IzvBio6_09SuntsovLO

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1 13ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ, 29, 6ｧ8 6, ｨC653 ｨｬ ｨC036.21: ｦ2036ｦｷ apple09fl apple0706: 0ｹ fl 08apple6 5apple6ｦｸ709fl fl0ｷ ｸ ｨC 06036ｦ fl, 036ｦ1 016ｦ apple6 76ｦ306fl 066ｦ fl0ｷ309070ｸ ｷ ｸ9 0ｸ803 0ｹ ｸ90ｷ309070ｸ ｸ : 06016ｦｹ apple ｸ fl0306fl ｹｸ apple6 76ｦ ｦ ｦ apple applefl , 6ｦｸ90ｸ ｦｹ06ｦ ｦ ｦ0 04apple ｦ ｸ ｷ apple ｦ0 06fl fl fl ｦ2-036ｦｷ apple6ｦ0 046ｦ ｸ apple apple0907 apple026ｦ0 096ｦ3030ｸ806 6ｦ16ｦ fl apple fl fl ｸ apple6ｦ36ｦ0 ｰ096ｦ0 0ｸ8030ｸ803 ｱ ｷ309070ｸ ｦ ｦ906 (emerging, reemerging diseases), apple0ｹ apple0ｸ ｦ ｹ06ｦ ｦ0fl 0ｹ4036ｦ0. ｨｨapple07 026ｦ0 6ｦｸ9-0ｸ ｦ fl 0ｸ8 066ｦ2036ｦｷ ｦ0, apple0ｸ8 08apple ｦ309apple ｦ0apple6ｦ ｦ apple ｦ (026ｦ3apple6ｦ ) 08apple07066ｦ (07ｨｨ07) (ｨ 6ｦ06 56ｦ0apple0909, 1974; ｨｨapple070907apple076 5, 20; Koonin et al., 21; Kunin et al., 25). ｨｨapple09fl ｹ9036ｦ3, 0ｹ ｦ fl0309 apple6ｦ90ｸ apple6ｦ60207apple036ｦｷ30909, 03apple6ｦ ｷ ｨB0ｸ96 76ｦｷ ｦ3070ｹ7 apple flapple07 apple6ｦ fl 04 08apple07016ｦ0apple ｨｦ 6ｦ ｸ8 6ｦｸ ｹｸ ｦ0 07apple6ｦ307apple0ｸ fl03, 6 5 0ｹ906030ｸ apple6ｦ apple0707apple6 76ｦ , 6ｦ ｦ30ｸ ｦ3020ｸ803 06apple 6ｦ3, 08apple ｷ303apple6ｦｷ309fl apple apple0ｷ apple apple09apple07-0ｸ ｸ ｦ3 08apple07016ｦ0apple ｸ803 07apple6 76ｦ ｨｨ070ｹ apple0706: 016ｦ ｸ apple07066ｦ ｨC ｦ306fl apple0ｸ8030ｸ ｷ , ｦｸ ｸ ｹ apple086ｦ fl0ｸ apple0409apple07ｨｪｨｦｨｺｨｪｨ ｨｮ050102ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｹ ｨｨｨｺｨ 0409ｨ ｨｨ 05ｨ 05ｨｦｨｦ03ｨｺ02ｨ ｨｦ050204ｨ ｨｹ 02ｨ 01ｨｺｨｦ0302 ｨｮｨｬ0201 Yersinia pestis: 07ｨｺ0205ｨｬ02ｨ ｨ 02 ｨ ｨ 0502ｨｲｨｪ0209ｨ ｨｦ04ｨ ｨ 02? ｷ3076 5*, ***, 04. ｨ ｷ ｦ0**, *** *ｨ apple ｹ ｹ ｸ90ｷ apple0ｷ ｦ0 ｨｺ0204, ｦ0, apple , 33 ** ｦ0apple ｸ apple ｦ0, ｨｴ01070ｷ303apple, ｦ0, apple0ｸ8 ***ｨｺ ｦ apple ｹ ｷ303apple, 6ｧ8 3, Street 3/2, 10 Distr., Hochimin, Vietnam [email protected] ｨｨ ｦ0 6 5 apple 6ｦ10ｷ3090ｸ flapple ｹ ｦ9fl fl fl, 08apple ｦ309apple040306fl 066ｦ2036ｦｷ apple fl 0ｹ4030ｸ8 Yersinia pestis apple apple07 6ｦ0 Y. pseudotuberculosis O:1b. ｨｨapple09 0ｹ apple070ｷ apple6ｦ ｦ9fl 066 5fl090ｸ86 56ｦｸ apple ｦ20ｸ apple6ｦ96 56ｦ ｦ0-0ｷ30603apple6ｦ apple030ｸ8 0ｹ ｸ ｹ apple apple, 6 5 0ｹ06ｦ ｷ ｹ ｦ90309 pfra 09 ppst. ｨｨapple 026ｦ06 76ｦ306fl 6ｦ203apple6ｦ ｦ0fl ｰ 6ｦ0apple ｦ0fl ｱ apple6ｦ0 046ｦ ｹ apple fl fl 0ｹ4030ｸ8, apple076ｦ0fl 6ｦ0 0ｹ ｹ ｨｨapple apple6ｦ ｹ ｸ90ｷ309070ｸ ｷ36ｦ0apple096 5 ｨC 066ｦ2036ｦｷ apple6ｦ0 046ｦ ｹ , 08apple 08070ｹ ｦ306fl apple6ｦ0 046ｦ ｦ2036ｦｷ ? ｨｴ ｸ apple apple090ｷ309086ｦ fl 09 apple6ｦ fl ｸ803 07apple6 76ｦ fl ｦ306fl ｦ06 50ｸ apple ｹ ｸ90ｷ ｹ909 (046ｦ96ｦ0apple076 5, 1991; ｨ 07apple 6ｦ , 21; ｹ , 22; ｨｹ , , 26). ｨｨ07 6 7apple6ｦ0 046ｦ ｹ ｸ90ｷ ｦｸ , 07 fl096ｦ30207 apple6ｦ apple07, 0207apple0309apple076 56ｦ ｸ803 08apple09096ｦ ｸ803 07apple6 76ｦ , ｦ2036ｦｷ ｨC 06016ｦｹ10707 apple6ｦ907, 016ｦ1 08apple6ｦ , ｦ ｹ ｸ90ｷ309070ｸ ｦｷ apple6ｦ ｦ apple030ｸ8. 056ｦ2036ｦｷ ｸ9 0ｹ ｸ90ｷ3090ｸ9 6 5apple 0ｸ ｸ ｦ ｹ apple ｸ90ｷ apple6ｦ0 046ｦ ｹ fl ｦ0 0ｹ ｸ90ｷ3090ｸ9 07apple- 6 76ｦ90ｹ , 0107apple apple0ｸ fl03 6ｦ ｦ3090ｹ7 08apple , 08apple09 apple ｦｸ90306fl ｦ ｦ606090ｹ ｹ ｸ90ｷ ｹ9fl ｨC 6ｦ0apple ｦ apple apple6ｦ ( ｹ apple0909 0ｹ ｸ90ｷ30909 ｨC 05ｨｪｨｴ) 08apple09fl fl0ｷ ｸ ｸ8 02fl ｸ803 07apple6 76ｦ ｦ10307apple0ｸ8 0ｹ ｸ90ｷ30909: 6ｦ ｦ0fl ｹ , fl0ｷ309fl, ｸ , 07apple 6ｦ0 096ｦ ｸ ｦ9, ｸ apple ｦ2040ｸ9 ( ｸ9) fl0ｷ3090ｸ9 060ｹ9036ｦｸ903 0ｹ ｦ0apple ｷ306 0ｹ ｸ90ｷ apple070ｷ306066ｦ0, apple apple fl fl0ｷ ｹ apple07 apple6ｦ ｦ0-09fl ｸ apple; ｹｸ8 07apple6 76ｦ909030ｸ8 fl6 502fl0ｸ90306fl 07 ｨB ｹ ｸ90ｷ30909 (026ｦ6apple, 1974). 046ｦ40ｹ ｦｸ ｦ2036ｦｷ ｦ ｸ ｸ ｹ ｸ90ｸ apple090ｸ ｦ apple apple ｦｸ90306fl ｹ ｦ ｨC flapple (0207) apple ｦ20ｸ

