Data analysis

in community and
landscape ecology
R.H.G.Jongman, C.J.F.ter Braak
and
O.F.R.van Tongeren
(Editors)

Published by the Press sindicate of the University of Cambridge

The Pitt Building, Trumpington Street, Cambridge CB2 IRP
40 West 20* Street, New York, Ny 10011-42,USA

Cambridge University Press

First published by Pudoc (Wageningen) 1987
New edition with corrections published by Cambridge University Press 1995
Reprinted 1996

Printed and bound in Great Britain by

Biddies Ltd, Guilford and King's Lynn

A catalogue record for this book is available from the British Library

Library of Congress cataloguing in publication data

Data analysis in community and landscape ecology. Translation from English. Editors of translation: A.N.Gelfan, N.M.Novikova, M.B.Shadrina. Moscow. 1999. 306 p.

Translation by N . B . L e o n o v a

Layout design:
V.S.Korshunova, G.Yu.Trofimova

Translation and scientific editing were made by Water Problems Institute of Russian Academy of
Sciences.
This publication was made possible through support provided by the UNESCO/BEO/BMBF Germany, under Grant 509/RAS/41 - Aral Sea, US Agency for International Development (Global
Bureau, Office of Agricultural and Food Security), the Global Livestock CRSP through the Management Entity, University of California, Davis, uncter Grant DAN-1328-G-00-0046-00.
The opinions expressed herein are those of the authors) and do not necessarily reflect the views of
the sponsors.

Translation and publication in Russia was permited by (Cambridge University Press, 16.03.1998).

ISBN
Cambridge University Press
TranslationfromEnglish to Russian and remarks of Water Problems Institute of Russian Academy of Sciences
Small Ruminant CRSP USA

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R. Jongman
S. . Braak

Wageningen

. . van Tongeren

September 1987

XI

Drs. A. Barendregt
Department of Environmental Studies
University of Utrecht
Heidelberglaan 2

IT. C. Looman
Research Institute for Nature Management
Ministry of Africulture and Fisheries
Present address:
Department of Social Health
Erasmus University
Dr. Molewaterplein 50
P.O.Box 1738
3000 DR Rotterdam
the Netherlands

P. O. Box 80.115
3508 TC Utrecht
the Netherlands

Professor Dr. P. Burrogh

Geographical Institute University of Utrecht
Heidelberglaan 2
P.O.Box80.115
3508 TC Utrecht
the Netherlands

Dr. Ir. Th.J. van de Nes

Department of Water Management
Province of Gelderland
P.O.Box 9090
6800 GX Arnhem
the Netherlands

Dr. J.C. Jager

Centre for Mathematical Methods
National Institute of Public Health

Drs. J.A.F. Oudhof

Department of Plant Ecology
University of Utrecht
Present address: RAET
Software & Computer Services
Endrachtlaan 10
Utrecht
the Netherlands

and Environment Hygiene

Anatomie van Leeuwenhoeklaan 9
P.O.Box 1
3720 BA Bilthoven
the Netherlands

Drs R.H.G. Jongman

Dpartment of Physical Planning
Wageningen Agricultural University
Generaal Foulkesweg 13
6703 BJ Wageningen
the Netherlands

Dr. C.J.F. ter Braak

Agricultural Mathematics Group
P.O.Box 100

6700AC Wageningen
the Netherlands
Dr. O.F.R. van Tongeren
Limnological Institute
Royal Netherlands Academy of Sciences
Rijksstratweg 6
3631 AC Nieuwersluis
the Netherlands

- ,
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R. Hegenveld, N.Gremmen, R.Barel, J.Oude Voshaar, P. Vereijken, L. Barendregt, J. Jansen,
M.Jansen, P.Verdonschot, H.van Dobben, H.Siepel, P.Opdam, J. Berdowski, F. Metselaar, G.
Hock, W. Gramer, F. de Hullu, K. van de Kewrkhof, P. Laan, A. van de Berg, E. LiefinkKoeijers, R. Webster, G. Oerlemans, W. Hockey-Wielinga, W. Slob and W.J. van der Steen.
M. Batterink, G. Wijffels P.Ketner Department of Vegetation
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, :
- (Hurlbert, 1984);
- t - (Innis, 1979);
- (Dawkins, 1983; Joh
nson & Berger, 1982; Mead & Pike, 1975);
- (Buckland, 1982; haetal., 1982);

23

-
(Meyers et al., 1982).
, ,
.
2.6.2.

.
. Mertz & McCauley (1980) ,

. ,
, . Hurlbert (1984),
, ,
"
". ""
, ,
. ,
,
. Tacha et al. (1982)
,
( )
( ).
,
,
.

, - . ,
, - ,
,
( , : "prophecy ex eventy").
2.6.3. ,
, " "
(Snedecor & Cochran, 1980). , , ,
,
( ) ,
(), .
, ,
(, ).
,
.
, .
24

2.6.4.
(Strong, 1980) .
,
, , : "
, "
(Strong, 1980, .271). : ", , ,
( ), - .
, ,
, . ...
.
() ,
, ,
" (Strong, 1980, .273).
,
( ) . ,

(Quinn & Dunham, 1983; Simberloff, 1983); , ,
.
(1980) (Toft & Shea, 1983,
. Cohen, 1977) ,
.
, "

, , .
,
,
, ".
(
)
" " " ".

() ,
.
- ,
. .
,

.
2.7.

, .
, :
25

- ;
- ;
- .
,
. ,
(, ),
. ,
, ,
, , , . ,
, . ,
, , , .
, ,
.
-
.
,
.

, .
.
, 2.3.1,
.
2.8.
.
, .
, *.
Sokal & Rohlf (1981),
. Clarke (1980)
.
, Snedecor & Cohran (1980), Box et al. (1978) & Snell (1981,
).
, , (Cohran & ,
1957; , 1958) (Cohran, 1977) .
Pielou
(1977), Green (1979), Southwood (1978). Slob (1987)
.

26

2.9.
2.1.
, ,
-
. ,
: - . , - .
/ -,
(1979) (. . 2.1).
.

N
N
N
N
S
S

g
h

s
s

[- JL

.
Calluna

Deschampsia

Calluna

Deschampsia

70
10
7
90
70
80
80
85

4
70
80
2
4
10
20
5

8
5
5
9
8
7
9
9

1
9
8
1
4
5
7
5

,
.
.
2.10.
2.1.
- . Deshampsia.
% (4 + 70 + 80 + 2) / 4 = 39%.
[(4 - 39)2 + (70 - 39)2 + (80 - 39)2 + (2 - 39)2 / 3]1/2 = 41.8%.
: (4 + 10 + 20 + 5) / 4 = 9.8%. : [(4 9.8)2 + (10 - 9.8)2+ (20 - 9.8)2 + (5 - 9.8)2/ ] = 7.3%. (
): , .

27

[ - .

s.d.

s.d

Calluna

Deschampsia

Calluna

Deschampsia

44.3
42.1
78.8
6.3

39.0
41.8
9.8
7.3

6.8
2.1
8.2
1.0

4.8
4.4
5.3
1.3

, Calluna
, . , Deshampsia, -
, Deshampsia .
,
.

28

3.
. J. F. ter Braak and . W. N. Looman

3.1.
3.1.1.

,

. , .
,
,
.
.
( , ),
- ( , ).
(response variable) ,
- ,
, .
, ,
. ,
(), ,
.
, , . ,
, () -
.
"
" (Whittaker, 1967). ,
, :
- , , ;
- ( ) ,
, ,
;
- ( -)
;
-
.
4.

29

3.1.2.
,
: ,
, ,
.
.
- . ,
, (.3.1):
= bo + bi + e ,

3.1

- ;
- ;
- ;
bo \ , ,
, .
() bo +b\x.
:
= bo + b\x.
-
.

.
6 \.
, -

>I

. 3.1. ,
. . 3.1.2.

30

 (LSR - - least-squares regression),

( 2.4.3). ,
,
(, - - 2.4.3),
.
(Williamson, 1972). ,
LSR - (,
- -
).
(LR - - logit regression),
.
3.7.5.

,
( 2.4.2). ,
, .
,
- .

. ,
. -

. 3 2 . . () ().
(: ), (: , : ); (d: ); (: , f:
, g: -); (h).

31

 ,
. ( ) , - , - .
(. 3.2):
- : - (. 3.2);
- ( ): ( )
. (. 3.2b-3.2d);
- : , .
(. 3.2) ,
(. 3.2f). , ,
. (
"" (. 3.2g)). (
), (. 3.2g).
- : , ,
, (. 3.2h);
- : .
.
.
,
, .
(. 3.3).
,
(. 3.3).
"",
(. 3.3).

. 33. , .
- f, - - d - f,
- b - - - - d. b -
( - ; - ).

32

3.1.4.

(LSR - , 3.2) - (LR - ,
3.3). ,
, ,
, , .
, ,
, ,
. - ,
- -
.
() () . 3.4
, , .
3.5 LSR - LR -
(- ).
,
-. 3.6
. -
(
- Gause, 1930),
.
3.2. LSR -
3.2.1. :

, ,
.
: , .
,
(
, , ,
). 3.4
.
.
,
, -
. 3.4 ,

. ,
.
(ANOVA - analysis of variance), LSR - .
33

8 *

s
a

34
9

"Ey

I-

fy i

a .
*

-~

3.4. () ,
( ).
(.3.1). 95%-
; - 95%- (
: : / , -
: , , ).

.
.
.
,
. ,
,
( . 3.4 ).
( . 3.4)
( . 3.4). , ,
;
.

, (ANOVA - ).
(. 3.1) ,
. (R2%dj) 0.25, . .
.
ANOVA- ,
, ,
, , .

34

 3.1. ANOVA -
3.4.

. (s.e.)

95% -,

1.70
3.17
2.33

0.33
0.38
0.38

(1.00; 2.40)
(2.37; 3.97)
(1.53; 3.17)

2
17
19

7.409
14.826
22.235

3.704
0.872
1.170

4.248

2.33

ANOVA

^^=0.25

F (. 3.1)
F- 2 17 ,
(2 17 - ANOVA - ).
( 5% ) 3.59 (. F- ,
, (Snedecor & Cochran, 1980).
(4.248) , 3.59. ,
5% . , ,
. ANOVA - ,
5% ( < 0.05; F ).
?
(.3.1).
. 95%-
( )
:
() tow () * ( )

3.2

( ). .
5% t - . t (v)
(v) t - (Snedecor & Co
chran, 1980). v = 17 . (17) = 2.11, ,
3.4 3.1.

35


.
, .
, ANOVA -
" ". 3.1
: ^(0.872) = 0.93. (s.d.)
.
, 95% , :
( ) to.os (v) J(*.<L2+*.e2)

3.3

s.d.- , s.e. - .
3.3. :
1.702.11/(0.931 +033') =(-0.38; 3.78).
, .
s.e.=0 .05 ( ) = 1.96 3.3 :
1.96 s.d. 3.4 .
ANOVA - , .
, ,
:
( ) = (, ) + () 3.4
, - 3.89 (
48%). 3.17 ( )
, , 3.89 - 3.17 = 0.72. , 3.89 = 3.17 +
0.72.
3.4 ;

(2.33)

() ,
, .
, ,
, 3.1 , s.s. ( ).
,
.
(d.f. . 3.1). -1 ( ), q - 1 (q -
, 1 ) - q
.

36

 - 20 q = 3. m.s. ( )
.
( , )
. ,
, . ,

/?idj= 1- ( / ),
.
2 8= 1- (0.872 /1.170) = 0.25.
, (2), ,
,
2= 1- ( / ).
, 2 1,
, , .
, ,
.
(F).
,
, , -
. .

3.2.2.

3.5 -
, .
:
= 0 +6 \,

3.5

- ,
- ( ),
bonbi- , :
bo - ;
\ - (.3.1).
, ANOVA ( 3.2.1), ..

37

t
a

* oT

Q.

6
4

. 3.5.
, .
, -
(*). - , ().

. 3.56. ,
3.5.

38

 , .
, ()
.
, :
bo \,
3.5
. ,
. 3.5 ,
,
.
, . ,

. b0 \

3.6
3.66

. Xi X i- X X,
.
3.2
. bo 4.411, 6i=-0.0370.

3.2. , :
ANOVA- 3.5.

bo

4.411

0.426

10.35

-0.0370

0.00705

-5.25

F
27.56

ANOVA-

R 2 40.58

13.45

13.45

18

8.78

0.488

19

22.23

1.170

39

ANOVA - ,
3.2.1 ( 3.4), ,
3.5. ANOVA - ,
3.2.1. V0.488 = 0.70,
VU70 = 1.08.
0.58.

(0.78 . 3. 5).
F
,
. F 5% 4.441,
1 (Snedecor & Cochran, 1980).
F (27.56) , -
. F - t \ ; \ , , . .
. t -
\ (. 3.2). (-5.25) (
) t - 5% ,
t -: t . (18) = 2.10; \, , 0,
.
95% bi
3.2. -0.37 2.10 0.00705 = (-0.052,
-0.022). , . ., , bi*0
(F - , t - )
- , , .
, ,
, . ,
:: 3.5,
:. 3.5 .

.
, 3.5, (.
. 3.56). ,
;
.
, , 3.1.
3.2.3.
3.2.2,
(. 3.5). , ?
3.5, (. 3.2)
40

 33. , :
ANOVA-
3.5.

3.988
-0.0187
-0.000169

0.819
0.0317
0.000284

4.88
-0.59
-0.59

2
17
19
0.57

13.63
8.61
22/23

6.815
0.506
1/10

13.97

(. )2
ANOVA-

*4

= bo + bix + 2

3.7

, 3.3,
. 0 \ 3.2;
2 . ,
( . 3.5), .
- ( 18 17),
, .
, , .
, /- (
3.2.2), . t (. 3.3)
2.11, ,
0. ,
,
.
t - 0 \ 3.3
- . , \
3.7 .
3.1 3.7 .

, 3.7
3, JC4,...,
.
, .
, , (^
) .
z=

cexp[-0.5(x-uf/i2l

3.8

41

. 3.6. : (),
() (). 4.

- ;
- ;
- ( , );
t - ( ).
, t :
( 3.2.2) .
, .
3.6 . ,
4*. 3.8, :
loge* = log*() - 0.5 ( - )21 = 0 + bix + 2

3.9

:
0 - / = 2 -2 + 2

*= logc(c) - 2/ (22);

\ = IZ2; 2 = -1 / (2?)

3.10

b0, b], b2y :
w = - ^i / (2)

3.11

t = 1/ V (-262)

3.116

= ( + \ + 2)

3.11

42

 < 0. ,
.

\
.
3.9.
, 3.2 (
3.2.8).
3.3. -pei -
3.3.1. : -
3.4 ,
Achillea ptarmica, , .
.
, Achillea ptarmica

.
3.2.1, ,
. , ^
. ,
, (. 3.4).
,
1 0. ,
3.2.1. , = 1 = 0,
, ..
, , .
, , .
3.4. , Achillea ptarmica

(
Kruijne . 1967). :
(ph), -(11), ()
().
Achillea ptarmica

ph

ph

37
109
146

40
356
396

27
402
429

9
558
567

113
1425
1538

0.254

0.101

0.063

0.016

0.073

43

 Achillea ptarmica

, ,
. -
. .3.4,
113/1538=0.073. , Achillea ptarmica
0.073 146 = 10.7, ^ 0.073
396 = 29.1 . , Achillea
ptarmica , , 146 - 10.7 = 135.3, - 396 - 29.1 = 366.9.
()
() -, {o-efle
3.4. : (37 -10.7) 2 / 10.7 + (109 -135.3) 2 /135.3 + (40 -29.1) 2
/ 29.1 + ... = 102.1. %()
- v , v = ( - 1 X - 1 )> - -
. v = 3 5%
2 (3) = 7.81 ( 2- , , Snedecor &
Cochran (1980)).
(102.1) 7.81, ,
Achillea ptarmica
. , -
:
Me.
- ,
. ,
1
, .
.
3.3.2.
, - ()
( -).
3.7. 3.3.1, -
.
. 0 1.

bo + b\x

3.12

, 0 + \ .

= 0 + bix)

3.13
44

0.5-

^ " - - ^ _ _ ^

?~^_^

^ ~
0

1
3

|
4

f
5

- _ _ _ _ _ _ _ _ _ _ _
,

II

1
8
-

3.7. , - ( =
1) ( = 0). ,
. () .

, 3.13 1,

=/> = [exp (6o+ b\x)] I [1 + (0+ \)]

3.14

, 0 1.
, - . ,
3.12 - 3.14, 3.8; 3.14
. ,
0 \. b0+bi x .
>" ( 3.14).
.
- ( ).
, ,
( 2.4.3). >> /? (1-). .

, .
- (LR - ). (McCullagh <&
Nelder, 1983) ,
(generalized linear model - GLM). "" -,

1

& \ I (1 - )]

3.15

:
= [ ( )] / [1 + ( )]

3.16

3.3 , 3.15 3.16 .

3.15 - GLM.
45

. 3.8, (), () (),

3.12, 3.13 3.14 .

GLM .

, .
, ,
.
, - 2 logc L, L . ,
.
( 3.2.1) ,
.
, , GLM. , GLM
;

,
, . LR (1 - />). ,

46

-X

3.9. (), () - (), 3.7,

3.8 3.17 .

" , , ;
. LR -
GLIM (Baker & Nelder, 1978), GENSTAT (Alvey et al, 1977),
BMDP (Dixon, 1981, PLR) SAS (Harrell, 1980). (1986)
GLIM.
, 3.14,
3.7 LR - . 3.5
; n-q, q
( 3.2.1). (. 3.7) ( > 0.05)
( - , \ 0).
, LR - , ,
( - 3.3.1).
, ,
3.2.2.

47

 3.5. , LR - :
- 3.7.

2.03
-0.484

33

1.98
0.357

1.03
-1.36
.

43.02

1.304

3.3.3. -
3.16 , (GLC - Gaussian logit curve)
= [exp (&o+ bw + b2x2)] I [i + exp (&<>+ bw + b2x2)]
= exp [-0.5 (x - uf I J2] / [+ exp(-0.5 (x - uf 111)]

3.17

3.8 - 3.10 GLC

( 3.8). , GLC 3.9
( . 3.8). GLC , ,
, GLC (<0.5). GLC
3.7 GENSTAT ( . 3.10). 3.6
b09 b\, 2,
3.11 , : = 5.28 t = 0.327.
3.10 ( max)

4.

0.5-

. _
/ ^ ^ ^

-fMi IIIHI

"1 f t

II

II

8
.

. -, LR- ( /? = 1) ( = 0) (). ,
3.7: = ; X = ; mtx= .

48

 3.6. -, LR -
-: . ( ,
. 3.7).

-128.8
49.4
4.68

51.1
19.8
1.90

-2.52
2.50
-2.47

32

23.17

0.724

3.17 (5.28)
\ 3.6. pmw= 0.858.
, GLC
, .
f- ( 3.2.3). t -2.47 (. 3.6),
. . GLC .
t - , , ..
< 0 (Snedecor & Cochran, 1980, 5.5). 0,
, . 95%-
(5.0; 5.8) (. 3.9).
GLM -

. ,
-
, - . ,
GLC (. 3.5 - 3.6) 43.02 - 23.17 = 19.85.
, Xo.os2(l) = 3.84, ,
. F-
LSR - .
, , 3.4-3.5.
3.4.
(.. )
. ,
, .
, . 0
,
, 0.1; 0.01 .
, ,
.
, 3.2.1, , 49

 ,
. ,
, (, . 3.4).
LSR - ,
- . ,
- 0. ,
- ( 2.4.3).
, , 0,
1, 2, 3,... , .
, , . ,
, . ,
, , 1.
, 3.13, .
- , , -,
GLM ( 3.3.2). -,
3.13
loge Ey =

3.18

b0 + b\x + ^2,
< 0 ( 3.8 - 3.11). ,
- .
. ,
0, \, , 3.2.3.
, ,
( 2.4.3). , , -
, - : ,
3ra4eHrao(McCullagh & Nelder, 1983). ,
, >. .

"" (/7/ . 1975). - -,
, , " " - .
, .
, ,
LR - .
,
, . ,
.
.
, ,

.
50


: "'', "", "".
: " ?" " ?", -
-.
3.5.
3.5.1.

. ,
.
.
( ).
,
, ..
.
, ,
, .
.
-,
.

.
3.5.2. :

(. 3.11):
= & + \\\ + \2,

3.19

Xi 2 - ;
bo, \2- .
0 \ = 0 xi = 0.
] 2, , bi 2. , \
xi % -
\.
, ..
.
(. 3 1),
,
.

51

. 3.11. , . , ..
() .

, ,
0, \9 ,
t (. 3.3).
, Xi = , = 2. t ,
( 3.2.1), ..
()
.
,
, :
- bo + b\Xi + 62X12 + 63X2 + >422

3.20

, "" - ,
*. t . ,
xi, (>
0) ( < 0) f,
, (. 3.3.3).
, * ,
2.
\
3.2.3 3.11, \ ,
3.20.
3.9.
\ &, *.
\,
0 , *. ,
F - ,

52

 . .
: - \ \ ,
2 , RSSi RSS2, ,
:
F= [(RSS,- RSS 2 )/(dfi- df2)] /(RSS 2 /df 2 )

3.21

dfi df2 - RSSi RSS2, .

