Hugo Kołłątaj University of Agriculture in Kraków, Cracow, Poland
HERPOTRICHIA NEEDLE BROWNING ON ABIES ALBA
IN THE LESKO FOREST DISTRICT STANDS:
DISEASE INTENSITY AND ASSOCIATED FUNGI1
T. Kowalski and K. Andruch
Abstract
Results of a survey on symptoms of the Herpotrichia needle browning and
studies on other fungi on dying needles in regenerated silver fir in the Lesko Forest
District are presented. In total, disease intensity on 450 naturally and 450 artificially regenerated trees of Abies alba, in 30 stands, was evaluated. Identification of
fungi on 600 dead attached needles was based on its morphology in vivo, including
the presence of whitish to plain brown hyphae covering the needles, sporulation
and fruiting bodies. Possible role of Rhizoctonia sp. in development of silver fir disease manifested by needle browning is pointed.
Key words: Abies alba, Herpotrichia parasitica, Nematostoma parasiticum, Rhizoctonia sp.
Introduction
Herpotrichia needle browning of silver fir is characterized by browning and dying of both the young and old needles, which do not fall for a long time. Instead
they hang loosely from the twigs, to which they remain attached by a superficial
mycelium (Butin 1995, Cech 1995, Freyer 1976 a, Hartmann et al. 2007, Mańka
2005, Talgø and Stensvand 2008).
The disease was observed for the first time in 1883, in the South of Germany. At
that time the novel fungal species, Trichosphaeria parasitica Hartig, was considered
to be a cause of the disease (Hartig 1884). Two new names, Acantostigma parasiticum (Hartig) Sacc. and Herpotrichia parasitica (Hartig) Rostr. were proposed for the
pathogen a few years later (Rostrup 1890, Saccardo 1891). Both names have been
1
This work was supported by the research project from the Polish Ministry of Science and Higher
Education (N N309 296634) for 2008–2011.
Phytopathologia 55: 21–34
© The Polish Phytopathological Society, Poznań 2010
ISSN 2081-1756
22
T. Kowalski and K. Andruch
commonly used in the phytopathological literature (Butin 1995, Freyer 1976 a,
1976 b, Munk 1957, Peace 1962, Sivanesan 1971, Talgø and Stensvand 2008).
Later detailed studies of the pseudothecium and subiculum morphology led to the
proposal of a new name, Nematostoma parasiticum (Hartig) M.E. Barr (Barr 1997).
Pyrenochaeta parasitica Freyer et van der Aa had been identified and described as an
anamorph stage (Freyer and van der Aa 1975).
Nematostoma parasiticum occurs in Europe, e.g. in Austria, Germany, Switzerland,
Denmark, Norway and Great Britain (Cech 1995, Freyer 1976 a, 1976 b, Hartig
1884, Holdenrieder 1993/94, Peace 1962, Roll-Hansen and Roll-Hansen 1995,
Talgø and Stensvand 2008) and sporadically in Japan and the North America (Barr
1997).
The fungus is a typical parasite of firs, seen mainly on Abies alba, but also on
other species (A. bornmuelleriana, A. nordmanniana, A. procera and A. veitchii). It can
also attack Picea species when infection potential is high (Butin 1995). It also occurs in Christmas tree plantations (Talgø and Stensvand 2008).
Symptoms characteristic of Herpotrichia needle browning have been observed
on A. alba in stands in southern Poland for many years (Kowalski, unpublished).
This paper presents results of a survey of symptoms of the Herpotrichia needle
browning in regenerated silver fir in the Lesko Forest District and studies on their
causal agent, Nematostoma parasiticum. Other fungi on dying needles of silver fir are
also included.
Materials and methods
Intensity of the Herpotrichia needle browning of silver fir
Analyses of the intensity of Herpotrichia needle browning were made in 30
stands with regenerated silver fir in the Lesko Forest District (N 49°28'28.0'',
E 22°19'39.6'') in August 2008. Stands were randomly chosen on the basis of the
Forest District inventory data. In 15 stands the fir was regenerated naturally, and
the species composition of the main stand included silver fir only or silver fir in admixture with other tree species (Table 1). In the other 15 stands the fir was regenerated artificially on forest or postagricultural land, and the species composition of
the main stand included pine (eight stands), silver fir (eight), larch (seven), beech
(three), spruce (two) and oak (one) (Table 1). Stands were growing in fresh highland broadleaved forest or in fresh mountain broadleaved forest. In most stands
(24) the fir crown density was discontinuous, in others moderate (four) or open
(two; Table 1). Thirty trees were analysed in each stand, including every second
tree in the biogroup located in the centre of the stand. In total, 900 trees were analysed.