2 ｷ3076 5, 05040ｷ ｦ0 6 56ｦ ｦ603090ｹ ｰ apple ｱ, ｦ (ｨｨ6ｦ , 25; Konstantinidis et al., 26). ｨｦ0ｷ ｦ0fl fl apple07066ｦ0 6 7apple6ｦ0 6ｦｷ ｦ2036ｦ0-0ｷ ｹ ｸ90ｷ ｹ909, apple09096ｦ3, 0ｹ apple030ｸ8 07apple ｦ20ｸ8 6ｦ0apple ｸ ｦ2036ｦｷ ｸ8 08apple09 apple ｦｸ90306fl 096ｦ303apple6ｦ207 08apple ｸ fl ｦ0 08appleapple0ｸ appleapple0ｸ ｸ806 08apple070ｷ ｹ ｸ90ｷ30909 (Kurland et al., 23). ｨｨapple09 0ｹ ｸ90306fl ｸ apple ｦ20ｸ apple ｸ803 apple040906, ｰ08apple ｹｸ803 ｱ , ｦ apple ｦ ｦｸ90ｸ apple090309apple09-030ｸ8 08apple ｦ fl. ｨｨapple ｸ9, 6ｦ06 702fl 040ｸ ｸ ｦ90ｸ ｷ3096ｦ ｦ309 apple6ｦ90ｸ803 08apple , apple6ｦ0fl ｦ3 apple04090ｸ ｸ8 6 7apple6ｦ0 046ｦ ｨC066ｦ2036ｦｷ , 08apple 06036ｦ06 502fl03 060ｷ36ｦ0apple apple fl 09 0ｹ ｸ90ｷ apple07 6ｦ0 0ｹ4030ｸ8 Y. pestis, ｦｸ806 6ｦ309 6ｦ ｹ ｹ ｦ ｹ fl ｸ ｦ ｦ fl 0307apple03090ｹ ｦ0-03apple apple ｨｪ6ｦ apple - 6ｦ96ｦｹ90ｸ9 0ｹ (06090ｹ ) fl 0307apple03090ｹ6016ｦ0fl 0807 apple08016ｦ ｰ0307apple- 030ｸ8 07fi 02fl ｱ, apple6ｦ0fl apple08 6ｦ3, 0ｹ apple 026ｦ3 016ｦ ｦ ｦ3090ｹ ｸ ｦｸ ｦ0 6ｦｹ9, apple6ｦ9apple ｸ apple6ｦ3016ｦ3 apple , 0ｹ06ｦｸ ｸ ｸ apple0903apple04, 016ｦ apple ｦ0 ｰ flapple0ｸ803 0ｹ06ｦ ｱ 6 50ｸ apple6 76ｦ ｸ803 08apple ｦ0 6 5 apple6ｦ3016ｦ3 066ｦ flapple , 036ｦ1 016ｦ1 096ｦ ｦ ｹ , 086ｦ ｦ3090ｹ apple ｹ ｦｸ apple ｦ406 apple6ｦ ｹ apple03090ｹ ｦ9 6ｦ ｦ3 036ｦ1: 08apple070ｷ306060ｸ flapple0706 0ｹ ｸ90ｷ fl 0ｹ4030ｸ ｸ8 0ｸ ｦ ｦｸ8 06 0ｹ ｸ90ｷ ｸ apple070ｷ306066ｦ309 6ｦ ｸ apple076 5fl03 07apple6 76ｦ ｦｷ ( ｹ703, 07apple6 76ｦ , fl0ｷ , 09070ｷ ｹ ) ｷ303apple6ｦ20ｸ apple07066ｦ3 0ｹ ｸ90ｷ ｦ306fl fl0ｷ apple076 5: fl0ｷ309fl fl6 502fl0306fl 0ｹ ｦ0apple ｷ306 0ｹ ｸ90ｷ apple070ｷ306066ｦ apple ｸ ｦ9fl ｦ0, 6ｦ ｨC 0ｹ apple036ｦ ｸ ｦ0-09fl (ｨｰ6 56ｦ0apple0ｷ3, 1980). ｨｴ ｹ ｦ9fl 0ｹ ｸ90ｷ309070ｸ803 08apple07-0ｷ apple6ｦ601apple0ｸ86 56ｦｸ ｸ7040ｸ9 016ｦ0apple ｦ fl0ｷ30906, ｷ apple6ｦ ｦ ｹ ｦ76 501, 09 0ｹ1036ｦ0 016ｦ0apple03096ｦ ｦ3 ｰ08apple ｸ9 ｱ 02fl 0107appleapple010309apple ｷ36ｦ0apple apple07 6ｦ0 ｰ08apple0701apple ｸ ｱ, ｦｸ90ｸ ｸ ｸ8, 6 50ｸ80307 fl0ｸ ｦ0 08apple 020ｸ ｹ (0ｹ ｹ ) ｨｨ07016ｦ ｦ302fl apple09096ｦ306fl 0107appleapple fl 03apple6ｦ0 090ｷ309070ｸ apple apple0401-0ｷ ｨｨapple fl apple6ｦ902090ｹ06ｦ3 ｰ036ｦ ｸ8 ｱ 09 ｰ6 70ｸ8 ｱ apple6 5fl: 08apple6 50ｸ ｸ ｦ ｹ apple apple04010ｷ309fl03 ( 096 5apple6 76ｦｷ ｦ ), apple0ｸ8 ｨC ｹ (apple6ｦ ｸ ｸ803 08apple09096ｦ1076 5). 05 apple6ｦ ｸ8 08apple apple6ｦ ｦ2036ｦｷ apple6ｦ0 046ｦ ｹ ｷ36ｦ0apple apple fl fl 0ｹ4030ｸ8. 056ｦ2036ｦｷ ｦ0fl apple076ｦ0 6ｦ0 6ｦｸ apple6ｦ fl fl (Achtman et al., 1999, 24; Pourcel et al., 24; Song et al., 24; Zhou et al., 24a, b, c; Cui et al., 28; Li et al., 28), apple03090ｹ fl6 502fl0306fl 086ｦ0apple6ｦ ｦ0 07ｨｨ07. 07apple6ｦ0 046ｦ ｹ6016ｦ0fl ｦ06ｦ0 6ｦ ｨｪｨｴ 6ｦ ｹ ｹ ｦ0-020ｸ ｦ0apple6ｦ ｹ ｦ3apple096ｦ2076 5, ｹｸ apple6ｦ2076ｦ9096ｦ (ｨｪ ) 08apple09apple apple ｹ apple0ｸ (05036ｦ ) 6ｦ3apple ｹ06ｦ06 76ｦ3 0ｹ4030ｸ ｹ4030ｸ8 ｨC ｦ30ｸ ｦ0apple6ｦ apple076 50ｸ apple0ｸ ｦ60ｷ307 apple6ｦ90ｸ ｦ ｦ96ｦ7, 0ｹ apple6 709apple076 56ｦ ｦ apple ｦ8apple fl apple ｦ0 Y. pseudotuberculosis O:1b, ｹ ｦ0apple6ｦ909036ｦ0 0409apple apple046 76ｦ0 066ｦ30ｸ803 apple6ｦ apple6ｦ90ｸ ｸ803, ｸ ｦ96ｦ0 (Achtman et al., 1999, 24; Skurnik et al., 20). ｨｦ ｹ fl 0ｹ4-030ｸ ｸ ｦ apple ｸ apple ｹ60109, 0ｹ ｹ60109, 0ｹ ｹ ｹ ｹ ｦｸ ｦ1030ｸ8 (05fl , 1989; 05040ｷ3076 5, 05040ｷ ｦ0, 26) ｦ0 6ｦ96ｦ0apple036ｦ0fl 6ｦ40ｹ06ｦ0fl 0903apple096 76ｦ0: 016ｦ ｸ80309fl 08apple ｦ0 6ｦ ｦ ｦ apple ｨC ｷ apple0907 6ｦ apple03fl, apple0ｸ8 08apple ｸ9 01apple6ｦ ｦ apple076 5fl ｦ0apple6ｦ909036ｦ ｦ apple ｹ apple07-6ｦ apple , 08070ｹ apple6ｦ608apple070603apple6ｦ0-090ｸ9; 016ｦ ｦ0 08apple09apple07 6ｦ ｸ ｦ10307apple076 5, 6 50ｸ ｦ apple6 70ｷ3090ｸ9 0ｹ apple ; ｦ apple6ｦ ｹ apple ? ｨｨ ｹ ｸ90ｷ309070ｸ apple apple07 6ｦ0 0ｹ4030ｸ ｹ06ｦ flapple0ｸ apple6ｦ3016ｦ ｷ3080ｷ ｨｨ apple0ｸ ｦ606090ｹ ｸ803 6ｦ401: , 0ｹ , 086ｦ0apple6ｦ , apple6ｦ20909, , 086ｦ , 086ｦ ｦ apple , apple03fl 016ｦ1 0ｷ ｦ apple ｸ803 08apple070ｷ ｹ ｸ90ｷ apple090ｷ ｦ apple6ｦ3016ｦ3 016ｦ apple6ｦ ｦ8apple6ｦ fl ｨｬ fl apple0ｹ906020ｸ ｦ606090ｹ ｦ ｦｹ06ｦ204 XXI ｸ ｦ ｹ ｦ , ｹ ｦ ｹ ｦ ｦｸ ｸ8 016ｦ0apple03090ｸ8 09-6ｦ apple6ｦ fl 08apple09apple07 0ｸ ｷ apple0907 6ｦ ｷ3 ｨｨ6ｦ26ｦ0apple ｨｴ ｦ ｦ90ｸ apple apple0309apple076 56ｦ0-09fl apple03apple apple030ｸ803 6ｦ0apple6ｦ apple apple apple0ｸ apple030ｸ apple0ｸ ｸ70403 ｨC ｸ fl6 5 0ｹ apple07 6ｦ0, ｹ , 066 5fl096ｦｸ ｦ ｹ06ｦ apple , 016ｦ1 ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