, 0,
F F- dfi - df2 df2
( 3.2.1). , F
, .
,
. F-
3.21, "
" ( ).
3.5,3. -: - -
, 3.3.2, ,
loge \p I (1-/?)] ,
3.19 -,
loge \ / (1 -)]~ + \\ + 2

3.22

,

. 3.12 xi 2 ,
. 3.22
GLM ( 3.3.2) , 3.7.
- , \, bj 0, . .
xjt 2. xi ( 3.14)
3.7.
\ ( = 0.16), (f \ 1.33).
, , ,
\. 2, ,
.
(. 3.7) , \ JC2
, , . ()
, (Witteker, 1964).
( .3.12) :
JCI,
*i 2. ,
45 3.12,
.
53

10

0-

. 3.12. , ,
. \ , ()
().

3.13. ,
. } 2 ,
() ().

54

 3.7. - 3.12
(fibres) -
)

*1
*i

*2

*1

1.53
0.16

-1.66

-0.15

d(res)

df.

2.98
1.33

-2.96

-1.17

23.99
45.25
45.69
47.11

32
33
33
34

3.8. - 3.13
,

\&
\

d(res)

df.

1.63
-2.85

-1.59

-2.88

17.47
20.57
20.82
41.88

32
33
33
34

<
-0.61
-0.94

-0.625

-1.016

3.13 . , \
, - ( . 3.8)
, \ 0. , \,
. ,
\9 2. | 2
, . ,
, , \ (, 1984).
3.13, xi ,
, , , \
, 2 - , .
. , , . ,

.
, .
,
.
3.12-3.13 . ,
,
, . , -

55

3.14. - {\
, .

\ 2 ,
. , , *i
, .

. ,
, , , ..
(,1984). ,

(Montgomery & Peck, 1982; Whittaker, 1984).
. 3.20
3.15, -, \
(. 3.14).
, *i 2 (. 3.146).
.
, F- (.
3.3.3).
3.5,4.
,
.
\.
3.19 :
= 0+ bjxj + ?2+ ]2= (0+ ) + (\ + ) \

56

3.23

 ,
. 3.146.
\ 2. \ 2.

3.23 , JCI
, , ,
, \ 2 , \.
\, > 3.23
( *=*1*2
).
- ( ).
3.20 ( ) :
loge [ I (1 -)] = 0+ &i*i + \2+ 2+ 22 + 5\2

3.24

+ &4<0, 4- > 0, 3.24

, 3.146,
. ,
(, Car
roll, 1972). , ()
3.24:
1 = (>- 2\) / d ,

3.25

= ( M i - 2) I d,

3.256

raerf = 4W>4-fc52.

57

. 3.15.
-.
2.
Xi, 2. 2 .

*i {\ + 65*2) / (262) , ,
2, bs 0.
3.25 3.10 3.11. 3.15
. , , t, $ .
3.5.5.

,
. ,
, ANOVA (.
3.2.1) . 3.2.1
, : , , .
"" ,
- *, 0, 1.
1, 0,
- . * 1,
0, -
. , , 0
. ,
3.2.1,
= \ + + * ,

3.26

\ ;
,

58

 - .
\,2 .
3.4. \ = 1,70; = 1.47; - 0.63.
\ = 1.70; b\ + hi- 3.17 \ + = 2.33,
3.1. ANOVA
3.1. ,
, - 1 (Montgomery &
Peck, 1982, 6).
- Equisetum
Jluviatile .
(mSm"1) -,
bge \ I (1 -)] = b0+ b\x\ + bixi2 + & + ,

3.27

\ - , * - ,
. by * (
).
3.16 ,
, .
- ;
, - .

, \ JCI2
3.27. - , -
, .
3.27,
3.11;
- 3.9. 3.6.

log EC

3.16. Equisetum fluviatile ,

-, ()
, (, , ). de Lange (1972).

59

3.6.
.
,
(), .
- (Belsley ., 1980; Cook & Weisberg, 1982;
Hocking & Pendleton, 1983). ,
, , ( )

. .
- .

. (Austin,
1980) , , .

. (1930), (Curtis & Mcin
tosh, 1951) (1956) ,
, .
,
:
,
.
, ,
(Fresco, 1982). (Hill, 1977)
, , (),
.
, ,

, 3.3.1 (Gounot 1969; Guillerm, 1971).

.

:
- , ;
- ;
- , ;
-
, ,
-, . ,
-, 3.3.1 - , t ( 3.3.2, 3.3.3 3.5)
.
, ,
60

. 3.17. - .
=7.6 ( ) . 3.10
.

. ,
? , , ,

.

.
, ,
-. , 1 3.10 (
= 7.6) (. 3.17).
,
3.12 - 3.13
. ,
, ,
;
.
, ,
( "
", (Efron, 1982).
.7.
(Ellenberg, 1982).

: , . ,
. 3.2.3 3.5.2 ,

.
(Ellenberg, 1948, 1979). ,
. -

61

 ,
. ,
- , .
,
, :
* = (\\+27+ ... +;ttvtn)/Oi+>'2+...+>'n)

3.28

*- ;
* - ;
\, *2,.,* - 1,2,.../.
.
,
(. 3.18).
(ter Braak & Looman, 1986)
-
- . ,
,
:
- ;
-
.

( 3.2.8). ;

,
, .
,
(
. 0.1)
. ,
0.1
. ,
(Aim praecox)
* = ( 2 2 + 5 ) / ( 2 + 3)=3.8
, -
(
). 3.9. Plantago lanceolata
, Ranunculus flammula - , a Alopecurus

62

0.6
0.4

<

0.2-

11#1#1^. ...

20]
10
JXX

20

10

+ ^

3.18. ();
80 , ()
(). .
. = , N ; - .

geniculates . 3.9
(>0) .
,
. 5.2.
3.8.

. 1920- .
(GLM) (Nelder &
Wedderburn, 1972) .
,
, Draper & Smith (1981), Seber (1977), Montgomery &
Peck (1982), Mosteller & Turkey (1977).
: Parker (1979), Sokal & Rohlf (1981) Snedecor & Cochran (1980). Dob-

63

 3.9. ,
0.1. ()
; () - (*)
. ,
. .
0.1.

26
18
28
1
6
7
23
17
9
11
13
5
20
16
27
2
13
24
4
15
3
25
2
14
8
10
12
21
22
30

Tri pra
Pla Lan
Vic

tat

Pch mil
B*I pQr

Rum
Lot

Cir
Ely
Poa
Pnt
Poa
Leo
Tri
Bra
Hyp
Sag

PI
Jun

Rir
Sal
Rgr

nor

pmr

arv
rep
pra
odo
tri
aut
rep
rut
ad
pro
gen
buf
pra

sto
art

Jun
Che
Ele
Emp
Pot
Ran

alb
pal
nig
pal
fla

tal

CUfi

11 11 1 111112 ..
12S67183407926345690
2S2
S5533 32
21 1
13222
42
32
2222
42 2
34
563
22
7S266726S6 2 4
2
444
44 6
442344354414 42
432
44
4
27645 654 5449 2
5333S52232223222 62
52S232216 332261 2
26246222 242 4434
2
2
5
2 5 2422
3
2
72 3855 4
2
44 3
2
3
3
35
48 3445447 5
4 4 33 4
1
4 458 4
2
22
22222 4
4 3 3

u*
1.0
1.2
1.3
1.4
1.5
1.5
1.7
1.7
2.0
2.0
2.0
2.1
2.6
2.6
2.7
2.3
3.4
3.5
3.7
3.8
3.8
3.9
4.1
4.8
5.0
5.0
S.O
5.0
5.0
5.0

11111112222445555555

son (1983), McCullagh & Nelder (1983) GLM.

(1956,
1967).
,
. , ""
.
. (Gleason, 1926), (1930) (1930),
, "",
. " (1928),
.
64

 . , ( ,
) .
(Mcintosh, 1981).
, (1982).
(1956, 1967) , ..
. (Gounot, 1969) (Guillerm, 1971)
, 3.3.1,
- .
, .
-
.
(Jowett & Scurfield, 1949).
- (Finney, 1964) -,
.
(Yarranton, 1969; 1970).
.
(Austin, 197I) .
,

. (Alderdice, 1972)
. (Gauch & Cha
se, 1974)
,
. GLM.
. (1984) GLM ,
- - .
, GLM ( 3.5)
. , ,
.
3.9. ;

hi 3.9, 3.17, 3.20 3.24 vn V22
v^. ,
:
var(u) = (vu +4uvl2 +42)/(4})

3.28

var(f) = v22 /(-&>23)

3.29

100(l-ot)%
(Fiellers) (Finney, 1964, . 27-29). ta - t , -3 , (,
65

t = 2.00 95% 63 ).
:
g = fa2 V22 / 2

3.30

D = \ var() - g(vn - vf2 / )

uiov^uuppcr ^\^05^/220^)/2\/(1-

3.31

fo (g > 1),
, .
.
3.1.


(van Dam ., 1981).
400 . Frustulia rhomboides var.saxonica
Srd. = [S0 4 "2J / ([] + [S0 4 "2] + ["]) 16 ,
1977 1978 . (. 2.5, van Dam .,1981):

Frustulia

5^.

V2

VI

Bl

D6

B7

D3

D2

Di

D5

D6

0
0.78

0
0.64

14
0.69

3
0.70

0
0.64

5
0.77

6
0.73

21
0.77

62
0.58

26
0.44

14
0.44

48
0.37

97
0.23

99
0.19

28
0.31

202
0.23

3.1.1. , loge [
Frustulia +1]. , .

66

 3.1.2. ,
loge [ Frustulia +1],
.
, .

Sri.

5.848
-5.96

0.806
1.41

7.26
-4.22

1
14
15

24.34
19.11
43.45

24.340
1.365
2.897

0
bx

ANOVA-

3.1.3.
, 3.1.2.
2.

3.1.4. ,
, .
?

3.1.5. 95%
\. \ ( < 0.05)?

3.1.6.
0.25; 0.50; 0.75.
. , , 0.49; 0.30; 0.42.
, Frustulia.

3.1.7. 95% ,
0.25,0.50 0.75.

67

 3.2. ,

,
Radiolaria (Lozano & Hays, 1976),
.
(%) Spogotrochus gracialis
( ) 34 .

i
12
0.8

2
14
1.1

3
13
1.6

4
22
1.8

5
18
1.7

6
19
2.0

7
7
1.6

8
8
1.9

9
11
2.0

10
15
2.5

11
12
3.7

12
14
4.2

13
16
4.1

14
21
5.8

15
35
6.1

16
30
6.6

17
34
7.9

18
48
10.2

19
47
11.0

20
63
11.9

21
54
12.8

22
62
14.8

23
56
15.9

24
52
18.1

25
41
16.9

26
38
17.1

27
30
18.0

28
18
18.5

29
25
20.0

30
35
21.0

31
37
19.4

32
38
19.8

33
42
19.0

34
41
21.6

3.2.1. ,
. , .
.
3.2.2.
. :

.2

2.119
0.2497
-0.00894

0.133
0.0356
0.00164

15.95
7.01
-5.46

2
31
33

9.42
3.06
12.48

4.7101
0.0988
0.3783

bj

b2

ANOVA-

3.2.3. 5, 10, 15 20
, ,
, , .
3.2.4.
?

68

 3.2.5. 95%
b-i. ?

3.2.6. 95% ,
3.31, , \ -0.00005704
.
,
.

3.2.7.
(. 3.2.3) .

3.2.8. ( 3.7)
Spongotrochus , ,
- . ,
. ?
3.3. -
3.15 3.16, , logc [p I (1 -/?)] = , ,
, = ( ) I (1 + ).
3.4. - -
160 Elymus
repens (
). , (Kruijne et al., 1967),
.

12
16
28

96
36
132

108
52
160

3.4.1. .repens .
3.4.2. ,
?
- (. 3.3.1) 5%.

69

 3.4.3. - -
- E.repens 160
.
,
"", 1, - 0 . - :

"1

-0.28
1.27

0.38
0.42

-0.74
3.02

158

192.9

1.221

, : loge [ pi (1 -)] = + \ ".

3.4.3.1. E.repens
, 3.3. 3.4.1.
3.4.3.2. f-
.
3.4.2.
3.4.3.3.

, loge [p I (1 - )] =

201.7 159 .
t - .
3.5. -
, ,
.
,
. E.repens
-. :

bo

-57.26
19.11
-1.55

15.4
5.3
0.44

-3.27
3.61
-3.52

157

176.3

1.123

70

 3.5.1. E.repens ?
.
3.5.2.
4.5; 5.0; 5.5; 6.0; 6.5; 7.0; 7.5 , ,
E.repens .
3.5.3.

( < 0.05) ; - ? t.
3.6. -
,
, E.repens
(. 3.4 3.5). , ;
.
, , E.repens
.
-.

1\(\-)]

= +1" + 1+112

""

-57.82
-0.04
19.30
-1.56

17.10
0.57
5.81
0.49

-3.38
-0.07
3.32
3.18

156

176.2

1.129

0
,

3.6.1.
= 5 = 6. ,
.
.3.5.1-3.5.2, .
3.6.2.

. ,
3.4.2?

71

 3.6.3. t - 3.6.2
. ?
3.6.4. ,
E.repens
.
, 3.5.3.
3.11.
3.1.
3.1.3. : = bQ +fciSrei.;
: ( - )
, SKL 2 14
( = 2.71) Srei. = 0.44; : 5.848 - 5.96 0.44 =
3.23; 2.71 - 3.23 =-0.52. Frustulia,
: (3.23) - 1 = 25 - 1 = 24.
3.1.4. ANOVA
19.11, 1.365; : Vl 365 = 1.17
1 - (1.365 / 2.897) = 0.529.
14 .
3.1.5. 3.2 . (14) = 2.145, \
-5.96 - (2.145 1.41) = -8.98
-5.96 + (2.145 1.41) = -2.94. , 95% -
\ (-8.98; -2.94). 0 .
/ \ (- 4.22) (2.45),
\ ( < 0.05) 0.
3.1.6. SKL = 0.25
5.848 - 5.96 0.25 = 4.36.
0.49 , 95% - (4.36 2.145 0.49; 4.36 +2.145 0.49) = (3.31; 5.41). S i. = 0.50 0.75 2.87 1.38
(2.23; 3.50) (0.47; 2.29), . ,

. 5rei. = 0.25; 0.50; 0.75
(4.36 - 1 ) = 77; 17 3, .

^ , . ,

.

72

3.0

2.0

15

to

20

. 3.19. ( ),
Spongotrochus gracialis ()
. 95% , - 95% .
. Lozano & Hays (1976).

1004

80
604

40
20

10

15

20

. 3.196. 95% 95% ,
. - Spongotrochus gracialis %.
- .

73

 . 95%
95% . S rei. = 0.25
(26,223).
3.1.7. 3.3 Si = 0.25
4.362.145xV(U7 2 +0.49 2 ) = 4.362.145 1.27 = (1.63, 7.08)
, 95%
4 1187.
, 400.
SreL= 0.50 0.75 95%
(0.28; 5.46) (-1.28; 4.05), .
3.2. ,
3.2.1. 3.19, .
3.2.3. 5, 2.119
+ 0.2497 5 - 0.00894 52 = 3.14. 10; 15 20,
3.72; 3.85 3.54. ,
5;10;150 20, 0.07; 0.10; 0.09 0.11.
3.11 3.116
= -, / (22) = -02497 / (-2 0.00894) = 14.0, 14
f = l / J(-2b2) = 7.48 ,
7.48. (. 3.19) 2.119 + 0.2497 14.0 0.00894 14.02 = 3.86.
3.2.4. V0.0988 = 0.314
1 - (0.0988 / 0.3783) = 0.739
(ANOVA).
3.2.5. 3.2 .5 (31) = 2.04,
2 (-0.00894 - 2.04 0.0164; 0.00894 + 2.04 0.00164) = (-0.0122; -0.0056).
, , 0 ( < 0.05) t
-5.46; , (&2=0)
(bj 0). ,
.
3.2.. 95% (12.8; 16.2).
3.2.7. Spongotrochus 5; 10;
15 20 - (3.14) = 23, 41, 47 34, .
95% 95% (
) 3.196.

74

 3.2.8. (12 0.8 + 14 1.1 + ... + 41 21.6) / (12 +

14 + ... + 41) = 12.7, , 12.7.
,
11. , (14),
, ;
,
, . ,
. :
95% ,
3.2.6.
3.3. -
log = [/(1 -)] = -+/(\

-) =

-> = ()(1 -) = - - + =
-> (1 + ) = -+ = ( ) I (1 + ).
; .
3.4. - -
3.4.1. : 12 / 28 = 0.43;
96 / 132 = 0.73.

3.4.2. ,
, 108 / 160 = 0.675. ,
28 0.675 28 = 18.9 E.repens, 28 - 18.9 = 9.1
. repens.
132 () : 89.1 .repens 42.9 -
. -
(12 - 18.9) 2 / 18.9 + ...+ (36 - 42.9)2 / 42.9 = 9.39;
5% - (2 - 1 ) (2 1) = 1 2.5(1) = 3.841. ( < 0.01),
.
3.4.3.1. : loge \p I (1 - )] = 0,
"" = 0 . 0 -0.28,
= (-0.28) / [1 + (-0.28)] = 0.43. loge \ I (1 -)]
= 0 + "" = 1 . c 0 +ci -0.28 + 1.27 = 0.99,
= 0.99 / (1 + 0.99) = 0.73. 3.4.1,
: ,
.

75

 3.4.3.2. "\" ""

( < 0.05), t = 3.02, t *.5 (158) = 1.98;
. 3.4.2;
.
3.4.3.3. , "
" , 201.7 -192.9 = 8.8,
%2 .
3.5.
3.5.1. 3.11,
=-19.11/(-2 1.55) = 6.16.
- 6.16 3.17,
=( 1.641)/(1 + 1.641) = 0.84 -57.26+ 19.11 6.16- 1.55
6.16 = 1.641. / = 0.57 ( 3.116).

3.5.2. =4.5; 5.0; 5.5; 6.0; 6.5; 7.0 7.5 3.17

0.07; 0.39; 0.72; 0.83; 0.81; 0.64 0.25.
3.5.3. ( < 0.05) 0, (-3.52)
, - (1.65
=0.05), , - , .
3.6. -
3.6.1. ( - 0) =5: loge \ I (1 -)] = -57.82 + (19.30 5) (1.56 52) =-0.32, = 0.421. ( = 1) = 5:1& \ I (1 - />)]= 0.32 - 0.04 =-0.36, =0.411.
= 6, ,
0.860 0.855. -19.30: (-2 1.56) = 6.18
0.57, .
, 0.867 0.862.
(. 3.20). ,
3.5 .

76

. 3.20. - E.repens (
) ( ) (),
-. E.repens = 5 0.421
0.411 , . Kruijne . (1967).

3.6.2. t \ ,
t 5%. , ,

. ,
, 3.4.

, .
..
176.2; ( "")
(.3.5), 176.3.
(0.1) , -
,
. , 3.6.2.
3.6.4.
176.2; ( 2)
(.3.4), 192.9.
, 16.2, -
2 cos (2) = 5.99.
,
.
, 3.5.3.

77

4.
. J. F. ter Braak

4.1.
3 ,
.
-
. , ,
.
,
- . () (),
.
, ,
, . . ,
,
,
. ,
.
, ,
.
( )
.
- ,
.
() .
, .
, ,
,
.
-, ,
- .
,
.
, ,
.
,
.
,
, .
78

 4.2) ,
- ,
( 4.3)
( 4.4).
4.2.

4.2.1.
{maximum likehood calibration MLC) .
, , ..
( 3) (
). , ,
. -
.
,
,
( 3.3.2).
,

4.2.2. 4.2.2
- .
.
5.5.5.
4.2.3
, -, -, , - .
,
. MLC
.
( 3),
, .
4.2.2.
, .
, : , , -
0.1; 0.2 0.4.
, ,
- "",

79

 . ,
- "", .
.
, (0.1 + 0.2) / (0.1 + 0.2
+ 0.4) = 0.43, , - (0.8 + 0.6) / (0.9 + 0.8 + 0.6) = 0.61, ..
*. ,
. , , ,
"".
, ,
. . ,
, , ,
, , .
"
" 0.6; 0.2; 0.2. , ,
, ,
- 0.6 0.1 = 0.06,
0.2 0.2 = 0.04 , 0.2 0.4 = 0.08 - . "
'' 0.06 + 0.04
+ 0.08 = 0.18 0.33; 0.22 0.44, .
- 0.44 - . ,
, : , ,
, . , , 0.8; 0.1 0.1,
- ",
. ,
. ,
. ,
. , , , ,
(Brown, 1979).

. . ,
, (, , )
, (, ).
(Lachenbruch, 1975).

.
. , ,
, ( = 0.01)
- ( = 0.98). ,
, , , , . ""
, . -

0.67. (..)

80

 " , " , ,
0.1 0.99 = 0.099; 0.198 0.008, .. - .
(0.35) , .
( ),
.
, " , ",
, , .
, ,
" " " "
. , ,
, .
, ,
.
, :
( = 0.2) ( = 0.4).
,
; .
,
. ,
, , .
,
(0.2 0.4 = 0.08). ,
.
:
- ;
- ;
- ;
- .
, 2 ,
.
, , , , 10,
. Lachenbruch (1975, .41-46)
.
,
, .
4.2.3.
-
,
. ,
0. , -

81

 . , , .
, : - +.
, ,
,
.
fc- /* (*) , /t- ,
: 1 - /*(*)
, ,
.
- ,
() . ,
:,
.
,
. ,
,
.
1 .
, , 1. ,
,
, .


