819,822 820,223 133 r 1,215,686 1,216,087 133 f conserved hypothetical protein 

Nmar_0942 CENSYa_1183 79 821,297 821,575 92 f 1,215,288 1,215,566 92 r PpiC-type peptidyl-prolyl cis-trans isomerase 

Nmar_0944 CENSYa_1182 65 821,861 823,369 502 r 1,213,605 1,215,281 558 f DEAD/DEAH box helicase domain protein 

Nmar_0947 CENSYa_1181 66 824,040 824,642 200 f 1,213,154 1,213,606 150 r CMP/dCMP deaminase zinc-binding 

Nmar_0948 CENSYa_1180 69 824,647 827,205 852 f 1,210,620 1,213,085 821 r DNA polymerase B region 


Nmar_0950 CENSYa_1177 56 827,535 828,191 218 f 1,209,091 1,209,744 217 f triosephosphate isomerase 

Nmar_0951 CENSYa_0473 67 828,219 830,882 887 f 396,302 398,956 884 f pyruvate, phosphate dikinase 

Nmar_0952 CENSYa_0474 67 830,925 831,482 185 f 398,970 399,527 185 f transcriptional regulator, XRE family 

Nmar_0953 CENSYa_0475 67 831,494 831,889 131 r 399,511 399,909 132 r glyoxalase/bleomycin resistance protein/dioxygenase 

Nmar_0954 CENSYa_0476 75 831,890 833,488 532 r 399,909 401,495 528 r methylmalonyl-CoA mutase, large subunit 

Nmar_0955 CENSYa_0477 59 833,495 834,412 305 r 401,500 402,423 307 r LAO/AO transport system ATPase 


Nmar_0957 CENSYa_0479 54 834,865 837,357 830 r 402,832 405,372 846 r peptidase M1 membrane alanine aminopeptidase 

Nmar_0958 CENSYa_0482 80 837,459 837,881 140 f 408,103 408,525 140 f cobalamin B12-binding domain protein 

Nmar_0959 CENSYa_1385 81 837,893 838,891 332 r 1,410,242 1,411,240 332 r ketol-acid reductoisomerase 

Nmar_0960 CENSYa_1386 59 839,050 839,580 176 r 1,411,439 1,411,969 176 r cytidyltransferase-related domain 

Nmar_0961 CENSYa_1387 49 839,615 840,535 306 r 1,412,000 1,412,932 310 r elongation factor Tu domain 2 protein 


Nmar_0963 CENSYa_1389 69 841,334 842,989 551 r 1,413,645 1,415,291 548 r methionyl-tRNA synthetase 

Nmar_0964 CENSYa_1390 73 843,027 844,268 413 r 1,415,334 1,416,584 416 r adenosylhomocysteinase 

Nmar_0965 CENSYa_1391 60 844,342 844,644 100 f 1,416,705 1,417,040 111 f Rieske (2Fe-2S) domain protein 

Nmar_0966 CENSYa_1345 52 844,775 845,488 237 f 1,350,107 1,350,922 271 r NAD+ synthetase 

Nmar_0967 CENSYa_1344 53 845,485 846,873 462 f 1,348,743 1,350,110 455 r cysteinyl-tRNA synthetase 




Nmar_0976 CENSYa_0436 43 852,580 853,506 308 f 367,004 367,903 299 r periplasmic binding protein 

Nmar_0978 CENSYa_1058 42 855,102 855,944 280 f 1,103,679 1,104,167 162 r response regulator receiver protein 

Nmar_0979 CENSYa_1332 40 856,416 857,327 912 f 1,340,590 1,341,576 269 r transcription initiation factor, TFIIB 

Nmar_0980 CENSYa_1342 68 857,503 858,714 403 r 1,347,103 1,348,323 406 f GTPase of unknown function domain protein 


Nmar_0982 CENSYa_1340 65 859,116 859,526 136 r 1,346,538 1,346,723 61 f CoA-binding domain protein 


Nmar_0984 CENSYa_1337 41 860,165 861,172 335 f 1,344,730 1,345,701 323 r biotin--acetyl-CoA-carboxylase ligase 


Nmar_0986 CENSYa_1335 48 861,646 863,085 479 f 1,342,821 1,344,224 467 r TPR repeat-containing protein 

Nmar_0987 CENSYa_1332 77 863,158 864,105 315 f 1,340,590 1,341,576 328 r transcription factor TFIIB cyclin-related 



Nmar_0991 CENSYa_1329 73 865,566 866,630 354 f 1,338,538 1,339,539 333 r poly(R)-hydroxyalkanoic acid synthase, class III, PhaC subunit 


Nmar_0993 CENSYa_1327 59 867,093 867,518 141 r 1,337,629 1,338,054 141 f transcriptional regulator, AbrB family 

Nmar_0994 CENSYa_1326 52 867,555 868,400 281 r 1,336,790 1,337,575 261 f alpha/beta hydrolase fold 

Nmar_0995 CENSYa_1324 66 868,653 869,105 150 r 1,335,811 1,336,263 150 f peptide methionine sulfoxide reductase 


Nmar_0997 CENSYa_0933 40 869,663 870,511 282 r 986,055 986,867 270 f methyltransferase type 11 

Nmar_1001 CENSYa_0120 41 876,009 876,260 252 f 94,871 95,185 105 r transcriptional regulator 

Nmar_1002 CENSYa_0064 37 876,296 876,688 393 f 45,734 46,117 128 f hypothetical membrane protein 

Nmar_1007 CENSYa_0971 55 881,616 882,884 1,269 r 1,029,131 1,030,387 419 f hemolysin 

Nmar_1017 CENSYa_1168 61 898,088 899,395 435 f 1,202,736 1,204,040 434 f aminotransferase class-III 

Nmar_1019 CENSYa_1169 46 899,956 900,777 273 f 1,204,093 1,204,899 268 f tetratricopeptide TPR_2 repeat protein 


Nmar_1021 CENSYa_1171 53 901,554 901,919 121 f 1,205,677 1,206,054 125 f hypothetical protein

Nmar_1022 CENSYa_1172 31 901,921 902,376 151 f 1,206,056 1,206,418 120 f cupin 2 conserved barrel domain protein 

Nmar_1024 CENSYa_0418 50 902,804 903,475 223 f 354,691 355,434 247 r heat shock protein DnaJ domain protein 

Nmar_1025 CENSYa_0417 53 903,476 903,883 135 r 354,274 354,690 138 f histidine triad (HIT) protein 



Nmar_1028 CENSYa_0413 75 904,421 905,560 379 f 351,794 352,891 365 r 3-hydroxybutyryl-CoA dehydrogenase 

Nmar_1030 CENSYa_0412 63 905,845 906,354 169 r 351,295 351,801 168 f Rossmann fold nucleotide-binding protein 


Nmar_1032 CENSYa_0408 65 907,229 907,432 67 r 350,199 350,420 73 f RNA polymerase Rbp10 


Nmar_1034 CENSYa_0406 90 907,775 909,073 432 r 348,543 349,853 436 f translation elongation factor EF-1, subunit alpha 

Nmar_1035 CENSYa_0564 78 909,195 910,340 381 f 492,785 494,302 505 f protein of unknown function DUF100 


Nmar_1037 CENSYa_1021 70 910,852 912,618 588 f 1,077,557 1,079,290 577 r DNA ligase I, ATP-dependent Dnl1 


Nmar_1039 CENSYa_1037 66 913,430 913,966 178 f 1,088,903 1,089,436 177 f tRNA intron endonuclease 

Nmar_1040 CENSYa_1258 36 914,012 916,327 2,316 r 1,281,178 1,282,698 507 f hypothetical protein 



Nmar_1043 CENSYa_1197 59 917,089 918,855 588 f 1,223,211 1,224,965 584 f short-chain dehydrogenase/reductase SDR 

Nmar_1047 CENSYa_1734 43 921,429 924,140 2,712 r 1,723,506 1,725,080 525 r hypothetical protein 




Nmar_1053 CENSYa_1201 70 927,691 928,155 154 f 1,226,888 1,227,256 122 f iron (metal) dependent repressor, DtxR family 

Nmar_1054 CENSYa_1202 62 928,142 929,959 605 r 1,227,253 1,229,088 611 r PilT protein domain protein 

Nmar_1056 CENSYa_0598 47 931,785 932,375 196 f 546,285 546,935 216 f phage SPO1 DNA polymerase-related protein 

Nmar_1057 CENSYa_0599 57 932,353 932,916 187 f 546,932 547,480 182 f DNA-3-methyladenine glycosylase 

Nmar_1058 CENSYa_0600 66 932,913 933,545 210 r 547,482 548,114 210 r SNARE associated Golgi protein 


Nmar_1061 CENSYa_1841 30 935,438 936,661 1,224 f 1,817,646 1,818,176 177 f hypothetical protein 

Nmar_1064 CENSYa_0602 63 937,290 938,225 311 r 548,575 549,555 326 r oligopeptide/dipeptide ABC transporter, ATPase subunit 

Nmar_1066 CENSYa_0611 68 938,826 940,112 428 f 556,181 557,458 425 f aspartyl-tRNA synthetase 


Nmar_1069 CENSYa_0612 63 940,960 941,973 337 r 557,455 558,453 332 r isopropylmalate/isohomocitrate dehydrogenase 

Nmar_1070 CENSYa_0613 73 942,007 943,524 505 r 558,460 559,977 505 r isopropylmalate/citramalate/homocitrate synthase 

Nmar_1071 CENSYa_0614 77 943,521 944,006 161 r 559,974 560,462 162 r acetolactate synthase, small subunit 

Nmar_1072 CENSYa_0615 86 944,015 945,715 566 r 560,465 562,120 551 r acetolactate synthase, large subunit, biosynthetic type 

Nmar_1078 CENSYa_0624 72 982,395 983,390 331 r 573,573 574,379 268 r fructose-bisphosphate aldolase 

Nmar_1079 CENSYa_0629 55 983,428 984,456 342 r 577,481 578,560 359 r alcohol dehydrogenase zinc-binding domain protein 

Nmar_1080 CENSYa_0630 76 984,493 984,786 97 r 578,670 578,960 96 r vacuolar H+transporting two-sector ATPase F subunit 



Nmar_1084 CENSYa_0768 46 985,813 986,562 249 f 719,908 720,600 230 f metallophosphoesterase 

Nmar_1085 CENSYa_0770 71 986,566 989,808 1080 r 721,706 723,676 656 r carbamoyl-phosphate synthase, large subunit 

Nmar_1086 CENSYa_0771 62 989,810 990,964 384 r 723,835 724,959 374 r carbamoyl-phosphate synthase, small subunit 

Nmar_1087 CENSYa_1874 29 991,041 992,597 1,557 r 1,846,587 1,848,137 517 r secreted periplasmic Zn-dependent protease 

Nmar_1088 CENSYa_0823 69 992,659 993,852 397 r 806,555 807,748 397 f AAA ATPase central domain protein 


Nmar_1091 CENSYa_0821 36 995,066 996,226 386 f 804,862 806,019 385 r transcriptional regulator, TrmB 

Nmar_1092 CENSYa_0818 50 996,256 996,711 151 f 763,435 763,908 157 r endoribonuclease L-PSP 

Nmar_1094 CENSYa_0298 43 998,322 998,714 130 f 256,809 257,180 123 r pyridoxamine 5'-phosphate oxidase family protein 

Nmar_1097 CENSYa_0260 39 1,000,385 1,000,831 447 f 228,794 229,225 144 r hypothetical protein 

Nmar_1098 CENSYa_0815 68 1,000,832 1,001,974 380 r 756,993 757,994 333 f phosphoesterase DHHA1 

Nmar_1099 CENSYa_0802 77 1,002,328 1,003,491 387 f 747,976 749,373 465 r 2-methylcitrate synthase/citrate synthase II 

Nmar_1100 CENSYa_0801 73 1,003,495 1,003,701 68 r 747,776 747,979 67 f hypothetical protein 

Nmar_1102 CENSYa_0033 35 1,004,439 1,005,395 318 f 24,575 25,207 210 f blue (type 1) copper domain protein 

Nmar_1103 CENSYa_0543 59 1,005,416 1,007,002 528 r 471,136 472,731 531 r excinuclease ABC, C subunit 

Nmar_1104 CENSYa_0544 70 1,006,999 1,009,818 939 r 472,728 475,535 935 r excinuclease ABC, A subunit 

Nmar_1105 CENSYa_0545 75 1,009,805 1,011,757 650 r 475,537 477,486 649 r excinuclease ABC, B subunit 

Nmar_1106 CENSYa_0547 59 1,011,807 1,012,457 216 r 478,638 479,270 210 r pyridoxamine 5'-phosphate oxidase-related FMN-binding 

Nmar_1109 CENSYa_0548 87 1,013,628 1,014,734 368 r 479,274 480,446 390 r luciferase family protein 

Nmar_1110 CENSYa_0549 82 1,014,829 1,015,773 314 r 480,454 481,398 314 r iron-containing alcohol dehydrogenase 

Nmar_1111 CENSYa_0799 46 1,015,826 1,017,103 425 f 746,118 747,377 419 f hydroxypyruvate reductase 


Nmar_1113 CENSYa_0118 61 1,017,395 1,018,360 321 r 93,587 94,630 347 f conserved hypothetical protein 

Nmar_1114 CENSYa_0117 65 1,018,479 1,019,054 191 f 92,960 93,535 191 r phosphoribosylglycinamide formyltransferase 

Nmar_1116 CENSYa_0278 45 1,019,373 1,019,657 94 r 241,761 241,916 51 r antibiotic biosynthesis monooxygenase 

Nmar_1117 CENSYa_0115 65 1,019,750 1,020,613 287 f 91,675 92,523 282 r methylenetetrahydrofolate dehydrogenase (NADP(+)) 

Nmar_1118 CENSYa_0114 46 1,020,603 1,021,166 187 f 91,134 91,655 173 r 5-formyltetrahydrofolate cyclo-ligase 

Nmar_1119 CENSYa_0113 61 1,021,157 1,022,422 421 r 89,903 91,144 413 f phosphoribosylamine--glycine ligase 

Nmar_1120 CENSYa_0112 63 1,022,506 1,025,703 1065 f 86,634 89,813 1059 r isoleucyl-tRNA synthetase 

