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.
Preface
This book relates ecology of fungi on woody plants. In it, role and
place of fungi in life of trees and shrubs are considered quite in a new fashion.
We avoided deliberately the stereotype account of the material that always is
present in such works. Unlike our predecessors, we consider fungi not
parasites of plants but their friends. The plant itself seems to us be a complex
of symbiotrophic micro-organisms and the fungi are a necessary, obligatory
component of this complex system. In the kingdom of fungi bound to plants
there is no chaos and disorder, all fungi occupy their own, strictly definite
ecological niches. The existent location order of fungi is in variable in range
of the plant species and genus. Each organ of a plant, each stage of
development, its condition is fixed by own species of fungi or by their
developmental stages. We accepted the conception of multidimensional
ecological niche proposed by J. Hutchinson which allowed to single out on
woody plants up to 100 ecological niches and to confirm their singling out by
corresponding fungi-indicators.
We have studied what processes occur on a plant or its parts after their
dying off. The established laws of succession changes of fungi are typical for
all plant species irrespective of their life form. This gave us an opportunity to
explain biological variety of fungi in a concrete ecological niche and also to
get new information on development of important phytopathogenic fungi.
Simultaneous study of questions of mycoflora formation in aboriginal
and introduced plants allowed to state that the condition of geographical
vicarisms widely spread in animate nature is also peculiar to fungi.
Knowledge of this law makes it possible to predict discovery of new species
on woody plants, appearance of quarantine species, to settle arguable points
on phylogeny of fungi and plants, introduction theory ets.
In the book, main principles of drawing up fungus or mycological
maps of woody plants are elucidated. These original methods allow to collect
and systematise mycological information on 80-100 fixed parameters
purposefully for all plants under studies. This makes it possible to give
sufficiently exact information about viability of any revealed fungus species
on a plant, or on their complex, on its adaptation degree, on influence of
environmental factors on the fungus condition, to determine causes and terms
of death of certain organs. The established laws of development and spreading
of fungi are typical for all plants irrespective of their provenance.
In the book, questions of biology and ecology of fungi on plants from
different regions of the world, types of life forms growing in different zones
of the area considered in detail. Original data on formation terms of fruit
bodies of micromycetes and polypores, on seasonal and daily development of
these fungi, on dissemination extent of spores in plantations and others are
presented. Author has conducted observations on such a rare phenomenon in
life of fungi as formation of anomalous spore-bearing and rise of sterile
forms of fungus development.
6
Disease is considered as a disturbance of ecological equilibrium.
Number of diseases on plants depends on number of ecological niches which
we can establish and confirm by isolated fungi-indicators. A brief critical
review of basis theories of plant immunity is given and causes hindering from
formulating common biological theory are expressed. On a basis of the
established laws of fungi development, we propose to reader our own view of
this problem.
In the book’s method part information on methods of account and
control of fungi in plantations and on woody plant, and also on the difficulties
which can arise at their study are elucidated.
The book was compiled taking into account psychological, emotional
perception of the material by an investigator who has been brought up on
settled ideas of this matter. For this purpose, philosophical, logical approaches
were used which allow to approach the question of fungi on trees without
prejudice. All propositions about fungi we expressed can be verified easily on
any plant and in any place.
In Supplement examples of drawing up of mycological maps of some
plant species are given and the reader himself can check correctness of the
established laws.
The book is intended for the very general public; for ecologists,
mycologists, phytopathologists, botanists, specialists of plant protection and
nature conservation, for nature-lovers who take an interest in the world of
fungi.
7
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X " A %=
>
- &
?>'
=
4
Cercis siliguastrum
Ficus carica
Fraxinus oxycarpa
Juglans regia
Malus spectabilis
Pistacia mutica
Platanus acerifolia
*Prunus divaricata
**Prunus divaricata
Salix babylonica
'& ''
min
/0
2,5
3,0
3,0
4,0
1,5
2,8
.
%
,
? A! " % ! B Diplodia X!
8
(B
!X " A %
)
I
4'>
opt
max
min
opt
R. 3.-.
R. 0/3/ R
3,0-6,0
9,0
4,0
5,0-7,0
4,0-7,0
9,0
4,5
6,0-8,0
4,0-6,0
6,5
4,6
6,0-8,0
5,0-6,0
7,5
5,0
6,0-8,0
2,0-5,0
6,0
4,0
6,0-7,0
3,0-6,0
10,5
3,0
5,0-7,0
2,0-4,0
6,5
6,0-8,0
3,0-4,5
3,5
5,0-7,0
1,5-3,5
1,5
5,0-7,0
1,5-2,5
5,0
3,0-5,0
2,9-5,1
7,5
3,8
5,3-7,2
-.1 /I.4. R. 3.-.
R. 0/3/ R
3,5
10,0
5,0
Arbutus andrachne
3,5-8,0
Eriobotrya japonica
5,0-7,0
Ilex aguifolium
2,3-5,0
7,0
5,0
6,0-8,0
Laurocerasus officinalis
2,0
3,0-6,0
9,0
1,5
Laurus nobilis
3,0-6,0
Photinia serrulata
6,0-7,0
Magnolia grandiflora
1,5-2,5
Olea europea
1,8-3,5
6,0
3,0
Quercus ilex
2,0
3,0-7,0
9,0
4,0
Phillyrea latifolia
2,0
3,3-5,6
8,2
3,7
'& ''
*
%
; **
.
+
2,0-2,8
- 2,2
. ?
: 7,5-8,2
,
,
"
7,7
47
.
6,0-8,0
5,0-6,0
2,5-6,0
9,0
6,0-7,5
4,0-10,0
6,0
5,0-6,0
6,0-8,0
5,2-7,3
max
8,0
9,5
11,0
7,5
10,0
12,0
9,0
6,5
9,2
9,0
10,0
7,5
8,9
D
.
/
3,3-5,6
2,9-4,9
,
2,9-5,1
3,0-6,0
,
,
,
3,2-4,9
3,1-5,3
.t
,
,
"
.
+
"
,
%
$
. +
,
,
5,3-7,2
5,2-7,0
.(
$
. /
$
5,2-7,3
4,3-6,1
,
,
,
,
4,9-6,9
,#
4,9-6,9
.2
,
"
"
$
. D
(
,
$
$
. 4-2). 9
!
,
,
!
,
.
E
8
1
2
3
4
5
. 4-2.
5
9
Diplodia # !#
2
!
'
6
6
7
/
L+
#
8
9
,
2
6
+
+ #
+ !
$!
,
%
"
$
,
.
48
"
4.4
2
A
!
% ""1
. D
,
,
+
6
,
,
$%
>
, "
Diplodia Phomopsis
Microdiplodia, Haplosporella, Coniothyrium
. D
Diplodia
(
$%
$
.
$
.
!
"
. 4-3). +
,
$
!
#
!
.
,
"
"
.
6
"
,
,
,
. B
"
.,
.
1
2
PHOMA
CAMAROSPORIUM
DIPLODIA
CYTOSPORA
1
2
. 4-3. . M
+ !6 $ #
3
4
'
6 !
2-
5
6 7 8
/
L #!K )L 6
): 1 - !
"$! !
" !#
(
+ #
9
10
,
2 6
+ # !
!
(
+ #;
A
. r
$
,
$
.
,
!
Cytospora leucosperma.
49
11
,
$
(
A
B
1
4
3
4
C
3
7
1
7
1
1
7
1
2
6
7
1
1
5
1
,
. 4-4).
