-

- -
Zygophyllum pinnatum Cham. -
Zygophyllaceae. , -
( ., 1987.). --
- ,
(), , (- ). -
( ..., 2011),
( ..., 2019) ( …, 2017b) , -
( …, 2017a), ( …,
2016). 2 - , .
.
, 4 : . . , -
-; . . , -; . . . , -
-; -, . . ( -,
.) (UFA, 2020).
, , -
-
, -
. -
,
( ., 2013).
.. , .. .. (2017)
Z. pinnatum,
-
, .
(- -, -
.).
, .. , .. , .
.. (- - ) -
16 Z. pinnatum
( -
) - ( ), -
2015-2019 .
, ,
3
. -
, ..,
, 16 (3, ),
, -
. -
(. 1)
,
, %
j im v g1 g2 g3 ss s ω I I
2,9 0,0 2,8 2,8 18,1 9,7 38,9 12,5 15,3 0,0 0,47 0,71 0,39 0,28
-
⁄ , σ
2 - ,
- ( -
11 ). ,
. IOd < 1 -
(); IOd > 1 – (-
, ), IOd = 1 (-
1969; , 2013). , -,
, .. , ,
-, , -
(..., 2014). -
Z. pinnatum
(Illian et al, 2008; ., 2009;
Gelfand et al, 2010; , , 2012; ..., 2014; Wiegand, Moloney,
2014; Baddeley et al, 2015). -

(-
., 2010a, 2010b, 2014; , 2013, 2015, 2016, 2018; , -
, 2016, 2019a, 2019b; ., 2017; , , 2017;
., 2017; , , 2020). -
-
, -
, ,

( ., 2010a, 2010b,
2014; , 2016, 2018).
,
,
. -
. , -
4
, -
. -

. -
-
(Gelfand et al, 2010; , , 2014; Wiegand, Moloney, 2014;
Baddeley et al, 2015). -
.
.. .. (2017)
Artemisia solsoloides,
(-
) k (-
, 2006; ., 2006; , 2012).
– -
-
Z. pinnatum.
1.
Z. pinnatum ( ).
Z. pinnatum -
k.
(- ).
4. -
.
.
-
, -
2018 .
Zygophyllum pinnatum Cham. (. 1) – -
.
, , 5-20 ,

. 1-2 , , -
-, , , -
, . 3-4 , -
, 5-12 , 3-6 , , -
, , .
, 2-3 7 , -
, . 6 , 3-5 ,
5
, . 1,5 ,
, , . –
, -, , , 12 -
, . -. -
. . , , . -
- -
, - ,
. ( ..., 2011; ..., 2020)
- 2020 . -
Zygophyllum pinnatum - . -
, . , .
( -, ). – 52,48725 .., 55,85400
..
- .
)
)
) , )

- 40°.
(. 2) ( - 80%, -
50 ) -
( - 25-30%, 35 ), -
.
, .
, : Eu-
phorbia seguieriana Neck., Agropyron pectinatum (M.
Bieb.) P. Beauv., Scabiosa isetensis L., Ar-
temisia marschalliana Spreng., A. lercheana Weber ex Stechm., -
A. salsoloides Willd., Psephellus car-
bonatus (Klokov) Greuter, Gypsophila altissima L.,
G. rupestris A.Kuprian., Ephedra distachya L.
, -
( 2.):
6
1 , -
, – , 30%;
2 , -
,
, 15%;
3 – - , - ,
– , 50%,
.
1) , 2) ,
3) 4) ( Z. pinnatum );
.
, , , -
, ,
. -
55 .
,
-.
()
, ,
. -
– (pre), (gene)
(post). ,
7
.
-
Zygophyllum pinnatum (spa-
tial point patterns), spatstat ver. 1.64-1 12.05.2020
(Baddeley et al, 2015) R (R Core Team, 2020).
(density) (moving
window) (Bailey, Gatrell, 1995) (kernel function)
(Silverman, 1986; Scott, 1992). -
-
(Illian et al, 2008; Wiegand, Moloney, 2014; Baddeley et
al, 2015), ..
⁄ , pre – ,
- , .
-
""
(weighted density) (Baddeley et al, 2015), .. -
.
() , -
-
. -
,
,
– , -
, ,
(, 1965). – -
( , ),