Nmar_1121 CENSYa_0485 74 1,025,889 1,026,476 195 f 411,900 412,481 193 r ferritin Dps family protein 


Nmar_1123 CENSYa_0110 73 1,026,949 1,028,055 368 f 85,041 86,138 365 r succinate--CoA ligase (ADP-forming) 

Nmar_1124 CENSYa_0109 75 1,028,052 1,028,969 305 f 84,130 85,044 304 r CoA-binding domain protein 

Nmar_1125 CENSYa_0108 76 1,029,001 1,029,168 55 f 83,933 84,100 55 r 50S ribosomal protein L40e 

Nmar_1129 CENSYa_1579 76 1,031,846 1,033,210 1,365 r 1,588,210 1,589,202 331 r hypothetical protein 

Nmar_1130 CENSYa_1581 66 1,033,565 1,034,743 392 r 1,592,406 1,593,530 374 r putative metal cation transporter 

Nmar_1131 CENSYa_1582 75 1,034,733 1,035,788 351 r 1,593,556 1,594,596 346 r multicopper oxidase type 3 

Nmar_1132 CENSYa_1201 59 1,035,937 1,036,419 483 r 1,226,888 1,227,256 123 f Mn-dependent transcriptional regulator 

Nmar_1139 CENSYa_1882 63 1,043,604 1,043,813 69 r 1,854,724 1,855,053 109 r hypothetical protein 

Nmar_1140 CENSYa_0223 39 1,044,115 1,044,795 681 r 201,595 202,233 213 r protein-disulfide isomerase 


Nmar_1143 CENSYa_0223 55 1,046,241 1,046,906 666 f 201,595 202,233 213 r protein-disulfide isomerase 

Nmar_1146 CENSYa_1238 54 1,048,465 1,049,589 1,125 r 1,263,583 1,264,740 386 f trypsin-like serine protease 



Nmar_1152 CENSYa_1266 45 1,053,652 1,053,921 270 f 1,286,943 1,287,272 110 r hypothetical protein 

Nmar_1161 CENSYa_0637 43 1,062,348 1,063,718 1,371 r 586,359 591,230 1,624 r hypothetical protein

Nmar_1165 CENSYa_0569 26 1,066,715 1,067,632 305 r 504,646 505,296 216 r MT-A70 family protein 

Nmar_1171 CENSYa_1291 40 1,071,457 1,071,837 126 f 1,306,457 1,306,843 128 f hypothetical protein 

Nmar_1174 CENSYa_0017 44 1,073,391 1,073,876 486 f 12,121 12,525 135 f hypothetical protein 

Nmar_1177 CENSYa_0484 64 1,075,014 1,076,786 590 f 409,898 411,667 589 f oligoendopeptidase, pepF/M3 family 


Nmar_1180 CENSYa_0488 80 1,078,768 1,079,319 183 r 413,884 414,450 188 r arginine decarboxylase, pyruvoyl-dependent 

Nmar_1181 CENSYa_0500 54 1,079,893 1,080,690 798 f 420,881 421,621 247 r protein-disulfide isomerase 

Nmar_1182 CENSYa_0491 69 1,080,792 1,082,114 440 f 415,609 416,913 434 f DEAD/DEAH box helicase domain protein 

Nmar_1183 CENSYa_0506 45 1,082,163 1,082,396 234 f 425,326 425,631 102 r transcriptional regulator 



Nmar_1187 CENSYa_0499 47 1,084,186 1,084,641 151 r 420,423 420,884 153 r hypothetical protein 




Nmar_1191 CENSYa_0504 60 1,086,413 1,088,440 675 r 422,726 424,756 676 r protein of unknown function DUF255 


Nmar_1193 CENSYa_0510 65 1,088,792 1,089,826 344 f 447,580 448,587 335 f Alcohol dehydrogenase GroES domain protein 



Nmar_1197 CENSYa_0075 40 1,092,124 1,092,354 231 r 52,786 53,025 80 r transcriptional regulator 

Nmar_1198 CENSYa_0515 51 1,092,421 1,093,617 398 r 450,723 451,910 395 r tRNA pseudouridine synthase D TruD 

Nmar_1199 CENSYa_0516 49 1,093,614 1,095,215 533 r 451,910 453,436 508 r conserved hypothetical protein 

Nmar_1200 CENSYa_0519 67 1,095,261 1,095,635 124 r 454,802 455,209 135 r aminoacyl-tRNA hydrolase 

Nmar_1201 CENSYa_0161 59 1,096,377 1,101,503 5,127 r 138,954 144,167 1,738 r hypothetical protein 

Nmar_1205 CENSYa_1796 37 1,103,274 1,104,188 915 f 1,775,021 1,776,016 332 f copper binding protein, plastocyanin/azurin family 

Nmar_1211 CENSYa_2031 31 1,110,705 1,111,259 184 r 1,995,184 1,995,735 183 f conserved hypothetical protein 

Nmar_1212 CENSYa_0956 44 1,111,485 1,112,732 415 f 1,015,051 1,016,304 417 r metallophosphoesterase 

Nmar_1213 CENSYa_0955 27 1,112,729 1,115,149 806 f 1,012,608 1,014,941 777 r SMC domain protein 

Nmar_1220 CENSYa_0520 51 1,121,044 1,122,180 378 f 455,208 456,329 373 f dehydrogenase (flavoprotein)-like protein 

Nmar_1221 CENSYa_0525 66 1,122,289 1,122,417 42 f 460,035 460,274 79 r protein of unknown function DUF1610 

Nmar_1222 CENSYa_0524 70 1,122,422 1,122,703 93 f 459,752 460,033 93 r translation elongation factor aEF-1 beta 

Nmar_1223 CENSYa_0523 78 1,122,715 1,124,880 721 f 457,571 459,745 724 r AAA family ATPase, CDC48 subfamily 


Nmar_1225 CENSYa_0521 74 1,125,183 1,126,052 289 r 456,459 457,274 271 f protoheme IX farnesyltransferase 


Nmar_1227 CENSYa_0531 48 1,127,264 1,127,656 130 f 464,099 464,497 132 f Mov34/MPN/PAD-1 family protein 

Nmar_1229 CENSYa_0970 80 1,128,090 1,128,347 85 f 1,028,713 1,028,970 85 r conserved hypothetical protein 


Nmar_1231 CENSYa_0968 83 1,128,591 1,130,630 679 r 1,026,431 1,028,470 679 f V-type H(+)-translocating pyrophosphatase 

Nmar_1232 CENSYa_0967 75 1,130,666 1,131,043 125 r 1,026,018 1,026,395 125 f hypothetical protein 

Nmar_1233 CENSYa_0966 39 1,131,195 1,131,659 154 r 1,025,433 1,025,888 151 f protein of unknown function DUF359 


Nmar_1235 CENSYa_0963 85 1,132,962 1,133,558 198 f 1,021,919 1,022,653 244 r DNA-directed RNA polymerase 

Nmar_1236 CENSYa_0962 83 1,133,560 1,133,748 62 f 1,021,729 1,021,917 62 r DNA-directed RNA polymerase subunit E, RpoE2 

Nmar_1237 CENSYa_0961 52 1,133,788 1,134,606 272 f 1,020,912 1,021,727 271 r nicotinate-nucleotide pyrophosphorylase 

Nmar_1239 CENSYa_0960 69 1,135,009 1,135,965 318 r 1,019,962 1,020,915 317 f quinolinate synthetase complex, A subunit 

Nmar_1240 CENSYa_0959 58 1,136,059 1,136,877 272 r 1,019,042 1,019,872 276 f aspartate dehydrogenase 


Nmar_1243 CENSYa_0957 16 1,138,810 1,140,342 510 f 1,016,301 1,017,860 519 r protein of unknown function DUF87 

Nmar_1250 CENSYa_0033 68 1,147,717 1,148,436 720 r 24,575 25,207 211 f copper binding protein, plastocyanin/azurin family 

Nmar_1251 CENSYa_0954 79 1,148,438 1,148,926 162 r 1,012,165 1,012,611 148 f PBS lyase HEAT domain protein repeat-containing protein 

Nmar_1252 CENSYa_0952 65 1,149,023 1,149,688 221 f 1,011,324 1,011,983 219 r alkyl hydroperoxide reductase/ Thiol specific antioxidant/ Mal allergen 

Nmar_1253 CENSYa_1795 52 1,149,923 1,150,696 774 r 1,774,086 1,774,961 292 f copper binding protein, plastocyanin/azurin family 

Nmar_1254 CENSYa_0949 69 1,150,749 1,151,213 154 r 999,655 1,000,113 152 f ribosomal protein L15e 

Nmar_1255 CENSYa_1580 33 1,151,421 1,152,080 660 f 1,589,264 1,592,332 1,023 r secreted periplasmic Zn-dependent protease 


Nmar_1258 CENSYa_0943 71 1,153,583 1,154,515 310 r 996,419 997,351 310 f D-isomer specific 2-hydroxyacid dehydrogenase NAD-binding 

Nmar_1260 CENSYa_0942 67 1,156,106 1,157,467 453 f 994,972 996,420 482 r fumarate lyase 

Nmar_1261 CENSYa_0941 63 1,157,471 1,158,739 422 f 993,703 994,968 421 r MiaB-like tRNA modifying enzyme 

Nmar_1262 CENSYa_0940 46 1,158,813 1,159,763 316 f 992,699 993,631 310 r tubulin/FtsZ GTPase 

Nmar_1263 CENSYa_0939 42 1,159,800 1,160,363 187 r 991,969 992,622 217 f ribosomal protein L31e 


Nmar_1265 CENSYa_0937 54 1,160,560 1,161,111 183 r 991,179 991,784 201 f NUDIX hydrolase 

Nmar_1266 CENSYa_0936 35 1,161,181 1,161,918 245 f 990,356 991,099 247 r 5 10-methylenetetrahydrofolate reductase-like protein 

Nmar_1267 CENSYa_0935 71 1,161,915 1,164,413 832 r 987,875 990,361 828 f methionine synthase 

Nmar_1268 CENSYa_0934 66 1,164,446 1,165,408 320 r 986,890 987,843 317 f homocysteine S-methyltransferase 

Nmar_1269 CENSYa_0932 59 1,165,474 1,166,622 382 f 984,780 985,925 381 r major facilitator superfamily MFS_1 

Nmar_1273 CENSYa_1796 50 1,168,392 1,169,255 287 r 1,775,021 1,776,016 331 f blue (type 1) copper domain protein 



Nmar_1282 CENSYa_0064 29 1,174,917 1,175,300 384 r 45,734 46,117 128 f hypothetical membrane protein 

Nmar_1283 CENSYa_0931 73 1,175,356 1,176,114 252 r 984,041 984,805 254 f binding-protein-dependent transport systems inner membrane component 

Nmar_1284 CENSYa_0930 71 1,176,120 1,176,884 254 r 983,286 984,044 252 f ABC transporter related 

Nmar_1285 CENSYa_0929 48 1,176,884 1,177,909 341 r 982,315 983,289 324 f aliphatic sulfonates family ABC transporter, periplsmic ligand-binding protein 

Nmar_1286 CENSYa_0928 69 1,178,045 1,179,244 399 f 981,022 982,206 394 r argininosuccinate synthase 

Nmar_1287 CENSYa_1455 67 1,179,241 1,179,408 55 f 1,467,880 1,468,047 55 f conserved hypothetical protein 

Nmar_1288 CENSYa_0927 67 1,179,408 1,180,265 285 f 980,160 981,005 281 r lysine biosynthesis enzyme LysX 

Nmar_1289 CENSYa_1203 85 1,180,294 1,181,340 348 f 1,229,148 1,230,194 348 f N-acetyl-gamma-glutamyl-phosphate reductase 

Nmar_1290 CENSYa_1204 69 1,181,343 1,182,146 267 f 1,230,197 1,230,997 266 f acetylglutamate kinase 

Nmar_1291 CENSYa_1205 65 1,182,139 1,183,320 393 f 1,231,005 1,232,156 383 f acetylornithine and succinylornithine aminotransferase 

Nmar_1292 CENSYa_1206 71 1,183,307 1,183,726 139 f 1,232,233 1,232,568 111 f transcriptional regulator, AsnC family 

Nmar_1293 CENSYa_1207 77 1,183,825 1,185,006 393 f 1,232,629 1,233,825 398 f pyruvate carboxyltransferase 

Nmar_1294 CENSYa_0123 72 1,185,028 1,185,195 55 f 99,306 99,473 55 f conserved hypothetical protein 

Nmar_1295 CENSYa_0124 76 1,185,192 1,186,034 280 f 99,470 100,318 282 f lysine biosynthesis enzyme LysX 

Nmar_1296 CENSYa_0125 74 1,186,038 1,187,165 375 f 100,362 101,444 360 f N-acetyl-ornithine/N-acetyl-lysine deacetylase 

Nmar_1297 CENSYa_0126 56 1,187,165 1,188,202 345 f 101,444 102,481 345 f diphthine synthase 

Nmar_1298 CENSYa_0127 58 1,188,243 1,188,863 206 f 102,510 103,118 202 f cytidyltransferase-related domain 


Nmar_1302 CENSYa_0129 71 1,191,966 1,193,108 380 r 103,800 104,867 355 r Toprim sub domain protein

Nmar_1303 CENSYa_0131 86 1,193,261 1,193,611 116 r 105,653 106,003 116 r iron-sulfur cluster assembly accessory protein 



Nmar_1307 CENSYa_0033 42 1,195,212 1,195,736 525 f 24,575 25,207 211 f copper binding protein, plastocyanin/azurin family 

Nmar_1308 CENSYa_0166 77 1,195,786 1,196,547 253 r 149,820 150,575 251 r enoyl-CoA hydratase/isomerase 

Nmar_1309 CENSYa_0167 75 1,196,584 1,198,701 705 r 150,613 152,679 688 r CoA-binding domain protein 

Nmar_1310 CENSYa_0168 59 1,198,744 1,199,427 227 r 152,821 153,453 210 r glutamine amidotransferase class-I 

Nmar_1312 CENSYa_0173 75 1,199,847 1,201,121 424 r 156,263 157,543 426 r Glu/Leu/Phe/Val dehydrogenase 

Nmar_1314 CENSYa_0175 73 1,201,690 1,202,412 240 f 157,814 158,530 238 f proteasome endopeptidase complex 

