Concurrent Acquisition of a Single Nucleotide Polymorphism in Diverse Influenza H5N1
Clade 2.2 Sub-clades
Henry L. Niman
1
, Mona M. Aly
2
, Abdel-Satar Arafa
2
, Nasr El-Sayed
3
, Ahmed E. Nayel
3
,
Ahmed S. Abdelghani
3
, Hala M. Esmat
3
, , Gregory A. Raczniak
4,5
, Mensah Agyen-
Frempong
6
, William K. Ampofo
6
, Bruce R. Boynton
7
1
Recombinomics, Inc., Pittsburgh, Pennsylvania, USA,
2
Central Laboratory for Veterinary
Quality Control, Giza, Egypt,
3
Ministry of Health and Population, Arabic Republic of Egypt,
Cairo, Egypt,
4
NAMRU-3 Ghana Detachment, Accra, Ghana
5
Ghana Veterinary Services,
Accra, Ghana,
6
Noguchi Memorial Institute for Medical Research, Accra, Ghana, ,
7
U.S.
Naval Medical Research Center, Silver Springs, Maryland, USA
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: 2236
6 Figures, 4 Supplemental Figures,1 Supplemental Table
Running Title: Concurrent Influenza SNP Acquisitions
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
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, Kuwait, 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. The recombination role is further supported by the high fidelity
replication in swine influenza
9
and aggregation of single nucleotide polymorphisms in
H5N1 clade 2.2 hemagglutinin
10
.
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
11,12
showed that isolates from migratory waterfowl
were easily distinguishable from previous isolates linked to poultry and human infections in
eastern and southeastern Asia
13,14
. The new strain was subsequently found in outbreaks
in Russia, Kazakhstan and Mongolia
15,16
. 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 bird
sanctuaries signaled the start of a major geographical expansion of H5N1. In the following
12 months, almost 50 countries west of China reported H5N1 for the first time. All
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
infections were clade 2.2. The geographical reach included Europe, the Middle East and
Africa. This expansion offered a unique opportunity to study the evolution of H5N1 as it
migrated into new regions, producing human cases in Turkey, Iraq, Azerbaijan, Egypt,
Djibouti in 2006 and Nigeria and Pakistan in 2007. Sequence analysis indicated all cases
were due to clade 2.2 infections. 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. The single nucleotide polymorphism
distribution pathways could then be used to predict vaccine targets.
We isolated H5N1 from patients and poultry
17
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. A subset of
these markers was also identified upstream in European isolates and downstream in west
African isolates. These isolates are listed in table S1.
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
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
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 (H3 numbering),, 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, as well as a case with a novel HA cleavage site,
GERRRRKR. 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 (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 (or the corresponding HA cladogram in figure 1S). The
location of the isolate on the tip of the branch indicated it was a recent acquisition. Indeed,
there were only two additions when compared to the most closely related human isolate.
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
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. These two isolates
had three new polymorphisms relative to isolates collected in June, 2006. 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 describe above. However, all sequenced,
plaque-purified clones had G743A (data not shown), confirming the acquisition of G743A
onto two distinct genetic backgrounds in Egypt.
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 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 supplemental figure 1. The isolates with G743A are also located at the tips of the
branches, supporting a recent acquisition of the polymorphism.
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
In February 2007 an H5N1 clade 2.2 outbreak was reported in Kuwait. The sequences in
Kuwait traced back to a June, 2006 massive wild bird outbreak at Uvs Lake in Mongolia,
which affected wild birds in northern Mongolia and southern Siberia. Related sequences
were subsequently isolated in the outbreak in South Korea in November, 2006. These
sequences have been designated clade 2.2.3 and are represented in an NA cladogram in
figure 2 or an HA cladogram in supplemental figure 2. The 2006 sequences did not have
G743A. However, G743A was present in all reported NA sequences from Kuwait. Related
sequences were subsequently reported in German outbreaks in Bavaria, Saxony, and
Thuringen during the summer of 2007, outbreaks in Krasnodar in the fall of 2007, and
outbreaks in Saudi Arabia in the winter of 2007. All reported NA sequences had G743A.
In February 2007, a distinct H5N1 clade 2.2.3 outbreak occurred near Moscow,
Russia. Isolates from infected chickens were most closely related to 2006 sequences from
Azerbaijan. The sequences formed a separate branch which is also seen in Figure 2.
Although these clade 2.2.3 isolates were collected from a wide geographical area, G743A
was not found in any of the 2006 isolates. The isolates from Moscow map onto the tip of
the branch formed by the Azerbaijan isolates. Subsequently, sequences related to the
Moscow sequences were reported from Rostov, Russia. As seen in figure 2, these
sequences had G743A. (see supplement figure 2 for HA phylogenetic tree or supplement
Table 1 for partial NA sequences with G743A).
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. Figure 3 contains the isolates from western
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
Africa, which include isolates from Nigeria collected in 2006 and 2007. Like the Egypt,
Russian, and German isolates above, the 2007 isolates extended branches rooted with
2006 isolates which did not have G743A. 2007 isolates from Ghana and Nigeria have
G743A appended onto the 2006 genetic backgrounds.
Although G743A was not present in the root sequences from the 2007 isolates which had
G743A, it was present in a subset of clade 2.2 sequences from Europe, which are shown in
Figure 4. G743A was found in a series of closely related sequences from southern
Germany, Switzerland, and France (additional partial sequences in Table 1).
The NA G743A polymorphism can be traced through public H5N1 sequences. Isolates with
full NA sequences are listed in the NA phylogram in figure 5. The polymorphism was
identified in the first reported sequences linked to the spread of H5N1 in Asia in 2003/2004
in South Korea
18
and Japan
19
. 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
20
(see figure 4)
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 the 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. The number of new acquisitions for the examples described was 2-6. Thus, in
each case the acquisition of G743A was included in a small number of new acquisitions,
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
which would be inconsistent with random mutations. All discussed isolates on the tips of
the branches were collected in 2007, and in five geographically distinct regions
representing 11 clade 2.2 genetic backgrounds. 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 four 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.
The example of G743A is dramatic, but not unique. Movement of three HA polymorphisms
are depicted in Figure 6. These three polymorphisms are found in the same set of 2006
isolates with G743A. One polymorphism is in all of the isolates with G743A. The HA
polymorphism, C689T, was initially reported in the first isolates from the outbreak in Siberia
in the summer of 2005. In 2007, the polymorphism is also in Egyptian isolates in one of the
branches which formed in the 2006/2007 and includes two of the first isolates from
Gharbiya which had NA G743A. This polymorphism has been stable and was present in
the 2008 isolate on the branch. A second example is the non synonymous change G754A,
which encodes the receptor binding domain change, M230I. This polymorphism is in one
of the German isolates and the isolate in the branch that links back to the other initial
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
isolate with G743A. G754A is also on additional 2007 isolates in Egypt. The third
polymorphism, C1614T, is in another 2006 isolate from Germany, but is found on two
Egyptian branches. One of the branches is a subset of the isolates with G754A. C1614T
is a recent acquisition by isolates on this branch. Recently, analysis of human influenza
found evidence for homologous recombination over small regions
21
. The majority of these
acquisitions were in NA. Detection of additional examples was limited, but the analysis did
not extend beyond sequences generated through an influenza sequencing program, which
limited parental sequences from early human isolates as well as isolates from other
species, including avian and swine.
In this paper the acquisitions that link through Germany represent a small subset of the
newly acquired polymorphisms, but help demonstrate mapping of the polymorphisms. A
more detailed map will be presented elsewhere. The origin of the acquisitions by clade 2.2
isolates is evolving. The initial regional polymorphisms frequently traced back to H5N1
isolates in Asia. Similar results for human influenza were recently described
22,23
. The
phylogenetic analysis identified Asian origins for seasonal flu, which subsequently spread
to Europe and North America, followed by South America.
