Concurrent Acquisition of a Single Nucleotide Polymorphism in Diverse Influenza
H5N1 Clade 2.2 Sub-clades
Niman HL*, and Boynton BR
+
Recombinomics, Inc., Pittsburgh, Pennsylvania, USA*, U.S. Naval Medical
Research Unit 3 (NAMRU-3), Cairo, EGY
+
Department of Influenza Recombination, Recombinomics, Inc, 648 Field Club
Road, Pittsburgh, PA 15238, USA
Correspondence to Henry L Niman, E-mail:
henry_niman@recombinomics.com
617.877.0987, 412.963.1362 FAX
Text word count: 1695
3 Figures 1 Supplemental Table
Running Title: Concurrent Influenza H5N1 Acquisitions
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Highly pathogenic Influenza A H5N1 was first identified in Guangdong Province
in 1996, followed by human cases in Hong Kong in 1997
1,2
. The number of
confirmed human cases now exceeds 300 and the associated Case Fatality Rate
exceeds 60%
3
. The genetic diversity of the serotype continues to increase. Four
distinct clades or sub-clades have been linked to human cases
4-7
. The gradual
genetic changes identified in the sub-clades have been attributed to copy errors
by viral encoded polymerases that lack an editing function, thereby resulting in
antigenic drift
8
. We report here the concurrent acquisition of the same
polymorphism by multiple, genetically distinct, clade 2.2 sub-clades in Egypt,
Russia and Ghana. These changes are not easily explained by the current theory
of "random mutation" through copy error, and are more easily explained by
recombination with a common source. This conclusion is supported by additional
polymorphisms shared by clade 2.2 isolates in Egypt and Germany.
The study of influenza evolution in nature has been aided by the emergence of a
new strain (clade 2.2) first identified at Qinghai Lake in central China in the spring
of 2005. Sequencing of all eight genes
9,10
showed that isolates from migratory
waterfowl were easily distinguishable from previous isolates linked to poultry and
human infections in eastern and southeastern Asia
11,12
. The new strain was
subsequently found in outbreaks in Russia, Kazakhstan and Mongolia
13,14
. Prior
to these clade 2.2 outbreaks, the highly pathogenic Asian version of H5N1 had
never been reported west of China. The detection of H5N1 in migratory waterfowl
in the summer of 2005 in Russian and Mongolian migratory bird sanctuaries
signaled the start of a major geographical expansion of H5N1. In the following 12
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
months, almost 50 countries west of China reported H5N1 for the first time. All
infections were clade 2.2. The geographical reach includes Europe, the Middle
East and Africa.
This expansion offered a unique opportunity to study the evolution of H5N1 as it
migrated into new regions, including human cases in Turkey, Iraq, Azerbaijan,
Egypt, Djibouti in 2006 and Nigeria in 2007. Sequence analysis indicated all
cases were due to clade 2.2. The outbreaks were due to multiple introductions
and isolates had region specific polymorphisms.
The sequences also allowed for the monitoring of new acquisitions of
discriminating polymorphisms. These new acquisitions created additional sub-
clades defined in phylogenetic analysis. Moreover, close monitoring of these
changes could provide insight into the mechanisms underlying the rapid evolution
of H5N1 in general, and sub-clade 2.2 in particular.
We isolated H5N1 from patients and poultry in Egypt. The first poultry isolates
were collected February 2006, and the first human cases developed symptoms in
March 2006. Analysis of the H5N1 isolates collected between February and May
2006 defined a series of HA and NA regional markers. These markers from
Egypt were also found in the human case from Djibouti, as well as in poultry
isolates from Israel and Gaza. These isolates are listed in Supplemental Table 1.
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
After a lull in reported infections over the summer, H5N1 re-emerged in Egypt in
September 2006. The more recent isolates had the same regional markers seen
in the previous season. However, both poultry and human isolates had acquired
a series of new polymorphisms. Non-synonymous polymorphisms were identified
in samples collected from a cluster of three family members from the Gharbiya
governorate in the Nile Delta. HA gene polymorphisms were identified in or near
the receptor binding domain, including V223I and M230I, as well as the
oseltamivir resistance polymorphism, N294S, in the NA gene. The patients first
developed symptoms in December 2006 and all three infections were fatal (a
detailed report on patients and polymorphism tracing will be described
elsewhere).
Additional cases in early 2007 included HA sequences with a 3 BP deletion of the
nucleotides encoding Ser at position 133 (H3 Numbering), as well as a case with
a novel HA cleavage site, REGRRRKR. The changes were found in multiple
patients in central and southern Egypt. The above non-synonymous changes
were associated with additional synonymous and non-synonymous changes in
the HA and NA sequences that created additional sub-clades of sub-clade
2.2. However, the isolates from the 2006/2007 season maintained the regional
markers seen in early 2006 in isolates from Egypt, Djibouti, Israel and Gaza.
Chicken isolates from Gharbiya samples collected on February 15, 2007 included
one sequence that was closely related to the sequences from the human
Gharbiya cluster. The sequence from this isolate, A/chicken/1892N3-HK49/2007
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
(HK49), had the regional markers previously seen in the 2006 and 2007 isolates,
as well as HA non-synonymous changes, V223I and M230I. Additionally, it also
had an NA synonymous polymorphism, G743A, appended onto the genetic
background of the human Gharbiya cluster, as seen in the NA cladogram in
Figure 1. The location of the isolate on the tip of the branch indicated it was a
recent acquisition.
This polymorphism was also found in two additional chicken isolates,
A/chicken/1890N3-HK45/2007 and A/chicken/1891N3-CLEVB/2007, collected
the same week in the Gharbiya governorate, but these two isolates fell onto a
separate branch of the tree, indicating the same polymorphism had been
appended onto two distinct sub-clades. Moreover, plaque-purified sub-clones of
HK49 were isolated because the original sequence had mixed signals in the NA
and HA sequences. These HK49 sub-clones fell into two major groups. The NA
consensus sequences for the two groups had 11 differences that matched the 11
differences between the two sets of chicken sequences. The major species were
closely related to the sequences from the human cluster, while the minor species
were closely related to the two additional chicken sequences. However, all
sequenced, plaque-purified clones had G743A (data not shown).
The G743A was subsequently found in human isolates from patients who
developed symptoms in April 2007. Included were siblings with HA sequences
that had the 3 BP deletion seen in earlier patients from central Egypt. Like the
chicken sequences above, the G743A polymorphism was appended onto
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
sequences identified earlier in Egypt. Similarly, distinct sequences from another
patient, A/Egypt/2630-NAMRU3/2007, that acquired G743A, also fell onto a
separate branch.
The distinct branches displayed in the NA cladogram were also seen in the HA
cladogram in Figure 1B. The isolates with G743A are also located at the tips of
the branches, supporting a recent acquisition of the polymorphism.
In February 2007, an H5N1 clade 2.2 outbreak occurred near Moscow,
Russia. Isolates from infected chickens were most closely related to 2006
sequences from Azerbaijan. Figure 2A is an expanded cladogram with isolates
from Europe, the Middle East and western Africa. Like the acquisitions in Egypt,
the isolates with G743A, mapped onto the tip of a branch composed of earlier
isolates that did not have the acquisition.