Nmar_1315 CENSYa_0176 69 1,202,461 1,203,153 230 f 158,584 159,264 226 f methyltransferase type 11 

Nmar_1316 CENSYa_0177 57 1,203,150 1,204,304 384 f 159,261 160,517 418 f major facilitator superfamily MFS_1 

Nmar_1317 CENSYa_1824 44 1,204,301 1,204,615 104 r 1,802,898 1,803,227 109 r nitrogen regulatory protein P-II 

Nmar_1318 CENSYa_0178 56 1,204,740 1,205,087 115 f 160,861 161,199 112 f sec-independent translocation protein mttA/Hcf106 

Nmar_1319 CENSYa_1368 56 1,205,088 1,205,864 258 r 1,398,175 1,398,969 264 r DNA methylase N-4/N-6 domain protein 

Nmar_1320 CENSYa_0180 57 1,206,022 1,208,469 815 f 161,579 164,041 820 f DEAD/DEAH box helicase domain protein 

Nmar_1321 CENSYa_1360 57 1,208,508 1,209,338 276 f 1,386,379 1,387,209 276 r short-chain dehydrogenase/reductase SDR 

Nmar_1322 CENSYa_0182 67 1,209,416 1,211,071 551 f 164,390 165,949 519 f 2-alkenal reductase 

Nmar_1328 CENSYa_0793 34 1,218,185 1,219,645 1,461 f 740,421 742,553 711 r hypothetical protein 

Nmar_1331 CENSYa_1176 46 1,221,245 1,222,054 269 f 1,208,048 1,208,812 254 r parB-like partition protein 

Nmar_1340 CENSYa_1332 54 1,227,903 1,228,823 921 r 1,340,590 1,341,576 329 r transcription initiation factor, TFIIB 

Nmar_1341 CENSYa_1825 40 1,229,190 1,230,119 930 f 1,803,378 1,804,292 305 f transcription initiation factor, TFIIB 

Nmar_1348 CENSYa_0779 56 1,234,431 1,234,745 315 f 731,412 731,969 186 f translation initiation factor 1 

Nmar_1349 CENSYa_0926 35 1,234,843 1,236,567 1,725 r 978,506 980,158 551 f adenylate cyclase 



Nmar_1354 CENSYa_1582 38 1,239,281 1,240,615 1,335 r 1,593,556 1,594,596 347 r multicopper oxidase 

Nmar_1360 CENSYa_1138 38 1,246,835 1,247,611 258 f 1,166,066 1,166,956 296 f protein of unknown function DUF541 



Nmar_1368 CENSYa_0186 44 1,254,350 1,255,168 272 f 167,673 168,464 263 f conserved hypothetical protein 


Nmar_1370 CENSYa_0574 12 1,255,511 1,256,839 442 r 510,093 511,292 399 f conserved hypothetical protein 


Nmar_1379 CENSYa_0471 64 1,264,147 1,265,178 343 f 394,117 395,085 322 r isocitrate dehydrogenase (NAD(+)) 

Nmar_1380 CENSYa_0470 48 1,265,179 1,265,715 178 r 393,630 394,127 165 f putative methylase 

Nmar_1381 CENSYa_0469 44 1,265,690 1,266,394 234 r 392,957 393,622 221 f ribosomal RNA adenine methylase transferase 

Nmar_1382 CENSYa_0468 59 1,266,391 1,266,957 188 r 392,352 392,927 191 f protein of unknown function DUF655 

Nmar_1383 CENSYa_0467 79 1,266,978 1,267,304 108 r 392,018 392,344 108 f RNA polymerase Rpb4 


Nmar_1386 CENSYa_0250 60 1,268,792 1,269,958 388 r 220,780 221,976 398 f DNA repair and recombination protein RadA 


Nmar_1388 CENSYa_0554 81 1,270,263 1,271,987 574 r 482,581 484,305 574 r radical SAM domain protein 

Nmar_1389 CENSYa_0555 60 1,272,039 1,272,851 270 r 484,468 485,247 259 r oxidoreductase FAD/NAD(P)-binding domain protein 

Nmar_1390 CENSYa_0556 65 1,272,835 1,273,746 303 r 485,261 486,106 281 r dihydroorotate dehydrogenase family protein 

Nmar_1391 CENSYa_0557 51 1,273,832 1,275,244 470 f 486,266 487,444 392 f pre-mRNA processing ribonucleoprotein, binding domain protein 

Nmar_1392 CENSYa_0558 58 1,275,231 1,275,905 224 f 487,518 488,123 201 f non-specific serine/threonine protein kinase 

Nmar_1393 CENSYa_0559 55 1,275,902 1,276,519 205 r 488,120 488,737 205 r ribonuclease HII 

Nmar_1394 CENSYa_0561 48 1,276,580 1,277,728 382 f 489,649 490,782 377 f tRNA (guanine-N(2)-)-methyltransferase 

Nmar_1395 CENSYa_0562 66 1,277,718 1,278,311 197 r 490,779 491,369 196 r ribosomal RNA methyltransferase RrmJ/FtsJ 




Nmar_1400 CENSYa_0792 57 1,284,188 1,284,700 170 f 739,797 740,309 170 r adenine phosphoribosyltransferase 

Nmar_1401 CENSYa_0791 68 1,284,700 1,285,491 263 f 739,022 739,744 240 r methylthioadenosine phosphorylase 






Nmar_1407 CENSYa_0784 86 1,287,346 1,288,458 370 r 735,490 736,410 306 f DNA topoisomerase (ATP-hydrolyzing) 

Nmar_1408 CENSYa_0783 63 1,288,445 1,290,325 626 r 734,298 735,191 297 f DNA topoisomerase VI, B subunit 

Nmar_1409 CENSYa_0781 61 1,290,312 1,290,884 190 r 732,767 733,351 194 f KH type 1 domain protein 

Nmar_1410 CENSYa_0780 39 1,290,881 1,291,660 259 r 732,017 732,691 224 f non-specific serine/threonine protein kinase 

Nmar_1411 CENSYa_0829 62 1,291,717 1,292,013 98 r 811,368 811,664 98 f protein of unknown function DUF424 

Nmar_1412 CENSYa_0828 84 1,292,013 1,292,435 140 r 810,946 811,368 140 f translation initiation factor IF2/IF5 




Nmar_1416 CENSYa_1470 75 1,294,697 1,295,488 263 f 1,476,506 1,477,294 262 r undecaprenyl diphosphate synthase 

Nmar_1417 CENSYa_1469 52 1,295,485 1,295,904 139 f 1,475,989 1,476,504 171 r NUDIX hydrolase 

Nmar_1418 CENSYa_1468 52 1,295,884 1,296,624 246 r 1,475,264 1,475,992 242 f orotidine 5'-phosphate decarboxylase 


Nmar_1420 CENSYa_1463 70 1,297,338 1,298,339 333 r 1,472,438 1,473,454 338 f Adenylosuccinate synthase 






Nmar_1427 CENSYa_1699 24 1,301,276 1,301,872 198 f 1,696,144 1,696,713 189 f Sua5/YciO/YrdC/YwlC family protein 

Nmar_1428 CENSYa_1384 43 1,301,869 1,302,390 173 f 1,409,614 1,410,144 176 r THUMP domain protein 

Nmar_1429 CENSYa_1383 39 1,302,382 1,303,155 257 r 1,408,801 1,409,625 274 f thiamineS protein 


Nmar_1431 CENSYa_0054 52 1,303,617 1,304,435 272 r 40,327 41,076 249 r tatD-related deoxyribonuclease 

Nmar_1432 CENSYa_0055 70 1,304,454 1,304,690 78 f 41,164 41,391 75 f transcription factor CBF/NF-Y/histone domain protein 




Nmar_1447 CENSYa_1078 53 1,317,902 1,319,200 432 f 1,119,536 1,120,801 421 r phosphoglucomutase/phosphomannomutase alpha/beta/alpha domain I 

Nmar_1448 CENSYa_1077 43 1,319,184 1,320,116 310 f 1,118,332 1,119,534 400 r thiamine-monophosphate kinase 

Nmar_1450 CENSYa_1076 93 1,320,746 1,321,072 108 f 1,117,961 1,118,287 108 r ribosomal protein S11



Nmar_1453 CENSYa_1073 73 1,321,778 1,323,226 482 r 1,115,891 1,117,270 459 f protein of unknown function UPF0027 

Nmar_1455 CENSYa_1069 66 1,323,582 1,324,943 453 f 1,113,242 1,114,597 451 r adenylosuccinate lyase 

Nmar_1456 CENSYa_1060 44 1,325,203 1,326,207 1,005 r 1,104,640 1,105,479 280 r secreted periplasmic Zn-dependent protease 

Nmar_1462 CENSYa_2060 37 1,329,980 1,331,845 1,866 r 2,030,326 2,034,024 1,233 r hypothetical protein 



Nmar_1466 CENSYa_1068 48 1,333,685 1,334,440 251 r 1,112,509 1,113,294 261 f asparagine synthase 

Nmar_1467 CENSYa_0688 29 1,334,547 1,335,326 259 f 649,353 651,266 637 f excalibur domain protein 

Nmar_1468 CENSYa_1064 62 1,335,364 1,335,639 91 f 1,110,183 1,110,506 107 r transcriptional coactivator/pterin dehydratase 


Nmar_1473 CENSYa_1631 23 1,340,205 1,341,287 360 r 1,635,661 1,636,638 325 r DNA-cytosine methyltransferase 

Nmar_1477 CENSYa_1062 66 1,345,206 1,347,521 771 r 1,106,918 1,109,239 773 r aminoacyl-tRNA synthetase class Ia 

Nmar_1478 CENSYa_0065 47 1,347,536 1,348,153 205 r 46,119 46,724 201 r SNO glutamine amidotransferase 

Nmar_1479 CENSYa_0066 76 1,348,150 1,349,118 322 r 46,721 47,689 322 r vitamin B6 biosynthesis protein 


Nmar_1481 CENSYa_1822 22 1,349,952 1,350,800 282 r 1,801,614 1,802,432 272 r putative signal transduction protein with CBS domains 

Nmar_1482 CENSYa_0071 70 1,350,892 1,353,156 754 r 49,155 51,419 754 r aconitate hydratase 



Nmar_1485 CENSYa_0075 96 1,354,520 1,354,759 79 r 52,786 53,025 79 r transcriptional regulator, AsnC family 


Nmar_1488 CENSYa_0077 54 1,355,702 1,357,345 547 r 53,729 54,940 403 r phenylalanyl-tRNA synthetase, beta subunit 

Nmar_1489 CENSYa_0079 61 1,357,336 1,358,724 462 r 55,338 56,711 457 r phenylalanyl-tRNA synthetase, alpha subunit 

Nmar_1490 CENSYa_0080 73 1,358,772 1,359,884 370 f 56,765 57,871 368 f tryptophanyl-tRNA synthetase 


Nmar_1494 CENSYa_0082 57 1,361,774 1,362,250 158 r 58,537 58,995 152 r cytidyltransferase-related domain 


Nmar_1496 CENSYa_0290 63 1,363,804 1,364,358 184 f 251,584 252,135 183 r alkyl hydroperoxide reductase/ Thiol specific antioxidant/ Mal allergen 

Nmar_1497 CENSYa_1117 34 1,364,467 1,365,354 295 r 1,147,995 1,149,071 358 f putative transcriptional regulator 


Nmar_1499 CENSYa_0403 58 1,366,322 1,367,164 280 f 346,746 347,606 286 r DNA adenine methylase 



Nmar_1502 CENSYa_0399 95 1,368,507 1,369,079 190 r 343,290 343,859 189 f ammonia monooxygenase/methane monooxygenase, subunit C 





Nmar_1508 CENSYa_0103 73 1,371,803 1,372,252 149 f 77,110 77,556 148 r ribosomal S13S15 domain protein 

Nmar_1509 CENSYa_0102 63 1,372,252 1,373,673 473 f 75,704 77,035 443 r phosphoesterase DHHA1 


Nmar_1511 CENSYa_0100 61 1,373,878 1,375,143 421 f 74,246 75,502 418 r seryl-tRNA synthetase 

Nmar_1512 CENSYa_0099 70 1,375,184 1,375,795 203 f 73,575 74,186 203 r ribosomal protein S3Ae 

Nmar_1513 CENSYa_0098 32 1,375,822 1,376,247 141 f 73,152 73,565 137 r protein of unknown function DUF54 

Nmar_1514 CENSYa_0097 71 1,376,438 1,377,895 485 f 71,565 73,016 483 r geranylgeranyl reductase 

Nmar_1515 CENSYa_0096 75 1,377,936 1,378,730 264 f 70,691 71,485 264 r ATPase associated with various cellular activities AAA_5 

Nmar_1516 CENSYa_0095 58 1,378,731 1,380,293 520 f 69,126 70,691 521 r von Willebrand factor type A 

Nmar_1517 CENSYa_0094 64 1,380,359 1,381,903 514 f 66,985 68,868 627 r pyridoxal-5'-phosphate-dependent protein beta subunit 

Nmar_1519 CENSYa_0092 87 1,382,425 1,382,988 187 f 65,855 66,415 186 r TATA-box binding family protein 


Nmar_1524 CENSYa_0075 83 1,385,420 1,385,659 240 r 52,786 53,025 80 r transcriptional regulator 

Nmar_1526 CENSYa_1777 54 1,386,804 1,388,972 722 r 1,760,307 1,763,201 964 r thrombospondin type 3 repeat 

Nmar_1527 CENSYa_1634 57 1,389,055 1,391,223 722 r 1,638,466 1,641,234 922 r thrombospondin type 3 repeat 

Nmar_1528 CENSYa_1634 45 1,391,235 1,393,613 2,379 r 1,638,446 1,641,234 930 r autotransporter adhesin 

Nmar_1530 CENSYa_1640 46 1,394,794 1,395,360 188 r 1,648,357 1,648,737 126 r non-canonical purine NTP pyrophosphatase, rdgB/HAM1 family 

Nmar_1531 CENSYa_1641 54 1,395,344 1,395,964 206 r 1,648,913 1,649,869 318 r Mn2+-dependent serine/threonine protein kinase 

