whole genome taxonomic classication for prokaryotic plant pathogens
TRANSCRIPT
Whole genome taxonomicclassification for prokaryoticplant pathogensNextGenBug, University of Edinburgh8th December 2015
Leighton PritchardInformation and Computational SciencesThe James Hutton Institute
Acceptable Use Policy
Recording of this talk, taking photos, discussing the content usingemail, Twitter, blogs, etc. is permitted (and encouraged),providing distraction to others during the presentation is minimised.
These slides will be available on SlideShare and the NextGenBugF1000 channel.
Table of Contents
IntroductionThe Insidious Dickeya MenaceqPCR Primer Design From Whole Genomes
ClassificationClassification is CriticalWhole-Genome Classification
Soft-Rot EnterobacteriaA Tangled TaxonomyThe Cracks Are Showing. . .
ConclusionFinal ThoughtsWithout Whom. . .
If you’re interested. . .Hidden diversity: pre-publication
Dickeya on crops and ornamentalsa
aCzajkowski et al. (2015) Ann. Appl. Biol. doi:10.1111/aab.12166
• Dickeya spp. are virulent enterobacterial soft-rottingpathogens of ornamental and crop plants
1 2
1Landesanst. f. Pflanzenbau und Pflanzenschutz, Mainz Archive
2Florida Division of Plant Industry Archive
Dickeya spp. moving across Europeab
aToth et al. (2011) Plant Path. doi:10.1111/j.1365-3059.2011.02427.x
bParkinson et al. (2015) Eur. J. Plant Path. doi:10.1007/s10658-014-0523-5
• D. dianthicola is established across Europe
• D. solani is an emerging, encroaching threat
Legislationa
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• European and Mediterranean Plant Protection Organisation(EPPO):member states should regulate D. dianthicola and E.amylovora as quarantine pests (A2 list)
• Seed Potatoes (Scotland) Amendment Regulations (2010):zero tolerance policy for all Dickeya spp. on potatoes inScotland to ensure production of ‘clean’ (disease-free) seedpotato production for export
• EUPHRESCO consortium: control and epidemiology
Why legislate by taxonomy?a
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• “Easy” to incorporate into legislation: binary classification
• Assumption: taxonomy can be determined precisely
• Assumption: taxonomy is a proxy for disease risk
Problems with legislating by taxonomya
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• Is bacterial taxonomy correct?
• Is a species concept even relevant for bacteria?
• Taxonomy is ‘vertical’, but pathogenicity may be ‘laterally’transferred (plasmid-borne, etc.)3
• Mapping from taxonomy to phenotype is not one-to-one4
• Testing for disease phenotypes not straightforward
• Relationship between gene complement and disease not fullyunderstood
3Toth et al. (2006) Ann. Rev. Phyto. doi:10.1146/annurev.phyto.44.070505.143444
4Deans et al. (2015) PLoS Biol. doi:10.1371/journal.pbio.1002033
Dickeya qPCR diagnosticsa
aPritchard et al. (2013) Plant Path. doi:10.1111/j.1365-3059.2012.02678.x
• To legislate on or quarantine contaminated materials, one hasto be able to identify and discriminate the pathogen
• qPCR is still cheaper, quicker and easier than bacterialgenome sequencing (for now, anyway. . . )5
• No qPCR primers existed to distinguish among Dickeya spp.
• Having sequenced 25 Dickeya isolates, we were approached todevelop diagnostic primers at the species/isolate level
5Czajkowski et al. (2015) Ann. Appl. Biol. doi:10.1111/aab.12166
qPCR Primer Designa
aPritchard et al. (2012) PLoS One doi:10.1371/journal.pone.0034498
1. Bulk predict primer sets on all chromosomes (Primer3)2. Predict cross-amplification in silico (primersearch)3. Evaluate in vitro against panel of previously “unseen” isolates
of known class
targets
o�-targets
classi�cation
V
IV
III
II
I
genomes
IIIIIIIVV
Table of Contents
IntroductionThe Insidious Dickeya MenaceqPCR Primer Design From Whole Genomes
ClassificationClassification is CriticalWhole-Genome Classification
Soft-Rot EnterobacteriaA Tangled TaxonomyThe Cracks Are Showing. . .
ConclusionFinal ThoughtsWithout Whom. . .
If you’re interested. . .Hidden diversity: pre-publication
Classification matters!a
aPritchard et al. (2012) PLoS One doi:10.1111/j.1365-3059.2012.02678.x
Primer design process:
1. Design large numbers of primers to training (draft) genomesfrom the target groups
2. Test cross-hybridisation of primer sets in silico against targetand off-target groups
3. Screen primers against broader set of off-target sequences
4. Classify primer sets according to in silico specificity
5. Evaluate specificity against unseen panel of target/off-targetorganisms
Classification is also a problem!a
aPritchard et al. (2013) Plant Path. doi:10.1111/j.1365-3059.2012.02678.x
• qPCR design gave no diagnostic primers for several species.• Misassigned species in GenBank made ‘training’ impossible
6
6ML tree, recA
Consequences of misclassificationa
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
Failed classification has real-world consequences:
• False positives (type I errors):clean samples rejected: economic costfarms quarantined/close: economic/social cost
• False negatives (type II errors):(irreversible) introduction of infectious materialpotential for novel host jumps and spread
“Gold-standard”, correctly classified training and test sets essentialto estimate classifier error rates.
MiSI: 18% of NCBI genomes: bacterial species misclassified7
7Varghese et al. (2015) Nucl. Acids Res. doi:10.1093/nar/gkv657
The cracks are showing. . .
Bacterial taxonomy?
DNA-DNA hybridisationa
aMorello-Mora and Amann (2001) FEMS Micro. Rev. doi:10.1016/S0168-6445(00)00040-1
• “Gold Standard” forprokaryotic taxonomy,since 1960s. “70%identity ≈ same species.”
• Denature DNA from twoorganisms.
• Allow to anneal.Reassociation ≈ similarity,measured as ∆T ofdenaturation curves.
Proxy for sequence similarity - replace with genome analysis8?
8Chan et al (2012) BMC Microbiol. doi:10.1186/1471-2180-12-302
Average Nucleotide Identity (ANIm)a
aRichter and Rossello-Mora (2009) Proc. Natl. Acad. Sci. USA doi:10.1073/pnas.0906412106
1. Align genomes(MUMmer)2. ANIm: Mean% identity of allmatches
• DDH:ANImlinear
• 70%ID ≈95%ANIb
ANIm
• Average identity of all ‘homologous’ regions
• Approximates a limiting case of MLST/MLSA/multigenecomparisons
• Straightforward principle
• Classification not dependent on dataset composition (unliketree methods)
Table of Contents
IntroductionThe Insidious Dickeya MenaceqPCR Primer Design From Whole Genomes
ClassificationClassification is CriticalWhole-Genome Classification
Soft-Rot EnterobacteriaA Tangled TaxonomyThe Cracks Are Showing. . .
ConclusionFinal ThoughtsWithout Whom. . .
If you’re interested. . .Hidden diversity: pre-publication
Tangled taxonomy of SREa
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
Gammaproteobacteria, Enterobacteria taxonomy difficult toresolve, in general9
Species classification mostly polyphasic/phenotypic.
• Soft rot enterobacteria (SRE) all originally Erwinia spp.
• SRE now three distinct genera (Dickeya, Pectobacterium,Erwinia)
• Pectobacterium used to be E. chrysanthemi
• Dickeya used to be P. chrysanthemi
• Old names hold over in the literature, collections, etc.
9Williams et al. (2010) J. Bact. doi:10.1128/JB.01480-09
Tangled taxonomy of SREa
aCzajkowski et al. (2015) Ann. Appl. Biol. doi:10.1111/aab.12166
D. solani ANIma
avan der Wolf et al. (2014) Int. J. Syst. Evol. Micr. doi:10.1099/ijs.0.052944-0
Defining D. solani as a new species, with ANIm:
34 Dickeya spp. ANIma
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• Ninespecies-levelgroups (twonovel)
• Three speciesmisidentifiedin GenBank
Dic
keya_s
ola
ni_
GB
BC
2040
Dic
keya_s
ola
ni_
IPO
2222
Dic
keya_s
ola
ni_
MK
10
Dic
keya_s
ola
ni_
MK
16
Dic
keya_s
ola
ni_
AM
YI0
1D
icke
ya_s
ola
ni_
AM
WE01
Dic
keya_d
ianth
icola
_NC
PPB
_3534
Dic
keya_d
ianth
icola
_GB
BC
2039
Dic
keya_d
ianth
icola
_NC
PPB
_453
Dic
keya_d
ianth
icola
_IPO
980
Dic
keya_s
pp_N
CPPB
_3274
Dic
keya_s
pp_M
K7
Dic
keya_d
adanti
i_N
CPPB
_2976
Dic
keya_d
adanti
i_3937_u
id52537
Dic
keya_d
adanti
i_N
CPPB
_3537
Dic
keya_d
adanti
i_N
CPPB
_898
Dic
keya_z
eae_A
PM
V01
Dic
keya_z
eae_A
JVN
01
Dic
keya_z
eae_N
CPPB
_3531
Dic
keya_z
eae_C
SL_
RW
192
Dic
keya_d
adanti
i_Ech
586_u
id42519
Dic
keya_z
eae_A
PW
M01
Dic
keya_z
eae_M
K19
Dic
keya_z
eae_N
CPPB
_3532
Dic
keya_z
eae_N
CPPB
_2538
Dic
keya_s
pp_N
CPPB
_569
Dic
keya_c
hry
santh
am
i_N
CPPB
_402
Dic
keya_c
hry
santh
am
i_N
CPPB
_516
Dic
keya_z
eae_E
ch1591_u
id59297
Dic
keya_c
hry
santh
am
i_N
CPPB
_3533
Dic
keya_a
quati
ca_D
W_0
440
Dic
keya_a
quati
ca_C
SL_
RW
240
Dic
keya_d
adanti
i_Ech
703_u
id59363
Dic
keya_p
ara
dis
iaca
_NC
PPB
_2511
Dickeya_solani_GBBC2040Dickeya_solani_IPO2222Dickeya_solani_MK10Dickeya_solani_MK16Dickeya_solani_AMYI01Dickeya_solani_AMWE01Dickeya_dianthicola_NCPPB_3534Dickeya_dianthicola_GBBC2039Dickeya_dianthicola_NCPPB_453Dickeya_dianthicola_IPO980Dickeya_spp_NCPPB_3274Dickeya_spp_MK7Dickeya_dadantii_NCPPB_2976Dickeya_dadantii_3937_uid52537Dickeya_dadantii_NCPPB_3537Dickeya_dadantii_NCPPB_898Dickeya_zeae_APMV01Dickeya_zeae_AJVN01Dickeya_zeae_NCPPB_3531Dickeya_zeae_CSL_RW192Dickeya_dadantii_Ech586_uid42519Dickeya_zeae_APWM01Dickeya_zeae_MK19Dickeya_zeae_NCPPB_3532Dickeya_zeae_NCPPB_2538Dickeya_spp_NCPPB_569Dickeya_chrysanthami_NCPPB_402Dickeya_chrysanthami_NCPPB_516Dickeya_zeae_Ech1591_uid59297Dickeya_chrysanthami_NCPPB_3533Dickeya_aquatica_DW_0440Dickeya_aquatica_CSL_RW240Dickeya_dadantii_Ech703_uid59363Dickeya_paradisiaca_NCPPB_2511
0.00
0.25
0.50
0.75
1.00
AN
Im_p
erc
enta
ge_i
denti
ty
55 Pectobacterium spp. ANIma
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• Tenspecies-levelgroups (fournovel)
• P. carotovorumsplit: severalspecies
• P. wasabiaesplit: twospecies
P. a
tros
eptic
um S
CR
I104
3P
. atr
osep
ticum
NC
PP
B 3
404
P. a
tros
eptic
um J
G10
08
P. a
tros
eptic
um 2
1AP
. atr
osep
ticum
CF
BP
627
6P
. atr
osep
ticum
NC
PP
B 5
49P
. atr
osep
ticum
ICM
P 1
526
P. c
arot
ovor
um P
C1
P. c
arot
ovor
um U
GC
32P
. bet
avas
culo
rum
NC
PP
B 2
793
P. b
etav
ascu
loru
m N
CP
PB
279
5P
. car
otov
orum
M02
2P
. was
abia
e C
FB
P 3
304
P. w
asab
iae
NC
PP
B 3
701
P. w
asab
iae
NC
PP
B37
02P
. was
abia
e C
FIA
1002
P. w
asab
iae
WP
P16
3P
. was
abia
e R
NS
08.4
2.1A
P. s
p. S
CC
3193
SC
C31
93P
. car
otov
orum
BC
D6
P. c
arot
ovor
um Y
C D
49P
. car
otov
orum
BC
S2
P. c
arot
ovor
um Y
C D
29P
. car
otov
orum
YC
D65
P. c
arot
ovor
um C
FIA
1001
P. c
arot
ovor
um P
CC
21P
. car
otov
orum
YC
D46
P. c
arot
ovor
um Y
C T
31P
. car
otov
orum
YC
D62
P. c
arot
ovor
um Y
C T
3P
. car
otov
orum
CF
IA10
09P
. car
otov
orum
YC
D52
P. c
arot
ovor
um Y
C D
21P
. car
otov
orum
YC
D64
P. c
arot
ovor
um Y
C D
60P
. car
otov
orum
CF
IA10
33P
. car
otov
orum
PB
R16
92P
. car
otov
orum
LM
G 2
1371
P. c
arot
ovor
um B
D25
5P
. car
otov
orum
ICM
P 1
9477
P. c
arot
ovor
um L
MG
213
72P
. car
otov
orum
KK
H3
P. c
arot
ovor
um N
CP
PB
3841
P. c
arot
ovor
um N
CP
PB
383
9P
. car
otov
orum
BC
S7
P. c
arot
ovor
um Y
C T
1P
. car
otov
orum
NC
PP
B 3
395
P. c
arot
ovor
um Y
C D
57P
. car
otov
orum
BC
T2
P. c
arot
ovor
um IC
MP
570
2P
. car
otov
orum
NC
PP
B 3
12P
. car
otov
orum
YC
D16
P. c
arot
ovor
um Y
C T
39P
. car
otov
orum
WP
P14
P. c
arot
ovor
um B
C T
5
P. atrosepticum SCRI1043P. atrosepticum NCPPB 3404P. atrosepticum JG1008P. atrosepticum 21AP. atrosepticum CFBP 6276P. atrosepticum NCPPB 549P. atrosepticum ICMP 1526P. carotovorum PC1P. carotovorum UGC32P. betavasculorum NCPPB 2793P. betavasculorum NCPPB 2795P. carotovorum M022P. wasabiae CFBP 3304P. wasabiae NCPPB 3701P. wasabiae NCPPB3702P. wasabiae CFIA1002P. wasabiae WPP163P. wasabiae RNS08.42.1AP. sp. SCC3193 SCC3193P. carotovorum BC D6P. carotovorum YC D49P. carotovorum BC S2P. carotovorum YC D29P. carotovorum YC D65P. carotovorum CFIA1001P. carotovorum PCC21P. carotovorum YC D46P. carotovorum YC T31P. carotovorum YC D62P. carotovorum YC T3P. carotovorum CFIA1009P. carotovorum YC D52P. carotovorum YC D21P. carotovorum YC D64P. carotovorum YC D60P. carotovorum CFIA1033P. carotovorum PBR1692P. carotovorum LMG 21371P. carotovorum BD255P. carotovorum ICMP 19477P. carotovorum LMG 21372P. carotovorum KKH3P. carotovorum NCPPB3841P. carotovorum NCPPB 3839P. carotovorum BC S7P. carotovorum YC T1P. carotovorum NCPPB 3395P. carotovorum YC D57P. carotovorum BC T2P. carotovorum ICMP 5702P. carotovorum NCPPB 312P. carotovorum YC D16P. carotovorum YC T39P. carotovorum WPP14P. carotovorum BC T5
0.00
0.25
0.50
0.75
1.00
AN
Im_p
erce
ntag
e_id
entit
y
Criticisms of ANIma
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• 95% threshold ‘arbitrary’
• Taxonomic classification, not phylogenetic reconstruction
• No functional (or gene-based) interpretation: still needpangenome classification and analysis
• Only considers ‘homologous’ regions:• Define ‘homologous’• misled by HGT/LGT?• misled by distant relationships/low extent of homology?
Coverage plots help interpretation: exclude HGT/LGT lowhomology bias.
55 Pectobacterium spp. ANIma
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• All isolatesalign over>50% ofwholegenome
P. c
arot
ovor
um Y
C T
31P
. car
otov
orum
YC
D21
P. c
arot
ovor
um Y
C T
3P
. car
otov
orum
CF
IA10
09P
. car
otov
orum
YC
D64
P. c
arot
ovor
um Y
C D
52P
. car
otov
orum
YC
D62
P. c
arot
ovor
um Y
C D
60P
. car
otov
orum
YC
D49
P. c
arot
ovor
um B
C D
6P
. car
otov
orum
YC
D65
P. c
arot
ovor
um Y
C D
46P
. car
otov
orum
BC
S2
P. c
arot
ovor
um Y
C D
29P
. car
otov
orum
PC
C21
P. c
arot
ovor
um C
FIA
1001
P. c
arot
ovor
um Y
C T
1P
. car
otov
orum
NC
PP
B 3
395
P. c
arot
ovor
um U
GC
32P
. car
otov
orum
KK
H3
P. c
arot
ovor
um B
C S
7P
. car
otov
orum
ICM
P 5
702
P. c
arot
ovor
um N
CP
PB
312
P. c
arot
ovor
um B
C T
2P
. car
otov
orum
YC
D16
P. c
arot
ovor
um Y
C T
39P
. car
otov
orum
YC
D57
P. c
arot
ovor
um W
PP
14P
. car
otov
orum
BC
T5
P. c
arot
ovor
um C
FIA
1033
P. c
arot
ovor
um P
C1
P. c
arot
ovor
um L
MG
213
71P
. car
otov
orum
BD
255
P. c
arot
ovor
um P
BR
1692
P. c
arot
ovor
um IC
MP
194
77P
. car
otov
orum
LM
G 2
1372
P. w
asab
iae
CF
BP
330
4P
. was
abia
e N
CP
PB
370
1P
. was
abia
e N
CP
PB
3702
P. s
p. S
CC
3193
SC
C31
93P
. was
abia
e W
PP
163
P. w
asab
iae
RN
S08
.42.
