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здеÑÑ - ÐÑÑÐ½Ð°Ð»Ñ ÐÐÐ Ð ÐÐ
здеÑÑ - ÐÑÑÐ½Ð°Ð»Ñ ÐÐÐ Ð ÐÐ
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;BHEH=BYFHJYlhf‹k462–472<br />
M>D<br />
;BHEH=BYBG>BM:EVGH=HJ:A
FHJNHEH=BQKDBHKH;GGHKLB<br />
463<br />
J:GaZe
464<br />
DHJGBGDHDHJG<br />
JbkIj_^ahwZ Pugettia quadridens:±dhglmjul_eZ;±Zgl_ggmeZ±fZdkbeebi_^,,±Z[^hf_g<br />
gZfZdkbeebi_^Zowf[jbhgZevgZydmlbdmeZg_bah[jZ`_gZFZkrlZ[:±ff;±±ff<br />
fu_ djmigu_ iZjgu_ ojhfZlhnhju jZkiheh`_gu ih<br />
\_gljZevghfmdjZxdZjZiZdkZHlf_q_ghg_kdhevdh[he__<br />
f_edbo ojhfZlhnhjh\ \i_j_^b ^hjkZevgh]h rbiZ<br />
ijbf_jgh \ p_glj_ [hdh\hc ih\_joghklb dZjZiZdkZ<br />
[eb`_dhkgh\ZgbxZ[^hf_gZgZi_j_^g_cih\_joghklbf_`^m]eZaZfbbhkgh\Zgb_fjhkljZevgh]hrbiZb<br />
ih^]eZaZfbQ_ldh\b^_gg_iZjgucojhfZlhnhjgZi_-<br />
j_^g_c ih\_joghklb hkgh\Zgby ^hjkZevgh]h rbiZ \<br />
p_gljZevghcqZklbrbiZib]f_glZg_l<br />
:gl_ggmeZ jbk < H^gh\_l\bklZy khklhbl ba<br />
wdahih^blZ g_kms_]h l_jfbgZevguo wkl_lZkdZ <br />
^ebgguobdhjhldbc^ebggmxbdhjhldb_l_jfbgZevgu_s_lbgdbWg^hih^blhlkmlkl\m_l<br />
:gl_ggZ jbk = >\m\_l\bklZy Wdahih^bl k<br />
^\mfyihqlbjZ\gufbkmijhlb\gufbkm[l_jfbgZevgufbs_lbgdZfbIjhlhih^blgZdhgp_ibevqZlucwg^hih^bl^hklb]Z_l^ebguijhlhih^blZ<br />
FZg^b[meujbk:kbff_ljbqgu_Wdahih^bl<br />
kam[qZlufbfheyjgufbj_aph\ufhljhkldZfbWg^hih^bliZeviZhlkmlkl\m_l<br />
FZdkbeemeZ jbk @ GZ dhdkhih^bl_ i_jbkluo<br />
s_lbghd GZ [Zabih^bl_l_jfbgZevgu_ i_jbklham[qZlu_<br />
aZhklj_ggu_ bkm[l_jfbgZevgu_ i_jbklham[qZlu_<br />
s_lbgdb Wg^hih^bl qe_gbdh\uc gZ<br />
ijhdkbfZevghfqe_gbd_gZ^bklZevghf±i_jbklham[qZlu_s_lbgdb<br />
FZdkbeeZjbkADhdkhih^bl^\mehiZklgucgZ<br />
dZ`^hcehiZklbihs_lbgdb;Zabih^bllZd`_^\mehiZklguc<br />
bf__ls_lbgdb Wg^hih^bl gZ ehiZklb<br />
g_jZa^_ey_lkyg_k_l±s_lbgdb
FHJNHEH=BQKDBHKH;GGHKLB<br />
465<br />
JbkAhwZ, Pugettia quadridens.:±\b^k[hdm;±\b^ki_j_^b±djZcdZjZiZdkZ±fZg^b[meu@±<br />
fZdkbeemeZA±fZdkbeeZB±fZdkbeebi_^,D±fZdkbeebi_^,,E±Z[^hf_gFZkrlZ[:;±ffebgZfdf DZ`^Zy nmjdZ<br />
^ebghchdhehfdfihdjulZam[qbdZfbbf__leZl_jZevgucrbi^ebgZdhlhjh]hhdhehfdfiZju<br />
s_lbghdjZkiheh`_guihaZ^g_fmdjZxl_evkhgZ<br />
AhwZ,,<br />
DZjZiZdk jbk :±< =eZaZ gZ g_[hevrbo kl_-<br />
[_evdZo GZ ih\_joghklb dZjZiZdkZ iZj s_lbghd<br />