Similarly, in April, 2007, an H5N1 clade 2.2 outbreak occurred near Tema,
Ghana. Sequences from three chickens were most closely related to turkey
isolates collected in December 2006 in the Ivory Coast. Like the Egypt and
Moscow isolates above, the recent isolates with G743A mapped to the tip of a
branch containing earlier isolates that did not have the G743A polymorphism.
Additional HA polymorphisms are noted in the HA phylogram in Figure 2B.
Isolates that had the NA polymorphism, G743A, also had a synonymous HA
polymorphism, C689T. This polymorphism was also in human and bird isolates
from the Nile Delta (see Supplemental Table 1). Another polymorphism, G754A,
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
that encodes M230I is in one of the German isolates
15
, A/eagle
owl/Germany/R166/2006, and maps to another branch with Egyptian human and
poultry isolates from the Nile Delta. A third polymorphism, C1614T, that encodes
T517I, is in another German isolate, A/mute swan/Germany/R797/2006, and
another branch with human isolates from southern Egypt. The isolates also have
the novel HA cleavage site initially found in whooper swan isolates in Mongolia in
2005. The polymorphisms found in German isolates in 2006 were in Russian
clade 2.2 isolates in 2005.
The NA G743A polymorphism can be traced through public H5N1 sequences.
Isolates with full NA sequences are listed in the NA phylogram in Figure 3. The
polymorphism was identified in the first reported sequences linked to the spread
of H5N1 in Asia in 2003/2004 in South Korea
16
and Japan
17
. The polymorphism
was subsequently identified in clade 1 isolates in southeast Asia, as well as clade
2.1 isolates in Indonesia and clade 2.3 isolates in China. The first reported clade
2.2 isolates were in wild birds in Germany (see Figure 2) collected in February
2006. The isolates in Germany formed distinct HA and NA branches due to a
series of regional markers in these isolates.
The concurrent acquisition of the same polymorphism by multiple sub-clades
challenges the current theory of influenza evolution that invokes random
mutations as a mechanism for the generation of antigenic drift. The isolates with
the newly acquired polymorphisms map to the tips of the branches of the
phylogenetic trees, indicating the acquisitions were recent. All discussed isolates
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
on the tips of the branches were collected over a short time frame between
February and April 2007, and in three geographically distinct regions. These data
do not support a common progenitor sequence because the most closely related
sequences to each of the respective recent isolates do not have this
change. Similarly, concurrent mutation / selection by eleven isolates that map to
six branches in three countries and were collected over a short time frame is also
unlikely.
An alternative explanation for the appending of these sequences is through
homologous recombination between closely related sequences. The newly
acquired polymorphisms in the recent isolates in Egypt are readily found in
recent H5N1 isolates, as noted in the three HA examples previously described
above. Moreover, the G743A polymorphism is found in these genetically distinct
sequences collected over a short time frame.
Polymorphism tracing demonstrates that most of the newly acquired
polymorphisms can be traced to the same serotype identified recently at
locations that are linked together by migratory bird flyways, raising the possibility
that the distribution and acquisition of the polymorphism is linked to
recombination between H5N1 sequences transmitted and transported by
migratory birds.
The individual polymorphisms recombine, generating sequences that create
antigenic drift. Mapping of these pathways and associations may be used to
develop novel vaccine targets representing rapidly evolving genomes.
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 1 NA and HA Phylograms of Egyptian Isolates
A. NA phylogram of positions 43-1337. Isolates with G743A marked with red
arrows. NA regional markers are C150T, C236T, A703G, A740G, T1088C and
G1280A.
Accession numbers and additional isolates with G743A listed in Table
S1.
B. HA phylogram of positions 93-1688. Isolates with G743A marked with red
arrows. HA regional markers are G196A, G466A, G496A, C661T, C727T,
C779T, A878G, C937T, G1018T, C1261T and C1686T.
Egyptian isolates, accession numbers, collection date and governorate in Table
S1. Trees generated using neighbor joining with 100 bootstrap repetitions.
Sequences generated as described previously
18
.
Figure 2 NA and HA phylogram of Clade 2.2 isolates.
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
A. NA phylogram of positions 43-1337. Isolates with G743A marked with red
arrows.
Accession numbers and additional isolates with G743A listed in Table
S1.
B. HA phylogram of positions 93-1688. Isolates with C628T marked with blue
arrows / bars. Isolates with G754A marked with green arrows / bars. Isolates
with C1614T marked with orange arrows / bars.
Accession numbers and additional isolates with HA polymorphisms listed in
Table S1
Figure 3 NA Phylogram of H5N1 isolates with G743A
Accession numbers and additional isolates with G743A listed in Table S1.
Table S1 Isolates and Accession Numbers
Names and accession numbers of HA and NA sequences used in figures 1-3.
Partial sequences
19-26
with polymorphisms in Figures 1-3 are listed.
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Acknowledgements
We thank
Magdi D. Saad, Jeff A. Tjaden, Ken C. Earhart, Marshall R. Monteville,
Moustafa M. Mansour, Elassai Labib, EA Ayoub from the US Naval Medical
Research Unit 3, Mona Aly and A-SA Arasa from the Central Laboratory from
Veterinary Quality Control, and Al Nayel, AS Abdelghani, Hala Essmat from
Ministry of Health, Arab Republic of Egypt, for the collection of human and animal
samples, and isolation and sequencing of H5N1.
