сравнительный анализ изменчивости митохондриальной днк у

;BHEH=BYFHJYlhf‹k289–295
M>D
FHE?DMEYJG:Y;BHEH=BY
KJ:<GBL?EVGUC:G:EBABAF?GQB<HKLB
FBLHOHG>JB:EVGHC>GDMDHJXRDH<UOJU;1
‹]E:KdmjbobgZ:=He_cgbdF<IZgvdh\Z
Bgklblml[bheh]bbfhjy><HJ:G<eZ^b\hklhd
e-mail: [email protected]
KlZlvyijbgylZdi_qZlb]
M iylb \b^h\ dhjxrdh\uo ju[ k_f_ckl\Z Osmeridae k bkihevah\Zgb_f j_kljbdpbhggh]h ZgZebaZ RFLP)
bkke_^h\ZgmqZklhdfblhohg^jbZevghc>GDdh^bjmxsbcND5/NDkm[t_^bgbpugZ^hdkb^^_]b^jh]_gZauZfiebnbpbjh\Zgguc \ ihebf_jZaghc p_ighc j_Zdpbb JKR Mjh\_gv ^b\_j]_gpbb gmde_hlb^guo ihke_^h\Zl_evghkl_cfblhohg^jbZevghc>GDf_`^m\b^Zfbe_`bl\ij_^_eZohl(Hypomesus nipponensis bH. japonicus)
^h (Osmerus mordax dentex b Mallotus villosus catervarius). Ijb wlhf mjh\_gv ]_g_lbq_kdhc ^b\_j]_gpbb
f_`^mihimeypbyfbH. nipponensis khklZ\ey_lH. japonicus – hl^h Osmerus mordax dentex –
Hlkmlkl\b_ h[sbo ]Ziehlbih\ iha\hey_l ^bnn_j_gpbjh\Zlv [ebadhjh^kl\_ggu_ \b^u dhjxr_d qlh
fh`_lihfhqvijbj_r_gbbkihjguo\hijhkh\kbkl_fZlbdbb[bh]_h]jZnbb^Zggh]hk_f_ckl\Zju[
Dexq_\u_keh\ZOsmeridaefblhohg^jbZevgZy>GDj_kljbdpbhggucZgZebaRFLPihebf_jZagZyp_igZyj_ZdpbyPCR]_g_lbq_kdZy^b\_j]_gpby
Comparative PCR-RFLP-analysis of the mitochondrial DNA variation in smelts (Osmeridae). L. A. Skurikhina, A. G. Oleinik, M. V. Pankova (Institute of Marine Biology, Far East Branch, Russian Academy of Sciences, Vladivostok 690041)
The mtDNA region coding for ND5/ND6 subunits of NADH-dehydrogenase mitochondrial DNA amplified in the
polymerase chain reaction (PCR) was studied, using the analysis of restriction fragment length polymorphism (RFLP), in
five species of smelts (the family Osmeridae). The divergence of mtDNA nucleotide sequences between the species was
in the range from 11.9 (Hypomesus nipponensis and H. japonicus) to 24.7% (Osmerus mordax dentex and Mallotus villosus catervarius). The level of genetic divergence between populations was 0.32% for H. nipponensis, 0.08 to 0.15% for
H. japonicus, and 0.025% for Osmerus mordax dentex. The absence of common haplotypes allows one to differentiate
between closely related species of smelts, which can be helpful in solving problems of systematics and biogeography of
the Osmeridae. (Biologiya Morya, Vladivostok, 2004, vol. 30, no. 4, pp. 289–295).
Key words: Osmeridae, mitochondrial DNA, restriction endonuclease analysis (RFLP), polymerase chain reaction
(PCR), genetic divergence.
[bgZlhjghc baf_gqb\hklb b h[eZ^Z_l [he__ \ukhdhc
kdhjhklvx gmde_hlb^guo aZf_g q_f y^_jguc ]_ghf
%URZQHWDOWlhiha\hey_lgZb[he__lhqghijh\h^blv^bnn_j_gpbZpbx[ebadhjh^kl\_gguolZdkhgh\
< ^Zgghc jZ[hl_ gZfb ij_^ijbgylZ ihiuldZ bkke_^h\Zlv baf_gqb\hklv fblhohg^jbZevghc >GD
fl>GD m iylb \b^h\ dhjxrdh\uo ju[ \oh^ysbo \
khklZ\gZa\Zgguo\ur_jh^h\k_f_ckl\Z2VPHULGDH
<\h^h_fZoZabZlkdh]hih[_j_`vyb\ijb[j_`guo
\h^Zo k_\_jhaZiZ^ghc qZklb Lboh]h hd_ZgZ h[blZxl
ij_^klZ\bl_eb gZb[he__ fZkkh\uo jh^h\ k_f_ckl\Z
Osmeridae – Osmerus, Hypomesus b Mallotus, \dexqZxsbo fhjkdb_ ijhoh^gu_ b `beu_ nhjfu ;_j] 0F$OOLVWHUDexdZgh\1HOVRQQ_j_rg_\ -RKQVRQ 3DWWHUVRQ J_r_lgbdh\ b ^j
Khdheh\kdZyb^jKhdheh\kdbcHdkxavyg
N_^hjh\ b ^j ?keb kbkl_fZlbdZ jh^h\
Osmerus b Mallotus g_ \uau\Z_l m bkke_^h\Zl_e_c hkh[uo jZagh]eZkbc lh lZdkhghfbq_kdb_ hlghr_gby b
