# Supplement 2008-2010 (no. 48) to the White-Kauffmann-Le Minor scheme.

Sylvie Issenhuth-Jeanjean, Peter Roggentin, Matthew Mikoleit, Martine Guibourdenche, Elizabeth de Pinna, Satheesh Nair, Patricia I Fields, François-Xavier Weill

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Sylvie Issenhuth-Jeanjean, Peter Roggentin, Matthew Mikoleit, Martine Guibourdenche, Elizabeth de Pinna, et al.. Supplement 2008-2010 (no. 48) to the White-Kauffmann-Le Minor scheme.. Research in Microbiology, 2014, 165 (7), pp.526-30. 10.1016/j.resmic.2014.07.004. pasteur-01104894

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1

2 Supplement 2008-2010 (no. 48) to the White-Kauffmann-Le Minor scheme

3

4

5 Sylvie Issenhuth-Jeanjean<sup>a</sup>, Peter Roggentin<sup>b</sup>, Matthew Mikoleit<sup>c</sup>, Martine Guibourdenche<sup>a</sup>,

6 Elizabeth de Pinna<sup>d</sup>, Satheesh Nair<sup>d</sup>, Patricia I. Fields<sup>c</sup>, François-Xavier Weill<sup>a*</sup>

7

8 <sup>a</sup>Institut Pasteur, WHO Collaborating Centre for Reference and Research on *Salmonella*,

9 Unité des Bactéries Pathogènes Entériques, Paris, France

10 <sup>b</sup>Salmonella-Zentrale, Institut für Hygiene und Umwelt, Hamburg, Germany

11 <sup>c</sup>Centers for Disease Control and Prevention, Atlanta, Georgia, USA

12 <sup>d</sup>Public Health England, Gastrointestinal Bacteria Reference Unit, Colindale, UK

13

14 In memory of Prof. Michel-Yvan Popoff (1945-2013) who headed the WHO Collaborating

15 Centre for Reference and Research on *Salmonella* at the Pasteur Institute between 1989 and

16 2003

17

18 * Corresponding author

19 WHO Collaborating Centre for Reference and Research on *Salmonella*, Unité des Bactéries

20 Pathogènes Entériques, Institut Pasteur, 28 rue du Dr Roux, 75724 Paris cedex 15, France.

21 Tel: +33 1 45 68 83 45. Fax: +33 1 45 68 88 37. Email: fxweill@pasteur.fr
22 **Abstract**

This supplement (no. 48) of the White-Kauffmann-Le Minor scheme reports on the characterization of 63 new *Salmonella* serovars and 25 new variants of previously described *Salmonella* serovars recognized by the WHO Collaborating Centre for Reference and Research on *Salmonella* between 2008 and 2010. Forty-four new serovars were assigned to *Salmonella enterica* subspecies *enterica*, 12 to subspecies *salamae*, two to subspecies *arizonae*, two to subspecies *diarizonae* and three to subspecies *houtenae*. All these new serovars or new variants are described with their multilocus sequence type.

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Keywords: *Salmonella*; Serovars; Taxonomy; MLST; White-Kauffmann-Le Minor scheme

32

33
The genus *Salmonella* consists of only two species, *Salmonella enterica* and *S. bongori*. *S. enterica* is divided into six subspecies: *S. enterica* subsp. *enterica*, *S. enterica* subsp. *salamae*, *S. enterica* subsp. *arizonae*, *S. enterica* subsp. *diarizonae*, *S. enterica* subsp. *houtenae*, and *S. enterica* subsp. *indica*. This nomenclature reflects present understanding of *Salmonella* taxonomy [1, 2]. The species called *S. subterranea* [3] does not belong in the genus *Salmonella* (Table 1, Figure).

Serovars belonging to *S. enterica* subsp. *enterica* are typically designated by a name usually related to the geographical place where the serovar was first isolated [7]. The serovar name is written in non-italicized Roman letters and the first letter capitalized. Serovars belonging to other subspecies are designated by their antigenic formulae, following the subspecies name.

