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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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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

- ,
, 1. Prentice,
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
Science, Plant Ecology and Weed Science of the Wageningen Agricultural University .
Kilic. J.van de Peppel, M de Vries, J. van Keulen. M. Mijling
W.Laoch-Gieskes
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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
0
50
0

*
25
58
60

17
60
*

63

29

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

5.3
*
4.9
2.5
6.3

4.2
5.7
3.8
4.7

5.3
2.5
6.3

4.9
2.5

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

205

. : pref = (SPOS - SNEO) / (SPOS +


SNEO).
d. ( )
.
. ( MIN) (MID =
(MAX + MIN)/2).
/. , , ,
.
g. b-f .

. a-g .

6.5.

- 1 2 (), 3, 4
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
4

2
3
4
5
6
2
3
4
5
6

(+-)

2
2
0
2
0
6
5
6
4
6


: (+-)

2
3
4
5
6

0.33
0.40
0.00
0.50
0.00

0.80
0.33
0.06
0.60

0.17
0.75
0.40

0.00
0.50

0.20

.lb. , ,
:

1
2
3
4
5

<

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

2,3
(2,3),5
(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
E,F
A.D
A,F
,
C,D

D,G
^
A,(D,G)
(A,D,G),(E,F)
(A,D,E,F,G),B
(A,B,D,E^,G),C

6. Id. .
,
.
2

5
62
1 5

2
6
6
4
1

3
2
2
6
5

5
3
3
2
3

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5
3
2

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BEADGCHjmCGAFEB
:
1

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1
1
2

1
1
2

3
1
1
1

1
1
4
3
1

2
3
5

1
4
1

208

6.2.
.
. 2. :
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. ,
. :
1: 4 6, - 63

2
3
(4,6)
5

(4,6)

29
40
0
50

50
25')
58

17")
25

0"')

*) min (25, 60) = 25


")mm(17,25)=17
*")min(0,29) = 0
2: 5 2, 58
:

(2.5)
3
(4.6)

1(2,5)

40
0

29')
50")
0

17
*) min (29,50) = 29
') min (50,60) -50

209

3 : (2,5) 3, - 50
:
1

(2,3,5)

(2,3,5)

29

(4,6)

0)

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

..
. .
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

.. ,
.

1
2
3
4
5

2.5
2.5
2.5
4.2
4.7

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

1,5
(1,5X3
4,6
(1,3,5),2
(1,2,3,5),(4,6)

4 6 - .
2 3 A, F G, .
1 5 - ..

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
6
4
5
1

1
4
3
3
4
5
0

-1
-0.33
-0.20
0.33
0
0
1

0
2
2
8
7
10
1

1
4
3
1
1
0
0

-1
-0.33
-0.20
0.78
0.75
1
1

0
1
1
6
7
10
1

1
5
4
3
1
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-0.67
-0.75
0.33
0.75
1
1

la. : 1,3 6 - , 2,4 5 -


.
lb-1. 6.8.
Id 1-6 : -1.73; 0.79; -0.86; 1;
0.13 0.66, .
le. MID - (-1.73 + 1)/2 - -0.36.
If. 1 3 , - .
2-2 . 6.8.
2d 1-6 : -1.73; 5.84; 0.34; 5.75; 1.03 8.81
.
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
.
. MID - (-3.17 + 7.91)/2 = 2.37.
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
20


, , (11 12) / 20 = 6.6 ( 'TWINSPAN'
, ).
(\ :


: 1+2
3+4+5

0
6.6
5.4
12

1
4.4
3.6
8

11
9
20

1+2+3
8.8
7.2
16

4
2.2
1.8
4

11
9
20

( ):
- = 4.4 ( - ef = 19.36 (TWINSPAN), - = 2.2 ( - ef = 4.84 (FLEXCLUS,
).
/ = 19.36 (1 / 6.6 + 1 / 4.4 + 1 / 5.4 + 1 / 3.6) = 16.30 (TWINSPAN)
*> = 4.84 (1 / 8.8 + 1 / 2.2 + 1 / 7.2 + 1 /1.8) = 6.11 (FLEXCLUS, ).
,
. , . . (
) 2 : v. - ,
( < 0.005 TWINSPAN 0.01 < < 0.025
FLEXCLUS ). -
, .
6.6.

,
1; , TWINSPAN
. ().
1
. 18 ,
.
. 7

10

20* 9

19

17

8*

16*

12* 13* 5*

14* 15

2.8

2..8

3.3

3.5

3.5

3.5

3.7

3.7

4.0

4.2

4.2

4.3

4.3

5.7

5.8

6.0

6.3

9.3

12

1.5

1.5

7.5

7.5

10.

10

12.

12.

14

15

16

17

18

19

10 12 10.5 12.5, . \ (
, ) : 105. = 8 (8 + 11 + 1) -105 =
55. (55),
. ,
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1.

212

7.
P. A. Burrogh

7.1.
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7.1. -. ,
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( . choros - ) - , ,
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213


Z = bo + bj+ s

7.1

0- , ;
bj - /, :,
(f bj = 0)\
- , ,
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Beckett, 1973) , -,
.

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214

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7.2.
7.2.1.

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( Z),
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jf, Z X , 7.2.
Z ,
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:
- Z ;
- Z X ;
- X .
Z - , X (
), . ,

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

215

. 7.2. . : Z. :
2 X. : X.
.

Z=*b0+ b]X + e,

7.2

. bo bj
( 3.2.2). t bj (. 3.2.2) Z X
, -
. () bt.
7.2.3.
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. , , ,
.
Z, ,
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.
7.2.4. ZomX
Z X
(. 7.3).

216

J - - -

. 7 3 . .

, , :
Z = bo +biX +b3X2 +

7.3

,
.
(. 3.2.1).
.
.
, . 3.2.3 ,
.
7.2.5.
Xi2 Z :
Z = S r + S i p {2

+ ,

7.4

.
:

0 + bjXj +

(1 );

bo + bjXi + + ]2 + biXiX2 + 22

(2 ).

7.4*. -
. 7.4 (+1)(+2)/2 .
bt -
(. 3.5.2). (Ripley, 1981) , -

. 7.4 1, 2 3 .

217

. 7.4. 1, 2 3- ,
.

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7.2.6.
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218

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7.2.7.

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219

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220

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120 .
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. 7.7. 1, 2 3- 4- , 150
.

221

7.7 1, 2, 3 4- ,
.
7.2.8.

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222

7.9. : - (); ,
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223

Z :
Z(x) = () + '() + ",

7.5

() - , ,
' ()- , ;
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*.
7.3.2.
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**.
Z(xi)t..,Z(xn) , Z (xi+h),...,Z(xn+h) */,...,*,
, .
,
h i xt + h.
, " ",
, .
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.
():

(..).
Z X. 2 "series " (,
). ,
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, . . .
224

7.6

E|Z(*)] = |i,

- Z ,
5 .
2 = I n Uxi)/n,

7.7

- .
. :
var Z () = E[Z(JC) - \i ]2

7.8

:
( 0 ) = Ii-i n ,) - z ]2/

7.9

:
(xi,x2) [Z (*;) - ji][Z (*) - ]

7.10

, xt = ,+, h - (),

C(fc) = ^i n - h [Z(x;+/,)- F][Z(x,)- z]/(n-h)

7.11

, 0, C(h)=C(0), .. .
,
, , , .
:
r(h)= (h)/C (0)

7.12

-1 +1
* Z(JC/) - Z(xj)
Y (h) = 1/2 var [Z (*i) -Z(xj)]

7.13

, ,
y(h) = E[Z ( + h) - Z (x)] 2

7.14

(. .).

225

. 7.10. /(h) C(h) ,


.

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

7.15

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

7.16

,
.
. ,
, (. 7.10).
7.3
,
().
, 7.11.
1. 24:8 = 3.
2.
(0)=[(1-3) 2 +(3-3) 2 +(4-3) 2 +(6-3) 2 +(4-3) 2 +(3-3) 2 +(3-1) 2 +(2-3) 2 ]/8 = : 20/8
= 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. ,
, 1.5 1.5 . 7.12
, 50 ,
200 , 4 . 7.12 . ,
, , ,
7.12.
.
7.12 ,
, . .
(. 7.12d) , ,
, .
, , 1 5 x 1 5 , ..
, 1/100 , -

227

(X 10 -)
30

29

I 28

| 27

26

-1

10

15

20

25
.