[bh]_h]jZnby jh^Z Hypomesus \gh\v b \gh\v h[km`^Zxlky b mlhqgyxlky +DPDGD 0F$OOLVWHU DexdZgh\ =jbp_gdhQmjbdh\Qmjbdh\DZji_gdh6DUXZDWDU\
HWDOQ_j_rg_\b^j
Ba\_klgh qlh \ ihke_^g__ \j_fy ^ey j_r_gby
fgh]bo lZdkhghfbq_kdbo ijh[e_f b ijh\_jdb jZaebqguo ]bihl_a ijb\e_dZxlky fhe_dmeyjgh]_g_lbq_kdb_
bkke_^h\Zgby Ijb wlhf [hevrh]h \gbfZgby aZkem`b\Z_l bamq_gb_ fblhohg^jbZevgh]h ]_ghfZ hj]Zgbafh\
dhlhjucgZke_^m_lkyihfZl_jbgkdhcebgbb[_aj_dhf1
F:L?JB:EBF?LH>BD:
FZl_jbZehfihkem`bebij_iZjZlu>GDhlju[kh[jZgguo \\_k_gg_e_lgbc i_jbh^ ] lZ[e <b^h\Zy
ijbgZ^e_`ghklvgZhkgh\_fhjnheh]bq_kdboijbagZdh\mklZgh\e_gZB:Q_j_rg_\uf
Bg^b\b^mZevgu_ ij_iZjZlu lhlZevghc >GD ihemqZeb
ba nbdkbjh\Zgguo ldZg_c i_q_gb b k_j^pZ ih klZg^Zjlghc
f_lh^bd_ 6DPEURRN HW DO Baf_gqb\hklv fl>GD ZgZebabjh\Zebbkihevamyj_kljbdpbhggucZgZeba5)/3mqZkldZ1'1'dh^bjmxs_]h^\_km[t_^bgbpugZ^hdkb^^_]b^jh]bgZauZfiebnbpbjh\Z\_]h\ihebf_jZaghcp_ighcj_Zdpbb3&5Ijhly`_gghklvwlh]hmqZkldZkhklZ\ey_liZj
hkgh\Zgbc _]h iheh`_gb_ gZ dZjl_ fl>GD ijZcf_jgu_ ihke_^h\Zl_evghklb b mkeh\by ZfiebnbdZpbb ijb\_^_gu \ jZ[hl_=Zj_llZkkhZ\lhjZfb*KDUUHWWHWDO
Bkke_^h\Zgb_hkms_kl\e_ghijbnbgZgkh\hcih^^_j`d_]jZglh\JNNBbJNNBIjbfhjv_‹b
289
KDMJBOBG:B>J
290
LZ[ebpZBkke_^h\Zggu_\b^ubihimeypbbdhjxrdh\uoju[k_f_ckl\ZOsmeridae
Ghf_j
1
2
3
4
5
6
7
8
9
<b^
Ihimeybpy
:fmjkdbcaZeb\k_\aZiqZklvaZeI_ljZ<_ebdh]hYihgkdh]hfhjy
[Hev]Zk_\aZiqZklvYihgkdh]hfhjy
H. japonicus (Brevoort, 1856)
[Wdki_^bpbbaZeIhkv_lx]haZiqZklv
H. japonicus (Brevoort, 1856)
aZeI_ljZ<_ebdh]hYihgkdh]hfhjy
jK_j_[jygdZk_\aZiqZklvYihgkdh]hfhjy
H. nipponensis McAllister, 1963
[. Hev]Z
H. nipponensis McAllister, 1963
jKZfZj]Zx]haZiqZklvLZlZjkdh]hijheb\Z
H. olidus (Pallas, 1814)
Mallotus villosus catervarius (Pennat, 1784) [GZ]Z_\Zk_\qZklvHohlkdh]hfhjy
Osmerus mordax dentex, Steindachner, 1870 [. Hev]Z
jFZdkbfh\dZk_\aZiqZklvYihgkdh]hfhjy
O. mordax dentex, Steindachner, 1870
Hypomesus japonicus (Brevoort, 1856)
H[t_f\u[hjdbwda
10
15
18
15
8
6
12
18
18
Khhl\_lkl\m_lgmf_jZpbbihimeypbc\lZ[e±
:fiebnbpbjh\ZggucnjZ]f_glZgZebabjh\ZebgZ[hjhf
baj_kljbdpbhgguon_jf_glh\\mkeh\byoj_dhf_g^h\Zgguoba]hlh\bl_e_f0%,)HUPHQWDV/LWKXDQLDKb[Wgabf
Jhkkby $VX, **1&& $YD,, **:&& %VW1, &&:**
BstUI (CG&* (FR59 *$7$7& +LQI, *$17& 0ER,
(*$7&0VS,&&**5VD,*7$&6W\,&&::**7DJ,
(TC*$ Ijh^mdlu j_kljbdpbb jZa^_eyeb \ ± Z]Zjhaghf ]_e_ kh^_j`Zs_f h^gm qZklv 8OWUD 3XUH™ agarose ("BRL
*LEFR*UDQG1<b^\_qZklb6\QHUJHO™ (Diversified Biotech
,QF %RVWRQ 0$ \ djZlghf ljbk[hjZlghf [mn_j_
6DPEURRN HW DO NjZ]f_glu >GD \ ]_e_ hdjZrb\Zeb
wlb^bmf[jhfb^hfbnhlh]jZnbjh\Zeb\ ijhoh^ys_f mevljZnbhe_lh\hfk\_l_<dZq_kl\_fhe_dmeyjguofZjd_jh\^ebgu
bkihevah\Zeb njZ]f_glu >GD nZ]h\ λ b φO ]b^jhebah\Zggu_ j_kljbdlZaZfb 3VW, b +LQI, khhl\_lkl\_ggh Z lZd`_
gZ[hjnjZ]f_glh\>GDdjZlguoiZjZfhkgh\Zgbc%5/
Gibco", Grand N.Y.).
=_l_jh]_gghklvf_`^m\u[hjdZfbhij_^_eyebkbkihevah\Zgb_f f_lh^Z Fhgl_DZjeh \ iZd_l_ ijh]jZff 5($3
0F(OUR\HWDODhebq_kl\haZf_ghkgh\ZgbcgZgmde_hlb^guckZcl(d)boklZg^Zjlgh_hldehg_gb_\_ebqbgm^b\_j]_gpbbf_`^m\u[hjdZfbδZlZd`_gmde_hlb^gmxπb]Ziehlbibq_kdmx (h) \_ebqbgu jZaghh[jZaby jZkkqblu\Zeb \
iZd_l_ijh]jZff5($30F(OUR\HWDOKlZlbklbq_kdZy
h[jZ[hldZ ihemq_gguo j_amevlZlh\ \dexqZeZ n_g_lbq_kdbc
ZgZeba ijbagZdh\ f_lh^hf g_\a\_r_gghc iZjgh]jmiih\hc
kj_^g_c k\yab 83*0$ k bkihevah\Zgb_f ijh]jZffu
NTSYS (Rohlf, 1990).
J?AMEVL:LUBH;KM@>?GB?