. 4-4. !
!
+ # !$ #
6 !
(6 ! '
6 K$
(: $"
, $ # , - !+
# $ . 1 - Phoma, 2 - Camarosporium, Hendersonia, 3 - Diplodia, 4 Phomopsis, 5 - Cytospora, 6 - Aphyllophorales, 7 !
#!
# $! !
)
+
( !+ .4-3).
7 "8 9 4-3. ' X!
!
VB
X % =!
& Y$ 8 A
$
?
!
> ! =8
A X!
S(
) F'?( =
@ +
&@ ! B % A! " %
% $! @ B! % &@
@ B #
')
> ;* & )
(>29D/A|t>2‹
1
Phoma
1
Hendersonia, Camarosporium
2-3
Sphaeropsis, Diplodia
4-8
Phomopsis, Cytospora
8-15
Cytospora (
! )
15
Aphyllophorales
+8D/D ‹t 9)#2t>(~
1
Phoma
50
S(
) F'?( =
@ + ')
> ;* & )
1-15
Cytospora, Phomopsis, Sphaeropsis
)4D9(qt>>‹t (/‹
1-15
#
Deuteromycotina
Ascomycotina
D
15-18
"
IV
!
. .
!
!
. #
!
I
,
! ,
,
( !+ . 4-4).
7 "8 9 4-4.
$!
?
"
9
% % A
B! % &@ !
S(
) F'?( '
@
)
9
0 ') IV + =&(
0 ') III + =&(
0 ') II + =&(
0 ') I + =&(
5
> ?+
9;' )
;
> B;
?
> B;
93
7
92
6
90
10
82
18
.
,
I
$
!
! Z &@ $ 8 ! Z &@
% &@ X " A @
>, %
5
Ascomycotina
> B;
?
> B;
73
27
67
33
59
41
44
56
$%
,
! ,
. )
%
!
.
"
$
:
!
.
B
"
,
$
.
"
.
,
"
"
"
4.5
.
3
1
B
!
). A
$
,
,
$
+ !
K
6 2
,
,
"
"
51
% "
!
$
. +
$
!
$!
+ #
,
$
I-II
. 2
: Phoma, Camarosporium,
Diplodia, Phomopsis, Cytospora. B
:
80 %
. A
,
$
( !+ . 4-5).
7 "8 9 4-5.
$
,
B $
! & ! B& A! " % X
X
Z ! B! % &@ !
> (]!& )
Y E '
=
=B
> ;* & )
) F'?( = @ +
I-II + =&(
a7'w (x 'w^baayz 'z^z ayz 'b 7za {
Cytospora
Phomopsis
S(
/
/
•
.
.
,
!
$%
Cytospora
+
Camarosporium
Phoma, Diplodia
zaayz <w'w y
#
Ascomycotina
8
bfw'
/
.
.
')
Cytospora
Diplodia
Cytospora
%
.
"
&@
,
,
#
Ascomycotina
D
$
..
$
$
,
"
,
2
" $%
"
10
$
,
.
20 %,
. A
"
!
!
,
52
"
"
. /
.
/
,
%
"
.B
.2
!
.t
!
"
(
, Amygdalus
, A. nana ),
,
$
"
!
,
Crataegus, Corylus, Elaeagnus, Rhamnus). A
communis .)
(
,
2
Cytospora.
"
,
9
Diplodia,
%
"
,
Cytospora
I-IV
Phomopsis,
$
% $
5-6
!
.
Phoma, Camarosporium,
. 2
$. D
,
,
+
"
%
,
.
2-3
,
Cytospora,
–
,
,
,
$
Diplodia, Camarosporium,
Cytospora, Phomopsis "
Phoma. >
$ . E
"
"
4.6
"
"
(
1
!
!
.
+
>
. .
!
, !
,
,
. B
.>
“
(
+
”
!
. 4-5).
())
!
,
%
. +
"
$
,
.D
,
,
,
,
%
. +
$
. 9
53
,
B
!
.
,
!
.
>
+
>
>
D
>
-
D
>
>
+
. 4-5. 0
9
#
'
6
"
( , , , )2
L
+ # !$ #
( )
#! ( “!
!
6 !
(6
6”
$
,
%
.
(Acer negundo)
. 9
54
.9
,
,
"
,
$
. +
) !
"
$
2
(
,
,
$
.
(#)
"
,
,
!
,
. D
,
.
,
. B
,
$
. r
,
"
r
,
,
!
$
$
.
(D)
$
.D
!
"
"
,
.B
,
. t
$
,
. +
"
,
$
"
C
. {
.
?
!
.>
"
%
$
,
.
?
"
Diplodia rhamni
Cucurbitaria
rhamni
Fusarium
lateritium
RHAMNUS ALATERNUS, 2-8
. 4-6. ! #
!
Diplodia rhamni
6# $ #
+ ##9
# K$
(6 2alaternus
55
>X " A
'
6
+
) L
+!
# Rhamnus
Diplodia
Cytospora
. 4-7. ! #
+ #
$! Cytospora # 9
!
+ #
L!6
CLERODENDRON
TRICUSPIDATA
CORIARIA JAPONICA
Phomopsis
Cytospora
Diaporthe
Phomopsis
. 4-8. !
!6 2
+ # Diplodia
$ 6 !
"4 ( !
(6 #
!
# $
+ #
+!
56
(
!
!
# 5
!$
$
6
q
"
"
"
(
4-6),
(
"
. 4-7),
$
%
(
$
,
!
$
$%
.
57
. 4-8).
:
.
,
5.
8
<
5.1
2; 4
4
!
% "
5
% !
+
,
$
. B
,
"
. 2
%
!
–
,
,
. D
,
,
"
.?
,
$ $
9
"
$%
.
,
. q
$
% $
,
"
" $
!? , ]! X %
Butcher, 1969).
%
.+
$
,
(|
`@ , 1965; _ 8
%, 1947;
% , 1954; Meredith, 1960; Hudson, 1968;
:
,
–
$%
,
, 1969; wA
9
$
,
! . +
,
"
.
,
"
+
%
,
,
.(
(Tomovic, Karadzic, 1988; Chapela, 1989; Chapela, Boddy,
1988).
?
III-IV
.
C
,
,
.
50
%
,
"
"
!
:
,
10
.D
"
>
50
(
"
)
$%
,
"
( $ ).
(40,
$
.
"
>
,
"
)
10
. D
,
,
. t"
.+
%
,
.>
(~
$
"
)
,
,
,
58
,
"
.
"
"
.
"
18
,
.
.
%
%
.
24
,
,
4
"
6
. .
!
:
,
. .
"
# %
"
"
,
,
-
.
-
!
"
Basidiomycotina
.
,
!
,
. D
,
!
"
,
$%
Ascomycotina,
.
50
!
Ascomycotina
(35,5
%).
32
(64,5 %)
$
.
Sphaeropsidales
17
.q
Diplodia
$
, Cytospora –
60 %, Phomopsis – 22 %, Phoma – 20 %.
Melanconiales
,
Pestalotia (5 %).
#
!
12
,
(35,5 %).
- Valsaceae, Diaporthaceae, Nectriaceae ,
. q
Valsa
Diaporthe
$
18 %
, Otthia – 20 %.
>
31
!