10

20

30

40

50

-1

10

15

20

25

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.
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.
7.3. .
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,
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, ,

* .
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228

. 7.13. .
C-D , -, , .

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Zfod-Zfa+h), h - . ,

<^. "" (..
).
,
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C(h)/C(0), .
,
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229

,
(
) *.
, Z(x+h) - Z(x)
, , ..
var [Z(x + h) - ZOO] = E + h) - Z(x)]2 = 2 ()

7.17

,
, .
Z(x). ,
7.10.
7.3.7. ,

( ),
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9
<rintrinsic hypothesis ( ),
, .

. - . (..)

230

), 7.14,
, Z(x)t
. , , s"
7.5.

.
. ,
(. 7.4),
.
, . (Webster &
Burgess, 1984) .

(Journel & Huijbregts, 1978) :
( )
- (. 7.14)
{) = + Ci[3/f / 2) - 0.5(/f / af]

0<h<a

= + ci

h> ,

7.18

- ,
-,
- ,
+ ci .
- ( , . 7.14)
y(h) = co+bh
= co+Ci

</
>

7.19

- (. 7.14)
Y () = + ci [1 - (-/? / )]

7.20

-
() = + ci [1- (-2/ 2)]

7.21

-
r(h)=co+ bh

7.22

-
(/*) = + ci ioge/i

7.23
231

100%
(. 7.14f) ,
.
Y(A) = co

7.24


(Webster & Burgess,
1984). ,
( ). , ,
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.

*,

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w

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> '

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6
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232

233

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234

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600 , 350 . ,
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235

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to
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30
10

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250

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500

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750

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1000 1250 1500 1750 m

()
. 7.16.
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236

, 100%
(. 7.16). ,

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7.4.2.
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2 (*) = 11 Z(xd I Wi,

7.25

<p(d(x,xi)) xt.
0 (< (d)->0, d -> <>).
. ,
<p(d) :
Z (Xj) = E H n [Z(*i) / d/] I (1 /

rf

7.26

xj - , (
, dtJ - xt xj).

237


(. 7.17) , .. ,
(Ripley, 1981). ,
, , .
,
.
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,
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, .
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- .
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.
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238

.
.
7.4.3. ,
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(1983) (1983).
(1985) (1985).

(Bullock & Stallybrass, 1977).
, , , w,
, ..
.
, ,
. , - ,
Z , . :
2 () = r.in >nZ(;ti)

7.27

%= 1.
, Z(XQ) * ,
.
Z(XQ)-Z(XQ) , wt ( i) :
Ij-in yvjf(xu xj) + v|/ = (*/,*),

7.28

( - Z { / (xit XQ} -
xt \

239

. 7.18. ,
: ; : .
e n d , . ,
.

\ - ( ,
). 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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, (Burrough,
1986). 7.18b-7.19b
. ,
, , ,
(. 7.18; 7.18d).
7.5

, Z
. ,
.
= faS21 (g-) 2

7.30

,
-g +g (/ - ).
, ,
,
, ,
, (19846) ,
, .
,
, ,
. .
, 1 ,
Z.
( ) 141 , , ,

242

.
. :
- , (= 1; 4; 9; 16; 25);
- .
= 9 1
(. 7.20).
,
. 7.21 ,
;
( )
(. , Webster & Burgess, 1984b).
, .

|
I

I
I

. 7.20.4 .

. 7.21.
.

243

s.e.

7.22. . :
. : .

,
, (. 7.20).
,
, .

,
. 7.22,
,
,
. (Webster & Nortkliff, 1984) ,
2-3 . (1984b) ,
,
,
.

244

7.6.
. -,

. -, ,
,
,
.

, ,
, .
, , ,
, . -,
, 50-100
(. , 1986).
. , ,
, .
7.7.
( )
, .

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

. ,
.

. , 1930-1940- , ,
,
. 1960- .
( , Shepard ,1968),
,
( ) (Davis, 1986).
,
, , (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.

.
,
.
7.2.
7.2 ,
( 3 ), ,

. 25 , 25 .
:
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 .
, ,
() 7 (700 ).
, 4
(400 ) , .
7.5 .

(. 7.6). 7.7,
, 1,
2 . ,
, 1 ,
.
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

-0.120

14

6.600

5.159

1.441

14.990

13.949

0.951

15

5.500

5.144

0.356

13.800

14.279

-0.479

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. ,
.

- ,
, 1-
( 5.2) .
( 45.7%
) , ,
: ,
, ,
,
.
, -
.
:
( 55 ),
, ,
, .
.
,
,
. 8.3.

254

e
i

lilil^
tx

. 8 3 . : :
( 55 ); b - ( 55 ); (18 ); d - (8 ).
; ,
, ,
55 .

255

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

. 8.4.
. 1: , 1
( , ). 2: 1 .
3: 1 . 4: , 1
( , ). ,
, ,
(. 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.), .
,
.
, , ,
,
,
.
. ,
.
TWINSPAN (Hill, 1976b), DECORANA (Hill, 1979a), FLEXCLUS
(van Tongeren, 1986). -
3000 1700 .
8.2.3.
(. . 8.5):
- ,
;
- ;
- .

259

150

,
*

!
|

1i

'

'

1

1

. 8.5. -
.

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. - ,
, ,
,
. ,
, ,
.
,
.
.
. .
, , . ,
,
. , ,
. ,
.
.
, ,
211 .
:
-
, . ,
. ,
, ;
-
, , ,
;
- , :
,
. 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 ,
- .
, . ,
.
.
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 .
.
.
. -
. ,
,
, , 7 .
, , 2
1, (. 8.8). ,
, , ,
, .
,
(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.


,
. , .
,
.
.
(/), ( , ).
() : , ,
. : , , ,
, , , .
.
,
. , ,

.
,
. 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

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

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

: *<


fiSSa"
E5SS
EZ23

CZ3

CZZ)

. 8.9. Gammarus :
. ( ).

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), ,
: , (
), ,
.
:
.
,
, .
,
,
. G1 G5 272

. , G3 Gammarus
. , Chironomus .
,

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

!
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
0.0449
0.0436
0.0420
0.0405
0.0395
0.0393
0.0383
0.0375

0.9262
0.7046
0.3566
0.5347
0.9180
0.4104
0.2539
0.1644
0.1488
0.1380
0.0329

17 15
18 27
19 23
20 22

Bcek Ordcrvccn (N)


LrSocrensc beck (N)
Loenen (Zilvcn)
Loencn (Stroobrock)

0.0335
0.0289
0.0281
0.0275

0.2925
0.5432
0.4430
0.9510

21
22
23
24
25
26
27

Meibcek (N)
Vlasbcck
Winkcwijert
M-Hccrdcr beck (W)
Loencn (Stecnput)
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274