Mbkke_^h\ZgguogZfbdhjxr_d\uy\e_gh
]Ziehlbih\ fl>GD m Hypomesus japonicus ± m
H. nipponensis – m H. olidus ± m Mallotus villosus
catervarius – m Osmerus mordax dentex ± lZ[e Wlb]ZiehlbiuhlebqZxlky\h\k_oZgZebabjm_fuolZdkhghfbq_kdbo]jmiiZo
:gZeba]_h]jZnbq_kdhc]_l_jh]_gghklbqZklhl]Ziehlbih\ f_lh^hf Fhgl_DZjeh 0F(OUR\ HW DO \uy\beklZlbklbq_kdbagZqbfu_jZaebqbyf_`^m^\mfy
bkke_^h\Zggufb ihimeypbyfb H. nipponensis (χ2 =
=J <<ZlZd`_f_`^mlj_fyihimeypbyfb
H. japonicus (χ2 J = JZaebqby f_`^m ihimeypbyfbO. mordax dentexhdZaZebkvg_^hklh\_jgufb
(χ2 J JZkkqblZgu lZd`_ agZq_gby χ2
f_`^mh[t_^bg_ggufb\u[hjdZfbH. japonicus, H. nipponensis, H. olidus, M. villosus catervarius b O. mordax
dentex JZaebqby f_`^m \b^Zfb hdZaZebkv \ukhdh^hklh\_jgufbJ << 0.001).
Kj_^gb_ agZq_gby jZaghh[jZaby ]Ziehlbih\ b
mjh\_gv gmde_hlb^ghc baf_gqb\hklb \ ihimeypbyo
H. japonicus (0.9231 ±bkhhl\_lkl\_ggh
hdZaZebkv \ur_ q_f \ ihimeypbyo H. nipponensis
(0.5845 ±bbO. mordax dentex (0.3105 ±
± b lZ[e Ihkdhevdm ba j KZfZj]Z
bkke_^h\Zgh g_[hevrh_ qbkeh hkh[_c H. olidus ihemq_ggu_ hp_gdb \_jhylgh g_ hljZ`Zxl j_Zevguc mjh\_gv ]Ziehlbibq_kdh]h b gmde_hlb^gh]h jZaghh[jZaby
^Zggh]h\b^Z
AgZq_gby \gmljb b f_`\b^h\hc ^b\_j]_gpbb ^ey
ZgZebabjm_fuolZdkhgh\1HL7DMLPD1HL0LOOHU
\ujZ`_ggu_\ijhp_glZogmde_hlb^guoaZf_ggZ
j_kljbdpbhgguc kZcl hij_^_e_gu k mq_lhf hlghkbl_evguoqZklhl]Ziehlbih\\ihimeypbyolZ[e<_ebqbgZ ^b\_j]_gpbb gmde_hlb^guo ihke_^h\Zl_evghkl_cfl>GDf_`^mH. nipponensis b H. japonicus khklZ\beZ hdheh H. nipponinsis b H. olidus – 14.1%,
H. japonicus b H. olidus ± Mjh\_gv ^b\_j]_gpbb
fl>GD f_`^m jh^Zfb Hypomesus b Osmerus jZ\gyeky
17.9%, Hypomesus b Mallotus – 18.0%, Osmerus b
Mallotus ± Ijb wlhf mjh\_gv ]_g_lbq_kdhc ^b\_j]_gpbb f_`^m ihimeypbyfb H. nipponensis khklZ\eye H. japonicus ± hl ^h O. mordax
dentex ± LZdbf h[jZahf mjh\_gv \gmljb\b^h\hcbaf_gqb\hklbfl>GD^ey\k_obkke_^h\ZgguolZdkhgh\g_ij_\urZe
GZ hkgh\_ ihemq_gguo hp_ghd ihkljh_gZ n_gh]jZffZ kbkihevah\Zgb_f f_lh^Z g_\a\_r_gghc iZjgh
]jmiih\hc kj_^g_c k\yab 83*0$ kf jbkmghd Bk
ke_^h\Zggu_ ihimeypbb H. nipponensis h[jZah\Zeb
dhfiZdlgmx]jmiimddhlhjhcijbkh_^bgy_lkydeZkl_j
h[t_^bgyxsbc ihimeypbb H. japonicus. >Ze__ d wlhfm
deZkl_jm ihke_^h\Zl_evgh ijbkh_^bgyxlky \_l\b
H. olidus, O. mordax dentexbgZb[he__^b\_j]bjh\Z\r_c
hlgboM. villosus catervarius.
Ke_^m_lhlf_lblvqlhjZ[hlihdhjxrdh\ufju[Zf \uiheg_gguo ]_g_lbdh[bhobfbq_kdbfb b fhe_;BHEH=BYFHJYlhf‹
KJ:<GBL?EVGUC:G:EBA
291
LZ[ebpZ<klj_qZ_fhklv]Ziehlbih\fl>GD\ihimeypbyojZaguo\b^h\dhjxrdh\uoju[k_f_ckl\Z2VPHULGDH
H[hagZIhjy^q_gb_
dh\uc
]Ziehlbghf_j
BstUI EcoRV RsaI
ih\
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
HJA
HJB
HJC
HJD
HJE
HJF
HJG
HJH
HJI
HJJ
HJK
HJL
HJM
HJN
HJO
HJP
HJQ
HJR
HJS
HJT
HNA
HNB
HNC
HND
HNE
HNF
HOA
MVA
MVB
MVC
MVD
MVE
MVF
OMA
OMB
OMC
OMD
OME
OMF
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
C
C
C
C
C
C
E
D
D
F
D
D
D
B
B
B
B
B
B
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
C
C
C
C
C
C
C
D
D
D
D
D
D
B
B
B
B
B
B
B
I
B
E
B
B
B
E
E
B
B
I
B
I
I
B
B
B
B
B
H
D
F
F
J
F
G
C
C
C
C
C
C
A
A
A
A
A
A
J_kljbdlZau
MspI
AsuI
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
C
C
C
C
C
C
E
G
D
D
D
D
D
B
B
B
B
B
B
B
B
B
B
B
F
B
F
B
B
H
G
B
B
B
B
B
B
G
F
C
C
C
C
C
C
E
D
D
D
D
D
D
A
A
A
J
A
A
Ihimeypby
HinfI AvaII BstNI MboI
B
B
B
B
B
B
B
B
B
D
B
B
D
B
D
K
B
L
D
M
G
C
C
C
C
C
F
E
E
E
J
E
H
A
A
A
A
I
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
F
A
A
A
C
C
C
C
C
A
E
D
D
D
D
D
D
B
B
B
B
B
B
A
A
A
H
A
A
A
H
A
A
A
A
A
A
A
A
A
A
A
A
F
C
C
C
C
C
E
D
D
D
D
D
D
B
B
B
B
B
G
B
B
B
B
B
B
G
B
B
B
B
B
B
B
B
B
B
B
G
B
F
C
C
C
C
C
E
D
D
D
D
D
D
A
A
A
A
A
A
StyI
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
C
C
C
C
C
C
E
D
D
D
D
F
D
B
B
B
B
B
B
TaqI 1 2 3 4 5 6 7 8 9
D
D
B
G
G
G
B
G
B
B
D
B
D
B
B
D
D
D
G
D
H
C
C
I
I
C
F
E
E
E
E
E
E
A
J
K
A
A
A
1
1
1
1
2
1
2
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
5
0
0
0
1
0
2
2
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
2
0
0
0
0
0
4
2
0
0
3
0
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
2
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
6
1
1
1
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
15
0
0
0
1
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
15
1
1
1
0
0
I_j\Zy[md\Z\h[hagZq_gbb]Ziehlbih\mdZau\Z_lgZijbgZ^e_`ghklvdjh^m+±Hypomesus, M – Mallotus, O – Osmerus.