Nmar_1533 CENSYa_1642 74 1,396,322 1,397,305 327 r 1,649,969 1,650,682 237 r putative metalloendopeptidase, glycoprotease family 

Nmar_1534 CENSYa_1644 66 1,397,307 1,398,503 398 r 1,650,955 1,652,142 395 r GTP-binding protein HSR1-related 

Nmar_1535 CENSYa_1645 41 1,398,567 1,399,115 182 f 1,652,305 1,652,766 153 f conserved hypothetical protein 

Nmar_1536 CENSYa_1646 55 1,399,135 1,400,658 507 f 1,652,780 1,654,306 508 f tRNA-guanine transglycosylase 

Nmar_1537 CENSYa_1647 95 1,400,699 1,401,001 100 r 1,654,313 1,654,615 100 r 4Fe-4S ferredoxin iron-sulfur binding domain protein 


Nmar_1539 CENSYa_1649 74 1,401,468 1,402,733 421 f 1,655,177 1,656,430 417 f beta-lactamase domain protein 

Nmar_1540 CENSYa_1651 47 1,402,714 1,403,577 287 f 1,656,651 1,657,328 225 f HhH-GPD family protein 



Nmar_1543 CENSYa_0002 66 1,404,590 1,406,185 531 r 769 2,397 542 f cytochrome b/b6 domain 

Nmar_1544 CENSYa_0001 72 1,406,169 1,406,774 201 r 42 650 202 f Rieske (2Fe-2S) domain protein 

Nmar_1545 CENSYa_2066 57 1,406,818 1,410,627 1269 r 2,041,909 2,045,052 1047 f peptidase S8 and S53 subtilisin kexin sedolisin 



Nmar_1548 CENSYa_2046 64 1,417,196 1,418,527 443 r 2,007,223 2,008,599 458 f UBA/THIF-type NAD/FAD binding protein 

Nmar_1549 CENSYa_2045 73 1,418,527 1,419,741 404 r 2,005,985 2,007,223 412 f threonine synthase 


Nmar_1551 CENSYa_2043 78 1,420,539 1,420,865 108 r 2,005,008 2,005,334 108 f phosphoribosyl-AMP cyclohydrolase 

Nmar_1552 CENSYa_2042 64 1,420,888 1,421,688 266 r 2,004,223 2,004,984 253 f imidazoleglycerol phosphate synthase, cyclase subunit 

Nmar_1553 CENSYa_2041 55 1,421,678 1,422,388 236 r 2,003,492 2,004,196 234 f 

1-(5-phosphoribosyl)-5-((5- phosphoribosylamino)methylideneamino)imidazole-4carboxamideisomerase 

Nmar_1554 CENSYa_2040 55 1,422,385 1,422,987 200 r 2,002,890 2,003,495 201 f imidazole glycerol phosphate synthase, glutamine amidotransferase subunit 

Nmar_1555 CENSYa_2039 63 1,422,987 1,423,574 195 r 2,002,309 2,002,890 193 f Imidazoleglycerol-phosphate dehydratase 

Nmar_1556 CENSYa_2038 46 1,423,603 1,424,523 306 r 2,001,344 2,002,309 321 f haloacid dehalogenase domain protein hydrolase 

Nmar_1557 CENSYa_2037 52 1,424,527 1,425,597 356 r 2,000,283 2,001,347 354 f aminotransferase class I and II 

Nmar_1558 CENSYa_2036 50 1,425,594 1,426,856 420 r 1,999,015 2,000,283 422 f histidinol dehydrogenase 

Nmar_1559 CENSYa_2035 82 1,426,856 1,427,833 325 r 1,998,044 1,999,018 324 f ATP phosphoribosyltransferase 

Nmar_1560 CENSYa_2026 61 1,427,870 1,429,126 418 r 1,990,738 1,991,985 415 f hydroxymethylglutaryl-CoA reductase, degradative 

Nmar_1561 CENSYa_2025 57 1,429,128 1,430,219 363 r 1,989,566 1,990,738 390 f glucose sorbosone dehydrogenase 

Nmar_1562 CENSYa_2023 44 1,430,379 1,431,731 450 r 1,987,870 1,989,396 508 f domain of unknown function DUF1743 

Nmar_1563 CENSYa_2018 72 1,431,790 1,432,785 331 f 1,983,977 1,984,972 331 r diphthamide biosynthesis protein

Nmar_1564 CENSYa_2017 44 1,432,780 1,433,379 199 r 1,983,345 1,983,965 206 f TPR repeat-containing protein 

Nmar_1565 CENSYa_1751 72 1,433,386 1,434,471 1,086 r 1,739,791 1,740,870 360 f Zn-dependent oxidoreductase 

Nmar_1566 CENSYa_2014 58 1,434,547 1,435,098 183 f 1,980,218 1,981,306 362 r RNA-binding protein (consists of S1 domain and a Zn-ribbon domain)-like protein 

Nmar_1567 CENSYa_2013 60 1,435,101 1,435,916 271 f 1,979,404 1,980,111 235 r prephenate dehydratase 


Nmar_1569 CENSYa_2008 78 1,436,416 1,437,846 476 r 1,975,735 1,977,159 474 f inosine-5'-monophosphate dehydrogenase 

Nmar_1570 CENSYa_2007 46 1,437,892 1,438,737 281 r 1,974,898 1,975,725 275 f dihydropteroate synthase 



Nmar_1573 CENSYa_2003 73 1,440,119 1,441,642 507 f 1,971,335 1,972,858 507 r GMP synthase, large subunit 



Nmar_1576 CENSYa_2002 58 1,442,282 1,443,151 289 r 1,970,486 1,971,334 282 f ROK family protein 

Nmar_1577 CENSYa_2001 78 1,443,189 1,444,811 540 r 1,968,870 1,970,474 534 f thermosome 

Nmar_1578 CENSYa_2000 58 1,444,856 1,446,571 571 r 1,967,039 1,968,868 609 f adenine deaminase 

Nmar_1579 CENSYa_1999 74 1,446,690 1,446,971 93 f 1,966,668 1,966,949 93 r ribosomal protein L44E 

Nmar_1580 CENSYa_1695 40 1,446,968 1,447,165 65 f 1,693,717 1,694,391 224 r ribosomal protein S27E 

Nmar_1581 CENSYa_1998 38 1,447,148 1,447,732 194 r 1,966,100 1,966,696 198 f 5-deoxyadenosylcobinamide phosphate nucleotidyltransferase 

Nmar_1582 CENSYa_1997 56 1,447,729 1,448,454 241 r 1,965,441 1,966,100 219 f cobalamin 5'-phosphate synthase 

Nmar_1583 CENSYa_1996 54 1,448,447 1,449,418 323 r 1,964,417 1,965,379 320 f cobalamin biosynthesis protein CobD 

Nmar_1584 CENSYa_1995 52 1,449,415 1,450,254 279 r 1,963,590 1,964,420 276 f cobyrinic acid ac-diamide synthase 

Nmar_1585 CENSYa_1994 42 1,450,251 1,451,330 359 r 1,962,436 1,963,593 385 f aminotransferase class I and II 

Nmar_1586 CENSYa_1993 80 1,451,414 1,452,490 358 f 1,961,358 1,962,437 359 r aspartate-semialdehyde dehydrogenase 

Nmar_1587 CENSYa_1992 72 1,452,654 1,453,028 124 f 1,960,834 1,961,292 152 r transcriptional regulator, AsnC family 

Nmar_1588 CENSYa_1991 75 1,453,031 1,453,960 309 f 1,959,891 1,960,823 310 r protoheme IX farnesyltransferase 


Nmar_1592 CENSYa_0091 58 1,456,838 1,457,221 127 f 65,453 65,809 118 r conserved hypothetical protein 


Nmar_1597 CENSYa_1735 67 1,459,613 1,460,944 443 r 1,725,195 1,726,466 423 r UbiD family decarboxylase 





Nmar_1602 CENSYa_1742 66 1,463,132 1,463,473 113 r 1,730,734 1,731,072 112 f ubiquitin-associated- domain-containing protein 

Nmar_1603 CENSYa_1741 62 1,463,470 1,463,943 157 r 1,730,258 1,730,734 158 f PUA domain containing protein 

Nmar_1604 CENSYa_1739 64 1,464,034 1,465,080 348 f 1,728,946 1,729,980 344 r phosphate uptake regulator, PhoU 

Nmar_1605 CENSYa_1738 65 1,465,136 1,466,251 371 f 1,727,846 1,728,928 360 r GTP1/OBG protein 


Nmar_1607 CENSYa_1760 87 1,466,801 1,467,316 171 f 1,746,840 1,747,355 171 f transcription factor TFIIE, alpha subunit 

Nmar_1610 CENSYa_1761 60 1,469,218 1,469,775 185 r 1,747,356 1,747,907 183 r hemerythrin HHE cation binding domain protein 

Nmar_1612 CENSYa_1759 42 1,470,160 1,470,816 218 r 1,746,161 1,746,814 217 f bifunctional deaminase-reductase domain protein 

Nmar_1613 CENSYa_1758 56 1,470,803 1,471,993 396 r 1,744,975 1,746,153 392 f amidohydrolase 

Nmar_1614 CENSYa_1757 42 1,471,987 1,472,742 251 r 1,744,421 1,744,978 185 f GTP cyclohydrolase IIa 

Nmar_1615 CENSYa_1756 67 1,472,739 1,473,155 138 r 1,743,846 1,744,256 136 f 6,7-dimethyl-8-ribityllumazine synthase 

Nmar_1616 CENSYa_1755 63 1,473,158 1,473,823 221 r 1,743,156 1,743,830 224 f 3,4-dihydroxy-2-butanone 4-phosphate synthase 

Nmar_1617 CENSYa_1754 60 1,473,902 1,474,495 197 r 1,742,512 1,743,099 195 f riboflavin synthase, alpha subunit 

Nmar_1618 CENSYa_1753 43 1,474,520 1,475,245 241 r 1,741,800 1,742,507 235 f TPR repeat-containing protein 

Nmar_1621 CENSYa_1752 54 1,476,524 1,477,378 284 r 1,740,877 1,741,743 288 f citryl-CoA lyase 

Nmar_1622 CENSYa_1751 83 1,477,386 1,478,462 358 r 1,739,791 1,740,870 359 f alcohol dehydrogenase zinc-binding domain protein 

Nmar_1623 CENSYa_1764 52 1,478,505 1,478,786 93 r 1,750,041 1,750,328 95 r DNA-binding TFAR19-related protein 

Nmar_1624 CENSYa_1765 79 1,478,794 1,479,246 150 r 1,750,336 1,750,770 144 r ribosomal protein S19e 

Nmar_1625 CENSYa_1768 52 1,479,729 1,480,397 222 r 1,751,289 1,751,957 222 r suppressor Mra1 family protein 


Nmar_1627 CENSYa_1770 75 1,481,570 1,484,224 884 r 1,753,072 1,755,717 881 r ribonucleoside-diphosphate reductase, adenosylcobalamin-dependent 

Nmar_1628 CENSYa_1772 71 1,484,428 1,486,599 723 r 1,756,597 1,758,057 486 r regulatory protein ArsR 


Nmar_1630 CENSYa_1779 67 1,486,969 1,487,355 128 r 1,763,411 1,763,782 123 r protein of unknown function DUF35 

Nmar_1631 CENSYa_1780 82 1,487,336 1,488,499 387 r 1,763,784 1,764,938 384 r propanoyl-CoA C-acyltransferase 

Nmar_1632 CENSYa_1782 54 1,488,548 1,489,006 152 f 1,765,747 1,766,373 208 f DNA topoisomerase type IA zn finger domain protein 

Nmar_1633 CENSYa_1783 54 1,488,990 1,489,796 268 r 1,766,363 1,767,172 269 r nitrilase/cyanide hydratase and apolipoprotein N-acyltransferase 

Nmar_1634 CENSYa_1784 70 1,489,962 1,490,138 58 r 1,767,203 1,767,379 58 r 4-oxalocrotonate tautomerase 

Nmar_1635 CENSYa_1785 79 1,490,172 1,490,930 252 r 1,767,418 1,768,185 255 r rhomboid family protein 



Nmar_1638 CENSYa_0049 30 1,494,591 1,496,462 1,872 r 35,479 36,987 503 r hypothetical protein 



Nmar_1641 CENSYa_0043 50 1,499,144 1,500,094 316 r 29,939 30,874 311 r von Willebrand factor type A 

Nmar_1642 CENSYa_0044 41 1,500,095 1,500,967 290 r 30,878 31,708 276 r conserved hypothetical protein 

Nmar_1643 CENSYa_0045 56 1,500,977 1,501,999 340 r 31,737 32,744 335 r ATPase associated with various cellular activities AAA_3 


Nmar_1645 CENSYa_1788 57 1,503,352 1,503,852 166 r 1,768,797 1,769,285 162 r transcriptional regulator, ArsR family 





Nmar_1650 CENSYa_1795 45 1,508,803 1,509,615 813 f 1,774,086 1,774,961 292 f copper binding protein, plastocyanin/azurin family 


Nmar_1652 CENSYa_1798 52 1,510,361 1,513,144 927 f 1,776,761 1,779,544 927 f copper resistance D domain protein 

Nmar_1653 CENSYa_1053 52 1,513,134 1,513,952 272 r 1,099,722 1,100,555 277 r ABC-3 protein 

Nmar_1654 CENSYa_1054 47 1,513,955 1,514,677 240 r 1,100,560 1,101,345 261 r ABC transporter related 

Nmar_1655 CENSYa_1055 26 1,514,671 1,515,591 306 r 1,101,351 1,103,033 560 r periplasmic solute binding protein 

Nmar_1656 CENSYa_1056 59 1,515,683 1,516,066 127 f 1,103,156 1,103,551 131 f putative transcriptional regulator, CopG family 

Nmar_1658 CENSYa_0500 42 1,516,880 1,517,671 263 f 420,881 421,621 246 r DSBA oxidoreductase 

Nmar_1659 CENSYa_0920 69 1,517,700 1,518,122 423 r 975,284 975,703 140 f universal stress protein 

Nmar_1662 CENSYa_1581 59 1,520,144 1,521,322 1,179 r 1,592,406 1,593,530 375 r divalent heavy-metal cation transporter 