( ) () .
= J(x), ,
= 0)
0.
,
. (. 3.1) ,

, ,
, . ,
(de Wit et al, 1984) ,
( ). , ,
, (Brown, 1982).
( 5.3.2) ,
jc0, 0 .
JC0.

82

 4.1

* = + &*

\ - fc-ro;
;
- .

,
( 3.6):
*0 = S k - i m ( n - a k ) b k / Z k . , m ^ 2

,
, .
(Brown, 1982).
4.3.
(
JC) -
. , .. ,
. (0) , ,
,
().
. ,
, (
). :
0 = Oiwi + 2**2 +

+ ) / ( + +...+)

4.3

:
,2, ~., - ,
Uo, 2,.., - .
, 3.7
, ,
( 3.28).

: (1948) (1956); (Zelinka & Marvan, 1961); (Chutter, 1972).
(1948; 1979) ,
(Bocker et al., 1983).
: 1- ,
, 9- - , .

83

 , , .

() .

. (1979), (Register, 1979), Aac (Vevle & Aase, 1980)

, .
, ..
, (.
4.2.3). , ,

, .
(Ellenberg, 1979) -
; (1961)
, . , , - ,
- , .
,
. ,
, - .
( Braak & Barendregt, 1986)
( 4.3). - ,
, , * 0 ,
:
- - ( 3.17) () (. . 3.6);
- ;
- ,
( 4.2.3);
- () ,
0 ( . 4.1);
- .
,

xQ = (w m >* Wk/ fk2) /(Ik-i m ^k/ '

4.4.

tk - ( 3.17).

(ter Braak & Barendregt, 1986).
(
3.8) , -

84

, .
( , )
(packing model - . 4.1).
,
> , . .
. (Christiansen & Fenchel, 1977, 3).
(
., 1973).
,
. , ,
.
,
. 4.4
, ,
.
. ,
,
, .
,
.

. 4.1. :
. -
( = 1), ( = 1) {- 0.5).
0 - .

85


, 0.1.
, -
3.7.
4.4
4.2.3. ,
.
.
, ,
, - . ,
, ,
. !>
, ,
(Brown, 1979).

.
,
.
. (Brown, 1982),
, ( 4.1).

. ,
(Bartlein et al., 1986). ,
,
.

(Howe & Webb, 1983). ,
(Bartlein & Webb, 1985)
, .
,
, .
, , .
. (Bartlein et al.,
1986): ,
( 4.2).

86

4.5.
,
, 4.3.
Sheenan (1984). Battarbee (1984)
,
(. Davis & Anderson, 1985).

,

. (Brown, 1979)
,
( 4.2.3.). ,
, Scheffe (1973), Schwartz (1977), Brown (1982). Williams
(1959, 3), Brown (1979), Brown (1982), Naes & Martens (1984)
-
, . ( ) Lachenbruch
(1975) . Titterington et al.(1981)
, Kanal (1974) -
.
4.6.

4.1.
- .
(Kruijne et al., 1967)
() , (1986)
.
. 4.2. - :

Agrostis ccmina
Stellaria graminea
Alopecurus geniculatus
Plantago major
Belli s perennis
Hordeum secalinum
Glechoma hederacea

AC
SO
AG
PM
BP
HS
OH

3.4
5.7
5.8
6.2
6.4
7.1
8.1

1.1
0.4
0.6
0.7
0.5
0.7
1.5

0.84
0.38
0.58
0.34
0.89
0.57
0.55

100 ,
.

. 1 AC, SG ; 2 - AG ;
3 - HS ( ).
, :
87

4.8

5.2

13

28

5.6

6.0

6.1

6.8

10

7.2

10

4.2. ()
, -. ,
( ). : Agrostis canina (AC), Stellaha graminea (SG), Alopecurus geniculates (AG), Plantago major (PM),
Bellis perennis (BP), Hordeum secalinum (HS^, Glechoma hederacea (GH).
no Heukels-van der Meijden (1983).

4.1.1.
.
4.1.2.
( 4.4)
4.1.3. . :
, , = 5.0; 5.5; 6.0; 6.5; 7.0; 7.5
) 0.1; () -
. 3.17 3
. : = max / (1 - max).
4.2.
3.1,
, Frustulia
rhomboides var. saxonica (70 ).

88

 4.3.
, 3.2,
, Spongotrochus glacialis
20% 60%.
4.7.
4.1.
-
4.1.1. ( 4.3) 1 :
* 0 =(1 3 . 4 + 1 5.7 + 0 x 5 . 8 + 0 x 6 . 2 + 1 6.4 + 0 x 7 . 1 + 0 x 8 . 1 ) / ( 1 + 1 + 0 + 0 + 1 + 0
+ 0)= 15.5/3 = 5.17.
, 5.17. ,
2 6.10 3 - 6.75.
4.1.2.
( 4.4) 1 :
jc0= (1 3.4/ 1.12+ 1 5.7/0.4 2 +0 5.8/0.6 2 +...+ 0 8.1 / 1.5 2 )/(1 / 1.12+ 1 /0.4 2 +...+ 0
/1.5 2 ) = 64.03 /11.08 = 5.78. 2 6.15 3 - 6.64.
4.1.3. 3.17
() = 5.0: - 0.646; SG - 0.117; AG - 0.362; -0.106;
-0.138; HS-0.015; GH - 0.126. , /r- ,
1 -/* = 5.0 1 (,
SG, ), 0.646 0.117 (1 - 0.362) (1 - 0.106) 0.138 =
(1-0.015) (1-0.126) = 0.0051.
5.5; 6.0; 6.5; 7.0 7.5 , :
0.0244; 0.0094; 0.0008; 0.0000; 0.0000. 0.0244
= 5.5. 5.4 5.6 ,
0.1. = 5.5
1.
2 = 5.0 0.0121;
(0.083) = 6.0. 5.9 6.1.
, 2- 6.0.
3 =5.0 0.0003;
=7.0. =7.1 , =7.2 -
. , 3- 7.1.
4.2. .
3.1 loge ( Frustulia + 1) =
5.848 - 5.96 5 70, . . = loge (70 + 1 )=

89

4.263. , Srd - (5.848 4.263)/5.96 = 0.27.

4.3.
20% S.glacialis, 0.00894 temp.2 + 0.247 temp. + 2.119 = loge (20) = 2.996. temp.=
4.1 temp. = 23.8. ,
, 0.8 21.6.
, 23.8 4.1.
60% S.glacialis,
. , 48%
14. , , 14.

90

5.
. J. F. terBraak

5.1.
5.7.7.
- ,
() ,
. ",> (Goodall, 1954)
"oninung", (1930)
.
,
. -
,
, , , .
- , , , 5.1,
. ,
, ,
.
5.1 ,
. :
.

. -, ,
() . ,

, -
. ,
, , (. 5.1).
(1967). ,
.
:
( - 3),
( - 4).

. -, , -
. ,
.

91

"

1 2 . . . .1. . I . . .

1 2 . . . J[. . 1 .

. 5.1.
, . () - .

, , .
, ,
, .
, :
,
.
-,

( 3) , , -
.
-, , ,
, , , , -

92

. ,
, , , .
( )
, .
-
.
5.1.2.
(Prentice, 1977).

.
,
. , ,
() , .. ,

( ) ( 3).
.
5.2 . 5.2
. 5.2
.

. ,
(. 5.2). ,
(. 5.2d). (. 5.2,)
(. 5.2b,d). Co
, ,
, "" , ,
. ,
,
.
, , .
, -
.
, .
, (principal components analysis - ),
{correspondence-analysis - ) , ,
.
,
. ,

93

"I

I '

"I

12

5.2. (,)
() (d).
. -
b d.

(. 5.2).
, , (. 5.2).

. ( )
. , ,
.
.
, ,

.
,

( 6.2.2).
, . , ,
,
.

94

5.1.3.
5.2 ()
5.3 - (). 5.4

( 5.5). (
5.6), 5.7
.
( 5.8) ( 5.9),
.
5.2. (DCA)
5.2.1.
() - ,
(1967) ( 3.7).
; 5.2.2.
, , ,
. ,

, , . ,
5.3, , ,
, D - . , ,
. 5.3 )
D: .
,
. ,
, - , ,
( 3.7). . 5.3
, - . ,
,
. , . .
,
. ,
. ,
, ,
.
, -
. ,
? -
, (. 5.3).
.5.3 , ,

QS

A B C

l so

. 5 3 . (-)
: - ; b - - ; : - , ""
, , 2 .
= 1, D , = 0, D . .
5.2.1 5.2.3.

, , D
, 5.3.
, ,
, , - .
- ,
, -
. ,
. ,
.
,
. , .
, ;

96


( 3.28).
5.2.2.
(Hill, 1973) ,
(reciprocal averaging algorithm - RA ). -
.
,
,
, .
(. 3.9).
- ,
, ,
(. 3.9). ,
, ,
, (
4.3), .
,
.
, (1973) " "
(RA - ) ,
.
, .
, , ( )
-.
.
RA- .
. 5.1
(. 0.1),
. 1
20, () 5.1.
"" "".
.
/ >, / {
i/k, ( 3.7):
= Z**in)M xj -\

5.1

Achillea millefolium 5.1 : - (1 1 + 2 + 2 5 + 2 6 + 2

7 + 4 10 + 2 17) / (1 + 3 + 2 + 2 + 2 + 4 + 2) =117 /16 = 7.31. ,
, 5.1.
, ..

97

^Qoro^ftQoo^mo^m^^r-Ninina^cotocooNoCNi
(^'--^
r~

J8

1 0

S 3

-to

'"

i
oco

r - t - t - r - r - r - r -

c- t-

r- r-

toco

CO
CM

*- t- r-

CM
STCO

CM

CO

sr

CO CM >T

N
I3IBBS$ "si" &$

sr

1
1 t >
cv to

1
N

QCM

ffl

*n

CMC
CM
_M

T N
* CM

- ' t. JO > _J ' - ^ . - ft ^ mft i a - ^ v

< -J

-7_]_1.__::11 - -<- > CQU

_
_ 0 - * N <"- - 10 (ON

^Nn>*u^Na3m*-^c-r-c-r-c-r-r-t-NNNr><NrMCMCMCMCMC0

98

t - tO

*"*;

r - CO
r- CM

r- r*

^
cO
N

CO
CM


>

sr sr
w

win

www

3*
sr

W
-

11

<r w

8$

*-

000

8N

<

ST sr W sr
W

WW

'"

- W

< sr

-
sf

tnin
NT sr <
sr

W
WIQ
NO
~

W ST ST N 1 *-

w$

W 3

sr SJ
sr
sr

wo

000

r- ID

I?9

c
c - D Hc i-<. H
- Lt -.2.: , ui -&' lu, "t -Hube~v' 8U2Q: ) U
c rQ my e^*a5 u} ca ec i 2it

CQ

( ~
<< W N m flDOON

W - W

r ( J r NCH

oSSoSintfNNcnow^Kin^wBoSc^
" **1(0(1)-*^ ^* 1<0<0

-w
r-^
t-
-N

0000
1253

3 S3
8 ?
1 gg
VO ~
*8*

WW
W

inffl
*?

r8 N
sT sr
st

<0

CD

'"

Q sr sr M
W
j^jN

in sr

sr
N sr

IP-

sf W
sr
sf

N
5
sr N fa W
sr W r-

)
.
W.
1

cOWn

WW

E$

( <r

sr

N
(0

- -

> o-

< t

S8-

U Id CO
o.i i
CD N f f l M D Cera.
imO
CMrW~ W

ONsrintomwcncofflsr^owoo m n w w
r-NCM--

99

-CO

st W

Q sr
N
CO

sr
irf

a
s
5

*"
W
I srWsr

?m
^CnN w
<rw
S3
S 8sr W1
*8n W"l

fflN^srsfsT
. ,
UlM
N

<r

<

w
w

MP)

WW

- W - W

<~ W

Xi = Zk-\myh

ukIY*-\myki

5.2

1 5.la xi = (l 7.31 + 4 4.38 + 7 6.31 + 4

7.25 + 2 7.25) / ( 1 + 4 + 7 + 4 + 2 ) = 112,5 / 18 = 6.25. 5.16
, .
() .
, ..
"".
.
,
. , , 19-
( 1 20), 14.36 6.25 = 8.11 (. 5. lb). , ,
.
.
. 0 100
0 100
.
(*, - 6.25) / 0.0811.
5.2. .
:
1 . ( ) (<).
2 . ()
( 5.1).
3 . (<)
( 5.2).
4 . - 5.
(/), ,
.
5 . (xj). . .
6 . . ,
, - ; 2.
:
4.1. /} xj.
4.2. v = ^ Xifi/y++
> - Ek- * *~ =
4.3. ^.- **.>- */<
4.4. 4.1-4.3 .
:
5.1. z, (xt) z - *\ * / ++
5.2.
5f2 = I i e l a y + i ( X i . Z ) 2 / ^

5.3. X(,He>B.=(xi^4,.-z)/.j.
, (), 5 .

100

 ,
() 5.2.
,

5 = I*i I ++

5.3

>>*+ - , &++ - .
,
10 .
.
.
,
. 5.1

.
.
5.1. 1 19
.
.
,
.
( )

( -/-).
(.. : . 3.1)
.
, 5.2. (
) .
, -
. 5.4
.
1 19 ,
. ,
. 5.2
.
5.1.
(eigenvectors;
"eigen" - ""). - ,
.
, <& (
5.1.3). ()
, , .
(Xi), -
101

Hyp rad
Air pra

Emp nfg

Sal rep

Ant odo x

2.0

leo iut

V*c )C

X
6ra n i t

Pla l a n #

- Cal cus

Pot pal
t i e pal
* Ran f l a

14

Ach .

Jun art
Sag pro

rb:
;2

:o#

x,

Rum ace

*"-'

Tr pra
Po
#

* Pra 2

Be) per*
Bro hor
,^Lol per

2.0
# A^r

12
9 X 13
<
X

sto

Po* t r i
Jun buf

Alo $en

l y rep
Cir arv

Che lb

-2.0 +

. 5.4. .
- , - ;
5.1;
0.1.

(\) . 0 1.
0.5 .
\ = 0.53; ,2= 0.40; = 0.26; ,4= 0.17. X*
Xi .2, ,
(. 5.4).

.
, ,
.
, ( )
(. 5.2).

102

 .
5.3, D
, , .
, (1979)
, , :
I*>V [L ( (Xi - Uff}2 />>*+] /++.

,
(.. , ).
DECORANA (Hill, 1979).
5.4
".
/(1 -)/

^/(l-X)A, .

(s.d.), , 4 s.d.,
(. 5.3). s.d.
5.2.4.
, .
( 2.4.4).
( ),
: 1& ( + 1) 3.3.1.
,
. ,
+++, - +/++-
, , , 3.

(ter Braak, 1987b).
5.2.3. :
5.3
. . 5.3

. ,
( - ,
,
( ) - ,
() ,
). ,
,

103

 .
(Gifi, 1981), (Heiser, 1981;
1986).
1;
, 5.3, \ = 0.87.
, ,
. RA- ,
; - .
.
(. 5.5). -,
(. 5.3, )
( -, ) , ,

. > ,
(. 5.5). -,
, , ,
. , %2 = 0.57
(. 5.5).
" ". "-", ,
.
53, , (- ). :
. :
(* xt) : Xi - 0.87.

F
G

12 3 4 5 6 7

1
I

F
D
G

1 I
.
_
-

1| _
- I
- I

I
|
|
-

1
l l - - I
. J_
I I 1- -

"A

17 2 4 6 5 3
j

i I
M l
- I I !
I l l I l l
1 I

1 1 0 0 0 11
4 0 6 0 6 0 4
0 8 0 0 0 8 0

104

-1.40
-1.24
-1.03
-0.56
0.00
0.56
1.03
1.24
1.40

8 ]
3

-3

5.5. 5.3. : " "

( ; 5.3; ), :
( , . :
DCA, - .

" '*. , -

( 5.3) ,
( 5.2.1).
( . 5.3), ,
, ""
(. 5.3). , ,
, , D,
. ""
, ,
, , "" . ,
, ,
, . , -
- " ".
5.2,4.
(Hill & Gauch, 1980) "
" (detrended correspondence analysis - DCA) ,
: (1) ,

105

 ; (2) ,
() (. 5.5).
" ".
" " - " ,
" (Hill & Gauch, 1980). ,
- ,
.
(),
(. 5.6).
DECORANA (, 1979) .
RA-
(. 5.2).
. ,
5.3, , .. " "
.
,
, (. 5.5), -
, . (1980)
. (1980) ,
, ("
"), () .
, ,
, .
; ,
,
.
;
, .
, , .
(1980) -

1
. 5.6. .
, - .
( )
(Hill &Gauch, 1980).

106

 , (s.d.),
4 s.d. (. 3.6). -
, , ,
, 4 s.d., .
- 5.3
(. 5.5); 6 s.d.
DCA , ,
(0.53)
(0.29 0.40 ). 3.7
3.1 s.d., . , 4 s.d.,
,
. 5.1 (
DC ): 17 16
, - .
DCA (. 5.7) , , 5.4, ,

Eap nig
Air pra
Hyp *

Sal rep

Sao. pro
Leo aut
Bra rut

#
t?

Vic U t

;
2

20
Ran fla
15

X
Tn rep

Juo but
X
B
y

Ca) cus
A*r *w^<
Cht alb

X
16

t i e P*i

Alo qcn

PU tan

Rum ace
x

Wi pra

Jon art

xx

Pot pal

10
Poa pra Z
-X
Bel ptr

lot per

* X

Po* t r j

4 (s.d. - )
8ro Kor

Ely rep

. 5.7. DCA . -
(s.d).

107

. , 5.4, .
17 19 . , Achillea millefolium
, 2, 5, 6, 7 10,
5.7 1;
,
(. 5.1). (Minchin, 1987) , DCA,
DECORANA, ,
.
, / . (Pielou, 1984, . 197) ,
DCA ''
, ".
DCA - ,

.
, , , Minchiun (1987). -, ,
.
. -, " " ,
,
Hill & Gauch (1980),
(detranding-by-polinomials-DpCA).
"" (. 5.3) , -
, - -
.. (, 1974). ,
,
(JCJ), (? ) , ,
(/3). " (detrending by
segments - DCA)", DpCA ,
. DpCA
(. 5.2) 4.
^ ,
. CANOCO (ter Braak, 1987b)
.
5.2.5.
(
). ( 5.2.2)
. ,
, ,
. , , ,
, , , , . ,
DCA

108

 ( 5.2.1 5.7), ..
, ,
(. 3.14).
DCA - ,

5.7 5.1.
Juncus bufonius - 12, 8, 13, 9, 18 4;
Juncus bufonius - ,
, 9,12, 13 7. Rumex acetosa : 5, 7, 6, 10, 2 11;
( ) - 6, 5, 7, 9
12. Ranunculus flammula: - 20, 14, 15, 16,
( ) 20, 14, 15, 16, 8 13.
, .
,
, DCA.
DCA,
. ,
,
, ,
.
. , ,
, , .
, - ,
,
, .
, ,
5.1, . ,
, .
5.2.6.
,
. , ,
(),
(. 5.4).
, , .
5.4 , ( )
1,
> . , , 1,
, ,
( 5.2.3). "
''
TWTNSPAN ( 6).

109

 5.4. . 1, 2, A3 4
, , (Xi = 1, = 1,
,=1).

0
Ai

, , , .
,
,
. ,
, .
" " (down-weighting) DECORANA (, 1979) ,
, ,
,
.
5.2.7. DCA
( 5.2.1), ,
,
.
,
(), (
). - - ,
,
. ,
.
,
( 3.3.2 4.2.1), ,
, . ,
DCA ,
, .
. 3 4.


, ,
( 3.1.2)
.
( 3.3.2). ,
:
=-21& ( ). ()
(. 3.9) , . 3.3.3
- (. ). ,
. , , (. 3.6),
. -
(),
-, ..
.

( 3.4), ,
. :
(), (. 3.6), tk
, . 3.8,
Eyjfc/ = Ck exp [- 0.5( - *)2 / /k2]

5.4

*i i (
/).
, ,
(. 5.2).
1: ,.
2: (
3.4). ^ tk <\.
3:
( 4.2.1).
4: , , ,
, 2, .

( 3) ( 4) ,
. ,
-
5.4. ,
,
, .., ""
, .

111

. ,
, .

.
. , (*) -
, , ( )

(,) - ,

.
3.7, (*)

(. 3.18):
1. .

4.3 , (,)
, (.4.1):
2. () \.
A3, U ( ; , 1985).
4. * ( ; , 1985).

, ,

( , 1985). -
- (
3.3.3). , , .
, 1 2
: ,
,
. :
( 5.2.3).
. ,
. (. 5.2)
, ,
,
. ( )
(. 5.2.),
, , ;
, DECORANA (, 1979).
.
,
,
. ,
3. (),
. 5.8 -

112

. 5.8. ,
(. 5.1) .
.

- ,
5.1.
( 3.4) 0 + \ + ( 3.18).

(. 3.14).
= < (- 0.5 [ (xi1 -i* 1 ) 2 + (*2 - nk2)2] / fk2

5..5

(wk1, Uk2) -
\ -
h -
(*i\ *i2) - i .
3.14, ,
, ,
.
5.5,
- :
113

1.
, ;
2. ;
A3, ( );
4. ( ).
, ,
( 5.2.3) . 5.9
,
(40 50 ), 5.5 = 5 h= 1

4.0
\

2.0 1"

8.0

'"" 1

'
*

".