The following characteristics were evaluated and recorded:
total height of each tree (±10 cm);
Herpotrichia needle browning on Abies alba...
23
Table 1
Characteristics of stands surveyed in the Lesko Forest District and share
of silver fir trees with symptoms of Herpotrichia needle browning
Compartment, Number of trees
Average height
Species
Density*
subcompartment
analysed
(m)
composition**
Number
(and percentage)
of trees infected
Natural regeneration
9d
30
b
2.57
Jd, Św, So, Db
20 (66.7)
109i
30
b
1.77
Jd
0 (0.0)
13f
30
b
3.00
Jd
2 (6.7)
14a
30
b
3.10
Jd
4 (13.3)
14m
30
b
2.07
Jd
4 (13.3)
88a
30
b
3.60
Bk, Gb, Jw, Jd
9 (30.0)
110f
30
a
2.27
Jd
4 (13.3)
110g
30
b
2.40
Jd
0 (0.0)
110h1
30
b
2.67
Jd
0 (0.0)
110h2
30
b
3.30
Jd
3 (10.0)
111b
30
b
3.10
Jd
21 (70.0)
118d
30
c
2.97
Jd
12 (40.0)
121c
30
b
2.00
Jd
0 (0.0)
128d
30
b
2.80
Jd, Bk, Db
10 (33.3)
30
b
2.73
Jd
LP2
Total/average
X = 2.85
450
2 (6.7)
91 (20.2)
Artificial regeneration
11a
30
b
2.63
Jd, Św, Bk
23 (76.7)
88f
30
b
3.87
Md, So
28 (93.3)
112a (P)
30
b
2.10
So, Jd
0 (0.0)
117h (P)
30
b
2.33
Md, So
25 (83.3)
117k (P)
30
a
2.93
Md
25 (83.3)
117m (P)
30
a
4.13
Md
30 (100.0)
117n
30
b
2.13
So
30 (100.0)
118a
30
a
2.37
So, Jd
30 (100.0)
127b
30
c
2.40
Jd, Św
30 (100.0)
127g
30
b
2.30
Jd
20 (66.7)
127h
30
b
2.17
Db, So, Md
19 (63.3)
175a (P)
30
b
2.67
Md, Bk, Jd, So
25 (83.3)
175b (P)
30
b
3.60
So, Jd, Bk
25 (83.3)
LP1
30
b
3.97
Md, So
23 (76.7)
LP3
30
b
2.37
Jd
Total/average
450
X = 2.66
1 (3.3)
334 (74.2)
LP – private forest, (P) – postagricultural land.
*a – moderately dense, b – discontinuous, c – open.
**Bk – Fagus sylvatica, Db – Quercus robur, Gb – Carpinus betulus, Jd – Abies alba, Jw – Acer pseudoplatanus,
Md – Larix decidua, So – Pinus sylvestris, Św – Picea abies.
24
T. Kowalski and K. Andruch
proportion of dead needles and of dying twig apices on branches with symptoms of Herpotrichia needle browning, according to the scale: a – 25% and less, b –
26–50%, c – 51–75%, d – more than 75% of dead needles or dying apices;
age of needles with symptoms of Herpotrichia needle browning; each tree
was included in one of three groups (a, b, c) depending on circumstances on the
most infected branch: a – tree with first- and second-year needles infected, b – tree
with first- to fourth-year needles infected, c – tree with the first- to seventh-year
needles infected;
part of crown with needles infected (bottom, centre, top).
Needles and branches with symptoms of damage or dieback resulting from infection by other fungi were not included in the analyses.
Fungi on needles of silver fir with symptoms
of Herpotrichia needle browning
Needles of silver fir were collected for mycological analysis on two occasions, in
November 2008 and April 2009, from two stands of artificial regeneration (comp.
88f, 175b) with symptoms of Herpotrichia needle browning. Needles were sampled from two branches with disease symptoms, one branch located at the bottom
and one in the centre of the crown, from each of 10 randomly chosen trees in each
stand (20 trees in two stands). Five needles with symptoms of browning were sampled from each of the five age groups (first- to fifth-year needles), taking every
third dead needle. Each sample of 200 needles from each of the five age groups was
placed in a separate envelope. In the laboratory each 200-needle sample was mixed
and 60 needles were randomly chosen and stored at 6oC. Three hundred needles
were analysed each year. Identification of fungi was based on their morphology in
vivo, including the presence of whitish to plain brown hyphae covering the needles,
sporulation (conidia and conidiophores) and fruiting bodies, particularly if they
were produced often and in aggregations. First analyses were made using a
stereomicroscope. Slides made from the sporulating structures, if present, were
examined using a light microscope. Identification of fungi was based on the available literature (Ellis and Ellis 1985, Gourbière and Morelet 1979, Grove 1935,
Munk 1957, Sutton 1975). The part of the needle (base, centre or apex) on which
sporulation occurred was recorded.