In this paper, more recent acquisitions trace back to other clade 2.2 isolates. Many of the
most recent acquisitions are present in isolates in Egypt, or in the newly emerging clade
2.2.3, which has become dominant in Europe
Recombination is further supported by the aggregation of clade 2.2 polymorphisms from
Germany, Egypt, and sub-Sahara Africa into a single human hemaglutinnin sequence in
Nigeria
10
.
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
This shuffling of single nucleotide polymorphisms creates new sequences which define
antigenic drift. Mapping of these pathways and associations may be used to develop novel
vaccine targets representing rapidly evolving genomes.
Figure 1 NA Phylogram of Egyptian Isolates
NA phylogram of positions 43-1337. Isolates with G743A are annotated. NA regional
markers are C150T, C236T, A703G, A740G, T1088C and G1280A.
Accession numbers
and additional isolates with G743A listed in Table S1.
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
24
.
Figure 2 NA Phylogram of Clade 2.2.3 Isolates.
NA phylogram of positions 43-1337. Isolates with G743A are annotated.
Accession
numbers and additional isolates with G743A listed in Table S1.
Trees generated using
neighbor joining with 100 bootstrap repetitions. Sequences generated as described
previously
24
.
Figure 3 NA Phylogram of West African Isolates.
NA phylogram of positions 43-1337. Isolates with annotated.
Accession numbers and
additional isolates with G743A listed in Table S1.
Trees generated using neighbor joining
with 100 bootstrap repetitions. Sequences generated as described previously
24
.
Figure 4 NA Phylogram of European Isolates.
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
NA phylogram of positions 43-1337. Isolates with G743A are annotated. Ac
cession
numbers and additional isolates with G743A listed in Table S1.
Trees generated using
neighbor joining with 100 bootstrap repetitions. Sequences generated as described
previously
24
Figure 5 NA Phylogram of H5N1 Isolates with G743A
Accession numbers and additional isolates with G743A listed in Table S1.
Trees generated
using neighbor joining with 100 bootstrap repetitions. Sequences generated as described
previously
24
Figure 6 HA Phylogram of Recent Egyptian Isolates
HA phylogram of positions 93-1688 in isolates with C689T, G754A, and C1614T. HA
regional markers are G467A, C661T, C727T, A880G, T937C, and G1018T. 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
24
.
Figure S1 HA Phylograms of Egyptian Isolates
HA phylogram of positions 93-1688. Isolates with G743A are annotated. HA regional
markers are G467A, C661T, C727T, A880G, T937C, and G1018T. 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
24
.
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
Figure S2 HA Phylogram of Clade 2.2.3 Isolates
HA phylogram of positions 93-1688. Isolates with G743A are annotated. Accession
numbers and additional isolates with HA polymorphisms listed in Table S1. Trees
generated using neighbor joining with 100 bootstrap repetitions. Sequences generated as
described previously
24
.
Figure S3 HA Phylogram of West African Isolates.
HA phylogram of positions 93-1688. Isolates with G743A are annotated. Accession
numbers listed in Table S. Trees generated using neighbor joining with 100 bootstrap
repetitions. Sequences generated as described previously
24
.
Figure S4 HA Phylogram of European Isolates.
HA phylogram of positions 93-1688. Isolates with G743A are annotated. Accession
numbers listed in Table S. Trees generated using neighbor joining with 100 bootstrap
repetitions. Sequences generated as described previously
24
.
Table S1 Isolates and Accession Numbers
Names and accession numbers of HA and NA sequences used in figures 1-6. Partial
sequences
25-32
with polymorphisms in figures 1-5 are listed.
Acknowledgements
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
We thank Magdi Saad, Jeffery Tjaden, Kenneth Earhart, Marshall Monteville, Emad Labib,
Ehab Ayoub, Moustafa Mansour, from US NAMRU-3 for sequences, Cecilia DeMattos from
US NAMRU-3 for plaque purification of avian isolates; Evelyn Yayra Bonney, Ivy Asante,
Ken-Edwin Aryee, Kofi Bonney, Jacob Barnor and Professor Alexander Nyarko from
Noguchi Memorial Institute for Medical Research; Mr. Jerry Odoi, Drs. Joseph Gaari-
Kweku, Joseph Adongo Awuni, John Niendow Karimu and Bashiru Kikimoto from the
Ghana Veterinary Services; Mr. Michael Adjabeng, Drs. Edward Antwi and Lawson
Ahadzie from the Ghana Health Service; for the collection, screening and testing of avian
and human samples for H5NI in Ghana.
The views expressed in this publication are those of the authors and do not necessarily
represent the official policy or position of the Department of the Navy.
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G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure 1 NA Phylogram of Egyptian Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure 2 NA Phylogram of Clade 2.2.3 Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure 3 NA Phylogram of West African Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure 4 NA Phylogram of European Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
