Äîêóìåíò âçÿò èç êýøà ïîèñêîâîé ìàøèíû. Àäðåñ îðèãèíàëüíîãî äîêóìåíòà : http://www.sevin.ru/dissertations/ecology/29.pdf Äàòà èçìåíåíèÿ: Fri Apr 18 14:04:00 2008 Äàòà èíäåêñèðîâàíèÿ: Tue Oct 2 11:04:39 2012 Êîäèðîâêà:

. ..





Capra (Bovidae, Artiodactyla): -

03.00.08 ­



, 2008


. ..

:

, ,

:





20 2008 . 14.00, 002.213.01 . .. : 119071, , , . 33, /: (495) 952-35-84, e-mail: admin@sevin.ru.

: 119071 , , 33.

18 2008 .

,
1

..


. (Capra) ­ . , Capra 1 13. Capra : , , , , . (Linnaeus, 1766; Gray, 1852; Lydekker, 1913; Schwarz, 1935; Ellerman, Morrison-Scott, 1951; .., 1953, 1959; ., 1961; Haltenorth, 1963; . ., 1979; .). , . , . , Capra. : , () . . , , , . , . - , , . , , . - (, 2005; , 2005; Pidancier et al., 2006). , Capra . : Capra. : 1. Capra. 2. (Y-) . 3. : · - ; · ; · .

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. - Capra. , . GenBank SRY 5 Capra. b . . Capra , , , ­ . - Capra . , Capra, , , . Capra. «» C. sibirica, , , , . , . . 7 , : «FEBS Forum for Young Scientists» (Praga, 2004), « » (, 2004, 2006), « » (, 2005), « » (, 2007), « » (, 2007), «- » (, 2007). «» (, 2007) , , , - , 28.02.2008. . 11 ( 1 - , 6 - , 4 - ). . , 7 , , , . 156 , 35 20 . 217 . . ..., . . . . ... . . , ... . . ... . . , . ... . .
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, ... . . , , , . ; ... .. , ... .. , ... .. , ... .. , .. , ; «», ... .. , .. , .. , .. , ... .. , ... .. , ... .. , .. , ... .. , . , , , . « » « », 04-07-01240- 06-04-49575 4534.2006.4.

1. Capra
, , , . , , , .

2. , Capra,
Capra (, , - .) . , - . , ( ., 1961; . ., 1979; .), 8 : (C. pyrenaica), (C. ibex), (C. aegagrus), (C. nubiana), (C. caucasica), (C. cylindricornis), (C. sibirica) (C. falconeri). , - (C. hircus).

3.
Capra ( ) (). 373 (. 1). 80 , 30 , 42 - 8 .
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1. .
C. C. C. C. C. C. aegagrus sibirica ibex caucasica cylindricornis falconeri 73 117 20 74 59 30 0-1,5 (I ) 7 2 - 4 2 6 2-3,5 (II ) 7 2 - 4 4 3 6 6 - 15 10 3 4-4,5 , (III ) 8 5 1 1 3 3 7 7 1 7 5 1 5-7,5 (IV ) 18 31 3 9 10 5 14 41 4 19 14 5 8-10,5 (V ) 1 1 - 3 - 1 3 14 7 10 10 2 11-16 (VI ) 1 3 - - - - 1 5 4 3 1 1