Five branches of silver fir with symptoms of disease were collected in November 2008 for isolation of fungi common in branches and needles, for comparison
purposes. Dead (36) and living (36) needles, with hyphae growing at the needle
base, were surface-disinfected either by rinsing in distilled, sterile water (3 × 30 s)
or by the method of Kowalski and Stańczykiewicz (2000), and dried in layers of
blotting paper. A piece 2 mm long was cut from the base of each needle. The pieces
were placed in Petri dishes, on 2% malt extract agar (MEA, Difco), each at a 45o angle, with its base sunk into the medium. The growing colonies were transferred
onto 2% MEA slopes and into separate Petri dishes containing 2% MEA. After incubation for three weeks at 20oC, mycelia from Petri dishes were compared with
mycelia growing in vivo, on branches and needles, for identification purposes.
Herpotrichia needle browning on Abies alba...
25
Results
Intensity of the Herpotrichia needle browning of silver fir
Symptoms of Herpotrichia needle browning were recorded on 425 silver fir
trees (47.2%) out of 900 trees assessed (Table 2). The intensity of disease differed
among stands. No infected trees were recorded in five stands. In seven stands less
than 20% of trees were infected, whereas nine stands had more than 80% of trees
infected. There were four times as many diseased trees in the artificially regenerated stands as in the naturally regenerated stands (Table 1). The average height of
fir trees regenerated naturally was 2.85 m and of those regenerated artificially was
2.66 m (Table 1). Trees less than 1 m tall were the least infected (Table 2). The infected trees had dying, brown needles that did not fall for a long time, but hung
loosely from twigs, attached by a superficial mycelium (Phots. 1, 2). The mycelium
was particularly abundant at the point of attachment of a needle to the twig. The
greatest number of trees had less than 25% of the crown needles infected. Only five
trees had 51–75% of needles infected (Fig. 1). Needles were infected most often in
the crown centre and most rarely in its top (Fig. 2). There were differences in
amounts of infection in different age-group ranges of needles. Infection of the
youngest, the first- to fourth-year needles, occurred in the greatest number of trees
(43.9%). Infection of the first- to seventh-year needles occurred on branches of
15.0% of trees (Fig. 3).
Table 2
Frequency of infection of Abies alba trees in relation to their height
Height range (m)
<1
Number of trees analysed
Number
(and percentage)
of trees infected
64
7 (10.9)
1.1–2.0
402
170 (42.3)
2.1–3.0
222
130 (58.6)
3.1–4.0
144
76 (52.8)
4.1–5.0
51
33 (64.7)
5.1–6.0
Total
17
9 (52.9)
900
425 (47.2)
The proportion of dying apices on branches with symptoms of the Herpotrichia
needle browning was small. Dying apices were recorded in 87 trees (20.5%). Most
often (12.7% of trees) there were only single cases recorded. No trees with more
than 25% of dying apices were observed.
26
T. Kowalski and K. Andruch
Phot. 1. Young silver fir with symptoms of Herpotrichia needle
browning (photo by K. Andruch)
Phot. 2. Branch of silver fir with symptoms of an early stage
of infection (photo by T. Kowalski)
Herpotrichia needle browning on Abies alba...
Fig. 1. Percentage of needles with disease symptoms
Fig. 2. Intensity of the needle infection in different parts of the crown
Fig. 3. Frequency of infection of needles of different age-groups
27
28
T. Kowalski and K. Andruch
Fungi on dead needles of silver fir identified on the basis
of fruiting bodies and mycelium
Ten species of fungi were identified on 600 dead needles of silver fir (Table 3).