Agterberg, F.D., 1982. Recent developments in Geomathematics. Geo-Processing


2: 1-32.
Aitchison. J. & J.A.C Brown, 1969. The lognormal distribution. Cambridge
University Press, Cambridge, 176 pp.
Alderdice, D.F., 1972. Factor combinations: responses of marine poikilotherms
to environmental factors acting in concert. In: O. Kinne (Editor): Marine
ecology, vol 1, part 3. Wiley, New York, p. 1659-1722.
Alvey, N.G. et al., 1977. GENSTAT: a general statistical program. Rothamsted
Experimental Station, Harpenden.
Ancot, J.P. & T. J. van de Nes, 1981. Integral evaluations of alternative water
management scenarios in East Gelderland. In: Water resources management
on a regional scale. Committee for hydrological research TNO, Proceedings
and Information no 27. The Hague, p. 129-164.
Anderberg, M.R., 1973. Cluster analysis for applications. Academic Press, London,
373 pp.
Anderson, A.J.B., 1971. Ordination methods in ecology. Journal of Ecology 59:
713-726.
Anderson, S., A. Auquier, W.W. Hunck, D. Oakes, W. Vandaele & H.I. Weisberg,
1980. Statistical methods for comparative studies. Wiley, New York, 302 pp.
Austin, M.P., 1971. Role of regression analysis in plant ecology. Proceedings of
the Ecological Society of Australia 6: 63-75.
Austin, M.P., 1976. Performance of four ordination techniques assuming three
different non-linear species response models. Vegetatio 33: 43-49.
Austin, M.P., 1980. Searching for a model for use in vegetation analysis. Vegetatio
42: 11-21.
Austin, M.P., R.B. Cunningham & P.M. Fleming, 1984. New approaches to direct
gradient analysis using environmental scalars and statistical curve-fitting
procedures. Vegetatio 55: 11-27.
Baker, R.J. & J.A. Nelder, 1978. The GLIM system, Release 3. Numerical
Algorithms Group, Oxford.
Ball, G.H., 1966. A comparison of some cluster seeking techniques. Stanford
Research Institute, California.
Barkman, J.J., H. Doing & S. Segal, 1964. Kritische Bemerkungen und Vorschlage
zur quantitativen Vegetationsanalyse. Acta Botanica Neerlandica 13: 394-419.
Barlow, R.E., D.J. Bartholomew, J.M. Bremner 8c H.D. Brunk, 1972. Statistical
inference under order restrictions. Wiley, New York, 388 pp.
Bartlein, P.J. & T. Ill Webb, 1985. Mean July temperature at 6000 yr B.T. in

275

Eastern North America: regression equations from fossil-pollen data. Syllogeus


55: 301-342.
Bartlein, P.J., I.C. Prentice & T. Ill Webb, 1986. Climatic response surfaces from
pollen data for some eastern North American taxa. Journal of Biogeography
13:35-57.
Battarbee, R.W., 1984. Diatom analysis and the acidification of lakes. Philosophical
Transactions of the Royal Society of London 305: 451-477.
Batterink M. & G. Wijffels, 1983. Een vergelijkend vegetatiekundig onderzoek
naar de typoiogie en invloeden van het beheer.van 1973 tot 1982 in de
Duinweiianden op Terschelling. Report Agricultural University, Department
of Vegetation Science, Plant Ecology and Weed Science, Wageningen. pp 101.
Beadle, N.C.W., 1954. Soil phosphate and the delimitation of plant communities
in eastern Australia. Ecology 35: 370-375.
Beals, E.W., 1985. Bray-Curtis ordination: an effective strategy for analysis of
multivariate ecological data. Advances in Ecological Research 14: 1-55.
Beckett, P.H.T. & P.A. Burrough, 1971. The relation between cost and utility
in soil survey. IV. Comparison of the utilities of soil maps produced by different
survey procedures and to different scales. Journal of Soil Science 22: 466480.
Becking, R.W. 1957. The Ziirich-Montpellier school of Phytosociology. Botanical
Review 23: 411-488.
Beisley, D.A., E. Kuh & R.E. Welsch, 1980. Regression diagnostics. Wiley, New
York, 292 pp.
Benzecri, J-P. et al., 1973. L'anaiyse des donnees. II: L'analyse des correspondances.
Dunod, Paris, 619 pp.
Bie, S.W. & P.H.T. Beckett, 1973. Comparison of four independent soil surveys
by air photo interpretation, Paphos Area (Cyprus). Photogrammetrica 29 (6):
189-202.
, R., I. Kowarik & R. Bornkamm, 1983. Untersuchungen zur Anwendung
der Zeigerwerte nach Ellenberg. Verhandlungen der Gesellschaft fur Okologie
11:35-56.
Box, G.E.P., 1976. Science and statistics. Journal of the American Statistical
Association 71: 791-799.
Box, G.E.P., W.G. Hunter 8c J.S. Hunter, 1978. Statistics for experimenters. Wiley,
New York, 671 pp.
Braun-Blanquet, J.J., 1964. Pflanzensoziologie, Grundziige der Vegetationskunde.
3rd Edition. Springer, Vienna, New York, 865 pp.
Bray, J.R. & J.T. Curtis, 1957. An ordination of the upland forest communities
of southern Wisconsin. Ecological Monographs 27: 325-349.
Brown, G.H., 1979. An optimization criterion for linear inverse estimation.
Technometrics 21: 575-579.
Brown, P.J., 1982. Multivariate calibration. Journal of the Royal Statistical Society,
Series 44: 287-321.
Buckland, ST., 1982. Statistics in ornithology. Ibis 124: 61-66.
Bullock, A. & O. Stallybrass, 1977. Fontana dictionary of modern thought. Fontana
Books, London, 694 pp.

276

Bunce, R.G.H., CJ. Barr & H. Whittaker, 1983. A stratification system for
ecological sampling. In: R.M. Fuller (Editor): Ecological mapping from ground,
air and space, I... symposium 10: 39-46.
Burrough, P.A., 1983a. Multiscale sources of spatial variation in soil I. The
application of fractal concepts to nested levels of soil variation. Journal of
Soil Science, 34: 577-597.
Burrough, P.A., 1983b. Multiscale sources of spatial variation in soil II. A nonBrownian fractal model and its application in soil survey. Journal of Soil
Science 34: 599-620.
Burrough, P. A., 1986. Principles of geographical information systems for natural
resource analysis. Oxford University Press, Oxford.
Burrough, P.A., L. Brown & E.C. Morris, 1977. Variations in vegetation and
soil pattern across the Hawkesbury Sandstone Plateau from Barren Grounds
to Fitzroy Falls, New South Wales. Australian Journal of Ecology 2: 137159.
Campbell, N.A. & W.R. Atchley, 1981. The geometry of canonical variate analysis.
Systematic Zoology 30: 268-280.
Carroll, J.D., 1972. Individual differences and multidimensional scaling. In: R.N.
Shepard, A.K. Romney & S.B. Nerlove (Editors): Multidimensional scaling.
Theory and application in the behavioral sciences. Vol. 1, Seminar Press, New
York, p. 105-155.
Chatfield, C, 1981. The analysis of time series. Chapman and Hall, London.
268 pp.
Christiansen, F.B. & T.M. Fenchel, 1977. Theories of populations in biological
communities. Springer Verlag, Berlin, 144 pp.
Chutter, F.M., 1972. An empirical biotic index of the quality of water in South
African streams and rivers. Water Research 6: 19-30.
Clarke, G.M., 1980. Statistics and experimental design. 2nd Edition. Edward
Arnold, London, 196 pp.
Clements, F.E., 1928. Plant succession and indicators. Wilson, New York, 453 pp.
Clymo, R.S., 1980. Preliminary survey of the peat-bog Hummell Knowe Moss
using various numerical methods. Vegetatio 42: 129-148.
Cochran, W.G., 1963. Sampling techniques. 2nd Edition. Wiley, New York, 430 pp.
Cochran, W.G., 1977. Sampling techniques. 3rd Edition. Wiley, New York, 444
PP.
Cochran, W.G., 1983. Planning and analysis of observational studies. Wiley, New
York, 158 pp.
Cochran, W.G. & G.M. Cox, 1957. Experimental Designs. New York, Wiley,
630 pp.
Coetzee, B.J. & M.J.A. Werger, 1975. On associatian analysis and the classification
of plant communities. Vegetatio 30: 201-206
Cohen, J., 1973. Statistical power and research results. American Educational
Research Journal 10: 225-229.
Cohen, J., 1977. Statistical power analysis for the behavioural sciences. Revised
edition. Academic Press, London, 489 pp.
Cook, R.D. & S. Weisberg, 1982. Residuals and influence in regression. Chapman

277

and Hall, London, 230 pp.