,
93 %
$
(34,4 %),
Ascomycotina (58,6 %),
7%
Basidiomycotina. #
$
Pyrenomycetes (67 %),
Helotiales (23 %)
Pseudosphaeriaceae. D %
17. /
$
Diatrypaceae
(21 %),
Diatrypella
24 %
.
/
$%
$ 10
(34 %),
:
Melanconiales (4
), Sphaeropsidales (3), Moniliales (3). /
$
Tubercularia (
18 %
), Coleophoma (26 %),
Fusarium (14 %).
/
$
(54,8 %),
90 %
q
59
q
Stereaceae,
$
Basidiomycotina
Peniophora, Lopharia. q
,
18
Lopharia.
>
,
(100 %). /
$
Sphaeropsidales,
$
Diplodia (100
%
) Cytospora (90 80 %).
- Diplodia, Cytospora, Phomopsis ! .
>
$
"
,
%
. q
Diplodia
$
(100 90 %), Cytospora - " ,
%
(60 50 %).
>
Ascomycotina
,
,
. +
!
% ,
.
Eutryblidiella
Cenangium
.
"
$ ?
,
$
Diplodia (80 90 %) Cytospora
(50
60 %). .
%
,
$
! ,
" ,
! .
?
$
"
. q
$
Diplodia (90
100 %),
Phoma (50 40 %), Cytospora (20
40 %). q
Cytospora
10
$
! ,
Diplodia
Phomopsis
.
2
,
$%
$
$%
:
Diplodia 90 %, Cytospora 6 0%, Phomopsis 22 %, Phoma 20 %,
Camarosporium 10 %;
: Diplodia 88 %, Cytospora 68 %, Phoma
18 %, Phomopsis 8 %.
10%.
$%
!
,
Sphaeropsidales.
!
$
$%
: Coleophoma 26 %, Tubercularia 18
%, Fusarium 14 %;
: Coleophoma 26 %,
Diatrypella 24 %. +
$%
.
$
Tubercularia 30 %,
Coleophoma 50 %, Diatrype 30 %. A
$
Coleophoma 60 %,
.
,
60
>
$
%
Nectria – 40 %,
oleophoma – 30 %,
Coleophoma -30 %,
Diatrypella –
%
Diatrypella – 40 %. 2
,
,
24
$
Sphaeropsidales. +
40 %. (,
,
Fusarium – 50 %,
Moniliales
" $
Sphaeropsidales Pyrenomycetes.
4
$
!
,
%
.
!
,
,
2,6:2,3. D
!
(Bolay, Spring, Monney, 1990).
A
!
,
14 %
! .
!
$
%–
42 %,
;
,
43
,
61 %,
12
!
,
26
20 %. 2
, 33
, 32
25 %,
,
-
%
;
%
.
!
(29
"
(42 %),
%).
>
( !+ . 5-1).
#
!
3,8
–
3,5
(
(
8,6 (
)
6,3
(
). +
$
)–
),
,
.
#
$ ,
. )
#
)
"
+
(Chapela, Boddy, 1988; Chapela, 1989). +
$
25
6
$
61
.
!
,
%
$%
.
7 "8 9 5-1. ! B %=% ?
`$9
> A! " % X 8
Y E '
B
?>'
=
& X $ =
8
8
B >
!
# X " A % (%
#9 @)
- ' =
>
=
=
+
/
') 9
E
'>
/
E
'>
/
E
'>
.
E
'>
.
E
'>
> &
1
2
3
7,2
9,6
12,8
3,8
5,6
8,5
6,8
8,6
13,4 14,5 17,3 18,0 22,0
5,1
8,5
14,2 14,6 21,8 23,0 23,0 23,0
8,6
13,1
14,7 15,6 19,0
6,3
10,0
11,4 14,3 20,0 23,0
3,8
8,6
15,1 20,1 23,0 23,0 23,0
24,0
6,4
8,9
13,7 17,0 17,0 18,0 20,0
20,0
7,3
10,0
11,0 12,8 14,0 15,7 20,5
3,5
8,0
8,0
+
!
? (Z'?? *
4
5
6
7
8
13,6 14,5 15,3 20,0 23,0
13,8 15,7 21,1 23,0
8,6
24,0
23,0 23,0
,
"
"
,
,
#
(8),
–
(5-6).
.t
$
$
,
,
"
"
(
62
. 5-1).
>;
? (Z'?;
?
=& 9
> &
I
? (Z'?? *
II
III
/
1
)
!
2
2
$%
!
/
3
$%
/
$%
/
. 5-1. !
+
!
#9
$%
$%
/
4
/
$%
!
/
2
IV
$%
"
'
6 ($ #
L!6
6
+ #
$
.E
$
!
,
! .B
"
5
15
. >
$
.
!
/
•
. 4
$
$%
!
!
20-23
$
75 %
Pinus, Juniperus, Cupressus, Platycladus,
,
.
.
"
.
2
,
"
.
$
$
,
!
!
$
. D
$
!
!
! . +
$%
63
,
23
. •
Seguoia, Seguoiadendron,
Metaseguoia, Taxodium, Cryptomeria, Torreja, Taxus,
"
,
$
!
: Ginkgo, Exochorda, Lagerstroemia, Acca, Aucuba,
Paulownia.
r
$
,
,
%
.
"
$%
Telephoraceae,
Stereaceae, Corticiaceae,
$
,
3
. B
..
Lonicera,
Weigela, Deutzia, Buddleija, Chimonanthus, Eucomya, Jasminum, Sambucus,
Staphylea
.
"
!
. +
,
,
.>
,
"
! ;
!
$
$%
;
!
!
$
! ,
$%
,
$%
. . )
$
.+
,
"
10
15
.
>
,
,
“
$% ”
,
,
.
/
"
,
%
,
. D
"
"
$
,
. +
,
$%
$%
100 %-
"
70-
80 %
. t
!
. B
100
. D
,
-
100 %50 %
!
$
,
"
,
. q
$
!
$%
.
64
+
–
. ?
,
,
,
!
% ,
, t
C
,
$
$
%
“
”
,
Phoma, Hendersonia, Camarosporium. Diplodia. +
,
,
$
.+
"
$
5-2).
. >
!
.
q
(
.
/
2
3
4
6
5
100%
+
70%
,
>
!
,
,
DIPLODIA SP.
. 5-2.
!
'
!$
!#
!#
$ !
( $
# $!
! + !
+! #
#
,
$
$
“
,
3
,
. +
(
”
8
%
( !+ . 5-2).
"
10-12
"
#
!
, 15,3
$
,
. B
,
65
)
. A
,
,
, 10,3
$
3
8
. .
"
,
,
.
7 "8 9 5-2. ! B
X $ =
8 ! $ % X #%8 #
X! B 8
8V
! =%
#"
! Z &@ $!
9
%
!
#@ ! =8 &@ X %
=
&@ Z ! , %
#9 @
(]!& , ($!
)
s
!
!( 2
(
!
!'! 2
$ # 6
#! (
+ #
! (
E
#
!$
%
$
K
$! ! #
+ #
(
7,2
'>
4,3
E
15,2
'>
13,2
7,7
5,5
13,5
11,9
10,6
7,5
11,5
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Celtis occidentalis
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Laburnum
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Phomopsis
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Microdiplodia
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Tubercularia
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Valsa cypri
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Podosphaera
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83
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D3
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Phyllactinia guttata
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Stereum hirsutum
Ustulina vulgaris
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Phellinus igniarius
C0
Podosphaera tridactyla
D2
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D3
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C2
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C3
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D1
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Phellinus igniarius
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9
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. schulzeri –
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:
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ytospora - 2
ytospora - 3, ytospora - 4.
ytospora - 3
ytospora - 4.
ytospora - 4
***********
Phomopsis - 1
Phomopsis - 2, Phomopsis - 3, Phomopsis - 4.