Figure 5 NA Phylogram of H5N1 Isolates With G743A
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
C689T
C689T
C689T
C689T
C689T
C689T
C689T
C689T
G754A
G754A
G754A
G754A
G754A
G754A
G754A
G754A
G754A
G754A
G754A
G754A
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C1614T
C689T
C689T
C689T
C689T
Figure 6 HA Phylogram of Recent Egyptian Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure S1 HA Phylogram of Egyptian Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure S2 HA Phylogram of Clade 2.2.3 Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure S3 HA Phylogram of West African Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
G743A
G743A
G743A
G743A
G743A
G743A
G743A
G743A
Figure S4 HA Phylogram of European Isolates
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
Table S1 H5N1 Isolates and Accession Numbers
Accession Name
Year Goverate
Figure
EF624250 A/chicken/Ghana/3158-NAMRU3/2007
2007 Tema
NA
3
EF624251 A/chicken/Ghana/3159-NAMRU3/2007
2007 Tema
NA
3
EF624252 A/chicken/Ghana/3160-NAMRU3/2007
2007 Tema
NA
3
A/chicken/Egypt/1079-NAMRU3/2007
2007 Beni Suef NA
1
EF486240 A/chicken/Egypt/1129N3-HK9/2007
2007 Fayoum
NA
1
A/chicken/Egypt/1300-NAMRU3/2007
2007 Gharbiya NA
1
EF486243 A/chicken/Egypt/1889N3-SM26/2007
2007 Giza
NA
1
EF486244 A/chicken/Egypt/1890N3-HK45/2007
2007 Gharbiya NA
1
EF486245 A/chicken/Egypt/1891N3-CLEVB/2007
2007 Gharbiya NA
1
EF486246 A/chicken/Egypt/1892N3-HK49/2007
2007 Gharbiya NA
1
A/duck/Egypt/1301-NAMRU3/2007
2007 Gharbiya NA
1
EF486249 A/duck/Egypt/1888N3-SM25/2007
2007 Giza
NA
1
EF382360 A/Egypt/0636-NAMRU3/2007
2007 Beni Suef NA
1
A/Egypt/1394-NAMRU3/2007
2007 Fayoum
NA
1
A/Egypt/1604-NAMRU3/2007
2007 Fayoum
NA
1
A/Egypt/1731-NAMRU3/2007
2007 Sharkia
NA
1
A/Egypt/1902-NAMRU3/2007
2007 Dakahlia NA
1
A/Egypt/2256-NAMRU3/2007
2007 Dakahlia NA
1
A/Egypt/2321-NAMRU3/2007
2007 Aswan
NA
1
A/Egypt/2331-NAMRU3/2007
2007 Aswan
NA
1
A/Egypt/2616-NAMRU3/2007
2007 Aswan
NA
1
A/Egypt/2620-NAMRU3/2007
2007 Menia
NA
1
A/Egypt/2621-NAMRU3/2007
2007 Qena
NA
1
A/Egypt/2629-NAMRU3/2007
2007 Qena
NA
1
A/Egypt/2630-NAMRU3/2007
2007 Sohag
NA
1
A/Egypt/2631-NAMRU3/2007
2007 Qalubiea NA
1
A/Egypt/2750-NAMRU3/2007
2007 Menia
NA
1
A/Egypt/2651-NAMRU3/2007
2007 Shoubra
NA
1
A/Egypt/4081-NAMRU3/2007
2007
NA
1
A/Egypt/4082-NAMRU3/2007
2007
NA
1
A/Egypt/4226-NAMRU3/2007
2007
NA
1
A/Egypt/6251-NAMRU3/2007
2007
NA
1
EF486241 A/chicken/Egypt/12378N3-CLEVB/2006
2006 Gharbiya NA
1
EF486242 A/chicken/Egypt/12379N3-CLEVB/2006
2006 Sharqiya NA
1
EF486247 A/duck/Egypt/12380N3-CLEVB/2006
2006 Gharbiya NA
1
EF486248 A/duck/Egypt/13010N3-CLEVB/2006
2006 Menoufiya NA
1
EF222324 A/Egypt/12374-NAMRU3/2006
2006 Gharbiya NA
1
EF222323 A/Egypt/14724-NAMRU3/2006
2006 Gharbiya NA
1
EF222322 A/Egypt/14725-NAMRU3/2006
2006 Gharbiya NA
1
A/Egypt/2991-NAMRU3/2006
2006 Kafr El-Sheikh
NA
1
EF486250 A/goose/Egypt/13009N3-SM2/2006
2006 Menoufiya NA
1
EF624253 A/chicken/Ghana/3158-NAMRU3/2007
2007 Tema
HA
S3
EF624254 A/chicken/Ghana/3159-NAMRU3/2007
2007 Tema
HA
S3
EF624255 A/chicken/Ghana/3160-NAMRU3/2007
2007 Tema
HA
S3
EF441277 A/chicken/Egypt/1079-NAMRU3/2007
2007 Beni Suef HA
S1, 6
EF469650 A/chicken/Egypt/1129N3-HK9/2007
2007 Fayoum
HA
S1, 6
EF441280 A/chicken/Egypt/1300-NAMRU3/2007
2007 Gharbiya HA
S1, 6
EF469653 A/chicken/Egypt/1889N3-SM26/2007
2007 Giza
HA
S1, 6
EF469654 A/chicken/Egypt/1890N3-HK45/2007
2007 Gharbiya HA
S1, 6
EF469659 A/chicken/Egypt/1891N3-CLEVB/2007
2007 Gharbiya HA
S1, 6
EF469660 A/chicken/Egypt/1892N3-HK49/2007
2007 Gharbiya HA
S1, 6
EF441281 A/duck/Egypt/1301-NAMRU3/2007
2007 Gharbiya HA
S1, 6
EF469657 A/duck/Egypt/1888N3-SM25/2007
2007 Giza
HA
S1, 6
EF382359 A/Egypt/0636-NAMRU3/2007
2007 Beni Suef HA
S1, 6
EF535817 A/Egypt/1394-NAMRU3/2007
2007 Fayoum
HA
S1, 6
EF535818 A/Egypt/1604-NAMRU3/2007
2007 Fayoum
HA
S1, 6
EF535819 A/Egypt/1731-NAMRU3/2007
2007 Sharkia
HA
S1, 6
EF535820 A/Egypt/1902-NAMRU3/2007
2007 Dakahlia HA
S1, 6
EF535821 A/Egypt/2256-NAMRU3/2007
2007 Dakahlia HA
S1, 6
EF535822 A/Egypt/2321-NAMRU3/2007
2007 Aswan
HA
S1, 6
EF535823 A/Egypt/2331-NAMRU3/2007
2007 Aswan
HA
S1, 6
EF535824 A/Egypt/2616-NAMRU3/2007
2007 Aswan
HA
S1, 6
EF535825 A/Egypt/2620-NAMRU3/2007
2007 Menia
HA
S1, 6
EF535826 A/Egypt/2621-NAMRU3/2007
2007 Qena
HA
S1, 6
EU095025 A/Egypt/2629-NAMRU3/2007
2007 Qena
HA
S1, 6
EU095026 A/Egypt/2630-NAMRU3/2007
2007 Sohag
HA
S1, 6
EU095027 A/Egypt/2631-NAMRU3/2007
2007 Qalubiea HA
S1, 6
EU095028 A/Egypt/2750-NAMRU3/2007
2007 Menia
HA
S1, 6
EU095029 A/Egypt/2651-NAMRU3/2007
2007 Shoubra
HA
S1, 6
EU095030 A/Egypt/4081-NAMRU3/2007
2007
HA
S1, 6
EU095031 A/Egypt/4082-NAMRU3/2007
2007
HA
S1, 6
EU095032 A/Egypt/4226-NAMRU3/2007
2007
HA
S1, 6
EU095033 A/Egypt/6251-NAMRU3/2007
2007
HA
S1, 6
EF441276 A/chicken/Egypt/1078-NAMRU3/2006
2006 Menoufiya HA
S1, 6
EF441278 A/chicken/Egypt/1080-NAMRU3/2006
2006 Damietta HA
S1, 6
EF441279 A/chicken/Egypt/1081-NAMRU3/2006
2006 Gharbiya HA
S1, 6
EF042622 A/chicken/Egypt/10845-NAMRU3/2006
2006
HA
S1, 6
EF469651 A/chicken/Egypt/12378N3-CLEVB/2006