V-

i /

-4.0

4.0

. 5.9. , : (). :
, , .
( 5.5) =5, fk=l ,
[-1,9] [-0.5;4.5]. b
.

114

 .
, 8 4
(. 5.9). ,
5.9.
, DCA, . DCA
Al, A2 A3: - ,
- .
DCA
.
DCA (. 5.7) 0.1.
, 4 , -
DC-
loge = bo + 6ikr,1 + &3k*i2 + kfc (*i12 + *i22)

5.6

*il ? / DCA- 1 2, .
&4<0, 5.5 ( 3.3.3).
5.6 :
wid= -lk/ (24), 2 = - 6 2 / (24) / = 1 / V(- 2&4k).
&&> 0 ,
DC . 5.10 ,
, DCA,
20 . , ,
Achillea millefolium,
(. 5.4). 5.10
h- ,
60% *.
, (- 0.5) = 0.60.
5.10 , , Trifolium repens
( ), Bromus hordaceus -
( ).
DCA .
:
V= 1 - (2* / V ) / (Xk Ac0)

5.7

Dko Du - / (
) , ( 5.6),
. ( . 3.7).
V - , .
( . 5.10) V = (1-360/987) = 0.64.
V- 0.51 ( .5.8).

115


n t9
Air pra

Hyp ra 4

S*1 rep

Sag P>
Leo *ut

Vic U t
Cir irv
Jwn art
Tri rep

Jun bvf
P

#C1

/
* (Ml

Pot pal

Ach nil

. 5.10. ,

DCA- (. 5.7). : DCA- (. 5.7)
. : .
, 60% .

, ,
(, 3.24), .
S.3.
5.3.1.

( - principal

components analysis)

. , -

116

41

. 5.106.

, .
,
, , , ,
.

, , , ..
(. 3.1;
3.2.2). , .
, , ,
.
,
.
,
,
.
(), . , ,
.

117

 5.11 .
.
30 .
.
. , 5.11 20
20x30=600 . 5.11 - , .
. ,
, , (, Agrostis stolonifera
. 5.11); , (,
Lolium pererme . 5.11). , 0, ,
() (, Sagina procumbens . 5.11).
,
(
3.5.2).
(. 3.11),
, , -
. ,

. , -
3- . ,
, ,
. , -

. 5.11. , ,
(. 5.1). Lolium pererme
. - . ,
, .
.
.

118

 ,
- ,
. (, Rao, 1973),
. - - ,
,
, .
,
.
5.3.2.
, ,
. ,
.
,
.
.
( 3.2.2).
, ,
( 4.2.3). ,
, , , (
5.2.2),
,
, ,
. ,
,
.
,

, 3=0 (^-jc) 2 = l-
( 3.6 ,) =0 \~ 1*{ 3.2.2.
= , =0 V (* - )2 = 1 ,
. & -
i (.. ^ = 0).

fa - Eni=i }

5.8

, 5.5 , ,

, 1
20 ( ). Achillea millifolium 0.80

119

 ai

-
O

--' -* c>idr-4 t s i d d <r a cvi OJ d dtrir- d d cvi cvi ci <-' r-dcsioi

i

) i

i i

9^5(0(1(|}))|1110
ONdrr'ododrV'oodoNcirNploddrooNQNo

-
N

en? R
ON

_ . a & a ". I Q I c<- i j o i a ^ Q 4 ' Q * t - i a < > a < . ^ i . u

3
s

ffiUUIiim

>

120

mu

S H

--- sr ss N mo in J N
ininW4Plinn(JnNNNNOOOOOQOOOrrn(n(T)<nOOO
i

i I

sr

m'

ST ST ST

ST

sf sr

" &( I

NT

CMHfflN
S \11

.8
sr

N I A N-YCM

I
I
|
1

J ST

::<
- -

~\
1ST
N

-
X

3ti?c?(![!c4mjdc8:>xuu^

"

(2

N<0(T)O
PIN
BJrffl
__
IDOJsfr-in
ONsrO
r-T-r-(M-C4CNIinN^^NU3N^^^<On-NNCVINsrC)N--N

8:8888&!8? ! 81?;
nNffllONNNrprrroooOOOOOOr
I I I

1 t

N N N M N n sr

NQ

sf

WNW(\jffi
sf
rST
N
Nu -.
srsr
sr N U )
(0
sT CM NCMl/5
CM
1
N
CMN

1CM

sr

5. *

P)sr
"^.Jtnjr
CO

8 8 S

st

sf
N.

ST

CM

_ -

&

_ja.ujo.fflo:cx(ifflH-H-CLuJi-)u^cicL y j a o f i f u K O i n u
NOT-
CO
NCflCO STin tO T-CMU)
T Q N 0) 1

> -JQ^ - +> +* - * - -*

*83 8S3j5*i!t3$*5*3m8.*>Ji$*5S*J

121

sr
c- sr

- - )
- -
ON

-
-
r-N

18 8?
I
88
|

fci= (1 - 0.80)x(-0.37) + (3 - 0.80)(-.) + (0 - 0.80)(-0.29) + ... ( - 0.80)(0.37) = -1,98.

, (. 5.5, ),

( 4.2 Ok = 0).
Xj = 1>~\

5.9

4.1 .
,
, 5.6. 1 5.5
5.9 \ = (1 - 0.80) (- 1,98) + (0 - 2,40)
(1,55) + (0 - 0.25) (1,49) + ...+ (0 - 0.50) (2,29) = -0.19. ,
. 5.5

. {Lolium )
, (Eleocharis palustris)
. - (),
. ,
(. 5.5)
. 5.5
. ,
(, Lolium perenne),
, ( , Sagina procumbens) , ,
(, Agrostis stolonifera).
5.11. 5.5
, (. 5.1),
.
(. 5.6) (
5.8 5.9) (. 5.2;
5.1 5.2). , (
),
, ,
(. 5.6). , , () - -
- .
, , .
( ,
). ,
( 5.2.2).
. .
,
( 3.1 3.2). :
} = +

5.10
122

 5.6. .
:
1. (;), .
2. ()
( 5.8).
3. (XJ)
( 5.9).
4. 5 . ,
(Xj) , .
5. (xj). .
6. ,
; - 2.
:
4.1. f>
**.
4.2. = EVi *ift 4.3. -= *4.>. -
4.4. 4.1-4.3 .
:
5.1. ^=2=i *
5.2. ^1 x-UCT^Js.
, , 5 .

> - ( ) &
.
, , (. 5.5)
Agrostis stolonifera = 8.67) 6 ( = -0.31)
16 (xi6= 0.45) : 8.67 (-0.31) = -2,75 8.67 0.45 = 3.99.
Agrostis stolonifera (2,40), -0.35 6.39,
, 0 7 6 16.
5.10 (
5.3.1). ,
.
( .5.12):
= {bi! x/ + bkW) +

5.11

* xt - / 1 2.
Agrostis stolonifera 6.10 6 16 -0.17
0.033 (. 5.12), 8.67 (-0.31) +
6.10 (-0.17) = -3.72 8.67 0.45 + 6.10 0.033 = 4.10.

123

Alo 9en

Agr sio

lot pr

b-scale
X-scale

. 5.12. PC . b () , () - . , ,
(0;0).

, , -3.72 +
2,40 = -1,3 4.10 + 2,40 = 6.5, , ,

, . Xi + X2. ,
( 2) . ,
, , (1+2) / (
). R2 3.2.
\ = 471; \

344 =1598. ,

(471 + 344)/1598 =51 % . ,

471 / 1598 = 29% , - 344 / 1598 = 22%.

124

5.3.3. -

.
.
. 5.1
25 ,
. -
(25) (15). ,
, .. (25; 15), ,
( 0.0) - (25,15) .

. - (1), - ,
2. , 1
(. 5.13). 5.13 ,
.
. 2 ,
1, , 2 . 5.13,
. 5.13 1.
, 5.13 b 5.13 ;
1. (. 5.13) 1
5.13. ,
, , 5.13 ,
( )
( ).
, , . ( > 3),
-
. :
. 1 - ,

( . 5.13 = 2). ,
5.1 ,
,
(. 3.1). , 5.3.1,
PC 1 2 (. 5.11).
, ,
; 2, 1
, . ,
-
; : (
: 1,0,0, ...; : 0,1,0,0,... ..), (. 5.13).

125

-20

--|--10

--+-XXJX

10

X-

--420

5.13. 25 , :
, (
34 . : , :
, . , .

126

5.3.4.
, ,
(Gabriel, 1971).
( 5.2.5),
. ,
.
.
, 5.3, 17.9,
- 0.8. , ,
,
. ,
( ).
. ,
(:- ) (- ). 5.12 5.15
.
5.3.1 ,
(. 5.11; 5.10) ,
, (. 3.11; 5.11). ,
, .
5.12.
, .. ,
,
.
.
,
, .. (
. 5.13). , ,
, 5.1 , , .
, 5.12,
Agrostis stolonifera 5.14. -
A. stolonifera
.
5.14 , , , A.stolonifera
16, - 13 6,
. ,
.
,
,
, , -

127

X 19

. 5.14. 5.12 Agrostis stolonifera.

.

. , ,
,
. , 3 20 Agrostis
stolonifera, 2 19 , ,
. , 5.5.
,
, 5.11,
(ter Braak, 1983). .
5.12 , A.stolonifera 10
. (. 5.14).
, , , 20 0.2
. 10 0.2 = 2.
A.stolonifera (2.4), 4.4

128

A.stolonifera 20; - 5. 51 %
. 5.3.2.
, .
, , ,
, . ,
80% Agrostis stolonifera, 78% trivialis, 25% Bromus hordaceus,
4% Brachythecium rutabulum 3% Empetrum nigrum.

. 5.11 , ,
, , 1
(^1, 5&2) (*// 3, ?15).
, .
,
(. 5.6).
.
(. 5.12)
:
= cos 9
- 6 - .
, , ,
, ,
, ,
.
(Corsten & Gabriel, 1976).

, ,
.
. 5.3.3
( 5.16), ..,
, ,
, - ,
.
(ter Braak, 1983). 5.15
.

. (\ )
(bkVVXi, ^)
, i (\\ ,2)
(/ VXi, Xi2 Vfo). 5.15 5.12,
VX,i VA.2 1.

129

tri

.JQ

Alo^gen

4
X

3
X

8
X

pra
Jun buf

Lol per

S49 pro
hor

Bel per

C1r try

,.
*X JTri rep
Rum *c
Vic U t ' ; ; .

)0
X X

**

10*

1#

,.

Jn * r t .

*
,'.,,

scale
X scale

0.5

*
to

Air pre
# Sal rep

I to *ut
P U J M

Aqr *tO
*

12
X

Ely rep
*

2
X

-0.5

13

*d#

Ant odo

* rut

"

|5

X
18
X

20
X

1?
X
19
X

. 5.15. .

5.3.5.

,
(, Morrison, 1967).
.
.
.
, , ,
, , . ,
, (
). , ,
, . , "
", " ".
, ,
, , , ,
. . (Noy Meir et al., 1975)
"" ""
.

130

 , ,
.
, -
, , , .
, ,
, , , .
,
,
.
CANOCO (ter Braak, 1987b). ,
.

.
- (Swaine & Greig-Smith, 1980).
5.3.6. R- Q-
(. 5.6)
Y .
CANOCO (ter Braak, 1987b)
,
(Noy Meir, 1973). ,
, .
, .
.
5.3.6 .
,
R- .

=?* (, 1=1, ...)

, , - .
, Q-

=?

(1.7=1,..., ).

Q- ,
5.6, 5.8 5.9. 2 3
:

131

 = Enj-i Cij Xj.

, , ,

5.9 , ,
5.8. , R- Q-
, . ,
, R - , Q- .
5.4.

,
.

, .
, , , ,
, ,
(. 5.7).
:
- ,
(. 5.7);
-
(. 5.16);
- (. 5.17);
-
(. 5.8);
-
(ANOVA, 3.2.1)
(. 5.16).
, ,
. ,
(, , )
, .

.

, . .
.
,
(Dargie, 1984). - , ZJ,
z}
(xj) ( .. -

132

 5.7. ,
.
DC A. A1: 1(), 9 9 ; :
1 = 5 = ; : 1 = , 2 = , 3 = ; : 0=
5 = .
: SF - , BF - , HF - , NM . F, R, N ,
, .

(i)
00001000010111111121
12350749636162485709

fkzh mil

11
7
17
6
19
20
26
23
21
18
4
8
10
27
2
15
30
14
9
22
28
29
16
5
24
25
13

Ely
Bro
Lot
Bel
Poa
Poa
Tri
Rum
Pot
Pla
Rio
Che
Ete
Tri
Ggr
Jun
Cat
Jun
Cir
Ran
Vic
Bra
Leo
Rnt
Sag
5at
Hyp

13 242 2
2
4444 46
4 2423
756266526 47
2
3222 2
2
445244443244
3 1
27664555494 24
2 2 5
5 3 26
2
2 2
535 5 3
27
23 55 48
1
4 6 4 5 4
52262135223 3621 2
4
83 54 744 4 5
2 4 3
4
3 4
3
4 4 3
3 4
2
22 2 2 2 4
1
2
1
2222226 2444 64 43
52333223235 2252226
442 3
4 4
52 222 4
3
3 53
2
2 5
2 3
2

rep
nor
per
per
pra
tri
pre
ace
pal
Ian
gen
aLb
pal
rep
sto
buf
COS

art
arv
fla
lat
rut
aut
odo
pro
rep
rad
3 pra
12 Emp nig
_W,

J
1
2
3
4
5
6
7

F RN

R1

SF
BF
HF

8 Ni

34463344464466959444
11212124155154515255
22211321223332312111
42421341233132000000
10100010010011000000
01001000000100000000
00010101101000000000
00000000000000111111

133

4
5
5
5
7

5
8
3
7
5
7

5
10 3 2
3
2
9 7 7
4
7
10
7
6
5
7 3
8

2
7
3 2
2 3 2
5
5
6 7 6
5

5 4 3
2 3 1
6 4

 5.8. (100 )
DC

1
2
3
4
5
6
7
8

SF
BF
HF
NM

1
58
76
35
6
22
-28
-22
21

2
24
57
-21
-68
-29
-24
-26
73

3
7
7
-3
-7
5
39
-55
17

4
9
-7
-5
-64
-60
22
-14
56

0.54

0.29

0.08

0.05

* 5.7.

NN

2t

* 8F
5

L4 * .

. 5.16. , DCA 5.7.

.
DCA- .
, , ,
.

134

2.44

1.2

-*

. 5.17. DCA- ,
.

\ hi ( 3.5.2)
5.12

Zj = bo + bjxi +&

0 ,
6 = arctg (/\)
( 3.2.1).
, , (),
5.16 . ;
,
( 5.3.4).
.
3 ,
.

. ,

. , (\)

5.13

JC/ = + \2\ + 22 + + C&q

- q / .
,
( 3.2.1.) , ,
, .
5.9.

135

 5.9.
, ,
, .
*

-2,32
0.14
0.38
0.31
-0.00

0.50
0.08
0.09
0.22
0.12

-4.62
1,71
4.08
1,37
-0.01

. .
(m.s.)

4.25
0.41
1,22

10.6

Ci
2

ANOV

(df)

4
15
19

17.0
6.2
23.2

(S.S.)

R2 = 0.75
R 2 ^ = 0.66
* . 5.7.

,
, , .
, , ,
. ,
, ,
,
. ,
, ,
.
5.5.
5.5.7.
, .
,
?
,
. ,
, ,
.
.

136


, .

, .
, ,
.
,
( )
,
. ,

.
5.5.2.
(canonical correspondance
analysis - CCA) ,
( 5.2.1). ( . 5.18),
, .

, .
(. 5.2),

.
,
. 5.2.1 (),
, 5.4

( 5.13). , 5.5.1,
, ,
. ,
? ,
. 5.18 ,
(3 + 2 ) ,
.
, , ..

, = + \ZH + 22 + ... + CqZqy

5.14

Zji - / i;
\ - ( ), ;
{ - /.

137

A B C

1t~

D E

"

2 x
3 x

D E

. 5.18. ( - )
' ,
, : . : , ,
: , .
=1, D , =0 , D .

,
(. 5.18; , 1987).
, (CJ) . .

, ,
( 5.2.1).
, .
,
- ; (,)
, ( 5.2.1).
, , ,

( 5.14).

138

 (. 5.2), .
,
( 3) ( - /++
). , ,
( 5.14) , ,
, 4 5.2. ,
,
.
. , ,
.
, ,
,
. ,
, ,
.
, :
.
,
, ,
, .
,
, . q <> -1,
: . , ,
, (, 1982).
(, , 1980),
CANOCO ( , 1987b).
"" . ,
( ),
. .
5.2.7 , ,
, 1-4.
,
,
( , 1986). ,
,
.
, - .
, ,
, , , .
,
(
). 20 , 30 (. 0.1) 5

139

 (. 0.2), - ,
4 .
( 3.5.5).
: (. 5.7) ,
.
, (0.46 0.54).
(. 5.4),
0.87.
, ,
0.96, .
: ,
, . ,
,
. . 0.29,
( 0.40), 0.89, (
0.83). 5.10 ,
.
,
, . (
, R-
). 5.10 ,
, - ,
, .
(. 5.19).
5.10. :
(100 ) (100 )
.
,
. SF
"' 3.5.5).
*

SF
BF
HF
NM

9
71
25
-7

-37
-29
5
-27

57
93
21
-30
16
-37
-36
56

-17
-14
-41
-79
-70
15
-12
76

-9
18
20

16
19
92

. 5.7.

140

 ( )
, :
( 5.2.5).

. ,
. 5.19
, .

.
(. 5.18) ,
, , .
, Cirsium arvense, Alopecurus geniculates, Elymus repens, Poa trivialis
, 5.19, ;
Agrostis stolonifera Trifolium repens - ;
Ranunculus Jlammula Anthoxanthum odoratum ,
. .
5.19 , ,
( ) ().

.
, ,
5.3.4.
,
. ,
, , ,
, . 5.3.2,
({+ ) ,
.
(. 5.19) 65 %. (
1.177).

. ( 5.2.1)
s r}S V $ (1-*), rfi - /
s Xs - s.
(1986). ,
, , ,
, , ,
.

( , 1986).
. , -

141

Hyp rad
A i r pra

2.010.5

Vic U t

Sal rep

n>9

Ant odo*

PU Ian*

Pot pat
-Cal cus

Ach *

8e1

T r i pra

10.
*6
4
5x , BF
x7

0.5

-2.0

hor
Rue ace*

Lo) pr
X1

*
2*

Ran f l a
I5 K X 0.6
Jun a r t * 2 . 0

scale
u

awisture

scale
*Ele pal

Ely rep
Agr s t o
Poa t r i

Jun buf

Cir arv

-2,0|-0.5

* A l o gen
Che alb

5.19. CCA , : ,
. - , - .
,
. :
, .

,
, (. 5.19). ,

. . 5.19, (
) ,
.
( , 1986) 133
125 . ,
(DCA),
( RO.60) :
(\ (),
(, ) (, , ).
-

142

. 5.196

: R 0.82 0.81 .
- 0.34 0.25
DCA 0.20 0.13 . ,
,
DCA,
. (. 5.20) ,
(> 0.8) . ,
, ,
- .
,
.
( = 0.9). , ,
,
. .

143

 rteny t r i f

Nympalba.

RdnU

-fldm

Cole.-sp
Stra

Pote p a U
^
Ranu 1ing #
&\

w Lyco

*l'.s
eur
Acor
calal

Pota perf*
Call obtt/

Trib bont)
Bide t r i p

Call hamu*

Pota dec

. 5.20. CCA (
).

5.5.3.
(redundancy analysis - RDA) .
Pao (Rao, 1964). RDA
,
.
( 5.3.1),
.
,
(. 5.11). , , ,
. ,
5.5.2, ,
. RDA - , ,
.
,
( 5.3.1). RDA
144

 , 5.5.2:
, 3 (. 5.6),
5.13
. (
).
, RDA
.
,
; RDA - ,
, ,
.
RDA ,
, 5.5.2. 29%
21% , . RDA
RDA, , ,
26% 17% .
0.95 0.89. ,
, .
, ,
. 1 (. 5.11), , , ,
- , , - .
RDA (. 5.21)
( 5.3.4).
( ); 5.11. :
(100 )
(100 ) RDA
.
,
. SF
""
( . 5.10).

SF
BF
HF
NM

-1
15
5
-8

-5
9
-6
16

54
92
15
-26
25
-48
-40
51

-6
12
29
86
76
-11
13
-79

-10
-10
-4

0
-2
-13

145

iri

. 5.21. RDA ,
. : , 0.067
0.4 .

.
( ),

. , , ,
, , ,
, , ,
, .
trivialis, Elymus repens, Cirsium arvense 5.21,
""; ,
.
trivialis, Cirsium arvense,
. ,
, ,
, . 5.21 , , , Salix repens,

146

Hypochaeris radicata Air praecox

.
(\ + ) , , . , 5.21
71% .
5.21 (
5.3.4)
.
(. 5.11) .
s
jS V (*/), jS - j s, n -
- - s.
,
, ( 5.15),
(
. , ter Braak (19876).
5.5.4.
RDA,
{"canonical correlation analysis" - COR)
. COR ,
() ,
. COR
RDA,
(,*), : ( *,*= )
(), ( - +
Ij Cj Zji). COR 5.12.