The antigenic formulae of *Salmonella* serovars are available in the White-Kauffmann-Le Minor scheme [7] at: http://www.pasteur.fr/sante/clre/cadrecnr/salmoms/WKLM_En.pdf

Updating this scheme is the responsibility of the WHO Collaborating Centre for Reference and Research on *Salmonella* (WHO-Salm), Institut Pasteur, Paris, France. The current edition (9th) issued in 2007 comprises antigenic formulae validated as of January 1st 2007. In 2010, Supplement 47 reported the characterization of 70 new *Salmonella* serovars recognized between January 1st 2003 and December 31st 2007 [8].

Supplement 48 reports the characterization of 63 new *Salmonella* serovars recognized between January 1st 2008 and December 31st 2010 by the WHO-Salm: 44 were assigned to *S. enterica* subsp. *enterica*, 12 to subsp. *salamae*, two to subsp. *arizonae*, two to subsp. *diarizonae* and three to subspecies *houtenae* (Table 2).

It is useful to note that strain 9736/07, erroneously described in the Supplement 47 [8] as the reference strain of *S. bongori* serovar 1,9,12:d:e,n,x has been reassigned to *S. enterica* subsp. *enterica* on the basis of its biochemical characteristics, its diphasic nature, *rpoB* sequence (Figure 1), and multilocus sequence type [6]. This strain is now considered as the
59 reference strain of the new serovar Viikki.

60
61 Among the 63 new serovars identified, 21 were received from the Gastrointestinal Bacteria
62 Reference Unit of Public Health England (PHE), Colindale, UK. These 21 new serovars were
63 isolated between 1994 and 2004 and correspond to all potential new serovars identified at
64 PHE during this period.

65
66 New variants of previously described *Salmonella* serovars are provided in Table 3

67
68 For the first time, the multilocus sequence type has been indicated for all the 63 new serovars
69 as well as for the 25 new variants of previously described *Salmonella* serovars. Multilocus
70 sequence typing (MLST) has been performed both at the WHO-CC or at the PHE according
71 to Achtman *et al.* [6]. The data have been submitted to the *Salmonella enterica* MLST
72 database (http://mlst.warwick.ac.uk/mlst/dbs/Senterica).

73
74 The present number of serovars per species and subspecies is given in Table 4.

75
76 The next issue of the White-Kauffmann-Le Minor scheme, which will contain MLST data,
77 is expected in December 2015.

78

79

80

81

82
83 Acknowledgements

84 We thank P. Dunker (Institut für Hygiene and Umwelt), S. Simington (CDC), and M.S. Van
85 Duyne (CDC), J. Bale (PHE) and D. Meunier (PHE) for their technical assistance.

86
87 We thank all the National Reference Centres that sent the new serovars or variants to the
88 WHO-Salm: Institut für Medizinische Mikrobiologie und Hygiene (IMED), Graz, Austria (C.
89 Kornschober); Institut scientifique de Santé Publique (ISSP), Brussels, Belgium (S. Bertrand);
90 Institut National des Laboratoires de la Santé (INLASA), La Paz, Bolivia (R.M. Ruiz); Public
91 Health Agency of Canada (PHAC), Winnipeg, Canada (H. Tabor), Korean CDC (KCDC),
92 Seoul, South Korea (D-Y. Lee); National Public Health Institute (NPHI), Helsinki, Finland
93 (S. Lukinmaa & A. Siitonen); Institut Pasteur (IP), Paris, France (F.-X. Weill & S. Le Hello);
94 Federal Institute for Risk Assessment (BfR), Berlin, Germany (I. Szabo & C. Dorn); Institut
95 für Hygiene and Umwelt (IHU), Hamburg, Germany (P. Roggentin); Robert Koch Institute
96 (RKI), Wernigerode, (W. Rabsch); Istituto Superiore di Sanità (ISS), Roma, Italy (I. Luzzi);
97 Norwegian Institute of Public Health (NIPH), Oslo, Norway (J. Lassen & A.-L. Wester);
98 Medical University of Gdansk (MUG), Gdansk, Poland (D. Kunikowska); Swedish Institute
99 for Infectious Disease Control (SMI), Solna, Sweden (R. Wollin); Institute for Food Safety
100 and Hygiene (IFSH), Zürich, Switzerland (H. Hächler); Public Health England (PHE),
101 Colindale, UK (E. de Pinna); Centers for Disease Control and Prevention (CDC), Atlanta,
102 USA (M. Mikoleit & P.I. Fields).