2006 Gharbiya HA
S1, 6
EF469652 A/chicken/Egypt/12379N3-CLEVB/2006
2006 Sharqiya HA
S1, 6
DQ837587 A/chicken/Egypt/5610NAMRU3-F3/2006
2006 Fayoum
HA
S1, 6
DQ837588 A/chicken/Egypt/5611NAMRU3-AN/2006
2006 Cairo
HA
S1, 6
DQ837589 A/chicken/Egypt/5612NAMRU3-S/2006
2006 Giza
HA
S1, 6
DQ447199 A/chicken/Egypt/960N3-004/2006
2006
HA
S1, 6
DQ666146 A/Djibouti/5691NAMRU3/2006
2006 Djibouti
HA
S1, 6
EF469655 A/duck/Egypt/12380N3-CLEVB/2006
2006 Gharbiya HA
S1, 6
EF469656 A/duck/Egypt/13010N3-CLEVB/2006
2006 Menoufiya HA
S1, 6
EF061116 A/Egypt/12374-NAMRU3/2006
2006 Gharbiya HA
S1, 6
EF200512 A/Egypt/14724-NAMRU3/2006
2006 Gharbiya HA
S1, 6
EF200513 A/Egypt/14725-NAMRU3/2006
2006 Gharbiya HA
S1, 6
EF042614 A/Egypt/2763-NAMRU3/2006
2006 Qaliubiya HA
S1, 6
DQ464377 A/Egypt/2782-NAMRU3/2006
2006 Qaliubiya HA
S1, 6
EF042615 A/Egypt/2783-NAMRU3/2006
2006 Qaliubiya HA
S1, 6
EF042616 A/Egypt/2786-NAMRU3/2006
2006 Gharbiya HA
S1, 6
EF042617 A/Egypt/2947-NAMRU3/2006
2006 Kafr El-Sheikh
HA
S1, 6
EF042618 A/Egypt/3105-NAMRU3/2006
2006 Sohaj
HA
S1, 6
EF042619 A/Egypt/3458-NAMRU3/2006
2006 Minufiyah HA
S1, 6
EF042620 A/Egypt/5494-NAMRU3/2006
2006 Sharabeya HA
S1, 6
EF042621 A/Egypt/5614-NAMRU3/2006
2006 Al Minya HA
S1, 6
EF469658 A/goose/Egypt/13009N3-SM2/2006
2006 Menoufiya HA
S1, 6
DQ837590 A/turkey/Egypt/5613NAMRU3-T/2006
2006 Menoufiya HA
S1, 6
EF042624 A/teal/Egypt/14051-NAMRU3/2005
2005 Damietta HA
S1, 6
AM773723 A/black-necked grebe/Germany/R1393/07
2007
NA
G743A
EU443569 A/chicken/Czech Republic/11242-38/07
2007
NA
2
EU163433 A/chicken/Krasnodar/300/07
2007
NA
2
CY029950 A/chicken/Kuwait/KISR2/2007
2007
NA
2
CY029958 A/chicken/Kuwait/KISR3/2007
2007
NA
2
CY029966 A/chicken/Kuwait/KISR5/2007
2007
NA
2
CY029974 A/chicken/Kuwait/KISR6/2007
2007
NA
2
CY029982 A/chicken/Kuwait/KISR7/2007
2007
NA
2
CY029990 A/chicken/Kuwait/KISR8/2007
2007
NA
2
CY029998 A/chicken/Kuwait/KISR9/2007
2007
NA
2
EF474448 A/chicken/Moscow/2/2007
2007
NA
2
EU148358 A/chicken/Nigeria/1071-1/2007
2007
NA
3
EU148406 A/chicken/Nigeria/1071-10/2007
2007
NA
3
EU148414 A/chicken/Nigeria/1071-15/2007
2007
NA
3
EU148422 A/chicken/Nigeria/1071-22/2007
2007
NA
3
EU148430 A/chicken/Nigeria/1071-23/2007
2007
NA
3
EU148438 A/chicken/Nigeria/1071-29/2007
2007
NA
3
EU148366 A/chicken/Nigeria/1071-3/2007
2007
NA
3
EU148446 A/chicken/Nigeria/1071-30/2007
2007
NA
3
EU148373 A/chicken/Nigeria/1071-4/2007
2007
NA
3
EU148382 A/chicken/Nigeria/1071-5/2007
2007
NA
3
EU148390 A/chicken/Nigeria/1071-7/2007
2007
NA
3
EU148398 A/chicken/Nigeria/1071-9/2007
2007
NA
3
EU402400 A/chicken/Rostov/22-1/2007
2007
NA
2
EU401753 A/chicken/Rostov-on-Don/35/2007
2007
NA
2
EF605599 A/chicken/Russia_Krasnodar/2/2007
2007
NA
2
EF447431 A/chicken/Russia_Moscow oblast_Odintsovo/1/2007 2007
NA
2
EU257632 A/Cygnus cygnus/Krasnodar/329/07
2007
NA
2
EU443570 A/Cygnus olor/Czech Republic/10732/07
2007
NA
2
EU445682 A/houbara bustard/Saudi Arabia/6732-1/2007
2007
NA
2
EU441928 A/muscovy duck/Rostov-on-Don/51/2007
2007
NA
2
AM749445 A/mute swan/Germany/R1349/07
2007
NA
2
I
SDN230944
A/Nigeria/6e/07
2007
NA
3
EU401799 A/peacock/Mansehra-Pakistan/NARC-7558/02/2007 2007
NA
2
EU441936 A/pigeon/Rostov-on-Don/6/2007
2007
NA
2
EU486854 A/starling/Rostov-on-Don/39/07
2007
NA
2
EU443568 A/turkey/Czech Republic/10309-3/07
2007
NA
2
EU401797 A/turkey/Islamabad-Pakistan/NARC-7871/02/2007
2007
NA
2
I
SDN140072
A/Azerbaijan/001-161/2006
2006
NA
2
I
SDN140073
A/Azerbaijan/002-115/2006
2006
NA
2
I
SDN140069
A/Azerbaijan/006-207/2006
2006
NA
2
I
SDN140070
A/Azerbaijan/008-208/2006
2006
NA
2
I
SDN140071
A/Azerbaijan/011-162/2006
2006
NA
2
EF165579 A/buzzard/Bavaria/13/2006
2006
NA
4
EF165572 A/buzzard/Bavaria/5/2006
2006
NA
4
EF523697 A/buzzard/Denmark/6370/06
2006
NA
4
DQ643984 A/cat/Germany/606/2006
2006
NA
4
DQ835390 A/cat/Iraq/820NAMRU3/2006
2006
NA
4
CY016813 A/chicken/Cote d'Ivoire/1787-34/2006
2006
NA
3
EF532639 A/chicken/Gaza/450/2006
2006
NA
1
EF532641 A/chicken/Gaza/713/2006
2006
NA
1
EF532642 A/chicken/Gaza/714/2006
2006
NA
1
EF362420 A/chicken/India/NIV33487/06
2006
NA
2
EF362428 A/chicken/India/NIV33491/06
2006
NA
2
EF532637 A/chicken/Israel/397/2006
2006
NA
1
EF532640 A/chicken/Israel/625/2006
2006
NA
1
CY021519 A/chicken/Ivory Coast/1787-35/2006
2006
NA
3
DQ887063 A/chicken/Jalgaon/8824/2006
2006
NA
2
EU233677 A/chicken/Korea/IS/2006
2006
NA
2
EU233685 A/chicken/Korea/IS2/2006
2006
NA
2
EU233701 A/chicken/Korea/IS3/2006
2006
NA
2
DQ676836 A/chicken/Krasnodar/01/2006
2006
NA
2
EF205173 A/chicken/Krasnodar/123/06
2006
NA
2
DQ676832 A/chicken/Mahachkala/05/2006
2006
NA
4
CY016941 A/chicken/Nigeria/1047-30/2006
2006
NA
3
CY016949 A/chicken/Nigeria/1047-34/2006
2006
NA
3
CY016925 A/chicken/Nigeria/1047-54/2006
2006
NA
3
CY016933 A/chicken/Nigeria/1047-62/2006
2006
NA
3
CY016909 A/chicken/Nigeria/1047-8/2006
2006
NA
3
CY016278 A/chicken/Nigeria/641/2006
2006
NA
3
DQ529296 A/chicken/Nigeria/641/2006
2006
NA
3
CY016286 A/chicken/Nigeria/957-20/2006
2006
NA
3
AM262564 A/chicken/Nigeria/BA210/2006
2006
NA
3
AM262565 A/chicken/Nigeria/BA211/2006
2006
NA
3
AM262560 A/chicken/Nigeria/SO300/2006
2006
NA
3
AM262561 A/chicken/Nigeria/SO452/2006