COR. 2
RDA: RDA
, COR ,
.
: COR
. , :
, .
, , ,
. RDA , COR,
, . COR
(Gittins, 1985). COR

147

 5.12. (COR).
1. (xt) ,
.
2.
. (6*) .
3. (xf)
( 5.9). , 2.
4.
(xt*) . (})
.
5. (xj)
, j=i4 cj z#. ,
4.
6. (*0 ,
5.6.
7. (\) 5.6.
8. , , ..
;
2.

,
, RDA ( 5.5.3).
5.9.3.
COR
(. 5.12). (Tso, 1981) ,
. COR
RDA,
() ( 5.10 5.14):
RDA COR.
, ,
COR. ,
.
COR, RDA .
5.5.5.
,
(canonical variate analysis - CVA),
( COR).
, , , ,
.
, ,
. 3.2.1
. , , ,
148

 , ,
. CVA
,

(
).
, , ANOVA
( 3.2.1).
, CVA COR,
, . ,
5.12 CVA. ,
CVA ,
, (Schaafsma & van Vark 1979; Varmuza 1980).
, CVA,
. (Green, 1979), noy(Pieloy, 1984),rHTTHHca(Gittins, 1985).
CVA, RDA
, .

Alo gen
C1r arv
Che alb

21 A g r # s t o

Sag pro
'Cal c u s

Jun buf
16 X SF

Ele pal

13xSF

Ran 'fla

12XSF
SF

20> NM

I*SF

Ely rep

5 X NM

HF

3xSF

1*"M

$HF

Poa'tri

8ra
-I

rut

Air
imp
Pot
2
Sal

NH

pra
nig
pal
rep

Jun art
1X

SF

19 NM

HHF

2XBF
Be) per
Lol*p*r

18*NM

11
*BF

BF

**HF

Hyp rad

Leo aut

MF

6 XHF

10 BF

Rum a c e

T r i pra Bro hor

I
Ach mil

Tri

I
Pla Ian

rep

W*NM

I
Vic l a t

Ant odo

. 5.22. C C A ,
( . 5.7).

149


( 3.5.5. " ",
Feoly & Orloci (1979). , 5.22
.
(,1 = 0.32) (NM)
, (.2 = 0.18) (
, SF) (HF) (BF),
. , ,
, , , ,
- .. 5.22 ,
5.19: ,
.
5.5.6.
,
.
5.14,
-
. , ,
.

, .

,
.
, ,
, ,
, .
,
. .,
, ,
.
, , ,
, .
3.5.3. ,
( 5.5.2),
. -

150

,
( CANOCO
). , .
,
.
. , ,
.
COR CVA
RDA. , RDA
, .
5.5.7.

RDA. 5.2.2.
5.3.5. COR CVA
, .
,
, .

(Loucks, 1962)
(Austin et al. 1984).
, 5.5.6.
5.6. ()
5.1 ,
, ,
, ,
- , .
.
, ( )
( 6.2.2),

.
().
, ,

. , , ,

. .
, ,

151

 , ,
.
,
.
-, - ,
5.3.3. - 52 i ;
52 = ++ Y*-\m Ob/v+i - ydy+j? />*+,

5.15

6
V = 2W" (* -jVkj)2

5.16

-
, - .

, , ,
.

(), (Gower, 1966; Pielou, 1977,
.290-395). , , ..
.
8
Cij = - 0.5 (5ij2 - 6i+2//i - cVj2/w + &-2/2)

5.17

+ .
Q - ( 5.3.6).
,
Q - .
( )

. ,
. , ,
-
( 5.2.7); ,
<
.

(1962) (1964).
. (5),
/
.
,
. , 152

 (Barlow et al., 1972).

(, 1
).
, ,
. , ,
.
.
,
. ,
,
, (Prentice, 1977).
- .
, ,
.
,

.
.
(Sibson, 1972). (Prentice 1977; 1980)
:
si} = 5* min (,

5.18

(Kendall, 1971) , ,
,
.

5.7.

5.13 , 3, 4 5
.
. (
COR) ,
. , -
. ,
, (
, ).
(GLM, 3),
.
, .
,
, ( 3.7),
. , DCA .

153

 5.13. ,
. MR - ; IR - ,
- , RDA- ; COR -
; WAE - ; GLM - ; ML-
; WAI - ; -
; DCA - ; - ;
DCCA - env.vars. - ; . grad.
- : , .

()

(env.vars.)

MR
IR

WAE, GLM, ML
WAI, ML
CA,DCA,ML

>1*
>1

RDA

CCA, DCCA, ML

COR

CCA, ML

(. grad)
I

, , WAE, WAI ML.

5.13, ,
. ()
,
.
, DCA, ,

(Clyrao, 1980; Gauch et al., 1981).
, (. 3.3),
,
.
( 3.2.3). , ,
, DCA . . , 2
, ( ) ,
RDA. RDA ,
,
, DCA (D)CCA, ,
( , 4 ).
,
3 , DCA
(. 5.7; 5.15).
DCA
( 5.2.5 5.3.5).

154

 5.13 , ,
, . ,
,
,
, .
, .
, ,
. ( 6)
, .
,
.
,
( 3.1.1. 5.1.1).
,
- .
,
.
,
. ,
. ,
, ,
. , ,
, ;
-
,
( 3.5.4). -
, ,
,
. ,
. .
, ; ,
-
- .
, , ,
,
- , .
5.5.2,
.
, ,
. , ,
. ,
(q) ().

155

 ,
1, , (
2 3.2.1). ,
, q -1.
, ,
,
. , , ,
DCA ,
,
. (D)CA
, , .
, ,
. , ,
.

, ..
,
() .
, .. ,
, .
. ( 5.3.5)
.
-
. ,
, .
, , , ,
,
.
.
.
.
,
. RDA
(Davies & Tso, 1982) (1988).
CANOCO ( , 1987b).
, -
.
5.8.
( &
Curtis, 1957; Gauch, 1982), (Beals, 1985).
. . (Mardia et al., 1979),
(Goodall, 1954) (Orloci, 1966).

156

 , 1935 , ,
. -
, ,
(Gifi, 1981, Gittins, 1985;
Greenacre, 1984; Nishisato, 1980). (Benzecri et al., 1973)
.
. (1973) ( 5.2.2).
5.2.1
- (2),
(Torgerson, 1958, 12.7) (Nishisato, 1980, . 23). RDA
(Israels 1984): (Rao, 1964),
PC , (Davies & Tso, 1982). (1986) CCA. COR 1935
(Gittins, 1985). (Campbell & Atchley, 1981)
CVA, (Williams, 1983)
. Gauch et al. (1974), Kooijman (1977), Goodall &
Johnson (1982), ,
,
.
, ,
. & van Groenewoud (1984) ter Braak
(1985) .
(Shepard, 1962) (Kniskal,
1964), a Schiffinan . (1981) . Meulman & Heiser
(1984) .
Anderson (1971),
Noy-Meir (1974), Austin (1976), Fasham (1977), Clymo (1980), Prentice (1977; 1980).
, ,
(Kruskal & Caroll, 1969; Dale 1975; de Sarbo & Rao,
1984; Heiser, 1987), Scmffinan et al., (1981)
, .
, , (Heiser, 1987).

,
(Gordon, 1981; Mardia et al., 1979; Greenacre, 1984; Rao, 1973; Gittins, 1985).
(Pielou, 1984). , 5,
( 5.9).
(, Gourlay &
Watson, 1973), , 5.6.
5,
, , .
, ,
. (1979) (1987b)
.
157

 5.2 5.6 (Gifi, 1981)

(Everitt, 1984).
(Wold, 1982).
, COR, CVA
. DCA DECORANA (
1979). CANOCO ( , 1987b)
DECORANA , , RDA, CCA, CVA
. ( 5.9).
GENSTAT (Alvey et al., 1977) SAS
(SAS Institute Inc., 1982), , ,
. . (Schiftman
et al., 1981) .
5 .
(Carroll, 1972)
( 5.11) ( 5.5)
( 3.24)
: , , (
Davison, 1983).
. , ,
(Prentice, 1980).
.
5.9.

. :
-
;
- , 5,
;
- ,

.
, 5.8.
5.9.1. ()
, Y =" {} ,
() (). -
, ..
, >\+ = 0 ( 5.3.1).
(singular-value decomposition, SVD) Y (Rao, 1973; Mardia et al., 1979;
Greenacre, 1984).
Y = P A 05 Q*

5.19
158

 Q -

, , = min

(,), .. F P = 1 Q'Q = I - A* (s = 1 ... ),

\^22.

...Z 0.

Q Y, >* 0 5 s -
Y. s - p s , ( m), a s Q qs, ( ), 5.19
Y= ^

X*05 ps q'

5.20

Y 5.11 5.3.2.
5.20
b, = X / 5 p , H x f = q, ( s = l , 2 ) .
- bi ( 6*i),a i- -xi -
(-*) . , \
.
( 5.3.4).
5.11:
Y
(Gabriel, 1971; 1978). 5.20 , 2
+ ... *, ,
S M [ ^ - ( ^ 1 X l l + ^ 2 X l 2 )] 2 =X 3 +?l4+... + Xr.
, , ( \ +
) . = I, Q'Q = I 5.20

b, = Yx,

5.21

8 = Y'bs.

5.22

, ,
' (. 5.6
5.3.2). 5.21 5.22 , bs $ -
YY' Y'Y, , , X - ;
R- Q- 5.3.6.
YY' \
5.19 , Q'Q = I. YY'
, , bib'i + '. YY' / (-1)
, 5 ,
(Corsten & Gabriel, 1976; ter Braak, 1983).

159

 Y'Y - Q AQ\
Y Y'Y, , bs 5 $=8 = X*0 5qs.

, 5.16;
.
Y'Y 5.17 ,
, .
5.9.2. ()
Y >, 0,
. = diag (+) - m x r a
, Y; N = diag () - ,
Y.
5.2.1,
, ,
( 5.1).
= (i*k) [ = 1,..., ] :
= "!Yx

5.23

:
8 = u'Mu / x'Nx = x'Y'M ]Yx / x'Nx

5.24

(. 5.2).
8

Y'M^Yx = X N

5.25

8 = X (Rao, 1973, lf.2 .74; Mardia et al., 1979, 9.2).

, , -
( = = 1). , ,
8 l'Nx = 0. ,
5.2, , ,
5.25, .. .
5.25
, . 5.25

X = 1 Y'u

5.26

160

 5.23 5.26 . ,
,
(. 5.2 . 5.1.3).
SVD
"05 Y N -0'5 = 05 Q'

5.27

us = *0,5 "5 $ xs = N^ 5qs, 8 qs s -

Q, (s = 1,..., ).
, is 5.23 5.26
. 5.2.7 , *
us x s , , , ( 5.2.2).
PC Y 5.27
, .

5.9.3.

5.9.1., Y - , /-
(.. + = 0) Z q ,
- (.. 25+ = 0).

S,2 = YZ', Sn = YY', S22 = ZZ' S21 = S'i2

5.28

COR b = (bk) [k =
1,..., m] = (CJ) = 1,..., q],
* = Y'b = Z'c. , b
:
S12 S22"1 S21 b = X SH b

5.29

S2i Si 1 Snc = X S22

5.30

X (Rao 1973; dia et al., 1979; Gittins, 1985).

, b

b = (YY')'1 Yx = Sif1 Sn

5.31

, , * ,
b

161

 = (ZZ') l Zx* = S221 S21 b

5.32

b 5.31 5.32
5.32 5.31, b 5.31 5.32
5.29 5.30. , .
5.31 5.32. 5.31 5.32
5.12. 7 (. . 5.12) :
X (. 5.6). , .,
i = Z'c ( 5).
, 5.12,
*, .
COR SVD
Sn* 5 S12 S2205 = 05 Q'

5.33

5.31 5.33
5.33 S 1i"0,5 Q
= Sn--5 P 05 = S22 ~'5 Q.

5.34

s -
, . 5.32 5.33
.
COR ,
.
, :
- ,

Si2.
Sn Sn, .
t G, -bqxt

S,i--5(Si2-GH')S220-51

= I S,,-0-5 S12 SM- 5 4SII' 0 - 5 GXS22'05Hy |

5.35

G , II II - (,

|| Y|| 2 =2W).
SVD ( 5.9.1) , , Sn*05 G fe"0"5
t 05 Q 5.33, .

162

 G
t Sn 0 5 05 S2205 Q, .
Q. ,
Si, 05 05 = S n Sn"03 05 = S n = YY'B = YX

5.36

S2205 Q = S22 S22"05 Q = S22 = ZX

5.37

X = Z'C. ' = I, , ,

(
, ). ,
, Si2
. ,
,
.
5.9.4. (RDA)
RDA Sn 5.9.3
(Rao, 1973, .594-595).
RDA, 5.5.3, S220 5Q 05, .
5.9.5.
5.24. Z ,
5.14 = Z'c, = () [/' = 0, 1,..., q]. = Z'c 5.24
(Y ):
S12 = YZ', Sn = = diag (yk+) S22 = ZNZ'

5.38

= ' S21 Sn"1 S12 / ' SM

5.39

, ,
5.30 (Sn, Sn S22 5.38).
d = (1, 0, 0, ..., 0), X = 1, = 1
8 TNx = TNZ'c = 0, 8
.
* = % y+t zJf I y++ (

 Z). Z
, 0: % y+i $ =0.
5.33 5.34
5.38.
5.5.2,
5.2. 1, 4, 5 6 ,
5.2.
b = 1 Y

5.40

x" = N l Y'b

5.41

36 = (ZNZ')1 ZN *

5.42

= Z'c

5.43

b = u - , Wk ( = 1,..., ).
, ,
= Z'c, ,,
(. 5.32). 5.42 (
5.32). , 5.30,
5.42, * 5.41
5.42, b 5.40, 5.43 , ,
S11, S12 S22 .
,

mxq
W = -1 YZ',
(kj) -
() j.
(+),
S22 ,
5 W S22"05 = Si I*05 S12 SM"0 5

5.44

5.44 5.33
W = (S,, 0 5 ) 0'5 (S2205 Q ) '

5.45

( 5.2.2),
, , -

164

 Su",5PA0-5 S2205 Q, . Sn"05 P 05

5.34. ,
, :
S225 Q = S22 S22-05 Q = S22 = ZNZ'C = ZNX

5.46

5.10.
5.1. :

(.5.2) , .

1
0
0
3

0
2
0

0
1
0
0

0
1
1

0
1
0
1

,
.
5.1.1 - 3, - 5.1.4 - 6.
5.1.1. 1, 2, ..., 5
.
, 5.2.
5.1.2.
(. 5.2).
. ( -
, - s).
, ,
4 .
. ,
.
5.1.3. 19 ()
0.101; -1.527; 1.998; -0.524; 1.113. (
)
. ? , 5.1
, 5.2 - .
5.2 , .

165

 5.1.4. ,
5.1.2.
, 5.2,
( - , VH s - ,
- ).
5.1.5.
. (
).
5.1.6. .
. ?
5.2. .
5.2.1.
. Hippophae rhamnoides,
9, 18 19 1, 2 1, .
. 5.1 .
.
? . 5.1?
annua, 1, 2, 3, 4, 7, 9, 10, 11, 13 18 3, 3, 6, 4, 2,
2, 3, 2, 3 4, , Ranunculus acris, 5, 6, 7,
9,14 15 2, 3 , 2 , 2 , 1 1, .
5.2.2. ,
.. , Bellis perennis, Poa
pmtensis, Rumex acetosa 5, 4 3,
( ). (: ,
5.1.3). ,
, ,
5.1. .
5.2.3. 5.1
, 5.4.
5.3.

5.2 PC 5.5.
. ?
5.5?

166

 5.4.
, DC
, 1.5 .
,
?
, 10 ?
5.5.
5.5.1.
Juncus bufonius 5.7, 5.15 5.1.
Eleocharis palustris.
5.5.2. 5.15
DCA, , ,
5.7 PC ,
. : ,
, , ?
5.6.
(Cramer, 1986)
. 1978 1984 .
135 1 2 .
. ,
( 5 ) ?
, - .
, , ,
63 ( ). 68
1000 .
,
(DCCA) , : 1984
( ) (0 1978, 6 1984).
-14 56 . 0.56 0.10, 4.4
0.9 , 0.95 0.74,
. 5.14 ,
,
.
.
5.6.1. 5.14 ,
, , 5.5.2* - :
= 0.054*; + 0.041z2

5.47

. (..)

167

 z\- (), z-i- () .

5.14.
: (100 ) (100 ) . .
(s.d.) ().

()
()

100(3)
12(3)

4(4)
-34(3)

99
7

19
-99

s.d

22
3

18.5
2,9

: , 5.14
, .. , z\* = (z\ -22)/
18.5 HZ 2 * = ( Z 2 - 3 ) / 2 , 9 .

, \ = 0.054 0.0016
= 0.041 0.010.
5.6.2. 5.47
(. 5.8; 5.18),
, .
5.47
1984 15 25 .
?
1978, 5.47?
5.6.3. 15 1984 5.47
1978 1984. ?
5.6.4. 5.47 ,
.
5.6.5. ,
?
5.6.6.
, ,
? : .
, ,
? : .

168

5.11.
5.1. :
5.1.1. - = (4 1 + 2 x 2 + 1 x 3 +
3 x 4 + 2 x 5 ) / 1 2 = 2,750, s2 = [4 (1 - 2.750)2 + ... + 2 (5 - 2.750)2] /
12 = 2.353, , = 1.5343.
\= (1- 2.750) / 1.5343 = -1.141.
5.15.
5.1.2. 2 , ,
[2 (-0.489) + 1 0.815)] / (2+1) = - 0.054, 5 3
(0.815 - 0.228) / (1 + 1) = 0.294. 5 =
(2 0.163 2 + 2 0.815 2 + 3 0.054 2 + 5 0.228 2) / 12 = 0/138.
5.15. ( 5.2.3).
5.1.3. , 19 20
, ; , (. 5.15) .
\ = 0.7799, s, .
5.2 '^" () .
, 3 - ,
20- s = 0.7799, "" .
s X, ,
X [ = 1*~ / Ek-i"1 *
3, (4) 5 ( xi), ,
. , 8 X
.
5.1.4. 4.2 : v= [4 (-1.141) 0.10 + 2 (-0.489) (-0.53)
+ 1 0.163 2.00 + 0.815 (-0.53) +2 1.466 1.11]/12 = 0.2771
1 4.3, -1.141 -0.277 1 0.10 = - 1.169.
5.16.
5.1.5. 5.16.
5.1.6. V, 1
2 3 . ( 5.2.3)
.
5.2.1. .
5.2.1. 5.1
. Hippophae rhamnoides [1 0.09 + 2

169

 5.15.
5.1. ( 1) ( 2).
. 1 - , 2 -
( 5.2), 3 : - - . 4 (5), (s).

-0.163

0.815

-0.54

0.228

(2)

(4)

(3)

0)

1.841

-0.283

-0.325

(4)

(3)

(2)

1 868

-0.465

-0.238

(4)

(2)

(3)

0
0
1
1
1
2
2
2
3
3
3
4
4
4
5
5
5

1
5
2
3
5
2
3
5
2
3
5
2
3
5
2
3
5

1.5343
0.1375

1.000
-1.141
-0.163

2.000
-0.489
0.815

3.000
0.163
-0.054

4.000
0.815
-0.228

5.000
1.466

-0.212
-0.704

-0.054
-0.179

0.815
2.706

-0.148
-0.491

0.294
0.976

-0.598
-0.393
-0.567

-0.283
-0.408

1.841
2.646

-0.402
-0.580

0.758
1.089

(1)
-0.574

-0.322
-0.448

-0.465
-0.646

1.868
2.597

-0.426
-0.592

0.815
1.113

)
-0.520

1.865

-0.628

-0.161

-0.251
-0.340

-0.628
-0.851

1.865
2.526

-0.436
-0.591

0.852
1.154

(2)

(4)

(3)

-0.466

1.840

-0.764

-0.091

-0.185
-0.246

-0.764
-1.015

1.840
2.444

0.440
-0.584

0.875
1.162

(2)

(4)

(1)

(3)

(3)

0)

(5)

(2)

(4)

-0.211

1.556

-1.193

0.178

0.081

-1.193

1.556

-0.409

0.867

(2)

(4)

(3)

0.104

-1.530

1.995

-0.524

1.112

(3)

(1)

(5)

(2)

(4)

X
U
X
X
U

0.3012
0.6885

X
X
U

0.6953
0.7171

X
X
U

0.7193
0.7342

X
X
U

0.7383
0.7498

X
X

20

20

20

0.7529
0.7606

0.7800

0.7799

(-0.31) + 1 (-0.68)] / (1+ 2+1) = -0.30, annua -0/33 Ranunculus acris -0/19.
5.1 Elymus repens Leontodon autumnalis.
Hippophae rhamnoides annua
- Ranunculus acris.
, .
, . ,
, ,
( 5.2.5).
5.2.2. [3 (-0.65) + 5 (-0.50)
+ 4 (-0.39)] / (3 + 5 + 4) = 0.50.
5.1.3 X (=0.536), , -0.93. -

170

 5.16.
5.1, .
(/).
( 3) ( 4). 5
- v ( . . 5.2), , 5.15.