Phomopsis - 2
Phomopsis - 3, Phomopsis - 4.
Phomopsis - 3
Phomopsis - 4.
Phomopsis - 4
************
Diplodia - 1
Diplodia - 2, Diplodia - 3, Diplodia - 4.
Diplodia - 2
Diplodia - 3, Diplodia - 4.
Diplodia - 3
Diplodia - 4.
Diplodia - 4
************
q : Cytospora - 1 -
,
Cytospora - 3 -
; Cytospora - 2 ; Cytospora - 4 -
"
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M A L U S
S Y L V E S T R I S
20
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4
1
3
2
1
2
3
4
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Cytospora leucostoma
Cytospora rubescens
Cytospora schulzeri
1
1
1
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1
1
1
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1
1
1
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1
1
1
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1
1
1
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2
1
1
2
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1
1
1
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2
1
1
2
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Sphaeropsis malorum
1
1
1
Diplodia malorum
1
1
2
1
1
3
1
1
4
1
1
5
1
1
6
1
2
7
1
1
8
1
2
9
10
1
1
1
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sylvestris #
2
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94
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PINUS SYLVESTRIS
20
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1
1
1
1
1
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2
1
1
2
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Cenangium abietis
1
Cytospora kunzei
2
1
1
2
Cytospora pinastri
Cytospora kunzei
1
1
1
Cytospora pinastri
2
1
1
2
Cytospora leucostoma
Cytospora pinastri
1
1
1
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2
1
1
2
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Cytospora leucostoma
1
1
1
Cytospora leucosperma
1
1
1
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1
Peniophora pini
1
2
2
1
1
1
1
3
1
2
4
1
1
5
1
2
6
1
1
7
1
2
8
1
1
9
1
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10
1
11
1
1
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!
2
sylvestris #
95
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4
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Ries, 1978; Tarr, 1972).
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!
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_`@ , 1978; 7
8 , 1964). +
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5-10%
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.
# $!
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,
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.
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97
! ,
100
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,
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Cytospora, Phomopsis,
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,
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Cytospora kunzei,
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. . B
% $
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–
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,
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Cytospora. D
$
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!
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17
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%
100-130
.
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160-170.
?
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%
,
,
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"
,
$
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,
%
,
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,
50
70
.
+
,
,
$ .
>
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,
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)
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213
,
107,
60 ( ` > %, 1989). q
50
,
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– 21
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+
200
(Grzywacz, 1990). (
$
99
(
$
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#
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( Acer negundo
) !# 69
( !6 !
!
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+ Polyporus squamosus,
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6
+ #
100
E
#
f.<.|`! $ % (1988)
98
. > 20(
26
(M.Lopez, R.Cuellar, 1987). w. . B !, <. .o % (1988)
$ 34
.
. 2
,
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!
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100 %
.
1
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$! 6 $ !
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!
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%
178
. 4
– 55 %, Ascomycotina – 20 %
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,
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,
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,
$
,
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"
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,
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Q. petreae
!
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,
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sargentii)
$
Cytospora pinastri.
2
,
,
" %
,
"
"
50
70 %
.
?
,
. #
%
$
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, " –
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.
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.
,
Hysterographium fraxini
Cytospora pruinosa.
. >
$
2
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,
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%
Phellinus, Ganoderma, Inonotus.
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,
25 %
101
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10
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2
5
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,
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A +! $ !
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Sphaeropsidales,
Camarosporium, Phomopsis, Microdiplodia. 2
Laburnum anagyroides,
†"
.
,
Camarosporium laburni. #
. t
(
),
"
.
t
"
(
),
%
,
,
%
.
(
$
"
Camarosporium crataegi,
.
Crataegus pojarkovae. E
$
"
(80 %)
.>
"
Microdiplodia
,
$
.>
104
microsporella. +
" ,
%
.
,
"
-
Phomopsis. A
.
-
,
. >
Eriobortya japonica,
.?
,
Phomopsis hughesi,
$
"
,
. D
.
%
9
,
$
,
–
$
,
!
. >
,
(1-5
),
Cytospora leucosperma
9
!
%
.)
10-15
"
,
! .
,
!
15-20
.
{
!
. t
,
,
!
$
%
! .
4
$
58 %,
!
71 %.
,
,
,
. t
!
Diplodia, Camarosporium,
3-4
,
–
Sphaeropsis,
“
$ ”
"
,
Cytospora, Phomopsis,
Hendersonia,
$
6-8
.(
,
,
A
,
+3 #
(Defago,
1935).
!
,
"
$
$%
#+2 1800-2000 #. +
,
–
,
"
% $
!
,
. E
"
. #
!
,
"
2-3
.
2!. #
"
"
.D
105
. # %
$
! .D
$ !
,
"
$
,
. D
!
–
,
!
!
-
.
#
,
! . (
Ascomycotina,
84 %
.
,
$
!
Taphrinaceae, Ophiostomataceae, Dothideaceae,
Coronophoraceae, Sphaeriaceae, Cerastomataceae, Diatrypaceae, Xylariaceae
. A
Cucurbitariaceae, Polystigmataceae, Valsaceae
! ,
! .+
,
Valsa .
!
! . /
.
Valsa ambiens. (
,
,
!
!
$
%
,
"
.2
Valsa .
$ ,
,
6-9,
9
!
$
! . 9
!
!
$
,
"
"
!
.
%
!
!
%
(42 %),
(29 %). +
!
2-3
% ,
"
!
. +
!
,
%
(61%)
(42 %). >
!
!
20-24
,
7000-9000 #
"
(#+2). A
!
"
, "
2-3
,
.
9
!
. >
,
$
,
$ $
.2
$#
.
!
! . t
Laburnum anagyroides
†"
.
Camarosporium laburni,
$
! ;
Ligustrum vulgare,
,
Cytospora
pruinosa
Valsa cypri. )
"
106
$
,
$
.
4
!
,
%
,
,
$%
, -
,
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,
,
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,
$
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$
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2!. #
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"
.
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107
,
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"
Sphaeropsidaceae Melanconiaceae.
2
. {
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,
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,
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!
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800
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700
600
500
400
300
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100
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22
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,
Phillyrea latifolia
!
. >
3-4 –
. {
C
,
,
"
$
.
.
178
6
>
$%
, 95 %,
500
L!
5000
100
5
1
. 12-12. !
"
2
"
12.4
#
% "
"
"!
!
"
,
,
,
,
,
"
.A
$
,
!
!
%
%
.
100
% %
!
"
"
!
4
+! Ganoderma applanatum.
!
#
3
,
,
$
$
..
"
. #
"
$
%
9
$%
-
(
10-20,
!
. 12-13).
,
"
30-40
.
. +
"
179
.
"
5
25
20 - 40
10 - 20
,%
9
!
. 12-13. !
5
10 - 20
20 - 40
,
!
+ # # ! !K$
# $! ! !#
$
t
25
"
,
!
!
. (
(9
$%
,
$
$
). ~
sylvestris
"
pyramidalis. E
%
,
,
,
Malus
Populus
$
.
,
4
!
Cytospora leucostoma. D
"
30
!
!