103

104
105 **References**

106 [1] Judicial Commission, The type species of the genus *Salmonella* Lignieres 1900 is
107 *Salmonella enterica* (ex Kauffmann and Edwards 1952) Le Minor and Popoff 1987, with
108 the type strain LT2<sup>T</sup>, and conservation of the epithet *enterica* in *Salmonella enterica* over
109 all earlier epithets that may be applied to this species. Opinion 80, Int J Syst Evol
110 Microbiol 55 (2005) 519-520.

111 [2] Tindall BJ, Grimont PAD, Garrity GM, Euzeby JP, Nomenclature and taxonomy of the
genus *Salmonella*, Int J Syst Evol Microbiol 55 (2005) 521-524.

113 [3] Shelobolina ES, Sullivan SA, O'Neill KR. Nevin KP, Lovley DR, Isolation, 114 characterization, and U(VI)-reducing potential of a facultatively anaerobic, acid-resistant 115 bacterium from low-pH, nitrate- and U(VI)-contaminated subsurface sediment and 116 description of *Salmonella subterranea* sp. nov, Appl Environ Microbiol 70 (2004) 2959-117 2965.

118 [4] Popoff MY, Guidelines for the preparation of *Salmonella* antisera, 6th Revision, WHO 119 Collaborating Centre for Reference and Research on *Salmonella*, Institut Pasteur, Paris, 120 2001.

121 [5] Ait Tayeb L, Ageron E, Grimont F, Grimont PA, Molecular phylogeny of the genus 122 Pseudomonas based on *rpoB* sequences and application for the identification of isolates, 123 Res Microbiol 156 (2005) 763-773.

124 [6] Achtman M, Wain J, Weill FX, Nair S, Zhou Z, Sangal V, Krauland MG, Hale JL, 125 Harbottle H, Uesbeck A, Dougan G, Harrison LH, Brisse S; S. *enterica* MLST Study 126 Group, Multilocus sequence typing as a replacement for serotyping in *Salmonella* 127 *enterica*, PLoS Pathog 8 (2012) e1002776.

128
129

[7] Grimont PAD, Weill FX, Antigenic formulae of the *Salmonella* serovars, ninth ed., WHO

130

Collaborating Centre for Reference and Research on *Salmonella*, Institut Pasteur, Paris,

131

2007. Available at :

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http://www.pasteur.fr/sante/clre/cadrecnr/salmoms/WKLM_En.pdf

133

[8] Guibourdenche M, Roggentin P, Mikoleit M, Fields PI, Bockemühl J, Grimont PA, Weill

134

FX, Supplement 2003-2007 (No. 47) to the White-Kauffmann-Le Minor scheme, Res

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Microbiol 161 (2010) 26-29.

136

137

138
139 **Table 1.** Atypical characteristics of the reference strain of "*Salmonella subterranea*" (strain

140 CIP 109002)

141

<table><thead><tr><th></th><th>Strain CIP109002</th></tr></thead><tbody><tr><td>Biochemical tests<sup>a</sup></td><td>Indole<sup>+</sup>, citrate<sup>-</sup>, LDC<sup>-</sup>, H2S<sup>-</sup></td></tr><tr><td>Serotyping<sup>b</sup></td><td>No agglutination</td></tr><tr><td><i>rpoB</i> sequencing<sup>c</sup></td><td>Not clustered with <i>Salmonella</i> strains</td></tr><tr><td>MLST<sup>d</sup></td><td>Not typable</td></tr></tbody></table>

142 <sup>a</sup>This strain was not pigmented in yellow in classical culture media as described by the authors

143 <sup>b</sup>Performed with antisera produced according to reference 4

144 <sup>c</sup>See Figure

145 <sup>d</sup>Performed according to reference 6. More than 2,300 strains belonging to >650 serotypes of *S. enterica* subsp. *enterica* (n=1,765), *salamae* (n=227), *arizonae* (n=51), *diarizonae* (n=165), *houtenae* (n=63), *indica* (n=15), and *S. bongori* (n=31) have been MLST typed at the WHO-CC without encountering not typable *Salmonella* strains