Corley, M.F.V., AC. Crundwell, R. Dull, M.O. Hill & A.J.E. Smith, 1981. Mosses
of Europe and the Azores; an annotated list of species, with synonyms from
the recent literature. Journal of Bryology 11: 609-689.
Corsten, L.C.A., 1985. Current statistical issues in agricultural research. Statistica
Neerlandica 39(2): 159-168.
Corsten, L.C.A. & K.R. Gabriel, 1976. Graphical exploration in comparing variance
matrices. Biometrics 32: 851-863.
Cox, D.R., 1958. Planning of experiments. Wiley, New York, 315 pp.
Cox, D.R. & E.J. Sneil, 1981. Applied statistics. Chapman and Hall, London,
197 pp.
Cramer, W, 1986. Vegetation dynamics on rising sea shores in eastern central
Sweden. Acta Universitatis Uppsaliensis. Comprehensive summaries of Uppsala
dissertations from the Faculty of Science 25. Almqvist & Wiksell International,
Stockholm, 34 pp.
Cuppen, H.P.J.J. & W. Oosterloo, 1980. Een orienterend onderzoek naar de
macrofauna van de Nijmolense beek (Gemeente Epe). Report Regionale
Milieuraad Oost Veluwe en Zuiveringsschap Veluwe, Apeldoorn, 50 pp.
Curtis, J.T. & R4P. Mcintosh, 1951. An upland forest continuum in the prairieforest border region of Wisconsin. Ecology 32: 476-496.
Dale, M.B., 1975. On objective methods of ordination. Vegetatio 30: 15-32.
Dargie, T.C.D., 1984. On the integrated interpretation of indirect site ordinations:
a case study using semi-arid vegetation in south-eastern Spain. Vegetatio 55:
37-55.
David, M., 1977. Geostatistical ore reserve estimation. Developments in Geomathematics 2. Elsevier, Amsterdam, 364 pp.
Davies, P.T. & M.K-S. Tso, 1982. Procedures for reduced-rank regression. Applied
Statistics 31:244-255.
Davis, J., 1986. Statistics and Data Analysis in Geology. 2nd Edition. Wiley,
New York, 550 pp.
Davis, R.B. & D.S. Anderson, 1985. Methods of pH calibration of sedimentary
diatom remains for reconstructing history of pH in lakes. Hydrobiologia 120:
69-87.
Davison, M.L., 1983. Multidimensional scaling. Wiley, New York, 242 pp.
Dawkins, H.C., 1983. Multiple comparisons misused: why so frequently in responsecurve studies? Biometrics 39: 789-790.
Dent, B.D., 1985. Principles of thematic map design. Addison-Wesley, Reading,
Massachusetts.
De Sarbo, W.S. & V.R. Rao, 1984. GENFOLD2: a set of models and algorithms
for the general unfolding analysis of preference/dominance data. Journal of
Classification 1: 147-186.
de Wit, A.E., J. Spuij & P.J.C. van Boheemen, 1984. Slootplanten als indicatoren:
een onderzoek naar indicatoren en hun bruikbaarheid in een biologisch meetnet.
Landschap 1:219-227. .
Dixon, W.J., 1981. BMDP statistical software. University of California Press,
Berkeley, 726 pp.

278

Dobson, A.J., 1983. introduction to statistical modelling. Chapman and Hall,


London, 125 pp.
Draper, N. & H. Smith, 1981. Applied regression analysis. 2nd Edition. Wiley,
New York, 709 pp.
Dunn, G. & B.S. Everitt, 1982. An introduction to mathemathical taxonomy.
Cambridge University Press, Cambridge.
Dupont, P., 1975. Les limites altitudinales des landes atlantiques dans les montagnes
cantabriques (Nord de FEspagne). In: J.M. Genu (Editor): La vegetation des
landes d'Europe occidentale Lille 1975, Colloques phytosociologiques 2: 4758. J. Cramer Vaduz.
Efron, ., 1982. The jackknife, the bootstrap and other resampling plans. SIAM,
Philadelphia, 92 pp.
Egerton, F.N., 1968. Leeuwenhoek as a founder of animal demography. Journal
of Historical Biology 1: 1-22.
Ellenberg, H., 1948. Unkrautgesellschaften als Mass fur den Sauregrad, die
Verdichtung und andere Eigenschaften des Ackerbodens. Berichte uber Landtechnik, Kuratorium fur Technik und Bauwesen in der Landwirtschaft 4: 130146.
Ellenberg, H., 1979. Zeigerwerte der Gef&sspflanzen Mitteleuropas. Scripta Geobotanica9, Gottingen, 121 pp.
Ellenberg, H., 1982. Vegetation Mitteleuropas mit den Alpen in Okologischer
Sicht. 3rd Edition. Ulmer Verlag, Stuttgart, 989 pp.
Everitt, B.S., 1980. Cluster analysis. 2nd Edition. Heinemann, London, 122 pp.
Everitt, B.S., 1984. An introduction to latent variable models. Chapman and Hall,
London, 107 pp.
Faith, D.P., 1983. Asymmetric binary similarity measures. Oecologia 57: 287-290.
Fasham, M.J.R., 1977. A comparison of nonmetric multidimensional scaling,
principal components and reciprocal averaging for the ordination of simulated
coenoclines and coenoplanes. Ecology 58: 551-561.
Feoli, E. & L. Orloci, 1975. Analysis of concentration and detection of underlying
factors in structured tables. Vegetatio 40: 49-54.
Finney, D.J., 1964. Statistical methods in biological assay. Griffin, London, 668
pp.
Fisher, R.A., 1954. Statistical Methods for Research Workers. 12th Edition/Oliver
& Boyd, London, 356 pp.
Fresco, L.F.M., 1982. An analysis of species response curves and of competition
from field data sets: some results from heath vegetation. Vegetatio 48: 175185.
Gabriel, K.R., 1971. The biplot graphic display of matrices with application to
principal component analysis. Biometrika 58: 453-467.
Gabriel, K.R., 1978. Least squares approximation of matrices by additive and
multiplicative models. Journal of the Royal Statistical Society, Series 40:
186-196.
Gauch, H.G., 1979. COMPCLUS - A FORTRAN program for rapid initial
clustering of large data sets. Cornell University, Ithaca, New York.
Gauch, H.G., 1982. Multivariate analysis in community ecology. Cambridge

279

University Press, Cambridge, 298 pp.


Gauch, H.G. & G.B. Chase, 1974. Fitting the Gaussian curve to ecological data.
Ecology 55: 1377-138!.
Gauch, H.G., G.B. Chase 8c R.H. Whittaker, 1974. Ordinations of vegetation
samples by Gaussian species distributions. Ecology 55: 1382-1390.
Gauch, H.G., R.H. Whittaker & S.B. Singer, 1981. A comparative study of
nonmetric ordinations. Journal of Ecology 69: 135-152.
Gause, G.F., 1930. Studies on the ecology of the Orthoptera. Ecology II: 307325.
i
Gift, A., 1981. Nonliriear multivariate analysis. DSWO-press, Leiden, 451 pp.
Giloi, W.K., 1978. Interactive Computer Graphics - Data Structures, algorithms,
languages. Prentice Hall, Englewood Cliffs, N.Y., 354 pp.
Giltrap, D., 1983. Computer Production of soil maps I. Production of grid maps
by interpolation. Geoderma 29: 295-311.
Gittins, R, 1985. Canonical analysis. A review with applications in ecology.
Springer-Verlag, Berlin, 351 pp.
Glacken, C.J., 1967. Traces on the Rhodian shore. Nature and culture in western
thought from ancient times to the end of the eighteenth century. University
of California Press, Berkely, 762 pp.
Gleason, H.A., 1926. The individualistic concept of the plant association. Bulletin
of the Torrey Botanical Club 53: 7-26.
Goodall, D.W., 1954. Objective methods for the classification of vegetation. III.
An essay in the use of factor analysis. Australian Journal of Botany 1: 3963.
Goodall, D.W., 1970. Statistical Plant ecology. Annual review of Ecology and
Systematics I: 99-124.
Goodall, D.W., 1973. Numerical methods of classification. In: R.H. Whittaker
(Editor): Handbook of vegetation science, Volume 5: Ordination and Classification of plant communities. Junk, the Hague: 575-615.
Goodall, D. W. & R. W. Johnson, 1982. Non-linear ordination in several dimensions.
A maximum likelihood approach. Vegetatio 48: 197-208.
Gordon, A.D., 1981. Classification. Methods for the exploratory analysis of
multivariate data. Chapman and Hall, London, 193 pp.
Gounot, M., 1969. M6thodes d'etude quantitative de la vegetation. Masson, Paris,
314 pp.
Gourlay, A.R. & G.A. Watson, 1973. Computational methods for matrix eigen
problems. Wiley, New York, 130 pp.
Gower, J.C., 1966. Some distance properties of latent root and vector methods
used in multivariate analysis. Biometrika 53: 325-338.
Gower, J.C. 1974. Maximal predictive classification. Biometrics 30: 643-654.
Green, R.H., 1979. Sampling design and statistical methods for environmental
biologists. Wiley, New York, 257 pp.
Greenacre, M.J., 1984. Theory and applications of correspondence analysis.
Academic Press, London, 364 pp.
Greig-Smith, P., 1971. Application of numerical methods to tropical forests. In:
G.P. Patil, E.C. Pielou & W.E. Waters (Editors): Statistical ecology. Pennsyl-

280

vania State University. 3: 195-204.