STATISTICA 7.0, SPSS 13.0, NCSS SYSTAT 11. , . ( ) . , . , , . ( 5 ) ( ) , ( ). . . F ( , F- ). () . . : ( ) . . , , . (Sokal, Rohlf, 1981). , ­ , .. (, , 2006). , , . « » ( ., 1989), () (Shepard 1962, Davison, Jones, 1983) «»
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(, 2001; ., 2003). , 1, 2 ... Em., ­ 1, 2 ... Km. ( , rs), - ( ) (GLM), (, ) . , , «» , PLS (Haenlein, Kaplan, 2004). (UPGMA) (, , 1986; ., 1989) . , (« », ). , , ( «» ). «» . : 1 ( . ., 2004), k- (Hartigan, Wong, 1979), (Kaufman, Rousseeuw, 1990). . 2 (Sokal, Rohlf, 1981). - Capra 5 Capra ( ). b, SRY, Y-. - (Sambrook et al., 1989) «Diatom DNA Prep 200» («», ). D- C. aegagrus (39) ( , ­ S-48300) (, 2001) PCR Purification Kit (QIAGEN). 48 (. 2). b GLU R, ­ PRO DLC (Ozawa et al, 1997; ., 2001). SRY , : SRYL1 (5` ­ GCA TGT AGC TCC AGA ATA TTT CAC T ­ 3`), SRYH1 (5` ­ ATA AAT C(T/C)(G/A) T(G/A)A GGC AAA CTT GAA A ­ 3`), SRYL2 (5` ­ TGC TTC TGC TAT GTT CAG AGT ATT G ­ 3`) SRYH3 (5` ­ GCA ATT TAC AAA GAG GTG GAA AGT A ­ 3`). «», ABI PRISM 310 ABI PRISM® BigDyeTM Terminator v. 3.1. 42

1

Fracdim ver.1.9 ( ., , 2004) 6


b (. 2), 28 , 19 37 ­ SRY. 2. - *



, SRY b b Y- 392 .. 1128 .. 402 .. 1832 .. 16 9 9 8 6 - - - 13 (12) 8 (8) 10 (8) 7 (2) 10 (6) 6 2 7 12 (11) 8 (8) 8 (6) 13 (7) 3 6 1 6 1 (1) 9 (9) 5 (5) 4 (3) 1 (1) - - 1 14 (14) 7 (7) 8 (8) 3 (3) 5 (5) - - 1

C. sibirica C. caucasica C. cylindricornis C. aegagrus C. falconeri C. pyrenaica C. ibex C. hircus

* ,

, GenBank. (Pseudois nayaur), (Nemorhaedus goral) (Ovis ammon). Bioedit (Hall, 1999). , ( ), (Net distance) (Neighbor-Joining) MEGA 3.1 (Kimura, 1980; Saitou, Nei, 1987; Kumar et al., 2004). (Net distance) D = dXY ­ (dX - dY)/2, dXY ­ X Y, dX, dY ­ . (, Maximum Parsimony) PAUP 4.10 (Swofford, 2000), (Maximum Likelihood) ­ METAPIGA (Lemmon, Milinkovitch, 2002). (Felsenstein, 1985). 1000 . Network (Bandelt et al., 1999).

4.
( ) .
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, 20,4. (<0,01). 0-3,5 . . . 31 (. 1). : (68), (70), (75), (77), (76), (72), ­ (73). 29 , 5 - (68, 70, 75, 72, 73). , .

. 1. Capra
, ­ . F

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C. C. C. C. C. C.

aegagrus sibirica ibex caucasica cylindricornis falconeri

. 2. Capra
() () 4- . .

, . «» , Capra , , . , , . . , : C. falconeri, , (22), (18), (53) . . , , . ( 0,88, <0,01, n=15). , , (. 2). , « » . : (C. caucasica, C. cylindricornis) , C. sibirica, C. ibex C. aegagrus. C. caucasica C. cylindricornis, C. aegagrus C. sibirica, C. ibex C. sibirica. . ( , ) Capra (), , , . ( E1 K1)
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. , : K1 ­ 89%, E1 ­ 74%, E2 ­ 26% E4 ­ 16%. , - E1 (24.5%) K1(30%). «» «» . , , « » : 48% K2 8% E5 ( ­ 25%). , , . . 3 , ­ 4 . 34 94% . E1 K1. (. 3). I C. caucasica, C. cylindricornis, C. falconeri, II ­ C. aegagrus, C. sibirica, C. ibex. C. falconeri ( I). . (-, ) , . , C. sibirica . , , , (37.5), (22.7). , .


. 3. ( UPGMA, ­ ) () () Capra
.

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, , () . . . , , I II, - . sibirica. I C. caucasica + C. cylindricornis . C. caucasica + C. cylindricornis C. falconeri. «» C. caucasica C. falconeri, 9 C. caucasica C. cylindricornis. I C. falconeri .