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Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 1A NA Egypt G743A
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 1B HA Egypt G743A
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 2A NA Phylogenetic Tree
G743A
G743A
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 2B HA Phylogenetic Tree
C689T
C689T
C1614T
C1614
T
G754A
C689T
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Figure 3 NA G743A
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007
Table S1 H5N1 Isolates and Accession Numbers
Accession
Name
Year Goverate
Sero
Figure
EF624253
A/chicken/Ghana/3158-NAMRU3/2007
2007 Tema
H5N1 HA
2B
EF624254
A/chicken/Ghana/3159-NAMRU3/2007
2007 Tema
H5N1 HA
2B
EF624255
A/chicken/Ghana/3160-NAMRU3/2007
2007 Tema
H5N1 HA
2B
EF441277
A/chicken/Egypt/1079-NAMRU3/2007
2007 Beni Suef
H5N1 HA
1B, 2B
EF469650
A/chicken/Egypt/1129N3-HK9/2007
2007 Fayoum
H5N1 HA
1B, 2B
EF441280
A/chicken/Egypt/1300-NAMRU3/2007
2007 Gharbiya
H5N1 HA
1B, 2B
EF469653
A/chicken/Egypt/1889N3-SM26/2007
2007 Giza
H5N1 HA
1B, 2B
EF469654
A/chicken/Egypt/1890N3-HK45/2007
2007 Gharbiya
H5N1 HA
1B, 2B
EF469659
A/chicken/Egypt/1891N3-CLEVB/2007
2007 Gharbiya
H5N1 HA
1B, 2B
EF469660
A/chicken/Egypt/1892N3-HK49/2007
2007 Gharbiya
H5N1 HA
1B, 2B
EF441281
A/duck/Egypt/1301-NAMRU3/2007
2007 Gharbiya
H5N1 HA
1B, 2B
EF469657
A/duck/Egypt/1888N3-SM25/2007
2007 Giza
H5N1 HA
1B, 2B
EF382359
A/Egypt/0636-NAMRU3/2007
2007 Beni Suef
H5N1 HA
1B, 2B
EF535817
A/Egypt/1394-NAMRU3/2007
2007 Fayoum
H5N1 HA
1B, 2B
EF535818
A/Egypt/1604-NAMRU3/2007
2007 Fayoum
H5N1 HA
1B, 2B
EF535819
A/Egypt/1731-NAMRU3/2007
2007 Sharkia
H5N1 HA
1B, 2B
EF535820
A/Egypt/1902-NAMRU3/2007
2007 Dakahlia
H5N1 HA
1B, 2B
EF535821
A/Egypt/2256-NAMRU3/2007
2007 Dakahlia
H5N1 HA
1B, 2B
EF535822
A/Egypt/2321-NAMRU3/2007
2007 Aswan
H5N1 HA
1B, 2B
EF535823
A/Egypt/2331-NAMRU3/2007
2007 Aswan
H5N1 HA
1B, 2B
EF535824
A/Egypt/2616-NAMRU3/2007
2007 Aswan
H5N1 HA
1B, 2B
EF535825
A/Egypt/2620-NAMRU3/2007
2007 Menia
H5N1 HA
1B, 2B
EF535826
A/Egypt/2621-NAMRU3/2007
2007 Qena
H5N1 HA
1B, 2B
A/Egypt/2629-NAMRU3/2007
2007 Qena
H5N1 HA
1B, 2B
A/Egypt/2630-NAMRU3/2007
2007 Sohag
H5N1 HA
1B, 2B
A/Egypt/2631-NAMRU3/2007
2007 Qalubiea
H5N1 HA
1B, 2B
A/Egypt/2750-NAMRU3/2007
2007 Menia
H5N1 HA
1B, 2B
A/Egypt/2651-NAMRU3/2007
2007 Shoubra
H5N1 HA
1B, 2B
EF441276
A/chicken/Egypt/1078-NAMRU3/2006
2006 Menoufiya
H5N1 HA
2B
EF441278
A/chicken/Egypt/1080-NAMRU3/2006
2006 Damietta
H5N1 HA
2B
EF441279
A/chicken/Egypt/1081-NAMRU3/2006
2006 Gharbiya
H5N1 HA
2B
EF042622
A/chicken/Egypt/10845-NAMRU3/2006
2006
H5N1 HA
2B
EF469651
A/chicken/Egypt/12378N3-CLEVB/2006
2006 Gharbiya
H5N1 HA
2B
EF469652
A/chicken/Egypt/12379N3-CLEVB/2006
2006 Sharqiya
H5N1 HA
2B
DQ837587
A/chicken/Egypt/5610NAMRU3-F3/2006
2006 Fayoum
H5N1 HA
2B
DQ837588
A/chicken/Egypt/5611NAMRU3-AN/2006
2006 Cairo
H5N1 HA
2B
DQ837589
A/chicken/Egypt/5612NAMRU3-S/2006
2006 Giza
H5N1 HA
2B
DQ447199
A/chicken/Egypt/960N3-004/2006
2006
H5N1 HA
2B
DQ666146
A/Djibouti/5691NAMRU3/2006
2006 Djibouti
H5N1 HA
2B
EF469655
A/duck/Egypt/12380N3-CLEVB/2006
2006 Gharbiya
H5N1 HA
2B
EF469656
A/duck/Egypt/13010N3-CLEVB/2006
2006 Menoufiya
H5N1 HA
1B, 2B
EF061116
A/Egypt/12374-NAMRU3/2006
2006 Gharbiya
H5N1 HA
1B, 2B
EF200512
A/Egypt/14724-NAMRU3/2006
2006 Gharbiya
H5N1 HA
1B, 2B
EF200513
A/Egypt/14725-NAMRU3/2006
2006 Gharbiya
H5N1 HA
1B, 2B
EF042614
A/Egypt/2763-NAMRU3/2006
2006 Qaliubiya
H5N1 HA
2B
DQ464377
A/Egypt/2782-NAMRU3/2006
2006 Qaliubiya
H5N1 HA
2B
EF042615
A/Egypt/2783-NAMRU3/2006
2006 Qaliubiya
H5N1 HA
2B
EF042616
A/Egypt/2786-NAMRU3/2006
2006 Gharbiya
H5N1 HA
2B
EF042617
A/Egypt/2947-NAMRU3/2006
2006 Kafr El-Sheikh H5N1 HA
2B
EF042618
A/Egypt/3105-NAMRU3/2006
2006 Sohaj
H5N1 HA
2B
EF042619
A/Egypt/3458-NAMRU3/2006
2006 Minufiyah
H5N1 HA
2B
EF042620
A/Egypt/5494-NAMRU3/2006
2006 Sharabeya
H5N1 HA
2B
EF042621
A/Egypt/5614-NAMRU3/2006
2006 Al Minya
H5N1 HA
2B
EF469658
A/goose/Egypt/13009N3-SM2/2006
2006 Menoufiya
H5N1 HA
2B
DQ837590
A/turkey/Egypt/5613NAMRU3-T/2006
2006 Menoufiya
H5N1 HA
2B
EF042624
A/teal/Egypt/14051-NAMRU3/2005
2005 Damietta
H5N1 HA
2B
EF624250
A/chicken/Ghana/3158-NAMRU3/2007
2007 Tema
H5N1 NA
2A, 3
EF624251