dmeyjgh]_g_lbq_kdbfb f_lh^Zfb \fbjh\hc gZmqghc
ebl_jZlmj_g_fgh]hh^gZdhhgbho\Zlu\Zxl\k_bkke_^m_fu_gZfbjh^uk_f_ckl\Z2VPHULGDHLZd^eyb^_glbnbdZpbb jZaguo \b^h\ dhjxr_d jh^Z Hypomesus m
dZebnhjgbckdh]h ih[_j_`vy ij_^eh`_gu Zeehabfgu_
fZjd_ju iha\heyxsb_ jZaebqZlv H. nipponensis b
H. transpacificus hij_^_e_gb_ dhlhjuo ih fhjnheh]bq_kdbfijbagZdZfaZljm^g_gh (Stanley et al., 1995; TrenKDPHWDOHlf_lbfqlhgZrbj_amevlZlulZd`_
;BHEH=BYFHJYlhf‹
iha\heyxl gZ^_`gh ^bnn_j_gpbjh\Zlv H. nipponensis,
H. japonicus b H. olidus \ lhf qbke_ b kbfiZljbqgu_
ihimeypbb \ [ Hev]Z ihkdhevdm ijZdlbq_kdb ih \k_f
bkihevah\Zgguf\ZgZeba_j_kljbdlZaZf^eygZa\Zgguo
\b^h\ ihemq_gu jZagu_ ]Ziehlbiu fl>GD lZ[e IjbwlhfH. japonicus bH. nipponensis ijhy\eyxlk_[y
dZd [ebadhjh^kl\_ggu_ \b^u Z H. olidus hl gbo h[hkh[e_gkfjbkmghdqlhkh]eZkm_lkykbf_xsbfbky\
ebl_jZlmj_ ^Zggufb \ qZklghklb k ihemq_ggufb gZ
KDMJBOBG:B>J
292
LZ[ebpZAgZq_gby]Ziehlbibq_kdh]hbgmde_hlb^gh]hjZaghh[jZaby\ihimeypbyodhjxrdh\uoju[k_f_ckl\ZOsmeridae
Ghf_j
1
2
3
4
5
6
7
8
9
<b^
Hypomesus japonicus
H. japonicus
H. japonicus
H. nipponensis
H. nipponensis
H. olidus
Mallotus villosus catervarius
Osmerus mordax dentex
O. mordax dentex
Ihimeypby
:fmjkdbcaZeb\
[. Hev]Z
[. Wdki_^bpbb
j. K_j_[jygdZ
[. Hev]Z
j. KZfZj]Z
[. GZ]Z_\Z
[Hev]Z
[FZdkbfh\dZ
=Ziehlbibq_kdh_
jZaghh[jZab_h
Gmde_hlb^gh_
jZaghh[jZab_ π
0.9556±0.0594
0.8857±0.0686
0.9281±0.0401
0.4190±0.1132
0.7500±0.1391
0.0000±0.0000
0.7576±0.1222
0.3072±0.1316
0.3137±0.1376
0.0058
0.0390
0.0059
0.0102
0.0422
0.0000
0.0035
0.0011
0.0092
H[t_f\u[hjhdijb\_^_g\lZ[e
hkgh\Zgbbfhjnheh]bq_kdh]hZgZebaZDexdZgh\
1975; Saruwatary et al., 1997).