(6
50 %.
"
"
.
q
,
,
! ,
%
$
pyramidalis.
(
. 12-14). #
!
!
!
%
$%
"
Populus
. E
!
. B
180
!
"
.+
%
%
. )
"
,
.
6
5
3
4
2
1
. 12-14. !
!
+! Cytospora leucostoma #
$ #
!$" Malus sylvestris # !#
!
K
(
'
#
2
:1- $
#
!
( Populus pyramidalis, 2 #
!
(, 3 !$
L 6
$ # 6 !
), 4 !K
) "'!
$ #
!$!, 5 $
!K
,6!+
9
,
./
!
,
$
$
.8
,
!
,
. 9
(
. 12-15). (
$
.>
$
!
,
. t
,
50
"
.
"
25 %. r
"
,
181
"
.
$
,
.
+
2%
"
35
10
10-20
10-20
20-40
. 12-15. !
!
9
Ceratocystis ulmi,
+ ##
,
%
!
182
.
$%
,
,
.
Scolytus scolytus. 9
,
)
13.
4 "
13.1 $
D
(Candolle,
!
9
%
“+
1832): “
!
(]`X! % , 1940;
Y$
1973
B!.). a.b. a `
$%
. “E
,
!
7
:
"
!
(”
"
”.
B
, !! !, 1959; ƒ`! %8 %, 1963; | !
% (1952)
"
-
"
"
! @ % (1962), “
$%
.
%,
,
"
. E
"
”. +
$ 7. .
,
$%
$%
!
" $%
$%
!
,
,
”.
D!
!
$%
.
“D
!
( –
! "L
!
+
! # 5
#
,
! #
!
(, #
!45
$ # (
2
!
!
+ !
(
6 " # )
$
# $(5
K
4
$"
#
!
)
) 6 +
.”
#
!
$%
!
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;
)
;
)
!
"
;
)
;
)
"
;
)
"
"
.
D
,
"
"
.
B
!
,
,
,
!
. # %
$%
!
,
$
$.
"
,
!
.2
,
,
$
!
,
!
$
"
. +
.
183
700
,
,
$
.
"
!
$
,
.
13.2 B
"
"
"
?
.2
Moniliaceae,
Basidiomycotina
Pucciniaceae.
D
+
) #
#.
1
!
) K . q
Monilia cinerea
.
. •
$
Rosaceae.
9
$
"
. >
“ " ”
. D
" $
,
Armeniaca vulgaris,
Cerasus tomentosa, Cydonia oblonga, Laurocerasus lusitanica, Photinia
serrulata, Persica vulgaris, Chaenomeles japonica
.
1"'
!(
!. q
Pseudoidium
" $
Lagerstroemia indica,
,
"
$ .9
.
!(
. q
Botrytis cinerea
$
,
,
.D
Rosaceae,
.
$ (100 %)
"
Eriobotrya japonica,
%
.
E
%
,
Laurocerasus
officinalis, L. lusitanica, Citisus elongatus, Lonicera fragrantissima,
Viburnum tinus.
K!#' !. q
Puccinia jasmini
Jasminum fruticans.
"
,
,
. ?
%
(†"
.
).
>
$!6 .
23
16
,
"
: Ascomycotina – 5, Basidiomycotina – 4
– 14
. D
$
"
( . 13-1).
. 4
!
Marsonina, Zythia, Phyllosticta, Sphaeropsis, Cytospora, Phoma. >
Marssonina,
$ $%
M. juglandis,
"
,
.
184
. 13-1. D
2
$ # $ ! #!
)
+ Fusicladium olivaceum #
! L " Sorbus domestica #
!
(
.
#!
+
$"
,
"
100%
. 9
Zythia
Punica granatum,
†"
.
,
! Phoma punica,
,
$
. q
Phyllosticta
Crataegus
Rosa
.
q
Sphaeropsis
Crataegus orientalis;
S. demersa
"
,
. >
Sphaeropsidales: Cytospora (Cupressus) Phoma (Juniperus, Pinus). D
$
.
185
%! L!. 4
Fusicladium. >
F. dendriticum, F. pyrinum.
$
$%
Rosaceae,
Crataegus, Sorbus, Pyracantha
. q
Malus, Pyrus,
. > $
$
,
. (
>
,
$%
"
Gloeosporium oleae,
" $
..
Olea europea,
.
,
,
"
% .
K!#' !. E
Gymnosporangium, Phragmidium. .
$%
.
$
Rosaceae,
Crataegus, Mespilus. q
Phragmidium
- Rosa canina,
$%
. #
"
Mespilus, Rosa.
1"'
!(
!.
Phyllactinia,
Sawadaea, Sphaerotheca. +
$
.
. 2 , Sawadaea bicornis
Acer campestre,
.>
"
"
. )
Fraxinus ornus,
"
%
. q
Sphaerotheca pannosa
Persica vulgaris Rosa canina.
! !L . E
,
“
”,
Taphrina pruni.
.
Prunus. D
$%
P.
divaricata.
!
,
90 %
" . > P. pissardii P. spinosa
" ,
!
.
% $ #!(
.
Monilia fructigena Botrytis
cinerea. D
$
Rosaceae,
%
Malus, Persica, Amygdalus.
Botrytis cinerea
,
$
.
13.3 B
>
!
"
.
337
. D
.q
– 76 %,
186
Ascomycotina –
15 %
"
$
$
!
$%
.
D" !( (
Cercospora, Marssonina, Entomosporium.
"
$%
,
.
,
Basidiomycotina
,
.
,
– 9 %. D
.
$
Phyllosticta,
Phyllosticta
.q
,
.
$
$
.
D !(
(
Septoria,
Ascochyta, Phleospora.
,
$
" ,
.
D %
"
.
10%. >
Berberis vulgaris, Clematis vitalba, Populus pyramidalis
. q
Septoria
Ascochyta %
$
,
( .
13-2).
! !( (
Polystigmina,
$
Prunus. .
Prunus spinosa
. (
,
"
"
.
•
!( (
Rhytisma, Ectostroma, Marssonina. A
"
. q
Melasmia acerina
.
%
Acer campestre
Acer steveni,
Acer platanoides Acer pseudoplatanus. >
11
. q
Ectostroma
liriodendri
$ (
"
)
$
Liriodendron tulipifera (>
). > r
"
.
,
,
. r
$
Marssonina rosae,
$
Rosa. (
. >
%
Rosa
.
'! "4
(
Clasterosporium
carpophillum.
.
: Amygdalus, Padus, Persica.
1"'
!(
!. B
Erysiphales,
$%
29
.?
20 %
, 37 %
187
. 13-2. D "4 (
#
#!
+
35
# $ +
!
"
$ # Septoria, Phyllosticta, Ascochyta
%
,
. >
Microsphaera, Phyllactinia,
.
$
Sphaerotheca,
$
188
?
-
,
>
! ,
Oidium, Pseudoidium
,
! .(
.
(
$
-
, .
!
, ~
,
11
,
K!#' !. .
,
" %
"
10 %
, 8 %
, (
,
.
.
$ 28
Basidiomycotina. 9"
,
13 %
, 20%
,
. >
$
Coleosporium,
Gymnosporangium, Melampsora, Pileolaria, Tranzschelia, Zaghouania. >
Cumminsiella,
Gymnosporangium, Phragmidium, Puccinia, Uromyces. >
Graphiola, Phragmidium, Puccinia,
Uromyces. 9
"
.