A/chicken/Ghana/3159-NAMRU3/2007
2007 Tema
H5N1 NA
2A, 3
EF624252
A/chicken/Ghana/3160-NAMRU3/2007
2007 Tema
H5N1 NA
2A, 3
A/chicken/Egypt/1079-NAMRU3/2007
2007 Beni Suef
H5N1 NA
1A, 2A
EF486240
A/chicken/Egypt/1129N3-HK9/2007
2007 Fayoum
H5N1 NA
1A, 2A
A/chicken/Egypt/1300-NAMRU3/2007
2007 Gharbiya
H5N1 NA
1A, 2A
EF486243
A/chicken/Egypt/1889N3-SM26/2007
2007 Giza
H5N1 NA
1A, 2A
EF486244
A/chicken/Egypt/1890N3-HK45/2007
2007 Gharbiya
H5N1 NA 1A, 2A, 3
EF486245
A/chicken/Egypt/1891N3-CLEVB/2007
2007 Gharbiya
H5N1 NA 1A, 2A, 3
EF486246
A/chicken/Egypt/1892N3-HK49/2007
2007 Gharbiya
H5N1 NA 1A, 2A, 3
A/duck/Egypt/1301-NAMRU3/2007
2007 Gharbiya
H5N1 NA
1A, 2A
EF486249
A/duck/Egypt/1888N3-SM25/2007
2007 Giza
H5N1 NA
1A, 2A
EF382360
A/Egypt/0636-NAMRU3/2007
2007 Beni Suef
H5N1 NA
1A, 2A
A/Egypt/1394-NAMRU3/2007
2007 Fayoum
H5N1 NA
1A, 2A
A/Egypt/1604-NAMRU3/2007
2007 Fayoum
H5N1 NA
1A, 2A
A/Egypt/1731-NAMRU3/2007
2007 Sharkia
H5N1 NA
1A, 2A
A/Egypt/1902-NAMRU3/2007
2007 Dakahlia
H5N1 NA
1A, 2A
A/Egypt/2256-NAMRU3/2007
2007 Dakahlia
H5N1 NA
1A, 2A
A/Egypt/2321-NAMRU3/2007
2007 Aswan
H5N1 NA
1A, 2A
A/Egypt/2331-NAMRU3/2007
2007 Aswan
H5N1 NA
1A, 2A
A/Egypt/2616-NAMRU3/2007
2007 Aswan
H5N1 NA
1A, 2A
A/Egypt/2620-NAMRU3/2007
2007 Menia
H5N1 NA
1A, 2A
A/Egypt/2621-NAMRU3/2007
2007 Qena
H5N1 NA 1A, 2A, 3
A/Egypt/2629-NAMRU3/2007
2007 Qena
H5N1 NA 1A, 2A, 3
A/Egypt/2630-NAMRU3/2007
2007 Sohag
H5N1 NA 1A, 2A, 3
A/Egypt/2631-NAMRU3/2007
2007 Qalubiea
H5N1 NA
1A, 2A
A/Egypt/2750-NAMRU3/2007
2007 Menia
H5N1 NA
1A, 2A
A/Egypt/2651-NAMRU3/2007
2007 Shoubra
H5N1 NA
1A, 2A
EF486241
A/chicken/Egypt/12378N3-CLEVB/2006
2006 Gharbiya
H5N1 NA
1A, 2A
EF486242
A/chicken/Egypt/12379N3-CLEVB/2006
2006 Sharqiya
H5N1 NA
1A, 2A
EF486247
A/duck/Egypt/12380N3-CLEVB/2006
2006 Gharbiya
H5N1 NA
1A, 2A
EF486248
A/duck/Egypt/13010N3-CLEVB/2006
2006 Menoufiya
H5N1 NA
1A, 2A
EF222324
A/Egypt/12374-NAMRU3/2006
2006 Gharbiya
H5N1 NA
1A, 2A
EF222323
A/Egypt/14724-NAMRU3/2006
2006 Gharbiya
H5N1 NA
1A, 2A
A/Egypt/14724a-NAMRU3/2006
2006 Gharbiya
H5N1 NA
1A, 2A
EF222322
A/Egypt/14725-NAMRU3/2006
2006 Gharbiya
H5N1 NA
1A, 2A
A/Egypt/14725a-NAMRU3/2006
2006 Gharbiya
H5N1 NA
1A, 2A
A/Egypt/2991-NAMRU3/2006
2006 Kafr El-Sheikh H5N1 NA
1A, 2A
EF486250
A/goose/Egypt/13009N3-SM2/2006
2006 Menoufiya
H5N1 NA
1A, 2A
EF474450
A/chicken/Moscow/2/2007
2007
H5N1 HA
2B
I
SDN230945 A/Nigeria/6e/07
2007
H5N1 HA
2B
I
SDN140057 A/Azerbaijan/001-161/2006
2006
H5N1 HA
2B
I
SDN140058 A/Azerbaijan/002-115/2006
2006
H5N1 HA
2B
I
SDN140054 A/Azerbaijan/006-207/2006
2006
H5N1 HA
2B
I
SDN140055 A/Azerbaijan/008-208/2006
2006
H5N1 HA
2B
I
SDN140056 A/Azerbaijan/011-162/2006
2006
H5N1 HA
2B
EF165057
A/buzzard/Bavaria/13/2006
2006
H5N1 HA
2B
EF165049
A/buzzard/Bavaria/5/2006
2006
H5N1 HA
2B
AM403474
A/Canada goose/Germany/R1207/06
2006
H5N1 HA
2B
AM403461
A/Canada goose/Germany/R71/06
2006
H5N1 HA
2B
EF395844
A/cat/Austria/649/2006
2006
H5N1 HA
2B
DQ864720
A/cat/Dagestan/87/06
2006
H5N1 HA
2B
DQ643982
A/cat/Germany/606/2006
2006
H5N1 HA
2B
AM403468
A/cat/Germany/R606/06
2006
H5N1 HA
2B
DQ864715
A/chicken/Adygea/203/06
2006
H5N1 HA
2B
AM400974
A/chicken/Burkina Faso/01.03/2006
2006
H5N1 HA
2B
AM400973
A/chicken/Burkina Faso/13.1/2006
2006
H5N1 HA
2B
CY016811
A/chicken/Cote d'Ivoire/1787-34/2006
2006
H5N1 HA
2B
EF532628
A/chicken/Gaza/450/2006
2006
H5N1 HA
1B, 2B
EF532630
A/chicken/Gaza/713/2006
2006
H5N1 HA
1B, 2B
EF532631
A/chicken/Gaza/714/2006
2006
H5N1 HA
1B, 2B
EF532626
A/chicken/Israel/397/2006
2006
H5N1 HA
1B, 2B
EF532629
A/chicken/Israel/625/2006
2006
H5N1 HA
1B, 2B
CY021517
A/chicken/Ivory Coast/1787-35/2006
2006
H5N1 HA
2B
DQ673901
A/chicken/Ivory Coast/1787/2006
2006
H5N1 HA
2B
DQ676834
A/chicken/Krasnodar/01/2006
2006
H5N1 HA
2B
EF205159
A/chicken/Krasnodar/123/06
2006
H5N1 HA
2B
DQ864718
A/chicken/Krasnodar/199/06
2006
H5N1 HA
2B
AM262541
A/chicken/Lagos.NIE/BA209/2006
2006
H5N1 HA
2B
AM262542
A/chicken/Lagos.NIE/BA210/2006
2006
H5N1 HA
2B
AM262543
A/chicken/Lagos.NIE/BA211/2006
2006
H5N1 HA
2B
AM262546
A/chicken/Lagos.NIE/SO300/2006
2006
H5N1 HA
2B
AM262547
A/chicken/Lagos.NIE/SO452/2006
2006
H5N1 HA
2B
AM262553
A/chicken/Lagos.NIE/SO493/2006
2006
H5N1 HA
2B
AM262572
A/chicken/Lagos.NIE/SO494/2006
2006
H5N1 HA
2B
DQ838517