2006
NA
3
AM262562 A/chicken/Nigeria/SO493/2006
2006
NA
3
AM262563 A/chicken/Nigeria/SO494/2006
2006
NA
3
CY016302 A/chicken/Sudan/1784-10/2006
2006
NA
3
CY016294 A/chicken/Sudan/1784-7/2006
2006
NA
3
CY020663 A/chicken/Sudan/1784-8/2006
2006
NA
3
CY021391 A/chicken/Sudan/2115-10/2006
2006
NA
3
CY020679 A/chicken/Sudan/2115-12/2006
2006
NA
3
CY020671 A/chicken/Sudan/2115-9/2006
2006
NA
3
DQ659680 A/common bussard/Bavaria/2/2006
2006
NA
4
EF165577 A/common buzzard/Bavaria/11/2006
2006
NA
4
EU152223 A/common coot/Switzerland/V544/2006
2006
NA
4
EU152223 A/common coot/Switzerland/V544/2006
2006
NA
4
AB284326 A/common goldeneye/Mongolia/12/06
2006
NA
2
EF165574 A/common pochard/Bavaria/7/2006
2006
NA
4
EU152221 A/common pochard/Switzerland/V505/2006
2006
NA
4
EU152225 A/common pochard/Switzerland/V592/2006
2006
NA
4
EU152226 A/common pochard/Switzerland/V762/2006
2006
NA
4
CY016781 A/Cygnus cygnus/Iran/754/2006
2006
NA
2
EU443572 A/Cygnus olor/Czech Republic/10814/06
2006
NA
4
EU443564 A/Cygnus olor/Czech Republic/6111/06
2006
NA
4
DQ534021 A/Cygnus olor/Czech Republic/6719/2006
2006
NA
4
CY017037 A/Cygnus olor/Italy/742/2006
2006
NA
2
CY020351 A/cygnus olor/Italy/808/2006
2006
NA
4
CY016805 A/duck/Cote d'Ivoire/1787-18/2006
2006
NA
3
EF532643 A/duck/Gaza/760/2006
2006
NA
1
EF532633 A/duck/Gaza/834/2006
2006
NA
1
AM403140 A/duck/Germany/R592/06
2006
NA
G743A
AM403141 A/duck/Germany/R603/06
2006
NA
G743A
EU233709 A/duck/Korea/Asan5/2006
2006
NA
2
EU233717 A/duck/Korea/Asan6/2006
2006
NA
2
CY017029 A/duck/Niger/914/2006
2006
NA
3
EU152217 A/duck/Switzerland/V389/2006
2006
NA
4
EU152218 A/duck/Switzerland/V426/2006
2006
NA
4
EU152219 A/duck/Switzerland/V487/2006
2006
NA
4
EF165576 A/eagle owl/Bavaria/10/2006
2006
NA
4
AM403151 A/eagle owl/Germany/R1166/06
2006
NA
G743A
EU233733 A/environment/Korea/W149/2006
2006
NA
2
EU233741 A/environment/Korea/W150/2006
2006
NA
2
EF165581 A/falcon/Bavaria/15/2006
2006
NA
4
AM403149 A/falcon/Germany/R899/06
2006
NA
G743A
EF165585 A/goldeneye duck/Bavaria/19/2006
2006
NA
4
EF165584 A/goosander/Bavaria/18/2006
2006
NA
4
EF165586 A/goosander/Bavaria/20/2006
2006
NA
4
AM403144 A/goose/Germany/R696/06
2006
NA
G743A
DQ835389 A/goose/Iraq/812NAMRU3/2006
2006
NA
4
EU401798 A/goose/Lahore-Pakistan/NARC-3321/4/2006
2006
NA
4
EF165588 A/great crested grebe/Bavaria/22/2006
2006
NA
4
EF523705 A/great crested grebe/Denmark/7498/06
2006
NA
4
AM403153 A/great crested grebe/Germany/R1226/06
2006
NA
G743A
EF523698 A/grey lag goose/Denmark/6692/06
2006
NA
4
CY017181 A/guinea fowl/Nigeria/957-12/2006
2006
NA
4
AM403148 A/gull/Germany/R882/06
2006
NA
G743A
I
SDN140068
A/Iraq/1/2006
2006
NA
4
I
SDN140067
A/Iraq/755/2006
2006
NA
4
EU152216 A/little grebe/Switzerland/V330/2006
2006
NA
4
DQ458993 A/mallard/Bavaria/1/2006
2006
NA
4
CY016797 A/mallard/Italy/835/2006
2006
NA
4
EU152222 A/mallard/Switzerland/V537/2006
2006
NA
4
EU152224 A/mallard/Switzerland/V558/2006
2006
NA
4
EU443566 A/Mergus albellus/Slovakia/Vh212/2006
2006
NA
4
EF165578 A/mute swan/Bavaria/12/2006
2006
NA
4
EF395821 A/mute swan/France/06299/2006
2006
NA
4
EU152215 A/mute swan/Switzerland/V68/2006
2006
NA
4
CY016917 A/ostrich/Nigeria/1047-25/2006
2006
NA
3
EF523699 A/peacock/Denmark/60295/06
2006
NA
4
EU443565 A/Peregrine falcon/Slovakia/Vh246/06
2006
NA
4
EF523700 A/peregrine/Denmark/6632/06
2006
NA
4
AM403139 A/pochard/Germany/R348/06
2006
NA
G743A
EU233725 A/quail/Korea/KJ4/2006
2006
NA
2
EF165580 A/swan/Bavaria/14/2006
2006
NA
4
EF165582 A/swan/Bavaria/16/2006
2006
NA
4
EF165583 A/swan/Bavaria/17/2006
2006
NA
4
EF165587 A/swan/Bavaria/21/2006
2006
NA
4
EF165573 A/swan/Bavaria/6/2006
2006
NA
4
DQ464355 A/swan/Germany/R65/2006
2006
NA
4
CY017045 A/swan/Slovenia/760/2006
2006
NA
4
EF165571 A/tufted duck/Bavaria/4/2006
2006
NA
4
EF165575 A/tufted duck/Bavaria/9/2006
2006
NA
4
EF523701 A/tufted duck/Denmark/6431/06
2006
NA
4
EF523703 A/tufted duck/Denmark/6540/06
2006
NA
4
AM403155 A/tufted duck/Germany/R1240/06
2006
NA
G743A
EU152220 A/tufted duck/Switzerland/V504/2006
2006
NA
4
I
SDN133097
A/Turkey/12/2006
2006
NA
4
I
SDN133104
A/Turkey/15/2006
2006
NA
4
I
SDN133358
A/Turkey/651242/06
2006
NA
4
I
SDN133366
A/Turkey/65596/06
2006
NA
4
EF532634 A/turkey/Israel/345/2006
2006
NA
1
EF532635 A/turkey/Israel/364/2006
2006
NA
1
EF532636 A/turkey/Israel/365/2006
2006
NA
1
EF532638 A/turkey/Israel/446/2006
2006
NA
1
CY020695 A/turkey/Ivory Coast/4372-2/2006
2006
NA
3
CY020703 A/turkey/Ivory Coast/4372-3/2006
2006
NA
3
CY020711 A/turkey/Ivory Coast/4372-4/2006
2006
NA
3
EF523702 A/whooper swan/Denmark/7224/06
2006
NA
4
EF523704 A/whooper swan/Denmark/7275/06
2006
NA
4
AB265202 A/whooper swan/Mongolia/2/06
2006
NA
2
EU443628 A/whooper swan/Scotland/1430/2006
2006
NA
4
DQ650661 A/chicken/Crimea/04/2005
2005
NA
2
DQ650665 A/chicken/Crimea/08/2005
2005
NA
2
DQ449634 A/chicken/Kurgan/05/2005
2005
NA
2
DQ323673 A/chicken/Kurgan/3/2005
2005
NA
2
EF205169 A/chicken/Omsk/14/05
2005
NA
2
EF205168 A/chicken/Suzdalka/06/05
2005
NA
4
EF205174 A/chicken/Tula/4/05
2005
NA
4
DQ449642 A/duck/Kurgan/08/2005
2005
NA
2
DQ864710 A/duck/Novosibirsk/02/05
2005
NA
2