149

150

151
<table><caption>1 **Table 2.** New *Salmonella* serovars recognized by the WHO Collaborating Centre for Reference and Research on *Salmonella*, 2008-2010.</caption><thead><tr><th>Serovar name</th><th>Antigenic formula</th><th>Other characters<sup>b</sup></th><th>Source</th><th>Area of isolation (or contamination)</th><th>Year</th><th>IP strain number</th><th>Received from<sup>c</sup></th><th>MLST type</th></tr></thead><tbody><tr><td colspan="9"><strong>Salmonella enterica subsp. enterica</strong></td></tr><tr><td>Viikki<sup>a</sup></td><td>1,9,12:d,e,n,x</td><td rowspan="10">ONPG<sup>+</sup>, Gal<sup>+</sup></td><td>Cattle</td><td>Finland</td><td>2007</td><td>9736/07</td><td>NPHI</td><td>ST1743</td></tr><tr><td>Bruxelles</td><td>41:y:-</td><td>Spice</td><td>Belgium</td><td>2007</td><td>9764/08</td><td>ISSP</td><td>ST1735</td></tr><tr><td>Forges</td><td>47:z<sub>4</sub>,z<sub>23</sub>:1,2</td><td>Human feces</td><td>France</td><td>2007</td><td>9765/08</td><td>IP</td><td>ST1700</td></tr><tr><td>Trumersee</td><td>1,6,14,25:k:e,n,x</td><td>Rosemary herb</td><td>Austria</td><td>2008</td><td>9789/08</td><td>IMED</td><td>ST1701</td></tr><tr><td>Pajala</td><td>13,23:z<sub>41</sub>:e,n,z<sub>15</sub></td><td>Bird</td><td>Sweden</td><td>2008</td><td>9806/08</td><td>SMI</td><td>ST1704</td></tr><tr><td>Chapuri</td><td>13,23:i:z<sub>6</sub></td><td>Human feces</td><td>USA</td><td>2007</td><td>9811/08</td><td>CDC</td><td>ST1705</td></tr><tr><td>Baltimore</td><td>35:y:1,w</td><td>Human feces</td><td>USA</td><td>2008</td><td>9812/08</td><td>CDC</td><td>ST1706</td></tr><tr><td>Landeskrone</td><td>39:-:1,6</td><td>Human feces</td><td>Germany</td><td>2007</td><td>9814/08</td><td>IHU</td><td>ST1707</td></tr><tr><td>Rosslyn</td><td>51:y:1,2</td><td>Snake</td><td>Niger</td><td>2008</td><td>9817/08</td><td>PHAC</td><td>ST1708</td></tr><tr><td>Kakikoka</td><td>30:b:z<sub>6</sub></td><td>Human feces</td><td>Benin</td><td>2008</td><td>9829/08</td><td>RKI</td><td>ST627</td></tr><tr><td>Parabiago</td><td>28:c:1,w</td><td>Human feces</td><td>Italy</td><td>2008</td><td>9837/08</td><td>ISS</td><td>ST1709</td></tr><tr><td>Durance</td><td>13,22:z:z<sub>6</sub></td><td>Human feces</td><td>France</td><td>2008</td><td>9843/08</td><td>IP</td><td>ST1710</td></tr><tr><td>Santaclara</td><td>3,10:z<sub>35</sub>:1,5</td><td rowspan="3">Mal<sup>+</sup>, Sal<sup>+</sup>, Gal<sup>+</sup></td><td>Human feces</td><td>USA</td><td>2007</td><td>9847/08</td><td>CDC</td><td>ST1711</td></tr><tr><td>Capetown</td><td>3,15,34:y:1,6</td><td>Human feces</td><td>USA</td><td>2008</td><td>9848/08</td><td>CDC</td><td>ST1712</td></tr><tr><td>Fortbend</td><td>1,3,19:z<sub>41</sub>:e,n,z<sub>15</sub></td><td>Human