A/chicken/Niger/2130-7/2006
2006
H5N1 HA
2B
DQ838516
A/chicken/Niger/2130-8/2006
2006
H5N1 HA
2B
CY016939
A/chicken/Nigeria/1047-30/2006
2006
H5N1 HA
2B
CY016947
A/chicken/Nigeria/1047-34/2006
2006
H5N1 HA
2B
CY016923
A/chicken/Nigeria/1047-54/2006
2006
H5N1 HA
2B
CY016931
A/chicken/Nigeria/1047-62/2006
2006
H5N1 HA
2B
CY016907
A/chicken/Nigeria/1047-8/2006
2006
H5N1 HA
2B
CY016276
A/chicken/Nigeria/641/2006
2006
H5N1 HA
2B
CY016284
A/chicken/Nigeria/957-20/2006
2006
H5N1 HA
2B
AM400975
A/chicken/Nigeria/SO227/2006
2006
H5N1 HA
2B
AM400976
A/chicken/Nigeria/SO294/2006
2006
H5N1 HA
2B
AM400977
A/chicken/Nigeria/SO457/2006
2006
H5N1 HA
2B
AM400978
A/chicken/Nigeria/SO461/2006
2006
H5N1 HA
2B
AM400979
A/chicken/Nigeria/SO470/2006
2006
H5N1 HA
2B
AM400980
A/chicken/Nigeria/SO478/2006
2006
H5N1 HA
2B
AM400981
A/chicken/Nigeria/SO485/2006
2006
H5N1 HA
2B
CY016300
A/chicken/Sudan/1784-10/2006
2006
H5N1 HA
2B
CY016292
A/chicken/Sudan/1784-7/2006
2006
H5N1 HA
2B
CY020661
A/chicken/Sudan/1784-8/2006
2006
H5N1 HA
2B
DQ862003
A/chicken/Sudan/1784/2006
2006
H5N1 HA
2B
CY021389
A/chicken/Sudan/2115-10/2006
2006
H5N1 HA
2B
CY020677
A/chicken/Sudan/2115-12/2006
2006
H5N1 HA
2B
CY020669
A/chicken/Sudan/2115-9/2006
2006
H5N1 HA
2B
DQ864719
A/chicken/Volgograd/236/06
2006
H5N1 HA
2B
DQ659679
A/common bussard/Bavaria/2/2006
2006
H5N1 HA
2B
EF165055
A/common buzzard/Bavaria/11/2006
2006
H5N1 HA
2B
AM403463
A/common buzzard/Germany/R306/06
2006
H5N1 HA
2B
AM408213
A/common buzzard/Germany/R870/06
2006
H5N1 HA
2B
EF110519
A/common coot/Switzerland/V544/06
2006
H5N1 HA
2B
EF165051
A/common pochard/Bavaria/7/2006
2006
H5N1 HA
2B
AM403469
A/coot/Germany/R655/06
2006
H5N1 HA
2B
AM408209
A/cormorant/Germany/R292/06
2006
H5N1 HA
2B
CY016779
A/Cygnus cygnus/Iran/754/2006
2006
H5N1 HA
2B
DQ515984
A/Cygnus olor/Czech Republic/5170/2006
2006
H5N1 HA
2B
CY017035
A/Cygnus olor/Italy/742/2006
2006
H5N1 HA
2B
CY020349
A/cygnus olor/Italy/808/2006
2006
H5N1 HA
2B
DQ435200
A/domestic cat/Iraq/820/2006
2006
H5N1 HA
2B
DQ435201
A/domestic goose/Iraq/812/2006
2006
H5N1 HA
2B
CY016803
A/duck/Cote d'Ivoire/1787-18/2006
2006
H5N1 HA
2B
EF532632
A/duck/Gaza/760/2006
2006
H5N1 HA
1B, 2B
EF532622
A/duck/Gaza/834/2006
2006
H5N1 HA
1B, 2B
AM403464
A/duck/Germany/R338/06
2006
H5N1 HA
2B
AM403466
A/duck/Germany/R592/06
2006
H5N1 HA
2B
AM403467
A/duck/Germany/R603/06
2006
H5N1 HA
2B
AM403470
A/duck/Germany/R751/06
2006
H5N1 HA
2B
CY017027
A/duck/Niger/914/2006
2006
H5N1 HA
2B
EF165054
A/eagle owl/Bavaria/10/2006
2006
H5N1 HA
2B
AM403473
A/eagle owl/Germany/R1166/06
2006
H5N1 HA
2B
EF165059
A/falcon/Bavaria/15/2006
2006
H5N1 HA
2B
AM408211
A/falcon/Germany/R899/06
2006
H5N1 HA
2B
EF165063
A/goldeneye duck/Bavaria/19/2006
2006
H5N1 HA
2B
EF165062
A/goosander/Bavaria/18/2006
2006
H5N1 HA
2B
EF165064
A/goosander/Bavaria/20/2006
2006
H5N1 HA
2B
EF110518
A/goosander/Switzerland/V82/06
2006
H5N1 HA
2B
AM408210
A/goose/Germany/R696/06
2006
H5N1 HA
2B
EF165066
A/great crested grebe/Bavaria/22/2006
2006
H5N1 HA
2B
AM408212
A/great crested grebe/Germany/R1226/06
2006
H5N1 HA
2B
CY017179
A/guinea fowl/Nigeria/957-12/2006
2006
H5N1 HA
2B
AM408215
A/gull/Germany/R882/06
2006
H5N1 HA
2B
AM400971
A/hooded vulture/Burkina Faso/1/2006
2006
H5N1 HA
2B
AM400972
A/hooded vulture/Burkina Faso/2/2006
2006
H5N1 HA
2B
EF090648
A/hooded vulture/Burkina Faso/5346-10/2006
2006
H5N1 HA
2B
DQ458992
A/mallard/Bavaria/1/2006
2006
H5N1 HA
2B
DQ449031
A/mallard/Italy/332/2006
2006
H5N1 HA
2B
CY016795
A/mallard/Italy/835/2006
2006
H5N1 HA
2B
EF165056
A/mute swan/Bavaria/12/2006
2006
H5N1 HA
2B
AM403460
A/mute swan/Germany/R65/06
2006
H5N1 HA
2B
AM403471
A/mute swan/Germany/R854/06
2006
H5N1 HA
2B
CY016915
A/ostrich/Nigeria/1047-25/2006
2006
H5N1 HA
2B
AM492165
A/stone marten/Germany/R747/2006
2006
H5N1 HA
2B
AM403475
A/stork/Germany/R1239/06
2006
H5N1 HA
2B
EF395845
A/swan/Austria/216/2006
2006
H5N1 HA
2B
EF165058
A/swan/Bavaria/14/2006
2006
H5N1 HA
2B
EF165060
A/swan/Bavaria/16/2006
2006
H5N1 HA
2B
EF165061
A/swan/Bavaria/17/2006
2006
H5N1 HA
2B
EF165065
A/swan/Bavaria/21/2006
2006
H5N1 HA
2B
EF165050
A/swan/Bavaria/6/2006
2006
H5N1 HA
2B
DQ464354
A/swan/Germany/R65/2006
2006
H5N1 HA
2B
DQ440535
A/swan/Iran/754/2006
2006
H5N1 HA