1AP
. was
abia
e C
FIA
1002
P. a
tros
eptic
um C
FB
P 6
276
P. a
tros
eptic
um N
CP
PB
340
4P
. atr
osep
ticum
NC
PP
B 5
49P
. atr
osep
ticum
ICM
P 1
526
P. a
tros
eptic
um S
CR
I104
3P
. atr
osep
ticum
JG
100
8P
. atr
osep
ticum
21A
P. c
arot
ovor
um N
CP
PB
3841
P. c
arot
ovor
um N
CP
PB
383
9P
. car
otov
orum
M02
2P
. bet
avas
culo
rum
NC
PP
B 2
793
P. b
etav
ascu
loru
m N
CP
PB
279
5
P. carotovorum M022P. carotovorum YC T1P. carotovorum KKH3P. carotovorum UGC32P. carotovorum CFIA1033P. carotovorum NCPPB3841P. carotovorum NCPPB 3839P. carotovorum BC S7P. carotovorum ICMP 19477P. carotovorum BD255P. carotovorum LMG 21372P. carotovorum PBR1692P. carotovorum LMG 21371P. carotovorum PC1P. carotovorum ICMP 5702P. carotovorum NCPPB 312P. carotovorum YC D57P. carotovorum BC T2P. carotovorum YC D16P. carotovorum WPP14P. carotovorum BC T5P. carotovorum YC D62P. carotovorum YC T31P. carotovorum YC D52P. carotovorum YC T3P. carotovorum YC D64P. carotovorum YC D21P. carotovorum YC D60P. carotovorum YC T39P. carotovorum CFIA1001P. carotovorum YC D46P. carotovorum YC D49P. carotovorum BC D6P. carotovorum YC D65P. carotovorum BC S2P. carotovorum YC D29P. carotovorum PCC21P. carotovorum CFIA1009P. atrosepticum ICMP 1526P. atrosepticum CFBP 6276P. atrosepticum SCRI1043P. atrosepticum NCPPB 3404P. atrosepticum NCPPB 549P. atrosepticum JG1008P. atrosepticum 21AP. wasabiae WPP163P. sp. SCC3193 SCC3193P. wasabiae RNS08.42.1AP. wasabiae CFIA1002P. wasabiae CFBP 3304P. wasabiae NCPPB 3701P. wasabiae NCPPB3702P. carotovorum NCPPB 3395P. betavasculorum NCPPB 2793P. betavasculorum NCPPB 2795
0.00
0.25
0.50
0.75
1.00
AN
Im_a
lignm
ent_
cove
rage
34 Dickeya spp. ANIma
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
• Most isolatesalign over>50% ofwholegenome
• Community:two outlierspecies arequestionableassignments
Dic
keya_d
adanti
i_Ech
703_u
id59363
Dic
keya_p
ara
dis
iaca
_NC
PPB
_2511
Dic
keya_a
quati
ca_D
W_0
440
Dic
keya_a
quati
ca_C
SL_
RW
240
Dic
keya_s
ola
ni_
MK
10
Dic
keya_s
ola
ni_
AM
YI0
1D
icke
ya_s
ola
ni_
AM
WE01
Dic
keya_s
ola
ni_
MK
16
Dic
keya_s
ola
ni_
GB
BC
2040
Dic
keya_s
ola
ni_
IPO
2222
Dic
keya_d
ianth
icola
_NC
PPB
_453
Dic
keya_d
ianth
icola
_GB
BC
2039
Dic
keya_d
ianth
icola
_IPO
980
Dic
keya_d
ianth
icola
_NC
PPB
_3534
Dic
keya_d
adanti
i_N
CPPB
_2976
Dic
keya_d
adanti
i_3937_u
id52537
Dic
keya_s
pp_N
CPPB
_3274
Dic
keya_s
pp_M
K7
Dic
keya_d
adanti
i_N
CPPB
_3537
Dic
keya_d
adanti
i_N
CPPB
_898
Dic
keya_z
eae_A
PM
V01
Dic
keya_z
eae_A
JVN
01
Dic
keya_z
eae_A
PW
M01
Dic
keya_d
adanti
i_Ech
586_u
id42519
Dic
keya_z
eae_C
SL_
RW
192
Dic
keya_z
eae_N
CPPB
_3532
Dic
keya_z
eae_N
CPPB
_2538
Dic
keya_z
eae_N
CPPB
_3531
Dic
keya_z
eae_M
K19
Dic
keya_s
pp_N
CPPB
_569
Dic
keya_c
hry
santh
am
i_N
CPPB
_402
Dic
keya_c
hry
santh
am
i_N
CPPB
_516
Dic
keya_z
eae_E
ch1591_u
id59297
Dic
keya_c
hry
santh
am
i_N
CPPB
_3533
Dickeya_dadantii_Ech703_uid59363Dickeya_paradisiaca_NCPPB_2511Dickeya_aquatica_DW_0440Dickeya_aquatica_CSL_RW240Dickeya_dianthicola_GBBC2039Dickeya_dianthicola_NCPPB_3534Dickeya_dianthicola_NCPPB_453Dickeya_dianthicola_IPO980Dickeya_solani_GBBC2040Dickeya_solani_IPO2222Dickeya_solani_MK10Dickeya_solani_MK16Dickeya_solani_AMYI01Dickeya_solani_AMWE01Dickeya_dadantii_NCPPB_3537Dickeya_dadantii_NCPPB_898Dickeya_spp_NCPPB_3274Dickeya_spp_MK7Dickeya_dadantii_NCPPB_2976Dickeya_dadantii_3937_uid52537Dickeya_zeae_APMV01Dickeya_zeae_AJVN01Dickeya_dadantii_Ech586_uid42519Dickeya_zeae_APWM01Dickeya_zeae_MK19Dickeya_zeae_NCPPB_3532Dickeya_zeae_NCPPB_2538Dickeya_zeae_NCPPB_3531Dickeya_zeae_CSL_RW192Dickeya_spp_NCPPB_569Dickeya_zeae_Ech1591_uid59297Dickeya_chrysanthami_NCPPB_3533Dickeya_chrysanthami_NCPPB_402Dickeya_chrysanthami_NCPPB_516
0.00
0.25
0.50
0.75
1.00
AN
Im_a
lignm
ent_
covera
ge
Looking wider
• What about species with high levels of recombination?• “Fuzzy species”: key example, Neisseria:
“The most challenging test of whether species can be clearlydelineated is provided by analysis of large populations ofclosely-related, highly recombinogenic, bacteria that colonise thesame body site”10
10Hanage et al. (2005) BMC Biol. doi:10.1186/1741-7007-3-6
719 Neisseria spp. ANIm
• AbsoluteseparationbetweenN. meningitidis,N. gonorrhoeae,N. lactamica
• Minor complexof possiblemisassign-ments/novelspecies
N. g
onor
rhoe
ae M
U_N
G9
N. g
onor
rhoe
ae N
YC
_201
1_05
_07
N. g
onor
rhoe
ae A
LB_2
011_
04_0
3N
. gon
orrh
oeae
MU
_NG
15N
. gon
orrh
oeae
MU
_NG
14N
. gon
orrh
oeae
MU
_NG
12N
. gon
orrh
oeae
MU
_NG
8N
. gon
orrh
oeae
MU
_NG
19N
. gon
orrh
oeae
MU
_NG
3N
. gon
orrh
oeae
MU
_NG
20N
. gon
orrh
oeae
SK
2302
0N
. gon
orrh
oeae
MIA
_201
1_05
10
N. g
onor
rhoe
ae S
K32
402
N. g
onor
rhoe
ae M
U_N
G25
N. g
onor
rhoe
ae M
IA_2
011_
051
5N
. gon
orrh
oeae
MIA
_201
1_02
_02
N. g
onor
rhoe
ae M
U_N
G4
N. g
onor
rhoe
ae A
LB_2
011_
03_0
3N
. gon
orrh
oeae
NY
C_2
011_
05_1
3N
. gon
orrh
oeae
AT
L_20
11_0
1_05
N. g
onor
rhoe
ae M
U_N
G21
N. g
onor
rhoe
ae M
U_N
G18
N. g
onor
rhoe
ae G
C1
182
N. g
onor
rhoe
ae A
TL_
2011
_01_
17N
. gon
orrh
oeae
CH
811
N. g
onor
rhoe
ae S
K16
942
N. g
onor
rhoe
ae 1
291
N. g
onor
rhoe
ae M
U_N
G17
N. g
onor
rhoe
ae N
OR
_201
1_03
06
N. g
onor
rhoe
ae S
K14
515
N. g
onor
rhoe
ae 0
9_10
_003
N. g
onor
rhoe
ae A
TL_
2011
_01_
03N
. gon
orrh
oeae
MIA
_201
1_03
09
N. g
onor
rhoe
ae M
IA_2
011_
051
6N
. gon
orrh
oeae
SK
7461
N. g
onor
rhoe
ae S
K36
809
N. g
onor
rhoe
ae S
K28
355
N. g
onor
rhoe
ae S
K17
973
N. g
onor
rhoe
ae A
TL_
2011
_01_
08N
. gon
orrh
oeae
MU
_NG
26N
. gon
orrh
oeae
MU
_NG
23N
. gon
orrh
oeae
MU
_NG
1N
. gon
orrh
oeae
GC
GS
006
N. g
onor
rhoe
ae G
CG
S14
8N
. gon
orrh
oeae
GC
GS
080
N. g
onor
rhoe
ae G
CG
S05
4N
. gon
orrh
oeae
GC
GS
196
N. g
onor
rhoe
ae G
CG
S14
0N
. gon
orrh
oeae
GC
GS
122
N. g
onor
rhoe
ae G
CG
S03
6N
. gon
orrh
oeae
GC
GS
222
N. g
onor
rhoe
ae G
CG
S00
4N
. gon
orrh
oeae
GC
GS
236
N. g
onor
rhoe
ae G
CG
S23
4N
. gon
orrh
oeae
GC
GS
099
N. g
onor
rhoe
ae N
CC
P11
945
N. g
onor
rhoe
ae G
CG
S23
2N
. gon
orrh
oeae
GC
GS
058
N. g
onor
rhoe
ae G
CG
S09
4N
. gon
orrh
oeae
GC
GS
238
N. g
onor
rhoe
ae A
LB_2
011_
010
2N
. gon
orrh
oeae
GC
GS
158
N. g
onor
rhoe
ae S
K9
267
9N
. gon
orrh
oeae
35/
02N
. gon
orrh
oeae
SK
1268
4N
. gon
orrh
oeae
SK
2947
1N
. gon
orrh
oeae
AT
L_20
11_0
513
N. g
onor
rhoe
ae S
K16
259
N. g
onor
rhoe
ae G
CG
S11
4N
. gon
orrh
oeae
GC
GS
066
N. g
onor
rhoe
ae G
CG
S16
0N
. gon
orrh
oeae
GC
GS
023
N. g
onor
rhoe
ae G
CG
S02
8N
. gon
orrh
oeae
GC
GS
119
N. g
onor
rhoe
ae G
CG
S07
6N
. gon
orrh
oeae
GC
GS
030
N. g
onor
rhoe
ae G
CG
S08
5N
. gon
orrh
oeae
GC
GS
166
N. g
onor
rhoe
ae G
CG
S15
7N
. gon
orrh
oeae
GC
GS
155
N. g
onor
rhoe
ae G
CG
S00
5N
. gon
orrh
oeae
GC
GS
235
N. g
onor
rhoe
ae G
CG
S12
4N
. gon
orrh
oeae
GC
GS
068
N. g
onor
rhoe
ae G
CG
S24
1N
. gon
orrh
oeae
GC
GS
147
N. g
onor
rhoe
ae G
CG
S14
5N
. gon
orrh
oeae
GC
GS
048
N. g
onor
rhoe
ae G
CG
S22
9N
. gon
orrh
oeae
GC
GS
075
N. g
onor
rhoe
ae G
CG
S11
7N
. gon
orrh
oeae
GC
GS
063
N. g
onor
rhoe
ae G
CG
S17
8N
. gon
orrh
oeae
GC
GS
179
N. g
onor
rhoe
ae G
CG
S08
1N
. gon
orrh
oeae
GC
GS
041
N. g
onor
rhoe
ae G
CG
S04
3N
. gon
orrh
oeae
GC
GS
061
N. g
onor
rhoe
ae G
CG
S13
1N
. gon
orrh
oeae
GC
GS
089
N. g
onor
rhoe
ae G
CG
S06
9N
. gon
orrh
oeae
GC
GS
189
N. g
onor
rhoe
ae G
CG
S05
7N
. gon
orrh
oeae
GC
GS
125
N. g
onor
rhoe
ae G
CG
S12
3N
. gon
orrh
oeae
GC
GS
001
N. g
onor
rhoe
ae G
CG
S10
4N
. gon
orrh
oeae
GC
GS
107
N. g
onor
rhoe
ae G
CG
S02
7N
. gon
orrh
oeae
GC
GS
129
N. g
onor
rhoe
ae G
CG
S04
9N
. gon
orrh
oeae
GC
GS
233
N. g
onor
rhoe
ae G
CG
S06
5N
. gon
orrh
oeae
GC
GS
037
N. g
onor
rhoe
ae G
CG
S18
5N
. gon
orrh
oeae
GC
GS
017
N. g
onor
rhoe
ae G
CG
S15
1N
. gon
orrh
oeae
GC
GS
159
N. g
onor
rhoe
ae G
CG
S03
5N
. gon
orrh
oeae
GC
GS
091
N. g
onor
rhoe
ae G
CG
S05
9N
. gon
orrh
oeae
GC
GS
109
N. g
onor
rhoe
ae G
CG
S16
5N
. gon
orrh
oeae
GC
GS
139
N. g
onor
rhoe
ae G
CG
S20
5N
. gon
orrh
oeae
GC
GS
053
N. g
onor
rhoe
ae G
CG
S19
5N
. gon
orrh
oeae
GC
GS
133
N. g
onor
rhoe
ae G
CG
S06
7N
. gon
orrh
oeae
GC
GS
055
N. g
onor
rhoe
ae G
CG
S17
7N
. gon
orrh
oeae
G28
91N
. gon
orrh
oeae
AT
L_20
11_0
125
N. g
onor
rhoe
ae M
U_N
G5
N. g
onor
rhoe
ae m
07.0
5N
. gon
orrh
oeae
GC
GS
046
N. g
onor
rhoe
ae G
CG
S10
5N
. gon
orrh
oeae
GC
GS
097
N. g
onor
rhoe
ae G
CG
S04
7N
. gon
orrh
oeae
GC
GS
237
N. g
onor
rhoe
ae G
CG
S08
8N
. gon
orrh
oeae
GC
GS
137
N. g
onor
rhoe
ae G
CG
S00
7N
. gon
orrh
oeae
GC
GS
021
N. g
onor
rhoe
ae G
CG
S10
1N
. gon
orrh
oeae
GC
GS
093
N. g
onor
rhoe
ae G
CG
S12
1N
. gon
orrh
oeae
GC
GS
135
N. g
onor
rhoe
ae G
CG
S03
3N
. gon
orrh
oeae
GC
GS
079
N. g
onor
rhoe
ae G
CG
S11
5N
. gon
orrh
oeae
GC
GS
019
N. g
onor
rhoe
ae G
CG
S11
3N
. gon
orrh
oeae
GC
GS
103
N. g
onor
rhoe
ae G
CG
S10
2N
. gon
orrh
oeae
GC
GS
025
N. g
onor
rhoe
ae G
CG
S23
1N
. gon
orrh
oeae
GC
GS
183
N. g
onor
rhoe
ae G
CG
S03
9N
. gon
orrh
oeae
GC
GS
143
N. g
onor
rhoe
ae G
CG
S04
5N
. gon
orrh
oeae
GC
GS
029
N. g
onor
rhoe
ae G
CG
S07
7N
. gon
orrh
oeae
GC
GS
031
N. g
onor
rhoe
ae G
CG
S05
1N
. gon
orrh
oeae
GC
GS
181
N. g
onor
rhoe
ae G
CG
S09
5N
. gon
orrh
oeae
GC
GS
227
N. g
onor
rhoe
ae G
CG
S12
7N
. gon
orrh
oeae
GC
GS
242
N. g
onor
rhoe
ae G
CG
S23
9N
. gon
orrh
oeae
GC
GS
187
N. g
onor
rhoe
ae G
CG
S16
3N
. gon
orrh
oeae
GC
GS
141
N. g
onor
rhoe
ae G
CG
S08
7N
. gon
orrh
oeae
GC
GS
003
N. g
onor
rhoe
ae F
A19
N. g
onor
rhoe
ae G
CG
S00
8N
. gon
orrh
oeae
GC
GS
146
N. g
onor
rhoe
ae G
CG
S18
6N
. gon
orrh
oeae
GC
GS
188
N. g
onor
rhoe
ae G
CG
S09
0N
. gon
orrh
oeae
MS
11N
. gon
orrh
oeae
GC
GS
210
N. g
onor
rhoe
ae G
CG
S21
2N
. gon
orrh
oeae
GC
GS
200
N. g
onor
rhoe
ae N
G05
N. g
onor
rhoe
ae n
01.0
8N
. gon
orrh
oeae
GC
GS
050
N. g
onor
rhoe
ae G
CG
S15
4N
. gon
orrh
oeae
GC
GS
072
N. g
onor
rhoe
ae F
62N
. gon
orrh
oeae
GC
GS
184
N. g
onor
rhoe
ae F
A 1
090
N. g
onor
rhoe
ae F
A61
40N
. gon
orrh
oeae
FA
19N
. gon
orrh
oeae
GC
GS
162
N. g
onor
rhoe
ae G
CG
S24
0N
. gon
orrh
oeae
GC
GS
190
N. g
onor
rhoe
ae G
CG
S17
0N
. gon
orrh
oeae
GC
GS
056
N. g
onor
rhoe
ae G
CG
S13
4N
. gon
orrh
oeae
GC
GS
078
N. g
onor
rhoe
ae G
CG
S16
8N
. gon
orrh
oeae
GC
GS
206
N. g
onor
rhoe
ae G
CG
S11
6N
. gon
orrh
oeae
GC
GS
044
N. g
onor
rhoe
ae G
CG
S03
2N
. gon
orrh
oeae
GC
GS
022
N. g
onor
rhoe
ae G
CG
S02
6N
. gon
orrh
oeae
GC
GS
060
N. g
onor
rhoe
ae G
CG
S18
0N
. gon
orrh
oeae
GC
GS
216
N. g
onor
rhoe
ae e
03.0
4N
. gon
orrh
oeae
GC
GS
208
N. g
onor
rhoe
ae A
TL_
2011
_05
08N
. gon
orrh
oeae
SK
3341
4N
. gon
orrh
oeae
MIA
_201
1_03
10
N. g
onor
rhoe
ae i1
9.05
N. g
onor
rhoe
ae G
CG
S15
0N
. gon
orrh
oeae
GC
GS
142
N. g
onor
rhoe
ae G
CG
S01
0N
. gon
orrh
oeae
GC
GS
144
N. g
onor
rhoe
ae S
K39
420
N. g
onor
rhoe
ae F
A19
N. g
onor
rhoe
ae M
U_N
G6
N. g
onor
rhoe
ae S
K9
310
35N
. gon
orrh
oeae
DG
I18
N. g
onor
rhoe
ae S
K22
871
N. g
onor
rhoe
ae G
CG
S17
6N
. gon
orrh
oeae
GC
GS
018
N. g
onor
rhoe
ae G
CG
S19
8N
. gon
orrh
oeae
GC
GS
202
N. g
onor
rhoe
ae G
CG
S22
8N
. gon
orrh
oeae
GC
GS
204
N. g
onor
rhoe
ae G
CG
S12
0N
. gon
orrh
oeae
GC
GS
138
N. g
onor
rhoe
ae G
CG
S19
2N
. gon
orrh
oeae
GC
GS
062
N. g
onor
rhoe
ae G
CG
S08
3N
. gon
orrh
oeae
GC
GS
084
N. g
onor
rhoe
ae G
CG
S00
9N
. gon
orrh
oeae
GC
GS
106
N. g
onor
rhoe
ae G
CG
S09
2N
. gon
orrh
oeae
GC
GS
098
N. g
onor
rhoe
ae G
CG
S21
8N
. gon
orrh
oeae
GC
GS
110
N. g
onor
rhoe
ae G
CG
S09
6N
. gon
orrh
oeae
GC
GS
118
N. g
onor
rhoe
ae G
CG
S17
4N
. gon
orrh
oeae
GC
GS
224
N. g
onor
rhoe
ae G
CG
S13
6N
. gon
orrh
oeae
GC
GS
156
N. g
onor
rhoe
ae G
CG
S12
8N
. gon
orrh
oeae
GC
GS
226
N. g
onor
rhoe
ae G
CG