J_kljbdpbhgguc ZgZeba fl>GD ZleZglbq_kdhc
fhc\uM. villosusijh\_^_gguc>h^khghfkkhZ\lhjZfb
'RGVRQ HW DO iha\hebe hijh\_j]gmlv fg_gb_ h
lhf qlh \ k_\_jhaZiZ^ghc :leZglbd_ h[blZ_l h^gZ
[hevrZy iZgfbdlbq_kdZy ihimeypby wlh]h \b^Z Kj_^gbcijhp_gl^b\_j]_gpbbihke_^h\Zl_evghkl_cfl>GD
f_`^m ^\mfy ]jmiiZfb ]Ziehlbih\ h[gZjm`_ggufb
kj_^bbkeZg^kdbob[Zj_gp_\hfhjkdboklZ^fhc\u[ue
^h\hevgh \ukhd b khklZ\eye AgZq_gb_ ]Ziehlbibq_kdh]hjZaghh[jZaby^eyZgZebabjm_fuoihimeypbc
dhe_[Zehkv hl ^h GZrb ^Zggu_ lZd`_ e_`Zl \ wlbo ij_^_eZo ihkdhevdm m bkke_^h\Zgghc gZfb
ihimeypbb ba [ GZ]Z_\Z mjh\_gv ]Ziehlbibq_kdh]h
jZaghh[jZaby^hklb]ZelZ[e
;he__ ihegh bamq_gu dhjxrdb jh^Z Osmerus
%DE\HWDO7D\ORU%HQW]HQb^j. Bkke_^h\Zgb_ jh^kl\_gguo hlghr_gbc ZgZ^jhfguo dhjxr_d
O. mordax, g_j_klmxsbo \ jZaguo ijblhdZo wklmZjby
aZeK\EZ\j_glby DZgZ^Z hkgh\Zggh_ gZ j_kljbdpbhgghf ZgZeba_ fl>GD ihdZaZeh bo ]_g_lbq_kdmx h^ghjh^ghklvMjh\_gv]Ziehlbibq_kdh]hjZaghh[jZaby\
bamq_gguo\u[hjdZodhe_[Zekyhl^h%DE\HW
DOIha^g__ijbbamq_gbb\u[hjhdO. mordax
ba wklmZjby b aZeK\EZ\j_glby k ihfhsvx wlh]h `_
ZgZebaZ mklZgh\e_gh qlh wdheh]bq_kdb_ ijhp_kku b
[bh]_h]jZnbq_kdb_ kh[ulby y\eyxlky nZdlhjZfb b]jZxsbfb[hevrmxjhev\nhjfbjh\Zgbbihimeypbhgghc ]_g_lbq_kdhc kljmdlmju m dhjxr_d wlh]h \b^Z \
^Zgghfj_]bhg_%HUQDWFKH]0DUWLQ
O. mordax, h[blZxsZy gZ k_\_jh\hklhd_ K_\_jghc :f_jbdb ij_^klZ\e_gZ fgh`_kl\hf ihimeypbc
Bgh]^Z ZgZ^jhfgu_ dhjxrdb aZoh^yl \ j_qgu_ kbkl_fu ]^_ `b\ml ij_kgh\h^gu_ ihimeypbb b g_j_^dh \
h^ghf b lhf `_ ha_j_ kh\f_klgh h[blZxl dhjxrdb
ghjfZevgh]h jZaf_jZ b bo dZjebdh\Zy nhjfZ 7D\ORU
%HQW]HQ Ijb 5)/3ZgZeba_ fl>GD ZgZ^jhfguo
bij_kgh\h^guodhjxr_d\uy\e_gu^\_nbeh]_g_lbq_kdb_ ]jmiiu jZaebqZxsb_ky gmde_hlb^guo ihke_^h\Zl_evghkl_c Mjh\_gv gmde_hlb^gh]h jZaghh[jZabymdZjebdh\uodhjxr_djZaguoihimeypbcdhe_[Zeky hl ^h \ kj_^g_f hg khklZ\be
mdhjxr_dghjfZevgh]hjZaf_jZ±hl
^hkj_^gbc±bmZgZ^jhfguoihimeypbc±hl^h=Ziehlbibq_kdh_
jZaghh[jZab_ \ p_ehf [ueh g_kdhevdh \ur_ q_f
mbkke_^h\Zgguo gZfb ihimeypbc ZabZlkdhc dhjxrdb
lZ[e b khklZ\eyeh \ kj_^g_f ^ey dZjebdh\ ^eyju[ghjfZevgh]hjZaf_jZ±b^eyZgZ^jhfguo
ihimeypbc±7D\ORU%HQW]HQ
Ijb bkke_^h\Zgbb ^ebgu j_kljbdpbhgguo njZ]f_glh\ fl>GD b ihebfhjnbafZ gmde_hlb^guo ihke_^h\Zl_evghkl_c mqZkldZ pblhojhfZ E 7D\ORU 'RGVRQ
LZ[ebpZAgZq_gby^b\_j]_gpbbfl>GDf_`^m\b^Zfbbihimeypbyfbdhjxrdh\uoju[k_f_ckl\ZOsmeridae
<_ebqbgZ^b\_j]_gpbbσ
Ghf_j
<b^
ihimeypby
1
2
3
4
5
6
7
8
9
1
2
3
4
5
6
7
8
9
Hypomesus japonicus:fmjkdbcaZeb\
H. japonicus ([. Hev]Z)
H. japonicus ([. Wdki_^bpbb)
H. nipponensis (j. K_j_[jygdZ)
H. nipponensis[Hev]Z
H. olidus (j. KZfZj]Z)
Mallotus villosus catervarius ([. GZ]Z_\Z)
Osmerus mordax dentex[Hev]Z
O. mordax dentexjFZdkbfh\dZ
––
0.0015
0.0008
0.1483
0.1118
0.1849
0.2193
0.1371
0.1363
––
0.0012
0.1143
0.0811
0.1706
0.1964
0.1331
0.1325
––
0.1479
0.1114
0.1887
0.2215
0.1380
0.1371
––
0.0032
0.1485
0.1390
0.2639
0.2644
––
0.1337
0.1356
0.2019
0.2025
––
0.1665
0.2006
0.1964
––
0.2451
0.2498
––
0.0003
––
;BHEH=BYFHJYlhf‹
KJ:<GBL?EVGUC:G:EBA
293
N_g_lbq_kdb_hlghr_gbyf_`^m\b^Zfbdhjxrdh\uoju[k_fOsmeridaeih^ZggufPCR-RFLPZgZebaZfblhohg^jbZevghc>GDDhwnnbpb_gldhn_g_lbq_kdhcdhjj_eypbbIhhkb±hp_gdb^b\_j]_gpbbNei, Tajima, 1981; Nei, Miller, 1990).
k^_eZgZ ihiuldZ \uykgblv aZdhghf_jghklb w\hexpbb jh^Z Osmerus. >b\_j]_gpby ihke_^h\Zl_evghkl_cfl>GDf_`^mlj_fynhjfZfbdhjxr_dlbohhd_ZgkdhZjdlbq_kdhc k_\_jhaZiZ^ghc ZleZglbq_kdhc b
k_\_jh\hklhqghc ZleZglbq_kdhc beb _\jhi_ckdhc dhe_[ZeZkvhl^h^b\_j]_gpby\gmljbnhjf[ueZ
agZqbl_evghgb`_bkhklZ\beZ\kj_^g_ff_g__M
bkke_^h\ZgguogZfbihimeypbcO. mordax dentex\_ebqbgZ ^b\_j]_gpbb gmde_hlb^guo ihke_^h\Zl_evghkl_c
mqZkldZ 1'1' fl>GD [ueZ lZd`_ g_\ukhdhc b khklZ\beZebrvlZ[eijbwlhfmjh\_gv^b\_j]_gpbbf_`^mjh^hfOsmerusb^jm]bfbjh^Zfbk_f_ckl\Z 2VPHULGDH \ ± jZaZ ij_\ukbe mjh\_gv ^b\_j]_gpbb nhjf bkke_^h\Zgguo Lwcehjhf b >h^khghf
(Taylor, Dodson, 1994).