3 13ｨｨｨｺｨ 0409ｨ ｨｨ 05ｨ 05ｨｦｨｦ03ｨｺ02ｨ ｨｦ050204ｨ ｨｹ 02ｨ 01ｨｺｨｦ0302 ｨｮｨｬ0201 Yersinia pestis , ｨC apple ｸ803 08apple09apple07 0ｸ ｹ06ｦ ｹ4030ｸ ｦ apple090ｹ apple apple ｦ apple07066ｦ3 08apple fl 09 0ｹ ｸ90ｷ apple07 6ｦ0 0ｹ4-030ｸ apple apple0ｹ ｸ ｦ606090ｹ apple apple09apple ｹ06ｦ ｹ4030ｸ8 ｨC ｹ apple fl ｨC ｸ ｦ ｦ ｦ103apple0906 apple07 6ｦ0 Yersinia, ｦ06 502fl0ｸ90ｸ ｦ fl ｹ ｨ 030ｸ90306fl 09 apple apple090ｹ90ｸ8. 046ｦ ｦ ｦｸ ｸ8fl apple ｹ apple07 6ｦ0, - 026ｦｸ90306fl 08apple fl apple apple016ｦｷ apple ｹ0apple apple apple6 70ｷ ｸ ｹ ( ｦ0apple076 5, ) ｦ0 Y. pestis ｦ9fl apple ｰ040107apple0ｸ8 ｱ 09 ｰ07apple090309apple076 56ｦｸ8 ｱ 6 76ｦ ｹ60109 apple6 5fl apple07 6ｦ0 Yersinia, fl 0ｸ apple0ｸ8, ｸ fl, ｦ ｸ apple6ｦ2090ｹ ｦ06ｦ209096ｦ apple apple6ｦ apple6ｦ909fl fl 0ｹ4030ｸ8, 0ｹ ｦ ｦ606090ｹ6016ｦ0fl 0307apple09fl 08apple09apple ｹ06ｦ ｹ ｸ ｸ ｦ apple fl apple07 6ｦ0 Y. pestis , 0ｹ307 apple ｦ30ｸ8 6ｦ06ｦ209096ｦ0, 08apple ｸ ｸ apple apple6ｦ3 ( ｦ0apple01apple6ｦ3, 6 7-0ｸ803 02apple6ｦ ｦ3), 08apple0706fl0306fl apple ｦ ｸ ｦ ｸ ｹ06ｦ apple ｸ8. ｨｴ apple apple090ｸ ｦ809fl03, ｹ06ｦ0, ｦ ｸ90306fl ｹｸ8 0ｹ flapple0ｸ ｸ8 07apple ｹ10307, ｹ06ｦ6030ｸ ｦ0 07apple ｸ apple , 036ｦ0apple6ｦ103apple ｹ309 02fl apple6ｦ apple6ｦ20909 (02 apple6ｦ30607, 27). 0ｸ ｦ , 0ｹ307 6ｦ ｸ ｹ06ｦ30ｸ ｹ apple ｦｹ fl ｦ90ｸ86 56ｦ3 6 50ｸ8 07apple flapple ｦ0apple01apple6ｦ0; ｦ ｦ30ｸ ｸ8 07apple 08apple 06036ｦ06 502fl ｸ9 08apple apple ｹ apple07 6ｦ ｸ8 06 apple6ｦ90ｸ ｦ0apple01apple6ｦ309 6ｦｸ ｦ30307 apple6ｦ902090ｹ06ｦｸ90ｸ709-06fl apple ｦ30ｸ8 (Achtman et al., 24; Pourcel et al., 24; Gu et al., 26; Torrea et al., 26; Vergnaud et al., 27; Cui et al., 28; Li et al., 28). ｨｺ ｦ30ｸ ｦ fl 0ｹ4-030ｸ apple0309apple076 56ｦ ｹ ｸ7 0309, 0ｹ ｦ ｸ apple ｦ , ｹ06ｦ60309fl ｦｷ30906, apple ｦｷ ｷ30906, apple6ｦ96 56ｦ9fl apple apple030ｸ8 0ｹ ｸ ｹ apple apple apple6ｦ apple6ｦ903apple6ｦ0 ｨC ｸ , 6 70ｸ , 03apple6ｦ , 08026ｦ90309 ｨC apple 0ｸ8 09/ ｦ ｹｸ apple0707apple6 76ｦ , 0807apple ｦ ｹ ｦ ｹ apple07-6ｦ0. ｨｦ 6ｦ apple6ｦ ｦ06ｦ ｦ90ｸ86 56ｦ306fl 6ｦ0 080ｸ ｹ apple6ｦ96 56ｦ30ｸ ｹ apple apple 09 apple6ｦ601apple0ｸ86 56ｦ306fl ｸ806 08apple070ｷ apple09fl apple6ｦ ｦ0-040ｸ9 0ｹ403040ｸ apple0903apple04, apple6ｦ601apple0ｸ86 56ｦ306fl 08apple090ｷ30908 ｰ6 5 apple09fl ｱ ｰ06080ｷ ｹ ｱ 08026ｦ90309 ppst 09 pfra: 06080ｷ ｹ ｦ ｸ8, apple ｸ9, 0207apple0309apple040ｸ90306fl 0ｹ ｨC ｷ ｹｸ8, apple ｦ ｦ909 (acquisition) ｸ ｦ0 0ｹ408apple07 0ｸ apple03, ｷ ｹ ｦ90ｸ86 56ｦ apple04080ｸ ｦ1030ｸ fl ｦ fl 0ｹ4030ｸ8, 01apple fl0ｸ90ｸ ｦ ｹ ｦ fl ｹ apple apple 0706, 0703apple6ｦ86ｦｸ ｦ2036ｦｷ apple apple6ｦ ｦ9fl 04 08apple07016ｦ0apple (apple ) apple07 0ｸ8 0ｹ4030ｸ8, 0ｷ309apple apple040ｸ90ｸ fl0ｷ309fl apple6ｦ0 036ｦ0 (ｨ 6ｦ0 6ｦ6016ｦ2, 026ｦｹ804apple09fl, fl), apple0ｸ (046ｦ0) ｸ (056ｦ ｦ0, 016ｦ ｦ9), ｦ0 6ｦｸ apple0ｸ ｹ ｦ309, ｦｸ90ｸ apple ｸ apple ( ｦ0apple6ｦ , 1998; Zhou et al., 24a, b). ｨｨ ｸ apple09096ｦ16ｦ ｦ ｦ ｦｸ apple apple07 6ｦ fl0309 0ｹ4030ｸ fl0ｷ ｦ ｦ96ｦ0, 05apple ｦ0fl ( , 20; ｦ0 09 apple., 22; Motin et al., 22; Achtman et al., 24; Song et al., 24; Zhou et al., 24a, b, c; Vergnaud et al., 27; Cui et al., 28; Li et al., 28) ｹ flapple0ｸ803 08apple09096ｦ16ｦ3 apple ｹ ｦ0apple 016ｦ1 08apple ｹ apple apple ｸ apple ｸ apple ｹ0apple0903 0ｹ4030ｸ803, 6ｦ apple ｹ ｹ06ｦ , 6ｦ apple6ｦ2090ｹ ｦ20ｸ803 apple apple046 76ｦ ｦ ｦ0apple ｨC ｹ304 (pestoides, microti, microtus, minor): 036ｦ apple6ｦ ｦ0apple6ｦ ｦ0 Y. pestis ｰpestoides ｱ 07016ｦ96ｦ206fl 03appleapple090307apple096ｦ207 apple6ｦ907apple6 56ｦｸ803 6ｦ ｰ070603apple ｱ, 6ｦ ｦ20ｸ803 apple apple046 76ｦ0. 05apple apple6ｦ90707 apple6ｦ90ｸ8 060ｷ36ｦ0apple apple6 70ｷ apple ｹ apple , ｦｸ8 6ｦ ｦ0, apple090ｷ ｸ8 08apple ｦ0 apple ｦ0apple6ｦ103apple0ｸ fl ｷ36ｦ0apple09fl 066ｦ2-036ｦｷ apple fl 09 0ｹ ｸ90ｷ apple07 6ｦ0 0ｹ4030ｸ ｦ96 56ｦ ｸ90ｸ ｹ06ｦ apple6ｦ9040ｸ90ｸ ｦｷ30909 (036ｦ1apple ｦｷ30909, ｦｷ30909, 07ｨｨ07) apple apple6ｦ2090ｹ ｦ0, ｦ06 56ｦ apple apple0ｸ8 06apple 0ｸ8 ( ｷ307096ｦ0). 2. ｨｦ03apple090ｷ36ｦ fl ｦ0- apple6ｦ103apple6ｦ0 0207apple0309apple076 56ｦ9fl 04 0ｹ apple07 6ｦ0 0ｹ ｸ9-0ｷ309070ｸ ｨｦ0307apple6 56ｦ apple ｸ8 0ｹ ｸ90ｷ fl ｹ ｸ90ｷ fl096ｦ apple07060ｹ9016ｦ0. ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