feces</td><td>USA</td><td>2008</td><td>9850/08</td><td>CDC</td><td>ST1713</td></tr><tr><td>Mali</td><td>51:r:e,n,x</td><td rowspan="6">H2S<sup>-</sup></td><td>Human feces</td><td>Germany (West Africa)</td><td>2009</td><td>9879/09</td><td>RKI</td><td>ST1714</td></tr><tr><td>Bergerac</td><td>44:c:e,n,x</td><td>Human feces</td><td>France (Guinea)</td><td>2009</td><td>9970/09</td><td>IP</td><td>ST1715</td></tr><tr><td>Salpetriere</td><td>21:k:1,6</td><td>Human feces</td><td>France</td><td>2009</td><td>9992/09</td><td>IP</td><td>ST1716</td></tr><tr><td>Wembleypark</td><td>17:g,m:-</td><td>Human feces</td><td>UK</td><td>1994</td><td>9997/09</td><td>PHE</td><td>ST1761</td></tr><tr><td>Noho</td><td>13,23:k:1,6</td><td>Human feces</td><td>UK</td><td>1996</td><td>9999/09</td><td>PHE</td><td>ST1762</td></tr><tr><td>Ashford</td><td>6,7:k:z<sub>35</sub></td><td>ONPG<sup>+</sup></td><td>Spice</td><td>UK</td><td>1996</td><td>10000/09</td><td>PHE</td><td>ST1760</td></tr><tr><td>Elmdon</td><td>51:i:1,2</td><td></td><td>Human feces</td><td>UK</td><td>1996</td><td>10002/09</td><td>PHE</td><td>ST1763</td></tr><tr><td>Torbay</td><td>3,10:l,z<sub>28</sub>:1,5</td><td></td><td>Human feces</td><td>UK (Gambia)</td><td>1997</td><td>10004/09</td><td>PHE</td><td>ST1764</td></tr><tr><td>Colchester</td><td>8,20:r:1,6</td><td></td><td>Sausage</td><td>UK</td><td>1997</td><td>10005/09</td><td>PHE</td><td>ST1765</td></tr><tr><td>Sandbanks</td><td>6,8:k:1,7</td><td></td><td>Human feces</td><td>UK</td><td>1997</td><td>10006/09</td><td>PHE</td><td>ST1766</td></tr><tr><td>Cork</td><td>9,46:y:1,2</td><td></td><td>Sausage</td><td>Republic of Ireland</td><td>1998</td><td>10008/09</td><td>PHE</td><td>ST1767</td></tr><tr><td>Frimley</td><td>39:r:1,5</td><td></td><td>Human feces</td><td>UK (Nigeria)</td><td>1999</td><td>10010/09</td><td>PHE</td><td>ST1769</td></tr><tr><td>Harrow</td><td>4,12:z<sub>10</sub>:1,7</td><td></td><td>Human feces</td><td>UK</td><td>1999</td><td>10011/09</td><td>PHE</td><td>ST1811</td></tr><tr><td>Goodmayes</td><td>44:z:e,n,z<sub>15</sub></td><td></td><td>Human feces</td><td>UK</td><td>1999</td><td>10012/09</td><td>PHE</td><td>ST1770</td></tr><tr><td>Poole</td><td>9,46:a:1,6</td><td></td><td>Human feces</td><td>UK (East Africa)</td><td>2001</td><td>10017/09</td><td>PHE</td><td>ST1773</td></tr><tr><td>Kingslynn</td><td>3,10:z:e,n,z<sub>15</sub></td><td></td><td>Human feces</td><td>UK (Gambia)</td><td>2004</td><td>10018/09</td><td>PHE</td><td>ST877</td></tr><tr><td>Maybush</td><td>11:z:e,n,z<sub>15</sub></td><td></td><td>Human