2B
CY017043
A/swan/Slovenia/760/2006
2006
H5N1 HA
2B
EF165048
A/tufted duck/Bavaria/4/2006
2006
H5N1 HA
2B
EF165052
A/tufted duck/Bavaria/8/2006
2006
H5N1 HA
2B
EF165053
A/tufted duck/Bavaria/9/2006
2006
H5N1 HA
2B
AM408216
A/tufted duck/Germany/R1240/06
2006
H5N1 HA
2B
AM403472
A/turkey/Germany/R1077/06
2006
H5N1 HA
2B
EF532623
A/turkey/Israel/345/2006
2006
H5N1 HA
1B, 2B
EF532624
A/turkey/Israel/364/2006
2006
H5N1 HA
1B, 2B
EF532625
A/turkey/Israel/365/2006
2006
H5N1 HA
1B, 2B
EF532627
A/turkey/Israel/446/2006
2006
H5N1 HA
1B, 2B
CY020693
A/turkey/Ivory Coast/4372-2/2006
2006
H5N1 HA
2B
CY020701
A/turkey/Ivory Coast/4372-3/2006
2006
H5N1 HA
2B
CY020709
A/turkey/Ivory Coast/4372-4/2006
2006
H5N1 HA
2B
AM403462
A/whooper swan/Germany/R88/06
2006
H5N1 HA
2B
AB263752
A/whooper swan/Mongolia/2/06
2006
H5N1 HA
2B
DQ449632
A/chicken/Kurgan/05/2005
2005
H5N1 HA
2B
DQ323672
A/chicken/Kurgan/3/2005
2005
H5N1 HA
2B
EF205155
A/chicken/Omsk/14/05
2005
H5N1 HA
2B
EF205154
A/chicken/Suzdalka/06/05
2005
H5N1 HA
2B
DQ864716
A/chicken/Tambov/570-2/05
2005
H5N1 HA
2B
EF205160
A/chicken/Tula/4/05
2005
H5N1 HA
2B
DQ449640
A/duck/Kurgan/08/2005
2005
H5N1 HA
2B
DQ864711
A/duck/Novosibirsk/02/05
2005
H5N1 HA
2B
DQ230522
A/duck/Novosibirsk/56/2005
2005
H5N1 HA
2B
DQ676840
A/goose/Krasnoozerka/627/2005
2005
H5N1 HA
2B
EF205156
A/goose/Suzdalka/10/05
2005
H5N1 HA
2B
DQ230521
A/grebe/Novosibirsk/29/2005
2005
H5N1 HA
2B
EF042624
A/teal/Egypt/14051-NAMRU3/2005
2005
H5N1 HA
2B
EF205158
A/turkey/Suzdalka/12/05
2005
H5N1 HA
2B
DQ407519
A/turkey/Turkey/1/2005
2005
H5N1 HA
2B
I
SDN138121 A/whooper swan/Mongolia/13/2005
2005
H5N1 HA
2B
AB233320
A/whooper swan/Mongolia/3/05
2005
H5N1 HA
2B
AB233321
A/whooper swan/Mongolia/4/05
2005
H5N1 HA
2B
AB233322
A/whooper swan/Mongolia/6/05
2005
H5N1 HA
2B
I
SDN138113 A/whooper swan/Mongolia/7/2005
2005
H5N1 HA
2B
I
SDN138105 A/whooper swan/Qinghai/1/2005
2005
H5N1 HA
2B
DQ864721
A/wild duck/Omsk/103-01/05
2005
H5N1 HA
2B
EF474448
A/chicken/Moscow/2/2007
2007
H5N1 NA
2A, 3
EF447431
A/chicken/Russia_Moscow oblast_Odintsovo/1/2007
2007
H5N1 NA
G743A
I
SDN230944 A/Nigeria/6e/07
2007
H5N1 NA
2A
I
SDN140072 A/Azerbaijan/001-161/2006
2006
H5N1 NA
2A
I
SDN140073 A/Azerbaijan/002-115/2006
2006
H5N1 NA
2A
I
SDN140069 A/Azerbaijan/006-207/2006
2006
H5N1 NA
2A
I
SDN140070 A/Azerbaijan/008-208/2006
2006
H5N1 NA
2A
I
SDN140071 A/Azerbaijan/011-162/2006
2006
H5N1 NA
2A
EF165579
A/buzzard/Bavaria/13/2006
2006
H5N1 NA
2A, 3
EF165572
A/buzzard/Bavaria/5/2006
2006
H5N1 NA
2A, 3
DQ643984
A/cat/Germany/606/2006
2006
H5N1 NA
2A
DQ835390
A/cat/Iraq/820NAMRU3/2006
2006
H5N1 NA
2A
CY016813
A/chicken/Cote d'Ivoire/1787-34/2006
2006
H5N1 NA
2A
EF532639
A/chicken/Gaza/450/2006
2006
H5N1 NA
1A, 2A
EF532641
A/chicken/Gaza/713/2006
2006
H5N1 NA
1A, 2A
EF532642
A/chicken/Gaza/714/2006
2006
H5N1 NA
1A, 2A
EF532637
A/chicken/Israel/397/2006
2006
H5N1 NA
1A, 2A
EF532640
A/chicken/Israel/625/2006
2006
H5N1 NA
1A, 2A
CY021519
A/chicken/Ivory Coast/1787-35/2006
2006
H5N1 NA
2A
DQ676836
A/chicken/Krasnodar/01/2006
2006
H5N1 NA
2A
EF205173
A/chicken/Krasnodar/123/06
2006
H5N1 NA
2A
DQ676832
A/chicken/Mahachkala/05/2006
2006
H5N1 NA
2A
CY016941
A/chicken/Nigeria/1047-30/2006
2006
H5N1 NA
2A
CY016949
A/chicken/Nigeria/1047-34/2006
2006
H5N1 NA
2A
CY016925
A/chicken/Nigeria/1047-54/2006
2006
H5N1 NA
2A
CY016933
A/chicken/Nigeria/1047-62/2006
2006
H5N1 NA
2A
CY016909
A/chicken/Nigeria/1047-8/2006
2006
H5N1 NA
2A
CY016278
A/chicken/Nigeria/641/2006
2006
H5N1 NA
2A
DQ529296
A/chicken/Nigeria/641/2006
2006
H5N1 NA
2A
CY016286
A/chicken/Nigeria/957-20/2006
2006
H5N1 NA
2A
AM262564
A/chicken/Nigeria/BA210/2006
2006
H5N1 NA
2A
AM262565
A/chicken/Nigeria/BA211/2006
2006
H5N1 NA
2A
AM262560
A/chicken/Nigeria/SO300/2006
2006
H5N1 NA
2A
AM262561
A/chicken/Nigeria/SO452/2006
2006
H5N1 NA
2A, 3
AM262562
A/chicken/Nigeria/SO493/2006
2006
H5N1 NA
2A
AM262563
A/chicken/Nigeria/SO494/2006
2006
H5N1 NA
2A
CY016302
A/chicken/Sudan/1784-10/2006
2006
H5N1 NA
2A
CY016294
A/chicken/Sudan/1784-7/2006
2006
H5N1 NA
2A
CY020663
A/chicken/Sudan/1784-8/2006
2006
H5N1 NA
2A
CY021391
A/chicken/Sudan/2115-10/2006
2006
H5N1 NA
2A
CY020679
A/chicken/Sudan/2115-12/2006
2006
H5N1 NA
2A
CY020671
A/chicken/Sudan/2115-9/2006