Grolle, R., 1976. Verzeichnis der Lebermoose Europas und benachbarter Gebiete.
Feddes Repertorium 87: 171-279.
Guillerm, J.L., 1971. Calcul de Information fournie par un profil ecologique
et valeur indicatrice des especes. Oecologia Plantarum 6: 209-225.
Hajdu, L.J., 1982. Graphical comparison of resemblance measures in
phytosociology. Vegetatio 48: 47-59.
Hardonk, R., 1968. Koornmullenaers, Pampiermaeckers en Coperslaghers. Korte
historie van de watermolens van Apeldoorn, Beekbergen en Loenen. Historisch
Museum Moerman, Apeldoorn, 256 pp.
Harrell, F, 1980. The LOGIST procedure. In: SAS supplemental library user's
guide, 1980 Edition. SAS Institute Inc., Cary, N.C., p. 83-89.
Hartigan, J.A., 1975. Clustering algorithms. Wiley, New York, 364 pp.
Harvey, ., 1981. Environmental intervention: The monitoring paradigm. I. The
monitoring concept and practice of descriptive monitoring. The environmen
talist I: 283-291.
Hawksworth, D.L., P.W. James & B.J. Coppins, 1980. Check-list of Britisch lichenforming, lichenocolous and allied fungi. Lichenologist 12: 1-115.
Heiser, W.J., 1981. Unfolding analysis of proximity data. Thesis. University of
Leiden, Leiden, 273 pp.
Heiser, W.J., 1986. Undesired nonlinearities in nonlinear multivariate analysis.
In: E. Diday et al. (Editors): Data analysis and Informatics 4. North Holland,
Amsterdam, p. 455-469.
Heiser, W.J., 1987. Joint ordination of species and sites: the unfolding technique.
In: P. Legendre & L. Legendre (Editors): Developments in numerical ecology.
Springer Verlag, Berlin, p. 189-221.
Hill, M.O., 1973. Reciprocal averaging: an eigenvector method of ordination.
Journal of Ecology 61: 237-249.
Hill, M.O., 1974. Correspondence analysis: a neglected multivariate method.
Applied Statistics 23: 340-354.
Hill, M.O., 1977. Use of simple discriminant functions to classify quantitative
phytosociological data. In: E. Diday, L. Lebart, J.P. Pages & R. Tomassone
(Editors): First International Symposium on Data Analysis and Informatics.
Vol 1: 181-199. Institut de Recherche d'Informatique et d'Automatique, Le
Chesnay, 181-199.
Hill, M.O., 1979a. DECORANA - A FORTRAN program for detrended cor
respondence analysis and reciprocal averaging. Cornell University Ithaca, N.Y.,
52 pp.
Hill, M.O., 1979b. TWINSPAN - A FORTRAN program for arranging mul
tivariate data in an ordered two-way table by classification of individuals and
attributes. Cornell University Ithaca, N.Y., 90 pp.
Hill, M.O., R.G.H. Bunce& M.W. Shaw, 1975. Indicator species analysis, a divisive
polythetic method of classification and its application to a survey of native
pinewoods in Scotland. Journal of Ecology 63: 597-613.
Hill, M.O. & H.G. Gauch, 1980. Detrended correspondence analysis, an improved
ordination technique. Vegetatio 42: 47-58:

281

Hoeking, R.R. & O.J. Pendleton, 1983. The regression dilemma. Communications
in Statistics - Theory and Methods 12: 497-527.
Hogeweg, P., 1976. Iterative character weighing in numerical taxonomy. Comput.
Biol. Med. 6: 199-211.
Howe, S.E. & T.HI Webb, 1983. Calibrating pollen data in climatic terms: improving
the methods. Quaternary Science Reviews 2: 17-51.
Hurlbert, S.H., 1984. Pseudoreplication and the design of ecological field experiments. Ecological Monographs 54: 187-211.
Ihm, P. & H. van Gro^enewoud, 1984. Correspondence analysis and Gaussian
ordination. COMPSTAT lectures 3: 5-60.
Uzerman, A.J., 1979. Sprengen en sprengenbeken op de Veluwe. Een onderzoek
naar beheer, onderhoud en watervoorziening in historisch perspectief. Rapport
ten behoeve van de commissie bestudering waterhuishouding Gelderland,
Arnhem, .110 pp.
Innis, G.S., 1979. Letters to the editor. Bulletin of the Ecological Society of America
60: 142.
Israels, A.Z., 1984. Redundancy analysis for qualitative variables. Psychometrika
49:331-346.
Jaccard, P., 1912. The distribution of the flora of the alpine zone. New Phytologist
11:37-50.
Janssen, J.G.M., 1975. A simple clustering procedure for preliminary classification
of very large datasets of phytosociological relev6es. Vegetatio 30: 67-71.
Johnson, S.B. & R.D. Berger, 1982. On the status of statistics in phytopathology.
Phytopathology 72: 1014-1015.
Jongman, R.H.G. & Th.J. van de Nes (Editors), 1982. Beken op de Veluwe. Een
onderzoek naar de mogelijkheden voor herstel en behoud. Begeleidingscommissie Proefgebied Nationaal Landschap Veluwe, Arnhem, 112 pp.
Journel, A.J. & C.J. Huijbregts, 1978. Mining gcostatistics. Academic Press, New
York, 600 pp.
Jowett, G.H. 8c G. Scurfield, 1949. A statistical investigation into the distribution
of Holcus mollis L. and Deschampsia flexuosa L. Trin. Journal of Ecology
37:68-81.
Kalkhoven, J. & P. Opdam, 1984. Classification and ordination of breeding bird
data and landscape attributes. In: J. Brandt & P. Agger (Editors): Methodology
in Landscape ecological research and planning. Universitetsforlag, Roskilde
Volume HI: 15-26.
Kanal, L., 1974. Patterns in pattern recognition 1968-74. IEEE Transactions on
Information Theory 20: 697-722.
Kant, G.R., 1982. Beken op de Veluwe. Hydrologische aspecten in relatie tot
de watervoering van de beken op de Oost- en Zuid Veluwe. Basisrapport 1
ten behoeve van de werkgroep Sprengen en beken. Provincie Gelderland,
Arnhem, 84 pp.
Kendall, D.G., 1971. Abundance matrices and seriation in archaeology. Zeitschrift
fur Warscheinlichkeitstheorie 17: 104-112.
Kilic, M., 1979. Soil variability near Follonica, Italy. MSc. Thesis. MSc Course
in Soil Science and Water Management, Agricultural University, Wageningen.