<gZklhys__\j_fyg_kms_kl\m_l_^bgh]hfg_gby
hlghkbl_evgh kbkl_fZlbq_kdh]h iheh`_gby dhjxrdh\uo ju[ ;_j] *RVOLQH DexdZgh\ 1HOVRQQ_j_rg_\J_r_lgbdh\b^j
N_^hjh\b^jb^jgh[hevrbgkl\hbkke_^h\Zl_e_c jZkkfZljb\Zxl bo \ dZq_kl\_ [eb`Zcrbo jh^kl\_ggbdh\ ehkhk_\uo < k\yab k wlbf bgl_j_kgh kjZ\gblv ihemq_ggu_ gZfb j_amevlZlu k ebl_jZlmjgufb
^Zggufbihk_f_ckl\m6DOPRQLGDHLZdijbbkke_^h\Zgbb baf_gqb\hklb fblhohg^jbZevgh]h ]_ghfZ m g_dhlhjuo \b^h\ jh^Z Oncorhynchus ih mqZkldm 1'1'
hij_^_e_gumjh\gb]Ziehlbibq_kdh]hbgmde_hlb^gh]h
jZaghh[jZaby ^ey ]hj[mrb O. gorbuscha – 0.6687 ±
±b^eyd_luO. keta – 0.4408 ± 0.08266
b ^ey g_jdb O. nerka – 0.6077 ± b
khhl\_lkl\_ggh&KXULNRYHWDO>eyk_\_jghcbx`ghcfZevfuSalvelinus malma wlbihdZaZl_ebkhklZ\beb±b±^eyk_\_jghc
b ± b ± ^ey x`ghc khhl\_lkl\_ggh He_cgbd b ^j LZdbf h[jZahf agZq_gby
]Ziehlbibq_kdh]h b gmde_hlb^gh]h jZaghh[jZaby ^ey
ehkhk_\uobdhjxrdh\uoju[e_`Zl\h^ghf^bZiZahg_
AgZq_gby^b\_j]_gpbbjZkkqblZggu_ ^ey ihimeypbcjZaguo\b^h\k_f_ckl\Z2VPHULGDHlZ[ekhihklZ\bfu k ebl_jZlmjgufb oZjZdl_jbamxsbfb \gmljb\b^h\mx baf_gqb\hklv fl>GD ehkhk_\uo ju[ Ih j_amevlZlZf5)/3ZgZebaZZfiebnbpbjh\ZgguomqZkldh\
;BHEH=BYFHJYlhf‹
]_gh\1'1'b1'1'mk_\_jhZf_jbdZgkdhcd_lu
mjh\_gv ^b\_j]_gpbb f_`^m ihimeypbyfb khklZ\be
± &URQLQ HW DO >b\_j]_gpby f_`^m
ihimeypbyfbk_\_jghcfZevfuSalvelinus malma malma,
jZkkqblZggZygZhkgh\Zgbbbaf_gqb\hklblj_omqZkldh\
fblhohg^jbZevgh]h ]_ghfZ \ khklZ\ dhlhjuo \oh^bl
j_]bhg dh^bjmxsbc 1'1' km[t_^bgbpu gZ^hdkb^^_]b^jh]_gZau khklZ\beZhdhehZf_`^mihimeypbyfb x`ghc fZevfu S. malma krascheninnikovi –
hl^hHe_cgbdb^jMjh\_gv
\gmljb\b^h\hc ]_g_lbq_kdhc ^b\_j]_gpbb m dhjxr_d
ih gZrbf ^Zgguf e_`bl \ ij_^_eZo hl (O. mordax dentex)^h(H. nipponensis).
< lh `_ \j_fy mjh\_gv ^b\_j]_gpbb f_`^m jZagufb \b^Zfb b jZagufb jh^Zfb dhjxr_d [ue agZqbl_evgh \ur_ b kms_kl\_ggh ij_\urZe lZdh\hc ^ey
ehkhk_\uo ju[ Ih ^Zgguf ;jmgg_jZ k khZ\lhjZfb
(BrunQHU HW DO ^b\_j]_gpby jZkFqblZggZy gZ
hkgh\_ k_d\_gbjh\Zgby mqZkldZ >i_leb fl>GD ^ey
Salvelinus alpinus complex b S. fontinalis khklZ\beZ
^eyS. alpinus complex bS. namaycush±Z
^ey S. alpinus complex b S. leucomaenis ± Mjh\_gvjZaebqbcf_`^mS. malma bS. leucomaenis ^hklb]Ze He_cgbd b ^j Ihemq_ggu_ gZfb
hp_gdb ^b\_j]_gpbb f_`^m jZagufb \b^Zfb jh^Z
Hypomesus e_`Zl\ij_^_eZohl^hZf_`^m
jh^Zfb±hl^h
Ba\_klgh qlh dhe_[Zgb_ mjh\gy f_`\b^h\hc ^b\_j]_gpbbih^Zggufj_kljbdpbhggh]hZgZebaZfl>GD
^ey fhjkdbo wklmZjguo b dZlZ^jhfguo \b^h\ ju[ e_`bl \ ij_^_eZo hl ^h Z ^ey ij_kgh\h^guo b
ZgZ^jhfguo\b^h\±hl^h%LOOLQJWRQ+HEHUW
IhdZaZgh lZd`_ qlh gbadbc mjh\_gv \gmljb\b^h\hc baf_gqb\hklb y\ey_lky lbibqguf ^ey \b^h\
ih^\_j]Z\rboky ^_ckl\byf he_^_g_gbc \ oh^_ w\hexpbb %LOOLQJWRQ +HEHUW :LOVRQ %HUQDWFKW] LZdbfh[jZahfgZrbj_amevlZlu\iheg_kh]eZkmxlkyk
ebl_jZlmjgufb^Zggufbbiha\heyxlk\ukhdbfmjh\g_f ^hklh\_jghklb b^_glbnbpbjh\Zlv \k_ bamq_ggu_
\b^u k_f_ckl\Z 2VPHULGDH. <i_j\u_ ihemq_gu dhebq_kl\_ggu_ hp_gdb ^b\_j]_gpbb fl>GD ^ey \k_o \b^h\
dhjxrdh\uo ju[ k_\_jhaZiZ^ghc qZklb Lboh]h hd_Z-
KDMJBOBG:B>J
294
gZG_khfg_gghqlhH. japonicus bH. nipponensis ±wlh
gZb[he__ [ebadh k\yaZggu_ \b^u jh^Z Hypomesus, Z
H. olidus agZqbl_evgh h[hkh[e_g hl gbo GZb[he__ ^b\_j]bjh\Z\rbcfblhohg^jbZevguc]_ghfkj_^b\k_obkke_^h\Zgguo\b^h\bf__lfhc\ZM. villosus catervarius.