4 ｷ3076 5, 05040ｷ ｦ0 Yersinia pestis pestoides Yersinia pestis ant med ori Yersinia pseudotuberculosis O:1b II ｹ ｸ80309 ｨC apple6ｦ96 56ｦ ｦ ｸ8 ppst ｷ3096ｦ0 I ｹ ｸ80309 ｨC apple6ｦ96 56ｦ ｦ ｸ8 pfra Fra1 ｨｺ ｦ0 066ｦ2036ｦｷ apple fl 09 0ｹ ｸ90ｷ fl 0ｹ4030ｸ ｰ apple6ｦ96 56ｦ9fl ｱ 06080ｷ ｹ ｦ90309 pfra 09 ppst ｦ0apple0ｸ8: ant ｨC antiqua; med ｨC medievalis; ori ｨC orientalis. 4. ｨｦ03016ｦ apple0ｹ apple apple6ｦ9040ｸ90ｸ706 apple apple6ｦ0 09 6ｦ0 6ｦ ｦ0apple6ｦ103apple6ｦ apple6ｦ ｦ apple070ｷ306066ｦ0; 0ｹ ｸ90ｷ309070ｸ806 08apple070ｷ30606 apple6ｦ606036ｦ3apple096 56ｦ306fl 016ｦ1 6ｦ0 6ｦ8036ｦｷ , apple fl ｸ706 apple apple ｹ4030ｸ8: ｸ ｦ ｦ0 6ｦ ｦ apple09fl030ｸ fl 09 06apple6ｦ904 6ｦ ｸ9 0ｹ ｸ9-0ｷ ｸ ｨｦ03016ｦ fl0ｷ ｦ0: ｦ0fl 09090ｷ36ｦ0 0ｹ ｸ90ｷ apple070ｷ306066ｦ0 ｨC 09090ｹ06ｦ0fl apple07 6ｦ0fl ｦ ｷ36ｦ0apple09fl ｹ apple 06036ｦ fl appleapple0ｸ applefl- 6ｦ0 apple03apple ｸ803 08apple ｹｸ apple03; 6ｦ06ｦ ｦｷ ｦ ｨC 01070ｷ308-0ｷ309fl 08appleapple0ｸ ｦ ｹ , apple08 6ｦｸ90ｸ706 apple6ｦ apple ｹ0-0ｸ apple03, 6ｦ fl09 0ｹ fl fl ｦ ｹ ｨｰ09apple apple09fl036ｦ0fl ｦ606090ｹ ｹ ｸ90ｷ ｸ apple09fl ｷ3080ｷ309fl 08apple6ｦ0 6ｦ8036ｦｷ ｹ ｷ36ｦ0apple0909 6ｦ apple fl. ｨ 036ｦ1, apple6ｦ ｦ fl 0ｹ4030ｸ8, 016ｦ fl flapple0ｸ , 08apple apple6 76ｦ ｦ ｦ apple07 6ｦ0 Microtus ( apple07 6ｦ0) ｹｸ apple07 0ｸ fl apple ｦ0 6ｦ ｹ06ｦ60ｸ803 01apple04080ｸ803 08apple07 apple6ｦ ｦ ｦ apple6ｦ ｸ8apple6ｦ8090ｸ ｦ309 6ｦ0apple ｦ0 ｨｺ ｦ0 ｨC ｦｹ ｸ apple076 5 (Goldscmidt, 1940) ｷ3080ｷ309fl 066ｦ2036ｦｷ apple6ｦ96 56ｦ9fl 01apple0408-0ｸ ｸ , 03apple6ｦ ｦ ｦ ｨB ｸ ｸ apple6 76ｦ909036ｦ0, ｹ apple07016ｦ0apple , fl6 502fl0306fl apple6ｦｷ309fl apple apple0ｸ803 0ｹ (08apple09096ｦ ) (Kurland, 20) ｦ ｦ606090ｹ apple ｦ fl03, 036ｦ1apple ｦｷ ｦ0 6ｦ ｸ8, 036ｦ1apple ｦ30ｸ ｸ apple ｦ0 6ｦ0apple040409fl 0307apple ｹ appleapple- 02fl0ｷ apple 096ｦｷ apple6 76ｦ76 5 (07apple6 76ｦ , 07apple6 76ｦ0-0202) 09 0ｷ ｸ803 07apple6 76ｦ ｨ ｦ ｦ , 0ｹ ｦ2036ｦｷ apple07 6ｦ0 0ｹ4030ｸ ｦ109 fl6 502fl0306fl apple apple0ｸ803 06apple flapple0ｸ ｨｪ6ｦ1, ｨｺ (Wren, 23) ｰ016 56ｦ ｸ ｦｹ ｦ0 (quantum leap in eye blink) ｱ apple apple07 6ｦ0 O:1b fl 0ｹ4030ｸ8 apple6ｦ606036ｦ3apple096 56ｦ ｹ ｦ604036ｦ0 6 5apple , ｦ ｹ06ｦ0, 6 50ｸ ｦ0 apple6ｦ fl0309fl apple6ｦ0 046ｦ ｦ fl ｦ ｦ ｦ06ｦ ｦ3fl ｸ ｦ flapple0ｸ , apple0706 6ｦ apple0ｸ8 08apple 06036ｦ apple0309apple076 56ｦ9fl ｹ apple apple, apple040ｸ90ｸ apple apple07 6ｦ0 0ｹ4030ｸ fl ｦ fl0ｷ apple apple07 6ｦ0 (Zhou, Yang, 29) ｦ3090ｹ7, 0ｹ applefl 6 5 0ｷ ｦ0 0ｹ ｹ apple apple fl 0ｹ4030ｸ8, 6 50ｸ8apple6ｦ fl ｦ ｹ ｸ ｦ0apple6ｦ ｸ8 07ｨｨ ｦ0, flapple0ｸ , 6ｦ1070ｷ3, 6ｦｹ96ｦｸ ｸ7040ｸ76ｦ fl fl apple03096ｦ3 05ｨｪｨｴ (08apple6ｦ0 6ｦ8036ｦｷ309fl, 6 5apple ｦ207 6ｦ ｦ9 08apple09096ｦ1076 5, apple , fl0ｷ309fl apple ｸ apple03, ｹ ｹ ｦ fl0ｷ30909, , 06- ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

5 13ｨｨｨｺｨ 0409ｨ ｨｨ 05ｨ 05ｨｦｨｦ03ｨｺ02ｨ ｨｦ050204ｨ ｨｹ 02ｨ 01ｨｺｨｦ0302 ｨｮｨｬ0201 Yersinia pestis apple ｸ , ｸ apple) apple ｦ0apple6ｦ ｨｨ07, 0207apple0309apple076 56ｦ9 0ｹ apple07 6ｦ apple fl03 05ｨｪｨｴ, apple , , 6 7apple6ｦ0 046ｦ207, fl096ｦ30207 apple6ｦ apple07. ｨｮ0307 0ｸ8 08apple09fl03 ｰ 6ｦ0apple ｱ apple6ｦ ｦ9fl fl 0ｹ4030ｸ8, ｦ96ｦ3 6ｦ2090ｹ9 08apple apple 0ｸ ｦ96ｦ ｹ ｦ ｸ ｦ9 08apple ｹｸ apple03 016ｦ1 apple6ｦ20ｸ apple03apple ｸ803 08apple09apple07 0ｸ803 fl ｦ606090ｹ6016ｦ0fl fl , 6ｦ0-0909apple040ｸ90ｸ76ｦ0fl06fl 6ｦ ｰ03apple ｦ0apple6ｦ ｦ0 ｱ, apple ｦ ｦ909 6ｦｸ803 06apple6ｦ ｦ06ｦ apple , 0ｹ103 apple ｦ , apple6ｦ apple6ｦ90ｹ0-0ｸ apple apple04010ｷ ｹ ｸ90ｷ309070ｸ803 08apple070ｷ apple07016ｦ0apple ｨｨ070ｹ , 036ｦ1 016ｦ1 6ｦ0 6ｦ8036ｦ0-0ｷ309fl 0ｹ apple07 6ｦ0 6ｦ8apple6ｦ ｦ apple ｦ9fl ｨC 086ｦ0apple6ｦ909036ｦ0apple apple0ｸ80904ｨC ｦ0, 02fl apple0709fl 060ｷ36ｦ0apple09fl apple fl 09 0ｹ ｸ9-0ｷ ｸ apple ｹ ｦ9 6ｦ0 0ｹ ｹ ｦ ｦ3 08apple ｸ8 ｨC ｦ fl fl apple030ｸ apple0ｸ809046ｦ0 ｨC apple016ｦ0-036ｦ0apple 6ｦ06 76ｦ06ｦ0 Marmota sibirica ｷ ｹ apple6ｦ apple Oropsylla silantiewi. 