3Zgl_jh^hjkZevgu_ihkl_jh^hjkZevgZybmhkgh\Zgby^hjkZevgh]hrbiZdZ`^uc\_gljZevgucdjZcg_k_l<br />
djmigmx b[he__ f_edbo s_lbghd JZkiheh`_gb_<br />
ojhfZlhnhjh\g_baf_gy_lky<br />
:gl_ggmeZjbk=Wdahih^blg_k_l^ebgguob<br />
dhjhldbcl_jfbgZevgucwkl_lZkdbfZe_gvdmxs_-<br />
lbgdmIhy\ey_lkyihqdZwg^hih^blZ<br />
:gl_ggZjbk>Wdahih^blgZdhgp_aZam[j_g<br />
Wg^hih^bl^hklb]Z_l^ebguijhlhih^blZ<br />
FZg^b[meujbkIhy\ey_lkyihqdZwg^hih^blZiZeviu<br />
FZdkbeemeZ jbk @ Dhdkhih^bl b wg^hih^bl<br />
dZd m ahwZ,GZ [Zabih^bl_s_lbghdl_jfbgZevguoi_jbklham[qZluoaZhklj_gguokm[l_jfbgZevgu_<br />
i_jbklham[qZlu_ beZl_jZevgu_ i_jbklu_ GZ wdahih^bl_i_jbklZys_lbgdZ<br />
FZdkbeeZjbkADhdkhih^blbwg^hih^bldZdm<br />
ahwZ,;Zabih^bl ^\mehiZklguc bf__li_jbklham[qZluos_lbghdIhdjZxkdZnh]gZlblZi_jbkluo<br />
s_lbghd lhekluc ^bklZevguc hljhklhd j_^mpbjm_lky<br />
^hjZaf_jh\s_lbgdb<br />
FZdkbeebi_^,jbkB>bklZevguck_]f_glwdahih^blZk<br />
6ieZ\Zl_evgufbs_lbgdZfb<br />
FZdkbeebi_^,,jbkD>bklZevguck_]f_glwdahih^blZk<br />
6ieZ\Zl_evgufbs_lbgdZfb<br />
FZdkbeebi_^,,,;_abaf_g_gbc<br />
I_j_hih^u;_abaf_g_gbc<br />
:[^hf_gjbkEKhklhblbakhfblh\ckhfblkiZjhceZl_jZevguo\ujhklh\±ckhfblukiZjhc<br />
^hjkhf_^bZevguo s_lbghd gZ ±f khfblZo khojZgyxlkydhjhldb_ihkl_jheZl_jZevgu_rbiugZ±f<br />
khfblZo \b^gu ihkl_jh^hjkZevgu_ s_lbgdbGZ ±f<br />
khfblZo ohjhrh aZf_lgu ie_hih^u hgbaZ bkdexq_-<br />
gb_fie_hih^Z]hkhfblZbf_xlihqdmwg^hih^blZ<br />
L_evkhgjbkE;_abaf_g_gbc<br />
;BHEH=BYFHJYlhf‹
466<br />
DHJGBGDHDHJG<br />
JbkAhwZ,, Pugettia quadridens.:±\b^k[hdm;±\b^ki_j_^b±Zgl_ggZ±fZg^b[meu@±<br />
fZdkbeemeZA±fZdkbeeZB±fZdkbeebi_^,D±fZdkbeebi_^,,E±Z[^hf_gFZkrlZ[:;±ff
FHJNHEH=BQKDBHKH;GGHKLB<br />
467<br />
JbkF_]ZehiZPugettia quadridens.:±\g_rgbc\b^;±Zgl_ggmeZ±fZdkbeemeZ±fZdkbeeZ<br />
FZkrlZ[:±ff;±±ff<br />
jhgu gZf qe_gbd_km[l_jfbgZevguo bdjZcg_<br />
j_^dhl_jfbgZevguc wkl_lZkd b ± am[pZ Wg^hih-<br />
^bl qe_gbdh\uc gZ ^bklZevghf qe_gbd_ l_jfb-<br />
gZevgu_bkm[l_jfbgZevgZys_lbgdZ;ZaZevguckl_-<br />
[_e_dqe_gbdh\uc k ^\mfy fZe_gvdbfb s_lbgdZfb<br />
gZfbfqe_gbdZo<br />
:gl_ggZjbk Wg^hih^bl g_k_]f_glbjh-<br />
\Zgguc bf__ll_jfbgZevgu_ bf_^bZevgmx i_jbklham[qZlmx<br />
s_lbgdm GZ [Zabih^bl_l_jfbgZevguo<br />
i_jbklham[qZluo aZhklj_gguo km[l_jfbgZevguo i_-<br />
jbklham[qZluos_lbghdb±eZl_jZevgu_ i_jbklham[-<br />