2006
H5N1 NA
2A
DQ659680
A/common bussard/Bavaria/2/2006
2006
H5N1 NA
2A
EF165577
A/common buzzard/Bavaria/11/2006
2006
H5N1 NA
2A, 3
AB284326
A/common goldeneye/Mongolia/12/06
2006
H5N1 NA
2A
EF165574
A/common pochard/Bavaria/7/2006
2006
H5N1 NA
2A, 3
EF124202
A/crested myna/Hong Kong/540/2006
2006
H5N1 NA
2A
CY016781
A/Cygnus cygnus/Iran/754/2006
2006
H5N1 NA
2A
DQ534021
A/Cygnus olor/Czech Republic/6719/2006
2006
H5N1 NA
2A
CY017037
A/Cygnus olor/Italy/742/2006
2006
H5N1 NA
2A
CY020351
A/cygnus olor/Italy/808/2006
2006
H5N1 NA
2A
CY016805
A/duck/Cote d'Ivoire/1787-18/2006
2006
H5N1 NA
2A
EF532643
A/duck/Gaza/760/2006
2006
H5N1 NA
1A, 2A
EF532633
A/duck/Gaza/834/2006
2006
H5N1 NA
1A, 2A
AM403140
A/duck/Germany/R592/06
2006
H5N1 NA
G743A
AM403141
A/duck/Germany/R603/06
2006
H5N1 NA
G743A
CY017029
A/duck/Niger/914/2006
2006
H5N1 NA
2A
EF165576
A/eagle owl/Bavaria/10/2006
2006
H5N1 NA
2A, 3
AM403151
A/eagle owl/Germany/R1166/06
2006
H5N1 NA
G743A
EF165581
A/falcon/Bavaria/15/2006
2006
H5N1 NA
2A, 3
AM403149
A/falcon/Germany/R899/06
2006
H5N1 NA
G743A
EF165585
A/goldeneye duck/Bavaria/19/2006
2006
H5N1 NA
2A, 3
EF165584
A/goosander/Bavaria/18/2006
2006
H5N1 NA
2A, 3
EF165586
A/goosander/Bavaria/20/2006
2006
H5N1 NA
2A, 3
AM403144
A/goose/Germany/R696/06
2006
H5N1 NA
2A
DQ835389
A/goose/Iraq/812NAMRU3/2006
2006
H5N1 NA
2A
EF165588
A/great crested grebe/Bavaria/22/2006
2006
H5N1 NA
2A
AM403153
A/great crested grebe/Germany/R1226/06
2006
H5N1 NA
G743A
CY017181
A/guinea fowl/Nigeria/957-12/2006
2006
H5N1 NA
2A
AM403148
A/gull/Germany/R882/06
2006
H5N1 NA
G743A
I
SDN140068 A/Iraq/1/2006
2006
H5N1 NA
2A
I
SDN140067 A/Iraq/755/2006
2006
H5N1 NA
2A
DQ458993
A/mallard/Bavaria/1/2006
2006
H5N1 NA
2A, 3
CY016797
A/mallard/Italy/835/2006
2006
H5N1 NA
2A
EF165578
A/mute swan/Bavaria/12/2006
2006
H5N1 NA
2A
CY016917
A/ostrich/Nigeria/1047-25/2006
2006
H5N1 NA
2A
AM403139
A/pochard/Germany/R348/06
2006
H5N1 NA
G743A
EF165580
A/swan/Bavaria/14/2006
2006
H5N1 NA
2A
EF165582
A/swan/Bavaria/16/2006
2006
H5N1 NA
2A, 3
EF165583
A/swan/Bavaria/17/2006
2006
H5N1 NA
2A, 3
EF165587
A/swan/Bavaria/21/2006
2006
H5N1 NA
2A, 3
EF165573
A/swan/Bavaria/6/2006
2006
H5N1 NA
2A, 3
DQ464355
A/swan/Germany/R65/2006
2006
H5N1 NA
2A
CY017045
A/swan/Slovenia/760/2006
2006
H5N1 NA
2A
EF165571
A/tufted duck/Bavaria/4/2006
2006
H5N1 NA
2A, 3
EF165575
A/tufted duck/Bavaria/9/2006
2006
H5N1 NA
2A
AM403155
A/tufted duck/Germany/R1240/06
2006
H5N1 NA
G743A
I
SDN133097 A/Turkey/12/2006
2006
H5N1 NA
2A
I
SDN133104 A/Turkey/15/2006
2006
H5N1 NA
2A
I
SDN133358 A/Turkey/651242/06
2006
H5N1 NA
2A
I
SDN133366 A/Turkey/65596/06
2006
H5N1 NA
2A
EF532634
A/turkey/Israel/345/2006
2006
H5N1 NA
2A
EF532635
A/turkey/Israel/364/2006
2006
H5N1 NA
2A
EF532636
A/turkey/Israel/365/2006
2006
H5N1 NA
2A
EF532638
A/turkey/Israel/446/2006
2006
H5N1 NA
2A
CY020695
A/turkey/Ivory Coast/4372-2/2006
2006
H5N1 NA
2A
CY020703
A/turkey/Ivory Coast/4372-3/2006
2006
H5N1 NA
2A
CY020711
A/turkey/Ivory Coast/4372-4/2006
2006
H5N1 NA
2A
AB265202
A/whooper swan/Mongolia/2/06
2006
H5N1 NA
2A
DQ650661
A/chicken/Crimea/04/2005
2005
H5N1 NA
2A
DQ650665
A/chicken/Crimea/08/2005
2005
H5N1 NA
2A
DQ449634
A/chicken/Kurgan/05/2005
2005
H5N1 NA
2A
DQ323673
A/chicken/Kurgan/3/2005
2005
H5N1 NA
2A
EF205169
A/chicken/Omsk/14/05
2005
H5N1 NA
2A
EF205168
A/chicken/Suzdalka/06/05
2005
H5N1 NA
2A
EF205174
A/chicken/Tula/4/05
2005
H5N1 NA
2A
DQ449642
A/duck/Kurgan/08/2005
2005
H5N1 NA
2A
DQ864710
A/duck/Novosibirsk/02/05
2005
H5N1 NA
2A
DQ230524
A/duck/Novosibirsk/56/2005
2005
H5N1 NA
2A
EF205171
A/goose/Krasnoozerskoe/627/05
2005
H5N1 NA
2A
EF205170
A/goose/Suzdalka/10/05
2005
H5N1 NA
2A
DQ230523
A/grebe/Novosibirsk/29/2005
2005
H5N1 NA
2A
I
SDN129516 A/turkey/Turkey/1/05
2005
H5N1 NA
2A
I
SDN138123 A/whooper swan/Mongolia/13/2005
2005
H5N1 NA
2A
AB239311
A/whooper swan/Mongolia/3/05
2005
H5N1 NA
2A
AB239318
A/whooper swan/Mongolia/4/05
2005
H5N1 NA
2A
AB239325
A/whooper swan/Mongolia/6/05
2005
H5N1 NA
2A
I
SDN138115 A/whooper swan/Mongolia/7/2005
2005
H5N1 NA
2A
I
SDN138107 A/whooper swan/Qinghai/1/2005
2005
H5N1 NA
2A
I
SDN212035 A/Chicken/Karo/BBPV1/2006
2006
H5N1 NA
3
DQ835315
A/China/GD01/2006
2006
H5N1 NA
3
EF124209
A/duck/Guangxi/89/2006