DQ230524 A/duck/Novosibirsk/56/2005
2005
NA
2
EF205171 A/goose/Krasnoozerskoe/627/05
2005
NA
2
EF205170 A/goose/Suzdalka/10/05
2005
NA
2
DQ230523 A/grebe/Novosibirsk/29/2005
2005
NA
2
I
SDN129516
A/turkey/Turkey/1/05
2005
NA
4
EU496400 A/chicken/Egypt/0836S-NLQP/2008
2008
HA
6
EU496398 A/chicken/Egypt/086Q-NLQP/2008
2008
HA
6
EU496399 A/chicken/Egypt/088S-NLQP/2008
2008
HA
6
AM773724 A/black-necked grebe/Germany/R1393/07
2007
HA
S2
EU443553 A/chicken/Czech Republic/11242-38/07
2007
HA
S2
EU496386 A/chicken/Egypt/07125-NLQP/2007
2007
HA
6
EU496387 A/chicken/Egypt/07181-NLQP/2007
2007
HA
6
EU496388 A/chicken/Egypt/07201-NLQP/2007
2007
HA
6
EU496389 A/chicken/Egypt/07202-NLQP/2007
2007
HA
6
EU496395 A/chicken/Egypt/07632S-NLQP/2007
2007
HA
6
EU496396 A/chicken/Egypt/07665S-NLQP/2007
2007
HA
6
EU496397 A/chicken/Egypt/07701S-NLQP/2007
2007
HA
6
EU373736 A/chicken/Egypt/2628-1/2007
2007
HA
6
EU371898 A/chicken/Egypt/3044NAMRU3-CLEVB59/2007
2007
HA
6
EU371899 A/chicken/Egypt/3045NAMRU3-CLEVB60/2007
2007
HA
6
EU371907 A/chicken/Egypt/3046NAMRU3-CLEVB62/2007
2007
HA
6
EU371903 A/chicken/Egypt/3049NAMRU3-CLEVB75/2007
2007
HA
6
EU371905 A/chicken/Egypt/3051NAMRU3-CLEVB78/2007
2007
HA
6
EU371906 A/chicken/Egypt/3052NAMRU3-CLEVB104/2007
2007
HA
6
EU373735 A/chicken/Egypt/452-2/2007
2007
HA
6
EU371908 A/chicken/Egypt/9383NAMRU3-CLEVB112/2007
2007
HA
6
EU371909 A/chicken/Egypt/9384NAMRU3-CLEVB118/2007
2007
HA
6
EU371910 A/chicken/Egypt/9385NAMRU3-CLEVB125/2007
2007
HA
6
EU371911 A/chicken/Egypt/9386NAMRU3-CLEVB136/2007
2007
HA
6
EU371912 A/chicken/Egypt/9387NAMRU3-CLEVB148/2007
2007
HA
6
EU371913 A/chicken/Egypt/9388NAMRU3-CLEVB149/2007
2007
HA
6
EU371914 A/chicken/Egypt/9389NAMRU3-CLEVB150/2007
2007
HA
6
EU371915 A/chicken/Egypt/9390NAMRU3-CLEVB157/2007
2007
HA
6
EU371916 A/chicken/Egypt/9391NAMRU3-CLEVB158/2007
2007
HA
6
EU371917 A/chicken/Egypt/9392NAMRU3-CLEVB167/2007
2007
HA
6
EU371918 A/chicken/Egypt/9396NAMRU3-CLEVB187/2007
2007
HA
6
EU371919 A/chicken/Egypt/9397NAMRU3-CLEVB188/2007
2007
HA
6
EU371922 A/chicken/Egypt/9400NAMRU3-CLEVB211/2007
2007
HA
6
EU183326 A/chicken/Egypt/F6/2007
2007
HA
6
EU183328 A/chicken/Egypt/R2/2007
2007
HA
6
EU183329 A/chicken/Egypt/R3/2007
2007
HA
6
EU183332 A/chicken/Egypt/R6/2007
2007
HA
6
EU163431 A/chicken/Krasnodar/300/07
2007
HA
S2
CY029948 A/chicken/Kuwait/KISR2/2007
2007
HA
S2
CY029956 A/chicken/Kuwait/KISR3/2007
2007
HA
S2
CY029964 A/chicken/Kuwait/KISR5/2007
2007
HA
S2
CY029972 A/chicken/Kuwait/KISR6/2007
2007
HA
S2
CY029980 A/chicken/Kuwait/KISR7/2007
2007
HA
S2
CY029988 A/chicken/Kuwait/KISR8/2007
2007
HA
S2
CY029996 A/chicken/Kuwait/KISR9/2007
2007
HA
S2
EF474450 A/chicken/Moscow/2/2007
2007
HA
S2
EU148356 A/chicken/Nigeria/1071-1/2007
2007
HA
S3
EU148404 A/chicken/Nigeria/1071-10/2007
2007
HA
S3
EU148412 A/chicken/Nigeria/1071-15/2007
2007
HA
S3
EU148420 A/chicken/Nigeria/1071-22/2007
2007
HA
S3
EU148428 A/chicken/Nigeria/1071-23/2007
2007
HA
S3
EU148436 A/chicken/Nigeria/1071-29/2007
2007
HA
S3
EU148364 A/chicken/Nigeria/1071-3/2007
2007
HA
S3
EU148445 A/chicken/Nigeria/1071-30/2007
2007
HA
S3
EU148372 A/chicken/Nigeria/1071-4/2007
2007
HA
S3
EU148380 A/chicken/Nigeria/1071-5/2007
2007
HA
S3
EU148388 A/chicken/Nigeria/1071-7/2007
2007
HA
S3
EU148396 A/chicken/Nigeria/1071-9/2007
2007
HA
S3
EU402398 A/chicken/Rostov/22-1/2007
2007
HA
S2
EU401751 A/chicken/Rostov-on-Don/35/2007
2007
HA
S2
EF605603 A/chicken/Russia_Krasnodar/2/2007
2007
HA
S2
AM498628 A/common pochard/France/06167-2.1/2006
2007
HA
S4
EU371907 A/crow/Egypt/9382NAMRU3-CLEVB111/2007
2007
HA
6
EU257631 A/Cygnus cygnus/Krasnodar/329/07
2007
HA
S2
EU443555 A/Cygnus olor/Czech Republic/10732/07
2007
HA
S2
EU496391 A/duck/Egypt/07264S-NLQP/2007
2007
HA
6
EU496392 A/duck/Egypt/07322S-NLQP/2007
2007
HA
6
EU373737 A/duck/Egypt/5169-1/2007
2007
HA
6
EU371921 A/duck/Egypt/9399NAMRU3-CLEVB202/2007
2007
HA
6
EU183331 A/duck/Egypt/R5/2007
2007
HA
6
EU496393 A/goose/Egypt/07364S-NLQP/2007
2007
HA
6
EU183330 A/goose/Egypt/R4/2007
2007
HA
6
EU424135 A/houbara bustard/Saudi Arabia/6732-1/2007
2007
HA
S2
EU441929 A/muscovy duck/Rostov-on-Don/51/2007
2007
HA
S2
AM749442 A/mute swan/Germany/R1349/07
2007
HA
S2
AM749443 A/mute swan/Germany/R1359/07
2007
HA
S2
I
SDN230945
A/Nigeria/6e/07
2007
HA
3S
EU401796 A/peacock/Mansehra-Pakistan/NARC-7558/02/2007 2007
HA
S2
EU441937 A/pigeon/Rostov-on-Don/6/2007
2007
HA
S2
EU496385 A/quail/Egypt/07120-NLQP/2007
2007
HA
6
EU486855 A/starling/Rostov-on-Don/39/07
2007
HA
S2
EU443554 A/turkey/Czech Republic/10309-3/07
2007
HA
S2
EU496394 A/turkey/Egypt/07444S-NLQP/2007
2007
HA
6
EU371920 A/turkey/Egypt/9398NAMRU3-CLEVB195/2007
2007
HA
6
EU401794 A/turkey/Islamabad-Pakistan/NARC-7871/02/2007
2007
HA
S2
I
SDN140057
A/Azerbaijan/001-161/2006
2006
HA
S2
I
SDN140058
A/Azerbaijan/002-115/2006
2006
HA
S2
I
SDN140054
A/Azerbaijan/006-207/2006
2006
HA
S2
I
SDN140055
A/Azerbaijan/008-208/2006
2006
HA
S2
I
SDN140056
A/Azerbaijan/011-162/2006
2006
HA
S2
EF165057 A/buzzard/Bavaria/13/2006
2006
HA
S4
EF165049 A/buzzard/Bavaria/5/2006
2006
HA
S4
EF523687 A/buzzard/Denmark/6370/06
2006
HA
S4
AM403474 A/Canada goose/Germany/R1207/06