feces</td><td>UK</td><td>2004</td><td>10019/09</td><td>PHE</td><td>ST878</td></tr><tr><td>Carshalton</td><td>47:g,m:s:-</td><td></td><td>Human feces</td><td>UK (Nigeria)</td><td>2004</td><td>10020/09</td><td>PHE</td><td>ST1774</td></tr><tr><td>Stafford</td><td>6,14,24:z:l,z<sub>13</sub>,z<sub>28</sub></td><td></td><td>Human feces</td><td>UK</td><td>2004</td><td>10023/09</td><td>PHE</td><td>ST557</td></tr><tr><td>Coppettswood</td><td>30:b:1,z<sub>13</sub>,z<sub>28</sub></td><td></td><td>Human feces</td><td>UK (Ghana)</td><td>2004</td><td>10024/09</td><td>PHE</td><td>ST1775</td></tr><tr><td>Welwyn</td><td>11:l,z<sub>13</sub>,z<sub>28</sub>:e,n,z<sub>15</sub></td><td></td><td>Dried ugu leaves</td><td>UK</td><td>2004</td><td>10026/09</td><td>PHE</td><td>ST887</td></tr><tr><td>Zollikerberg</td><td>6,7:z<sub>4</sub>,z<sub>24</sub>:1,5</td><td></td><td>Human feces</td><td>Switzerland</td><td>2007</td><td>10031/09</td><td>IFSH</td><td>ST1717</td></tr></tbody></table>
<table><tbody><tr><td>Yverdon</td><td>1,4,12:z<sub>38</sub>:1,5</td><td>Human feces</td><td>Switzerland (Burkina Faso)</td><td>2009</td><td>10033/09</td><td>IFSH</td><td>ST1718</td></tr><tr><td>Nuorikkala</td><td>8:z<sub>4</sub>,z<sub>24</sub>:e,n,x</td><td>Feed factory environment</td><td>Finland</td><td>2010</td><td>10084/10</td><td>NPHI</td><td>ST1719</td></tr><tr><td>Gostenhof</td><td>1,42:d:z<sub>6</sub></td><td>Human feces</td><td>Germany</td><td>2009</td><td>10085/10</td><td>IHU</td><td>ST1720</td></tr><tr><td>Owerri</td><td>6,14,25:l,z<sub>13</sub>,z<sub>28</sub>:1,6</td><td>Human feces</td><td>Norway (Nigeria)</td><td>2010</td><td>10105/10</td><td>NIPH</td><td>ST1721</td></tr><tr><td>Braeswood</td><td>47:c:z<sub>6</sub></td><td>Human feces</td><td>USA</td><td>2009</td><td>10113/10</td><td>CDC</td><td>ST1722</td></tr><tr><td>Saintes</td><td>13,23:d:e,n,x</td><td>Human feces</td><td>France (Guinea)</td><td>2010</td><td>10142/10</td><td>IP</td><td>ST1723</td></tr><tr><td>Perrosguirec</td><td>4,12:z<sub>4</sub>,z<sub>23</sub>:e,n,z<sub>15</sub></td><td>Human feces</td><td>France (Cote d'Ivoire)</td><td>2010</td><td>10146/10</td><td>IP</td><td>ST1724</td></tr><tr><td colspan="9"><strong>subsp. salamae</strong></td></tr><tr><td></td><td>3,15:z<sub>87</sub>:e,n,x,z<sub>15</sub></td><td>Sal<sup>+</sup>, ONPG<sup>+</sup></td><td>Reptile</td><td>Germany</td><td>2007</td><td>9822/08</td><td>BfR</td><td>ST1349</td></tr><tr><td></td><td>11:z<sub>41</sub>:e,n,x</td><td>ONPG<sup>+</sup></td><td>Chamaeleon</td><td>Germany</td><td>2008</td><td>9827/08</td><td>BfR</td><td>ST1365</td></tr><tr><td></td><td>3,10:a:z<sub>87</sub></td><td></td><td>Human