S06
4N
. gon
orrh
oeae
GC
GS
172
N. g
onor
rhoe
ae G
CG
S01
2N
. gon
orrh
oeae
GC
GS
100
N. g
onor
rhoe
ae G
CG
S16
4N
. gon
orrh
oeae
GC
GS
040
N. g
onor
rhoe
ae G
CG
S02
4N
. gon
orrh
oeae
GC
GS
082
N. g
onor
rhoe
ae S
K29
344
N. g
onor
rhoe
ae S
K69
87N
. gon
orrh
oeae
SK
1545
4N
. gon
orrh
oeae
SK
7842
N. g
onor
rhoe
ae N
G_8
69N
. gon
orrh
oeae
FA
6140
N. g
onor
rhoe
ae P
ID33
2N
. gon
orrh
oeae
DG
I2N
. gon
orrh
oeae
SK
8976
N. g
onor
rhoe
ae S
K70
8N
. gon
orrh
oeae
GC
GS
220
N. g
onor
rhoe
ae G
CG
S18
2N
. gon
orrh
oeae
GC
GS
214
N. g
onor
rhoe
ae G
CG
S08
6N
. gon
orrh
oeae
GC
GS
002
N. g
onor
rhoe
ae G
CG
S02
0N
. gon
orrh
oeae
GC
GS
152
N. g
onor
rhoe
ae G
CG
S05
2N
. gon
orrh
oeae
GC
GS
132
N. g
onor
rhoe
ae G
CG
S13
0N
. gon
orrh
oeae
GC
GS
042
N. g
onor
rhoe
ae G
CG
S19
4N
. gon
orrh
oeae
GC
GS
108
N. g
onor
rhoe
ae G
CG
S07
0N
. gon
orrh
oeae
GC
GS
038
N. g
onor
rhoe
ae G
CG
S23
0N
. gon
orrh
oeae
GC
GS
126
N. g
onor
rhoe
ae G
CG
S17
5N
. gon
orrh
oeae
GC
GS
073
N. g
onor
rhoe
ae G
CG
S21
3N
. gon
orrh
oeae
GC
GS
171
N. g
onor
rhoe
ae G
CG
S20
9N
. gon
orrh
oeae
GC
GS
169
N. g
onor
rhoe
ae G
CG
S20
3N
. gon
orrh
oeae
GC
GS
201
N. g
onor
rhoe
ae G
CG
S07
1N
. gon
orrh
oeae
GC
GS
173
N. g
onor
rhoe
ae G
CG
S19
7N
. gon
orrh
oeae
GC
GS
167
N. g
onor
rhoe
ae G
CG
S21
9N
. gon
orrh
oeae
GC
GS
153
N. g
onor
rhoe
ae G
CG
S21
1N
. gon
orrh
oeae
GC
GS
221
N. g
onor
rhoe
ae G
CG
S14
9N
. gon
orrh
oeae
GC
GS
161
N. g
onor
rhoe
ae G
CG
S19
9N
. gon
orrh
oeae
GC
GS
207
N. g
onor
rhoe
ae G
CG
S19
1N
. gon
orrh
oeae
GC
GS
217
N. g
onor
rhoe
ae G
CG
S19
3N
. gon
orrh
oeae
GC
GS
215
N. g
onor
rhoe
ae G
CG
S22
3N
. gon
orrh
oeae
GC
GS
225
N. g
onor
rhoe
ae G
CG
S03
4N
. gon
orrh
oeae
GC
GS
074
N. g
onor
rhoe
ae 3
5/02
N. g
onor
rhoe
ae G
CG
S01
1N
. gon
orrh
oeae
PID
241
N. g
onor
rhoe
ae P
ID1
N. g
onor
rhoe
ae M
S11
N. g
onor
rhoe
ae S
K19
02N
. gon
orrh
oeae
AT
L_20
11_0
121
N. g
onor
rhoe
ae P
ID18
N. m
enin
gitid
is Z
2491
N. m
enin
gitid
is 6
5014
N. m
enin
gitid
is 6
3006
N. m
enin
gitid
is 6
3041
N. m
enin
gitid
is 6
4182
N. m
enin
gitid
is 9
7027
N. m
enin
gitid
is 6
3049
N. m
enin
gitid
is 6
5012
N. m
enin
gitid
is 9
7018
N. m
enin
gitid
is 9
4018
N. m
enin
gitid
is 9
6024
N. m
enin
gitid
is 9
8005
N. m
enin
gitid
is 7
5689
N. m
enin
gitid
is 9
7008
N. m
enin
gitid
is 7
5643
N. m
enin
gitid
is 8
8050
N. m
enin
gitid
is 9
7020
N. m
enin
gitid
is 9
6023
N. m
enin
gitid
is W
UE
259
4N
. men
ingi
tidis
NM
2254
N. m
enin
gitid
is N
M25
24N
. men
ingi
tidis
200
0080
N. m
enin
gitid
is N
M28
10N
. men
ingi
tidis
NM
1364
N. m
enin
gitid
is N
M18
26N
. men
ingi
tidis
NM
1758
N. m
enin
gitid
is N
M23
94N
. men
ingi
tidis
NM
1893
N. m
enin
gitid
is N
M15
49N
. men
ingi
tidis
NM
1976
N. m
enin
gitid
is N
M15
50N
. men
ingi
tidis
NM
2393
N. m
enin
gitid
is N
M27
17N
. men
ingi
tidis
NM
2382
N. m
enin
gitid
is N
M28
14N
. men
ingi
tidis
NM
2606
N. m
enin
gitid
is N
M24
41N
. men
ingi
tidis
200
3022
N. m
enin
gitid
is N
M60
7N
. men
ingi
tidis
200
4085
N. m
enin
gitid
is N
M14
71N
. men
ingi
tidis
NM
2033
N. m
enin
gitid
is 2
0030
22N
. men
ingi
tidis
NM
2933
N. m
enin
gitid
is N
M22
37N
. men
ingi
tidis
NM
2809
N. m
enin
gitid
is 2
0020
07N
. men
ingi
tidis
NM
1829
N. m
enin
gitid
is N
M22
63N
. men
ingi
tidis
NM
1901
N. m
enin
gitid
is N
M18
05N
. men
ingi
tidis
NM
1360
N. m
enin
gitid
is N
M14
82N
. men
ingi
tidis
NM
1919
N. m
enin
gitid
is N
M26
17N
. men
ingi
tidis
NM
2187
N. m
enin
gitid
is N
M17
57N
. men
ingi
tidis
NM
2008
N. m
enin
gitid
is N
M13
61N
. men
ingi
tidis
NM
2813
N. m
enin
gitid
is N
M15
61N
. men
ingi
tidis
NM
1910
N. m
enin
gitid
is N
M60
6N
. men
ingi
tidis
200
0063
N. m
enin
gitid
is N
M18
91N
. men
ingi
tidis
NM
1931
N. m
enin
gitid
is N
M19
37N
. men
ingi
tidis
NM
1938
N. m
enin
gitid
is N
M13
59N
. men
ingi
tidis
NM
1264
N. m
enin
gitid
is N
M22
39N
. men
ingi
tidis
NM
1362
N. m
enin
gitid
is N
M28
12N
. men
ingi
tidis
NM
2431
N. m
enin
gitid
is N
M29
35N
. men
ingi
tidis
NM
2439
N. m
enin
gitid
is N
M22
32N
. men
ingi
tidis
NM
2193
N. m
enin
gitid
is N
M23
32N
. men
ingi
tidis
NM
2228
N. m
enin
gitid
is N
M22
64N
. men
ingi
tidis
NM
1895
N. m
enin
gitid
is N
M27
18N
. men
ingi
tidis
NM
2188
N. m
enin
gitid
is N
M21
81N
. men
ingi
tidis
NM
2369
N. m
enin
gitid
is N
M23
89N
. men
ingi
tidis
NM
2811
N. m
enin
gitid
is N
M24
32N
. men
ingi
tidis
NM
2335
N. m
enin
gitid
is N
M22
44N
. men
ingi
tidis
NM
1928
N. m
enin
gitid
is N
M28
08N
. men
ingi
tidis
NM
2381
N. m
enin
gitid
is N
M27
00N
. men
ingi
tidis
NM
2932
N. m
enin
gitid
is N
M14
83N
. men
ingi
tidis
NM
604
N. m
enin
gitid
is 5
1061
2N
. men
ingi
tidis
NM
1673
N. m
enin
gitid
is N
M20
32N
. men
ingi
tidis
NM
1666
N. m
enin
gitid
is N
M15
78N
. men
ingi
tidis
NM
1963
N. m
enin
gitid
is N
M17
97N
. men
ingi
tidis
NM
1837
N. m
enin
gitid
is N
M31
29N
. men
ingi
tidis
NM
2602
N. m
enin
gitid
is N
M28
57N
. men
ingi
tidis
NM
2206
N. m
enin
gitid
is N
M15
44N
. men
ingi
tidis
NM
1921
N. m
enin
gitid
is N
M18
45N
. men
ingi
tidis
NM
2856
N. m
enin
gitid
is N
M17
79N
. men
ingi
tidis
NM
2934
N. m
enin
gitid
is N
M18
92N
. men
ingi
tidis
NM
1325
N. m
enin
gitid
is N
M15
73N
. men
ingi
tidis
NM
3128
N. m
enin
gitid
is N
M27
01N
. men
ingi
tidis
NM
2009
N. m
enin
gitid
is N
M18
31N
. men
ingi
tidis
NM
1446
N. m
enin
gitid
is N
M20
25N
. men
ingi
tidis
NM
1363
N. m
enin
gitid
is N
M16
72N
. men
ingi
tidis
NM
2433
N. m
enin
gitid
is 2
0070
56N
. men
ingi
tidis
NM
3652
N. m
enin
gitid
is 2
0040
90N
. men
ingi
tidis
200
1212
N. m
enin
gitid
is N
M36
42N
. men
ingi
tidis
Nm
1140
N. m
enin
gitid
is a
lpha
704
N. m
enin
gitid
is N
M27
95N
. men
ingi
tidis
LN
P27
256
N. m
enin
gitid
isN
. men
ingi
tidis
NM
586
N. m
enin
gitid
is M
6190
N. m
enin
gitid
is S
0108
N. m
enin
gitid
isN
. men
ingi
tidis
ES
1490
2N
. men
ingi
tidis
NM
1482
N. m
enin
gitid
is N
M95
N. m
enin
gitid
is N
M13
3N
. men
ingi
tidis
NM
1482
N. m
enin
gitid
is M
2059
9N
. men
ingi
tidis
K12
07N
. men
ingi
tidis
S01
08N
. men
ingi
tidis
NM
94N
. men
ingi
tidis
NM
82N
. men
ingi
tidis
NM
3680
N. m
enin
gitid
is N
M36
81N
. men
ingi
tidis
NM
3683
N. m
enin
gitid
is N
M36
82N
. men
ingi
tidis
M71
24N
. men
ingi
tidis
200
5040
N. m
enin
gitid
is M
1020
8N
. men
ingi
tidis
NM
3686
N. m
enin
gitid
is N
M36
85N
. men
ingi
tidis
M70
89N
. men
ingi
tidis
M71
24N
. men
ingi
tidis
NM
174
N. m
enin
gitid
is N
M36
87N
. men
ingi
tidis
NM
3684
N. m
enin
gitid
is N
M36
88N
. men
ingi
tidis
NM
126
N. m
enin
gitid
is F
AM
18N
. men
ingi
tidis
NM
762
N. m
enin
gitid
is L
9154
3N
. men
ingi
tidis
M12
611
N. m
enin
gitid
is 2
0012
13N
. men
ingi
tidis
NM
43N
. men
ingi
tidis
M14
12N
. men
ingi
tidis
200
1073
N. m
enin
gitid
is 2
0010
68N
. men
ingi
tidis
NM
32N
. men
ingi
tidis
NM
23N
. men
ingi
tidis
NM
35N
. men
ingi
tidis
NM
36N
. men
ingi
tidis
737
04N
. men
ingi
tidis
200
0175
N. m
enin
gitid
is 2
0020
04N
. men
ingi
tidis
200
4264
N. m
enin
gitid
is 2
0000
81N
. men
ingi
tidis
200
5079
N. m
enin
gitid
is 2
0010
72N
. men
ingi
tidis
NM
1495
N. m
enin
gitid
is N
M31
3N
. men
ingi
tidis
NM
3147
N. m
enin
gitid
is 7
0012
N. m
enin
gitid
is 7
0030
N. m
enin
gitid
is 6
9166
N. m
enin
gitid
is 6
1103
N. m
enin
gitid
is 6
8094
N. m
enin
gitid
is 6
9096
N. m
enin
gitid
is 6
3023
N. m
enin
gitid
is 6
1106
N. m
enin
gitid
is 7
0021
N. m
enin
gitid
is 6
9155
N. m
enin
gitid
is 7
0082
N. m
enin
gitid
is 6
9100
N. m
enin
gitid
is 9
6060
N. m
enin
gitid
is 6
9176
N. m
enin
gitid
is N
M47
7N
. men
ingi
tidis
NM
3141
N. m
enin
gitid
is M
1326
5N
. men
ingi
tidis
NM
3173
N. m
enin
gitid
is 2
0020
20N
. men
ingi
tidis
200
2030
N. m
enin
gitid
is N
M14
76N
. men
ingi
tidis
H44
/76
N. m
enin
gitid
is H
44/7
6N
. men
ingi
tidis
MC
58N
. men
ingi
tidis
LN
P21
362
N. m
enin
gitid
is N
M41
8N
. men
ingi
tidis
M13
255
N. m
enin
gitid
is 9
506
N. m
enin
gitid
is C
U38
5N
. men
ingi
tidis
128
88N
. men
ingi
tidis
411
9N
. men
ingi
tidis
NM
422
N. m
enin
gitid
is 9
757
N. m
enin
gitid
is 3
2050
3N
. men
ingi
tidis
100
514
N. m
enin
gitid
is 3
7060
1N
. men
ingi
tidis
100
572
N. m
enin
gitid
is 3
6062
4N
. men
ingi
tidis
421
007
N. m
enin
gitid
is 4
4050
1N
. men
ingi
tidis
340
552
N. m
enin
gitid
is 1
0053
0N
. men
ingi
tidis
321
114
N. m
enin
gitid
is 3
4173
N. m
enin
gitid
is 3
1111
2N
. men
ingi
tidis
053
442
N. m
enin
gitid
is 1
0060
1N
. men
ingi
tidis
Nm
1100
3N
. men
ingi
tidis
341
215
N. m
enin
gitid
is 3
2050
1N
. men
ingi
tidis
220
601
N. m
enin
gitid
is 4
4090
2N
. men
ingi
tidis
130
803
N. m
enin
gitid
is 1
3114
8N
. men
ingi
tidis
420
718
N. m
enin
gitid
is 1
0060
3N
. men
ingi
tidis
330
505
N. m
enin
gitid
is N
m27
32N
. men
ingi
tidis
818
58N
. men
ingi
tidis
801
3N
. men
ingi
tidis
Nm
1025
9N
. men
ingi
tidis
Nm
9418
N. m
enin
gitid
is N
m69
38N
. men
ingi
tidis
920
45N
. men
ingi
tidis
NM
255
N. m
enin
gitid
is 9
8002
N. m
enin
gitid
is 2
0030
51N
. men
ingi
tidis
736
96N
. men
ingi
tidis
NM
134
N. m
enin
gitid
is N
m81
87N
. men
ingi
tidis
Nm
3127
N. m
enin
gitid
is 2
0010
01N
. men
ingi
tidis
200
7461
N. m
enin
gitid
is 2
0040
32N
. men
ingi
tidis
992
008
N. m
enin
gitid
is a
lpha
710
N. m
enin
gitid
is O
X99
.303
04N
. men
ingi
tidis
NM
151
N. m
enin
gitid
is N
Z0
5/33
N. m
enin
gitid
is D
E96
86N
. men
ingi
tidis
DE
9622
N. m
enin
gitid
is D
E99
38N
. men
ingi
tidis
M05
79N
. men
ingi
tidis
M01
240
149
N. m
enin
gitid
is M
0579
N. m
enin
gitid
is N
M26
57N
. men
ingi
tidis
NM
576
N. m
enin
gitid
is N
M27
81N
. men
ingi
tidis
NM
140
N. m
enin
gitid
is N
M18
3N
. men
ingi
tidis
Nei
sser
ia m
enin
gitid
is C
HU
VN
. men
ingi
tidis
M13
399
N. m
enin
gitid
is M
012
4001
3N
. men
ingi
tidis
M04
240
196
N. m
enin
gitid
is 7
7221
N. m
enin
gitid
is a
lpha
14N
. men
ingi
tidis
NM
3001
N. m
enin
gitid
is M
012
4035
5N
. men
ingi
tidis
930
03N
. men
ingi
tidis
930
04N
. men
ingi
tidis
NM
3081
N. m
enin
gitid
is 9
8008
N. m
enin
gitid
is 8
0179
N. m
enin
gitid
is A
TC
C 1
3091
N. m
enin
gitid
is N
M04
5N
. men
ingi
tidis
NM
2866
N. m
enin
gitid
is N
M00
3N
. men
ingi
tidis
NM
3139
N. m
enin
gitid
is N
M05
52N
. men
ingi
tidis
NM
518
N. m
enin
gitid
is N
M32
30N
. men
ingi
tidis
960
37N
. men
ingi
tidis
N15
68N
. men
ingi
tidis
200
2038
N. m
enin
gitid
is 9
7021
N. m
enin
gitid
is 2
0060
87N
. men
ingi
tidis
970
14N
. men
ingi
tidis
200
8223
N. m
enin
gitid
is 2
0051
72N
. men
ingi
tidis
NM
220
N. m
enin
gitid
is N
M27
1N
. men
ingi
tidis
NM
115
N. m
enin
gitid
is N
M31
44N
. men
ingi
tidis
NM
27N
. men
ingi
tidis
NM
3131
N. m
enin
gitid
is N
M31
58N
. men
ingi
tidis
NM
3222
N. m
enin
gitid
is N
M31
64N
. men
ingi
tidis
Nm
6756
N. m
enin
gitid
is N
m86
63N
. men
ingi
tidis
NM
27N
. men
ingi
tidis
NM
90N
. men
ingi
tidis
NM
233
N. m
enin
gitid
is N
S44
N. m
enin
gitid
is N
M80
N. m
enin
gitid
is N
M16
5N
. men
ingi
tidis
NM
51N
. men
ingi
tidis
NM
3042
N. m
enin
gitid
is N
M32
23N
. men
ingi
tidis
B61
16/7
7N
. men
ingi
tidis
G21
36N
. men
ingi
tidis
980
80N
. men
ingi
tidis
872
55N
. men
ingi
tidis
NM
BN
. men
ingi
tidis
961
594
5N
. pol
ysac
char
ea A
TC
C 4
3768
N. p
olys
acch
area
NS
342
N. l
acta
mic
a N
S19
N. l
acta
mic
a 02
006
N. l
acta
mic
a Y
921
009
N. l
acta
mic
a A
TC
C 2
3970
N. l
acta
mic
a A
TC
C 2
3970
N. c
iner
ea A
TC
C 1
4685
N. f
lave
scen
s N
RL3
0031
/H21
0N
. sic
ca V
K64
N. s
icca
AT
CC
292
56N
. men
ingi
tidis
433
_NM
EN
N. m
enin
gitid
is 1
044_
NM
EN
N. m
enin
gitid
is 8
40_N
ME
NN
. men
ingi
tidis
776
_NM
EN
N. s
ubfla
va N
J970
3N
. men
ingi
tidis
104
5_N
ME
NN
. men
ingi
tidis
768
_NM
EN
N. m
ucos
a C
6AN
. lac
tam
ica
595_
NLA
CN
. lac
tam
ica
583_
NLA
CN
. lac
tam
ica
338.