050ｸ8-07apple 0ｹ ｨB ｸ ｦ96ｦ ｦ906, 6ｦ ｸ apple6ｦ902090ｹｸ803 6ｦ ｦ3 0307apple0909 ｨｨ6ｦ (1946) 07 08apple09apple ｹ06ｦ apple6ｦ ｸ ｹ ｦ ｸ ｦ ｹ ｹ ｦ0 020ｸ9 09fl apple6ｦ ｨｦ ｹ apple016ｦ0-036ｦ0apple 6ｦ06 76ｦ06ｦ O. silantiewi ｦ apple 06036ｦ fl ｦ3, 08apple apple0ｸ803 0ｹ apple03030ｸ apple ｦｸ90ｸ ｦ96ｦ ｹ ｦ ｹｸ apple apple ｹ ｸ90ｷ309070ｸ806 08apple apple090ｷ ｦ ｸ9 06apple ｦ9fl, ｸ9 6ｦ0 6ｦ ｸ ｨｨ07 apple ｹ ｷ36ｦ0apple ｦ0 6ｦ309 apple6ｦ0 (05040ｷ apple., 1997; 05040ｷ3076 5, 05040ｷ ｦ0, 20, 26, 28). ｨｦ 0703apple6ｦ86ｦ3 0ｹ ｹ apple apple04010ｷ apple07 6ｦ0 Yersinia ｦ0 Y. pestis 6ｦ ｦ06ｦ209096ｦ0 6ｦ0 6ｦ ｸ fl6 5, apple07060ｹ ｦ fl 6ｦ ｦ20ｸ ｦ3090ｹ apple09apple07 0ｸ8. ｨｺ ｦ30ｸ8 6ｦ ｸ ｦ0apple6ｦ ｹ ｹ ｹ ｦ apple090ｷ ｦ ｷ3080ｷ apple fl 0ｹ4030ｸ8: apple07 Y. pestis 096 5apple6 709apple076 56ｦ ｹ ｦ0apple6ｦ909036ｦ0 ｨC apple apple07 6ｦ apple ｦ0 (O:1b) ｷ ｷ3-6ｦ0 ( ｦ ｦ ｦ0apple036ｦ apple03fl, 22ｨC15 030ｸ ｦ96ｦ0 ) apple6ｦ (apple ) ｦ0apple6ｦ909036ｦ0apple ｰ apple0701 ｨC ｦ0 O. silantiewi ｱ, apple040ｸ apple 06036ｦ ｦ apple6ｦ apple6ｦ304apple ｹ60107 ｰ0ｹ ｸ90ｷ ｱ ｦ86ｦ apple6ｦ304apple 08apple0903apple- 07 5ｨC37 紊; apple apple6ｦ ｦ9fl apple0703apple096ｦ9 6 7apple04036ｦ0 6ｦ ｦ26 76ｦ9fl ｹｸ ｦ3apple apple016ｦ0, , 6 50ｸ ｦ ｸ80ｹ7 02fl 02090ｹ apple070ｹ ｨC 08apple ｸ ｦ fl0ｸ apple016ｦ ｦ ｦ ｸ ｦ307026ｦ ｸ ｦ ｦ0applefl 0ｹ ｹ ｦｷ apple fl0ｸ apple016ｦ0 0ｹ0apple ｦ0apple ｦｷ ｦ ｰ070301apple0ｸ apple07036ｦ0 ｱ 02fl ｹ apple6ｦ ｦ3090ｹ (03036ｦ90ｹ ) 6 5 apple09fl apple apple07 6ｦ apple076 5fl07 apple ｦ2-0ｸ803 0ｹ06ｦ603090ｷ3, ｦ ｦ0 6ｦｸ90ｸ apple ｸ apple03fl apple ｦ ｹ706 08apple ｦ403 0ｹ ｹ ｷ36ｦ0apple apple fl 09 0ｹ ｸ90ｷ fl 0ｹ4030ｸ8 6 50ｸ8 020ｸ8 6 56ｦ0 0ｹ1036ｦ86ｦ0: 036ｦ1apple apple070ｹ ｸ90ｷ309070ｸ ｦ1apple070ｹ ｸ90ｷ309fl, apple070ｹ ｸ90ｷ30906, 08apple 06036ｦ3 016ｦ fl0ｷ ｹ , 6ｦ apple090ｹ ( ｹ ) 08apple070ｷ30606 (apple ). ｨｴ apple070ｷ apple ｦ2 6ｦ0 6 7apple6ｦ apple0309 6ｦ0apple0708 6ｦｸ90ｸ70606fl fl0ｷ ｦ0 08apple09 0ｹ apple036ｦ20ｸ fl apple04086ｦｸ90ｸ706 06apple 0ｸ8, 0ｹ ｹ06ｦ apple070ｹ ｸ90ｷ30909 ｨC ｸ90ｸ706 03fl ｰ08apple090309apple ｱ apple030ｸ ｸ803 ( ｦ20ｸ803) fl ｸ ｦ9fl. 076ｦ0apple6ｦ103apple0ｸ apple6ｦ0 046ｦ ｹ ｷ36ｦ0apple09fl fl ｦ9fl 060ｷ36ｦ0apple09fl ｦｸ8 6ｦｸ ｸ803 6ｦ401: 0ｹ , apple6ｦ , 086ｦ , 086ｦ ｦ , apple , 086ｦ0apple6ｦ , 0ｹ , flapple apple , 0ｹ ｦ apple6ｦ ｸ ｰ0ｹ ｸ803 ｱ. 2. ｨｨ apple016ｦ0-036ｦ0apple 6ｦ06 76ｦ06ｦ0, ｦｸ90ｸ ｦ0 6ｦ0apple09 0ｸ803 08apple070603apple6ｦ ｦ3 0903apple6ｦ , 066 5fl096ｦ fl0ｹ1, fl ｸ apple ｦ ｹ apple070ｹ ｨC apple ｦ8apple07026ｦ ( 03apple ｦ ) ｦ0apple6ｦ ｨC apple6ｦ606036ｦ3apple096 56ｦｸ90306fl 016ｦ ｸ8 0ｹ ｹ60109 (0ｹ ｹ60109) 026ｦ10307apple0ｸ8 06apple 0ｸ8 ( 09070ｷ ｦ0), ｦ apple ｸ fl 0207apple0309apple076 56ｦ9fl 0ｹ ｹ ｸ8 0ｹ4030ｸ8. 3. ｨｨapple apple090ｹ fl fl 0ｹ4030ｸ ｦ ｦ36ｦ apple , apple apple6ｦ ｷ ｷ3; ｦ fl 01apple6ｦ6 0604apple apple6ｦ ｦ ｦ36ｦ0 0ｹ ｸ8 0903apple6ｦ ｸ ｸ86 5 0ｸ apple6ｦ fl; 08apple ｸ80603apple0ｸ8 ( apple0ｸ ｷ3016ｦ3 ｨC 016ｦ36ｦ603apple ｹ60109) fl ｦ3090ｹ , 08apple ｸ ｸ ｦ ｹ apple070ｹ ｨC 08apple ｦ ｦ ｦ0apple6ｦ apple076 56ｦ90ｸ9, ｸ fl0ｷ ｦ0 08apple 020ｸ8 07apple030ｸ8 apple6ｦ10ｷ30909; 6 5 apple ｦ3 036ｦ ｰ03036ｦ ｹ ｱ ｦ apple070ｹ ｦ0 6ｦ ｷ303apple6ｦ2076ｦ9096ｦ apple ｷ307096ｦ fl ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

6 05040ｷ3076 5, 05040ｷ ｦ ｦ3 6ｦ ｦ9 22ｨC ｸ ｦ96ｦ ｦ ｦ ｦ0 6ｦ ｦ36ｦ0 07ｨC ｦｲ ｦ0 09 6ｦｲ ｦ ｨｺ ｨｺ6ｦ ｲ ｦ0 0ｹ4030ｸ ｦ ｨC ｦｲ ｦ0 6 50ｹ ｸ8; 2 ｨC 6ｦ9 0ｸ8 6ｦ6070ｸ ｹ ｦ90ｲ4; 3 ｨC 6ｦ ｲ ｦ0 0ｹ4030ｸ8 6ｦ ｸ ｦ36ｦ ｦ ｸ ｸ ｸ ; 4 ｨC 6ｦ ｦ9 036ｦ ｦ ｹ ｹ (066ｦ36ｦ ) ｦ90ｲ ｦ90ｷ ( ); 5 ｨC 6ｦ06ｦ ｹｸ ｸ ｹ06ｦ ｹ4030ｸ ｦ9 0ｸ ｦ9 0ｸ803 6ｦ6076ｦ3 6 56ｦ ｦ ｦ96ｦｸ8 6ｦ86ｦ ｲ ｹ ｦ ｦ ｲ40ｷ ｦ0-036ｦ0 6ｦ06 76ｦ06ｦ ｷ ｷ36ｦ ｷ3. 