1 2
0
0
0
0
1
1
1
1
2
2
2
2
3
3
3
3
4
4
4
4

4.1
4.1
4.3
5.3
2
3
4
5
2
3
4
5
2
3
4
5
2
3
4
5

0.10
-1.141
-1.169
-1.216

-1.53
0.489
0.065
-0.068

2.00
0.163
0.391
-0.407

-0.53
0.815
0.962
1.001

1.11
1.466
1.158
1.205

0.0001

-0.243
-0.243
-1.114

0.288
0.288
1.320

0.399
0.399
1.829

-0.036
-0.036
-0.165

0.056
0.056
0.257

-0.0011

-0.647
-0.647
-1.084

0.825
0.823
1.379

1.043
1.045
1.751

-0.155
-0.156
-0.261

0.197
0.198
0.332

-0.0014

-0.645
-0.645
-1.078

0.832
0.830
1.387

1.042
1.045
1.747

-0.159
-0.160
-0.267

0.203
0.205
0.343

-.0016

-0.642
-0.642
-1.073

0.836
0.834
1.393

1.045
1.048
1.751

-0.156
-0.157
-0.262

0.206
0.208
0.348

f
X

0.2771

X
X
U

0.9612
0.0837

X
X
X
U

0.2182
0.5956

X
X
X
U

0.5967
0.5980

X
X
X
U

0.5982
0.5984

X
X
X

0.5985

-0.107

-0.399

0.288

-0 288

-0.639

1.043

0.825

-0.650

-0.673

1.042

0.832

-0.636

-0.672

1.045

0.836

-0.632

,
5 (. 5.4).
5.2.3. 5.1.
^(\-)/
JX(l-X)

= ^/(0.464/0.536) = 0.93
= ^/(0536 0.464) = 050 ( 5.2.2). 20, ,

1.95 / 0.93 = 2.10 Juncus articulatus 1.28 / 0.50 = 2.56.

5.2, 5.1
5.2 5.1.3.
5.3.
Hippophae rhamnoides 0.2. 5.8
(0 - 0.2) (-0.31) + (0 -0.2) (-0.30) + ... + (2-0.2) (-0.04) + (1-0.2) 0.00 + ... + (00.2) 0.45 = - 0.03. -3.22 annua -1.48
Ranunculus acris.
. ,
( 5.3.1).

171

 5.5. , annua Bromus hordaceus.

( 5.9) 3.90 / 471 = 0.008.
5.4. DCA
DCA ,
1 1. , ,
, 4 (. 3.6;
. 5.3). 1.5 ,
. ,
, ,
(. 3.3). 10 ,
,
.
5.5. .
5.5.1. Juncus bufortius,
5.7 (DCA): 12>8>13>9>4 18;
5.15 (): 13 3 > 4 > 19 12;
5.1: 9 = 12> 13 > 7 - Eleocharis palustris
5.7 (DCA): 16 > 14 15 > 20 > 8
5.15 (): 16 > 20 > 15 > 14 > 19
5.1: 16 > 1 5 > 8 = 14 = 20.
5.5.2. ,
. DCA
,

. , ,
, .
, , DCA -
; ,
(Eleocharis palustris .5.7).
5.6.
5.6.1. 5.14 ,
- 1.00 zi* + 0.12 zi*. = 1.00
(zi -22) / 18.5 + 0.12 (22 - 3) /2.9 = (1.00 /18.5) + (0.12 /2.9]z2 - 22/ 18.5 - 0.12 3 / 2.9 =
0.054 z\ + 0.041 z2 - 1.31. \ 0.03/18.5 = 0.00162 2
0.03/2.9 = 0.010.

172

 5.6.2. JC 15 1984
= 0.054 15 + 0.041 6 = 1.056 . 25
= 0.054 25 + 0.041 6 = 1.596
. , 1.596 - 1.056 = 0.54
. 5.47
: 0.054 (25 - 15) = 0.54
, . ,
10 , 0.54 .
5.6.3. 15 1984 1.056
1984 ( 5.6.2) 0.054 15 + 0.041 0 = 0.81
1978. 1.056 - 0.81 = 0.246
0.041 6 s.d., .. . ,
6 .
5.6.4. 0.041
, ,
0.054 . ,
0.041 ,
0.041/0.054 = 0.76 . 95 %
\ ,
(Finney, 1964). .
, (0.4 , 1.1 ).
5.6.5. 5.6.4 ,
, 0.76
. ( 0.5 ) , ,
, .
, . ,
0 ,
. ,
.
5.6.6. 4.4 .
5.6.3 , 0.25 6 .
, ,
(4.4/0.25) 6 100 . 0.9 ,
, ,
. ,
-0.34/2.9 = - 0.117. , 0.117 6 s.d.=
0.70 ,
0.25 . ,
, .

173

.
. F. R. van Tongeren

6.1.
6.

, . . - :
, , .
, , -

, (Westhoff &
vander Maarel, 1978; Bering, 1957).
.
, , : , -
, , ,
.

.
:
- ( ),
- ( ),
- , ,

( ).
6 .
, , 1
. - .
- ,
. ,
;
.

174

6.1.2.

,
. , ,
(Lance & Williams, 1967; Dunn & Everitt, 1982).
.
1
. ,
. ,
,
" ".
, ,
: . ,
,
, , . ""
.
6.3, - 6.2.
.
6.2.3.
- .
,
. ,
, ,
.
.
:
- : , ,
, ;
- :
" ";
- : ,
.
6.2.
6.2.1.

. ,

. , ""

{(
. (. ).

175

 .
. .
6.2.2.

: MOiyr ,
. , ,
, .

(
). . ,
. , ,
. , ,
. , ,
.
,
, .

. ,

, ,
.
, , -
.

.
-

(, ),
.
, ,
, , ,
. , .
, (Jaccard, 1912). ,
, :
SJ = c/(a + b + c)

6.1

176


SJ- ;
- , ;
- .
:
SJ = c/(A +B-c)

6.2

- , - :
= + = + .
(Serensen, 1948) ,
().
= 2/(+)2/(

+ + 2)

6.3

,
. (Faith, 1983)
-.


(.. ); ,
, 3.4.
-
.
, : (Ball, 1966)
(, Gauch, 1982).
:
SRV = 1* /(I* * 2 + Sb yif - ? )

6.4

yh - - /', j .
- (0 - , 1 - ),
6.1, .. .
( 6.3) :
PSy = 2001*\(

,) /(Zkyh + *>^

6.5

min (. - %.
.
( ,

177

. 6.1. (1-5) , , .
1 .
2 , - , . 3 4
. 5 .

). (. 6.1),
(. 5.3.3).
,
(. 6.1),
. (ED)
:
6.6

0=^*(-#)*

6.1 ,
: 1 2,
, , 1 3,
.
,
, (. 6.2). ,
- . ^
.
( Ochiai):

cos = OS = /

Vbytfay2

6.7

, ,
, :

178

. . , 6.1. . , ( ).
, 2* , 3".
1 3 .

CD-

Vl*\yb/(bykt2)in-yb/(Zky*2)in]2

6.8

,
( 6.2).


. , , ,
.
, ,
, ,
, .
:
- .
. ( & tiss, 1957), ,
- . ,
(IA) .
,
, PD :
6.9

PD = 1A-PS

179

 , . ,
, .
6.2,3.
(, (Williams et al., 1966; Hajdu,
1982 ), .
,
, . (Dann
& Everitt, 1982) (Sneath & Socal, 1973)
, ,
.
, - , ,
.
(Lambert & Dale, 1964; Greig-Smith, 1971).
, , (
?)
( ?).
-
,
. ,
, , , .

(), ,
(. 6.1-6.4).
(. 6.3-6.6) 0 1. 6.1. ( ).
10, (-),
.

10

1.00
1.00
1.00
1.00
1.00
1.00
1.00

1.25
1.25
1.25
1.25
1.25
1.25
1.25

1.00
1.00
1.00

1.25

I
J

1.67
1.67
1.67
1.67
1.67
1.67

1.43
1.43
1.43
1.43
1.43
1.43
1.43

180

2.00
2.00
2.00
2.00
2.00

2.50
2.50
2.50
2.50

3.33
3.33
3.33

5.00
5.00

10.00

Jacc
SR
PS

. 63. ()
6.1. .
, .

6.2. ( ).
,
.

10

I
J

1
1
1
1
1
1
1
1
1
1

Jacc cos

2
2
2
2
2
2
2
2
2
2

3
3
3
3
3
3
3
3
3
3

4
4
4
4
4
4
4
4
4
4

5
5
5
5
5
5
5
5
5
5

6
6
6
6
6
6
6
6
6
6

7
7
7
7
7
7
7
7
7
7

8
8
8
8
8
8
8
8
8
8

9
9
9
9
9
9
9
9
9
9

10
10
10
10
10
10
10
10
10
10

. 6.4. ()
6.2. .
, (),
.

181

 6. ( ). !
(2) (10) ,
.

1
9

2
8

3
7

4
6

5
5

6
4

7
3

8
2

9
1

fli

. 6.5. ()
6.3.
. 6.4:
.

6.4. ( ).
.
.

1
1
1

F
G

1
1
1
1
1

I
J

1
1
1
1

182

 .
6.1 - 6.4 6.3 - 6.6,
. 6.5 , , ,
.

6.6. ()
6.3. .
(ED)
( ), : . : .

6.5. .
: - ; + ; ++ +++
.

! 1

"-

|
SR
PS
Cos
SJ

<

>

++

++

*
>
>

+
++
+

CD
PD
ED
ED2

*
*
*

*
*
*

++
++
+++

183

++
+
+

+
+
++
+++

++
+

~
+
++
+++

6.2.4. ,
,
, .
, ,
.

.
:
, , -
.

( ) * = loge ( + 1)

6.10

:
- ( 2.4.4);
- ,
;
-
.

* =

6.11

:
- , (,
, - );
- .

* = ^

6.12

1, .
. .
, ,
. ,
.
- (1/0).
184

 ,
, .
( -
) .

2.4.4.
, .

( ) ()
. -
:
.

.
"" ( ,
). ,
,
: , ,
.
.

. .
. ,
.
.
- .
, .
,
, .
,

.
. ,
, (
) .
, ,
.
(,
).
.
,
.
.
185


.
.
. ,
. ,
,
() .
, .
( ), (
; ,
, , , ,
. .).
.
6.2.5.

() ( ) .
, ,
.
( )
( , , . 6.8).
.

()
(. 6.7).
,
, 6.8.

'

6.7. ( ) : ,
, .

186

0.2+

0.4

0.61

O.Bf

1.01

10 5

I I 18 t? 13 lb <!0 16

1 14 17 19

. 6.8. ,
.

. 6.9.
.

, ,
.. .
, .
,
, :
(. 6.9).

187


()
.
,
(. 6.10). (. 6.11)
; , 6.8,
, .
:
, .

6.10. ( ) : ,
, .

0.0

.
0.4f

0.6

0.8}

6.11. ,
.

188

0.0

0.2

0.4f

0.6

0.8

1.0J
i

io s 6

/ n i J 4

8 9 12 u 14 is 16 w

. 6.12. ,
.

( ).

(). ,
( , Sneath & Socal, 1973). (. 6.12 ),
.
(UPGMA - unweighted-pair groups method, Socal &
Michener, 1958). ,
(Lance & Williams, 1967).


. ,
,
(. 6.13). 6.14
. ,
, ,
(. 6.12).

189

 6.13. ( ) : ,
, ;
.

<><>

6.14. . ,
, .
.
, , .

190


, (Orloci, 1967), .
,
,
.
, ,
, ,
. :

, .
,
( 3.2.1,
), (dE) :
Ep+q Eq ,

- ;
Ep+q - ,
q\
- q.
:
, = / / # ! , * *(;->* )2>
, -
.
,
,
, 6.15.
6.3.
6.3.1.
,
,
, .
(Williams & Lambert, 1960) -
. -

191

riifl

10 4

( I I

1/

I M

4 ]

N IS 20 It

12

1 1 10 S 7 I I ! 17 19 J

1 I? U IS 16

25j
201

16}

12{

I I U I / 19 2 10 *

3 9

U 14 W 16

192

 . ,
(, Kirkpatrick et al., 1985).
6.2.3.
, (Roux & Roux, 1967)
, . .
(, 1976b) ,
, .
(Hill et al. 1975; Hill, 1979) , .

, 6.3.3.
6.3.2.
( , 1959; 1960; 1961)
, .
.
. , ,
: , - .
,
. :
- . .
, , ,
, , ,
(Coetzee & Werger, 1975; Hill et al., 1975).
6.3.3.

(Two Way Indicator Spieces Analysis - TWTNSPAN). TWINSPAN,
(1976b) ,
.
,
TWINSPAN. , ,
.

. 6.15. ,
: (), ()
(). () () .
. ,
, \* .

193


TWINSPAN ,
, ,
,
. , , ,
(Hill et al., 1975)
, (. 3.4).
.
, .
- .

(Heiser, 1981). ,
,
(. 3.4), , ,
.
;
.
TWINSPAN, , (, 1973; 5.2)
.
() ()
. ,
(Hogeweg, 1976)
(Gower, 1974), (Hill, 1977).
.
.

. (,
) .
,
, - .
() , , ,
.
.
,
( 5, 5.9). ,
1.
(
, 5, 5.2).
. ,
.
, .
194

 1,
-1 1.
, , ,
(. Hill, 1979b). "" ,
. ( ,
), . -
,
, .
(1979b) < ",
TWTNSPAN, ,
TWINSPAN, . "
"
(, POLYDIV (Williams, 1976 b)).

:
.
, - ,
. 6.16.
, 4, 5, 6 7 .
.
, 8 9,10 -11 ..
(, 10 11) 11,10, 11
4, 10 , , 3, 10.
(10, 11). ,
, .. ,
.

/ \
4

/ \
5

// 1\ \\

10

6.16, TWINSPAN
TWINSPAN.

12

13

14

15

195

 6.6. TWINSPAN .
TWINSPAN, ,
, 1, 2, 3,..., 9.
.

1111

111112
17895670123489234560

12
13
28
5
16
1
26

7
3
11
17
19
23
16
20
27
29
4
24
25
2
10
21
22
30
14
8
15

Pir

Hyp
Vic
Rnt
Pla
Och
Tri
Bel
Bro
Cir
Ely
Lot
Poa
Rum
Leo
Poa
Tri
Bra
PLo
5ag
Sal
Rgr
Ele
Pot
Ran
Cal
Jun
Che
Jun

nig
rad
lat
odo
Ian
mil
pea
per
hor
arv
rep
per
pra
ace
aut
tri
r*p
rut
gen
pro
rep
to
pal
pal
fla
cue
art
alb
buf

.2.3
00000

... 2
000
22.S
00000
2. 1.... 1
00000
.4 . 44324
00001
323.5553
00010
.2. .222413
000110
2S2
000110
.2.2..2.322
000111
2.24.4.3
000111
2
000111
. . . .4. . .4444.6
001
7.2.2666756542
001
413.2344445444.2
001
....563
22
001
52563333.522322222.2 01
645427654549. .2. 01
3.222526.521233261. . 01
4.632622..22224..444 01
2725385. .4. 10
2. .3
52242
10
. .33
5 10
48434S447S 110
4. ..4564 11100
22.. 11100
2. .22224 11100
4.33 11100
44. .334 11101
1
1111
2
443
1111
00000000000011111111
00001111111100001111
00001111

TWINSPAN

.
. .
,
, " "
. TWINSPAN,
, 6.6.

196

6.4.
,
,
. . (Gauch, 1979)

, .
COMPCLUS ,
. ,
, , .
(Janssen, 1975) ,
, .
CLUSLA (Louppen, van der Maarel, 1979). -
,
.
.
, .
CLUSLA ""
. :
, ,
, . COMPCLUS, CLUSLA
,
. , , FLEXCLUS
(van Tongeren, 1986).
.
.

. , , ,
.
, , (Ben-zekri, 1973), (Salton) (Wong, 1978) (Swain, 1978).
- -
, , ,
.
,
. , ,
, .
- ,
. , 6.17.
,
-
-.
TABORD (van der Maarel et al., 1978), CLUSLA (Louppen, van der Maarel, 1979)
FLEXCLUS (van Tongeren, 1986).

197

6.17. , : . :
, : .
.
.

6.5.
,
, . .
.
.
.
- (Westhoff, van der Maarel, 1978),
, ,
, .
- ,
, ..
(. , 3.3.1).
- ,
, .
, ,
. ,
, .
,
, . .
: ,
, .
( Popma et al., 1983)
:
- ( );
- (
- ).

198

 .
, ,
. ,
,
(Hogeweg, 1976; Popma et al., 1983).
6.6.
.
:
- -,
.

.
- - ,
.
, -
, .
. .
- (Braun-Blanquet)

/ . 8.3
. 8.3 I V
(0-20, 20-40, 40-60, 60-80, 80-100%, ).
. ,
-

.
-
.
.
(. . 8.7).
6.7.
,
,
. :

- .
.
- ( 6.7.1)
( 6.7.2).

199

 6.7. FLEXCLUS , ,
.
1, ,
.
.

11:1112;
11 1 11 J
79617065! 123469: 23:4560:
Leo aut
Bra rut
Tri rep
Rgr sto
Qch mil
nt odo
Pta Ian
Poa pra
Lot per
Bel per
Ely rep
Rio gen
Poa tri
SaQ pro
Jun buf
Jun art
Cal cus
4e pal

! 265535 52232!
! 3942262; 2222 i
5 2S32622J 521231
4843:
!
;
; 2 2 24 2! 13
:443 24 4!
J 2 535335;
! 1 3444321 44S44 I
t 676622! 75642 ;
222; 322 J
i
5
4i 4444 !
27253!
!
4 54 6J 276545:
J
! 2
; 522.
4 :
2

45!44751
4 : 444;
32:61 :
4S:447S:

2 !

as! 4 :
49;

2 :

42;

:
43 :
. 434 ;
I 433 ;
14584;
4
2 . 212224:

44

Ran
Qir
Bro
Hyp
Pot
Rum
Sat
Tri
Vic
Che

Ha
pra 23
hor '
24 2 4 3
ad 26 2
pal
ace
6 3 5
2
rep
3
3
pra
5 2 2
lat
2 11
alb
Cir arv
;
2

:22

2 s
J

:
5j

1:

Emp nig I 2

:
* OKpy^l

; 4.0 : .:5.! 7.s!

:
i.i
; .:0.1s 3.6:
% rF
:
: :o : 0 :
Ni
:
; :0 J 75 :

cl
cl
cl
cl
cl

1
2
3
4
5

: *****
: **

i *
: **

; *
: *

; * ;
j *

;****:

cl
cl
cl
cl

1
2

: *****
: **
: *

:
;
:
i

*
*
*
***

: * :*

200

6.7.1.

, ,
"" - . 6.7

1. , 1.

.
,
,
. 6.7 ,
, .

. , ,

, 4,
.

. ,

, DISCRIM (ter Braak, 1982; 1986). DISCRIM
,
. 6.18 TWINSPAN
(. 6.6) , ,
DISCRIM.

1(12)

0(4)

3(4)

2+

7(8)

(8)

3+

0(4)

3(4)

0(4)

6.18. TWINSPAN, 6.6,

. , DISCRIM.
.

201

. 7.2.
3 ( 3.2). , ,
- (, ,
, ),
, .
, -
.

3.2.1.
.

( ), () -
.
.
, , (. 2.4.4).
-
(. 3.3.1)
, .
,
(. 3.3.1).
.
t - (. 3.2.1) "
" . .
, ,
. , (Wilcoxon),
- (Mann-Whitney),
.
(
, ) () (
1).
.
.
(/). 7^ :
= ;( ; +,+ 1 ) - 7 ,

202

 ]2- , .
Tj 7 ,
, Z
:

-(|-)-03)/

// = (/1 7 +2+1)/2= /( / , 2 ^/).

(
4 4, 3 5, 2 8) -
.
(, Snedecor & Cochran, 1980).
6.8.
(von Hum
boldt, 1807). . (1912) ,
. (Sorensen, 1948),
, .

. ,
, , (Socal & Sneath, 1963).
70-
( ). (Pielou, 1969), (Goodall, 1970) (Wil
liams, 1976) .
, (Goo
dall, 1973) - (Mueller-Dombois & Ellenberg, 1974).

(Everitt, 1980) (Dunn & Everitt, 1982). (Gauch, 1982)

.

203

6.9.

6.1.

. 1.
, . *
1,
(. . 6.1 ).

2
1

1
2

1
1
3
1

1
1
3

4
1

5
3
1

. lb. .

. 1. :

F
G

0
0
0.60
0.40
0.60
0.60

0
0
0.33
0.25
0

0.25
0
0
0.25

0.17
0.33
1.0

0.75
0.17

0.33

. Id
. .

204

 6.2.

. 2. ( *)
( ) 6.1.

2
3
4
5
6

29
40
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25
58
60

17
60
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63

29

6.2. .
6.3.
6.3. ( *) 6.1.
1
2
3
4
5
6

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4.9
2.5
6.3

4.2
5.7
3.8
4.7

5.3
2.5
6.3

4.9
2.5

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6.1.
6.4.
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. TWINSPAN (Hogeweg, 1976). 6.1.
. - .
, - .
1, 3 6 5, 2 4
.
. (SPOS
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205

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g. b-f .

. a-g .

6.5.

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5 () :

6.5. TWINSPAN (. 6.6)

6.5. 6.7 ()
( ). ?
? - ?

6.6.

1 TWINSPAN
(. 6.6).

206

6.10.

6.1.