feces</td><td>USA</td><td>2008</td><td>9849/08</td><td>CDC</td><td>ST1348</td></tr><tr><td></td><td>9,12:g,m,t:z<sub>39</sub></td><td></td><td>Tea leaves</td><td>Germany</td><td>2008</td><td>9880/09</td><td>IHU</td><td>ST1725</td></tr><tr><td></td><td>13,23:g,s,t:e,n,x</td><td></td><td>Human</td><td>Germany</td><td>2009</td><td>9944/09</td><td>IHU</td><td>ST1726</td></tr><tr><td></td><td>17:d:z<sub>6</sub></td><td>Dul<sup>-</sup></td><td>Human</td><td>Germany</td><td>2009</td><td>9991/09</td><td>RKI</td><td>ST1727</td></tr><tr><td></td><td>6,7:z:e,n,x,z<sub>15</sub></td><td></td><td>Spice</td><td>France</td><td>2009</td><td>9994/09</td><td>IP</td><td>ST1728</td></tr><tr><td></td><td>35:g,z<sub>62</sub>:e,n,x</td><td></td><td>Human feces</td><td>UK</td><td>1994</td><td>9996/09</td><td>PHE</td><td>ST1812</td></tr><tr><td></td><td>6,7:b:1,5</td><td></td><td>Human feces</td><td>UK</td><td>2000</td><td>10014/09</td><td>PHE</td><td>ST1772</td></tr><tr><td></td><td>13,22:g,m,t:z<sub>42</sub></td><td></td><td>Human feces</td><td>UK</td><td>2001</td><td>10016/09</td><td>PHE</td><td>ST1813</td></tr><tr><td></td><td>1,9,12,46,27:b:z<sub>39</sub></td><td></td><td>Lizard</td><td>Germany</td><td>2010</td><td>10090/10</td><td>BfR</td><td>ST1729</td></tr><tr><td></td><td>43:m,t:-</td><td>ONPG<sup>+</sup></td><td>Chamaeleon</td><td>Germany</td><td>2010</td><td>10134/10</td><td>BfR</td><td>ST1730</td></tr><tr><td colspan="9"><strong>subsp. arizonae</strong></td></tr><tr><td></td><td>51:z<sub>36</sub>:-</td><td></td><td>Human sputum</td><td>USA</td><td>2008</td><td>9813/08</td><td>CDC</td><td>ST1414</td></tr><tr><td></td><td>17:g,z<sub>51</sub>:-</td><td></td><td>Bobcat lung</td><td>USA</td><td>2007</td><td>9846/08</td><td>CDC</td><td>ST1375</td></tr><tr><td colspan="9"><strong>subsp. diarizonae</strong></td></tr><tr><td></td><td>48:k:z<sub>57</sub></td><td></td><td>Human</td><td>Poland</td><td>2007</td><td>9785/08</td><td>MUG</td><td>ST1731</td></tr><tr><td></td><td>11:i:z</td><td></td><td>River</td><td>USA</td><td>2007</td><td>9809/08</td><td>CDC</td><td>ST1265</td></tr><tr><td colspan="9"><strong>subsp. houtenae</strong></td></tr><tr><td></td><td>30:g,m,t,z<sub>51</sub>:-<sup>d</sup></td><td>Sal<sup>-</sup></td><td>Sea water</td><td>Bolivia</td><td>1978</td><td>2193/78</td><td>INLASA</td><td>ST1200</td></tr><tr><td></td><td>48:z<sub>4</sub>,z<sub>24</sub>:-</td><td></td><td>Human feces</td><td>USA</td><td>2007</td><td>9810/08</td><td>CDC</td><td>ST1732</td></tr><tr><td></td><td>21:z<sub>38</sub>:-</td><td></td><td>Toucan bird</td><td>Brazil</td><td>2008</td><td>9828/08</td><td>BfR</td><td>ST1733</td></tr></tbody></table>