, d –
∑
– k,
(, 2006; ., 2006; , 2012). -
, k -
-

.
K(r) (Ripley, 1976, 1977).
L(r)-r, ( ) √ ( ) ⁄ (Besag, 1977). -
L(r) LCSR(r),
LCSR(r) – (complete spatial
randomness, CSR). L(r) LCSR(r)
- (Besag, 1977) 299 CSR.
8
( 199 , r -
p-value = 2/(199+1) = 0.01. , -
(-
..., 2014) ).
– -
, ,
adult (.. adult=gene+post). ,
-
- ,
. 2
, .. -
.
. 2 ,
, -
. -
, 5-6 ,
– 10,
3, 25 1,2,3 .
, -
, 2 .
.
2 0,36 ./ 2 , 2 , 1,
0,64 ./ 2 . 3, ,
2 , 1, 1,36 ./ 2 . -

.
2.
(.), (./ 2 ).
S,

pre gene post pre gene post min max mean
1
25,0
16 10 5 1 0,64 0,40 0,20 0,04 0,30 1,20 0,58
2 9 3 6 0 0,36 0,12 0,24 0,00 0,19 0,53 0,34
3 34 25 6 3 1,36 1,00 0,24 0,12 0,00 5,30 1,39
-
128128 "" (pixel),
0,0390,039 ( 16384 ). . 2
(min, max) (mean) -
. (: -
, -
9
– -
. .)
, -
. 3.
1
L (r
– ( -
,
pre, gene, post); – ;
– ( ,
LCSR(r)–r, - ).
1 0,30 1,20
./ 2 , 0,58 ./
2 .
, -
– 0,30 0,90 ./ 2 , -
. 2,
0,19 0,53 ./ 2 ,
0,34 ./ 2 , 0,22-0,48 ./
2 .
3
0,00 5,30 ./ 2 . , -
10
1,0 ./ 2 ,
1,39 ./ 2 .
, 4,0 -
. -
3
. -
(CSR) " " (...,
2014),
.
(. 3),
Z. pinnatum 1-2 -
0,2 . (, 1969; -
, 1976; , 1994),
. -
3 CSR 0,3-
0,7, .. .
-
-
, - (-
, 1969). 1-2
-
.

, ,
3 . -

– 3 -
.
, LCSR,
(. 4). ,
,
, , ,
-
. (: -
3
, -
. , CSR
, -
. Z. pinnatum
, ..
.)
,
r <0,1-0,2 , -
.
11
r,
. 4. 3 : – .
. 3, ,
(min, man) (mean) .
(S), -
k,
. . 5.
3.
k pre gene post
min max mean min max mean min max mean min max mean
1 5,0 12,0 7,5 12,0 20,0 16,0 14,0 14,0 14,0 0,010 0,007 0,003 0,013 0,006
2 9,0 13,0 11,0 16,0 41,0 32,0 — — — 0,063 0,023 0,008 0,047 0,021
3 1,0 13,0 6,0 10,0 23,0 15,7 13,0 18,0 14,7 0,008 0,010 0,000 0,061 0,012
1 2 3
. 5. :
k,
, ( . 3).

1,0 13,0 . 2 ,
3 . -
10,0 41,0 . 2
2 1 3, -
. -
12
. , 2
, 1 3. k, -
3
.
, -
–
(pre) (adult=gene+post), 2
. . 6. - 1 3.
-
. 1 -
, 3 0,4 - 0,7 . -
, -
. -
, -
, ..
.
. ,

. (: ,
, -
1 3
)- r
)- r
13
. ,
3 ,
(. . 4),
.)
pre
adult
,
(ppre>0,5),
.
1,
.
2, . ,
, .
, –
,
. , k. -
- , -
( ).
3, , -
.
, -
. , -
, -
k. ,
. -
"" , -
.
.

, -
. , -
, , .
. -
. -
.
1. -
Zygophyllum pinnatum
(CSR). ,
, , Z. pinnatum
. ,
, .
0,00 5,30 ./ 2 , -
, -
.
k 0,007-0,023. k
, -
.
3. . -