. 4. () () . caucasica + . cylindricornis

. 5. , C. caucasica (1) C. cylindricornis (2)
C. caucasica () C. cylindricornis () : ., 1961 , , 1993

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C. caucasica + C. cylindricornis (. 4, 5). 10 (20%). (12 , 24%). , , , . II 4 . , C. aegagrus C. ibex, ­ C. sibirica. 13% , C. aegagrus C. ibex. . sibirica : (, B), (. 6, 7).


. 6. () () (A, B) . sibirica . 7. , (A, B) . sibirica
1 - . sibirica : Fedosenko, Blank, 2001

. Capra . 4 . 74% (30­94%). E1 K1. , , , . , . .
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, . I (. 3) C. aegagrus C. falconeri. . «» . II 90% C. sibirica, 1 C. ibex 3 - C. aegagrus. C. ibex, C. sibirica. , A B (. 6, 7). IIb (C. ibex + C. caucasica + C. cylindricornis) C. ibex. , ( 70%) , C. caucasica, C. cylindricornis. , C. caucasica - C. cylindricornis. (. 4, 5). 6 (27%), . (32% «» ). C. cylindricornis C. caucasica Capra. , . C. falconeri. C. aegagrus, C. caucasica C. cylindricornis . C. aegagrus ­ 19 62. C. cylindricornis , C. caucasica. , C. aegagrus . 30% , . ... Capra. . , Capra . . , . sibirica, «» «» . aegagrus «» C. falconeri ­ C. cylindricornis . sibirica C. caucasica.
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5.
b 1128 .. 227, 20,1%, 179 . ­ 53 (7, 5, 41) 4 (1, 0, 3). , b, , . 7 (. 8). , . . sibirica, . nubiana (C. ibex C. pyrenaica). , . , . C. aegagrus . Capra (. 3). . aegagrus C. aucasica, C. falconeri . aegagrus, . aegagrus C. cylindricornis, . aegagrus C. hircus, C. ibex C. pyrenaica. , . (, ) (0.037). 3. (Net distance) Capra, b (1128 ..) *
net 1 2 3 4 5 6 7 8 9 S.E. distance C. sibirica C. falconeri C. cylindricornis C. pyrenaica C. ibex C. nubiana C. aegagrus C. caucasica C. hircus * 1 ­ 0.0659 0.0610 0.0680 0.0668 0.0582 0.0568 0.0662 0.0669 2 0.0085 ­ 0.0264 0.0414 0.0447 0.0339 0.0171 0.0280 0.0289 3 0.0078 0.0053 ­ 0.0314 0.0366 0.0270 0.0066 0.0195 0.0271 4 0.0084 0.0068 0.0056 ­ 0.0181 0.0393 0.0268 0.0364 0.0385 5 0.0082 0.0073 0.0062 0.0043 ­ 0.0451 0.0304 0.0387 0.0421 6 0.0080 0.0062 0.0053 0.0065 0.0070 ­ 0.0256 0.0322 0.0413 7 0.0075 0.0037 0.0014 0.0047 0.0052 0.0047 ­ 0.0077 0.0115 8 0.0086 0.0053 0.004 0.0061 0.0063 0.0054 0.0015 ­ 0.0231 9 0.0087 0.0055 0.005 0.0065 0.0067 0.0065 0.0022 0.0044 ­

. C. sibirica (14, 20), C. caucasica AF034738 C. cylindricornis, C. cylindricornis 12 13 ­ C. caucasica

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. 8. Capra, b (1128 ..) (Neighbor-Joining)
­ -. , . ­ . 1­7 ­ Capra

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. 9. , b (1128 ..)
, ,

b, Capra (. 8, 9) ­ C. sibirica. , b , . (C. pyrenaica C. ibex), , , ­ . - , «C. ibex», - (Ellerman, Morrison-Scott, 1951) . (Haltenorth, 1963): , , . , C. caucasica C. cylindricornis. : -, . . C. aegagrus 4- (. 9). C. aegagrus C. aegagrus . , , . , , b. 402 165 ( 41%), 94 . ­ 37 6 .
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C. cylindricornis - 17 .., C. caucasica, . ( 1 ..), . aegagrus C. caucasica. : ( ), . , b, ( ). ... , Capra . , , . , .