+
"
$%
.2 ,
Melampsora
Salicaceae; Cronartium, Coleospora –
Pinaceae;
Gymnosporangium, Tranzschelia, Phragmidium –
Rosaceae; Zaghouania –
Oleaceae; Uromyces –
Fabaceae; Pileolaria Anacardiaceae;
Cumminsiella –
Berberidaceae. A
Puccinia
,
"
8
.
" '!#
.B
$
Taphrina,
10
.
9
,
,
$%
.
9
.
.2
– T. acerina, T.
carpini, T. cerasi –
$
,
!
Armeniaca vulgaris
#
.
(
. 13-3). > Acer campestre
Carpinus
$
" ,
,
!
. #
"
T. pruni,
. D
$
.
T.
deformans Persica vulgaris.
%! L!.
Fusicladium.
.
7
10
.q
.#
$
Pyrus
189
. 13-3. D
“# $
2
)
+ Taphrina cerasi #
” ! !+
#
190
#!
+
"
communis
Sorbus,
P. elaeagnifolia,
#($!
" $
$%
"
$
100 %
. ( 5
.
,
Sorbus graeca.
. •
.
Moniliales
Melanconiales,
Monilia, Botrytis, Volutella, Cladosporium, Alternaria.
,
>
. >
"
Gloeosporium platani,
Platanus. D
.
70-80 %,
P. acerifolia
A
$
"
.
(
!
Alternaria. .
: A. alternata, A. tenuissima, A. longissima, A. state. B
$
,
"
$%
. +
,
Platycladus,
Betula, Viburnum, Diospyros, Malus, Pyrus, Rosa. .
!
43
,
.
, ~
, #
. >
$
% (75%),
(10%)
(15 %).
{4
. 2
,
.
4
- Lophodermium
pinastri Herpotrichia nigra.
$
Pinus pallasiana P.
sylvestris
.(
.
2
,
!
11
( !+ . 13-1).
13.4 B
% ! "
>
913
,
$
624
!
"
512
Ascomycotina – 100, Basidiomycotina – 12. B
( !+ . 13-2).
191
.
,
7 "8 9 13-1. w
+;
% &
X& " 8 = > 8
X ! B
'E '*
3 &; )
V % B! %
&@
9
+
Marssonina, Entomosporium, Phyllosticta, Cercospora,Coniothyrium,
Gloeosporium
Septoria, Ascochyta
Polystigma
Rhytisma, Ectostroma, Marssonina
Clasterosporium
Microsphaera, Erysiphe, Phyllactinia, Podosphaera, Sphaerotheca,
Uncinula, Sawadaea, Uncinuliella, Arthocladiella
Coleosporium, Cumminsiella, Melampsora, Puccinia, Gymnosporangium,
Phragmidium, Tranzschelia, Zaghouania
Taphrina
Fusicladium
Gloeosporium, Monilia, Alternaria, Botrytis
Lophodermium, Herpotrichia
?
9"
.
+
A
:$
7 "8 9 13-2. w
+
>
2
)
9"
A
'E
% &
X& " 8 = > X " A % B! %
!
>
1 ?> >
9?> 'F ' ?> ,%
80-95
3 &)
Camarosporium, Cytospora, Diplodia, Hendersonia,
Melanconium, Microdiplodia, Pestalotia, Phoma,
Phomopsis, Seiridium, Coniothyrium, Dothichiza,
Cucurbitaria, Valsa, Eutryblidiella, Leucostoma,
Cenangium, Clithris, Hysterographium, Otthia,
Pseudovalsa, Melanconis, Hercospora, Prosthecium,
Tubercularia
Ophiostoma, Verticillium
Gloeosporium, Fusicladium, Coryneum
Gymnosporangium, Puccinia, Uromyces
Monilia, Botrytis, Alternaria
.B
&@
1-3
3-5
1-3
1
,
.
,
Sphaeropsidaceae,
Melanconiaceae,
"
$,
$%
!
.
!6
. B
Ophiostoma, Verticillium,
192
,
Moniliaceae.
(
"
.
,
$
. B
!
Ophiostoma
. q
Ulmaceae. +
9
,
, #:)
!
!
Moniliaceae
!
ulmi,
$
,/
,(
.
Melanconiaceae. q
IV
,
A
,
4
,
.
" $ ,
,
Platanus,
Salix, Quercus.
K!#' !.
"
Gymnosporangium, Puccinia, Uromyces. •
Crataegus, Juniperus, Genista. q
$
(
! $)
"
. >
$
$
%
(Juniperus excelsa).
#($!
. 4
.
Moniliaceae. +
"
,
.
P
(#!
2 6
+ #. >
.
25
,
$%
( !+ . 13-3).
4
Melanconiales Sphaeropsidales. q
–
Ascomycotina
–
Basidiomycotina. +
–
.D
!
Pestalotia neglecta
Euonymus japonica
4
% $ .^.^ %, _.a.7 % (1990). q
Cytospora
.
!
30
.
,
Cytospora
Ephedra procera Paeonia cluinensis ( %!
?% 8 , 1982). >
>
Laurocerasus
officinalis, Osmanthus fragrans, Punica granatum, Quercus agrifolia,
Lonicera fragrantissima, Cotoneaster serotina. +
$
,
,
!
$
. >
Cytospora leucosperma,
Cytospora
schulzeri.
193
7 "8 9 13-3. ! "& X " A %, >B &
!
> % ]!& `
- &)
ytospora leucosperma
Cytospora schulzeri
Diplodia sp.
Diplodia sp.
Phoma berberidis
Phoma juniperi
Phoma pinicola
Phomopsis sambucina
Phomopsis sp.
Macrophoma candollei
Macrophoma mirbelii
Sphaeropsis visci
Coryneum corni-albae
Coryneum microstictus
Pestalotia diospyri
Pestalotia elaeagni
Pestalotia eugeniae
Pestalotia funerea
Pestalotia nerii
Seiridium viburni
Stigmatea nicholsoni
Pleospora herbarum
Pleospora sp.
Leptosphaeria rusci
Microstroma juglandis
Microstroma album
.
8
V#@ B! %
3 ?>' ' - E=
Laurocerasus officinalis
Lonicera fragrantissima
Osmanthus fragrans
Punica granatum
Quercus agrifolia
Cotoneaster serotina
Daphne laureola
Cotoneaster serotina
Berberis oblonga
Juniperus foetidissima
Pinus pallasiana
Sambucus nigra
Mahonia bealii
Buxus sempervirens
Buxus balearica
Buxus sempervirens
Viscum album
Cornus mas
&@
S( > +
C2
C2
C2
C2
C2
C1
C2
C2
C2
C2-n
B0
C2
C2
C2
C1
C2
D1
C2
C2
C2
C2
C2
C3
C2
C1
C1
C2
B0
C3
C2
C2
B2
Viburnum tinus
Diospyros kaki
Elaeagnus argentea
Myrtus communis
Cephalotaxus harringtonica
Cryptomeria japonica
Nerium oleander
Viburnum tinus
Laurocerasus officinalis
Juniperus oxycedrus
Viburnum tinus
Ruscus ponticus
Juglans regia
Quercus robur
Cytospora
Pestalotia,
$%
"
. 9
.( $
visci, Leptosphaeria rusci, Macrophoma mirbelii,
.
2
,
,
"
"
!
,
.
194
$
"
. 2
"
, %
Sphaeropsis
$
12.5
B
"
>
"
$
,
.