2006
H5N1 NA
3
DQ842490
A/Guangzhou/1/2006
2006
H5N1 NA
3
EF473077
A/chicken/Cambodia/022LC2b/2005
2005
H5N1 NA
3
EF473078
A/chicken/Cambodia/022LC3b/2005
2005
H5N1 NA
3
DQ493015
A/chicken/Dairi/BPPVI/2005
2005
H5N1 NA
3
DQ321030
A/chicken/Guangxi/604/2005
2005
H5N1 NA
3
DQ321059
A/chicken/Salatiga/BBVet-I/2005
2005
H5N1 NA
3
DQ189241
A/chicken/Thailand/Kamphaengphet-3-01/2005
2005
H5N1 NA
3
DQ189242
A/chicken/Thailand/Kamphaengphet-3-02/2005
2005
H5N1 NA
3
DQ251794
A/chicken/Thailand/Kamphaengphet-3-03/2005
2005
H5N1 NA
3
DQ251795
A/chicken/Thailand/Kamphaengphet-3-04/2005
2005
H5N1 NA
3
EF124275
A/duck/Guangxi/2775/2005
2005
H5N1 NA
3
EF124287
A/duck/Guangxi/4016/2005
2005
H5N1 NA
3
DQ321031
A/duck/Guangxi/793/2005
2005
H5N1 NA
3
DQ321033
A/duck/Guangzhou/20/2005
2005
H5N1 NA
3
DQ321043
A/duck/Hunan/1265/2005
2005
H5N1 NA
3
DQ321044
A/duck/Hunan/1608/2005
2005
H5N1 NA
3
DQ321045
A/duck/Hunan/1652/2005
2005
H5N1 NA
3
DQ493046
A/duck/Vietnam/286/2005
2005
H5N1 NA
3
DQ493043
A/duck/Vietnam/S640/2005
2005
H5N1 NA
3
EF473067
A/goose/Cambodia/022b/2005
2005
H5N1 NA
3
DQ321047
A/goose/Shantou/2216/2005
2005
H5N1 NA
3
EF124296
A/goose/Yunnan/4804/2005
2005
H5N1 NA
3
I
SDN128027 A/Indonesia/6/2005
2005
H5N1 NA
3
DQ321029
A/quail/Guangxi/575/2005
2005
H5N1 NA
3
DQ321065
A/quail/Malaysia/6309/2005
2005
H5N1 NA
3
DQ493075
A/Vietnam/CL105/2005
2005
H5N1 NA
3
AY770992
A/chicken/Ayutthaya/Thailand/CU-23/04
2004
H5N1 NA
3
DQ083605
A/chicken/Ayutthaya/Thailand/CU-24/04
2004
H5N1 NA
3
AY609314
A/chicken/Guangdong/174/04
2004
H5N1 NA
3
DQ017340
A/chicken/Kamphaengphet-2-01/2004
2004
H5N1 NA
3
DQ017339
A/chicken/Kamphaengphet-2-03/2004
2004
H5N1 NA
3
DQ017330
A/chicken/Kamphaengphet-2-05/2004
2004
H5N1 NA
3
DQ017335
A/chicken/Kamphaengphet-2-06/2004
2004
H5N1 NA
3
DQ096567
A/chicken/Malaysia/01/2004
2004
H5N1 NA
3
DQ321066
A/chicken/Malaysia/5858/2004
2004
H5N1 NA
3
DQ017333
A/chicken/Nakornsawan-2-01/2004
2004
H5N1 NA
3
DQ017328
A/chicken/Nakornsawan-2-04/2004
2004
H5N1 NA
3
DQ017338
A/chicken/Nakornsawan-2-05/2004
2004
H5N1 NA
3
DQ017336
A/chicken/Phetchabun-2-01/2004
2004
H5N1 NA
3
DQ017329
A/chicken/Phetchabun-2-02/2004
2004
H5N1 NA
3
DQ017341
A/chicken/Phitsanulok-2-01/2004
2004
H5N1 NA
3
DQ017342
A/chicken/Phitsanulok-2-02/2004
2004
H5N1 NA
3
DQ017334
A/chicken/Phitsanulok-2-04/2004
2004
H5N1 NA
3
DQ083618
A/chicken/Prachinburi/Thailand/CU-104/04
2004
H5N1 NA
3
DQ083608
A/chicken/Saraburi/Thailand/CU-27/04
2004
H5N1 NA
3
AY553813
A/chicken/Thailand/Nakornsawan-01/2004
2004
H5N1 NA
3
AY553815
A/chicken/Thailand/Sukhothai-01/2004
2004
H5N1 NA
3
AY553831
A/chicken/Thailand/Sukhothai-02/2004
2004
H5N1 NA
3
AY553812
A/chicken/Thailand/Tak-01/2004
2004
H5N1 NA
3
AY553819
A/chicken/Thailand/Uttaradit-02/2004
2004
H5N1 NA
3
DQ017332
A/chicken/Uthaithani-2-01/2004
2004
H5N1 NA
3
DQ017344
A/chicken/Uthaithani-2-02/2004
2004
H5N1 NA
3
DQ017345
A/chicken/Uthaithani-2-03/2004
2004
H5N1 NA
3
DQ017337
A/chicken/Uthaithani-2-04/2004
2004
H5N1 NA
3
AY651455
A/Chicken/Viet Nam/C57/2004
2004
H5N1 NA
3
DQ493050
A/chicken/Vietnam/147/2004
2004
H5N1 NA
3
DQ884460
A/chicken/Vietnam/486A/2004
2004
H5N1 NA
3
AY818142
A/chicken/Vietnam/C58/04
2004
H5N1 NA
3
DQ321068
A/chicken/Vietnam/DT171/2004
2004
H5N1 NA
3
DQ083606
A/crow/Bangkok/Thailand/CU-25/04
2004
H5N1 NA
3
AB189063
A/crow/Osaka/102/2004
2004
H5N1 NA
3
DQ530175
A/dog/Thailand-Suphanburi/KU-08/04
2004
H5N1 NA
3
DQ017326
A/duck/Phichit-2-02/2004
2004
H5N1 NA
3
DQ017331
A/duck/Phitsanulok-2-01/2004
2004
H5N1 NA
3
AY553822
A/duck/Thailand/Sukhothai-01/2004
2004
H5N1 NA
3
DQ017343
A/duck/Uthaithani-2-01/2004
2004
H5N1 NA
3
DQ493056
A/goose/Vietnam/264/2004
2004
H5N1 NA
3
EF512562
A/Prachinburi/6231/2004
2004
H5N1 NA
3
DQ083607
A/rollers/Bangkok/Thailand/CU-26/04
2004
H5N1 NA
3
AY842936
A/tiger/Thailand/CU-T3/2004
2004
H5N1 NA
3
AY972543
A/tiger/Thailand/CU-T4/04
2004
H5N1 NA
3
AY972544
A/tiger/Thailand/CU-T5/04
2004
H5N1 NA
3
AY972545
A/tiger/Thailand/CU-T6/04
2004
H5N1 NA
3
AY866476
A/tiger/Thailand/CU-T7/2004
2004
H5N1 NA
3
AY972546
A/tiger/Thailand/CU-T8/04
2004
H5N1 NA
3
EF541464
A/chicken/Korea/es/2003
2003
H5N1 NA
3
AY676044
A/duck/Korea/ESD1/03
2003
H5N1 NA
3
Nature Precedings : hdl:10101/npre.2007.459.2 : Posted 16 Jul 2007