Nmar_1663 CENSYa_1582 79 1,521,312 1,522,373 1,062 r 1,593,556 1,594,596 347 r multicopper oxidase 

Nmar_1664 CENSYa_1201 57 1,522,483 1,522,974 492 r 1,226,888 1,227,256 123 f Mn-dependent transcriptional regulator 


Nmar_1670 CENSYa_0500 52 1,529,002 1,529,724 723 f 420,881 421,621 247 r protein-disulfide isomerase

Nmar_1671 CENSYa_0516 39 1,530,045 1,531,619 1,575 f 451,910 453,436 509 r streptogramin lyase 

Nmar_1673 CENSYa_1238 53 1,532,270 1,533,415 1,146 r 1,263,583 1,264,740 386 f trypsin-like serine protease 



Nmar_1678 CENSYa_1796 40 1,537,367 1,538,377 1,011 r 1,775,021 1,776,016 332 f copper binding protein, plastocyanin/azurin family 



Nmar_1686 CENSYa_1440 73 1,546,135 1,547,064 309 r 1,454,784 1,455,713 309 r aspartate carbamoyltransferase 

Nmar_1687 CENSYa_1441 75 1,547,106 1,547,567 153 f 1,455,753 1,456,214 153 f aspartate carbamoyltransferase, regulatory subunit 

Nmar_1688 CENSYa_1442 69 1,547,603 1,547,911 102 r 1,456,242 1,456,541 99 r H+transporting two-sector ATPase C subunit 

Nmar_1689 CENSYa_1444 83 1,548,095 1,548,724 209 r 1,456,722 1,457,087 121 r V-type ATPase, D subunit 

Nmar_1690 CENSYa_1445 88 1,548,729 1,550,114 461 r 1,457,356 1,458,744 462 r sodium-transporting two-sector ATPase 

Nmar_1691 CENSYa_1446 86 1,550,111 1,551,889 592 r 1,458,741 1,460,519 592 r H+transporting two-sector ATPase alpha/beta subunit central region 

Nmar_1692 CENSYa_1447 50 1,551,891 1,552,487 198 r 1,460,521 1,461,117 198 r H+transporting two-sector ATPase E subunit 

Nmar_1693 CENSYa_1451 66 1,552,577 1,554,676 699 f 1,462,539 1,464,614 691 f V-type ATPase 116 kDa subunit 

Nmar_1697 CENSYa_1452 51 1,557,401 1,558,177 258 r 1,464,598 1,465,389 263 r rhodanese domain protein 

Nmar_1698 CENSYa_1453 81 1,558,251 1,559,816 521 r 1,465,453 1,467,018 521 r ammonium transporter 


Nmar_1700 CENSYa_1456 59 1,560,670 1,561,704 344 f 1,468,122 1,469,162 346 f proton-transporting two-sector ATPase 




Nmar_1705 CENSYa_1531 64 1,563,854 1,564,537 227 r 1,545,226 1,545,912 228 f uridylate kinase, putative 

Nmar_1706 CENSYa_1530 56 1,564,534 1,565,769 411 r 1,544,003 1,545,226 407 f metal dependent phosphohydrolase 

Nmar_1707 CENSYa_1529 40 1,565,766 1,566,347 193 r 1,543,420 1,543,980 186 f thymidylate kinase 

Nmar_1708 CENSYa_1510 52 1,566,629 1,567,009 126 f 1,522,043 1,522,426 127 r heat shock protein Hsp20 

Nmar_1709 CENSYa_1509 64 1,567,013 1,568,557 514 r 1,520,453 1,521,997 514 f ABC-1 domain protein 



Nmar_1713 CENSYa_1506 68 1,570,929 1,571,552 207 r 1,517,927 1,518,607 226 f ABC transporter related 


Nmar_1715 CENSYa_1504 56 1,572,933 1,573,835 300 f 1,515,644 1,516,588 314 r putative transcriptional regulator, AsnC family 

Nmar_1717 CENSYa_1503 53 1,574,377 1,575,357 326 r 1,514,679 1,515,656 325 f phosphate uptake regulator, PhoU 

Nmar_1718 CENSYa_1502 49 1,575,422 1,576,333 303 f 1,513,742 1,514,614 290 r GHMP kinase 

Nmar_1719 CENSYa_1501 52 1,576,344 1,577,108 254 f 1,512,977 1,513,735 252 r Ppotein of unknown function DUF137 

Nmar_1720 CENSYa_1500 57 1,577,101 1,577,940 279 f 1,512,166 1,512,954 262 r 3-methyl-2-oxobutanoate hydroxymethyltransferase 

Nmar_1721 CENSYa_1499 57 1,577,930 1,579,174 414 f 1,510,896 1,512,083 395 r phosphopantothenoylcysteine decarboxylase/phosphopantothenate--cysteine ligase 

Nmar_1722 CENSYa_1498 43 1,579,343 1,580,185 280 f 1,510,052 1,510,873 273 r methyltransferase type 11 

Nmar_1724 CENSYa_1497 63 1,580,685 1,581,749 354 r 1,508,989 1,510,050 353 f peptidase M24 

Nmar_1725 CENSYa_1496 74 1,581,822 1,582,835 337 f 1,507,907 1,508,917 336 r tRNA (5-methylaminomethyl-2-thiouridylate)-methyltransferase 


Nmar_1727 CENSYa_1545 79 1,583,179 1,585,119 646 r 1,554,047 1,555,984 645 r beta-lactamase domain protein 

Nmar_1728 CENSYa_1546 82 1,585,127 1,585,759 210 r 1,555,992 1,556,579 195 r proteasome endopeptidase complex 

Nmar_1729 CENSYa_1547 64 1,585,837 1,586,901 354 f 1,556,693 1,557,757 354 f 3-dehydroquinate synthase 

Nmar_1730 CENSYa_1548 45 1,586,903 1,587,649 248 r 1,557,762 1,558,547 261 r peptidylprolyl isomerase FKBP-type 

Nmar_1731 CENSYa_1549 70 1,587,692 1,588,837 381 r 1,558,576 1,559,706 376 r serine--pyruvate transaminase 

Nmar_1732 CENSYa_1550 41 1,588,879 1,589,373 164 r 1,559,764 1,560,099 111 r cytidyltransferase-related domain 

Nmar_1733 CENSYa_1552 42 1,589,437 1,590,453 338 f 1,560,320 1,561,339 339 f RNA-3'-phosphate cyclase 



Nmar_1736 CENSYa_1556 66 1,591,671 1,592,381 236 f 1,562,476 1,563,285 269 f cob(II)yrinic acid a,c-diamide reductase 

Nmar_1737 CENSYa_1558 60 1,592,549 1,593,073 174 f 1,563,477 1,564,001 174 f NADPH-dependent FMN reductase 



Nmar_1740 CENSYa_1438 75 1,594,446 1,594,922 158 f 1,453,532 1,454,011 159 r peptidylprolyl isomerase 



Nmar_1745 CENSYa_1434 75 1,597,262 1,598,155 297 f 1,450,743 1,451,636 297 r PfkB domain protein 

Nmar_1746 CENSYa_1433 66 1,598,195 1,598,542 115 f 1,450,340 1,450,642 100 r dUTPase 

Nmar_1747 CENSYa_1432 43 1,598,547 1,599,014 155 r 1,449,921 1,450,346 141 f tetrahydromethanopterin S-methyltransferase 23 kD subunit 

Nmar_1748 CENSYa_1431 52 1,599,007 1,600,140 377 r 1,448,791 1,449,873 360 f glutamine--scyllo-inositol transaminase 


Nmar_1751 CENSYa_1428 63 1,601,298 1,602,344 348 f 1,446,783 1,447,820 345 r tyrosyl-tRNA synthetase 

Nmar_1752 CENSYa_1427 75 1,602,341 1,603,135 264 r 1,446,017 1,446,802 261 f precorrin-3B C17-methyltransferase 


Nmar_1754 CENSYa_1423 58 1,604,682 1,606,085 467 r 1,441,118 1,442,497 459 f aspartate kinase 

Nmar_1755 CENSYa_1422 77 1,606,121 1,606,867 248 r 1,440,313 1,441,059 248 f proliferating cell nuclear antigen PcnA 

Nmar_1756 CENSYa_1421 60 1,606,902 1,607,219 105 r 1,439,929 1,440,240 103 f transcription termination factor Tfs 




Nmar_1760 CENSYa_1297 58 1,608,424 1,608,933 169 r 1,309,462 1,309,962 166 f dual specificity protein phosphatase 

Nmar_1761 CENSYa_0285 50 1,608,962 1,609,603 213 r 248,375 249,004 209 r cyclase family protein 

Nmar_1762 CENSYa_0286 64 1,609,679 1,609,936 85 f 249,085 249,345 86 f transcriptional regulator, AsnC family 

Nmar_1763 CENSYa_0287 50 1,609,931 1,611,076 381 r 249,340 250,464 374 r aminotransferase class V 


Nmar_1765 CENSYa_0289 92 1,611,819 1,612,118 99 f 251,173 251,472 99 f 4Fe-4S ferredoxin iron-sulfur binding domain protein 

Nmar_1766 CENSYa_0926 44 1,612,286 1,614,061 591 f 978,506 980,158 550 f adenylate/guanylate cyclase with integral membrane sensor 

Nmar_1767 CENSYa_0925 55 1,614,058 1,615,212 384 r 977,223 978,425 400 f heat shock protein DnaJ domain protein 

Nmar_1768 CENSYa_0920 73 1,615,312 1,615,734 140 r 975,284 975,703 139 f UspA domain protein 


Nmar_1771 CENSYa_1872 79 1,617,863 1,619,308 481 r 1,844,447 1,845,874 475 f glutamine synthetase, type I 

Nmar_1772 CENSYa_1871 70 1,619,396 1,620,742 448 r 1,843,004 1,844,374 456 f lysine 2,3-aminomutase related protein 

Nmar_1773 CENSYa_1870 65 1,620,969 1,622,612 547 f 1,841,072 1,842,844 590 r MscS Mechanosensitive ion channel 

Nmar_1774 CENSYa_1869 48 1,622,652 1,623,530 292 f 1,840,087 1,840,938 283 r ribose-phosphate pyrophosphokinase 

Nmar_1776 CENSYa_1867 59 1,624,474 1,625,373 299 f 1,838,172 1,839,068 298 r ribonuclease Z 


Nmar_1780 CENSYa_1864 57 1,627,309 1,627,902 197 r 1,836,923 1,837,453 176 f dephospho-CoA kinase, CoaE 

Nmar_1781 CENSYa_1863 38 1,627,930 1,628,472 180 f 1,836,293 1,836,820 175 r 2'-5' RNA ligase 

Nmar_1782 CENSYa_1862 45 1,628,469 1,629,800 443 f 1,834,968 1,836,296 442 r tRNA cytidylyltransferase

Nmar_1783 CENSYa_1861 52 1,629,775 1,630,536 253 f 1,834,247 1,834,936 229 r putative serine/threonine protein kinase 


Nmar_1786 CENSYa_1857 77 1,632,027 1,632,767 246 r 1,832,578 1,833,309 243 f short-chain dehydrogenase/reductase SDR 

Nmar_1787 CENSYa_1856 75 1,632,871 1,633,092 73 f 1,832,252 1,832,497 81 r AN1-type Zinc finger protein 



Nmar_1790 CENSYa_1853 82 1,634,005 1,635,477 490 f 1,829,910 1,831,358 482 r glutamine synthetase, type I 


Nmar_1792 CENSYa_1851 82 1,635,808 1,637,520 570 r 1,827,824 1,829,524 566 f thermosome 

Nmar_1793 CENSYa_1849 77 1,637,624 1,638,946 440 f 1,825,900 1,827,225 441 r glycine hydroxymethyltransferase 

Nmar_1794 CENSYa_1827 70 1,638,948 1,642,325 1125 r 1,804,970 1,808,308 1112 r DNA polymerase II, large subunit DP2 

Nmar_1795 CENSYa_1828 68 1,642,337 1,643,089 250 r 1,808,313 1,809,098 261 r geranylgeranylglyceryl phosphate synthase 



Nmar_1798 CENSYa_1836 56 1,644,373 1,644,534 53 f 1,813,052 1,813,207 51 f transcriptional regulator, AbrB family 


N. maritimus ORFs highlighted in blue and bold represent genes implicated in the proposed archaeal ammonia 

oxidation pathway.

Supplemental Table 5: Predicted distribution pattern of proteins diagnostic for major archaeal phyla. 

Protein COG N. maritimus C. symbiosium Euryarchaeota Crenaracheota 

Cell division protein FtsZ COG0206 YP_001582596 YP_875876 + - 

Predicted ATP-dependent protease COG1067 - - + - 

ERCC4-type nuclease COG1111 YP_001582278 YP_876110 + - 

MiaB family, Radical SAM enzyme COG1244 - - + - 

Archaeal DNA polymerase D, small 

subunit 

COG1311 YP_001581340 YP_876754 + - 

Archaeal DNA polymerase D, large 

subunit 

COG1933 YP_001583128 YP_876741 + - 

Predicted membrane protein COG1422 YP_001581738 YP_875263 + - 

Uncharacterised MobA-related protein COG1873 -/? 1 -/? 1 + - 

Uncharacterised conserved protein COG2450 YP_001583060 YP_876421 + - 

Predicted membrane protein COG4243 - - - + 

ribosomal protein L13E COG4352 - - - + 

Uncharacterised protein conserved in 

archaea 

COG4755 - - - + 

Uncharacterised conserved protein COG4353 - - - + 


archaea 

COG4879 - - - + 

Predicted aminopeptidase, Iap family COG4882 - - - + 

Uncharacterized Zn ribbon-containing 

protein 

COG4888 - - - + 

Predicted metallopeptidase COG4900 - - - + 

Ribosomal protein S25 COG4901 

-/? 