, ,
. -
, .. -
.
4. ,
. -
-
– -
, .
.
(
), Zygophyllum pinnatum -
. : -
, ,
; -
; ,
Z. pinnatum ,
; ; -
. -
, -
.
-
-
. -
–
, -
15
. -
. . , ,
Koeleria sclerophylla P. Smirn.,
Tulipa patens Agardh ex Schult. & Schult. fil., -
Bupleurum multinerve DC., Hedysarum
grandiflorum Pall., Hedysarum razoumovianum Fisch. et
Helm, Iris pumila L., Artemisia
salsoloides Willd., Gypsophila rupestris A. Kuprian.,
Ephedra distachya L. -
-
– -
, .
-
(..., . . . -
, - - )
, -
(..., ..., . , ) -
, (...,
., . ),
(..., ., . ),
(..-. ., ..., . .. ), -
, -
R,
.
(Trollius europaeus L.). . . . .
. , 2006. – 19 .
.. . .: , 1969. – 273 .
.., .., .., .. -
// -
1 II -
. -: . -, 2006. – . 248-249.
.. -
// .
2012. . 14, 1(6) – . 1462-1465
.. -
// . . . . . -, 1994. – 90 .
.., .. -
(Halocnemum
strobilaceum (Pall.) Bieb.) «» // -
.. .
. 25. 2020. – . 321-329.
16
(Artemisia salsoloides Willd., Asteraceae) //
, 2017. – . 6. 4 (21). – . 41-47.
.., .. -
(Stipa korshinskyi Roshev.,
Poaceae)// . 2019a. – . 8, 1 (26) – .26-30
.., .. -
(Hedysarum grandiflorum Pall., Fabaceae) -
. :
. 2019b. – . 28, 1. – . 55-62.
.., .., .. :
. : ,
2013. – 439 .
( )// , 2017, 23, 1
(70). – . 51-59
: 2 . . 1:
/ . . .. ,
.. . – , 2016. – 292 .
: -
, . , 2019. - 488 .
: / . . . . – -
, 2017a. – 267 .
: 2 . . 1 : /
. -. . , . . . . - 2- ., . . –
: , 2011. – 384 .
: , , / . .
.. . – .:, 2017b. – 504 .
.., .., ..
. – .:, 1987. – 204 .
.. . – : - , 1976. –
155.
: http://www.plantarium.ru, . – . . (30.10.2020)
.., ..
-
// : ,
: III -
, 85- -
. «
», , 21-26 2017 . - : : -
, 2017. – . 189-192.
17
«» // -
: XII .
(1939-2016). .
.., .., .. – -:
«», 2017. – . 189-192.
( R): / .:
.. , .. , .. , .. . – :
- . -, 2014. – 146 .
.. // . 1965. . 1. –
. 251-254.
Orchis militaris L. (Orchidaceae Juss.) // . . .
2013. – . 15, 3(1). – . 352-357.
.. - -
// : -
- (1-5 2015 . , -
) / . . .. . – : .. , 2015. – . 244-
249
// -
, 5, 2016. – . 761-773.
.. -
Neottianthe ucullata (L.)
Schlechter // , 16 (46), 2018. – . 75-85
.., .. -
(Vaccinium myrtillus L.) // -
: : . .
. . , 2009. – . 307-312.
.., .. -
// . .48, . 4, 2012.
– . 597-613
.., .., .., .., ..
-
// . 4, 2009. – . 249-257.
.., .., ..

Cypripedium calceolus L. // . -. :
. . 2010a. . 152, . 3. – . 159-173.
18
- -
// . -
: , , (V ) /
. .-. .. . .. . – :
, 2010b. – .195-201.
Vaccinium myrtillus (Ericaceae)
( ) // . .50, .
3, 2014. – . 376-397.
R. – : -
, 2014. – 314 .
UFA, , : 454054, . , . -
, 71, . – -
2020 .
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Series B, 1977. – V. 39. – P. 193-195.
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19
1 2 3
r1, r2, Age d, r1, r2, Age d, r1, r2, Age d,
507 113 pre 7 164 518 gene 40 212 528 gene 15
478 180 gene 20 104 440 gene 36 204 485 pre 5
310 250 pre 8 112 400 gene 41 236 464 pre 8
19 482 pre 7 206 368 pre 13 343 307 pre 13
56 457 pre 9 253 459 gene 38 340 281 pre 10
55 466 pre 9 415 103 gene 16 310 261 pre 5
175 460 pre 12 435 387 gene 23 337 222 pre 7
490 267 pre 5 476 606 pre 9 463 257 gene 13
503 283 gene 15 491 628 pre 11 462 196 gene 23
526 324 gene 17 498 167 gene 15
456 367 pre 5 478 130 pre 8
571 507 pre 7 417 281 gene 10
490 619 gene 12 360 325 pre 3
193 500 post 14 346 322 pre 2
415 123 gene 16 353 280 pre 4
630 490 pre 6 434 370 pre 9
447 371 pre 5
465 375 pre 6
466 387 pre 1
451 423 pre 7
445 433 pre 9
618 390 pre 6
588 376 pre 3
582 457 post 13
533 498 pre 3
414 473 pre 10
446 523 post 18
456 507 pre 6
477 537 pre 4
500 527 pre 4
508 522 pre 7
455 576 pre 5
308 240 post 13
339 202 gene 18
: r1, r2 – (
, ,
5,0 ); Age – ; d – .