rep1
_NLA
CN
. men
ingi
tidis
338
.rep
2_N
ME
NN
. fla
vesc
ens
SK
114
N. m
enin
gitid
is 7
82_N
ME
NN
. men
ingi
tidis
121
0_N
ME
NN
. muc
osa
C10
2N
. men
ingi
tidis
127
9_N
ME
NN
. elo
ngat
a A
TC
C 2
9315
N. e
long
ata
AT
CC
293
15N
. sp.
ora
l tax
on 0
14 F
0314
N. m
enin
gitid
is 4
31_N
ME
NN
. men
ingi
tidis
404
_NM
EN
N. l
acta
mic
a 91
9_N
LAC
N. m
enin
gitid
is 3
13_N
ME
NN
. men
ingi
tidis
379
_NM
EN
N. m
enin
gitid
is 3
78_N
ME
NN
. sic
ca D
S1
N. s
p. G
T4A
_CT
1 G
T4A
_CT
1N
. mac
acae
AT
CC
339
26N
. men
ingi
tidis
127
3_N
ME
NN
. sp.
HM
SC
06F
02 H
MS
C06
F02
N. m
ucos
a A
TC
C 2
5996
N. m
enin
gitid
is 4
80_N
ME
NN
. sic
ca 4
320
N. s
p. 8
3E34
83E
34N
. sp.
74A
18 7
4A18
N. w
adsw
orth
ii 97
15N
. sp.
KH
1503
KH
1503
N. s
haye
gani
i 871
N. w
eave
ri A
TC
C 5
1223
N. w
eave
ri LM
G 5
135
N. m
enin
gitid
is 4
91_N
ME
NN
. sp.
ora
l tax
on 0
20 F
0370
N. m
enin
gitid
is 2
03_N
ME
NN
. lac
tam
ica
914_
NLA
CN
. bac
illifo
rmis
AT
CC
BA
A1
200
N. gonorrhoeae MU_NG9N. gonorrhoeae NYC_2011_05_07N. gonorrhoeae ALB_2011_04_03N. gonorrhoeae MU_NG15N. gonorrhoeae MU_NG14N. gonorrhoeae MU_NG12N. gonorrhoeae MU_NG8N. gonorrhoeae MU_NG19N. gonorrhoeae MU_NG3N. gonorrhoeae MU_NG20N. gonorrhoeae SK23020N. gonorrhoeae MIA_2011_0510N. gonorrhoeae SK32402N. gonorrhoeae MU_NG25N. gonorrhoeae MIA_2011_0515N. gonorrhoeae MIA_2011_02_02N. gonorrhoeae MU_NG4N. gonorrhoeae ALB_2011_03_03N. gonorrhoeae NYC_2011_05_13N. gonorrhoeae ATL_2011_01_05N. gonorrhoeae MU_NG21N. gonorrhoeae MU_NG18N. gonorrhoeae GC1182N. gonorrhoeae ATL_2011_01_17N. gonorrhoeae CH811N. gonorrhoeae SK16942N. gonorrhoeae 1291N. gonorrhoeae MU_NG17N. gonorrhoeae NOR_2011_0306N. gonorrhoeae SK14515N. gonorrhoeae 09_10_003N. gonorrhoeae ATL_2011_01_03N. gonorrhoeae MIA_2011_0309N. gonorrhoeae MIA_2011_0516N. gonorrhoeae SK7461N. gonorrhoeae SK36809N. gonorrhoeae SK28355N. gonorrhoeae SK17973N. gonorrhoeae ATL_2011_01_08N. gonorrhoeae MU_NG26N. gonorrhoeae MU_NG23N. gonorrhoeae MU_NG1N. gonorrhoeae GCGS006N. gonorrhoeae GCGS148N. gonorrhoeae GCGS080N. gonorrhoeae GCGS054N. gonorrhoeae GCGS196N. gonorrhoeae GCGS140N. gonorrhoeae GCGS122N. gonorrhoeae GCGS036N. gonorrhoeae GCGS222N. gonorrhoeae GCGS004N. gonorrhoeae GCGS236N. gonorrhoeae GCGS234N. gonorrhoeae GCGS099N. gonorrhoeae NCCP11945N. gonorrhoeae GCGS232N. gonorrhoeae GCGS058N. gonorrhoeae GCGS094N. gonorrhoeae GCGS238N. gonorrhoeae ALB_2011_0102N. gonorrhoeae GCGS158N. gonorrhoeae SK92679N. gonorrhoeae 35/02N. gonorrhoeae SK12684N. gonorrhoeae SK29471N. gonorrhoeae ATL_2011_0513N. gonorrhoeae SK16259N. gonorrhoeae GCGS114N. gonorrhoeae GCGS066N. gonorrhoeae GCGS160N. gonorrhoeae GCGS023N. gonorrhoeae GCGS028N. gonorrhoeae GCGS119N. gonorrhoeae GCGS076N. gonorrhoeae GCGS030N. gonorrhoeae GCGS085N. gonorrhoeae GCGS166N. gonorrhoeae GCGS157N. gonorrhoeae GCGS155N. gonorrhoeae GCGS005N. gonorrhoeae GCGS235N. gonorrhoeae GCGS124N. gonorrhoeae GCGS068N. gonorrhoeae GCGS241N. gonorrhoeae GCGS147N. gonorrhoeae GCGS145N. gonorrhoeae GCGS048N. gonorrhoeae GCGS229N. gonorrhoeae GCGS075N. gonorrhoeae GCGS117N. gonorrhoeae GCGS063N. gonorrhoeae GCGS178N. gonorrhoeae GCGS179N. gonorrhoeae GCGS081N. gonorrhoeae GCGS041N. gonorrhoeae GCGS043N. gonorrhoeae GCGS061N. gonorrhoeae GCGS131N. gonorrhoeae GCGS089N. gonorrhoeae GCGS069N. gonorrhoeae GCGS189N. gonorrhoeae GCGS057N. gonorrhoeae GCGS125N. gonorrhoeae GCGS123N. gonorrhoeae GCGS001N. gonorrhoeae GCGS104N. gonorrhoeae GCGS107N. gonorrhoeae GCGS027N. gonorrhoeae GCGS129N. gonorrhoeae GCGS049N. gonorrhoeae GCGS233N. gonorrhoeae GCGS065N. gonorrhoeae GCGS037N. gonorrhoeae GCGS185N. gonorrhoeae GCGS017N. gonorrhoeae GCGS151N. gonorrhoeae GCGS159N. gonorrhoeae GCGS035N. gonorrhoeae GCGS091N. gonorrhoeae GCGS059N. gonorrhoeae GCGS109N. gonorrhoeae GCGS165N. gonorrhoeae GCGS139N. gonorrhoeae GCGS205N. gonorrhoeae GCGS053N. gonorrhoeae GCGS195N. gonorrhoeae GCGS133N. gonorrhoeae GCGS067N. gonorrhoeae GCGS055N. gonorrhoeae GCGS177N. gonorrhoeae G2891N. gonorrhoeae ATL_2011_0125N. gonorrhoeae MU_NG5N. gonorrhoeae m07.05N. gonorrhoeae GCGS046N. gonorrhoeae GCGS105N. gonorrhoeae GCGS097N. gonorrhoeae GCGS047N. gonorrhoeae GCGS237N. gonorrhoeae GCGS088N. gonorrhoeae GCGS137N. gonorrhoeae GCGS007N. gonorrhoeae GCGS021N. gonorrhoeae GCGS101N. gonorrhoeae GCGS093N. gonorrhoeae GCGS121N. gonorrhoeae GCGS135N. gonorrhoeae GCGS033N. gonorrhoeae GCGS079N. gonorrhoeae GCGS115N. gonorrhoeae GCGS019N. gonorrhoeae GCGS113N. gonorrhoeae GCGS103N. gonorrhoeae GCGS102N. gonorrhoeae GCGS025N. gonorrhoeae GCGS231N. gonorrhoeae GCGS183N. gonorrhoeae GCGS039N. gonorrhoeae GCGS143N. gonorrhoeae GCGS045N. gonorrhoeae GCGS029N. gonorrhoeae GCGS077N. gonorrhoeae GCGS031N. gonorrhoeae GCGS051N. gonorrhoeae GCGS181N. gonorrhoeae GCGS095N. gonorrhoeae GCGS227N. gonorrhoeae GCGS127N. gonorrhoeae GCGS242N. gonorrhoeae GCGS239N. gonorrhoeae GCGS187N. gonorrhoeae GCGS163N. gonorrhoeae GCGS141N. gonorrhoeae GCGS087N. gonorrhoeae GCGS003N. gonorrhoeae FA19N. gonorrhoeae GCGS008N. gonorrhoeae GCGS146N. gonorrhoeae GCGS186N. gonorrhoeae GCGS188N. gonorrhoeae GCGS090N. gonorrhoeae MS11N. gonorrhoeae GCGS210N. gonorrhoeae GCGS212N. gonorrhoeae GCGS200N. gonorrhoeae NG05N. gonorrhoeae n01.08N. gonorrhoeae GCGS050N. gonorrhoeae GCGS154N. gonorrhoeae GCGS072N. gonorrhoeae F62N. gonorrhoeae GCGS184N. gonorrhoeae FA 1090N. gonorrhoeae FA6140N. gonorrhoeae FA19N. gonorrhoeae GCGS162N. gonorrhoeae GCGS240N. gonorrhoeae GCGS190N. gonorrhoeae GCGS170N. gonorrhoeae GCGS056N. gonorrhoeae GCGS134N. gonorrhoeae GCGS078N. gonorrhoeae GCGS168N. gonorrhoeae GCGS206N. gonorrhoeae GCGS116N. gonorrhoeae GCGS044N. gonorrhoeae GCGS032N. gonorrhoeae GCGS022N. gonorrhoeae GCGS026N. gonorrhoeae GCGS060N. gonorrhoeae GCGS180N. gonorrhoeae GCGS216N. gonorrhoeae e03.04N. gonorrhoeae GCGS208N. gonorrhoeae ATL_2011_0508N. gonorrhoeae SK33414N. gonorrhoeae MIA_2011_0310N. gonorrhoeae i19.05N. gonorrhoeae GCGS150N. gonorrhoeae GCGS142N. gonorrhoeae GCGS010N. gonorrhoeae GCGS144N. gonorrhoeae SK39420N. gonorrhoeae FA19N. gonorrhoeae MU_NG6N. gonorrhoeae SK931035N. gonorrhoeae DGI18N. gonorrhoeae SK22871N. gonorrhoeae GCGS176N. gonorrhoeae GCGS018N. gonorrhoeae GCGS198N. gonorrhoeae GCGS202N. gonorrhoeae GCGS228N. gonorrhoeae GCGS204N. gonorrhoeae GCGS120N. gonorrhoeae GCGS138N. gonorrhoeae GCGS192N. gonorrhoeae GCGS062N. gonorrhoeae GCGS083N. gonorrhoeae GCGS084N. gonorrhoeae GCGS009N. gonorrhoeae GCGS106N. gonorrhoeae GCGS092N. gonorrhoeae GCGS098N. gonorrhoeae GCGS218N. gonorrhoeae GCGS110N. gonorrhoeae GCGS096N. gonorrhoeae GCGS118N. gonorrhoeae GCGS174N. gonorrhoeae GCGS224N. gonorrhoeae GCGS136N. gonorrhoeae GCGS156N. gonorrhoeae GCGS128N. gonorrhoeae GCGS226N. gonorrhoeae GCGS064N. gonorrhoeae GCGS172N. gonorrhoeae GCGS012N. gonorrhoeae GCGS100N. gonorrhoeae GCGS164N. gonorrhoeae GCGS040N. gonorrhoeae GCGS024N. gonorrhoeae GCGS082N. gonorrhoeae SK29344N. gonorrhoeae SK6987N. gonorrhoeae SK15454N. gonorrhoeae SK7842N. gonorrhoeae NG_869N. gonorrhoeae FA6140N. gonorrhoeae PID332N. gonorrhoeae DGI2N. gonorrhoeae SK8976N. gonorrhoeae SK708N. gonorrhoeae GCGS220N. gonorrhoeae GCGS182N. gonorrhoeae GCGS214N. gonorrhoeae GCGS086N. gonorrhoeae GCGS002N. gonorrhoeae GCGS020N. gonorrhoeae GCGS152N. gonorrhoeae GCGS052N. gonorrhoeae GCGS132N. gonorrhoeae GCGS130N. gonorrhoeae GCGS042N. gonorrhoeae GCGS194N. gonorrhoeae GCGS108N. gonorrhoeae GCGS070N. gonorrhoeae GCGS038N. gonorrhoeae GCGS230N. gonorrhoeae GCGS126N. gonorrhoeae GCGS175N. gonorrhoeae GCGS073N. gonorrhoeae GCGS213N. gonorrhoeae GCGS171N. gonorrhoeae GCGS209N. gonorrhoeae GCGS169N. gonorrhoeae GCGS203N. gonorrhoeae GCGS201N. gonorrhoeae GCGS071N. gonorrhoeae GCGS173N. gonorrhoeae GCGS197N. gonorrhoeae GCGS167N. gonorrhoeae GCGS219N. gonorrhoeae GCGS153N. gonorrhoeae GCGS211N. gonorrhoeae GCGS221N. gonorrhoeae GCGS149N. gonorrhoeae GCGS161N. gonorrhoeae GCGS199N. gonorrhoeae GCGS207N. gonorrhoeae GCGS191N. gonorrhoeae GCGS217N. gonorrhoeae GCGS193N. gonorrhoeae GCGS215N. gonorrhoeae GCGS223N. gonorrhoeae GCGS225N. gonorrhoeae GCGS034N. gonorrhoeae GCGS074N. gonorrhoeae 35/02N. gonorrhoeae GCGS011N. gonorrhoeae PID241N. gonorrhoeae PID1N. gonorrhoeae MS11N. gonorrhoeae SK1902N. gonorrhoeae ATL_2011_0121N. gonorrhoeae PID18N. meningitidis Z2491N. meningitidis 65014N. meningitidis 63006N. meningitidis 63041N. meningitidis 64182N. meningitidis 97027N. meningitidis 63049N. meningitidis 65012N. meningitidis 97018N. meningitidis 94018N. meningitidis 96024N. meningitidis 98005N. meningitidis 75689N. meningitidis 97008N. meningitidis 75643N. meningitidis 88050N. meningitidis 97020N. meningitidis 96023N. meningitidis WUE 2594N. meningitidis NM2254N. meningitidis NM2524N. meningitidis 2000080N. meningitidis NM2810N. meningitidis NM1364N. meningitidis NM1826N. meningitidis NM1758N. meningitidis NM2394N. meningitidis NM1893N. meningitidis NM1549N. meningitidis NM1976N. meningitidis NM1550N. meningitidis NM2393N. meningitidis NM2717N. meningitidis NM2382N. meningitidis NM2814N. meningitidis NM2606N. meningitidis NM2441N. meningitidis 2003022N. meningitidis NM607N. meningitidis 2004085N. meningitidis NM1471N. meningitidis NM2033N. meningitidis 2003022N. meningitidis NM2933N. meningitidis NM2237N. meningitidis NM2809N. meningitidis 2002007N. meningitidis NM1829N. meningitidis NM2263N. meningitidis NM1901N. meningitidis NM1805N. meningitidis NM1360N. meningitidis NM1482N. meningitidis NM1919N. meningitidis NM2617N. meningitidis NM2187N. meningitidis NM1757N. meningitidis NM2008N. meningitidis NM1361N. meningitidis NM2813N. meningitidis NM1561N. meningitidis NM1910N. meningitidis NM606N. meningitidis 2000063N. meningitidis NM1891N. meningitidis NM1931N. meningitidis NM1937N. meningitidis NM1938N. meningitidis NM1359N. meningitidis NM1264N. meningitidis NM2239N. meningitidis NM1362N. meningitidis NM2812N. meningitidis NM2431N. meningitidis NM2935N. meningitidis NM2439N. meningitidis NM2232N. meningitidis NM2193N. meningitidis NM2332N. meningitidis NM2228N. meningitidis NM2264N. meningitidis NM1895N. meningitidis NM2718N. meningitidis NM2188N. meningitidis NM2181N. meningitidis NM2369N. meningitidis NM2389N. meningitidis NM2811N. meningitidis NM2432N. meningitidis NM2335N. meningitidis NM2244N. meningitidis NM1928N. meningitidis NM2808N. meningitidis NM2381N. meningitidis NM2700N. meningitidis NM2932N. meningitidis NM1483N. meningitidis NM604N. meningitidis 510612N. meningitidis NM1673N. meningitidis NM2032N. meningitidis NM1666N. meningitidis NM1578N. meningitidis NM1963N. meningitidis NM1797N. meningitidis NM1837N. meningitidis NM3129N. meningitidis NM2602N. meningitidis NM2857N. meningitidis NM2206N. meningitidis NM1544N. meningitidis NM1921N. meningitidis NM1845N. meningitidis NM2856N. meningitidis NM1779N. meningitidis NM2934N. meningitidis NM1892N. meningitidis