:\lhju \ujZ`Zxl ]em[hdmx ijbagZl_evghklv
K< Njheh\m Bgklblml [bheh]bb fhjy ><H J:G b
B:Q_j_rg_\mBgklblml[bheh]bq_kdboijh[e_fK_\_jZ><HJ:GaZg_hp_gbfmxihfhsvhdZaZggmxijb
k[hj_ fZl_jbZeZ _]h hij_^_e_gbb b h[km`^_gbb ihemq_gguo j_amevlZlh\ Z lZd`_ FX AZkuidbgm Bgklblml [bheh]bq_kdbo ijh[e_f K_\_jZ ><H J:G
XB=Z\j_gdh\m BA IZjimjZ b :X K_f_gq_gdh
LBGJHp_gljaZihfhsvijbk[hj_fZl_jbZeZ
KIBKHDEBL?J:LMJU
;_j]EKJu[uij_kguo\h^KKKJbkhij_^_evguokljZgF
EBa^\h:GKKKJQk
=jbp_gdh HN Qmjbdh\ :: Kbkl_fZlbdZ fZehjhluo dhjxr_d jh^Z Hypomesus 6DOPRQLIRUPHV 2VPHULGDH ZabZlkdh]h ih[_j_`vy Lboh]h hd_ZgZ Ahhe `mjg L\uiK±
DexdZgh\ <: Gh\u_ ^Zggu_ h jZkijhkljZg_gbb fZehjhluo
dhjxr_d\\h^ZoKKKJ>hde:GKKKJL
‹K±
DexdZgh\ <: Fhjnheh]bq_kdb_ hkgh\u kbkl_fZlbdb dhjxr_d jh^Z Hypomesus 2VPHULGDH Ahhe `mjg L\uiK±
DexdZgh\<:Kbkl_fZlbdZbjh^kl\_ggu_hlghr_gbydhjxr_djh^h\Osmerus bHypomesus 2VPHULGDHbbojZkk_e_gb_Ahhe`mjgL\uiK±
DexdZgh\ <: Ijhbkoh`^_gb_ jZkk_e_gb_ b w\hexpby dhjxrdh\uo2VPHULGDHHkgh\udeZkkbnbdZpbbbnbeh]_gbb ehkhk_\b^guo ju[ E ABG :G KKKJ K 13–27.
He_cgbd:=KdmjbobgZE:;judh\<e:b^j>b\_j]_gpby
fblhohg^jbZevghc >GD ^\mo ih^\b^h\ fZevfu
Salvelinus malma 6DOPRQLGDH 3LVFHV >hde J:G
L‹K±
He_cgbd := KdmjbobgZ E: ;judh\ <e: >b\_j]_gpby
fZevfu Salvelinus malma \ ZabZlkdhc qZklb k_\_jghc
IZpbnbdb ih ^Zgguf 3&55)/3ZgZebaZ fblhohg^jbZevghc >GD =_g_lbdZ L ‹ K ±
1401.
J_r_lgbdh\XK;h]mpdZyG=<Zkbev_\Z?>b^jKibkhd
ju[hh[jZaguobju[ij_kguo\h^Jhkkbb<hijbolbheL‹K±
Khdheh\kdZyL=Khdheh\kdbc:KKh[he_\kdbc?BKibkhd
ju[ aZeb\Z I_ljZ <_ebdh]h Yihgkdh_ fhj_ <hij
bolbheL‹K±
Khdheh\kdbc :K Hdkxavyg ?; Mlhqg_gby b ^hiheg_gby d
kibkdm bolbhnZmgu [Zkk_cgZ j_db LmfZgghc Wdheh]bq_kdh_ khklhygb_ b [bhlZ x]haZiZ^ghc qZklb aZeb\Z
I_ljZ <_ebdh]h b mklvy j_db LmfZgghc <eZ^b\hklhd
>ZevgZmdZK±
N_^hjh\ << Q_j_rg_\ B: GZaZjdbg F< b ^j DZlZeh]
fhjkdbobij_kgh\h^guoju[k_\_jghcqZklbHohlkdh]h
fhjy<eZ^b\hklhd>ZevgZmdZk
Q_j_rg_\B::gghlbjh\Zgguckibkhdju[hh[jZaguobju[
ij_kguo \h^ :jdlbdb b khij_^_evguo l_jjblhjbc <hijbolbheL‹K 597–608.
Q_j_rg_\B:R_klZdh\:<Kdhi_pF;HjZkijhkljZg_gbbfZehjhluodhjxr_djh^ZHypomesus 2VPHULGDH\
k_\_jghc qZklb Hohlkdh]h fhjy <hij bolbhe L‹K±
Q_j_rg_\ B: R_klZdh\ :< Njheh\ K< D kbkl_fZlbd_
fZehjhluodhjxr_djh^ZHypomesus 2VPHULGDHaZeb\Z
I_ljZ <_ebdh]h Yihgkdh]h fhjy ;bhe fhjy L‹K±
Qmjbdh\ :: DZji_gdh <B Gh\u_ ^Zggu_ h jZkijhkljZg_gbb fhjkdhc fZehjhlhc dhjxrdb Hypomesus japonicus
%UHYRRUW \ \h^Zo KKKJ <hij bolbhe L ‹K±
Baby M.-C., Bernatchez L., Dodson J.J. Genetic structure and relationships among anadromous and landlocked populations
of rainbow smelt, Osmerus mordax, Mitchill, as revealed by
mtDNA restriction analysis // J. Fish Biol. 1991. Vol. 39.