282

Kirkpatrick, J.B., P.R. Minchin & J.B. Davies, 1985. Floristic composition and
macroenvironmental relationships of Tasmanian vegetation containing bolster
plants. Vegetatio 63: 89-96.
Kooijman, S.A.L.M., 1977. Species abundance with optimum relations to en
vironmental factors. Annals of System Research 6: 123-138.
Krebs, J., 1978. Ecology, the experimental analysis of distribution and abun
dance. Harper and Row, New York, 648 pp.
Kruijne, A.A., D.M. de Vries & H. Mooi, 1967. Bijdrage tot de oecologie van
de Nederlandse graslandplant (with english summary). Verslagen Landbouwkundige Onderzoekingen 696. Pudoc, Wageningen, 65 pp.
Kruskal, J.B., 1964. Multidimensional scaling by optimizing goodness of fit to
a nonmetric hypothesis. Psychometrika 29: 1-27.
Kruskal, J.B. & J.D. Carroll, 1969. Geometric models and badness-of-fit functions.
In: P.R. Krishnaiah (Editor): Multivariate analysis II. Academic Press, New
York, p. 639-671.
Laboratory for Computer Graphics, 1968. SYMAP Manual, Graduate School
of Design, Harvard.
Lachenbruch, P.A., 1975. Discriminant analysis. Hafner Press, New York, 128
pp.
Lam, N.S., 1983. Spatial interpolation methods: a review. The American Car
tographer 10: 129-149.
Lambert, J.M. & M.B.Dale, 1964. The use of statistics in phytosociology. Advances
in Ecological Research 2: 59-99.
Lambert, J.M., S.E. Meacock, J. Barrs & P.F.M. Smartt, 1973. AXOR and
MONIT: two new polythetic-divisive strategies for hierarchial classification.
Taxon 22: 1733-176.
Lance, G.N. & W.T. Williams, 1967. A general theory of classificatory sorting
strategies. I Hierarchical systems. Computer Journal 9: 373-380.
Loucks, O.L., 1962. Ordinating forest communities by means of environmental
scalars and phytosociological indices. Ecological Monographs 32: 137-166.
Louppen, J.W.M. & E. van der Maarel, 1979. CLUSLA: a computer program
for the clustering of large phytosociological data sets. Vegetatio 40: 107-114.
Lozano, J.A. & J.D. Hays, 1976. Relationship of Radiolarian assemblages to
sediment types and physical oceanography in the Atlantic and Western Indian
Ocean. In: R.M. Cline & J.D. Hays (Editors): Investigation of late Quaternary
paleoceanography and paleoclimatology. Geological Society of America Me
moir 145: 303-336.
Mandelbrot, B.B., 1982. The fractal geometry of nature. Freeman, San Francisco,
460 pp.
Mardia, K.V., J.T. Kent & J.M. Bibby, 1979. Multivariate analysis. Academic
Press, London, 521 pp.
Matheron, G., 1971. The theory of regionalized variables and its applications.
Les Cahiers du Centre de Morphologie Mathematique de Fontainebleu. Ecole
Nationale Superieure des Mines de Paris.
McBratney, A.B., R. Webster, R.G. McLaren & R.B. Spiers, 1982. Regional
variation of extractable copper and cobalt in the topsoil of south-east Scotland.

283

Agronomie 2 (10): 969-982.


McCullagh, P. & J.A. Nelder, 1983. Generalized linear models. Chapman and
Hall, London, 261 pp.
Mcintosh, R.P., 1981. Succession and ecological theory. In: D.C. West, H.H.
Shugart & D.B. Botkin (Editors): Forest succession. Concepts and application.
Springer Verlag, New York, p. 10-23.
Mead, R. & D.J.P. Pike, 1975. A review of response surface methodology from
a biometric viewpoint. Biometrics 31: 803-851.
_.. .
Meijers, E.t 1986. Defining confusions-confusing definitions. Environmental monitoring and assessment 7: 157-159.
Meijers, E., W.J. ter Kcurs & E. Meelis, 1982. Biologische meetnetten voor het
beleid. Mededelingen van de Werkgemeenschap Landschapsecoiogisch Onderzoek9:51-58.
Mertz, D.B. & D.E. McCauley, 1980. The domain of laboratory ecology. Synthese
43:95-110.
Meulman, J. & W.J. Heiser, 1984. Constrained multidimensional scaling: more
directions than dimensions. COMPSTAT 1984, Physica-Verlag, Vienna, p. 137142.
Minchin, P.R., 1987. An evaluation of the relative robustness of techniques for
ecological ordination. Vegetatio 67: 1167-1179.
Montgomery, D.C. & E.A. Peck, 1982. Introduction to linear regression analysis.
Wiley, New York, 504 pp.
Morrison, D.F., 1976. Multivariate statistical methods. 2nd Edition. McGrawHill, Tokyo, 415 pp.
Mosteiler, F. & J.W. Tukey, 1977. Data analysis and regression. Addison-Wesley,
Reading, 588 pp.
Mueller-Dombois, D. & H. Ellenberg, 1974. Aims and methods of vegetation
ecology. Wiley, New York, 547 pp.
Naes, T. & H. Martens, 1984. Multivariate calibration. II Chemometric methods.
Trends in analytical chemistry 3: 266-271.
Neef, E., 1982. Stages in development of landscape ecology. In: S. Tjallingi &
A. de Veer (Editors): Perspectives in Landscape Ecology. Proceedings of the
International symposium of Landscape ecology at Veldhoven Netherlands.
Pudoc, Wageningen, p. 19-27.
Nelder, J.A. & R.W.M. Wedderburn, 1972. Generalized linear models. Journal
of the Royal Statistical Society, Series A 135: 370-384.
Nielsen, D.R. & J. Bouma (Editors), 1985. Soil spatial variability. Proceedings
of a workshop of the ISSS and the SSSA, Las Vegas, USA, 30 Nov. - 1 Dec.
1984. Pudoc, Wageningen, 243 pp.
Nishisato, S., 1980. Analysis of categorical data: dual scaling and its applications.
Toronto University Press, Toronto, 276 pp.
Noy-Meir, I., 1973. Data transformation in ecological ordination. I. Some
advantages of non-centering. Journal of Ecology 61: 329-341.
Noy-Meir, 1., 1974. Catenation: quantitative methods for the definition of coenoclines. Vegetatio 29: 89-99.
Noy-Meir, 1., D. Walker & W.T. Williams, 1975. Data transformations in ecological

284

ordination. II. On the meaning of data standardization. Journal of Ecology


63:779-800.
Odum, E.P., 1971. Fundamentals of ecology. 3rd Edition. Saunders, Philadelphia.
Orl6ci, L., 1966. Geometric models in ecology. I. The theory and application
of some ordination methods. Journal of Ecology 54: 193- 215.
Orl6ci, L., 1967. An agglomerative method for the classification of plant com
munities. Journal of Ecology 55: 193-206.
Parker, R.E., 1979. Introductory statistics for biology. Studies in biology,
No 43. Arnold, London, 122 pp.
Pavlidis, ., 1982. Algorithms for graphics and image processing. Springer Verlag,
Berlin, 416 pp.
Pielou, E.C., 1969. An introduction to mathematical ecology. Wiley, New York,
286 pp.
Pielou, E.C., 1976. Population and community ecology. Gordon and Breach, New
York, 432 pp.
Pielou, E.G., 1977. Mathematical ecology. 2nd Edition. Wiley, New York, 385
pp.
Pielou, E.C., 1984. The interpretation of ecological data. A primer on classification
and ordination. Wiley, New York, 263 pp.
Popma, J., 1982. Beken op de Veluwe. Numerieke classificatie van macrofauna
gemeenschappen van sprengen en beken op de Veluwe. Basisrapport 2 ten
behoeve van de werkgroep sprengen en beken. Provincie Gelderland, Arnhem,
132 pp.
Popma, J., L. Mucina, O. van Tongeren & E. van der Maarel, 1983. On the
determination of optimal levels in phytosociological classification. Vegetatio
52: 65-76.
Prentice, I.C., 1977. Non-metric ordination methods in ecology. Journal of Ecology
65: 85-94.
Prentice, I.C., 1980. Vegetation analysis and order invariant gradient models.
Vegetatio 42: 27-34.
Quinn, J.F. & A.E. Dunham, 1983. On hypothesis testing in ecology and evolution.
The American Naturalist 122: 602-617.
Ramensky, L.G., 1930. Zur Methodikder vergleichenden Bearbeitungund Ordnung
von Pflanzenlisten und anderen Objekten, die durch mehrere, verschiedenartig
wirkende Faktoren bestimt werden. Beitrage zur Biologie der Pflanzen 18:
269-304.
Rao, C.R., 1964. The use and interpretation of principal component analysis in
applied research. Sankhya A 26: 329-358.
Rao, C.R., 1973. Linear statistical inference and its applications. 2nd Edition.
Wiley, New York, 625 pp.
Ripley, ., 1981. Spatial statistics. Wiley, New York, 252 pp.
Risser, P.G., J.R. Karr & R.T.T. Forman, 1984. Landscape ecology directions
and approaches. A workshop held at Allerton Park, Piatt. County Illinois
1983. Illinois Natural History Survey Special Publication Number 2, 16 pp.
Rivas-Martinez, S., 1979. Brezales jarales de Europa occidental. Lazaroa 1:
5-127.