NM1325N. meningitidis NM1573N. meningitidis NM3128N. meningitidis NM2701N. meningitidis NM2009N. meningitidis NM1831N. meningitidis NM1446N. meningitidis NM2025N. meningitidis NM1363N. meningitidis NM1672N. meningitidis NM2433N. meningitidis 2007056N. meningitidis NM3652N. meningitidis 2004090N. meningitidis 2001212N. meningitidis NM3642N. meningitidis Nm1140N. meningitidis alpha704N. meningitidis NM2795N. meningitidis LNP27256N. meningitidisN. meningitidis NM586N. meningitidis M6190N. meningitidis S0108N. meningitidisN. meningitidis ES14902N. meningitidis NM1482N. meningitidis NM95N. meningitidis NM133N. meningitidis NM1482N. meningitidis M20599N. meningitidis K1207N. meningitidis S0108N. meningitidis NM94N. meningitidis NM82N. meningitidis NM3680N. meningitidis NM3681N. meningitidis NM3683N. meningitidis NM3682N. meningitidis M7124N. meningitidis 2005040N. meningitidis M10208N. meningitidis NM3686N. meningitidis NM3685N. meningitidis M7089N. meningitidis M7124N. meningitidis NM174N. meningitidis NM3687N. meningitidis NM3684N. meningitidis NM3688N. meningitidis NM126N. meningitidis FAM18N. meningitidis NM762N. meningitidis L91543N. meningitidis M12611N. meningitidis 2001213N. meningitidis NM43N. meningitidis M1412N. meningitidis 2001073N. meningitidis 2001068N. meningitidis NM32N. meningitidis NM23N. meningitidis NM35N. meningitidis NM36N. meningitidis 73704N. meningitidis 2000175N. meningitidis 2002004N. meningitidis 2004264N. meningitidis 2000081N. meningitidis 2005079N. meningitidis 2001072N. meningitidis NM1495N. meningitidis NM313N. meningitidis NM3147N. meningitidis 70012N. meningitidis 70030N. meningitidis 69166N. meningitidis 61103N. meningitidis 68094N. meningitidis 69096N. meningitidis 63023N. meningitidis 61106N. meningitidis 70021N. meningitidis 69155N. meningitidis 70082N. meningitidis 69100N. meningitidis 96060N. meningitidis 69176N. meningitidis NM477N. meningitidis NM3141N. meningitidis M13265N. meningitidis NM3173N. meningitidis 2002020N. meningitidis 2002030N. meningitidis NM1476N. meningitidis H44/76N. meningitidis H44/76N. meningitidis MC58N. meningitidis LNP21362N. meningitidis NM418N. meningitidis M13255N. meningitidis 9506N. meningitidis CU385N. meningitidis 12888N. meningitidis 4119N. meningitidis NM422N. meningitidis 9757N. meningitidis 320503N. meningitidis 100514N. meningitidis 370601N. meningitidis 100572N. meningitidis 360624N. meningitidis 421007N. meningitidis 440501N. meningitidis 340552N. meningitidis 100530N. meningitidis 321114N. meningitidis 34173N. meningitidis 311112N. meningitidis 053442N. meningitidis 100601N. meningitidis Nm11003N. meningitidis 341215N. meningitidis 320501N. meningitidis 220601N. meningitidis 440902N. meningitidis 130803N. meningitidis 131148N. meningitidis 420718N. meningitidis 100603N. meningitidis 330505N. meningitidis Nm2732N. meningitidis 81858N. meningitidis 8013N. meningitidis Nm10259N. meningitidis Nm9418N. meningitidis Nm6938N. meningitidis 92045N. meningitidis NM255N. meningitidis 98002N. meningitidis 2003051N. meningitidis 73696N. meningitidis NM134N. meningitidis Nm8187N. meningitidis Nm3127N. meningitidis 2001001N. meningitidis 2007461N. meningitidis 2004032N. meningitidis 992008N. meningitidis alpha710N. meningitidis OX99.30304N. meningitidis NM151N. meningitidis NZ05/33N. meningitidis DE9686N. meningitidis DE9622N. meningitidis DE9938N. meningitidis M0579N. meningitidis M01240149N. meningitidis M0579N. meningitidis NM2657N. meningitidis NM576N. meningitidis NM2781N. meningitidis NM140N. meningitidis NM183N. meningitidis Neisseria meningitidis CHUVN. meningitidis M13399N. meningitidis M01240013N. meningitidis M04240196N. meningitidis 77221N. meningitidis alpha14N. meningitidis NM3001N. meningitidis M01240355N. meningitidis 93003N. meningitidis 93004N. meningitidis NM3081N. meningitidis 98008N. meningitidis 80179N. meningitidis ATCC 13091N. meningitidis NM045N. meningitidis NM2866N. meningitidis NM003N. meningitidis NM3139N. meningitidis NM0552N. meningitidis NM518N. meningitidis NM3230N. meningitidis 96037N. meningitidis N1568N. meningitidis 2002038N. meningitidis 97021N. meningitidis 2006087N. meningitidis 97014N. meningitidis 2008223N. meningitidis 2005172N. meningitidis NM220N. meningitidis NM271N. meningitidis NM115N. meningitidis NM3144N. meningitidis NM27N. meningitidis NM3131N. meningitidis NM3158N. meningitidis NM3222N. meningitidis NM3164N. meningitidis Nm6756N. meningitidis Nm8663N. meningitidis NM27N. meningitidis NM90N. meningitidis NM233N. meningitidis NS44N. meningitidis NM80N. meningitidis NM165N. meningitidis NM51N. meningitidis NM3042N. meningitidis NM3223N. meningitidis B6116/77N. meningitidis G2136N. meningitidis 98080N. meningitidis 87255N. meningitidis NMBN. meningitidis 9615945N. polysaccharea ATCC 43768N. polysaccharea NS342N. lactamica NS19N. lactamica 02006N. lactamica Y921009N. lactamica ATCC 23970N. lactamica ATCC 23970N. cinerea ATCC 14685N. flavescens NRL30031/H210N. sicca VK64N. sicca ATCC 29256N. meningitidis 433_NMENN. meningitidis 1044_NMENN. meningitidis 840_NMENN. meningitidis 776_NMENN. subflava NJ9703N. meningitidis 1045_NMENN. meningitidis 768_NMENN. mucosa C6AN. lactamica 595_NLACN. lactamica 583_NLACN. lactamica 338.rep1_NLACN. meningitidis 338.rep2_NMENN. flavescens SK114N. meningitidis 782_NMENN. meningitidis 1210_NMENN. mucosa C102N. meningitidis 1279_NMENN. elongata ATCC 29315N. elongata ATCC 29315N. sp. oral taxon 014 F0314N. meningitidis 431_NMENN. meningitidis 404_NMENN. lactamica 919_NLACN. meningitidis 313_NMENN. meningitidis 379_NMENN. meningitidis 378_NMENN. sicca DS1N. sp. GT4A_CT1 GT4A_CT1N. macacae ATCC 33926N. meningitidis 1273_NMENN. sp. HMSC06F02 HMSC06F02N. mucosa ATCC 25996N. meningitidis 480_NMENN. sicca 4320N. sp. 83E34 83E34N. sp. 74A18 74A18N. wadsworthii 9715N. sp. KH1503 KH1503N. shayeganii 871N. weaveri ATCC 51223N. weaveri LMG 5135N. meningitidis 491_NMENN. sp. oral taxon 020 F0370N. meningitidis 203_NMENN. lactamica 914_NLACN. bacilliformis ATCC BAA1200
0.00
0.25
0.50
0.75
1.00
ANIm
_per
cent
age_
ident
ity
719 Neisseria spp. ANIm
719 Neisseria spp. ANIm
• Almost allNeisseria alignto over 50% ofgenome
• Minor complexof possiblemisassign-ments/novelspecies
N. g
onor
rhoe
ae S
K70
8N
. gon
orrh
oeae
MU
_NG
1N
. gon
orrh
oeae
PID
1N
. gon
orrh
oeae
MIA
_201
1_03
09
N. g
onor
rhoe
ae P
ID18
N. g
onor
rhoe
ae D
GI2
N. g
onor
rhoe
ae S
K9
310
35N
. gon
orrh
oeae
MS
11N
. gon
orrh
oeae
FA
19N
. gon
orrh
oeae
SK
92
679
N. g
onor
rhoe
ae P
ID33
2N
. gon
orrh
oeae
SK
3341
4N
. gon
orrh
oeae
MS
11N
. gon
orrh
oeae
GC
GS
033
N. g
onor
rhoe
ae N
CC
P11
945
N. g
onor
rhoe
ae M
IA_2
011_
031
0N
. gon
orrh
oeae
FA
19N
. gon
orrh
oeae
GC
GS
227
N. g
onor
rhoe
ae G
CG
S24
2N
. gon
orrh
oeae
GC
GS
061
N. g
onor
rhoe
ae G
CG
S08
9N
. gon
orrh
oeae
GC
GS
103
N. g
onor
rhoe
ae G
CG
S09
5N
. gon
orrh
oeae
GC
GS
189
N. g
onor
rhoe
ae G
CG
S18
1N
. gon
orrh
oeae
GC
GS
131
N. g
onor
rhoe
ae G
CG
S07
9N
. gon
orrh
oeae
GC
GS
115
N. g
onor
rhoe
ae G
CG
S23
9N
. gon
orrh
oeae
GC
GS
135
N. g
onor
rhoe
ae G
CG
S06
9N
. gon
orrh
oeae
GC
GS
113
N. g
onor
rhoe
ae G
CG
S20
5N
. gon
orrh
oeae
GC
GS
039
N. g
onor
rhoe
ae G
CG
S19
5N
. gon
orrh
oeae
GC
GS
133
N. g
onor
rhoe
ae G
CG
S05
1N
. gon
orrh
oeae
GC
GS
067
N. g
onor
rhoe
ae G
CG
S05
5N
. gon
orrh
oeae
GC
GS
127
N. g
onor
rhoe
ae G
CG
S01
9N
. gon
orrh
oeae
GC
GS
166
N. g
onor
rhoe
ae G
CG
S05
3N
. gon
orrh
oeae
GC
GS
141
N. g
onor
rhoe
ae G
CG
S14
3N
. gon
orrh
oeae
GC
GS
102
N. g
onor
rhoe
ae G
CG
S18
3N
. gon
orrh
oeae
GC
GS
140
N. g
onor
rhoe
ae G
CG
S00
4N
. gon
orrh
oeae
GC
GS
194
N. g
onor
rhoe
ae G
CG
S15
2N
. gon
orrh
oeae
GC
GS
182
N. g
onor
rhoe
ae G
CG
S21
4N
. gon
orrh
oeae
GC
GS
222
N. g
onor
rhoe
ae G
CG
S05
4N
. gon
orrh
oeae
GC
GS
034
N. g
onor
rhoe
ae G
CG
S20
9N
. gon
orrh
oeae
GC
GS
171
N. g
onor
rhoe
ae G
CG
S16
4N
. gon
orrh
oeae
GC
GS
073
N. g
onor
rhoe
ae G
CG
S21
3N
. gon
orrh
oeae
GC
GS
169
N. g
onor
rhoe
ae n
01.0
8N
. gon
orrh
oeae
GC
GS
173
N. g
onor
rhoe
ae G
CG
S07
1N
. gon
orrh
oeae
GC
GS
080
N. g
onor
rhoe
ae G
CG
S17
0N
. gon
orrh
oeae
GC
GS
056
N. g
onor
rhoe
ae G
CG
S03
6N
. gon
orrh
oeae
GC
GS
177
N. g
onor
rhoe
ae G
CG
S19
0N
. gon
orrh
oeae
GC
GS
052
N. g
onor
rhoe
ae G
CG
S23
4N
. gon
orrh
oeae
GC
GS
240
N. g
onor
rhoe
ae G
CG
S23
6N
. gon
orrh
oeae
FA
19N
. gon
orrh
oeae
GC
GS
203
N. g
onor
rhoe
ae G
CG
S15
3N
. gon
orrh
oeae
GC
GS
197
N. g
onor
rhoe
ae G
CG
S20
0N
. gon
orrh
oeae
GC
GS
175
N. g
onor
rhoe
ae G
CG
S16
7N
. gon
orrh
oeae
GC
GS
201
N. g
onor
rhoe
ae G
CG
S02
2N
. gon
orrh
oeae
GC
GS
108
N. g
onor
rhoe
ae G
CG
S12
2N
. gon
orrh
oeae
GC
GS
196
N. g
onor
rhoe
ae G
CG
S07
0N
. gon
orrh
oeae
GC
GS
238
N. g
onor
rhoe
ae A
LB_2
011_
010
2N
. gon
orrh
oeae
MU
_NG
5N
. gon
orrh
oeae
GC
GS
078
N. g
onor
rhoe
ae G
CG
S22
1N
. gon
orrh
oeae
GC
GS
011
N. g
onor
rhoe
ae G
CG
S20
6N
. gon
orrh
oeae
GC
GS
116
N. g
onor
rhoe
ae G
CG
S21
1N
. gon
orrh
oeae
GC
GS
217
N. g
onor
rhoe
ae G
CG
S10
6N
. gon
orrh
oeae
GC
GS
191
N. g
onor
rhoe
ae G
CG
S10
0N
. gon
orrh
oeae
GC
GS
026
N. g
onor
rhoe
ae G
CG
S23
2N
. gon
orrh
oeae
GC
GS
023
N. g
onor
rhoe
ae G
CG
S04
2N
. gon
orrh
oeae
GC
GS
148
N. g
onor
rhoe
ae G
CG
S00
6N
. gon
orrh
oeae
GC
GS
132
N. g
onor
rhoe
ae G
CG
S09
4N
. gon
orrh
oeae
GC
GS
130
N. g
onor
rhoe
ae G
CG
S08
4N
. gon
orrh
oeae
GC
GS
118
N. g
onor
rhoe
ae G
CG
S19
8N
. gon
orrh
oeae
GC
GS
060
N. g
onor
rhoe
ae G
CG
S22
8N
. gon
orrh
oeae
GC
GS
202
N. g
onor
rhoe
ae G
CG
S01
8N
. gon
orrh
oeae
GC
GS
176
N. g
onor
rhoe
ae G
CG
S03
8N
. gon
orrh
oeae
GC
GS
009
N. g
onor
rhoe
ae G
CG
S06
2N
. gon
orrh
oeae
GC
GS
010
N. g
onor
rhoe
ae G
CG
S07
4N
. gon
orrh
oeae
GC
GS
096
N. g
onor
rhoe
ae G
CG
S17
4N
. gon
orrh
oeae
GC
GS
120
N. g
onor
rhoe
ae G
CG
S22
4N
. gon
orrh
oeae
GC
GS
204
N. g
onor
rhoe
ae G
CG
S02
0N
. gon
orrh
oeae
GC
GS
235
N. g
onor
rhoe
ae G
CG
S13
7N
. gon
orrh
oeae
GC
GS
003
N. g
onor
rhoe
ae G
CG
S06
8N
. gon
orrh
oeae
GC
GS
007
N. g
onor
rhoe
ae G
CG
S09
1N
. gon
orrh
oeae
GC
GS
220
N. g
onor
rhoe
ae G
CG
S10
4N
. gon
orrh
oeae
GC
GS
037
N. g
onor
rhoe
ae G
CG
S18
7N
. gon
orrh
oeae
GC
GS
017
N. g
onor
rhoe
ae G
CG
S24
1N
. gon
orrh
oeae
GC
GS
125
N. g
onor
rhoe
ae G
CG
S10
7N
. gon
orrh
oeae
GC
GS
157
N. g
onor
rhoe
ae G
CG
S04
9N
. gon
orrh
oeae
GC
GS
027
N. g
onor
rhoe
ae G
CG
S23
3N
. gon
orrh
oeae
GC
GS
151
N. g
onor
rhoe
ae G
CG
S05
9N
. gon
orrh
oeae
GC
GS
123
N. g
onor
rhoe
ae G
CG
S14
7N
. gon
orrh
oeae
GC
GS
076
N. g
onor
rhoe
ae G
CG
S12
9N
. gon
orrh
oeae
GC
GS
001
N. g
onor
rhoe
ae G
CG
S05
7N
. gon
orrh
oeae
GC
GS
035
N. g
onor
rhoe
ae G
CG
S15
5N
. gon
orrh
oeae
GC
GS
030
N. g
onor
rhoe
ae G
CG
S18
5N
. gon
orrh
oeae
GC
GS
160
N. g
onor
rhoe
ae G
CG
S16
5N
. gon
orrh
oeae
GC
GS
159
N. g
onor
rhoe
ae G
CG
S08
5N
. gon
orrh
oeae
GC
GS
087
N. g
onor
rhoe
ae G
CG