(YP_001582264) 

-/? (YP_875201) - + 

Predicted nucleotidyltransferase COG4914 - - - + 

Ribosomal protein S30 COG4919 YP_001582721 YP_875487 - + 

Predicted membrane protein COG4920 - - - + 


archaea 

COG5399 - - - + 

Uncharacterized conserved protein 

containing a coiled-coil 

COG5493 - - - + 

Predicted thioredoxin/glutaredoxin COG5494 - - - + 

EMAP domain RNA-binding protein COG0073 YP_001581931 - + + 

Phosphoribosylanthranilate isomerase COG0135 - - + + 

Molybdopterin biosynthesis enzyme COG0303 - - + + 

Hydrogenase maturation factor COG0309 - - + + 

Molybdenum cofactor biosynthesis 

enzyme 

COG0315 - - + + 

Dihydrodipicolinate synthase/Nacetylneuraminate 

lyase 

COG0329 - - + + 

Membrane protease subunit, 

stomatin/prohibitin homolog 

COG0330 -:? - + + 

NAD(FAD)-dependent dehydrogenase COG0446 -/? - + + 

Molybdopterin biosynthesis enzyme COG0521 - - + + 

Topoisomerase IA COG0550 - - + + 

CDP-diglyceride synthetase COG0575 - - + + 

Methionine synthase II (cobalaminindependent) 

COG0620 - - + + 

Deoxycytidine deaminase COG0717 - - + + 

ABC-type molybdate transport system, 

periplasmic component 

COG0725 - - + + 

Molybdopterin-guanine dinucleotide 

biosynthesis protein A 

COG0746 - - + + 

Predicted transcriptional regulator COG1395 - - + + 

NMD protein affecting ribosome 

stability and mRNA decay 

COG1499 - - + +

Molybdopterin-guanine dinucleotide 

biosynthesis protein 

COG1763 - - + + 

Archaeal S-adenosylmethionine 

synthetase 

COG1812 - - + + 

Predicted transcription factor, homolog 

of eukaryotic MBF1 

COG1813 - - + + 

Transcriptional regulators COG1846 - - + + 

Zn-dependent protease COG1994 - - + + 

Ribosomal protein S24E COG2004 - - + + 

Zn-dependent hydrolase of the betalactamase 

fold 

COG2220 - - + + 

Molybdenum cofactor biosynthesis 

enzyme 

COG2896 - - + + 

1 - Proteins present in N. maritmius and C. symbiosum showing similarity with the euryarchaeal protein. 

? - Proteins present in N. maritimus or/and C. symbiosium but (i) showing a weak similarity or (ii) sharing only a domain with 

euryarchaeal or crenarchaeal proteins

Supplemental Table 6: Gene predictions conserved between the N. maritimus genome and crenarchaeal fosmids and Sargasso 

Sea contigs. 

CRENARCHAEAL GENOME FRAGMENTS 

ORF Genome fragment Strand Left bp Right bp Annotation of top BLAST hit 

Nmar0007 DeepAntEC39 f 5,639 7,168 Peptidylprolyl isomerase 

Nmar0129 DeepAntEC39 r 119,812 119,660 hypothetical protein 

Nmar0221 54d9 r 196,972 196,736 Like-Sm ribonucleoprotein core 

Nmar0271 54d9 f 239,256 239,546 Putative uncharacterized protein 


Nmar0344 4B7 r 308,168 307,311 hypothetical protein 

Nmar0375 54d9 f 341,630 342,262 protein of unknown function UPF0126 

Nmar0413 54d9 f 370,822 372,732 

2-oxoglutarate ferredoxin 

oxidoreductase, alpha subunit 

Nmar0414 54d9 f 372,722 373,684 


oxidoreductase, beta subunit 

Nmar0429 54d9 f 383,085 383,438 hypothetical protein 

Nmar0438 74A4 f 388,048 388,542 Tetratricopeptide TPR_2 repeat protein 


Nmar0527 4B7 f 469,717 469,908 Cold-shock protein DNA-binding 

Nmar0570 54d9 f 507,793 508,344 DNA-3-methyladenine glycosylase I 

Nmar0601 4B7 r 536,325 537,218 Methionine aminopeptidase 

Nmar0602 4B7 f 537,506 537,751 Zn-ribbon protein 

Nmar0616 4B7 f 554,316 557,606 Putative uncharacterized protein 

Nmar0617 4B7 f 557,641 558,132 Protein of unknown function DUF367 

Nmar0624 54d9 r 565,880 564,972 Transcription factor TFIIB cyclin-related 

Nmar0639 4B7 f 579,023 579,685 DSBA oxidoreductase 

Nmar0640 4B7 f 579,686 580,165 hypothetical protein 


Nmar0642 4B7 r 582,430 581,336 DNA-directed DNA polymerase 

Nmar0672 DeepAntEC39 r 613,656 612,652 thioredoxin reductase 

Nmar0726 4B7 f 659,913 661,637 SNF2-related protein 

Nmar0735 DeepAntEC39, 54d9 r 668,933 667,620 hypothetical protein 


Nmar0737 4B7 r 669,981 669,433 heat shock protein DnaJ domain protein 

Nmar0748 4B7 f 675,576 677,768 translation elongation factor aEF-2 


Nmar0850 DeepAntEC39 f 750,195 751,031 

adenylyl cyclase class-3/4/guanylyl 

cyclase 

Nmar0878 4B7 r 772,098 771,370 conserved hypothetical protein 

Nmar0921 DeepAntEC39 r 807,076 806,078 NAD(+) kinase 

Nmar0922 DeepAntEC39 r 807,987 807,118 PfkB domain protein 

Nmar0923 DeepAntEC39 f 808,250 808,537 Ribosomal protein S26E 

Nmar0924 DeepAntEC39 r 809,112 808,540 CDP-alcohol phosphatidyltransferase 

Nmar0925 DeepAntEC39 r 810,065 809,190 putative agmatinase 

Nmar0926 DeepAntEC39 r 811,934 810,069 threonyl-tRNA synthetase 


Nmar0928 DeepAntEC39 f 812,404 813,468 deoxyhypusine synthase 


Nmar0930 DeepAntEC39 f 813,793 814,104 small subunit ribosomal protein S25e 


Nmar0932 DeepAntEC39 f 814,486 815,034 RNA polymerase Rpb6 

Nmar0933 DeepAntEC39 f 815,086 815,400 Alba, DNA/RNA-binding protein 

Nmar0934 DeepAntEC39 r 815,397 815,813 Transcriptional regulator

Nmar0935 DeepAntEC39 f 815,847 816,866 

Asparagine synthase (glutaminehydrolyzing) 

Nmar0937 54d9 r 817,489 818,589 Putative uncharacterized protein 

Nmar0938 DeepAntEC39 r 818,620 819,510 

1,4-dihydroxy-2-napthoate 

octaprenyltransferase 

Nmar0939 DeepAntEC39 r 819,822 820,223 Putative uncharacterized protein 

Nmar0949 54d9 f 827,214 827,495 conserved hypothetical protein 

Nmar0966 74A4 f 844,775 845,488 NH3-dependent NAD+ synthetase 

Nmar0967 74A4 f 845,485 846,873 Cysteinyl-tRNA synthetase 

Nmar0970 4B7, 74A4 f 847,835 848,116 hypothetical protein 

Nmar0971 74A4 r 848,935 848,117 MscS Mechanosensitive ion channel 

Nmar0972 74A4 r 849,716 848,940 conserved hypothetical protein 

Nmar0975 4B7, 74A4 r 852,485 851,328 Protein of unknown function DUF650 

Nmar0976 4B7, 74A4 f 852,580 853,506 periplasmic binding protein 

Nmar0981 74A4 r 858,749 859,114 Putative uncharacterized protein 

Nmar0982 74A4, 54d9 r 859,116 859,526 Coenzyme A-binding protein 

Nmar0983 4B7, 74A4 f 859,607 860,128 Putative uncharacterized protein 

Nmar0984 4B7, 74A4 f 860,165 861,172 Bifunctional protein; biotin repressor 


Nmar0986 4B7, 74A4 f 861,646 863,085 Zn-dependent metalloprotease 

Nmar0987 4B7 f 863,158 864,105 Transcription initiation factor TFIIB 

Nmar0988 74A4 r 864,515 864,108 phosphoribosyltransferase-like 

Nmar0992 54d9 r 867,096 866,668 hypothetical protein 

Nmar0996 4B7 r 869,170 869,568 Hypothetical membrane protein 

Nmar0997 74A4 r 870,511 869,663 Methyltransferase type 11 

Nmar1011 4B7 r 887,979 886,990 nitroreductase 

Nmar1017 4B7, 74A4 f 898,088 899,395 aminotransferase class-III 

Nmar1019 74A4, DeepAntEC39 f 899,956 900,777 TPR-repeat protein 

Nmar1020 74A4, DeepAntEC39 r 900,772 901,476 Double-stranded beta-helix fold enzyme 

Nmar1022 74A4, DeepAntEC39 f 901,921 902,376 Cupin 2 conserved barrel domain protein 

Nmar1023 74A4 f 902,379 902,741 Cupin 2 conserved barrel domain protein 

Nmar1024 74A4, DeepAntEC39 f 902,804 803,475 Molecular chaperone 

Nmar1025 74A4, DeepAntEC39, 54d9 r 903,476 903,883 HIT superfamily hydrolase 



Nmar1028 74A4 f 904,421 905,560 3-hydroxyacyl-CoA dehydrogenase 

Nmar1030 74A4, DeepAntEC39 r 905,845 906,354 Rossman fold nucleotide-binding protein 

Nmar1031 74A4 f 906,617 907,027 Multiple Zn-finger protein 

Nmar1032 74A4 r 907,432 907,229 RNA polymerase Rbp10 

Nmar1033 74A4 r 907,455 907,763 Ribosomal protein S10 

Nmar1034 74A4 r 907,775 909,073 Elongation factor 1-alpha 

Nmar1035 74A4 f 909,195 910,340 Putative uncharacterized protein 

Nmar1036 74A4 f 910,408 910,815 Secreted protein 

Nmar1037 74A4 f 910,852 912,618 DNA ligase 

Nmar1038 74A4 f 912,685 913,338 Coil protein 

Nmar1039 74A4 f 913,430 913,966 tRNA intron endonuclease 


Nmar1043 74A4 f 917,089 918,855 Rossman-fold oxidoreductase 

Nmar1048 74A4 r 924,387 925,160 Permease 

Nmar1049 74A4 r 925,828 925,196 hypothetical protein 

Nmar1050 74A4 r 926,360 925,869 protein of unknown function DUF192 


Nmar1098 54d9 r 1,001,974 1,000,832 phosphoesterase DHHA1

Nmar1121 4B7 f 1,025,889 1,026,476 Ferritin Dps family protein 

Nmar1182 4B7 f 1,080,792 1,082,114 

DEAD/DEAH box helicase domain 

protein 

Nmar1183 4B7 r 985,476 985,778 Putative uncharacterized protein 

Nmar1185 4B7 r 986,566 989,808 Carbamyl phosphate synthetase 

Nmar1186 4B7 r 989,810 990,964 

Carbamoylphosphate synthase, small 

subunit 

Nmar1189 4B7 f 1,085,567 1,085,911 hypothetical protein 

Nmar1190 4B7 f 1,086,156 1,086,416 hypothetical protein 

Nmar1191 4B7 r 1,088,440 1,086,413 protein of unknown function DUF255 

Nmar1275 4B7 r 1,170,906 1,170,133 Rhodanese domain protein 

Nmar1310 54d9 r 1,199,427 1,198,744 glutamine amidotransferase class-I 

Nmar1352 DeepAntEC39 r 1,238,660 1,237,860 regulatory protein ArsR 

Nmar1357 4B7 f 1,243,214 1,243,858 nitroreductase 

Nmar1455 54d9 f 1,323,582 1,324,943 Adenylosucinnate lyase 

Nmar1466 54d9 r 1,333,685 1,334,440 


Nmar1469 54d9 f 1,335,702 1,336,520 hypothetical protein 

Nmar1477 54d9 r 1,345,206 1,347,521 Valyl-tRNA synthetase 

Nmar1500 54d9 r 1,367,890 1,367,240 

hypothetical protein: putative archaeal 

ammonia monoxygenase subunit A 

Nmar1501 54d9 r 1,368,387 1,368,025 hypothetical protein 

Nmar1503 54d9 f 1,369,326 1,369,895 

hypothetical protein: putative archaeal 

ammonia monoxygenase subunit B 

Nmar1524 DeepAntEC39 r 1,385,659 1,385,420 regulatory protein AsnC/Lrp family 

Nmar1622 54d9 r 1,478,462 1,477,386 

Alcohol dehydrogenase zinc-binding 

domain protein 

Nmar1649 54d9 f 1,507,916 1,508,743 hypothetical protein 

Nmar1667 54d9 r 1,526,542 1,525,130 hypothetical protein 

Nmar1685 4B7 r 1,545,979 1,545,383 hypothetical protein 

SARGASSO SEA CONTIGS 

ORF Accession number Strand Left bp Right bp Annotation of top BLAST hit 

Nmar0037 EQ086597 f 28,946 29,659 Organic radical activating enzyme 

Nmar0083 EQ085329 r 70,282 71,367 Cobalt-precorrin-6A synthase 

Nmar0084 EQ085329 f 71,458 72,327 Nucleoside-diphosphate-sugar epimerase 

Nmar0085 EQ085329 r 72,304 73,470 S-adenosylmethionine synthetase 

Nmar0087 EQ085329 f 73,840 74,493 RecA/RadA recombinase related protein 

Nmar0088 EQ085329 f 74,592 77,333 Lhr-like helicase 

Single-stranded DNA-binding 

Nmar0089 EQ085329 r 77,330 78,730 replication protein A, large (70 kD) 