6. Y-
SRY (727 1105 ..) (1832 ..). . 1832 26 (1,4%), 25 . 5 3 . Capra , (. 10, 11). : C. falconeri + C. aegagrus + C. hircus C. sibirica, C. caucasica C. cylindricornis. C. sibirica . C. sibirica, «14» «20», , . , : (C. caucasica) ( - -), (C. cylindricornis) 4 ( -, ). (. 10, 11, . 4): , ; . , . , (. 10) , , . , , .
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. 10. Capra, SRY Y (1832 . .) (NeighborJoining)
­ -. , . ­

. 11. , SRY (1832 ..)
. . 9

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4. (Net distance) Capra, SRY Y- (1832 ..) *
S.E.
1 2 3 4 5 6 7 1 2 3 4 5 6 7

net distance C. falconeri 0.0015 0.0020 - C. aegagrus 0.0033 0.0017 - C. sibirica 0.0077 0.0055 - C. sibirica 0.0088 0.0066 0.0033 C. caucasica 0.0078 0.0056 0.0033 C. cylindricornis 0.0083 0.0061 0.0039 C. hircus 0.0039 0.0005 0.0061 *

0.0023 0.0019 0.0014 - 0.0040 0.0050 0.0072

0.0020 0.0017 0.0013 0.0016 - 0.0019 0.0062

0.0022 0.0019 0.0014 0.0017 0.0010 - 0.0066

0.0014 0.0005 0.0018 0.0020 0.0018 0.0019 -



C. falconeri C. aegagrus. , , . C. nubiana C. sibirica, C. hircus C. aegagrus C. sibirica. , C. sibirica, C. caucasica, C. ibex, C. nubiana ( , , ), «C. ibex», Y-. C. pyrenaica C. cylindricornis , , «ibex» . C. falconeri C. aegagrus , .

7. Capra
Capra Capra . (Pidancier et al., 2006) . («C. sibirica» «C. aegagrus» ), «C. sibirica» «C. aegagrus». «C. aegagrus» «» C. caucasica, C. cylindricornis, C. nubiana, C. ibex C. pyrenaica. , . (Pidancier et al., 2006) , 12. Y- (. 10) C. sibirica (. 8, 9). «C. aegagrus» Y «C. sibirica» C. caucasica, C. cylindricornis, C. nubiana, C. ibex, C. pyrenaica C. aegagrus (. 8) C. sibirica Y- (. 10).
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. 12. Capra