S10
9N
. gon
orrh
oeae
GC
GS
066
N. g
onor
rhoe
ae G
CG
S03
1N
. gon
orrh
oeae
GC
GS
163
N. g
onor
rhoe
ae G
CG
S13
9N
. gon
orrh
oeae
GC
GS
029
N. g
onor
rhoe
ae G
CG
S06
5N
. gon
orrh
oeae
GC
GS
025
N. g
onor
rhoe
ae G
CG
S04
5N
. gon
orrh
oeae
GC
GS
231
N. g
onor
rhoe
ae G
CG
S07
7N
. gon
orrh
oeae
GC
GS
075
N. g
onor
rhoe
ae G
CG
S14
5N
. gon
orrh
oeae
GC
GS
119
N. g
onor
rhoe
ae G
CG
S23
7N
. gon
orrh
oeae
GC
GS
117
N. g
onor
rhoe
ae G
CG
S09
7N
. gon
orrh
oeae
GC
GS
005
N. g
onor
rhoe
ae G
CG
S04
8N
. gon
orrh
oeae
GC
GS
028
N. g
onor
rhoe
ae G
CG
S12
4N
. gon
orrh
oeae
GC
GS
178
N. g
onor
rhoe
ae G
CG
S10
5N
. gon
orrh
oeae
GC
GS
101
N. g
onor
rhoe
ae G
CG
S22
9N
. gon
orrh
oeae
GC
GS
114
N. g
onor
rhoe
ae G
CG
S04
6N
. gon
orrh
oeae
GC
GS
047
N. g
onor
rhoe
ae G
CG
S06
3N
. gon
orrh
oeae
GC
GS
088
N. g
onor
rhoe
ae G
CG
S04
1N
. gon
orrh
oeae
GC
GS
043
N. g
onor
rhoe
ae F
A61
40N
. gon
orrh
oeae
MU
_NG
18N
. gon
orrh
oeae
DG
I18
N. g
onor
rhoe
ae A
TL_
2011
_01_
05N
. gon
orrh
oeae
AT
L_20
11_0
1_08
N. g
onor
rhoe
ae M
U_N
G26
N. g
onor
rhoe
ae A
LB_2
011_
03_0
3N
. gon
orrh
oeae
AT
L_20
11_0
1_03
N. g
onor
rhoe
ae M
U_N
G4
N. g
onor
rhoe
ae M
U_N
G21
N. g
onor
rhoe
ae A
TL_
2011
_01_
17N
. gon
orrh
oeae
PID
241
N. g
onor
rhoe
ae 1
291
N. g
onor
rhoe
ae 3
5/02
N. g
onor
rhoe
ae i1
9.05
N. g
onor
rhoe
ae N
YC
_201
1_05
_07
N. g
onor
rhoe
ae M
U_N
G15
N. g
onor
rhoe
ae M
U_N
G14
N. g
onor
rhoe
ae M
U_N
G12
N. g
onor
rhoe
ae N
G_8
69N
. gon
orrh
oeae
CH
811
N. g
onor
rhoe
ae e
03.0
4N
. gon
orrh
oeae
MU
_NG
19N
. gon
orrh
oeae
AT
L_20
11_0
121
N. g
onor
rhoe
ae N
OR
_201
1_03
06
N. g
onor
rhoe
ae N
YC
_201
1_05
_13
N. g
onor
rhoe
ae G
C1
182
N. g
onor
rhoe
ae M
U_N
G23
N. g
onor
rhoe
ae M
U_N
G17
N. g
onor
rhoe
ae S
K15
454
N. g
onor
rhoe
ae M
IA_2
011_
051
6N
. gon
orrh
oeae
FA
109
0N
. gon
orrh
oeae
m07
.05
N. g
onor
rhoe
ae S
K22
871
N. g
onor
rhoe
ae S
K74
61N
. gon
orrh
oeae
MU
_NG
9N
. gon
orrh
oeae
MIA
_201
1_02
_02
N. g
onor
rhoe
ae A
LB_2
011_
04_0
3N
. gon
orrh
oeae
MU
_NG
6N
. gon
orrh
oeae
SK
1902
N. g
onor
rhoe
ae M
IA_2
011_
051
0N
. gon
orrh
oeae
GC
GS
142
N. g
onor
rhoe
ae S
K29
344
N. g
onor
rhoe
ae S
K28
355
N. g
onor
rhoe
ae S
K17
973
N. g
onor
rhoe
ae S
K69
87N
. gon
orrh
oeae
SK
1694
2N
. gon
orrh
oeae
SK
7842
N. g
onor
rhoe
ae S
K12
684
N. g
onor
rhoe
ae S
K29
471
N. g
onor
rhoe
ae G
CG
S21
0N
. gon
orrh
oeae
GC
GS
212
N. g
onor
rhoe
ae G
CG
S04
0N
. gon
orrh
oeae
SK
3942
0N
. gon
orrh
oeae
MU
_NG
8N
. gon
orrh
oeae
MU
_NG
3N
. gon
orrh
oeae
MU
_NG
20N
. gon
orrh
oeae
SK
8976
N. g
onor
rhoe
ae F
62N
. gon
orrh
oeae
SK
3240
2N
. gon
orrh
oeae
GC
GS
064
N. g
onor
rhoe
ae S
K36
809
N. g
onor
rhoe
ae G
CG
S18
4N
. gon
orrh
oeae
GC
GS
082
N. g
onor
rhoe
ae G
CG
S02
4N
. gon
orrh
oeae
AT
L_20
11_0
513
N. g
onor
rhoe
ae G
2891
N. g
onor
rhoe
ae A
TL_
2011
_05
08N
. gon
orrh
oeae
MIA
_201
1_05
15
N. g
onor
rhoe
ae M
U_N
G25
N. g
onor
rhoe
ae G
CG
S05
0N
. gon
orrh
oeae
SK
2302
0N
. gon
orrh
oeae
GC
GS
012
N. g
onor
rhoe
ae G
CG
S21
5N
. gon
orrh
oeae
GC
GS
225
N. g
onor
rhoe
ae G
CG
S16
8N
. gon
orrh
oeae
GC
GS
199
N. g
onor
rhoe
ae G
CG
S21
9N
. gon
orrh
oeae
GC
GS
193
N. g
onor
rhoe
ae 0
9_10
_003
N. g
onor
rhoe
ae A
TL_
2011
_01
25N
. gon
orrh
oeae
GC
GS
072
N. g
onor
rhoe
ae G
CG
S16
2N
. gon
orrh
oeae
FA
6140
N. g
onor
rhoe
ae 3
5/02
N. g
onor
rhoe
ae G
CG
S15
0N
. gon
orrh
oeae
GC
GS
216
N. g
onor
rhoe
ae G
CG
S18
6N
. gon
orrh
oeae
GC
GS
090
N. g
onor
rhoe
ae G
CG
S18
8N
. gon
orrh
oeae
GC
GS
008
N. g
onor
rhoe
ae S
K16
259
N. g
onor
rhoe
ae S
K14
515
N. g
onor
rhoe
ae N
G05
N. g
onor
rhoe
ae G
CG
S18
0N
. gon
orrh
oeae
GC
GS
208
N. g
onor
rhoe
ae G
CG
S15
4N
. gon
orrh
oeae
GC
GS
146
N. g
onor
rhoe
ae G
CG
S21
8N
. gon
orrh
oeae
GC
GS
128
N. g
onor
rhoe
ae G
CG
S22
6N
. gon
orrh
oeae
GC
GS
032
N. g
onor
rhoe
ae G
CG
S12
6N
. gon
orrh
oeae
GC
GS
092
N. g
onor
rhoe
ae G
CG
S09
8N
. gon
orrh
oeae
GC
GS
058
N. g
onor
rhoe
ae G
CG
S00
2N
. gon
orrh
oeae
GC
GS
086
N. g
onor
rhoe
ae G
CG
S20
7N
. gon
orrh
oeae
GC
GS
161
N. g
onor
rhoe
ae G
CG
S13
4N
. gon
orrh
oeae
GC
GS
223
N. g
onor
rhoe
ae G
CG
S14
9N
. gon
orrh
oeae
GC
GS
138
N. g
onor
rhoe
ae G
CG
S11
0N
. gon
orrh
oeae
GC
GS
083
N. g
onor
rhoe
ae G
CG
S19
2N
. gon
orrh
oeae
GC
GS
136
N. g
onor
rhoe
ae G
CG
S15
6N
. gon
orrh
oeae
GC
GS
172
N. g
onor
rhoe
ae G
CG
S14
4N
. gon
orrh
oeae
GC
GS
230
N. g
onor
rhoe
ae G
CG
S04
4N
. gon
orrh
oeae
GC
GS
081
N. g
onor
rhoe
ae G
CG
S17
9N
. gon
orrh
oeae
GC
GS
121
N. g
onor
rhoe
ae G
CG
S02
1N
. gon
orrh
oeae
GC
GS
158
N. g
onor
rhoe
ae G
CG
S09
9N
. gon
orrh
oeae
GC
GS
093
N. p
olys
acch
area
AT
CC
437
68N
. pol
ysac
char
ea N
S34
2N
. lac
tam
ica
NS
19N
. cin
erea
AT
CC
146
85N
. lac
tam
ica
020
06N
. lac
tam
ica
AT
CC
239
70N
. lac
tam
ica
AT
CC
239
70N
. lac
tam
ica
Y92
100
9N
. men
ingi
tidis
NM
233
N. m
enin
gitid
is N
m11
003
N. m
enin
gitid
is K
1207
N. m
enin
gitid
is S
0108
N. m
enin
gitid
is N
M27
01N
. men
ingi
tidis
NM
2264
N. m
enin
gitid
is N
M15
49N
. men
ingi
tidis
NM
1550
N. m
enin
gitid
is N
M18
31N
. men
ingi
tidis
NM
1325
N. m
enin
gitid
is N
M19
76N
. men
ingi
tidis
NM
2254
N. m
enin
gitid
is N
M20
32N
. men
ingi
tidis
NM
2933
N. m
enin
gitid
is N
M23
94N
. men
ingi
tidis
NM
1893
N. m
enin
gitid
is N
M25
24N
. men
ingi
tidis
NM
1779
N. m
enin
gitid
is N
M16
72N
. men
ingi
tidis
NM
1826
N. m
enin
gitid
is N
M15
73N
. men
ingi
tidis
NM
3128
N. m
enin
gitid
is N
M28
56N
. men
ingi
tidis
NM
2932
N. m
enin
gitid
is N
M17
97N
. men
ingi
tidis
NM
1937
N. m
enin
gitid
is N
M17
58N
. men
ingi
tidis
NM
2228
N. m
enin
gitid
is N
M20
09N
. men
ingi
tidis
NM
1892
N. m
enin
gitid
is N
M16
73N
. men
ingi
tidis
NM
2617
N. m
enin
gitid
is N
M14
46N
. men
ingi
tidis
NM
2187
N. m
enin
gitid
is N
M24
33N
. men
ingi
tidis
NM
1921
N. m
enin
gitid
is N
M13
63N
. men
ingi
tidis
NM
2857
N. m
enin
gitid
is N
M18
45N
. men
ingi
tidis
NM
2934
N. m
enin
gitid
is N
M24
41N
. men
ingi
tidis
NM
2700
N. m
enin
gitid
is N
M26
02N
. men
ingi
tidis
NM
2393
N. m
enin
gitid
is N
M19
10N
. men
ingi
tidis
NM
1471
N. m
enin
gitid
is N
M13
64N
. men
ingi
tidis
NM
1901
N. m
enin
gitid
is N
M28
09N
. men
ingi
tidis
NM
1483
N. m
enin
gitid
is N
M18
29N
. men
ingi
tidis
NM
2718
N. m
enin
gitid
is N
M12
64N
. men
ingi
tidis
NM
2813
N. m
enin
gitid
is N
M22
37N
. men
ingi
tidis
NM
2181
N. m
enin
gitid
is N
M13
59N
. men
ingi
tidis
NM
1963
N. m
enin
gitid
is N
M22
39N
. men
ingi
tidis
NM
1938
N. m
enin
gitid
is N
M15
61N
. men
ingi
tidis
NM
1891
N. m
enin
gitid
is N
M20
08N
. men
ingi
tidis
NM
2381
N. m
enin
gitid
is N
M15
78N
. men
ingi
tidis
NM
2193
N. m
enin
gitid
is N
M24
31N
. men
ingi
tidis
NM
1928
N. m
enin
gitid
is N
M22
06N
. men
ingi
tidis
NM
1757
N. m
enin
gitid
is N
M28
12N
. men
ingi
tidis
NM
2432
N. m
enin
gitid
is N
M22
32N
. men
ingi
tidis
NM
2025
N. m
enin
gitid
is N
M28
08N
. men
ingi
tidis
NM
1837
N. m
enin
gitid
is N
M28
11N
. men
ingi
tidis
NM
2717
N. m
enin
gitid
is N
M19
31N
. men
ingi
tidis
NM
2263
N. m
enin
gitid
is N
M28
10N
. men
ingi
tidis
NM
2332
N. m
enin
gitid
is N
M18
05N
. men
ingi
tidis
NM
1360
N. m
enin
gitid
is N
M23
82N
. men
ingi
tidis
NM
1482
N. m
enin
gitid
is N
M23
35N
. men
ingi
tidis
NM
1895
N. m
enin
gitid
is N
M26
06N
. men
ingi
tidis
NM
2389
N. m
enin
gitid
is N
M13
62N
. men
ingi
tidis
NM
1361
N. m
enin
gitid
is N
M23
69N
. men
ingi
tidis
NM
1919
N. m
enin
gitid
is N
M22
44N
. men
ingi
tidis
NM
3129
N. m
enin
gitid
is N
M29
35N
. men
ingi
tidis
NM
1666
N. m
enin
gitid
is N
M28
14N
. men
ingi
tidis
NM
2439
N. m
enin
gitid
is N
M21
88N
. men
ingi
tidis
NM
1544
N. m
enin
gitid
is 2
0000
63N
. men
ingi
tidis
200
4085
N. m
enin
gitid
is N
M31
58N
. men
ingi
tidis
Nm
8663
N. m
enin
gitid
is N
M43
N. m
enin
gitid
is 2
0010
73N
. men
ingi
tidis
NM
1482
N. m
enin
gitid
is 4
2071
8N
. men
ingi
tidis
131
148
N. m
enin
gitid
is 2
0012
13N
. men
ingi
tidis
NM
95N
. men
ingi
tidis
Nm
9418
N. m
enin
gitid
is N
m10
259
N. m
enin
gitid
is N
m67
56N
. men
ingi
tidis
340
552
N. m
enin
gitid
is 4
4090
2N
. men
ingi
tidis
M13
265
N. m
enin
gitid
is 3
6062
4N
. men
ingi
tidis
Nei
sser
ia m
enin
gitid
is C
HU
VN
. men
ingi
tidis
100
603
N. m
enin
gitid
is 1
3080
3N
. men
ingi
tidis
330
505
N. m
enin
gitid
is 1
0051
4N
. men
ingi
tidis
NM
220
N. m
enin
gitid
is N
S44
N. m
enin
gitid
is N
M36
80N
. men
ingi
tidis
NM
36N
. men
ingi
tidis
NM
3684
N. m
enin
gitid
is N
m11
40N
. men
ingi
tidis
992
008
N. m
enin
gitid
is N
M36
85N
. men
ingi
tidis
NM
3688
N. m
enin
gitid
is 3
4121
5N
. men
ingi
tidis
100
601
N. m
enin
gitid
is N
M32
30N
. men
ingi
tidis
S01
08N
. men
ingi
tidis
200
2004
N. m
enin
gitid
is M
0579
N. m
enin
gitid
is N
M14
76N
. men
ingi
tidis
NM
2866
N. m
enin
gitid
is N
M31
47N
. men
ingi
tidis
NM
3164
N. m
enin
gitid
is 7
3696
N. m
enin
gitid
is N
M82
N. m
enin
gitid
is 2
0000
81N
. men
ingi
tidis
M01
240
355
N. m
enin
gitid
is N
M04
5N
. men
ingi
tidis
NM
0552
N. m
enin
gitid
is 2
0020
20N
. men
ingi
tidis
630
23N
. men
ingi
tidis
200
3051
N. m
enin
gitid
is 9
6060
N. m
enin
gitid
is 2
0074
61N
. men
ingi
tidis
NM
271
N. m
enin
gitid
is 8
1858
N. m
enin
gitid
is a
lpha
14N
. men
ingi
tidis
AT
CC
130
91N
. men
ingi
tidis
M04
240
196
N. m
enin
gitid
is H
44/7
6N
. men
ingi
tidis
NZ
05/
33N
. men
ingi
tidis
NM
606
N. m
enin
gitid
is 9
7018
N. m
enin
gitid
is 6
4182
N. m
enin
gitid
is 9
7027
N. m
enin
gitid
is N
M60
4N
. men
ingi
tidis
970
08N
. men
ingi
tidis
700
21N
. men
ingi
tidis
611
06N
. men
ingi
tidis
700
82N
. men
ingi
tidis
691
00N
. men
ingi
tidis
200
3022
N. m
enin
gitid
is 7
5643
N. m
enin
gitid
is 6
5012
N. m
enin
gitid
is 9
4018
N. m
enin
gitid
is 7
3704
N. m
enin
gitid
is C
U38
5N
. men
ingi
tidis
NM
2795
N. m
enin
gitid
is M
1261
1N
. men
ingi
tidis
LN
P27
256
N. m
enin
gitid
is D
E96
22N
. men
ingi
tidis
NM
2781
N. m
enin
gitid
is 3
2050
1N
. men
ingi
tidis
100
572
N. m
enin
gitid
is 3
1111
2N
. men
ingi
tidis
NM
BN
. men
ingi
tidis
M14
12N
. men
ingi
tidis
NM
140
N. m
enin
gitid
is 8
0179
N. m
enin
gitid
is D
E96
86N
. men
ingi
tidis
100
530
N. m
enin
gitid
is 3
2050
3N
. men
ingi
tidis
321
114
N. m
enin
gitid
is 2
2060
1N
. men
ingi
tidis
421
007
N. m
enin
gitid
is 3
7060
1N
. men
ingi
tidis
H44
/76
N. m
enin
gitid
is 6
9155
N. m
enin
gitid
is N
M32
23N
. men
ingi
tidis
200
4032
N. m
enin
gitid
is D
E99
38N
. men
ingi
tidis
Nm
3127
N. m
enin
gitid
is 4
4050
1N
. men
ingi
tidis
341
73N
. men
ingi
tidis
Nm
8187
N. m
enin
gitid
is N
m27
32N
. men
ingi
tidis
alp
ha70
4N
. men
ingi
tidis
Nm
6938
N. m
enin
gitid
isN
. men
ingi
tidis
NM
003
N. m
enin
gitid
is N
M16
5N
. men
ingi
tidis
NM
51N
. men
ingi
tidis
N. m
enin
gitid
is N
M13
3N
. men
ingi
tidis
NM
3141
N. m
enin
gitid
is 9
6037
N. m
enin
gitid
is N
M15
1N
. men
ingi
tidis
NM
3042
N. m
enin
gitid
is N
M32
22N
. men
ingi
tidis
NM
90N
. men
ingi
tidis
NM
1482
N. m
enin
gitid
is N
M31
39N
. men
ingi
tidis