subunit 

Nmar0090 EQ085329 f 78,944 79,882 

ABC-type multidrug transport system, 

ATPase component 

Nmar0091 EQ085329 f 79,866 80,627 

ABC-type multidrug transport system, 

permease component 

Nmar0093 EQ085329 r 81,548 82,600 

Uncharacterized conserved archaeal 

protein 

Nmar0094 EQ085329 f 82,664 83,803 Putative uncharacterized protein 

Nmar0095 EQ085329 f 83,839 84,402 Molecular chaperone GrpE 

Nmar0096 EQ085329 f 84,405 86,315 Molecular chaperone 

Nmar0097 EQ085329 f 86,365 87,450 DnaJ-class molecular chaperone 

Nmar0099 EQ085329 r 88,308 90,212 

Archaeal Glu-tRNA Gln 

amidotransferase subunit E 

Nmar0100 EQ085329 r 90,221 91,510 

Archaeal Glu-tRNA Gln 

amidotransferase subunit D/asparaginase

Nmar0101 EQ085329 r 91,543 93,711 AAA ATPase 

Nmar0102 EQ085329 f 93,816 94,550 Ribosomal protein L2 

Nmar0104 EQ085329 r 94,876 95,208 

Nmar0105 EQ085329 r 95,588 96,502 ATPase of the PP-loop superfamily 

Nmar0107 EQ085329 r 97,656 98,852 

Membrane-associated Zn-dependent 

protease 

Nmar0108 EQ085329 f 98,896 99,207 

Uncharacterized protein involved in 

tolerance to divalent cations 

Nmar0168 EQ086682 f 155,950 156,876 Hypothetical UDP-glucose 4-epimerase 

Nmar0173 EQ084738 r 160,619 161,266 Short-chain alcohol dehydrogenase 

Nmar0198 EQ086546 r 178,988 179,500 Putative uncharacterized protein 

Nmar0231 EQ086498, EQ085222 r 203,816 205,486 Putative uncharacterized protein 


Nmar0270 EQ085249 f 238,081 239,238 

Replicatin factor C/ATPase involved in 

DNA replication 


Nmar0272 EQ085249 f 239,653 241,200 

Acetyl-CoA carboxylase, alpha and beta 

subunits 

Nmar0273 EQ085249 f 241,206 242,693 Biotin carboxylase 

Nmar0274 EQ085249 f 242,700 243,212 

Acetyl/proprionyl-CoA carboxylase, 

alpha subunit 

Nmar0275 EQ085249 r 243,229 243,693 Peroxiredoxin 

Nmar0276 EQ085249 f 243,850 244,182 

NADH-ubiquinone oxidoreductase, 

subunit A 

Nmar0277 EQ085249 f 244,223 244,747 


subunit B 

Nmar0278 EQ085249 f 24,474 245,349 


subunit C 

Nmar0279 EQ085249 f 245,352 246,488 


subunit D 

Nmar0280 EQ085249 f 246,489 247,787 


subunit H 

Nmar0281 EQ085249 f 247,787 248,284 


subunit I 

Nmar0282 EQ085249 f 248,277 248,789 


subunit J 

Nmar0283 EQ085249 f 248,770 249,075 


subunit 4L 

Nmar0284 EQ085249 f 249,075 250,628 

Formate hydrogenlyase, subunit 

3/multisubunit Na/H antiporter 

Nmar0285 EQ085249 f 250,630 252,708 


subunit L 

Nmar0286 EQ085249 f 252,721 254,205 


subunit N 

Nmar0287 EQ085249 r 254,195 255,046 

Geranylgeranyl pyrophosphate 

synthase/geranyltransferase 

Nmar0289 EQ085222 r 256,122 256,727 Orotate phosphoribosyltransferase 


Nmar0291 EQ085222 r 257,852 258,958 

Membrane-associated Zn-dependent 

protease 

Nmar0292 EQ085222 f 259,014 259,529 Acetyltransferase 

Nmar0294 EQ085222 r 259,787 260,614 Methyltransferasae 

Nmar0295 EQ085222 r 260,620 262,407 ATPase, Rnase L inhibitor 

Nmar0352 EQ085222 f 319,896 320,219 Ribosomal protein L30E 

Nmar0353 EQ085222 f 320,271 320,735 Transcription elogation factor 

Nmar0354 EQ085222 f 320,739 321,176 30S ribosomal protein S12P 

Nmar0355 EQ085222 f 321,179 321,778 Ribosomal protein S7

Nmar0356 EQ085222 r 321,937 322,266 


Nmar0359 EQ085222 r 323,446 324,444 Dehydrogenase 

Nmar0367 EQ085222, EP918968 r 330,772 333,639 Leucyl-tRNA snthetase 

Nmar0368 EQ085222, EP918968 r 333,646 336,315 Alanyl-tRNA synthetase 


Nmar0374 EQ085222, EP918968 f 341,230 341,547 

Ribosomal protein 

L12E/L44/L45/RPP1/RPP2 

Nmar0375 EP918968 f 341,630 342,262 Putative uncharacterized protein 

Nmar0381 EP918968 r 345,558 346,424 Ribosomal protein L10 

Nmar0382 EQ085222 r 346,417 347,079 Ribosomal protein L1 

Nmar0383 EQ085222 f 347,271 347,708 Tranascriptional reglator 

Nmar0385 EQ085222, EP918968 r 348,548 349,027 Ribosomal protein L11 

Nmar0386 EQ085222, EP918968 r 349,062 349,520 Ribosomal prrotein L24A 

Nmar0387 EQ085222, EP918968 f 349,736 350,512 


protein 

Nmar0388 EQ085222, EP918968 f 350,509 350,808 

Methylated DNA-protein cysteine 

methyltransferase 

Nmar0390 EQ085222, EP918968 r 351,250 351,705 Ribosomal protein L19E 

Nmar0391 EQ085222, EP918968 r 351,689 352,093 Ribosomal protein L32E 

Nmar0392 EQ085222, EP918968 f 352,349 353,848 

Phosphoenolpyruvate carboxykinase 

(ATP) 