(. 12) , C. sibirica: C. ibex - , C. nubiana - C. caucasica - . , «C. sibirica», (C. aegagus + C. falconeri) . C. falconeri Capra. , , . , Y-. C. falconeri , , , . C. sibiric , , 13 () . sibirica. .. (1935), , : . sibirica sibirica ( ) C. s. sakeen ( ). , , . . . (1950), , , . . (1959): . sibirica sibirica (, , ), . sibirica hagenbeckii ( , ), . sibirica alaiana ( -, ), . sibirica formosovi ( -), . sibirica dementievi ( , ). . . ( ., 1961) , , . . , . s. sibirica . s. alaiana . . sibirica,
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.. (1935). « » 43 , Y- «» (. 10), . C. sibiric (14 20, . 8, 9). C. sibiric SRY Y- , , . . b (. 9) , «» C. falconeri, C. aegagus, C. hircus. C. ibex C. pyrenaica . C. pyrenaica , . C. ibex «»: C. caucasica, C. nubiana , , . sibirica. , C. ibex C. pyrenaica , , . C. aegagrus , (. 2). Y- C. aegagrus , , C. hircus C. falconeri (. 11). , , , (. 9). . -, , , , C. aegagrus ( «» ). «» C. aegagrus C. caucasica, C. cylindricornis C. hircus. C. aegagrus ­ . C. aegarus , ( C. caucasica, C. cylindricornis C. hircus), . C. aegagrus , -, , . , - (, ) , , (, 1949, 1959; ., 1961; , 1978). C. aegagrus , . , , , , b C. aegagrus C. caucasica C. cylindricornis - - . b, C. aegagrus C. cylindricornis, C. caucasica.
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C. caucasica C. cylindricornis , C. caucasica C. cylindricornis . , , «» C. sibirica, C. nubiana C. ibex. , . , «» (Ibex cebennarum, I. priscus, I. cenomanus, Capra prisca) , (, 1935, 1948; , 1948; , 1959). (, 1959), « - ». ( 350 -) «» - ( 600 ). , «» C. caucasica, C. ibex C. pyrenaica (Rivals, 2006), , «C. caucasica» . , C. cylindricornis, C. caucasica ­ , C. ibex. , C. caucasica (C. ibex C. pyrenaica), C. sibirica C. nubiana. «C. caucasica» C. caucasica C. cylindricornis, : C. caucasica C. cylindricornis , , C. caucasica C. cylindricornis. - ( ., 1961; , , 1993; , 2005; .). , , : - C. cylindricornis Y- C. caucasica. , , . Capra Capra . , (C. falconeri, C. aegagrus ). , Capra (Net distance) b (. 3), : 1,8­10,4 - Procapra; 2­5,33 ­ Ovis; 1,76­9,03 ­ Gazella; 3,4-6,4 ­ Moschus; 0,41­0,62 ­ Cervus elaphus C. nippon; 4,12 - Capreolus capreolus C. pygargus (Arctander et al., 1996; Randi et al., 1996, 1998; Boyce et al., 1999; Nagata et al., 1999; Vernesi et al., 2002; Lei et al., 2003; Ludt et al., 2004; , 2005; Zhang, 2005; Kholodova et al., 2006; , 2007; Chao-Ting Xiao, 2007; .). SRY Capra , 1,06±0,27 1000
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.. (Meadows et al., 2004). Capra (. 11, . 4). (C. falconeri, C. aegagrus, . sibirica , . sibirica , C. nubiana, C. caucasica C. cylindricornis) . , Capra , ( , ), . - , , : C. falconeri, C. pirenaica, C. cylindricornis. , , , , , , , , . . sibirica sibirica . s. sakeen (, 1935), , (, 1979; , 2003) . , , Capra, , . - . , , , , , , , , , « » (, 1982). . , Capra , , , . , , . , Capra, . , Capra, . , «» , , ( , ) .

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: 1. Capra 4- . «» ­ . 2. b Capra ( 20,1 41% ). SRY, , (1,4% ), . 3. , Capra . 4. Capra . ­ , C. aegagus , . sibirica. 5. C. falconeri . , . - C. aegagrus. 6. Capra , . . C. caucasica C. cylindricornis . 7. . sibirica ( , ) . 8. Capra C. ibex C. pyrenaica .

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, : .. ( ..), .., .., 2007. Capra (Bovidae, Artiodactyla) // . . 43, 2. . 245­253. .. ( ..), 2004. Capra // . . . . . 20­ 21 . .: . .65­68. .. ( ..), . ., .., 2005. Capra // . . .-. . . . 188­191. .. ( ..), . ., .., .., 2005. Capra // . : . . . 80­84. .. ( ..), 2006. Capra (Artiodactyla, Bovidae) // . . . . . 5­6 . .: . . 116­123. .., 2007. Capra (Bovidae, Artiodactyla) // . . . . 26­30 . . .: . C. 86­95. .., .., .., 2007. (Capra caucasica C. cylindricornis) // . . . . 15­18 . . .: . . 124­127. : Kuznetsova M.V., Kazanskaya E.J. (Zvychaynaya E.Y), Danilkin A.A., 2004. Analysis of Capra samples from Caucasus region based on mitochondrial D-loop seguences // The FEBS Forum for Young Scientists. 12­15 july. V. 271, S. 1. P. 32. .. ( ..), 2007. (Capra caucasica, Bovidae, Artiodactyla) // . (VIII . -). .: . . 179. Zvychaynaya E.Y., 2007. Genetic differentiation of Capra aegagrus (Bovidae, Artiodactyla) // Biodiversity. Ecology. Adaptation. Evolution. Mater. III Intern. Young Scientists conf. 15­18 May, Odessa. P. 236. Zvychaynaya E.Y., 2007. Role of the ancient and recent interspecific hybridization in forming of the genus Capra (Bovidae, Artiodactyla) // 13th Annual European Meeting of PhD Students in Evolutionary Biology. 14­17 of August. Sweden. P. 26.

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