200
8223
N. m
enin
gitid
is N
M27
N. m
enin
gitid
is 2
0051
72N
. men
ingi
tidis
NM
23N
. men
ingi
tidis
NM
94N
. men
ingi
tidis
200
1068
N. m
enin
gitid
is 2
0050
79N
. men
ingi
tidis
200
4264
N. m
enin
gitid
is N
M47
7N
. men
ingi
tidis
NM
35N
. men
ingi
tidis
200
1001
N. m
enin
gitid
is N
M51
8N
. men
ingi
tidis
B61
16/7
7N
. men
ingi
tidis
G21
36N
. men
ingi
tidis
M10
208
N. m
enin
gitid
is N
M36
86N
. men
ingi
tidis
NM
3682
N. m
enin
gitid
is N
M31
3N
. men
ingi
tidis
NM
115
N. m
enin
gitid
is N
M31
73N
. men
ingi
tidis
NM
134
N. m
enin
gitid
is 9
8002
N. m
enin
gitid
is N
M27
N. m
enin
gitid
is N
M80
N. m
enin
gitid
is 2
0020
30N
. men
ingi
tidis
M01
240
149
N. m
enin
gitid
is L
NP
2136
2N
. men
ingi
tidis
M71
24N
. men
ingi
tidis
FA
M18
N. m
enin
gitid
is N
M36
83N
. men
ingi
tidis
200
0175
N. m
enin
gitid
is 2
0050
40N
. men
ingi
tidis
NM
1495
N. m
enin
gitid
is N
M32
N. m
enin
gitid
is 2
0010
72N
. men
ingi
tidis
756
89N
. men
ingi
tidis
NM
607
N. m
enin
gitid
is N
M20
33N
. men
ingi
tidis
960
24N
. men
ingi
tidis
200
3022
N. m
enin
gitid
is 2
0020
07N
. men
ingi
tidis
980
05N
. men
ingi
tidis
200
0080
N. m
enin
gitid
is Z
2491
N. m
enin
gitid
is 6
9176
N. m
enin
gitid
is N
M31
31N
. men
ingi
tidis
NM
3144
N. m
enin
gitid
is 2
0012
12N
. men
ingi
tidis
630
49N
. men
ingi
tidis
611
03N
. men
ingi
tidis
053
442
N. m
enin
gitid
is N
M57
6N
. men
ingi
tidis
NM
2657
N. m
enin
gitid
is N
M30
01N
. men
ingi
tidis
411
9N
. men
ingi
tidis
920
45N
. men
ingi
tidis
L91
543
N. m
enin
gitid
is 9
3003
N. m
enin
gitid
is N
M18
3N
. men
ingi
tidis
N15
68N
. men
ingi
tidis
M61
90N
. men
ingi
tidis
ES
1490
2N
. men
ingi
tidis
NM
174
N. m
enin
gitid
is 8
7255
N. m
enin
gitid
is O
X99
.303
04N
. men
ingi
tidis
NM
586
N. m
enin
gitid
is 9
8080
N. m
enin
gitid
is M
7089
N. m
enin
gitid
is N
M25
5N
. men
ingi
tidis
510
612
N. m
enin
gitid
is W
UE
259
4N
. men
ingi
tidis
630
06N
. men
ingi
tidis
680
94N
. men
ingi
tidis
630
41N
. men
ingi
tidis
970
20N
. men
ingi
tidis
NM
3642
N. m
enin
gitid
is 2
0040
90N
. men
ingi
tidis
880
50N
. men
ingi
tidis
960
23N
. men
ingi
tidis
691
66N
. men
ingi
tidis
700
12N
. men
ingi
tidis
NM
3652
N. m
enin
gitid
is 2
0070
56N
. men
ingi
tidis
650
14N
. men
ingi
tidis
961
594
5N
. men
ingi
tidis
MC
58N
. men
ingi
tidis
930
04N
. men
ingi
tidis
801
3N
. men
ingi
tidis
128
88N
. men
ingi
tidis
NM
3081
N. m
enin
gitid
is N
M41
8N
. men
ingi
tidis
NM
422
N. m
enin
gitid
is M
0579
N. m
enin
gitid
is M
012
4001
3N
. men
ingi
tidis
alp
ha71
0N
. men
ingi
tidis
950
6N
. men
ingi
tidis
M13
399
N. m
enin
gitid
is N
M36
81N
. men
ingi
tidis
NM
3687
N. m
enin
gitid
is 9
7014
N. m
enin
gitid
is 6
9096
N. m
enin
gitid
is 9
757
N. m
enin
gitid
is 9
7021
N. m
enin
gitid
is 2
0060
87N
. men
ingi
tidis
M20
599
N. m
enin
gitid
is 7
7221
N. m
enin
gitid
is M
1325
5N
. men
ingi
tidis
980
08N
. men
ingi
tidis
NM
762
N. m
enin
gitid
is M
7124
N. m
enin
gitid
is N
M12
6N
. men
ingi
tidis
700
30N
. men
ingi
tidis
200
2038
N. m
enin
gitid
is 4
80_N
ME
NN
. sp.
HM
SC
06F
02 H
MS
C06
F02
N. s
icca
432
0N
. sic
ca D
S1
N. m
ucos
a A
TC
C 2
5996
N. m
enin
gitid
is 7
82_N
ME
NN
. fla
vesc
ens
SK
114
N. m
ucos
a C
102
N. m
enin
gitid
is 3
38.r
ep2_
NM
EN
N. l
acta
mic
a 33
8.re
p1_N
LAC
N. m
ucos
a C
6AN
. fla
vesc
ens
NR
L300
31/H
210
N. s
icca
VK
64N
. men
ingi
tidis
379
_NM
EN
N. m
enin
gitid
is 3
78_N
ME
NN
. men
ingi
tidis
313
_NM
EN
N. s
p. o
ral t
axon
014
F03
14N
. sub
flava
NJ9
703
N. l
acta
mic
a 59
5_N
LAC
N. l
acta
mic
a 58
3_N
LAC
N. m
enin
gitid
is 4
33_N
ME
NN
. men
ingi
tidis
104
5_N
ME
NN
. men
ingi
tidis
104
4_N
ME
NN
. men
ingi
tidis
840
_NM
EN
N. m
enin
gitid
is 7
76_N
ME
NN
. men
ingi
tidis
121
0_N
ME
NN
. men
ingi
tidis
768
_NM
EN
N. s
p. G
T4A
_CT
1 G
T4A
_CT
1N
. mac
acae
AT
CC
339
26N
. sic
ca A
TC
C 2
9256
N. m
enin
gitid
is 1
273_
NM
EN
N. l
acta
mic
a 91
9_N
LAC
N. m
enin
gitid
is 1
279_
NM
EN
N. m
enin
gitid
is 4
31_N
ME
NN
. men
ingi
tidis
404
_NM
EN
N. e
long
ata
AT
CC
293
15N
. elo
ngat
a A
TC
C 2
9315
N. s
p. o
ral t
axon
020
F03
70N
. bac
illifo
rmis
AT
CC
BA
A1
200
N. l
acta
mic
a 91
4_N
LAC
N. m
enin
gitid
is 2
03_N
ME
NN
. men
ingi
tidis
491
_NM
EN
N. s
p. 7
4A18
74A
18N
. wea
veri
AT
CC
512
23N
. wea
veri
LMG
513
5N
. sp.
KH
1503
KH
1503
N. s
p. 8
3E34
83E
34N
. sha
yega
nii 8
71N
. wad
swor
thii
9715
N. cinerea ATCC 14685N. lactamica ATCC 23970N. lactamica 02006N. lactamica ATCC 23970N. lactamica Y921009N. polysaccharea NS342N. polysaccharea ATCC 43768N. lactamica NS19N. gonorrhoeae 09_10_003N. gonorrhoeae SK7461N. gonorrhoeae SK16259N. gonorrhoeae SK14515N. gonorrhoeae PID1N. gonorrhoeae PID332N. gonorrhoeae DGI2N. gonorrhoeae MU_NG9N. gonorrhoeae MU_NG1N. gonorrhoeae MIA_2011_0309N. gonorrhoeae MIA_2011_02_02N. gonorrhoeae ALB_2011_04_03N. gonorrhoeae NYC_2011_05_07N. gonorrhoeae MU_NG15N. gonorrhoeae NYC_2011_05_13N. gonorrhoeae PID18N. gonorrhoeae GC1182N. gonorrhoeae MU_NG14N. gonorrhoeae MU_NG12N. gonorrhoeae MU_NG21N. gonorrhoeae ATL_2011_01_17N. gonorrhoeae ATL_2011_01_08N. gonorrhoeae MU_NG26N. gonorrhoeae 35/02N. gonorrhoeae MU_NG23N. gonorrhoeae MU_NG17N. gonorrhoeae ALB_2011_03_03N. gonorrhoeae ATL_2011_01_03N. gonorrhoeae MU_NG4N. gonorrhoeae MU_NG20N. gonorrhoeae 1291N. gonorrhoeae SK92679N. gonorrhoeae MU_NG19N. gonorrhoeae DGI18N. gonorrhoeae FA6140N. gonorrhoeae PID241N. gonorrhoeae SK931035N. gonorrhoeae MU_NG8N. gonorrhoeae MU_NG18N. gonorrhoeae MU_NG3N. gonorrhoeae ATL_2011_01_05N. gonorrhoeae MS11N. gonorrhoeae GCGS033N. gonorrhoeae GCGS227N. gonorrhoeae GCGS242N. gonorrhoeae GCGS061N. gonorrhoeae GCGS089N. gonorrhoeae GCGS135N. gonorrhoeae GCGS069N. gonorrhoeae GCGS095N. gonorrhoeae GCGS181N. gonorrhoeae GCGS189N. gonorrhoeae GCGS131N. gonorrhoeae GCGS240N. gonorrhoeae GCGS036N. gonorrhoeae GCGS103N. gonorrhoeae GCGS127N. gonorrhoeae GCGS113N. gonorrhoeae GCGS079N. gonorrhoeae GCGS239N. gonorrhoeae GCGS115N. gonorrhoeae GCGS177N. gonorrhoeae FA19N. gonorrhoeae GCGS102N. gonorrhoeae GCGS166N. gonorrhoeae GCGS053N. gonorrhoeae GCGS183N. gonorrhoeae GCGS019N. gonorrhoeae GCGS066N. gonorrhoeae GCGS143N. gonorrhoeae GCGS031N. gonorrhoeae GCGS163N. gonorrhoeae GCGS141N. gonorrhoeae GCGS205N. gonorrhoeae GCGS039N. gonorrhoeae GCGS051N. gonorrhoeae GCGS067N. gonorrhoeae GCGS133N. gonorrhoeae GCGS055N. gonorrhoeae GCGS195N. gonorrhoeae FA19N. gonorrhoeae GCGS075N. gonorrhoeae NCCP11945N. gonorrhoeae GCGS028N. gonorrhoeae GCGS124N. gonorrhoeae GCGS237N. gonorrhoeae GCGS097N. gonorrhoeae GCGS005N. gonorrhoeae GCGS117N. gonorrhoeae GCGS119N. gonorrhoeae GCGS241N. gonorrhoeae GCGS145N. gonorrhoeae GCGS048N. gonorrhoeae GCGS087N. gonorrhoeae GCGS085N. gonorrhoeae GCGS160N. gonorrhoeae GCGS165N. gonorrhoeae GCGS159N. gonorrhoeae GCGS129N. gonorrhoeae GCGS001N. gonorrhoeae GCGS027N. gonorrhoeae GCGS107N. gonorrhoeae GCGS157N. gonorrhoeae GCGS109N. gonorrhoeae GCGS125N. gonorrhoeae GCGS049N. gonorrhoeae GCGS151N. gonorrhoeae GCGS233N. gonorrhoeae GCGS035N. gonorrhoeae GCGS059N. gonorrhoeae GCGS155N. gonorrhoeae GCGS076N. gonorrhoeae GCGS057N. gonorrhoeae GCGS123N. gonorrhoeae GCGS147N. gonorrhoeae GCGS209N. gonorrhoeae GCGS171N. gonorrhoeae GCGS164N. gonorrhoeae GCGS054N. gonorrhoeae n01.08N. gonorrhoeae GCGS173N. gonorrhoeae GCGS071N. gonorrhoeae GCGS170N. gonorrhoeae GCGS056N. gonorrhoeae GCGS222N. gonorrhoeae GCGS213N. gonorrhoeae GCGS169N. gonorrhoeae GCGS200N. gonorrhoeae GCGS190N. gonorrhoeae GCGS236N. gonorrhoeae GCGS052N. gonorrhoeae GCGS234N. gonorrhoeae GCGS175N. gonorrhoeae GCGS197N. gonorrhoeae GCGS167N. gonorrhoeae GCGS203N. gonorrhoeae GCGS153N. gonorrhoeae GCGS201N. gonorrhoeae GCGS220N. gonorrhoeae GCGS007N. gonorrhoeae GCGS068N. gonorrhoeae GCGS030N. gonorrhoeae GCGS185N. gonorrhoeae GCGS235N. gonorrhoeae GCGS137N. gonorrhoeae GCGS020N. gonorrhoeae GCGS094N. gonorrhoeae GCGS130N. gonorrhoeae GCGS023N. gonorrhoeae GCGS232N. gonorrhoeae GCGS042N. gonorrhoeae GCGS132N. gonorrhoeae GCGS148N. gonorrhoeae GCGS006N. gonorrhoeae GCGS091N. gonorrhoeae GCGS003N. gonorrhoeae GCGS104N. gonorrhoeae GCGS122N. gonorrhoeae GCGS238N. gonorrhoeae GCGS070N. gonorrhoeae GCGS108N. gonorrhoeae GCGS196N. gonorrhoeae GCGS022N. gonorrhoeae GCGS034N. gonorrhoeae GCGS073N. gonorrhoeae GCGS194N. gonorrhoeae GCGS152N. gonorrhoeae GCGS080N. gonorrhoeae GCGS214N. gonorrhoeae GCGS182N. gonorrhoeae GCGS140N. gonorrhoeae GCGS004N. gonorrhoeae GCGS037N. gonorrhoeae GCGS187N. gonorrhoeae GCGS017N. gonorrhoeae GCGS025N. gonorrhoeae GCGS065N. gonorrhoeae GCGS139N. gonorrhoeae GCGS029N. gonorrhoeae GCGS045N. gonorrhoeae GCGS231N. gonorrhoeae GCGS077N. gonorrhoeae GCGS002N. gonorrhoeae GCGS121N. gonorrhoeae GCGS058N. gonorrhoeae GCGS134N. gonorrhoeae GCGS223N. gonorrhoeae GCGS149N. gonorrhoeae GCGS161N. gonorrhoeae GCGS215N. gonorrhoeae GCGS012N. gonorrhoeae GCGS207N. gonorrhoeae GCGS086N. gonorrhoeae GCGS081N. gonorrhoeae GCGS178N. gonorrhoeae GCGS105N. gonorrhoeae GCGS101N. gonorrhoeae GCGS046N. gonorrhoeae GCGS047N. gonorrhoeae GCGS099N. gonorrhoeae GCGS093N. gonorrhoeae GCGS158N. gonorrhoeae GCGS229N. gonorrhoeae GCGS041N. gonorrhoeae GCGS114N. gonorrhoeae GCGS179N. gonorrhoeae GCGS063N. gonorrhoeae GCGS043N. gonorrhoeae GCGS088N. gonorrhoeae GCGS021N. gonorrhoeae GCGS142N. gonorrhoeae GCGS092N. gonorrhoeae GCGS098N. gonorrhoeae GCGS032N. gonorrhoeae F62N. gonorrhoeae GCGS126N. gonorrhoeae MIA_2011_0510N. gonorrhoeae GCGS210N. gonorrhoeae GCGS040N. gonorrhoeae GCGS226N. gonorrhoeae GCGS212N. gonorrhoeae GCGS128N. gonorrhoeae ALB_2011_0102N. gonorrhoeae GCGS072N. gonorrhoeae MU_NG5N. gonorrhoeae GCGS162N. gonorrhoeae GCGS206N. gonorrhoeae GCGS116N. gonorrhoeae GCGS191N. gonorrhoeae GCGS217N. gonorrhoeae GCGS100N. gonorrhoeae GCGS026N. gonorrhoeae GCGS221N. gonorrhoeae GCGS106N. gonorrhoeae GCGS211N. gonorrhoeae GCGS078N. gonorrhoeae GCGS011N. gonorrhoeae 35/02N. gonorrhoeae FA6140N. gonorrhoeae GCGS184N. gonorrhoeae GCGS110N. gonorrhoeae GCGS082N. gonorrhoeae G2891N. gonorrhoeae GCGS050N. gonorrhoeae GCGS225N. gonorrhoeae GCGS219N. gonorrhoeae GCGS199N. gonorrhoeae GCGS168N. gonorrhoeae GCGS150N. gonorrhoeae GCGS193N. gonorrhoeae ATL_2011_0125N. gonorrhoeae SK36809N. gonorrhoeae SK32402N. gonorrhoeae GCGS224N. gonorrhoeae GCGS204N. gonorrhoeae GCGS174N. gonorrhoeae GCGS120N. gonorrhoeae i19.05N. gonorrhoeae GCGS064N. gonorrhoeae GCGS138N. gonorrhoeae GCGS156N. gonorrhoeae GCGS136N. gonorrhoeae GCGS192N. gonorrhoeae GCGS230N. gonorrhoeae GCGS144N. gonorrhoeae GCGS172N. gonorrhoeae GCGS083N. gonorrhoeae SK39420N. gonorrhoeae e03.04N. gonorrhoeae GCGS218N. gonorrhoeae GCGS024N. gonorrhoeae GCGS044N. gonorrhoeae SK23020N. gonorrhoeae MU_NG25N. gonorrhoeae ATL_2011_0508N. gonorrhoeae MIA_2011_0515N. gonorrhoeae SK33414N. gonorrhoeae GCGS154N. gonorrhoeae GCGS146N. gonorrhoeae GCGS096N. gonorrhoeae GCGS060N. gonorrhoeae GCGS228N. gonorrhoeae GCGS202N. gonorrhoeae GCGS188N. gonorrhoeae