020ｲ ｦ ｹ ｦ06ｦ909036ｦ ｸ ｦ90ｸ ｦ ｲ4096ｦ ｦ ｦ0: ｦ ( ｦ06ｦ86ｦ ｸ ｸ ｦｷ ｸ ｸ ｦ3046ｦ 紊), 6ｦ2090ｹ ｦ3 6ｦ309026ｦ ｷ309036ｦｸ ｦ , ｦ ｦ3060ｲ ｦ107026ｦ06 76ｦ3, ｦ90ｸ ｲ4 6ｦ (Zhou, Yang, 29) ｨC ｦ0 6ｦ8036ｦｷ ｦ90ｸ9 6 5 ｰ ｱ ｲ40ｸ ｦ36ｦ0 6ｦ06 76ｦ06ｦ ｦ , , ｦ ｦ90ｸ ｲ4 0ｹ4030ｸ8. ｨｨ ｸ , 056ｦ ｦ ｦ , ｹ ｹ903, ｹ ｦ ｸ8 (Eppinger et al., 27). ｨｨ , 0ｹ ｸ ｦ ｷ ｹ ｦ ｹ ｦ0 ppst 09 pfra ｦ1070ｹ ｸ90ｷ309070ｸ ｦ0 046ｦ ｹ ｦ06ｦ ｦ ｦ90ｲ ｦ ｲ4 0ｹ4030ｸ8: 0ｹ ｸ90ｷ ｸ90ｸ ｸ ｦ06ｦ909036ｦ ｸ8 ｰ6 70ｸ80904 (08090ｸ76 56ｦ903020ｸ ｦ103)ｨC6 5040ｲ40ｲ4 06 6ｦ0 (08070ｹ06 56ｦ0)ｨC6 70ｸ80904 ｱ ｸ9, 096ｦ ｸ ｰ6 70ｸ80904 ( ｸ )ｨC ｦ0 (08090ｸ76 56ｦ903020ｸ ｦ103)ｨC6 70ｸ80904 ｱ, ｹ ｦ ｹ ｷ306060ｸ ｦ ｷ ｦ20ｸ ｲ4; ｸ ｦ0 0ｸ802 6ｦ86ｦ ｦ ｦ 紊 ( ｲ ｦ ｦ0, ｦ ｦ0) 紊 ( ｹ ｹ ｦ ｲ409 0ｹ ｦ ｦ ｲ4096ｦ0) ｦｲ4 0ｹ ｸ90ｷ3090ｲ ｦ0 0ｹ4030ｸ ｦ ｹ ｸ90ｷ ｷ ｦ0 6ｦ ｦ0 096ｦｷ ｷ36ｦ ｦ ｲ403 0ｹ10ｷ ｹ601040ｸ9 0ｹ ｸ90ｷ ｸ , 6ｦ ｦ606036ｦ ｦ3 09 6ｦ06ｦ ｦ ｸ8 0ｹ ｸ90ｷ : ｦ ｹ ｸ90ｷ30909, 6 76ｦ0 046ｦ ｦ ｦ2036ｦｷ ｦ0, 036ｦ1070ｹ ｸ90ｷ ｦ0, ; , , ｦ ｹ ｸ90ｷ30909; ｦ ( ｦ90ｹ706 0ｹ ｸ90ｷ30909) ｦ0 6ｦ8036ｦｷ ｦ1070ｹ ｸ90ｷ309070ｸ ｸ803090ｲ403, 086ｦ06ｦ ｸ8, ｷ ｦ ｦ ｸ ｸ ｦ ｦ90ｸ86 56ｦ3, 0ｹ307 6ｦ ｸ ｹ ｸ90ｷ ｸ ｹ06ｦｲ403 ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ86 29

7 13ｨｨｨｺｨ 0409ｨ ｨｨ 05ｨ 05ｨｦｨｦ03ｨｺ02ｨ ｨｦ050204ｨ ｨｹ 02ｨ 01ｨｺｨｦ0302 ｨｮｨｬ0201 Yersinia pestis 651 ｨｨ ｷ ｷ3 ｨｪ ｨｪ ｨC5 030ｸ ｦ96ｦ ｸ ｦ96ｦ ｸ ｦ96ｦ ｸ ｦ96ｦ ｷ3096ｦ0, apple016ｦｷ309fl 08apple ｹ0apple apple 紊 0703apple0703apple036ｦ0fl 6 703apple ｦ0fl 06apple 6ｦ0 +5 紊 Yersinia pestis ssp. Ma Mj LBEm AcMf 02 6ｦ ｦ0fl apple6ｦ0 096ｦｷ309fl, 07 apple6ｦ ｦ ｦ20ｸ 紊 apple ｦ0fl 096 5apple6 70ｷ309fl apple6ｦ2090ｹ6016ｦ0fl 0ｹ ｦ609fl Yersinia (Yersinia) pestis tarbagani Fra 1, ｦ0fl 06080ｷ3096ｦ209096ｦｷ309fl, 06036ｦ ｦｷ309fl 08apple09096ｦ ｦ0apple6ｦ6036ｦ apple ｦ0 ｨ fl0ｷ309fl ｦ0, apple ｨｺ ｦ ｦ36ｦ0, apple0703apple096ｦ9 6 7apple04036ｦ0, 08apple ｹ apple070ｹ ｦ8apple07026ｦ ｦ ｦ ｦ307026ｦ ｦ0apple ｦｷ ｦ apple apple016ｦ0 Yersinia (Pfeifferia) pseudotuberculosis O:1b apple070ｹ ｸ90ｷ309fl 026ｦ1apple070ｹ ｸ90ｷ309fl ｨｨapple apple090ｷ ｦ ｸ9 06apple ｦ9fl, apple6ｦ ｦ apple09096ｦ16ｦ apple07 6ｦ ｦ0. ｨｺ apple6ｦ0 046ｦ ｹ6016ｦ0fl ( 6ｦ0apple ｦ0fl) apple fl 09 0ｹ ｸ90ｷ fl 0ｹ4030ｸ8. ｨｦ03 020ｸ fl096ｦ0-0ｸ8 08applefl ｹ apple07 6ｦ0 0ｹ4030ｸ8, 0ｷ309apple apple040ｸ90ｸ fl0ｷ309fl03 apple6ｦ90ｸ : Lb ｨC Lasiopodomys brandti; Mf ｨC Microtus fuscus; Em ｨC Eothenomys melanogaster; Mj ｨC Microtus juldaschi; Ma ｨC Microtus arvalis; Ac ｨC Apodemus chevrieri ｹ4 6ｦｹ96ｦ fl0ｷ ｹ ｦ1apple070ｹ ｸ90ｷ30909, apple6ｦ0fl 096ｦ06 5apple046ｦ306fl apple6ｦ ｦ903 (6 5 6ｦ ｸ apple09-096ｦ apple07 6ｦ ｦ0), 6ｦ0 096ｦ303 08apple apple070ｹ ｸ90ｷ30909 ｨC 08apple07 apple6ｦ ｦ90ｸ fl0ｷ ｹ ｦ ｹ ; 036ｦ apple6ｦ90703, ｦ ｹ1036ｦ ｹ ｸ90ｷ309070ｸ803 08apple07 apple6ｦ ｦ ｸ , 0ｹ ｦ0apple09apple ｨｪｨｴ apple6ｦ0 046ｦ ｹ6016ｦ0fl fl ｹ apple 06036ｦ ｦ0apple6ｦ103apple ｦ3 apple03apple ｸ8 08apple ｹｸ8 0207apple030ｸ fl apple ｦ0 09 0ｹ4030ｸ8, apple0ｸ ｸ ｦ fl0ｷ apple0703apple030ｸ apple ｦｹ06ｦ26ｦ ｦ0 6ｦ0apple0708 6ｦｸ90ｸ fl 0ｹ4030ｸ apple ｦ ｦ ｷ3096ｦ0, ｹ ｹ ｦ30406; ｹ apple ｹ0-0ｸ8 0207apple030ｸ ｸ8 0ｸ applefl03 03apple ｦ0-09fl apple apple ｦ0 ( ｹ ｹ apple060909) ｷ36ｦ0apple0906 6ｦ06 702fl apple09apple0403 apple6ｦ2-040ｸ apple0703apple03040ｸ9 (6 703apple ｸ9) 08apple ｹ40ｸ9 06apple 04, ｸ9 02fl apple ｹ9fl ｨC 086ｦ0apple6ｦ909036ｦ0apple040ｸ ｰ apple0701ｨC ｦ0 O. silantiewi ｱ, apple apple apple07 apple6ｦ ｦ apple apple07 6ｦ fl 0ｹ4030ｸ8; 036ｦ apple6ｦ90703, apple apple fl03 6ｦ309 6ｦ0apple fl 6 50ｸ ｦ ｸ ｹ apple07 apple6ｦ ｦ fl0ｷ30906, 0ｹ apple070ｷ30606 apple6ｦ ｸ ; ｸ9 6ｦ apple fl 05ｨｪｨｴ ｦ1apple apple070ｹ ｸ90ｷ309070ｸ fl0ｷ ｹ apple070ｷ ｦ1070ｷ3, 08apple ｸ ｷ36ｦ0apple ｹ apple6ｦ96 56ｦ306fl ｸ ｷ36ｦ0apple fl ｦ36ｦ ｸ apple apple6ｦ apple03fl apple0907 6ｦ ｷ ｷ36ｦ ; 016ｦ apple ｹ ｦ06 56ｦ3, ｸ80309apple6ｦ9 036ｦ apple ｸ ｦ apple ｦ ｦ0- apple apple ｹ ｦ0apple6ｦ90ｸ8 ｨC fl 0ｹ4030ｸ8. ｨ 09 08apple 06036ｦ ｸ ｷ36ｦ0apple apple fl fl, apple0706 ｨC 6 7apple6ｦ0 046ｦ ｹ ｨC 6 50ｸ8090ｸ86 56ｦ apple09fl, fl ｸ803. ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