2006
HA
S4
AM403461 A/Canada goose/Germany/R71/06
2006
HA
S4
EF395844 A/cat/Austria/649/2006
2006
HA
S4
DQ864720 A/cat/Dagestan/87/06
2006
HA
S4
DQ643982 A/cat/Germany/606/2006
2006
HA
S4
AM403468 A/cat/Germany/R606/06
2006
HA
S4
DQ864715 A/chicken/Adygea/203/06
2006
HA
S3
AM400974 A/chicken/Burkina Faso/01.03/2006
2006
HA
S3
AM400973 A/chicken/Burkina Faso/13.1/2006
2006
HA
S3
CY016811 A/chicken/Cote d'Ivoire/1787-34/2006
2006
HA
S3
EU372943 A/chicken/Egypt/06207-NLQP/2006
2006
HA
6
EU372944 A/chicken/Egypt/06459-3-NLQP/2006
2006
HA
6
EU372945 A/chicken/Egypt/06495-NLQP/2006
2006
HA
6
EU372946 A/chicken/Egypt/06541-NLQP/2006
2006
HA
6
EU496383 A/chicken/Egypt/06553-NLQP/2006
2006
HA
6
EU496384 A/chicken/Egypt/06612-NLQP/2006
2006
HA
6
EU372947 A/chicken/Egypt/06959-NLQP/2006
2006
HA
6
CY020645 A/chicken/Egypt/2253-1/2006
2006
HA
6
EU183323 A/chicken/Egypt/F3/2006
2006
HA
6
EU183324 A/chicken/Egypt/F4/2006
2006
HA
6
EU183327 A/chicken/Egypt/R1/2006
2006
HA
6
EF532628 A/chicken/Gaza/450/2006
2006
HA
S1, 6
EF532630 A/chicken/Gaza/713/2006
2006
HA
S1, 6
EF532631 A/chicken/Gaza/714/2006
2006
HA
S1, 6
EF362418 A/chicken/India/NIV33487/06
2006
HA
S2
EF362426 A/chicken/India/NIV33491/06
2006
HA
S2
EF532626 A/chicken/Israel/397/2006
2006
HA
S1, 6
EF532629 A/chicken/Israel/625/2006
2006
HA
S1, 6
DQ673901 A/chicken/Ivory Coast/1787/2006
2006
HA
S3
CY021517 A/chicken/Ivory Coast/1787-35/2006
2006
HA
S3
EU233691 A/chicken/Korea/CA7/2006
2006
HA
S2
EU233675 A/chicken/Korea/IS/2006
2006
HA
S2
EU233683 A/chicken/Korea/IS2/2006
2006
HA
S2
EU233699 A/chicken/Korea/IS3/2006
2006
HA
S2
DQ676834 A/chicken/Krasnodar/01/2006
2006
HA
S2
EF205159 A/chicken/Krasnodar/123/06
2006
HA
S2
DQ864718 A/chicken/Krasnodar/199/06
2006
HA
S2
AM262541 A/chicken/Lagos.NIE/BA209/2006
2006
HA
S3
AM262542 A/chicken/Lagos.NIE/BA210/2006
2006
HA
S3
AM262543 A/chicken/Lagos.NIE/BA211/2006
2006
HA
S3
AM262546 A/chicken/Lagos.NIE/SO300/2006
2006
HA
S3
AM262547 A/chicken/Lagos.NIE/SO452/2006
2006
HA
S3
AM262553 A/chicken/Lagos.NIE/SO493/2006
2006
HA
S3
AM262572 A/chicken/Lagos.NIE/SO494/2006
2006
HA
S3
DQ838517 A/chicken/Niger/2130-7/2006
2006
HA
S3
DQ838516 A/chicken/Niger/2130-8/2006
2006
HA
S3
CY016939 A/chicken/Nigeria/1047-30/2006
2006
HA
S3
CY016947 A/chicken/Nigeria/1047-34/2006
2006
HA
S3
CY016923 A/chicken/Nigeria/1047-54/2006
2006
HA
S3
CY016931 A/chicken/Nigeria/1047-62/2006
2006
HA
S3
CY016907 A/chicken/Nigeria/1047-8/2006
2006
HA
S3
CY016276 A/chicken/Nigeria/641/2006
2006
HA
S3
CY016284 A/chicken/Nigeria/957-20/2006
2006
HA
S3
AM400975 A/chicken/Nigeria/SO227/2006
2006
HA
S3
AM400976 A/chicken/Nigeria/SO294/2006
2006
HA
S3
AM400977 A/chicken/Nigeria/SO457/2006
2006
HA
S3
AM400978 A/chicken/Nigeria/SO461/2006
2006
HA
S3
AM400979 A/chicken/Nigeria/SO470/2006
2006
HA
S3
AM400980 A/chicken/Nigeria/SO478/2006
2006
HA
S3
AM400981 A/chicken/Nigeria/SO485/2006
2006
HA
S3
DQ862003 A/chicken/Sudan/1784/2006
2006
HA
S3
CY016300 A/chicken/Sudan/1784-10/2006
2006
HA
S3
CY016292 A/chicken/Sudan/1784-7/2006
2006
HA
S3
CY020661 A/chicken/Sudan/1784-8/2006
2006
HA
S3
CY021389 A/chicken/Sudan/2115-10/2006
2006
HA
S3
CY020677 A/chicken/Sudan/2115-12/2006
2006
HA
S3
CY020669 A/chicken/Sudan/2115-9/2006
2006
HA
S3
DQ864719 A/chicken/Volgograd/236/06
2006
HA
S4
DQ659679 A/common bussard/Bavaria/2/2006
2006
HA
S4
EF165055 A/common buzzard/Bavaria/11/2006
2006
HA
S4
AM403463 A/common buzzard/Germany/R306/06
2006
HA
S4
AM408213 A/common buzzard/Germany/R870/06
2006
HA
S4
EF110519 A/common coot/Switzerland/V544/06
2006
HA
S4
EF165051 A/common pochard/Bavaria/7/2006
2006
HA
S4
EU016355 A/common pochard/Switzerland/V505/2006
2006
HA
S4
EU016358 A/common pochard/Switzerland/V592/2006
2006
HA
S4
EU016359 A/common pochard/Switzerland/V762/2006
2006
HA
S4
AM403469 A/coot/Germany/R655/06
2006
HA
S4
AM408209 A/cormorant/Germany/R292/06
2006
HA
S4
CY016779 A/Cygnus cygnus/Iran/754/2006
2006
HA
S2
EU443541 A/Cygnus olor/Czech Republic/10662/06
2006
HA
S4
EU443542 A/Cygnus olor/Czech Republic/10814/06
2006
HA
S4
EU443542 A/Cygnus olor/Czech Republic/10814/06
2006
HA
S4
DQ515984 A/Cygnus olor/Czech Republic/5170/2006
2006
HA
S4
EU443543 A/Cygnus olor/Czech Republic/5583/06
2006
HA
S4
EU443547 A/Cygnus olor/Czech Republic/5761/06
2006
HA
S4
EU443545 A/Cygnus olor/Czech Republic/5963/06
2006
HA
S4
EU443549 A/Cygnus olor/Czech Republic/6111/06
2006
HA
S4
EU443544 A/Cygnus olor/Czech Republic/6206/06
2006
HA
S4
EU443551 A/Cygnus olor/Czech Republic/6461/06
2006
HA
S4
EU443546 A/Cygnus olor/Czech Republic/7185/06
2006
HA
S4
EU443548 A/Cygnus olor/Czech Republic/7208/06
2006
HA
S4
CY017035 A/Cygnus olor/Italy/742/2006
2006
HA
S4
CY020349 A/cygnus olor/Italy/808/2006
2006
HA
S4
DQ435200 A/domestic cat/Iraq/820/2006
2006
HA
S4
DQ435201 A/domestic goose/Iraq/812/2006
2006
HA
S4
CY016803 A/duck/Cote d'Ivoire/1787-18/2006
2006
HA
S4
CY016899 A/duck/Egypt/2253-3/2006
2006
HA
6
EU183325 A/duck/Egypt/F5/2006
2006
HA
6
EF532632 A/duck/Gaza/760/2006
2006
HA
S1, 6
EF532622 A/duck/Gaza/834/2006
2006
HA
S1, 6
AM403464 A/duck/Germany/R338/06
2006
HA
S4
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008