¹ *Serovar* previously described as belonging to *S. bongori* in Supplement 47 of the White-Kauffman-Le Minor scheme [8] but reassigned to *S. enterica* subsp. *enterica*

² *enterica*

³ *b*The differential phenotypical characters of *Salmonella* species and subspecies are indicated in reference 7. Only the atypical results are mentioned. Tests

4 used : β-galactosidase, ONPG ; galacturonate, Gal ; malonate, Mal ; salicin, Sal ; dulcitol, Dul ; L(+) tartrate (= d-tartrate), Dtar ;
<table><tr><td>9780/08</td><td>47:c:e,n,x,z15:Rz84</td><td>Snake</td><td>Canada</td><td>2007</td><td>PHAC</td><td>47:c:e,n,x,z15: [Rz57], [Rz84]</td></tr><tr><td>9874/09</td><td>48: l,v:1,5,7:Rz90</td><td>Reptile</td><td>Germany</td><td>2009</td><td>BfR</td><td>48:l,v:1,5,(7): [Rz47], [Rz50], [Rz70], [Rz89], [Rz90]</td></tr><tr><td>9965/09</td><td>47:r:z:Rz50</td><td>Human feces</td><td>Korea</td><td>2009</td><td>KCDC</td><td>47:r:z:[Rz50]</td></tr><tr><td>9973/09</td><td>60:r:e,n,x,z15:Rz55</td><td>Snake</td><td>Germany</td><td>2009</td><td>BfR</td><td>60:r:e,n,x,z15:[Rz55]</td></tr><tr><td>9974/09</td><td>61:i:z:Rz54</td><td>Snake</td><td>Germany</td><td>2009</td><td>BfR</td><td>61:i:z:[Rz54]</td></tr><tr><td>9993/09</td><td>48:r:z:Rz50</td><td>Poultry environment</td><td>La Réunion<sup>c</sup></td><td>2009</td><td>IP</td><td>48:r:z:[Rz50]</td></tr><tr><td>10028/09</td><td>(6),14:z10:z:Rz57</td><td>Poultry</td><td>France</td><td>2009</td><td>IP</td><td>(6),14:z10:z:[Rz56], [Rz57], [Rz90]</td></tr><tr><td>10082/10</td><td>38:k:z35:z65</td><td>Human feces</td><td>Finland</td><td>2010</td><td>NPHI</td><td>38:k:z35:[z65]</td></tr><tr><td>10112/10</td><td>48:l,v:1,5,7:Rz70</td><td>Snake</td><td>USA</td><td>2009</td><td>CDC</td><td>48:l,v:1,5,(7): [Rz47], [Rz50], [Rz70], [Rz89], [Rz90]</td></tr></table>

1. <sup>a</sup>The differential phenotypical characters of *Salmonella* species and subspecies are indicated in reference 7. Only the atypical results are mentioned. Tests

2. used: Gas production from glucose, Gas; β-galactosidase, ONPG; L(+) tartrate (= d-tartrate), Dtar; +, positive reaction; -, negative reaction

3. <sup>b</sup>The National Reference Centres that sent the isolates are indicated in the acknowledgments section

4. <sup>c</sup>French island in the Indian Ocean

5. <sup>d</sup>This antigenic formula was erroneously mentioned in the 9th edition of the White-Kauffmann-Le Minor scheme

6.
<table><caption>1. Table 4. Present number of serovars in each species and subspecies of *Salmonella*.</caption><thead><tr><th></th><th></th><th>n</th></tr></thead><tbody><tr><td>4</td><td><i>S. enterica</i></td><td></td></tr><tr><td>5</td><td>subsp. <i>enterica</i></td><td>1586</td></tr><tr><td>6</td><td>subsp. <i>salamae</i></td><td>522</td></tr><tr><td>7</td><td>subsp. <i>arizonae</i></td><td>102</td></tr><tr><td>8</td><td>subsp. <i>diarizonae</i></td><td>338</td></tr><tr><td>9</td><td>subsp. <i>houtenae</i></td><td>76</td></tr><tr><td>10</td><td>subsp. <i>indica</i></td><td>13</td></tr><tr><td>11</td><td></td><td></td></tr><tr><td>12</td><td><i>S. bongori</i></td><td>22</td></tr><tr><td>13</td><td></td><td></td></tr><tr><td>14</td><td>Total</td><td>2 659</td></tr><tr><td>15</td><td></td><td></td></tr><tr><td>16</td><td></td><td></td></tr></tbody></table>
<figure><img src="image_2.png" /><figcaption>rpoB</figcaption></figure>

**Figure. Phylogenetic analysis of partial rpoB sequences of 103 strains of Enterobacteriaceae, including strains 9736/07 and CIP109002.** The analysis was done by the neighbour-joining method using BioNumerics version 6.6 software package (Applied Maths, Sint-Martens-Latem, Belgium). The partial rpoB sequences (about 1000 bp) were obtained by PCR using primers VIC4 (5'GGCGAAATGGCDGARAACCA-3') and VIC6 (5'-GGTTACAACTTCGARGAYTC-3') with conditions (annealing at 50°C) described previously [5]. The different *S. enterica* subspecies are indicated by symbols, I (subsp. *enterica*), II (subsp. *salamae*), IIIa (subsp. *arizonae*), IIIb (subsp. *diarizonae*), IV (subsp. *houtenae*), and VI (subsp. *indica*). The strains 9736/07 and CIP109002 are indicated with an asterisk.