8 ｷ3076 5, 05040ｷ ｦ apple6ｦ ｦ3090ｹ ｦ0 ｰ 6ｦ0apple ｦ0fl ｱ ｦ ｦｹ9 02fl 08apple apple6ｦ apple6ｦ0-09fl: ｸ ｦｸ903 08apple09 apple ｸ ｦ3apple ｸ fl apple04086ｦｸ9-0ｸ706 06apple 0ｸ apple 020ｸ803 apple6ｦ6076ｦ3 08apple ｹ apple 020ｸ ｦ3090ｹ , 0ｹ ｷ ｹ , 6ｦ apple090ｹ apple03fl ｷ3096ｦ207-0ｹ ｹ apple6ｦ2090ｹ ; 08apple09 0ｹ fl0ｷ309fl fl 08apple ｹ ｸ90ｷ309070ｸ apple apple ｹ706 06apple 0ｹ0apple apple020ｸ8 08apple0307 0ｸ8 0207apple030ｸ ｦ ｹ06ｦ0, fl apple apple 0408apple fl fl ｦ1020ｸ90ｹ ｸ8 6 50ｸ8apple6ｦ ｦ , 0ｹ307 6ｦ ｦ ｹ ｦ60ｸ80ｸ apple 06036ｦ ｸ803 0ｹ ｸ90ｷ309070ｸ ｷ36ｦ0apple ｦ ｸ ｦ ｸ ｦ apple ｦ ｦ80ｸ apple ｹ ｸ9-0ｷ apple0909: 6 7apple6ｦ0 046ｦ ｦ2036ｦｷ ? ｨｨapple ｸ ｦ , apple0ｸ apple 07016ｦ80306fl ｰ apple070ｸ803 ｱ ｸ7040ｸ9 0307apple090ｸ9 0ｹ ｸ90ｷ30909, 0807apple apple6ｦ96 5apple ｸ ｦ ｸ9 08apple076 7apple6ｦ ｨｨ07016ｦ0 036ｦ16ｦ0fl 08apple076 7apple6ｦ3036ｦ ｦ8apple6ｦ06 502fl03 fl ｹｸ803, 076ｦ0-04fl ｦ0 02fl 01apple ｨｦ apple fl ｦ ｹ0 6 50ｸ8fl6 502fl fl030ｸ ｦ1036ｦ3, 0ｹ307 0ｹ06ｦ apple6ｦ ｸ ｦ401, 6ｦ0 06apple6ｦ ｦ0fl ｸ apple0909 0ｷ apple apple fl 09 0ｹ ｸ90ｷ fl 0ｹ4030ｸ8 036ｦ apple , , ｸ ｦ0! 05 apple apple07-0ｸ8, 08apple09 apple6ｦ apple0909 apple , apple ｷ36ｦ0apple09fl ｦ fl , 6 50ｸ804 6ｦ309 0ｸ802 08apple apple0903apple ｦ ｦ flapple apple0309apple076 56ｦ ｹ apple apple apple07 6ｦ0 0ｹ4030ｸ8, ｸ ｦ0 08apple07fl ｸ apple ｦ (Zhou, Yang, 29), apple0ｸ fl ｸ86 702fl ｹ ｹ , 03036ｦ90ｹ ｨB 09fl0ｸ90ｸ ｨｨ ｦ0, 06036ｦ2 016ｦ1 0ｸ8 08apple ｹｸ apple6ｦ0 046ｦ ｨC 066ｦ2036ｦｷ ｨ ｦ ｹ06ｦ0, , 0ｹ307 08apple 06036ｦ ｸ8 060ｷ36ｦ0apple ｦ ｸ90ｹ06ｦｸ903 apple046 7 apple046 76ｦ0, 6ｦ ｸ903 08apple090ｷ : ｰ apple6 5fl ｱ ｦ ｸ903 ｰ apple- 6 5fl03 036ｦ ｱ ｦ ｹ109 09apple09fl, 02fl apple07-09fl apple07 6ｦ0 Y. pestis 07 ｨB ｦ8apple6ｦ ｦ306fl ｹ ｹ ｦ0, ｦ ｹ903 08apple090ｹ90ｸ8 apple6ｦ ｹ apple6ｦ ｦ ｷ36ｦ0apple09fl apple ｸ apple apple04090ｸ ｸ90ｹ0-07, 0ｹ307 0ｹ apple ｦ apple appleapple0ｸ apple6ｦ fl (punctuated equilibrium). 05ｨｨｨ 05ｨｦ01 ｨ ｨｪｨｺ02ｨｪｨｬｨｺ01 02 apple6ｦ ｨ apple6ｦ209fl: , 08apple ｸ8, 08apple ｸ8 // ｨｦ ｨｪ ｧ ｨC ｦ0apple6ｦ ｨ.05. ｨｮ04036ｦ0. 02.: ｷ3096ｦ0, fl ｨ. ｨｴ ｸ90ｷ309070ｸ8 6ｦ ｸ apple09apple ｹ06ｦ ｹ4030ｸ ｦ06 5apple : 05036ｦ06 5apple , ｨ 6ｦ06 56ｦ0apple ｹ6016ｦ0fl ｦ309016ｦ0 6ｦ103apple0906. ｨｨapple070603apple6ｦ ｹ : 046ｦ4-016ｦ0, ｨ 07apple 6ｦ ｨｴ ｸ90ｷ309fl : 02016ｦ0 0309fl, ｹ ｨｴ.ｨ ｦ36ｦ603apple ｷ : 6 50ｹ06ｦ0fl ｦ ｦ0? ｨ apple apple ｹ ｹ0apple ｨｨ.: 046ｦ4016ｦ0, ｦ0.02., ｨｨapple070ｷ30107 ｨｦ.02., ｸ8apple apple- 030ｸ ｷ3080ｷ fl fl0309 0ｹ4-030ｸ apple ｦ0 // ｦ0, apple fl apple fl ｧ ｨC7. 026ｦ6apple ｨｴ. ｨｨ fl0ｷ30909, ｸ8 09 0ｹ ｸ90ｷ309fl. 02.: 0209apple, ｦ96ｦ0apple ｨ. ｨｬ0ｹ ｦ1apple070ｹ ｸ90ｷ ｦ : 046ｦ4016ｦ0, ｨｨ6ｦ ｨｦ ｸ8 040ｹ9fl 07 08apple09apple ｹ06ｦ apple6ｦ ｸ ｹ ｦ0 // 0504apple. 07 0ｸ ｨｪ. 7. 6ｧ ｨｨ6ｦ ｨ.ｨｹ apple03040ｸ : , ｨｨapple070907apple apple ｦ20ｸ806 08apple ｦ1-03apple0906 // ｨｬ apple ｨｪ ｧ ｨC ｷ , 05040ｷ ｦ0 04.ｨ., ｸ ｷ ｹ6016ｦ0fl 0603apple apple6ｦ0 070ｹ06ｦ ｹ4030ｸ ｦ apple07060ｹ90109 // 05. apple6ｦ ｸ9 ｨｪapple ｷ303apple6ｦ ｨｮ. II. ｨｪapple ｹ6016ｦ0fl ｷ3096ｦ0 / ｨｦ apple. 0107apple0904.ｨｨ., 01apple ｷ , 046ｦ , ｦ0; 076ｦ706: ｨｪapple0708-0ｷ303apple, ｨC ｷ , 05040ｷ ｦ0 04.ｨ. ｨｴ ｹ ｦ ｸ8 0ｹ ｸ90ｷ apple07 6ｦ0 0ｹ4030ｸ8 Yersinia pestis apple09apple07 0ｸ ｹ06ｦ // ｨ ｨｺ apple ｧ ｪ ｷ , 05040ｷ ｦ0 04.ｨ. ｨｮ04036ｦ0. ｨｨapple ｹ ｸ90ｷ309fl 0ｹ ｹ ｸ8 (0ｹ ｹ , apple6ｦ2090ｹ ｷ3096ｦ20ｸ8 6ｦ ｸ8). 02.: , c ｷ , 05040ｷ ｦ0 04.ｨ. 026ｦ1apple apple070ｹ ｸ90ｷ309fl apple apple fl apple ｹ ｦ ｹ apple07 6ｦ0 Yersinia pestis // ｨ ｨｺ apple ｧ ｨC ｸ ｹ ｹ ｸ8 086ｦ ｸ803 09apple // ｨｨapple ｸ ｦ60ｸ ｷ ｸ ｨC88. ｨｰ6 56ｦ0apple0ｷ ｨｴ ｹ ｦ apple ｹ ｸ9-0ｷ : 046ｦ4016ｦ0, ｨｹ , ｨｴ ｸ90ｷ ｹ9. 02.: 050ｸ , Achtman M., Zurth K., Morelli G. et al. Yersinia pestis, the cause of plague, is a recently emerged clone of Yersinia pseudotuberculosis // PNAS V ｧ8 24. P ｨC ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29