. 1. () (+-)
( 6.2.2) :

( )

4
2
3
3

1
3
2

0
2

5
6
5
5

6
4
6

6
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3
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6
2
3
4
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6

(+-)

2
2
0
2
0
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5
6
4
6

: (+-)

2
3
4
5
6

0.33
0.40
0.00
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0.00

0.80
0.33
0.06
0.60

0.17
0.75
0.40

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0.20

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:

1
2
3
4
5

<

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3,5
2,6
1,5
4,6

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(2,3,5),6
1.(2,3,5,6)
(1,2,3,5,6),4

0.80
0.75
0.60
0.50
0.50

207

 . I .

1
2
3
4
5
6

1.0
0.75
0.60
0.60
0.33
0.25

D,G
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208

 6.2.
.
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PSW: 2:10
3: 6
: = 1,B0,C0,D1,E0,F 1.G-1
: 1 + 0 + 0 + 1 + 0 + 1 + 1 - 4
PS u = 200x4/(10 + 6) = 50
PS3.6:ce 1+0 + 0 + 0 + 0+ 1=2
PS3<6-200x2/(6+10)=25
PS4.5: : 0 PS^ e 0
:

1 2
2
3
4
5
6

29
40
0
50
0

50
25
58
60

17
60
25

0
63

29

6.2. ,
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2
3
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5

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29
40
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50

50
25')
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17")
25

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")mm(17,25)=17
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2: 5 2, 58
:

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3
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1(2,5)

40
0

29')
50")
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17
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209

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:
1

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(2,3,5)

29

(4,6)

0)

4 : (2,3,5) 1, - 29
: (1,2,3,5) (4,6), - 0

..
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EDi 3 = [(0 - 1)2+ (0 - 0) 2 + (1 - 0) 2 + (2 - 1) 2 + (0 - 0) 2 + (0 - 1) 2 + (1 - 3 ) 2 ]
= (1+0+1 + 1+0+1+4)

= 8 = 2.8

ED5,6= [(2 - 0) 2 + (0 - 0) 2 + (0 - 5) 2 + (0 - 3) 2 + 91 - ) 2 ] 1/2 = (4 + 1 + 25 + 9 + 1) 1/2 =

= 401/2 = 6.3

.. ,
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1
2
3
4
5

2.5
2.5
2.5
4.2
4.7

1,5
3,5
4,6
23
2,6

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4,6
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4 6 - .
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210

 6.4.
a-g. - ( 6.8)
6.1.
6.8. -, 6.1
SPOS, SNEG PREF, b 6.4.

1
2

Jb

_2

Jb

SPOS SNEG

PREF

SPOSSNEG

PREF

SPOS SNEG

PREF

1
1
2

3
1
1
1

1
1
4
3
1

2
3
5

1
4
1

2
2
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1

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2
8
7
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1
4
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0.75
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1

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lb-1. 6.8.
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2-2 . 6.8.
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.
2. MID - (-1.73 + 8.81)/2 = 3.56.
2f. 1,3 5 , - .
-. . 6.8.
3d 1-6 : -3.17; 2.15; -1.68; 5.75; 0.76 7.91
.
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3f. To , 2f, .
h. .

6.5.

():
TWINSPAN (. 6.6)

: 1+2
3+4+5

0
11
1
12

1
0
8
8

211

11
9
20

FLEXCLUS(Ta6.6.7)
1+2+3
11
5
16

4
0
4
4

11
9
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6.6
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1
4.4
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*> = 4.84 (1 / 8.8 + 1 / 2.2 + 1 / 7.2 + 1 /1.8) = 6.11 (FLEXCLUS, ).
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17

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1.5

7.5

7.5

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10

12.

12.

14

15

16

17

18

19

10 12 10.5 12.5, . \ (
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212

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P. A. Burrogh

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215

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216

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217

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7.7

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7.8

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7.9

:
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7.10

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7.11

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7.12

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7.13

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7.14

(. .).

225

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.

(h) = 1 11 * , - Z(*y)]2 f[2(n-h)]

7.15

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C(0)-C(h)n
r(h) = 1-yW 1 (0)

7.16

,
.
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,
().
, 7.11.
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2.
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= 2.5
3. () 1
( 1 ) = [ ( 3 - 3 ) ( 1 - 3 ) + ( 4 - 3 ) ( 3 . 3 ) + (6-3)(4-3) + (4-3)(6-3) + (3-3)(4-3) + (1-3)
(3 - 3) + (2 - 3) (1 - 3)] / 7 = 8 / 7 = 1 /143.
226

7.11. .

1, (1) = 1.143 / 2.5 = 0.46. 2

(?(2)=[(4-3)(1-3) + (-3)(3-3) + (4.3)(4-3) + ( 3 . 3 ) ( 6 . 3 ) + (1-3)(4-3) + (2-3)

( 3 - 3 ) ] / 6 = - 3 / 6 = -0.5.
2 , (2) = - 0.5 / 2.5 = - 0.2.
4. 1
(1)= [(1 - 3)2+ (3 - 4)2 + (4 - 6) 2 + (6 - 4) 2 + (4 - 3) 2 + (3 -1) 2 + (1 - 2)2] / ( 2 7) = 19 / 1 4 =
1.357
2
( 2 ) = [ ( 1 - 4 ) 2 + ( 3 - 6 ) 2 + ( 4 - 4 ) 2 + ( 6 - 3 ) 2 + ( 4 - 1 ) 2 + ( 3 - 2 ) 2 ] / ( 2 6 ) = 3 7 / 1 2 = 3.
.
7.3.5.


(Wagenin, Swifterbant, Flevopold, Netherlands, ten Berge
., 1983). ( )
1500 26 19... 13.00. ,
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229

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7.18

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7.19

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7.20

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7.21

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7.22

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7.23
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rf

7.26

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238

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7.27

%= 1.
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.
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Ij-in yvjf(xu xj) + v|/ = (*/,*),

7.28

( - Z { / (xit XQ} -
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239

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.

\ - ( ,
). 7.28 5^in wt = 1 +1
+ 1 wt
(i=]t...,n) \\f .
:
2= Zj!11 Wj (xj,x0) + v|/

7.29

(1985) 0)
.
,
,
.

240

. 7.19. , 7.18, .

.
. </

.

241

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( 7.28 7.29) ,
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.
,
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,
, (. 7.18-7.19).
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Z ( - ).
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, (Burrough,
1986). 7.18b-7.19b
. ,
, , ,
(. 7.18; 7.18d).
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, Z
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= faS21 (g-) 2

7.30

,
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242

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. :
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- .
= 9 1
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, .

|
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. 7.21.
.

243

s.e.

7.22. . :
. : .

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,
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,
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,
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.

244

7.6.
. -,

. -, ,
,
,
.

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.

. , 1930-1940- , ,
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,
, , (Matheron, 1971)

245

 ,
(. David, 1977 Joumel &
Huibregts, 1978).
(1984,1985) , (Nielsen &
, 1985).
- Agterberg (1982),
- Burrough (1986), - Lam (1983)
- Ripley (1982).
, ,
PASCAL 16- , MS-DOS
PC-DOS. ,
(Geographical Institute, Utrecht University, P.O. Box 80.115, 3508
TC Utrecht, the Netherlands).
7.8.
7.1. ,
7.1 ,
20 , 100 . :
- ;
-
().
7.1. ()
20 , 100
.

1
2
3
4
5
6
7
8
9
10

12
13
14
15
16
17
18
19
20

5.2
5.4
6.1
3.1
5.4
6.4
7.2
6.5
4.5
3.3
2.2
5.5
6.2
6.6
5.5
4.3
6.3
5.5
4.2
4.8

8.3
9.8
11.2
12.2
14.2
13.2
12.4
11.5

8.7
9.8
10.5
12.6
13.5
14.9
13.8
14.3
15.7
16.7
15.4
17.2

246

 . ,

. , 5%-
18 0.444.

.
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.
7.2.
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. 25 , 25 .
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7.2.

- -3
.

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

-3

29
28
29
33
38
33
36
28
25
24
22
23
30
34
38
40
46
44
51
45
46
43
36
32
25

20
22
21
21
32
36
22
16
24
27
26
31
29
38
35
19
27
62
52
61
39
34
32
26
22

247

- ,
- .
,
.
7.9.
7.1, ,
7.3 , ,
. 7.4 .
, ,
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7.5 .

(. 7.6). 7.7,
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2 . ,
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7 3 . .

1
2

5.21
12.80

1.30
2.57

2.20
8.30

7.20
17.20

7.4. .

1
2

1.171
2.177
2.781
2.484
1.762
1.640
1.390
1.952
1.959
1.492

0.993
2.289
4.022
5.525
5.811
5.626
6.024
6.490
7.670
8.211

3
4
5
6
7
8
9
10

248

 7.5. .

i
2
3
4
5
6
7
8
9
10

0.151
-0.130
-0.287
-0.217
0.024
0.083
0.134
-0.020
0.021
0.031

0.346
0.237
0.090
-0.020
0.026
0.015
0.001
0.001
-0.101
-0.148

7.6. .

1
2

-0.015
0.330

5.363
9.332

-0.067
0.760
2

1
Z

Z-Z

Z-Z

5.200

5.349

-0.149

8.300

9.661

-1.361

5.400

5.334

0.066

9.800

9.991

-0.191

6.100

5.319

0.781

11.200

10.321

0.879

3.100

5.305

-2.205

12.200

10.651

1.549

5.400

5.290

0.110

14.200

10.981

3.219

6.400

5.276

1.124

13.200

11.311

1.889

7.200

5.261

1.939

12.400

11.641

0.759
-0.470

6.500

5.246

1.254

11.500

11.970

4.500

5.232

-0.732

8.700

12.300

-3.600

12.630

-2.830

10

3.300

5.317

-1.917

9.800

11

2.200

5.203

-3.003

10.500

12.960

-2.460

12

5.500

5.188

0.312

12.600

13.290

-0.690

13

6.200

5.174

1.026

13.500

13.620

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14

6.600

5.159

1.441

14.990

13.949

0.951

15

5.500

5.144

0.356

13.800

14.279

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16

4.300

5.130

-0.830

14.300

14.609

-0.309

17

6.300

5.115

1.185

15.700

14.939

0.761

18

5.500

5.101

0.399

16.700

15.269

1.431

19

4.200

5.086

-0.886

15.400

15.599

0.199

20

4.800

5.071

-0.271

17.200

15.929

1.271

249

 7.7.
(.. . 7.6).

1
2
3
4
5
6
7
8
9
10

1.171
2.174
2.777

0.893
1.975
3.347

2.490
1.765
1.638
1.383
1.941
1.957
1.474

4.457
4.576
4.164
3.948
3.661
3.278
2.666

7.2.
7.8. .
1 - 95% 2,
2 140% 2, .
1 - 6 150 . 2 - 3 75 .
,
, , 7.23.

. 7.23. .

250

 7.8. (% 2)

-3

1
2
3
4
5
6
7
8
9
10

10.208
22.696
44.659
64.452
79.225
93.974
95.500
100.176
95.906
92.767

59.542
110.826
142.432
157.095
162.425
140.684
147.389
99.000
122.031
178.300

251

8. :

8.1. ,
, , -

P. A. Burrough
8,1.1.
, , -
. ,

.
25x10 ,
(. 8.1). ,
, 5 .
,
2500 1500 - .
, ,
600 , .
, ,
:
, - .

. :

.
,
,

(Beadle, 1954).
1970- , ,
. ,
. (1977),
,
.
- - , ,
.
,
, , 55
.

252

. 8.1. ,
.

.
. 18
, 30 ( ).
111 18 .
, 8 ,
. 8.2.
.

253

. 8.2. ,
.

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, 1-
( 5.2) .
( 45.7%
) , ,
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254

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tx

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( 55 ); b - ( 55 ); (18 ); d - (8 ).
; ,
, ,
55 .

255

 8.3 ,
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( ,
).
, .
R2 (. 8.3) ,
.
8.1.2.
,
8.3,
. 8.4,
,
, .
,
.
, .

. 8.4.
. 1: , 1
( , ). 2: 1 .
3: 1 . 4: , 1
( , ). ,
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(. 8.3).

256

 ,
, - ,
200 , , . ,
, , , , ,
; -, ,
, .

. (Budderoo)
, .
-
.
, ,
, ,
(Wianamatta Shales).
8.1.3.
,

, ,
.
, , ,
, . ,
,
, .
,

, ().

257

8.2.
-
J. . F. Oudhof & A.Barendregt
8.2. L
1981 .
.
200 250 - .
,
.. 1966 .: 135 100 - .
, (-, , 1974),

(-, 1979) , , .
,
,
-, 700

, 600 .
, , ,
- Ulicion
minoris P. Duvign (1944). ,
- ,
.
, , ,
.. (300 ).
, , 1000
. ,
.
8.2.2.
,
.
.
.
.
.
-
.

258

 , ,
.
, , .
5 2 200 2. , ,
.
, -
(, 1975).
.
.
,
, .
,
.
(Tutin et al., 1964-1980), - (Corley et al.,
1981); - (Grolle, 1976); -
. (Hawksworth et al., 1980).
8 , 4
. ()
X, ,
.
. ,
0 9 ( , 19766, .
2.4.2.), .
,
.
, , ,
,
,
.
. ,
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TWINSPAN (Hill, 1976b), DECORANA (Hill, 1979a), FLEXCLUS
(van Tongeren, 1986). -
3000 1700 .
8.2.3.
(. . 8.5):
- ,
;
- ;
- .

259

150

,
*

!
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1

1

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.

260

,
, ,
. ,
,
. ,
( , 5.2) DECORANA
(, 1979).
.
(. 8.6) 150 , , ,
,
.
. 150

. , , .

TWINSPAN (, 1979b). , -

-40 - -

.,<- 300 400 .500 600

700 800 900 1000 1100 1200 1300 1400

!
V-'
.

. 8.6. 3000 .

261

 , 1000 .
, 600-700
.
, .
. , ,
,
Erica umbellata L., .
, ,
. 500 ,
FLEXCLUS ( , 1986).


FLEXCLUS. - ,
, ,
,
. ,
, ,
.
,
.
.
. .
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,
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211 .
:
-
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;
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,
. 262

 -,
.

211 (. 6.6)
- (1964)
, ,
.
.
.
.
,
(
,
).
8.2.4.
8.7.
211 . 8
.
-, , Erica tetralix Sphagnum sp.
-
. , , -,
.

, () (
), Erica austmlis ( ) Colluna vul
garis Erica cinerea .
Ulex .
, .
8.1.
8.2.5.
,

.
, ,
(. 8.5). , ,
. .
, , , ,
.
? 9 , , -

263

 , 3000 ,
211 ,
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, . ,
.
.
8.1. , 8
. - , I - , F - , S - ,
-.
:

E,I,F,S,P

a)

1,F,S,P

b)

E.J,F,S,P

1]

a)

( )

b)

( )

c)

( )

E,F,S

III Ulexgalii E,F,S

IV Calluna- Erica cinema

E,F

Erica tetralix
Molinia caerulea
(Erica ciliaris)
Narthecium ossifragum
Drosera rotundifolia
Sphagnum species
Carex binervis
Ulex gallii
Erica cinerea
Holcus lanatus
Festuca rubra
Lotus corniculatus
Daucus carota
Scilla verna
Plantago coronopus
Armeria maridma
Leontodon taraxacoides
Cirsium filipendulum
Erica vagans
Brachypodium pinnatum
Ulex gallii
Erica ciliaris

Calluna vulgaris
Erica cinerea.
Ulex minor.

,

,
I.
cupressiforme
Dicranum scoparium
Pleurozium schreberi
Pseudoscleropodium purum

264

Erica umbellata - Ulex minor

VI
Erica umbellata.
Chamaespartium

S.P

VII
Erica umbellata - Halimium oevnoides

VIII

Erica australis

S,P

Erica umbellata
Halimium alysoides
Ulex minor
Erica umbellata
Halimium alysoides
Ulex micrantha
Chamaespartium tridentatum
Genista triacanthos
Tuberaria globular ifolia
Erica umbellata
Halimium alysoides
Halimium ocymoides
Erica australis
Chamaespartium tridentatum
Erica australis
Deschampsia Jlexuosa
Vaccinium myrtillus

*-

8.7. 211 . 1- -, 2 - - .
. : , , , . ,
, .

265

8.3.
, (Veluwe, the Netherlands)
R. H. G. Jongman, . J. van de Nes
8.3.1.
:
, .
.
- .
, .
.
, , .
, ,
, . ,
. 60-100 .
.
.
. -
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1, (. 8.8). ,
, , ,
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,
(Ijzerman, 1979). ,
.
40 .
17 (donk, 1968). -
. 19
. ,
, .
.

.

.
, - .
1950- ., ,
, . -

266

"

8.8.

. - , -
1968 1978 (Kant, 1982).
,
, ..
. -
, .
-
. , ,
.

, 30 .
.
1981
(Sprengen en Beken op de Veluwe).
(Jongman & van de Nes, 1982):
-
;
- ,
;
-
;
- ,
, .
.

.
.
267

8.3.2.

,
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. : , , ,
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.
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.
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. 181 93
.
8.3.3.
.
,
. 1 999.
, Polycelisfelina Tubifex.
,
. , (.
8.2), (. 2.1),
.
, , .
8. ,
(Cuppen & Oosterloo, 1980).

1-3
4-10
11-20
21-40
50

1
2
3
4
5

268

- ;
- ,
, ;
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vareigatus, Lumbricus sp. Lumbriculidae, )
. 24 ,
475. .
,
( ).
.
.
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8.3.4.

. ,
Chironomus (), Melius (A), Gammarus (G). ,
Chironomus , Asellus - Gammarus (, 1982). Gammarus. -

: *<

fiSSa"
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CZ3

CZZ)

. 8.9. Gammarus :
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269

 8 3 . Gammarus. (I-V)
- , .
: , , .

G3
57
21.5

G5
20
21.4

GI
19
20.3

Gammarus pulex
Prodiamesa olivacea
Dicranota spp.

V 4.45
IV 1.54
HI 0.32

IV 1.66. Y_JL33
0.29 IV 0.79
HI 0.44 III 0.27

Paraiendipes spp.
Paratanytarsus spp.

III 0.67
11 0.28

Herpobdellu octoculata
Dendrocoelum lacieum
Gammarus fossarum

IV 0.88
II 0.19
III 1.50

Pofyceiis felina
Dixa macula t a
Agapetus /uscipes
Eusimutium costaium
Micropterna sequax
Simulium ornatum
Thaumaleidae spp.
Crunoecia irrorata
Elmis aenea
Tipulidae spp.
Baetis vernus
Psychodidae spp.
Krenopefopia spp.

G2
26
26.7

G4
31
19.3

V 2.36
V 2.01
II 0.09

II 0.13
HI 0.29
II 0.08

0.06
11 0.07
IV 2.25
V
IV
HI
11
11
1!
II
II
11
II
11
II
11

2.26
1.19
0.33
0.32
0.15
0.14
0.14
0.13
0.10
0.10
0.06
0.06
0.06

V 1.46
111 0.49
II 0.04

Heiodes spp.
Sericostoma spp.
Haiesus spp.

II 0.10
II 0.07
11 0.25
11 0.28
11 0.12

Anabolia nervosa
Lymnaea perega

11 0 14
II 0.09
111
III
II
11
11
II
11
11

Tubificidae spp.
Paraiendipes gr. albimanus
Micropsectra irivialb
SiempellineUa brevis
Sialis/uliginosa
Ceraiopogonidae spp.
Rheocricotopus gr. /uscipes
Phaenopseclra spp.

270

0.48
0.46
0.37
021
0.14
0.09
0.08
0.06

111
II
III
11
11
II
11
II
II

LimnephWdae spp.
Corynoneura spp.
Tanytursus spp.
Agabus spp.
Heiophorus brevipalpis
Zavrelimyia spp.
Heterotanyiarsus apicalis
Heterotrissicladus marcidus
Polypedilium gr. nubucuhsa
Micropsecira gr. praecox

V 4.25

Procladius spp.

III 0.47

Hydracarina spp.

III 0.56

11 0.15

V 2.75
11 0 38
11 0.06

II 0.06
II 0.31
II 0.12

Ghssiphon'w

III 0.55

11 0.07

Sialis luturia

IV 1.24

Chironomus spp.

II 0.28

Chaetopieryx spp.
Pisidium spp.
PolyceJis tenuis/nigra

III 0.36
III 0.50
11 028

Asellus aqualicus
Umnephilus iunatus
Macropeiopia spp.

IV 1.57
IV 1.01
IV 0.65

/ spp.

11 0.10
II 0.16
11 0.16
21 0.05
III 0.27

Conchapelopia spp.
Apsecirotanypus trifascipennis
Nemoura cinerea

Anacaena globulus
Orthocladiinae spp.
r///m modesta
Plectrocnemia conspersa

III 0.39

111 0.58
II 0.12
II 0.10

11 0.11
II 0.06
II 0.29
HI 0.22
III 0.35

271

II

0.13

II 0.09

11 0.06

IV 1.92

II 0.30

III 0.22
III 0.48
111 0.74

V 2.14
IV 1.21
IV 0.93

II 0.21
11 0.26
V 2.73

III 0.63
11 0.26
II 0.07

III 0.46
11 0.12
11 0.12

111 0.38
III 0.75
11 0.23

HI 0.26
IV 0.78
II 0.25

II 0.18
11 0.10
IV 0.99

IV
II
II
HI
11

HI
11
II
II
II

IV

II
III
IV

11 0.24

Micropsecira spp.