qZlu_ s_lbgdbDhdkhih^blkl_jfbgZevgufbkm[-<br />
l_jfbgZevgufb b eZl_jZevgufb i_jbklham[qZlufb<br />
s_lbgdZfb<br />
FZdkbeeZjbkWg^hih^blh[uqgh]heucgh<br />
bgh]^ZgZg_fkhojZgyxlky±s_lbgdbgZ[Zabih^bl_<br />
±gZdhdkhih^bl_±i_jbklham[qZluos_lbghd<br />
M ebqbghd ihemq_gguo hl jZaguo kZfhd dhebq_kl\h<br />
i_jbkluos_lbghdihdjZxkdZnh]gZlblZ[uehjZaebqgufmebqbghdhl^\mokZfhd±±hlh^ghc±±<br />
32. Djhf_lh]hhlf_q_guiZjZs_lbghdjZkiheh`_gguo<br />
;BHEH=BYFHJYlhf‹
468<br />
DHJGBGDHDHJG<br />
JbkFZdkbeebi_^uf_]Zehiu Pugettia quadridens.:±fZdkbeebi_^,;±fZdkbeebi_^,,Zdlbemk<br />
i_j_hih^h\ gZ \gmlj_gg_c klhjhg_ g_k_libevqZluo<br />
rbiZ dh]hlv bah]gmluc ke_]dZ ibevqZluc
FHJNHEH=BQKDBHKH;GGHKLB<br />
469<br />
Ie_hih^u jbk ,±9 Ijbkmlkl\mxl gZ ±f<br />
khfblZo Z[^hf_gZ Wdahih^bl ie_hih^h\]h khfb-<br />
lh\Z[^hf_gZkb^ebggufbi_jbklufbieZ-<br />
\Zl_evgufb s_lbgdZfb l_jfbgZevgu_ s_lbgdb ^ebgg__q_fkm[l_jfbgZevgu_gZwg^hih^bl_ih^\ZdjxqdZ<br />
Ie_hih^]h khfblZ Z[^hf_gZ kieZ\Zl_evgufb<br />
s_lbgdZfbgZwdahih^bl_wg^hih^blZg_l<br />
K_ahggZy^bgZfbdZqbke_gghklbebqbghd<br />
\ieZgdlhg_<br />
470<br />
DHJGBGDHDHJG<br />
Jbk:[^hf_gbie_hih^u I–V f_]ZehiuPugettia quadridens. FZkrlZ[ff<br />
ihdZaZeh qlh m ihke_^g_]h gZ wg^hih^bl_ fZdkbeeu \<br />
jy^_kemqZ_\khojZgyxlky±s_lbgdbdZdbmP. marissinica\lh\j_fydZdmP.<br />
intermediabP. quadridens<br />
badhj_ckdbo\h^wlbs_lbgdbiheghklvxj_^mpbjmxlkyHimr_gghklvfZdkbeeumP.<br />
quadridensbaaZe
FHJNHEH=BQKDBHKH;GGHKLB<br />
471<br />
LZ[ebpZFhjnheh]bq_kdb_hkh[_gghklbebqbghdjZaguo\b^h\jh^ZPugettia<br />
DZjZiZdk<br />
bdhg_qghklb<br />
DZjZiZdk<br />
:gl_ggmeZ<br />
wdahih^bl<br />
:gl_ggZ<br />
wg^hih^bl<br />
:gl_ggmeZ<br />
wdahih^bl<br />
Pugettia quadridens gZrb<br />
^Zggu_)<br />
P. quadridens (ih: Ko,<br />
Hwang, 1997; Ko, 1998)<br />
AhwZI<br />
klvojhfZlhnhjmhkgh- klvojhfZlhnhjumhkgh-<br />
\Zgby^hjkZevgh]hrbiZ \Zgbyb\p_gljZevghc<br />
ojhfZlhnhj\p_gljZevghc qZklb^hjkZevgh]hrbiZ<br />
qZklb^hjkZevgh]hrbiZ<br />
hlkmlkl\m_l<br />
Hdheh^ebguijhlhih-<br />
^blZ<br />
F_]ZehiZ<br />
^ebgguodhjhldbo<br />
wkl_lZkdZ^ebggZy<br />
dhjhldZyl_jfbgZevgZy<br />
s_lbgdZ<br />
Hdheh^ebguijhlhih^blZ<br />
qe_gbdh\ucgZfqe_-<br />
gbd_wkl_lZkdh\bs_-<br />
lbgdZgZfqe_gbd_ + 1<br />
wkl_lZkdZb±l_jfbgZevguoam[pZ<br />