285

Rogister, J.E., 1978. De ecologische mR- en mN-waarden van de kruidlaag en


de humuskwaliteit van bosplantengezelschappen. Werken Proefstalion Waters
en Bossen (Groenendaal- Hoeilaart) reeks A 20: 1-29.
Roux, G. & M. Roux, 1967. A propos de quelques m6thodes de classification
en phytosociologie. Revue de statistique appliquee 15: 59-72.
Saaty, T.L., 1977. A scaling method for priorities in hierarchical structures. Journal
of mathematical psychology 14: 234-281.
Salton, G. & A. Wong, 1978. Generation and search oj" cluster files. Association
for computing machinery transactions on database systems 3: 321-346.
SAS Institute Inc., 1982. SAS user's guide: statistics, 1982 Edition. SAS Institute
Inc., Cary, N.C., 584 pp.
Schaafsma, W. & G.N. van Vark, 1979. Classification and discrimination problems
with application, part Ha. Statistica Neerlandica 33: 91-126.
Scheffe, H., 1973. A statistical theory of calibration. Annals of Statistics 1:
1-37.
Schiffman, S.S., M.L. Reynolds & F.W. Young, 1981. Introduction to multi
dimensional scaling. Theory, methods and applications. Academic Press,
London, 413 pp.
Schwartz, L.M., 1977; Nonlinear calibration. Analytical chemistry 49: 2062-2068.
Seber, G.A.F., 1977. Linear regression analysis. Wiley, New York, 465 pp.
Sheenan, P.J., 1984. Effects on community and ecosystem structure and dynamics.
In: P.J. Sheenan, D.R. Miller, G.C. Butler & Ph. Bourdeau (Editors): Effects
of pollutants at the ecosystem level. Wiley, New York, p. 51-99.
Shepard, R.N., 1962. The analysis of proximities: multidimensional scaling with
an unknown distance function. II. Psychometrika 27: 219-246.
Shepard, D., 1968. A two-dimensional interpolation function for irregularly spaced
data. In: Proc. Ass. Comput. Mach. 517-523.
Sibson, R., 1972. Order invariant methods for data analysis. Journal of the Royal
Statistical Society, Series 34: 311-349.
Simberloff, D., 1983. Competition theory, hypothesis-testing, and other community
ecological buzzwords. The American Naturalist 122: 626-635.
Slob,. W., 1987. Strategies in applying statistics in ecological research. Free
University Press, Amsterdam, 112 pp.
Sneath, P.H.A. & Sokal, R.R., 1973. Numerical taxonomy. W.H. Freeman, San
Fransisco.
Snedecor, G.W. & W.G. Cochran, 1980. Statistical methods. 7th Edition. The
Iowa State University Press, Ames, 507 pp.
Sokal, R.R. & C D . Michener, 1958. A statistical method for evaluating systematic
relationships. University of Kansas Science bulletin 38: 1409-1438.
Sokal, R.R. 8L F.J. Rohlf, 1981. Biometry. 2nd Edition. Freeman, San Fransisco,
877 pp.
Serensen, ., 1948. A method of establishing groups of equal amplitude in plant
sociology based on similarity of species content. Det. Kong. Danske Vidensk.
Selsk. Biol. Skr. (Copenhagen) 5 (4): 1-34.
Southwood, T.R.E., 1978. Ecological methods, with particular reference to the
study of insect populations. 2nd Revised edition. Methuen, London, 391 pp.

286

Strong jr., DR., 1980. Null hypotheses in ecology. Synthese 43: 271- 285.
Swain, P.H., 1978. Fundamentals of pattern recognition in remote sensing. In:
P.H. Swain & S.M. Davies (Editors): Remote sensing: the quantitative
approach. McGraw-Hill, New York.
Swaine, M.D. & P. Greig-Smith, 1980. An application of principal components
analysis to vegetation change in permanent plots. Journal of Ecology 68:
33-41.
Tacha, T.C., W.D. Warde & K.P. Burnham, 1982. Use and interpretation of statistics
in wildlife journals. Wildlife Society Bulletin 10: 355-362.
Taylor, C.C. & P.A. Burrough, 1986. Multiscale sources of spatial variation in
soil III. Improved methods for fitting the nested model to one-dimensional
semivariograms. Mathematical Geology 18: 811-821.
ten Berge, H.F.M., L. Stroosnijder, P.A. Burrough, A.K. Bregt & M.J. de Heus,
1983. Spatial variability of physical soil properties influencing the temperature
of the soil surface. Agricultural Water Management 6: 213-216.
ter Braak, C.J.F., 1982. DISCRIM - A modification of TW1NSPAN (Hill, 1979)
to construct simple discriminant functions and to classify attributes, given
a hierarchical classification of samples. Report 82 ST 10756. TNO Institute
of Mathematics, Information processing and Statistics, Wageningen.
ter Braak, C.J.F., 1983. Principal components biplots and alpha and beta diversity.
Ecology 64: 454-462.
ter Braak, C.J.F., 1985. Correspondence analysis of incidence and abundance data:
properties in terms of a unimodal reponse model. Biometrics 41: 859-873.
ter Braak, C.J.F., 1986a. Canonical correspondence analysis: a new eigenvector
technique for multivariate direct gradient analysis. Ecology 67: 1167-1179.
ter Braak, C.J.F., 1986b. Interpreting a hierarchical classification with simple
discriminant functions: an ecological example. In: E. Diday et al. (Editors):
Data analysis and informatics 4. North Holland, Amsterdam, p. 11-21.
ter Braak, C.J.F., 1987a. The analysis of vegetation-environment relationships
by canonical correspondence analysis. Vegetatio 69: 69-77.
ter Braak, C.J.F., 1987b. CANOCO - a FORTRAN program for canonical
community ordination by [partial] [detrended] [canonical] correspondence
analysis, principal components analysis and redundancy analysis (version 2.1)
Agricultural Mathematics Group, Wageningen, 95 pp.
ter Braak, C.J.F., 1988. Partial canonical correspondence analysis. In: H.H. Bock
(Editor): Classification and related methods of data analysis. North Holland,
Amsterdam, p. 551-558.
ter Braak, C.J.F. & L.G. Barendregt, 1986. Weighted averaging of species indicator
values: its efficiency in environmental calibration. Mathematical Biosciences
78: 57-72.
ter Braak, C.J.F. & C.W.N. Looman, 1986. Weighted averaging, logistic regression
and the Gaussian response model. Vegetatio 65: 3-11.
Titterington, D.M., G.D. Murray, L.S. Murray, D.J. Spiegelhalter, A.M. Skene,
J.D.F. Habbema & G.5. Gelpke, 1981. Comparison of discrimination techniques
applied to a complex data set of head injured patients (with discussion). Journal
of the Royal Statistical Society, Series A 144: 145-175.