0.12
0.64
0.85
0.10
0.10
0.34
0.33
0.36
0.25

III 0.63

Oligochaeia spp.
Eukurfferiella gr. discoloripes
Limnophila spp.
complanata

II
III
111
11
11
II
II
II
11

11 0.10

II 0.08

Lumbriculidae spp.

0.46
0.06
0.60
0.15
0.07
0.24
0.12
0.18
0.29

0.68
0.09
0.11
0.21
0.13

0.31
0.05
0.20
0.19
0.10

0.81
0.08
0.33
0.28
1.34

 ( , 6.2.5)
(G1-G5). ,
, 8.3. ,
20%.
.
Gammarus ,
8.9. G1
. G5 - . G4, , . G2 G3
, G3
, G2. -, G3 G1 G2.
G1 . G5 - ,
. G2, G3, G4
. G1
, G3 .
8.3.5.

. ,
.
,
:
- ?
- ?

(van de Nes, 1980).
, (Saaty, 1977),
GELPAM (Gelders Policy Analysis Model),
, .
(Ancot & van de Nes, 1981).
,
. ,
(1982), ,
: , (
), ,
.
:
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,
, .
,
,
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. , G3 Gammarus
. , Chironomus .
,

.
, .
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.
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.
8.4. . 1
(), 4 - .
, ; ,
.

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2
3
4
5

N-Horsth. beck (N)

Tongercnse bcek
M-Heerder bcek (0)
Stccnbcck
M-Horsth. beck

0,0603
0.0509
0.0486
0.0481
0.0479

0.9127
0.8349
0.9535
0.5499
0.9513

6 26 Ecrbcckse beck (W)

7 16 Beck Ordcrvccn (Z)
8 18 Koppelsprengen
9
5 Z-Heerder beck
10 7 N~Horslh. beck (Z)
11 21 Oosierhuizensprcng
12 2 N-Heerdcr beck
13 9 E-Horsth. bcek
14 19 Ugchelense bcek
15 12 Hartense molenbeek
16
1 Molecaiense beck

0.0458
0.0455
0.0450
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:
- . . .: . 1967. 356.
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292

, , ,
.

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)

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214,224,225
226
227,247

autocovariance
autocovaruogram
autocorrellogram (see also
autocvariogram)
autocorrelation
agglomerative clustering

()

93-94,116-132, 144, 152, 154-160,

166, 193
152,158

()

(.
)

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7, 225-229, 245,248
175, 186-191
97,100, 101, 106,108,138,165,
170
119,123,131,144-145

150
7,215-222,254, 257
93, 95-116, 154, 157,160-161, 165166,194,254,261
93,105-109, 115-117,154, 158,
167, 263, 265

two-way weighted averaging

algorithm
two-way weighted summation
algorithm
PCA, see principal
components analysis
principal component analysis
(PCA)
principal coordinate analysis
(PCO)
RDA, see redundancy analysis
analysis of concentration
trend-surface analysis
correspondence analysis (CA)

191-193
191-196

detrended correspondence
analysis
power, see statistical power
analysis
association analysis
divisive methods

229,234-235,247
80
93
147, 159, 162,164

anisotropy
posterior distribution
data approximation
least squares approximations

80,86
18-19,31,33,216
84,104, 177

prior distribution
skewed distribution
asymmetry between presence
and absence
autoecology
binary variable (see also
dummy variable)
binominal distribution
biomonitoring
biotic index
block structure
block kriging
vector
vector model
| probability for occurrence
weight (see also indicator
weight)
weights for species or sites

1,2
268
19,20,45,157
3,24,78
78,83,87
109-110
242
101,125,159,178-179,223
158
43-44,79, 82, 84, 88-89,109
122,137,239-240,273
103,130-131,164, 184-186
7.Q3

97,157, 254

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)

122, 137, 145, 147, 159

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62,83,97,100,139,141,160, 164,
194
61-64, 68, 83-88,93, 97,112, 154
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174

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4, 7-8,10, 15
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33, 41-43,48, 50,68, 73, 84,112113

33,48-49, 84
53,56, 57
110-115, 139,157
52,113,116
41-43,48-50,52-59
19
230
118, 122, 125,254
64-65, 155
4,91
213
191
5,62,97-102,107,113,115-117,
120-124,127-130, 133-136,139,
145-146,149-150,166,187-193,
196,200-201
8, 18, 31, 43-49, 53-62, 65,79-85,
97,104,112,142,176-177,184
157
32,60,109,170
193-196
186-192,199
30, 33, 37, 45, 67
63, 97, 103, 196
18
201
79, 87,148
35,37, 40
33-37,58,148,202,214
37,130,225
230-233,237,242, 244
37
106,154,167

294

reciprocal averaging (see also

correspondence analysis)
weighted sum (see also linear
combination)
weighted average
weighted averaging
weighted moving averages
species composition (see also
community composition)
internal data structure (see
also indirect gradient analysis)
within-group sum of squares
choice of methods
selection of variables
sample
bulk sample
outlier
Gaussian response curve
Gaussian curve, see Gaussian
response curve
| Gaussian logit curve (see also
logit regression)
Gaussian logit surface
Gaussian ordination
Gaussian response surface
Gaussian regression
geometric mean
intrinsic hypothesis
principal component
gradient
gradient analysis (see also
regression and ordination)
boundaries
minimum variance clustering
Dune Meadow Data

presence-absence data
dual scaling
bimodal curve
Two Way INdicator SPecies
ANalysis
dendrogram
systematic part of response
model
diagonal structure
discret variable
discriminant functions
[ oMscriminant analysis (see also
canonical variate analysis)
variance ratio
analysis of variance
variance
nugget variance
residual variance
| length of ordination axes






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53
96,97,112,118,125,127,129,
132,139,145,159

95,97,112,118,122,137,155,
159, 166
175
239
245
234
176-177,181-183,204
178, 1883
177,181-183
61
78, 195
17,179
23-26, 53-56, 71, 132, 150, 156,
167,199-203,227,235-237
8-9,60,61,63-65, 156-157,174,
191,203,245
46,100,111,123,129,145,148,
152, 157,162,165, 194,197
5,29,78-90,91,111,119,154,155
6,93,136-151,154,155
148-149, 151, 157
137-144, 149-151, 154-157, 163165, 167
147-148,151,154,157,161-163
139, 140, 145, 150-151,162,167,
168
141
213
242
3,6,174,213,245
258
8,10,23,174,258
152
4,6, 8, 23, 155,174-212, 258,269
131
146,159,225,229
17,37,44,65,81,213

91
47, 158,193,197,201, 245-246
60
220,247
131, 163
157

295

confidence interval
dominant species
supplementary point
complementary variables
site score
score, see site score and
species score
species score
hierarchical methods
isolines
isoplcths
isotropy
Jaccard index
cosine coefficient
Sorensen coefficient
species - indicator value (see
also indicator value)
indicator species
interval scale
interpolation, see spatial
interpolation
interpretation of results
historical notes
iterative algorithm
calibration
cannonical ordination
cannonical variate analysis
(CVA)
cannonical correspondence
analysis (CCA)
canonical correlation analysis
(COR)
canonical coefficient
cannonical eigenvalue (see
also eigenvalue)
choropleth map
error map
mapping
classification system
classification (see also cluster
analysis)
classical scaling
cluster analysis
covariance matrix
covariance
quantitative variable
bell-shaped response curve
(see also unimodal response
curve)
component analysis (see also
principal component analysis)
computer programs
competition between species
autocorrelated residuals
correlation matrix
correlation ratio

139,140, 145,147, 156,167

, .

24,26,37, 40,81, 129,132,147

19
37
156,256

37,40,217

14,22,29, 34-35, 40,44,49, 53,

156, 198,202

, .

(.
)

(.
)





-
-
-

202
274

DCA

177,181-183
37
108,221
7,239-242, 245
32, 60, 94

43-44,60,69,202
35,40,72,75
215-216,219,229,252

225,231,247
1,2,4,8,11,12,26
93
91,93
122,137, 144,147,167
93,154
45
19,21,41,184,254
44-49, 53-59,62, 65,69-71
50,65,113,115
19,31,50,72
222
111,152
48, 71, 84
45-46,63,79-82,84,87,112-115,
154, 157
106,108,115
102-103,106, 127-129,141,147,
166
157,158,240
19,72
215,222,229,252
7-8,24,29,78,91,175,268
263
191-193
272
91,151-154,157-158
53-59,71
51-59, 134-135, 139, 140, 147,154,
161,215,254

296

species-environment
correlation
correlation, see also regression
coefficient of variation
coefficient of determination
R2 (see also coefficient of
determination)
multiple correlation coefficient
coefficient of community
regression coefficient
edge effects
| kriging
response curve (see also
unimodal response curve)
significance tests
test of significance, see
significance test
rank sum test
concordance measure
association measure, see
similarity
chi-square test
critical value
long-range spatial variation
(see also trend)
lag
landscape ecology
latent variable (see also
ordination axis)
latent structure
linear combination
linear model
linear predictor
logarithmic transformation
logit regression
log-linear regression
log-normal distribution
local trend surfaces
local minimum
maximum probability of
occurrence
maximum likelihood
reseating in DCA (see also
scaling of ordination axes)
scaling of ordination axes
matrix algebra
median
short-range variation
site
Braun-Blanquet method
Ward's method
multicriteria decision-making
multidimensional scaling
multiple logit regression
multiple regression (see also
regression, least-squares reg
ression and logit regression)



, .

, .

(.

)

, .


(. )

, .

, .

(.
)
, .

, .

51-53

30,93,153-155,158
231-232, 245
58, 140, 150
157,158
224
225
3,24
55,150-151
122,127,254
216,229
175,197
63,155,216

151, 184
18, 19,202
91, 136
228-229
17, 33,43, 58,65, 140,142,149,
202,206,268

168
21,140,145,150,185
103, 141,161,166,171
13-14,25, 35,44, 202
5,8,33,49,62,83,97,117,177,194
197
9,45-46,50,53,63,65,154
154, 155
94, 108-109,127,141,167
86,87,154
37
117, 144, 159
11,12,26,29-30,51-59,154
186-187, 204,205
32, 50, 62, 81, 95-97, 138,153,167

222,238, 239
198
198
29,32,42,43, 47-50, 52, 57, 59,
60-62,65,68,71, 81-82, 84-85,95,
109,111-117
91, 102,108,127, 132,142,146,
149,167,261,265
5,91-173,258
46,49,53,111,115
36

297

multiple least-squares
regression
mode, see optimum
response model
semivariogram model
dummy variable
alternating least-squares
first-order moment
second-order moments
moments, see statistical
moments
monitoring
mutticollinearity
loading
non-normal distribution
non-hierarchical clustering
non -linear model (see also
generalized linear model)
non-linear transformation
continuous variable, see also
quantitative variable
indirect gradient analysis (see
also ordination)
non-stationarity
nominal variable
nominal scale, see nominal
variable
standardized canonical
coefficient
standardization of variables
Hill's scaling
null hypothesis
abundance
composite clustering
generalized linear model
composite gradient
joint plot
inverse regression
total variance
total residual sum of squares
explanatory variable
single - linkage clustering
unimodal curve, see unimodal
response model (see also
Gaussian response curve)
uniformity, see stationarity
window
cluster optimality
optimality of a clustering
optimum of response curve

ordination diagram
ordination
axis, see ordination axis
residual deviance
residual standart deviation


, .

(.
-)

96,101,118,122,132,135,138,
152,156,257

194, 205
177,181-183, 187,200,262
31,82,84,104,177
34,37,41,124,130,141

(. )

(.
)







, .

14,30,34,39,45,51
30, 148, 216,223,239
33,40-42,48,52,68,73
37,83,119
12,37,42

R2 (.
)

(. )

41,217
80
80, 81,206
80
34,37

177,181-183
30, 37-40,66, 82, 86, 87,117-119

184
11,12,26,29-30,83
231
78,86
161,165
233, 234
51-59

91,175-183,196,198,204
34,36, 40, 67, 122-124
2,3,4,15,24
41,65,87,217
108
188,192,193,204
226,230-237, 239, 242, 243, 246
156
17-18,31,84,206,268

37
34
12, 272-274
22,26,29,155
259
79-81, 86
214,215,222,224,225,239
219,224-229,245,248
218,221,237-242
220
215,223-224,245
214,228
125,178
100,123,165
100-101,122-123,166

298

ordination axis
deviance, see residual
deviance
preference score
similarity ratio
absence of species (see also
presence-absence data)
fraction of variance accounted
for
estimation of parameters
error (see also residual)
parabola
slope parameter
parameters (see also
estimation of parameters)
square root transformation
response variable
transition model
transfer function
transition equations of CA
periodic variation
response surface
surface, see response surface
and trend surface
similarity
fitted value
field survey
polynomial regression
detrending-by-polynomials
complete-linkage clustering
semivariogram
polar ordination
ordinal variable
ordinal scale, see ordinal
variable
order of a polynomial
classification rule
assignment rule
rule of assignment
adjusted R2 (see also fraction
of variance accounted)
adjusted coefficients of
determination
least squares principle
decision making
causality
checking of data
training set
spatial (auto)covariance (see
also semivariogram)
spatial (auto) correlation
spatial interpolation
spatial independence
spatial structure
spatial scale
species space
standardization procedure
orthogonalization procedure
(see also detrending)
percentage similarity
straight line

 (.

)

(.

)
* .





-

, .

29,64,91

50,194
237
19-20,50,111,114,157,184
94,151-152,176,178-183
157

16,18-20
95,101,110,137-138,157,160,
169-171
100,170
159
34,36, 67, 73
224, 246
5,29-77,79,86,91,93,111,115,
119, 125,132, 135,140,145, 154155
30,31,33-43,46, 51-53,63,65,
216
82,109-110,130,183
15

3,4,7,10,15-16,242-244
229
259
31-32,44-47,65

157,159
157,158
157,158
174,198,258,262,265
1
262
30, 33

100,101, 122,139, 141,147, 159,

160-164,169
101, 122, 161,169

(.
)

4,15,24
110,186,194

127-130, 135,141-148,159,162164,167
124,129,159
62, 64, 86, 98-99, 102, 120-121,
133,174,193,195-196,200
64,78,174
263
17,18,20,27,224-225
109
189
37

299

direct gradient analysis (see

also regression and canonical
ordination)
pseudo-species
pseudo-cyclic variation
Poisson distribution
dissimilarity
Bernoulli distribution (see
also binominal distribution)
distribution, see frequency
distribution and also response
curve
frequency distribution
dispersion of species scores
dispersion of site scores
chi-square distance
prediction interval
regionalized variables
regression
least-squares regression
rare species
retrospective study
CA, see correnspondencc
analysis
sampling
smoothing
seasonal aspects
sigmoid curve (see also
monotonic response curve)
singular value
singular-value decomposition
singular vector
syntaxonomy
synecology
fusion of clusters
error part of response model
bias
down-weighting of rare
species
eigenvalue
eigenvector (see also singular
vector)
biplot
covariance biplot
table arrangement
community composition
composite sample
mean
expected abundance (see also
expected response)
average-linkage clustering
mean square
E, see expected response

CCA, .

,
. ,

,
(.
)
, .

(.
)



-



-

, .

(.
)
, .

(.
)




(.
)

F - (. F
-)

CCA, see canonical analysis

35,65
21,36

103,106-107, 154, 168,173

224-225
13,14,25,60
224,228-230
35-36,44,49, 53
15,253
127,134-135,141-142,146
167
36
34,36-37,41,53,117,149
36,149
231-232
199
34-37,39,41,67,68,136
95,118,137,155
202
202
174,273
42, 47-48, 50, 52, 65, 71, 84-85, 88,
106,111-115
31,93
242
240
20-21,49,61,103,130,151,184186,216,229
214, 223, 227, 234,246, 252
106,108,115,139
30

91
151,153
29,41,79,82
151
178, 181-183,185
100, 142,169
20, 130
189
156
156
53

300

standard error
standard deviation
standard deviation of species
turnover, see standard
deviation, response curve
standard deviation, response
curve (see also tolerance and
length of ordination axis)
statistical tests, see
significance test
statistical moments
statistical power analysis
stationarity
degrees of freedom
stratified random sampling
arrow in ordination diagram
succession
sum of squares
error sum of squares, see
residual sum of squares
residual sum of squares
regression sum of squares
spherical model
species-by-sites table (see also
table arrangement)
ANOVA table
theoretical variable
Wilcoxon test
Mann-Whitney test
community types
tolerance
scatter diagram
point kriging
exact interpolator
transformation, see data
transformation
data transformation
trend
detrending
regression equation (see also
response model)
cut-level, see pseudo-species
factor asnalysis (see also
principal component analysis)
stress function
response function
loss function
chord distance
centroid (see also weighted
average)
centring
centroid clastering
partial ordination
partial PCA
partial canonical ordination
partial Ftest (see also F test)

(.
)

"** (.
)

ANOVA - .

DC , .

F -
F -
Q-
R-
s.d, .
s.e., .
SVD, .
t , .

/ -
/ (. / -)
t , . t

43
17-18,21,27
17,179
17-18,27
175,224
152,178,181-183,185,205
42,78,84,115,203
1,2,8,9
1.2
12-15,22,24
33,40,44
231
144
104
104-108,114,139,169

35,53,219
35,53
131,152,159
131,159

23, 35, 40, 41-42, 47, 49, 52-55, 5758,60,70,202

40,216

301

frequency of occurrence
van der Maarel's scale
ratio scale
Braun-Blanquet scale
noise
Euclidean Distance
ecological amplitude (see also
tolerance)
ecology
community ecology
experiments
exponential curve
exponential model for
semivariogram
exponential transformation
horse-shoe effect (see also arch
effect)
arch effect
ANOVA see analysis of
variance
DCA, see detrended canonical
correspondence analysis
Ftest
F distribution
Q-mode algorithm
R-mode algorithm
s.d, sec standard deviation
s.e., see standard error
SVD, see singular value
decomposition
t of Gaussian curve, see
tolerance
ftest
t value (see also t test)
t of a parameter, see t value

XIV
XVII

1.
1.1.
1.2.
1.3.
1.4.
1.5.

2.
2.1.
2.2.
2.3.
2.3.1.
2.3.2.
2.3.3.
2.4.
2.4.1.
2.4.2.
2.4.3.
2.4.4.
2.5.
2.6.
2.6.1.
2.6.2.
2.6.3.
2.6.4.
2.7.
2.8.
2.9.
2.10.

-
: ,

10
10
11
12
12
14
15
16
16
16
18
20
22
23
23
24
24
25
25
26
27
27

3.
3.1.
3.1.1.

29
29
29
302

1
1
2
4
7
8
w

3.1.2.
3.1.3.
3.1.4.
3.2.
3.2.1.
3.2.2.
3.2.3.
3.3.
3.3.1.
3.3.2.
3.3.3.
3.4.

LSR -
:


- -
: -

-

:

- : -
-

30
31
33
33
33
37
40
43
43
44
48
49

78
78
79

4.2.1.
4.2.2.
4.2.3.
4.3.
4.4.
4.5.
4.6.
4.7.

5.
5.1.
5.1.1.
5.1.2.

3.5.
3.5.1.
3.5.2.
3.5.3.
3.5.4.
3.5.5.
3.6.
3.7.
3.8.
3.9.
3.10.
3.11.
4.
4.1.
4.2.

51
51
51
53
56
58
60
61
63
65
66
72

79
79
81
83
86
87
87
89
91
91
91
93

5.1.3.
5.2.

5.2.4.

(DCA)

5.2.5.
5.2.6.
5.2.7.

DCA

5.3.
5.3.1.

5.2.1.
5.2.2.
5.2.3.

5.3.2.
5.3.3.
5.3.4.

5.3.5.

5.3.6.
5.4.
5.5.
5.5.1.

R- Q-

5.5.2.

5.5.3.
5.5.4.
5.5.5.
5.5.6.

5.5.7.
5.6.

()

5.7.
5.8.

95
95
95
97
103

105
108
109

116
116
119
125
127
130
131
132

5.9.2.

()
()

5.9.3.
5.9.4.
5.9.5.
5.10.

(RDA)

5.11.

136
136
137
144
147
148
150
151
151
153
156
158
158
160
161
163
163
165
169

6.

( ..)

174

6.1.

174

6.1.1.

174

6.1.2.

175

5.9.
5.9.1.

304

6.2.
6.2.1.

6.2.2.
6.2.3.
6.2.4.
6.2.5.
6.3.
6.3.1.
6.3.2.
6.3.3.
6.4.

6.5.
6.6.
6.7.
6.7.1.

6.7.2.
6.8.

6.9.
6.10.

7.

7.1.
7.2.
7.2.1.
7.2.2.
7.2.3.

7.2.4.

Z X

7.2.5.

7.2.6.
7.2.7.

7.2.8.
7.3.
7.3.1.
7.3.2.

7.3.3.

7.3.4.

7.3.5.
7.3.6.
7.3.7.

7.3.8.
7.3.9.

7.310.
7.4.

305

175
175
176
180
184
186
191
191
193
193
197
198
199
199
201
202
203
204
207
213
213
215
215
215
216
216
217
218
219
222
222
222
224
224
226
227
228
230
234
235
235
237

1.
2.
3.

1.
2.
3.

1.
2.
3.
4.
5.
1.
2.
3.
4.
5.

,
, ,
-

, (Veluwe, the Netherlands)

ISBN 5 - 85941 - 124 - 3

. 25.03.99 70 100/16
19,25 ..
300
45
115598, , . , 12

237
237
239
242
245
245
246
248
252
252

252
256
257
258
258
258
259
263
263
266
266
268
268
269
272
275
291
293
302