qe_gbdh\ucgZfqe_-<br />
gbd_wkl_lZkdh\bs_-<br />
lbgdZgZfqe_gbd_<br />
wkl_lZkdZgZfqe_gbd_<br />
l_jfbgZevgucwkl_lZkd<br />
P. marissinica (ih: Ko,<br />
Hwang, 1997; Ko, 1998)<br />
klvojhfZlhnhjmhkgh-<br />
\Zgby^hjkZevgh]hrbiZ<br />
ojhfZlhnhj\p_gljZevghcqZklb^hjkZevgh]h<br />
rbiZhlkmlkl\m_l<br />
^ebgguodhjhldbcwkl_lZkd^ebggZydhjhldb_l_jfbgZevgu_s_-<br />
lbgdb<br />
Hdheh^ebguijhlhih-<br />
^blZ<br />
^ebgguodhjhldbcwkl_lZkd^ebggZydhjhldb_l_jfbgZevgu_s_-<br />
lbgdb<br />
^ebgguodhjhldbcwkl_lZkd^ebggZydhjhldZyl_jfbgZevgZys_-<br />
lbgdZ<br />
Hdheh^ebguijhlhih-<br />
^blZ<br />
qe_gbdh\ucgZfqe_-<br />
gbd_wkl_lZkdh\bs_-<br />
lbgdZgZfqe_gbd_<br />
km[l_jfbgZevguobl_jfbgZevgucwkl_lZkd<br />
P. intermedia (ih: Ko,<br />
Hwang, 1997; Ko, 1998)<br />
OjhfZlhnhjumhkgh\Zgby<br />
b\p_gljZevghcqZklb<br />
^hjkZevgh]hrbiZhlkmlkl\mxl<br />
qe_gbdh\ucgZf<br />
qe_gbd_wkl_lZkdh\b1<br />
s_lbgdZgZfqe_gbd_<br />
wkl_lZkdZgZfqe_gbd_<br />
l_jfbgZevgucwkl_lZkd<br />
:gl_ggZ<br />
wdahih^bl<br />
bl_jfbgZevgu_s_lbgdb<br />
bl_jfbgZevgu_s_lbgdb<br />
bl_jfbgZevgu_s_lbgdb<br />
bl_jfbgZevgu_s_lbgdb<br />
FZdkbeemeZ Wg^hih^bl[Zabih-<br />
^bls_lbghd<br />
Wg^hih^bl[Zabih-<br />
^bls_lbghd<br />
Wg^hih^bl[Zabih-<br />
^bls_lbghd<br />
Wg^hih^bl[Zabih-<br />
^bls_lbghd<br />
FZdkbeeZ Wg^hih^bl±[Zabih^bl<br />
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^bldhdkhih^blkdZnh]gZlblbs_lbgdb<br />
gZih\_joghklb<br />
Wg^hih^bl[Zabih^bl<br />
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gZlblbs_lbgdbgZ<br />
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kdZnh]gZlblbs_-<br />
lbgdbgZih\_joghklb<br />
\_joghklb<br />
FZdkbeebi_^I Wg^hih^bl]heucbeb Wg^hih^bl]heuc[Zabih- Wg^hih^blkm[l_jfbgZevgZyl_jfbgZevgu_ih^bldhdkhih^bl<br />
Wg^hih^bl]heuc[Zab-<br />
km[l_jfbgZevgZys_lbgdZ ^bldhdkhih^blwib-<br />
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dhdkhih^bl±wibih^bl±^bklZevguc<br />
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lbgdb<br />
gbdwdahih^blZs_lbghd wibih^bl^bklZevguc<br />
qe_gbdwdahih^blZs_-<br />
lbghd<br />
qe_gbdwdahih^blZ<br />
5s_lbghd<br />
FZdkbeebi_^,,<br />
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lbghd^bklZevgucqe_gbd<br />
wdahih^blZs_lbgdb<br />
FZdkbeebi_^III Wg^hih^bl±±±<br />