287

Toft, . . & PJ. Shea, 1983. Detecting community-wide patterns: estimating power
strengthens statistical inference. The American Naturalist 122: 618-625.
Torgerson, W.S., 1958. Theory and methods of scaling. Wiley, New York, 460
PP.
Tso, M.K-S., 1981. Reduced-rank regression and canonical analysis. Journal of
the Royal Statistical Society Series 43: 89-107.
Tutin, T.G. et al. (Editor) 1964-1980. Flora Europaea. Cambridge University Press,
Cambridge,
van Dam, H., G. Suurmond & C.J.F ter Braak, 1981. Impact of acidification
on diatoms and chemistry of Dutch moorland pools. Hydrobiologia 83: 425459.
van de Nes, T.J., 1980. Introduction to a system approach for water management
in Gelderland, The Netherlands. In: Water resources management on a regional
scale. Committee for hydrological research TNO, Proceedings and Information
no 27, p. 9-22.
van der Maarel, E., 1979a. Multivariate methods in phytosociology, with reference
to the Netherlands. In: M.J.A. Werger (Editor):The study of vegetation. Junk,
The Hague, .61-226.
van der Maarel, E., 1979b. Transformation of cover-abundance values in phy
tosociology and its effects on community similarity. Vegetatio 39: 97-114.
van der Maarel, E., L. Orl6ci & S. Pignatti, 1976. Data processing in phyto
sociology, retrospect and anticipation. Vegetatio 32: 65-72.
van der Maarel, E., J.G.M. Janssen & J.M.W. Louppen, 1978. TABORD, a
program for structuring phytosociological tables. Vegetatio 38: 143-156.
van der Meijden, R., E.J. Weeda, F.A.C.B. Adema & G.J. de Joncheere, 1983.
Flora van Nederland. 20th Edition. Wolters-Noordhoff, Groningen, 583 pp.
van der Steen, W.J., 1982. Algemene methodologie voor biologen. Bohn, Scheltema
and Holkema, Utrecht, 115 pp.
van Tongeren, O., 1986. FLEXCLUS, an interactive program for classification
and tabulation of ecological data. Acta Botanica Neerlandica 35: 137-142.
Varmuza, K., 1980. Pattern recognition in chemistry. Springer Verlag, Berlin, 217
PP.
Vevle, O. & K. Aase, 1980. Om bruk ov oekologiske faktortall i norske plantesamfunn. In: K. Baadsvik, T. Klokk & O.I. R6nning (Editors): Fagmoete
i vegetasjonsoekologi pa Kongvoll 16- 18.3.1980. Kgl. Norske Videnskabers
Selskab Museet, Botanisk serie 1980-5, p. 178-201.
Vink, A.P.A., 1963. Planning of Soil Surveys in Land Development. Publication
of the International Institute for Land Reclamation and Improvement (ILRI)
10, 53 pp.
von Humboldt, A., 1806. Ideen zu einer Physiognomik der Gewachse. Cotta,
Stuttgart.
Webster. R., 1984. Elucidation and characterization of spatial variation in soil
using regionalized variable theory. In: G. Verly et al. (Editors): Geostatistics
for Natural Resources Characterization, Part 2, Reidel, Dordrecht, p. 903913.
Webster, R., 1985. Quantitative spatial analysis of soil in the field. Advances in

288

Soil Science, Volume 3. Springer-Verlag, New York.


Webster, R. & T.M. Burgess, 1983. Spatial variation in soil and the role of kriging.
Agricultural Water Management 6: 111-122.
Webster, R. & T.M. Burgess, 1984. Sampling and bulking strategies for estimating
soil properties in small regions. Journal of Soil Science 35: 127-140.
Webster, R. & S. Nortcliff, 1984. Improved estimation of micronutrients in hectare
plots of the Sonning Series. Journal of Soil Science 35: 667-672.
Westhoff V. &. E. van der Maarel, 1978. The Braun Blanquet approach, In: R.H.
Whittaker (Editor): Classification of plant communities. Junk, The Hague,
p. 287-399.
Whittaker, J., 1984. Model interpretation from the additive elements of the
likelihood function. Applied Statistics 33: 52-65.
Whittaker, R.H., 1956. Vegetation of the Great Smoky Mountains. Ecological
Monographs 26: 1-80.
Whittaker, R.H., 1967. Gradient analysis of vegetation. Biological Reviews 49:
207-264.
Whittaker R.H., 1973. Approaches of classifying vegetation. In: Whittaker R.H.
(Editor): Ordination and classifiction of communities. Junk, The Hague, p.
325-354.
Whittaker, R.H., S.A. Levin & R.B. Root, 1973. Niche, habitat and ecotope.
American Naturalist 107: 321-338.
Wiens, J. A. & J.T. Rotenberry, 1981. Habitat associations and community structure
of birds in shrub steppe environment. Ecological Monographs 51: 21-41.
Williams, B.K., 1983. Some observations on the use of discriminant analysis in
ecology. Ecology 64: 1283-1291.
Williams, E.J., 1959. Regression analysis. Wiley, New York, 211 pp.
Williams, W.T. (Editor), 1976a. Pattern analysis in agricultural science. CSIRO
Melbourne, Australia.
Williams, W.T., 1976b. Hierarchical divisive strategies. In: W.T. Williams (Editor):
Pattern analysis in agricultural science. CSIRO, Melbourne, Australia.
Williams, W.T. & J.M. Lambert, 1959. Multivariate methods in plant ecology.
I. Association analysis in plant communities. Journal of Ecolology 47: 83101.
Williams, W.T. & J.M. Lambert, 1960. Multivariate methods in plant ecology.
II The use of an electronic digital computer for association-analysis. Journal
of Ecology 48: 689-710.
Williams, W.T. & J.M. Lambert, 1961. Multivariate methods in plant ecology.
III Inverse association-analysis. Journal of Ecology 49: 717-729.
Williams, W.T, J.M. Lambert & G.N. Lance, 1966. Multivariate methods in plant
ecology. V. Similarity-analysis and information-analysis. Journal of Ecology
54:427-445.
Williamson, M., 1972. The analysis of biological populations. Arnold, London,
180 pp.
Wishart, D., 1978. CLUSTAN user manual. Program library unit. Edinburgh
University Press, Edinburgh, 175 pp.
Wold, H., 1982. Soft modelling: the basic design and some extensions. In: K.G.

289

Joreskog 8L H. Wold (Editors): Systems under indirect observation, il. North


Holland, Amsterdam, p. 1-54.
Yarranton, G.A., 1969. Plant ecology: a unifying model. Journal of Ecology 57:
245-250.
Yarranton, G.A., 1970. Towards a mathematical model of limestone pavement
vegetation. III. Estimation of the determinants of species frequencies. Canadian
Journal of Botany 48; 1387-1404.
Zelinka, M. & P. Marvan, 1961. Zur Przisierung der biologischen {Classification
der ReinheitfliessenderGewasser. Archiv fCir HydroBTologie 57: 389-407.
Zhang, L., 1983. Vegetation ecology and population biology of Fritillaria meleagris
L. at the Kungsangen nature reserve, eastern Sweden. Acta Phytogeographica
Suecica 73: 1-92.

290

:
- . . .: . 1967. 356.
. : . : . 1981.251.
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. . . .. 1972. 486 .
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. ., . , . . :
. .: . 1985. 487.
. ., . , . . :
. .: . 1983. 412.
. ., 3. ., . . .
.: . 1974. 240.

291

. ., . . .
: . 1983.464.
., . :
. : . 1980. 510.
., . :
. .: . 1981. 627.
. ., . . .:
. 1976. 736.
. ., . . .: . 1973. 899.
. . .:
. 1988. 349.
. . .:
. 1982. 344.

292

, , ,
.

(.
)

()

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

()




(.
)



(. )

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

(.
)

122, 137, 145, 147, 159


(.
)

62,83,97,100,139,141,160, 164,
194
61-64, 68, 83-88,93, 97,112, 154
237-238
150,168
174


, .

- (.
-)
-





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)




(.

)

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5-7,153-156,180-183,
219-220,
191
245
13, 56,139,201
4, 7-8,10, 15
242
16, 60-61, 175, 197, 202, 220
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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)


, .




(.
)

()



(.
)
(.
)




35,40,65-66, 67,72-74,173
103,131,183
132-134,166
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