#
#.
!
Sphaeropsidales, Melanconiales, Diaporthales.
A
20
,
,
50
100 %. A
,
. #
. D
,
$
"
$
. .
,
,
Populus,
$
. 4
Cytospora. 9
Ficus carica (.
,
A
)
!
Phomopsis cinerea. D
,
.9
"
!
Cytospora,
Phomopsis,
Sphaeropsis,
Camarosporium,
$
. +
,
%
!
,
.
9
$
,
$%
,
$
. 9
,
"
,
,
40-50
.
K!#' !. 4
"
Gymnosporangium, Cronartium. +
.D
$
,
"
,
.
,
,
, "
$
.+
,
,
.>
"
Juniperus excelsa (.
).
,
,
"
""
7090 %. B
.+
"
Juniperus excelsa
,
"
,
Pyrus,
$
. Juniperus excelsa
"
"
,
. 9"
,
!
$ .t
"
,
"
.4
"
, "
.
!
195
.
,
"
"
ribicola. /
Pinus strobus,
,
"
$
Cronartium
Juniperus excelsa. B
$
,
,
$
,
.
, Cronartium flacidium,
. q
.(
"
"
$
/
"
Pinus sylvestris
$$
,
"
,
"
"
$
.
#
,
F
.
!(
!
.
B
$
,
!
,
$
|
"
.
,
,
$
.
>
"
#
$
$
Pyrofomes, Antrodia, Ganoderma. D
“
”
.
Phellinus,
$
. . "
$
,
. >
Pinus
Phellinus pini,
Juniperus excelsa –
Pyrofomes demidofii,
Quercus – Phellinus robustus, Populus tremula – Phellinus tremulae.
t
,
$%
$
$
$
,
%
,
,
,
"
,
"
!
!
.
!
$
$
$
Inonotus, Laetiporus, Polyporus,
"
! ,
,
Fomes fomentarius, Phellinus igniarius,
.
"
2-5-10
" ,
"
. (
3-5
,
$%
,
10-15
. +
,
10-15
"
. +
$
!
196
%
.
9
$
$
,
"
$
. D
%
2
5-10
' !(
,
$%
,
,
$
.
"
. B
"
"
!
.q
,
$
%
,
, "
.B
Coriolus, Funalia, Lentinus, Flammulina, Daedalea
! .D
%
,
$
.B
$
$
. /
"
,
,
%
+
,
F#
#
"
.
Loranthaceae. >
! !
,
..
$%
!
.
$
$
$
%
.
.D
Viscum album,
. 2 ,
100
,
A
. D
.
"
,
Stereum,
,
"
,
"
. 9
V. album
$
!
" $%
. ?
!
>
. .
1-2
"
"
% $
. +
,
$
,
,
$
%
!
.
$
Arceuthobium oxycedri,
Cupressaceae.
"
.
20
Cupressus, Platycladus, Juniperus. r %
.+
.
t%
, Loranthus
t
,
#
"
!+
13-4
.
197
!
"
,
5-10
europea,
Quercus.
.
7 "8 9 13-4. w
+
'E
/?
,
>
9 ;' )
X& " 8 = >
!
>
% 8 % B! %
++; )
9, 9;E;9 ba
'E ?>9
9
Sphaeropsidales, Melanconiales,
q
Diaporthales
Gymnosporangium, Cronartium
Phellinus, Ganoderma, Inonotus, Laetiporus,
Polyporus, Daedalea, Antrodia, Coriolellus,
Fomes, Ischnoderma, Lentinus, Oxyporus,
Spongippelis, Pyrofomes, Lenzites, Tyromyces
Stereum, Coriolus, Funalia, Schizophyllum,
Cerrena, Flammulina, Gloeophyllum, Irpex,
Hapalopilus, Hirschioporus, Hhyphodontia,
Lopharia, Odontia, Peniophora, Pholiota,
Pleurotus, Pulcherricium, Vuilleminia,
Auricularia, Byssomerulius, Bjerkandera,
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40
Valsa (1)
39
Cytospora (2)
38
Valsa (2)
37
Cytospora (3)
36
Valsa (3)
35
Sphaeropsis (1)
34
Botryosphaeria (1)
Sphaeropsis (2)
32
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30
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226
29
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9
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28
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27
Diplodia (2)
26
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25
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24
Otthia (3)
23
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22
Cucurbitaria (1)
Camarosporium (2)
21
20
Cucurbitaria (2)
Camarosporium (3)
19
18
Cucurbitaria (3)
Hendersonia (1)
17
16
Ascospora (1)
15
Hendersonia (2)
14
Ascospora (2)
13
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12
Ascospora (3)
11
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10
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9
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8
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7
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6
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5
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227
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20
19
18
17
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15
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230
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37
36
35
34
33
32
31
30
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29
28
27
26
25
24
23
22
21
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!
!
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20
19
18
17
16
15
14
13
12
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!
!
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!
!
!
#
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11
10
9
8
7
6
5
4
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42
41
#
40
39
Basidiomycotina
#
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!
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!
Stereum
!
38
37
36
35
34
33
32
31
30
29
Peniophora
28
!
27
26
Odontia
25
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24
23
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22
236
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21
20
Corticium
19
!
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18
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Schizophyllum
17
Crepidotus
16
Exidia
15
#
Ascomycotina
Xylariales
14
Helotiales
13
12
11
Hysteriales
10
9
Hypocreales
8
7
Sphaeriales
6
5
Pseudosphaeriales
4
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2
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237
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238
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Basidiomycotina
42
Phellinus, Pyrofomes,
Antrodia
Inonotus, Piptoporus
41
#
Basidiomycotina
40
39
Phellinus
Inonotus
#
!
Basidiomycotina
Phellinus
,
,
Lenzites, Antrodia
,
241
38
37
36
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9
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9
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35
,
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Gloeophyllum
Funalia, Cerrena
!
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,
,
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Coriolus, Hirschioporus
,
,
Inonotus
Panellus
,
,
34
33
32
31
,
,
30
29
28
27
,
,
26
25
24
23
,
,
22
21
20
19
,
,
18
17
16
15
,
,
Polyporus
Laetiporus
,
,
!
Lentinus
,
,
Schizophyllum
#
Basidiomycotina
-
!
Oudemansiella
Pholiota
14
13
Pleurotus
12
11
!
Lentinellus
Flammulina
#
10
9
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Auricularia
Tremella
!
#
!
!
#
!
!
Ascomycotina
?
Sphaeropsidales
Melanconiales
Moniliales
242
8
7
6
5
4
3
2
1
. 15-6. " # )
excelsa #
+ Antrodia juniperina #
#
! ! ! !
243
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$
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. ( . 15-10).
#
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$
!
$
244
,
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246
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9
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Basidiomycotina
33
32
31
30
#
Basidiomycotina
Inonotus
#
c
29
28
Basidiomycotina
Fomitopsis, Gloeoporus
27
26
Abortiporus, Phylloporia
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Melanconium, Gloeosporium.
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Camarosporium laburni.
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296
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298
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341
Summary
Dendromycology
A plant is rather complicated biological system with quite definite
structural organisation¸In the process of evolution, formerly indivisible stem
has differentiated into organs and shoots of different orders which are adapted
to environment conditions quite differently. The rigiden conditions, the
stronger is expressed shoot differentiation as main vegetative mass on a plant.
On the area boundaries, the kife form changes as an adaptive sign in a plant.