9 13ｨｨｨｺｨ 0409ｨ ｨｨ 05ｨ 05ｨｦｨｦ03ｨｺ02ｨ ｨｦ050204ｨ ｨｹ 02ｨ 01ｨｺｨｦ0302 ｨｮｨｬ0201 Yersinia pestis 653 Achtman M., Morelli G., Zhu P. et al. Microevolution and history of the plague bacillus, Yersinia pestis // PNAS. 24. V ｧ8 51. P ｨC Cui Y., Li Y., Gorge O., Platonov M.E. et al. Insight into microevolution of Yersinia pestis by clustered regularly interspaced short palindromic repeats // PloS ONE. 28. V. 3. 6ｧ8 3. P. 1ｨC10. Eppinger M., Rosovitz M.J., Fricke F.W. et al. The complete genome sequence of Yersinia pseudotuberculosis IP31758, the causative fgent of far east Scarlet-like fever // PLoS Genet. 27. V. 3. 6ｧ8 8. ｨｺ. E142. Goldschmidt R. The material basic of evolution. N. Haven; Connecticut: Yale Chiv. Press, P. 205ｨC206. Gu J., Neary J.L., Sanchez M. et al. Genome evolution and functional divergence in Yersinia // J. Exp. Zool. (Part B. Mol Dev Evol). 26. V ｧ8 1. P. 37ｨC49. Konstantinidis K.T., Ramette A., Tiedje J.M. The bacterial species definition in the genomic era // Phil. Trans. 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Bacteriol. 22. V P. 1019ｨC1027. Pourcel C., Andre-Mazeaud F., Neubauer H. et al. Tandem repeats analysis for the high resolution phylogenetic analysis of Yersinia pestis // Skurnik M., Peippo A., Ervela E. Characterization of the O-antigen gene cluster of Yersinia pseudotuberculosis and the cryptic O-antigen gene cluster of Yersinia pestis shows that the plague bacillus is most closely related to and has evolved from Y. pseudotuberculosis serotype O:1b // Mol. Microbiol. 20. V ｧ8 2. P. 316ｨC330. Song Y., Tong Z., Wang J. et al. Complete Genome Sequence of Yersinia pestis Strain 911, an Isolate Avirulent to Humans // DNA Research. 24. V ｧ8 3. P. 179ｨC 197. Torrea G., Chenal-Francisque V., Leclercq A. et al. Efficient Tracing of Global Isolates of Yersinia pestis by restriction fragment Length Polymorphism Analisis Using Three Insertion Sequences as Probes // J. Clinical Microbiol. 26. V ｧ8 6. P. 2084ｨC2092. Vergnaud G., Li Y., Gorge O. et al. Analysis of the Three Yersinia pestis CRISPR Loci Provides New Tools for Phylogenetic Studies and Possibly for the Investigation of Ancient DNA // Adv. Exp. Med. Biol. 27. V P. 327ｨC338. Wren B.W. The Yersinia ｨC a model genus to study the rapid evolution of bacterial pathogens // Nat. Rev. Microbiol. 23. V. 1. 6ｧ8 1. P. 55ｨC64. Zhou D., Han Y., Song Y. et al. DNA microarray analysis of genome dynamics in Yersinia pestis: insights into bacterial genome microevolution and niche adaptation // J. Bacteriol. 24a. V ｧ8 15. ｨｺ. 5138ｨC5146. Zhou D., Han Y., Song Y. et al. Comparative and evolutionary genomics of Yersinia pestis // Microbes Infect. 24b. V. 6. 6ｧ8 13. P. 1226ｨC1234. Zhou D., Tong Z., Song Y. et al. Genetics of metabolic variations between Yersinia pestis biovars and the proposal of a new biovar, microtus // J. Bacteriol. 24c. V ｧ8 15. P. 5147ｨC5152. Zhou D., Yang R. Molecular Darwinian evolution of virulence in Yersinia pestis // Infect. Immun org/cgi/content/abstract/iai.01477ｨc08v1. Principles of Speciation of the Plague Causative Agent Versinia Pestis: Gradualism or Saltation? V. V. Suntsov a, c and N. I. Suntsova b, c a Severtsov Institute of Problems of Ecology and Evolution, Russian Academy of Sciences, Leninskii pr. 33, Moscow, Russia b Moscow State University, Ecological Center, Leninskie gory, Moscow, c Russian and Vietnam Tropical Center, No. 3 Street 3/2, 10 Distr., Ho Chi Minh City, Vietnam [email protected] Abstract ｪThe saltation origin of the causative agent of the plague Yersinia pestis from the pseudotuberculosis microbe Y. pseudotuberculosis O:1b has been proclaimed in recent investigations on molecular genetics. The speciation process in this case is proposed to be connected with horizontal inclusion of exogenous genetic structures (such as specific plasmids pfra and ppst) into the genome of the ancestral form. The alternative ｰDarwinian ｱ model of the gradual origin of the plague agent is proposed based on ecological factors. The comparison of two evolutionary scenarios, saltation and gradual, is performed; the latter seems more likely. ｨ ｨ 0505ｨｪｨ ｨｹ ｨｺ ｨｺｨ ｨｹ 03ｨ ｨｦｨｦ07ｨ ｨｮ050102ｨｹ 6ｧ8 6 29