Suppl. A. P. 61–68.
Bernatchez L., Martin S. Mitochondrial DNA diversity in anadromous rainbow smelt, Osmerus mordax Mitchill: a genetic assessment of the member-vagrant hypothesis // Can.
J. Fish. Aquat. Sci. 1996. Vol. 53, no. 2. P. 424–433.
Billington N., Hebert P.D.N. Mitochondrial DNA diversity in
fishes and its implications for introductions // Can. J. Fish.
Aquat. Sci. 1991. Vol. 48. Suppl. 1. P. 80–94.
Brown W.M., George M. Jr., Wilson A.C. Rapid evolution of animal mitochondrial DNA // Proc. Natl. Acad. Sci. USA.
1979. Vol. 76. P. 1967–1971.
Brunner P.C., Douglas M.R., Osinov A. et al. Holarctic phylogeography of arctic charr (Salvelinus alpinus L.) inferred
from mitochondrial DNA sequences // Evolution. 2001.
Vol. 55, no. 3. P. 573–586.
Churikov D., Matsuoka M., Luan X. et al. Assessment of concordance among genealogical reconstructios from various
mtDNA segments in three species of Pacific salmon (genus
Oncorhynchus) // Mol. Ecol. 2001. Vol. 10. P. 2329–2339.
Cronin M.A., Spearman W.J., Wilmot R.L. et al. Mitochondrial
DNA variation in chinook salmon (Oncorhynchus
tshawytscha) and chum salmon (O. keta) detected by restriction enzyme analysis of polymerase chain reaction
(PCR) products // Can. J. Fish. Aquat. Sci. 1993. Vol. 50,
no. 4. P. 708–715.
Dodson J.J., Carscadden J.E., Bernatchez L., Colombani F. Relationship between spawning mode and phylogeographic
structure in mitochondrial DNA of North Atlantic capelin
Mallotus villosus // Mar. Ecol. Prog. Ser. 1991. Vol. 76, no.
2. P. 103–113.
Gharrett A.J., Gray A.K., Brykov V.A. Mitochondrial DNA variation in Alaskan coho salmon, Oncorhynchus kisutch // Fish.
Bull. 2001. Vol. 99. P. 528–544.
Gosline W.A. Contributions toward a classification of modern isospondylous fishes // Bull. Brit. Mus. Zool. 1960. Vol. 6, no.
6. P. 327–365.
Hamada K. Revision of Hypomesus olidus (Pallas) and Hypomesus japonicus (Brevoort) of Hokkaido, Japan // Bull.
Fac.Fish. Hokkaido Univ. 1954. Vol. 4, no. 4. P. 256–261.
Hamada K. A new osmerid fish, Hypomesus sakhalinus new species, obtained from Lake Taraika, Sakhalin // Jap. J. Ichthyol. 1957. Vol. 5, no. 3–6. P. 136–140.
Hamada K. Taxonomic and ecological studies of the genus Hypomesus of Japan // Mem. Fac. Fish. Hokkaido Univ. 1961.
Vol. 9, no. 1. P. 1–55.
Johnson G.D., Patterson C. Relationships of lower euteleostean
fishes // Interrelationships of fishes. San Diego; London:
Acad. Press. 1996. XIII, 496 p.
McAllister D.E. A revision of the smelt family, Osmeridae // Bull.
Nat. Mus. Can. 1963. No. 191. Biol. Ser. No. 71. P. 1–53.
;BHEH=BYFHJYlhf‹
KJ:<GBL?EVGUC:G:EBA
McElroy D., Moran P., Bermingham E., Kornfield I. REAP: an
integrated environment for the manipulation and phylogenetic analysis of restriction data // J. Hered. 1992. Vol. 83.
P. 153–158.
Nei M., Miller J.C. A simple method for estimating average number of nucleotide substitutions within and between populations from restriction data // Genetics. 1990. Vol. 125.
P. 873–897.
Nei M., Tajima F. DNA polymorphism detectable by restriction
endonucleases // Genetics. 1981. Vol. 97. P. 145–163.
Nelson J.S. Fishes of the World. New York: John Wiley and Sons,
Inc. 1994. XVII, 600 p.
Rohlf F.J. NTSYS-pc: Numerical taxonomy and multivariate
analysis system. Version 1.60. New York: Exter Publ., Ltd.
1990.
Sambrook J., Fritsch E.F., Maniatis T. Molecular cloning. A laboratory manual. New York: Cold Spring Harbor Lab. Press.
1989. 1626 p.
Saruwatari T., Lopez J.A., Pietsch T.W. A revision of the osmerid
genus Hypomesus Gill (Teleostei: Salmoniformes), with the
description of a new species from the southern Kuril Islands // Species Div. 1997. Vol. 2. P. 59–82.
;BHEH=BYFHJYlhf‹
295
Stanley S.E., Moyle P.B., Shaffer H.B. Allozyme analysis of delta
smelt, Hypomesus transpacificus, and longfin smelt, Spirinchus thaleichthys, in the Sacramento-San-Joaquin estuary,
California // Copeia. 1995. Vol. 2. P. 390–396.
Taylor E.B., Bentzen P. Evidence for multiple origins and sympatric divergence of trophic ecotypes of smelt (Osmerus) in
northeastern North America // Evolution. 1993. Vol. 47,
no. 3. P. 813–832.
Taylor E.B., Dodson J.J. A molecular analysis of relationships
and biogeography within a species complex of holarctic
fish (genus Osmerus) // Mol. Ecol. 1994. Vol. 3, no. 3.
P. 235–248.
Trenham P.C., Shaffer H.B., Moyle P.B. Biochemical identification and assessment of population subdivision in morphologically similar native and invading smelt species (Hypomesus) in the Sacramento San Joaquin estuary, California // Trans. Amer. Fish. Soc. 1998. Vol. 127, no. 3.
P. 417–424.
Wilson C.C., Bernatchtz L. The ghost of hybrids past: fixation of
arctic charr (Salvelinus alpinus) mitochondrial DNA in an
introgressed population of lake trout (S. namaycush) // Mol.
Ecol. 1998. Vol. 7. P. 127–132.
;BHEH=BYFHJYlhf‹k289–295
290