Moreover, in plant crown portion of less adapted shoots increases and
population becomes more heterogeneous. Fungi play very important role in
plant life. No one plant species or specimen occurs in nature where fungi
would be absent. Variety of fungi is connected with their close relation to
different plant organs. Fungi are symbiotrophic organisms having come to
existence in the course of long co-evolution with plants. Owing to this, mutual
relations of fungi and plants must be built on cellular level. Existence of life
forms of fungi is possible as organelles. Environment influences on shoot
adaptability and disturbs homeostasis of fungus and plant. After stopping of
physiological functions of plant organs or following their serious disturbances,
there appear fungi as indicators of such disturbances.
Development of fungi on plants proceeds according the same laws as
development of every living thing on the Earth. The basic law of life is the law
of competition elimination ensuring variety of fungi in nature and their
refuges. Plants represent a whole complex of such refuges we call ecological
niches. The idea of the multidimensional ecological niche is confirmed by the
example of woody plants. Not only plant organs but also their age, state,
growth conditions ets. are econiches. All selected econiches are confirmed by
appropriate fungi-indicators. In each ecological niche, succession changes take
place for some time. Some fungi are substituted by others which pave the way
for development of the rest of fungi groups. The order of succession changes
in the ecological niche is common for all plants. Depending on environment
conditions, development of several variants of fungi succession rows in
possible. The phenomenon of geographical vicarism or substitution of fungi in
econiches at introduction with closely related species is inherent in all plants.
Both fungi species and their development stages can be vicarious. Knowing
this law makes it possible to forecast discovery of new fungi species, to
establish connection of anamorph with teleomorph, to settle various practical
tasks. Life of fungi on plants is regulated, in the main, by three enumerated
general biological laws, without knowledge of which one can not comprehend
the order in the world of fungi. Formerly, these laws have been never used in
investigations of fungi.
Mycoflora of plants belonging to one species and genus is identical.
Insignificant species composition differences of fungi can depend upon
342
different age, condition of the plant, upon effects of environment factors on
them. In range of a plant family, similar species number to 50%. Rate of the
fungi on a plant or their standard is established in the zone of an area
optimum. In is fixed according to econiches and quantitative parameters of
fungi development. Formation of a fungus sterile form is evidence of the fact
that the plant is on the boundary of natural adaptation of the species. One can
determine limits of the adaptive zones by development stages of the fungi. On
the boundaries anomalous spore-bering or predominance of teleomorph over
anamorph is observable more often. Development of fungus fruit bodies
occurs at positive temperatures only. Development of many species of fungi
livens up in the periods of change of plant phenophases by seasons. Transition
from darkness to light and vice versa is a stress factor stimulating development
of fungi. Spreading of spores of many specialised pathogens and
symbiotrophic fungi species occurs within plant crown, spore scattering does
not exceed two or three heights of plants.
Mycobiota of aboriginal plants is five to ten times riches than in
introduced ones. On native plants there are more specialised fungi species
which occupy narrow ecological niches. Each plant species has only its own
fungi characteristic for given taxon. And in introduced plants, as a rule, fungi
of five or seven genera dominate in entire tree crown. The circle of food plants
of such fungi is within entire plant families. In lees adapted species, a whole
plant is one ecological niche for one fungus species. Such plants are
vulnerable and perish soon. Types of plant life forms, planting types, plant age
ets. can be ecological niches for the fungi. According to species composition
of the fungi and nature of their development one can make up an objective
viability estimation of a plant or a whole plantation. Development of fungi in
man-made plantations depends entirely upon human activities. An introduced
plant species or a breeding hybrid produced artificially needs thorough nursing
during all life. Climatic anomalies in nature practically destroy the introduced
plants in the area discomfort zone or else cause outbreaks of epiphytoties in
zone of introduction and adaptation. In the area optimum zone, they do not
result in bad consequences. Spreading of fungi spores and diseases on plants
occur slower than in mountain areas. In mountains a disease develops more
intensively below focus of infection.
A plant disease is a disturbance of homeostasis between organisms and
a symbiotrophic fungus on cellular level. It rises as a result of disturbing
plant’s physiological functions as influenced by abiotic, anthropogenic factors,
natural processes of organism ageing, as well under influence of the infection
background which change the balance of organism structures in favour of the
fungus. At present, there exist 17 different theories of plants’ immunity to
fungus diseases. To our mind, non-recognition of fungi on plants as
symbiotrophic organisms and non-use of basic laws of development in their
study is the main mistake of many investigators. So immunity became as if an
economical and not biological evaluation of a plant. A new, biological theory
of immunity is proposed which explains the cause of plant resistance from a
343
viewpoint of basic, above-mentioned biological laws. All plants are affected,
to one or another extent, by diseases. There are no fungus-free plants in nature
at all. “Resistant” species are natural or artificial mutants having lost
specialised species of fungi and, respectively, natural protection of certain
organs and entire organism as a whole. The “resistant” plants are short-lived,
they are injured severely by climatic factors, affected by saprotrophicspolyphages. Adaptation process, in result of which the plant “returns” its
specialised fungi, is the cause of resistance loss in plants. This process
proceeds in opposite order with increased activity of the fungus, its
development in an unusual ecological niche and, as a result of this, the
qualities for which entire breeding process was carried out are lost.
Drawing up of mycological or fungal maps is the final stage of
studying mycobiota of any plant species. The complete information will
depend on number of ecological niches singled out on a plant. Good
knowledge of morphology and biology of plant and fungus allow to fix a large
number of ecological niches. However, as practice has shown, to decide
important practical and theoretical problems, selection of about 100
fundamental ecological niches would be sufficient. Creation of mycological
models includes two stages: drawing-up of a map on fungi species revealed
actually and drawing-up of a theoretical map proceeding from the laws of
fungi distribution in crowns of plants. The mycological models are a matrix of
a plant for its whole life period in given geographic area. Knowledge of
development laws of the fungi makes it possible to make up objectively a
biological estimation of a specimen, species and population of a plant by each
revealed fungus species or its development stage. Methods of forecasting new
fungus species on plants by dominant fungi in plant crown was developed. A
new principle of counting up the fungi on plants was proposed.
Study of species composition of fungi starts with collecting them on
plants. Owing to some objective reasons, an investigator can examine only a
small part of crown. However, knowing the laws of fungi distribution in
plants’ crowns one can already draw up the fungus map. It is necessary to
know well that there always are fungi on each plant, so the examination
always should be carried out thoroughly. Necrotrophic fungi are formed on
shoots for a long time, to one year after their dying-off. The shoots whose age
does not exceed 20 years are an age barrier for this group of fungi. Specialised
bracket-fungi do not occur on young plants. On the shoots which died during
dormancy period fruit bodies of fungi nature better than on shoots died off in
summer. Spores of many fungi on plants are covered with oily pellicle and are
caught easily on clean glass plates. When studying sporulation of polypores
causing root and stem rots, the spore traps must be masked from birds and
tourist who always destroy them for incomprehensible reasons. The author
came across these and other problems more than once, so communicates
readily his experience to colleagues.
344
P
.................................................................................................4
Preface ...........................................................................................................6
1.
........................................................................8
1.1
........................................................ 8
1.2
............................ 11
1.3
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1.4
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1.5
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345
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346
17. :.
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$%
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; /
.....................................................................................................338
Summary....................................................................................................342
Dendromycology ........................................................................................342
347