AM403466 A/duck/Germany/R592/06
2006
HA
S4
AM403467 A/duck/Germany/R603/06
2006
HA
S4
AM403470 A/duck/Germany/R751/06
2006
HA
S4
EU233707 A/duck/Korea/Asan5/2006
2006
HA
S2
EU233715 A/duck/Korea/Asan6/2006
2006
HA
S2
CY017027 A/duck/Niger/914/2006
2006
HA
S3
EU016352 A/duck/Switzerland/V389/2006
2006
HA
S4
EU016353 A/duck/Switzerland/V426/2006
2006
HA
S4
EU016354 A/duck/Switzerland/V487/2006
2006
HA
S4
EF165054 A/eagle owl/Bavaria/10/2006
2006
HA
S4
AM403473 A/eagle owl/Germany/R1166/06
2006
HA
S4
EU233731 A/environment/Korea/W149/2006
2006
HA
S2
EU233739 A/environment/Korea/W150/2006
2006
HA
S2
EF165059 A/falcon/Bavaria/15/2006
2006
HA
S4
AM408211 A/falcon/Germany/R899/06
2006
HA
S4
EF523696 A/fowl/Denmark/60296/06
2006
HA
S4
EF165063 A/goldeneye duck/Bavaria/19/2006
2006
HA
S4
EF165062 A/goosander/Bavaria/18/2006
2006
HA
S4
EF165064 A/goosander/Bavaria/20/2006
2006
HA
S4
EF110518 A/goosander/Switzerland/V82/06
2006
HA
S4
EF110518 A/goosander/Switzerland/V82/06
2006
HA
S4
AM408210 A/goose/Germany/R696/06
2006
HA
S4
EU401795 A/goose/Lahore-Pakistan/NARC-3321/4/2006
2006
HA
S2
EF165066 A/great crested grebe/Bavaria/22/2006
2006
HA
S4
EF523695 A/great crested grebe/Denmark/7498/06
2006
HA
S4
AM408212 A/great crested grebe/Germany/R1226/06
2006
HA
S4
EF523688 A/grey lag goose/Denmark/6692/06
2006
HA
S4
AM498631 A/greylag goose/France/06310/2006
2006
HA
S4
CY017179 A/guinea fowl/Nigeria/957-12/2006
2006
HA
S3
AM408215 A/gull/Germany/R882/06
2006
HA
S4
AM400971 A/hooded vulture/Burkina Faso/1/2006
2006
HA
S3
AM400972 A/hooded vulture/Burkina Faso/2/2006
2006
HA
S3
EF090648 A/hooded vulture/Burkina Faso/5346-10/2006
2006
HA
S3
EU016351 A/little grebe/Switzerland/V330/2006
2006
HA
S4
DQ458992 A/mallard/Bavaria/1/2006
2006
HA
S4
DQ449031 A/mallard/Italy/332/2006
2006
HA
S4
CY016795 A/mallard/Italy/835/2006
2006
HA
S4
EU016356 A/mallard/Switzerland/V537/2006
2006
HA
S4
EU016357 A/mallard/Switzerland/V558/2006
2006
HA
S4
EU443550 A/Mergus albellus/Slovakia/Vh212/2006
2006
HA
S4
EF165056 A/mute swan/Bavaria/12/2006
2006
HA
S4
EF395820 A/mute swan/France/06299/2006
2006
HA
S4
AM498630 A/mute swan/France/06303/2006
2006
HA
S4
AM498632 A/mute swan/France/06631a/2006
2006
HA
S4
AM403460 A/mute swan/Germany/R65/06
2006
HA
S4
AM403471 A/mute swan/Germany/R854/06
2006
HA
S4
EU016350 A/mute swan/Switzerland/V68/2006
2006
HA
S4
CY016915 A/ostrich/Nigeria/1047-25/2006
2006
HA
S3
EF523689 A/peacock/Denmark/60295/06
2006
HA
S4
EU443552 A/Peregrine falcon/Slovakia/Vh246/06
2006
HA
S4
EF523690 A/peregrine/Denmark/6632/06
2006
HA
S4
EU233723 A/quail/Korea/KJ4/2006
2006
HA
S2
AM492165 A/stone marten/Germany/R747/2006
2006
HA
S4
AM403475 A/stork/Germany/R1239/06
2006
HA
S4
EF395845 A/swan/Austria/216/2006
2006
HA
S4
EF165058 A/swan/Bavaria/14/2006
2006
HA
S4
EF165060 A/swan/Bavaria/16/2006
2006
HA
S4
EF165061 A/swan/Bavaria/17/2006
2006
HA
S4
EF165065 A/swan/Bavaria/21/2006
2006
HA
S4
EF165050 A/swan/Bavaria/6/2006
2006
HA
S4
DQ464354 A/swan/Germany/R65/2006
2006
HA
S4
DQ440535 A/swan/Iran/754/2006
2006
HA
S2
CY017043 A/swan/Slovenia/760/2006
2006
HA
S4
EF165048 A/tufted duck/Bavaria/4/2006
2006
HA
S4
EF165052 A/tufted duck/Bavaria/8/2006
2006
HA
S4
EF165053 A/tufted duck/Bavaria/9/2006
2006
HA
S4
EF523691 A/tufted duck/Denmark/6431/06
2006
HA
S4
EF523692 A/tufted duck/Denmark/6540/06
2006
HA
S4
AM408216 A/tufted duck/Germany/R1240/06
2006
HA
S4
EF547197 A/tufted duck/Switzerland/V504/06
2006
HA
S4
CY020653 A/turkey/Egypt/2253-2/2006
2006
HA
S1, 6
EU183321 A/turkey/Egypt/F1/2006
2006
HA
6
EU183322 A/turkey/Egypt/F2/2006
2006
HA
6
AM498629 A/turkey/France/06222-1.1/2006
2006
HA
S4
AM403472 A/turkey/Germany/R1077/06
2006
HA
S4
EF532623 A/turkey/Israel/345/2006
2006
HA
S1, 6
EF532624 A/turkey/Israel/364/2006
2006
HA
S1, 6
EF532625 A/turkey/Israel/365/2006
2006
HA
S1, 6
EF532627 A/turkey/Israel/446/2006
2006
HA
S1, 6
CY020693 A/turkey/Ivory Coast/4372-2/2006
2006
HA
2B
CY020701 A/turkey/Ivory Coast/4372-3/2006
2006
HA
2B
CY020709 A/turkey/Ivory Coast/4372-4/2006
2006
HA
2B
EF523693 A/whooper swan/Denmark/7224/06
2006
HA
S4
EF523694 A/whooper swan/Denmark/7275/06
2006
HA
S4
AM403462 A/whooper swan/Germany/R88/06
2006
HA
S4
AB263752 A/whooper swan/Mongolia/2/06
2006
HA
S3
EU443629 A/whooper swan/Scotland/1430/2006
2006
HA
S4
EU443549 Cygnus olor/Czech Republic/6111/06
2006
HA
S4
DQ449632 A/chicken/Kurgan/05/2005
2005
HA
S2
DQ323672 A/chicken/Kurgan/3/2005
2005
HA
S2
EF205155 A/chicken/Omsk/14/05
2005
HA
S2
EF205154 A/chicken/Suzdalka/06/05
2005
HA
S2
DQ864716 A/chicken/Tambov/570-2/05
2005
HA
6
EF205160 A/chicken/Tula/4/05
2005
HA
6
DQ449640 A/duck/Kurgan/08/2005
2005
HA
S2
DQ864711 A/duck/Novosibirsk/02/05
2005
HA
S2
DQ230522 A/duck/Novosibirsk/56/2005
2005
HA
S2
DQ676840 A/goose/Krasnoozerka/627/2005
2005
HA
S2
EF205157 A/goose/Krasnoozerskoe/627/05
2005
HA
S2
EF205156 A/goose/Suzdalka/10/05
2005
HA
S2
DQ230521 A/grebe/Novosibirsk/29/2005
2005
HA
S2
EF042624 A/teal/Egypt/14051-NAMRU3/2005
2005
HA
6
EF205158 A/turkey/Suzdalka/12/05
2005
HA
S2
DQ407519 A/turkey/Turkey/1/2005
2005
HA
S4
Nature Precedings : hdl:10101/npre.2008.459.4 : Posted 7 May 2008