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wdahih^blZs_lbgdb<br />
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wibih^bls_lbghd<br />
^bklZevgucqe_gbdwdahih^blZs_lbghd<br />
Wg^hih^blbs_-<br />
lbghd^bklZevgucqe_gbd<br />
wdahih^blZs_lbghd<br />
Wg^hih^blb<br />
s_lbgdbdhdkh[Zabih^bl<br />
wibih^bls_lbghd<br />
^bklZevgucqe_gbdwdahih^blZs_lbghd<br />
Wg^hih^blbs_-<br />
lbghd^bklZevgucqe_gbd<br />
wdahih^blZs_lbgdb<br />
Wg^hih^blb<br />
s_lbgdbdhdkh[Zabih^bl<br />
wibih^bls_lbghd<br />
^bklZevgucqe_gbdwdahih^blZs_lbghd<br />
kdZnh]gZlblZmf_]Zehiihemq_gguohljZaguokZfhd<br />
\_jhylgh hljZ`Z_l \gmljb\b^h\u_ jZaebqby ihlhfm<br />
wlhl ijbagZd g_ fh`_l y\eylvky ^hklZlhqguf hkgh\Zgb_f^eyhij_^_e_gby\b^h\<br />
Ihke_ f_lZfhjnhaZ \ f_]Zehim wg^hih^bl fZdkbeebi_^Z,klZgh\blkyg_k_]f_glbjh\Zggufijbwlhf<br />
mP. intermediabP. quadridensbadhj_ckdbo\h^s_lbgdbgZwg^hih^bl_j_^mpbjmxlkyiheghklvxZmP.<br />
marissinica<br />
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s_lbgdb .R +ZDQJ M [hevrbgkl\Z<br />
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wibih^bl_\gZr_fwdki_jbf_gl_kh\iZ^Z_lklZdh\uf<br />
mP. marissinica\lh\j_fydZdmP. quadridensba<br />
Dhj_b bo \k_]hDhebq_kl\h s_lbghd gZ h[t_^bg_gghf<br />
dhdkh[Zabih^bl_ fZdkbeebi_^Z ,,, m f_]Zehiu<br />
hibkZgghcgZfbf_gvr_q_f[uehihdZaZghmf_]Zehi<br />
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f_]ZehiuP. quadridensbaDhj_b<br />
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472<br />
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h^bgZdh\h_bjZ\ghlj_fIe_hih^uf_]ZehiuP. quadridens<br />
gZfb hibkZgu \i_j\u_ \ ebl_jZlmj_ _klv ebrv<br />
mihfbgZgb_hie_hih^_]hkhfblZgZwdahih^bl_dhlhjh]hbf__lkys_lbghd.XUDWD<br />
LZdbfh[jZahfijbg_khfg_gghfh[s_fkoh^kl\_<br />
m ebqbghd P. quadridens ba jhkkbckdbo \h^ h[gZjm`_g<br />
jy^fhjnheh]bq_kdbohkh[_gghkl_chlebqZxsbobohl<br />
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G_dhlhju_ ijbagZdb \ kljh_gbb f_]Zehiu k[eb`Zxl<br />
P. quadridens kjh^kl\_ggufb\b^Zfbjh^ZPugettia.<br />
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fZj_gdhaZihfhsvijbk[hj_`b\hlguobdmevlb\bjh-<br />
\ZgbbebqbghdZlZd`_E:Rdhe^bghcaZij_^hklZ\-<br />
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