GCGS216N. gonorrhoeae GCGS176N. gonorrhoeae GCGS018N. gonorrhoeae GCGS186N. gonorrhoeae GCGS090N. gonorrhoeae GCGS118N. gonorrhoeae GCGS198N. gonorrhoeae GCGS008N. gonorrhoeae GCGS084N. gonorrhoeae SK22871N. gonorrhoeae GCGS010N. gonorrhoeae GCGS074N. gonorrhoeae GCGS208N. gonorrhoeae NG05N. gonorrhoeae GCGS180N. gonorrhoeae GCGS062N. gonorrhoeae GCGS009N. gonorrhoeae GCGS038N. gonorrhoeae SK7842N. gonorrhoeae NOR_2011_0306N. gonorrhoeae ATL_2011_0121N. gonorrhoeae SK29471N. gonorrhoeae SK29344N. gonorrhoeae SK6987N. gonorrhoeae SK8976N. gonorrhoeae SK12684N. gonorrhoeae MS11N. gonorrhoeae FA19N. gonorrhoeae SK16942N. gonorrhoeae SK28355N. gonorrhoeae SK17973N. gonorrhoeae CH811N. gonorrhoeae NG_869N. gonorrhoeae ATL_2011_0513N. gonorrhoeae SK708N. gonorrhoeae MIA_2011_0516N. gonorrhoeae SK1902N. gonorrhoeae MU_NG6N. gonorrhoeae SK15454N. gonorrhoeae MIA_2011_0310N. gonorrhoeae FA 1090N. gonorrhoeae m07.05N. meningitidis 98008N. meningitidis 93004N. meningitidis NM3081N. meningitidis 70012N. meningitidis 61106N. meningitidis M12611N. meningitidis NM576N. meningitidis CU385N. meningitidis NM2657N. meningitidis 96023N. meningitidis 63049N. meningitidis 2007056N. meningitidis 69166N. meningitidis 61103N. meningitidis 2001212N. meningitidis 70030N. meningitidis WUE 2594N. meningitidis NM3652N. meningitidis 65014N. meningitidis 2004090N. meningitidis NM3642N. meningitidis 68094N. meningitidis 69096N. meningitidis 63006N. meningitidis 97020N. meningitidis 63041N. meningitidis 88050N. meningitidis NM126N. meningitidis 97014N. meningitidis 12888N. meningitidis NM418N. meningitidis 8013N. meningitidis M13255N. meningitidis M20599N. meningitidis 77221N. meningitidis 9757N. meningitidis 2002038N. meningitidis NM255N. meningitidis NM422N. meningitidis 92045N. meningitidis NM586N. meningitidis M01240013N. meningitidis 4119N. meningitidis NM2244N. meningitidis NM1919N. meningitidis NM2935N. meningitidis NM2369N. meningitidis NM1362N. meningitidis NM1361N. meningitidis NM3129N. meningitidis NM2335N. meningitidis NM1895N. meningitidis NM1482N. meningitidis NM1891N. meningitidis NM1561N. meningitidis NM2382N. meningitidis NM1666N. meningitidis NM2814N. meningitidis NM1931N. meningitidis NM1360N. meningitidis NM2813N. meningitidis NM2181N. meningitidis NM1359N. meningitidis NM2008N. meningitidis NM1938N. meningitidis NM1963N. meningitidis NM2808N. meningitidis NM2237N. meningitidis NM2381N. meningitidis NM2431N. meningitidis NM2232N. meningitidis NM2025N. meningitidis NM2812N. meningitidis NM2193N. meningitidis NM1928N. meningitidis NM2206N. meningitidis NM1757N. meningitidis NM1264N. meningitidis NM2239N. meningitidis NM2432N. meningitidis NM2718N. meningitidis NM1578N. meningitidis NM2811N. meningitidis NM1544N. meningitidis NM2439N. meningitidis NM2188N. meningitidis NM2606N. meningitidis NM2389N. meningitidis NM2932N. meningitidis NM2433N. meningitidis NM1921N. meningitidis NM1837N. meningitidis NM2717N. meningitidis NM2700N. meningitidis NM2441N. meningitidis NM2810N. meningitidis NM2263N. meningitidis NM2332N. meningitidis NM233N. meningitidis Nm11003N. meningitidis K1207N. meningitidis S0108N. meningitidis NM80N. meningitidis alpha704N. meningitidis alpha14N. meningitidis NM220N. meningitidis NS44N. meningitidis 100514N. meningitidis NM2933N. meningitidis NM1779N. meningitidis NM1910N. meningitidis NM1364N. meningitidis NM1901N. meningitidis NM1471N. meningitidis NM2524N. meningitidis NM2394N. meningitidis NM1672N. meningitidis NM1826N. meningitidis NM1573N. meningitidis NM1893N. meningitidis NM1325N. meningitidis NM2701N. meningitidis NM1976N. meningitidis NM1550N. meningitidis NM1758N. meningitidis NM2009N. meningitidis NM1892N. meningitidis NM2602N. meningitidis NM2857N. meningitidis NM1363N. meningitidis NM2393N. meningitidis NM1845N. meningitidis NM1446N. meningitidis NM2264N. meningitidis NM2187N. meningitidis NM2617N. meningitidis NM2856N. meningitidis NM2228N. meningitidis NM2934N. meningitidis NM1805N. meningitidis NM3128N. meningitidis NM1937N. meningitidis NM1797N. meningitidis NM1549N. meningitidis NM1673N. meningitidis NM2032N. meningitidis NM1831N. meningitidis NM2254N. meningitidis Nm1140N. meningitidis ATCC 13091N. meningitidis 341215N. meningitidis S0108N. meningitidis 34173N. meningitidis Nm8663N. meningitidis NM36N. meningitidis NM3684N. meningitidis NM3688N. meningitidis 992008N. meningitidis NM3685N. meningitidis 100603N. meningitidis 130803N. meningitidis 330505N. meningitidis NM3680N. meningitidis 360624N. meningitidis 131148N. meningitidis 440902N. meningitidis 420718N. meningitidis 340552N. meningitidis 100601N. meningitidis Nm10259N. meningitidis Nm6756N. meningitidis NM1483N. meningitidis NM1829N. meningitidis NM51N. meningitidis 73704N. meningitidis NM165N. meningitidis NM3131N. meningitidis 2005079N. meningitidis M10208N. meningitidis M04240196N. meningitidis H44/76N. meningitidisN. meningitidis B6116/77N. meningitidis G2136N. meningitidis NM003N. meningitidis NM3223N. meningitidis 2004032N. meningitidis 220601N. meningitidis 100530N. meningitidis Nm6938N. meningitidis 321114N. meningitidis 421007N. meningitidis Z2491N. meningitidis 440501N. meningitidis Nm8187N. meningitidis Nm9418N. meningitidis 320503N. meningitidis Nm3127N. meningitidis NM2809N. meningitidis 2004085N. meningitidis 2000063N. meningitidis 64182N. meningitidis 69155N. meningitidis 70021N. meningitidis 97027N. meningitidis NM3141N. meningitidis 96037N. meningitidis NM151N. meningitidis NM477N. meningitidis NM90N. meningitidis NM3230N. meningitidis NM95N. meningitidis NM133N. meningitidis NM43N. meningitidis 2001073N. meningitidis NM3158N. meningitidis M13265N. meningitidis Neisseria meningitidis CHUVN. meningitidis Nm2732N. meningitidis NM23N. meningitidis NM1482N. meningitidis NM3139N. meningitidis 81858N. meningitidis DE9938N. meningitidisN. meningitidis 2001213N. meningitidis NM1482N. meningitidis 053442N. meningitidis NM762N. meningitidis M7124N. meningitidis MC58N. meningitidis alpha710N. meningitidis M0579N. meningitidis NM183N. meningitidis 97021N. meningitidis 2006087N. meningitidis NM3001N. meningitidis 9615945N. meningitidis N1568N. meningitidis NM174N. meningitidis ES14902N. meningitidis 93003N. meningitidis 9506N. meningitidis M13399N. meningitidis M6190N. meningitidis M7089N. meningitidis 98080N. meningitidis 2004264N. meningitidis 2001068N. meningitidis NM518N. meningitidis 2001001N. meningitidis NM35N. meningitidis NM3042N. meningitidis NM3222N. meningitidis NM27N. meningitidis NM94N. meningitidis 2008223N. meningitidis 2005172N. meningitidis NM604N. meningitidis 97008N. meningitidis 65012N. meningitidis 70082N. meningitidis 69100N. meningitidis 69176N. meningitidis 94018N. meningitidis NM606N. meningitidis 97018N. meningitidis DE9686N. meningitidis DE9622N. meningitidis H44/76N. meningitidis 370601N. meningitidis 100572N. meningitidis 311112N. meningitidis 320501N. meningitidis NM045N. meningitidis NM2795N. meningitidis NMBN. meningitidis M1412N. meningitidis 2000175N. meningitidis M7124N. meningitidis FAM18N. meningitidis NM3683N. meningitidis NM82N. meningitidis NM32N. meningitidis NM2781N. meningitidis 80179N. meningitidis LNP27256N. meningitidis OX99.30304N. meningitidis NM3681N. meningitidis NM3687N. meningitidis L91543N. meningitidis M01240149N. meningitidis M01240355N. meningitidis NM140N. meningitidis 87255N. meningitidis 2002004N. meningitidis NM1476N. meningitidis 2000081N. meningitidis 2007461N. meningitidis NM271N. meningitidis NM3164N. meningitidis NM2866N. meningitidis NM3147N. meningitidis 2003051N. meningitidis NM3144N. meningitidis NM115N. meningitidis 2002020N. meningitidis LNP21362N. meningitidis NM313N. meningitidis 2001072N. meningitidis 2005040N. meningitidis NM1495N. meningitidis NZ05/33N. meningitidis NM3686N. meningitidis NM3682N. meningitidis NM27N. meningitidis NM134N. meningitidis NM3173N. meningitidis 2002030N. meningitidis 73696N. meningitidis M0579N. meningitidis 96060N. meningitidis 98002N. meningitidis NM0552N. meningitidis 2002007N. meningitidis 63023N. meningitidis 2000080N. meningitidis 510612N. meningitidis 75689N. meningitidis NM607N. meningitidis 96024N. meningitidis 2003022N. meningitidis 98005N. meningitidis NM2033N. meningitidis 2003022N. meningitidis 75643N. mucosa ATCC 25996N. sicca 4320N. meningitidis 480_NMENN. sicca DS1N. sicca ATCC 29256N. meningitidis 1273_NMENN. lactamica 919_NLACN. meningitidis 378_NMENN. meningitidis 313_NMENN. macacae ATCC 33926N. sp. GT4A_CT1 GT4A_CT1N. sicca VK64N. meningitidis 379_NMENN. sp. HMSC06F02 HMSC06F02N. flavescens NRL30031/H210N. sp. oral taxon 014 F0314N. meningitidis 776_NMENN. lactamica 595_NLACN. lactamica 583_NLACN. meningitidis 1045_NMENN. meningitidis 840_NMENN. meningitidis 1044_NMENN. subflava NJ9703N. meningitidis 1210_NMENN. meningitidis 782_NMENN. mucosa C6AN. meningitidis 433_NMENN. flavescens SK114N. mucosa C102N. meningitidis 338.rep2_NMENN. lactamica 338.rep1_NLACN. meningitidis 768_NMENN. sp. KH1503 KH1503N. sp. 83E34 83E34N. shayeganii 871N. wadsworthii 9715N. meningitidis 491_NMENN. weaveri ATCC 51223N. weaveri LMG 5135N. meningitidis 1279_NMENN. meningitidis 431_NMENN. meningitidis 404_NMENN. elongata ATCC 29315N. elongata ATCC 29315N. sp. 74A18 74A18N. sp. oral taxon 020 F0370N. meningitidis 203_NMENN. lactamica 914_NLACN. bacilliformis ATCC BAA1200
0.00
0.25
0.50
0.75
1.00
ANIm
_alig
nmen
t_co
vera
ge
719 Neisseria spp. ANIm
Table of Contents
IntroductionThe Insidious Dickeya MenaceqPCR Primer Design From Whole Genomes
ClassificationClassification is CriticalWhole-Genome Classification
Soft-Rot EnterobacteriaA Tangled TaxonomyThe Cracks Are Showing. . .
ConclusionFinal ThoughtsWithout Whom. . .
If you’re interested. . .Hidden diversity: pre-publication
Final thoughts
• Diagnostics and large-scale analyses require accuratetaxonomic classification
• Historical collections/public databases have inaccuracies
• Bacterial taxonomy can be messy!
• Whole-genome classification with ANI works
• Retrospective sequencing and classification: clean things up
• Misclassification and hidden diversity may be difficult news formetagenomics
• Accurate MinION classification in-the-field is possible?
Recent papera
aPritchard et al. (2015) Anal. Methods doi:10.1039/c5ay02550h
pyani softwarea
ahttp://widdowquinn.github.io/pyani
• Python ANImodule
• Activedevelopment
• Preprintcoming soon(withsimulatedgenomes andlots ofbacterialtaxonomy!)
Acknowledgements
James Hutton InstituteEmma CampbellPeter CockNicola HoldenAshleigh HolmesSonia HumphrisPeter ThorpeIan TothNUI GalwayFlorence AbramFiona BrennanUniversity of AberdeenKen ForbesNorval Strachan
FeraValerie BertrandJohn ElphinstoneRachel GloverNeil ParkinsonUniversity of MunsterMartina BielaszewskaHelge KarchSASAGerry SaddlerGitHubBenjamin LeopoldMichael Robeson