Nmar0393 EQ085222 r 353,850 355,832 Vpr 

Nmar0394 EQ085222 f 355,951 356,571 Superoxide dismutase 

Nmar0395 EQ085222 f 356,654 357,091 Prefoldin, molecular chaperone 

Nmar0396 EQ085222 f 357,095 358,636 Signal recognition particle GTPase 

Nmar0409 EQ086635 r 367,332 368,645 Dehydrogenase 


Nmar0412 EQ086635 r 369,607 370,575 Lactate dehydrogenase 

Nmar0413 EQ086635 f 370,822 372,732 


oxidoreductase, alpha subunit 

Nmar0414 EQ086635 f 372,722 373,684 


oxidoreductase, beta subunit 





Nmar0420 EQ085799 f 376,584 377,852 

3-isopropylmalate dehydratase, large 

subunit 

Nmar0421 EQ085799 f 377,852 378,334 

3-isopropylmalate dehydratase, small 

subunit 

Nmar0422 EQ085799 f 378,343 379,758 

3-isopropylmalate dehydratase, large 

subunit 

Nmar0423 EQ085799 f 379,760 380,344 

3-isopropylmalate dehydratase, small 

subunit 

Nmar0425 EQ085799 r 381,045 381,494 30S ribosomal protein S9P 


Nmar0427 EQ085799 r 381,948 382,295 Ribosomal protein L18E 

Nmar0428 EQ085799 r 382,339 383,004 

DNA-directed RNA polymerase, alpha 

subunit 

Nmar0430 EQ085799 f 383,542 384,228 Exosome complex subunit 

Nmar0431 EQ085799 f 384,228 384,908 Exosome complex RNA-binding protein 

Nmar0432 EQ085799 f 384,908 385,642 Rnase PH 

Nmar0433 EQ085799 f 385,645 386,463 Rnase PH-related exoribonuclease 

Nmar0435 EQ085799 f 386,661 386,906

Nmar0436 EQ085799 f 386,930 387,319 Prefolding, chaperonin cofactor 

Nmar0437 EQ085799 f 387,372 388,058 

Helicase-associated endonuclease for 

fork-structured DNA 

Nmar0439 EQ085799 r 388,545 389,024 Nucleic acid-binding protein 

Nmar0440 EQ085799 r 389,014 389,850 

Inorganic polyphosphate/ATP-NAD 

kinase 

Nmar0443 EQ085799 f 391,108 392,055 Ferredoxin 

Nmar0444 EQ085799 f 392,111 394,177 Glycosyltransferase 

Nmar0445 EQ085799 r 394,174 395,703 Methyl-accepting chemotaxis protein 

Nmar0446 EQ085799 f 396,135 397,460 

Kef-type K transport system, membrane 

component 


Nmar0520 EQ085799 r 464,980 465,363 Ribosomal protien S8E 

Nmar0521 EQ085799 r 465,432 466,010 

Transcriptional regulator containing 

HTH domain 

Nmar0523 EQ085799 f 466,837 467,889 Zn-dependent alcohol dehydrogenase 

Nmar0526 EQ086555 f 469,360 469,671 Translation initiation factor 1 


Nmar0529 EQ085799 f 470,881 471,291 

Translational elongation factor P (EF- 

P)/translation initiation factor 5A 

Nmar0530 EQ085799 f 471,402 472,091 ATPase of the PP-loop superfamily 

Nmar0531 EQ085799 f 472,084 473,412 Signal recognition particle GTPase 

Nmar0532 EQ086635 f 473,435 474,580 Pseudouridylate synthase 




Nmar0537 EQ086635 r 476,694 477,980 Phosphoglycerate mutase 

Nmar0538 EQ086635 f 478,209 479,225 

Galactose-1-phosphate 

uridylyltransferase 


Nmar0562 EQ086635 f 503,038 503,463 

Nmar0578 EQ085355 r 513,299 514,888 Lysyl-tRNA synthetase 

Nmar0579 EQ085355 r 514,892 516,475 Histone acetyltransferase 

Nmar0581 EQ085355 r 516,887 518,137 

Kef-type K transport system, membrane 

component 

Nmar0582 EQ085355 r 518,212 519,234 

Phosphoribosylformylglycinamidine 

cyclo-ligase 


Nmar0585 EQ085355 r 522,090 522,437 

Nmar0590 EQ085355 f 525,903 526,793 Ca2/Na antiporter 

Nmar0592 EQ085355, EQ086504 r 528,167 529,426 Thiamine biosynthesis enzyme 

Nmar0593 EQ086504 r 529,423 530,595 Thiamine biosynthesis enzyme 

Nmar0594 EQ086504 r 530,634 531,773 Trypsin-like serine protease 

Nmar0595 EQ086504 r 531,812 533,020 

Glycosyltransferase involved in cell wall 

biogenesis 


Nmar0597 EQ086504 f 533,768 534,178 Methionyl-tRNA synthetase 

Nmar0598 EQ086504 r 534,324 534,884 

TATA-box binding protein (TBP), 

compoenent of TFIID and TFIIIB 

Nmar0599 EQ086504 f 535,023 536,144 


protein 

Nmar0601 EQ086504 r 536,325 537,218 Methionine aminopeptidase 

Nmar0602 EQ086504 f 537,506 537,751 Zn-ribbon protein 


Nmar0627 EQ086546 f 567,675 569,051 Putative uncharacterized protein

Nmar0652 EQ086546 r 591,731 592,699 

ABC-type oligopeptide transport system, 

ATPase component 



Nmar0674 EQ086597 f 614,251 614,685 Universal stress protein 

Nmar0684 EQ086597 r 624,050 625,159 GTPase 

Nmar0685 EQ086597 f 625,481 626,176 Hydrolase of the HAD superfamily 

Nmar0686 EQ086597 f 626,178 628,064 Arginyl-tRNA synthetase 

Nmar0690 EQ086597 f 631,181 632,062 

L-isoaspartate methyltransferase/tRNA 

(1-methyladenosine) methyltransferase 

Nmar0691 EQ086597 f 632,107 632,970 Sugar kinase 

Nmar0694 EQ086597 f 634,836 635,432 20S proteasome, alpha and beta subunit 

Nmar0695 EQ086597 r 635,442 635,918 RNA-binding protein 

Nmar0696 EQ086597 r 635,915 636,925 Homoserine dehydrogenase 


Nmar0698 EQ086597 r 637,462 639,099 Thymidylate synthase 



Nmar0701 EQ086597 f 640,471 642,387 acetate-CoA ligase 

Nmar0703 EQ086597 r 642,950 643,972 5'-3' exonuclease 

Nmar0788 EQ086597 r 703,382 703,780 Bacterioferritin comigratory protein 

Nmar0795 EQ086466 r 708,081 708,473 Ribosomal protein S8 


Nmar0798 EQ086466 r 709,185 709,901 Ribosomal protein S4E 

Nmar0804 EQ086466 r 711,620 712,375 30S ribosomal protein S3P 




Nmar0809 EQ086466 r 714,340 715,335 50S ribosomal protein L3 


Nmar0811 EQ086466 r 716,385 717,122 20S proteasome, alpha and beta subunit 


Nmar0816 EQ086466 r 720,467 721,117 

Conserved protein implicated in 

secretion 

Nmar0817 EQ086466 r 721,209 722,318 Thiol-disulfide isomerase 

Nmar0818 EQ086466 r 722,323 723,057 Cytochrome c biogenesis protein 


Nmar0889 EP710054 f 780,605 780,922 

Nmar0934 EQ086597 r 815,397 815,813 Transcriptional regulator 

Nmar0935 EQ086597 f 815,847 816,866 



Nmar0938 EQ086597 r 818,620 819,510 

1,4-dihydroxy-2-napthoate 

octaprenyltransferase 


Nmar0940 EQ086597 f 820,283 820,573 


Nmar0963 EQ086597, EP710054 r 841,334 842,989 Methionyl-tRNA synthetase 

Nmar0964 EQ086597, EP710054 r 843,027 844,268 Adenosylhomocysteinase 

Nmar0965 EP710054 f 844,342 844,644 Dioxygenase ferredoxin protein 

Nmar0966 EP710054 f 844,775 845,488 NH3-dependent NAD+ synthetase 

Nmar0967 EP710054 f 845,485 846,873 Cysteinyl-tRNA synthetase 

Nmar0980 EP710054 r 857,503 585,714 GTPase 

Nmar0981 EP710054 r 858,749 859,114 Putative uncharacterized protein

Nmar0982 EP710054 r 859,116 859,526 Coenzyme A-binding protein 


Nmar0984 EP710054 f 860,165 861,172 Bifunctional protein; biotin repressor 

Nmar0985 EP710054 r 861,173 861,535 Putative uncharacterized protein 

Nmar0986 EP710054 f 861,646 863,085 Zn-dependent metalloprotease 

Nmar0987 EP710054 f 863,158 864,105 Transcription initiation factor TFIIB 

Nmar0989 EP710054 r 864,552 864,944 Putative uncharacterized protein 


Nmar0991 EP710054 f 865,566 866,630 Poly(3-hydroxyalkanoate) synthetase 

Nmar0993 EP710054 r 867,093 867,518 Transcriptional regulator 

Nmar0994 EP710054 r 867,555 868,400 Acyltransferase 

Nmar0996 EP710054 r 869,170 869,568 Hypothetical membrane protein 

Nmar1019 EQ086597 f 899,956 900,777 TPR-repeat protein 

Nmar1020 EQ086597 r 900,772 901,476 Double-stranded beta-helix fold enzyme 

Nmar1024 EQ086597 f 902,804 803,475 Molecular chaperone 

Nmar1025 EQ086597 r 903,476 903,883 HIT superfamily hydrolase 

Nmar1028 EQ086597 f 904,421 905,560 3-hydroxyacyl-CoA dehydrogenase 

Nmar1030 EQ086597 r 905,845 906,354 Rossman fold nucleotide-binding protein 

Nmar1031 EQ086597 f 906,617 907,027 Multiple Zn-finger protein 


Nmar1034 EQ086597 r 907,775 909,073 Elongation factor 1-alpha 


Nmar1036 EQ086597 f 910,408 910,815 Secreted protein 

Nmar1037 EQ086597 f 910,852 912,618 DNA ligase 

Nmar1038 EQ086597 f 912,685 913,338 Coil protein 

Nmar1039 EQ086597 f 913,430 913,966 tRNA intron endonuclease 


Nmar1043 EQ086597 f 917,089 918,855 Rossman-fold oxidoreductase 

Nmar1044 EQ086597 f 918,858 919,307 

Nmar1048 EQ086597 r 924,387 925,160 Permease 

Nmar1057 EQ086597 f 932,353 932,916 3-methyladenine DNA glycosylase 

Nmar1058 EQ086597 r 932,913 933,545 

Uncharacterized membrane-associated 

protein 

Nmar1066 EQ086498 f 938,826 940,112 Aspartyl-/asparaginyl-tRNA synthetase 

Nmar1069 EQ086498 r 940,960 941,973 

Isocitrate/isopropylmalate 

dehydrogenase 

Nmar1070 EQ086498 r 942,007 943,524 Isopropylmalate dehydrogenase 

Nmar1071 EQ086498 r 943,521 944,006 Acetolactate synthase 

Nmar1072 EQ086498 r 944,015 945,715 Acetolactate synthase 

Nmar1077 EQ086498 f 981,989 982,393 

Nmar1078 EQ086498 r 982,395 983,390 

DhnA-type fructose-1,6-bisphosphate 

aldolase 

Nmar1079 EQ086498 r 983,428 984,456 

L-iditol 2-dehydrogenase/threonine 





Nmar1084 EQ086498 f 985,813 986,562 ICC-like phosphoesterase 

Nmar1085 EQ086498 r 986,566 989,808 Carbamyl phosphate synthetase 

Nmar1086 EQ086498 r 989,810 990,964 

Carbamoylphosphate synthase, small 

subunit 

Nmar1088 EQ086498 r 992,659 993,852 AAA ATPase 

Nmar1092 EQ086498 f 996,256 996,711 Translation initiation inhibitor 

Nmar1098 EQ086498 r 1,000,832 1,001,974 Single-stranded DNA-specific

Nmar1099 EQ086498 f 1,002,328 1,003,491 

exonuclease 

Citrate synthase 

Nmar1101 EQ086498 f 1,003,778 1,004,125 

Nmar1103 EQ086498 r 1,005,416 1,007,002 Excinuclease nuclease subunit 

Nmar1104 EQ086498 r 1,006,999 1,009,818 Excinuclease ATPase subunit 

Nmar1105 EQ086498 r 1,009,805 1,011,757 UvrABC system protein B 

Nmar1106 EQ086498 r 1,011,807 1,012,457 Flavin-nucleotide-binding protein 

Nmar1109 EQ086498 r 1,013,628 1,014,734 Flavin-dependent oxidoreductase 

Nmar1110 EQ086498 r 1,014,829 1,015,773 Alcohol dehydrogenase 

Nmar1111 EQ086498 f 1,015,826 1,017,103 Hydroxypyruvate reductase 

Nmar1113 EQ086498 r 1,017,395 1,018,360 Exonuclease of the beta-lactamase fold 

5,10-methylene-tetrahydrofolate 

Nmar1117 EQ086498 f 1,019,750 1,020,613 dehydrogenase/methenyltetrahydrofolase 

cyclohydrolase 

Nmar1119 EQ086498 r 1,021,157 1,022,422 Phosphoribosylamine-glycine ligase 

Nmar1120 EQ086498 f 1,022,506 1,025,703 Isoleucyl-tRNA synthetase 

Nmar1122 EQ086498 f 1,026,579 1,026,908 Putative uncharacterized protein 

Nmar1123 EQ086498 f 1,026,949 1,028,055 Succinyl-CoA synthetase, beta subunit 

Nmar1124 EQ086498 f 1,028,052 1,028,969 Succinyl-CoA synthetase, alpha subunit 

Nmar1254 EQ086682 r 1,150,749 1,151,213 Ribosomal protein L15E 

Nmar1255 EQ086682 f 1,151,421 1,152,080 

Secreted periplasmic Zn-dependent 

protease 

Nmar1257 EQ086682 r 1,152,589 1,153,482 Putative uncharacterized protein 

Nmar1258 EQ086682 r 1,153,583 1,154,515 Phosphoglycerate dehydrogenase 

Nmar1259 EQ086682 r 1,154,583 1,155,995 

Putative copper-containing nitrite 

reductase 

Nmar1260 EQ086682 f 1,156,106 1,157,467 Fumarate hydratase/fumarase 

Nmar1261 EQ086682 f 1,157,471 1,158,739 2-methylthioadenine synthetase 

Nmar1262 EQ086682 f 1,158,813 1,159,763 Cell division GTPase 

Nmar1263 EQ086682 r 11,589,800 1,160,363 Ribosomal protein L31E 

Nmar1265 EQ086682 r 1,160,560 1,161,111 NTP pyrophosphohydrolase 


Nmar1267 EQ086682 r 1,161,915 1,164,413 

Methionine synthase I, cobalaminbinding 

domain 

Nmar1268 EQ086682 r 1,164,446 1,165,408 

Methionine synthase I (cobalamindependent), 

methyltransferase domain 

Nmar1269 EQ086682 f 1,165,474 1,166,622 Permease 

Nmar1283 EQ086682 r 1,175,356 1,176,114 

ABC-type nitrate/sulfonate/bicarbonaet 

transport system, permease component 

Nmar1284 EQ086682 r 1,176,120 1,176,884 


transport system, ATPase component 

Nmar1285 EQ086682 r 1,176,884 1,177,909 


transport system, periplasmic component 

Nmar1286 EQ086682 f 1,178,045 1,179,244 Argininosuccinate synthase 

Nmar1288 EQ086682 f 1,179,408 1,180,265 

Glutathione synthase/ribosomal protein 

S6 modification enzyme 

Nmar1289 EQ086682 f 1,180,294 1,181,340 N-acetyl-gamma-glutamyl-phosphate 

reductase 

Nmar1291 EQ086682 f 1,182,139 1,183,320 

Pyridoxal-phosphate-dependent 

aminotransferase 

Nmar1292 EQ086682 f 1,183,307 1,183,726 Transcriptional regulator 

Nmar1293 EQ086682 f 1,183,825 1,185,006 2-isopropylmalate synthase 

Nmar1295 EQ086682 f 1,185,192 1,186,034 

Glutathione synthase/ribosomal protein 

S6 modification enzyme 

Nmar1296 EQ086682 f 1,186,038 1,187,165 

Acetylornithine deacetylase/succinyldiaminopimelate 

desuccinylase

Nmar1297 EQ086682 f 1,187,165 1,188,202 Dipthine synthase 

Nmar1298 EQ086682 f 1,188,243 1,188,863 Cytidylyltransferase 

Nmar1301 EQ086682 r 1,191,274 1,191,957 

Transcriptional regulator of a 

riboflavin/FAD biosynthetic operon 

Nmar1302 EQ086682 r 1,191,966 1,193,108 Bacteral-type DNA primase 

Nmar1303 EQ086682 r 1,193,261 1,193,611 

Putative copper-containing nitrite 

reductase 

Nmar1308 EQ086682 r 1,195,786 1,196,547 

Enoyl-CoA hydratase/carnithine 

racemase 

Nmar1309 EQ086682 r 1,196,584 1,198,701 Acyl-CoA synthetase 

Nmar1312 EQ086682 r 1,199,847 1,201,121 Glutamate/leucine dehydrogenase 

Nmar1314 EQ086682 f 1,201,690 1,202,412 20S proteasome, alpha and beta subunits 

Nmar1315 EQ086682 f 1,202,461 1,203,153 

Methylase involved in 

ubiquinone/menaquinone biosynthesis 

Nmar1316 EQ086682 f 120,150 1,204,304 

Permease of the major facilitator 

superfamily 

Nmar1321 EQ086635 f 1,208,508 1,209,338 Short-chain alcohol dehydrogenase 

Nmar1322 EQ086635 f 1,209,416 1,211,071 DNA topoisomerase IB 


Nmar1378 EQ085799 r 1,263,279 1,264,037 Metallophosphoesterase 

Nmar1379 EQ086635 f 1,264,147 1,265,178 

Isocitrate/isopropylmalate 


Nmar1380 EQ086635 r 1,265,179 1,265,715 

Methylase of polypeptide chain release 

factor 

Nmar1381 EQ086635 r 1,265,690 1,266,394 

Dimethyladenosine transferase (rRNA 

methylation) 

Nmar1382 EQ086635 r 1,266,391 1,266,957 RNA-binding protein 


Nmar1384 EQ086635 r 1,267,307 1,267,606 Ribosomal proteini L21E 

Nmar1386 EQ086635 r 1,268,792 1,269,958 

DNA repair and recombination protein, 

radA 

Nmar1388 EQ086635 r 1,270,263 1,271,987 Fe-S oxidoreductase 

Nmar1389 EQ086635 r 1,272,039 1,272,851 2-polyprenylphenol hydroxylase 

Nmar1390 EQ086635 r 1,272,835 1,273,746 Dihydroorotate dehydrogenase 

Nmar1391 EQ086635 f 1,273,832 1,275,244 

Ribosomal bioogenesis protein/nucleolar 

protein 

Nmar1392 EQ086635 f 1,275,231 1,275,905 Fibrillarin-like rRNA methylase 

Nmar1393 EQ086635 r 1,275,902 1,276,519 Ribonuclease 

Nmar1394 EQ086635 f 1,276,580 1,277,728 

N2,N2-demthylguanosine tRNA 

methyltransferase 

Nmar1395 EQ086635 r 1,277,718 1,278,311 rRNA large subunit methyltransferase J 

Nmar1396 EQ086635 r 1,278,308 1,279,183 Transcriptional regulator 

Nmar1397 EQ086635 r 1,279,233 1,279,709 

Nmar1398 EQ086635 f 1,279,813 1,281,906 ATP-dependent endonuclease 

Nmar1407 EQ086555 r 1,287,346 1,288,458 DNA topoisomerase VI, subunit A 

Nmar1408 EQ086555 r 1,288,445 1,290,325 DNA topoisomerase VI, subunit B 

Nmar1409 EQ086555 r 1,290,312 1,290,884 RNA-binding protein 

Nmar1410 EQ086555 r 1,290,881 1,291,660 Serine/threonine protein kinase 


Nmar1412 EQ086555 r 1,292,013 1,292,435 

Translation initiation factor 2, beta 

subunit/elF-5 N-terminal domain 


Nmar1415 EQ086555 f 1,293,472 1,294,617 Membrane protein 

Nmar1416 EQ086555 f 1,294,697 1,295,488 Undecaprenyl pyrophosphate synthase 

Nmar1417 EQ086555 f 1,295,485 1,295,904 

Diadenosine 5'5'''-P1,P4-tetraphosphate 

pyrophosphohydrolase

Nmar1418 EQ086555 r 1,295,884 1,296,624 Orotidine-5'phosphate decarboxylase 

Nmar1420 EQ086555 r 1,297,338 1,298,339 Adenylosuccinate synthetase 



Nmar1447 EQ084738 f 1,317,902 1,319,200 Phosphomannomutase 

Nmar1448 EQ084738 f 1,319,184 1,320,116 Thiamine monophosphate kinase 

Nmar1450 EQ084738 f 1,320,746 1,321,072 30S ribosomal protein S11P 




Nmar1455 EQ084738 f 1,323,582 1,324,943 Adenylosucinnate lyase 

Nmar1466 EQ084738 r 1,333,685 1,334,440 


Nmar1468 EQ084738 f 1,335,364 1,335,639 Pterin-4a-carbinolamine dehydratase 

Nmar1477 EQ084738 r 1,345,206 1,347,521 Valyl-tRNA synthetase 

Nmar1479 EQ084738 r 1,348,150 1,349,118 

Pyridoxine/pyridoxal 5-phosphate 

biosynthesis enzyme 


Nmar1482 EQ084738 r 1,350,892 1,353,156 

3-isopropylmalate isomerase/aconitase 

A 




Nmar1488 EQ084738 r 1,355,702 1,357,345 

Phenylalanyl-tRNA synthetase, beta 

subunit 

Nmar1489 EQ084738 r 1,357,336 1,358,724 

Phenylalanyl-tRNA synthetase, alpha 

subunit 

Nmar1490 EQ084738 f 1,358,772 1,359,884 Tryptophanyl-tRNA synthetase 

Nmar1491 EQ084738 f 1,359,881 1,360,162 


Nmar1494 EQ084738 r 1,361,774 1,362,250 

Pantetheine-phosphate 

nucleotidyltransferase 

Nmar1496 EQ084738 f 1,363,804 1,364,358 Thiol-disulfide isomerase 

Nmar1592 EP710054 f 1,456,838 1,457,221 Putative uncharacterized protein 


Nmar1618 EQ086597 r 1,474,520 1,475,245 TPR repeat protein 


Nmar1641 EQ086546 r 1,499,144 1,500,094 BatA 


Nmar1643 EQ086546 r 1,500,977 1,501,999 MoxR-like ATPase 

Nmar1644 EQ086682, EP710054 r 1,502,127 1,503,287 

Secreted Zn-dependent protease 

containing TPR repeats 

Nmar1650 EQ086498 f 1,508,803 1,509,615 

Copper binding protein, 

pastocyanin/azurin family

Nitrosopumilus maritimus genome reveals unique - de la Torre ...

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