Rhizoctonia sp. was the most common (on 92.0% of needles). This fungus was identified on the basis of its morphology, including the vegetative mycelium growing
on the surface of diseased needles in vivo and from diseased and healthy needles on
MEA in vitro. Thysanophora penicillioides, Rhizosphaera oudemansii and R. macrospora
were also frequent occurring, respectively, on 52.2%, 21.0% and 7.2% of needles
(Table 3, Phots. 3, 4). Fruitbody of Nematostoma parasiticum were rarely produced
on needles (3.2%). The anamorph of the fungus was recorded more often, pseudothecia developed only sporadically and were immature. A number of crowded bristles borne on the apical halves of pseudothecia helped to distinguish the fungus
(Phots. 5, 6). Sporulation of most fungi was recorded with similar frequency in autumn and spring. Only fruitbodies of N. parasiticum were eight times more frequent
on needles in spring than in autumn (Table 3). The more common fungi were recorded from all needle age-groups, and differences in density and diversity of
mycobiota between needles of various age-groups were small. Production of fruiting bodies by some fungi occurred more rarely on first-year needles than on older
ones (Table 3).
Different parts of needles were colonized with different frequency by some
fungi (Table 3). Mycelium of Rhizoctonia sp. was more than three times more fre-
Phot. 3. Conidiophores of Thysanophora penicillioides on silver fir needles (bar = 0.5 mm)
(photo by T. Kowalski)
3.0
Toxosporium camptospermum
Number of needles analysed 300. 0
51.7
89.3
Rhizoctonia sp.
17.3
5.0
Phomopsis occulta
Thysanophora penicillioides
0.7
Nematostoma parasiticum
Rhizosphaera oudemansii
0.3
Lirula nervisequia
7.3
0.3
Rhizosphaera macrospora
0.3
Epicoccum nigrum
November
2008
Alternaria alternata
Fungi
300.0
2.7
52.7
24.7
7.0
94.7
3.0
5.7
1.0
0.7
0.3
April
2009
600.0
2.8
52.2
21.0
7.2
92.0
4.0
3.2
0.7
0.5
0.3
total
Needles colonized
by fungi (%)
600.0
0.0
6.5
1.1
3.2
17.7
1.5
0.7
0.0
0.0
0.0
first-year
needles
I
600.0
1.0
9.8
1.7
5.3
19.7
0.3
0.2
0.3
0.3
0.2
second-year
needles
II
600.0
1.0
10.5
1.3
4.5
19.0
0.5
1.3
0.0
0.2
0.2
third-year
needles
III
600.0
0.5
12.7
1.8
3.7
17.9
0.8
0.3
0.3
0.0
0.0
fourth-year
needles
IV
Needles in age-group (%)
600.0
0.3
12.7
1.1
4.3
17.9
0.8
0.7
0.0
0.0
0.0
fifth-year
needles
V
600.0
0.8
6.0
0.7
1.7
91.5
2.2
2.5
0.7
0.0
0.0
base
600.0
2.5
32.7
3.8
10.0
65.7
3.0
2.0
0.7
0.5
0.2
centre
Needles (%)
600.0
2.3
41.2
6.3
12.5
24.5
0.8
0.8
0.7
0.3
0.2
apex
Table 3
Fungi identified on the basis of sporulation and vegetative mycelium on dead needles of silver fir, in vivo,
in relation to sampling time, needle age and position on needle with fungus
29
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T. Kowalski and K. Andruch
Phot. 4. Pycnidia of Rhizosphaera oudemansii on silver fir needles (bar = 0.5 mm)
(photo by T. Kowalski)
Phot. 5. Pycnidia of Nematostoma parasiticum on silver fir needles (bar = 200 m)
(photo by T. Kowalski)
Herpotrichia needle browning on Abies alba...
31
Phot. 6. Pseudothecium of Nematostoma parasiticum on silver fir needles (bar = 50 m)
(photo by T. Kowalski)
quent in needle bases than in needle apices (Table 3). Thysanophora penicillioides, R.
macrospora and R. oudemansii were most common in needle apices, and their frequency decreased towards the base (Table 3).
Discussion
The symptoms of disease observed in regenerated silver fir in the Lesko Forest
District are consistent with those described as typical for Herpotrichia needle
browning of silver fir (Butin 1995, Cech 1995, Freyer 1976 a, Holdenrieder
1993/94, Mańka 2005, Stocka 1997, Talgø and Stensvand 2008).
The brown hanging needles attached to twigs by superficial mycelium are very
characteristic symptoms that distinguish this particular disease from other needle-casts of which quick fall of needles is typical, e.g. needle-cast of fir caused by
Lirula nervisequia or needle-cast of pine caused by Lophodermium seditiosum (Butin
1995, Butin and Kowalski 1989, Mańka 2005). The proportion of infected trees,
which amounted to 47.2%, was clearly high. The intensity of disease in the crown
of particular trees was not great, however, and only 1.2% of trees had more than
50% of needles infected. Such a situation may have resulted from the moderate
density of the stands. Biogroups of trees grew with discontinuous or, sporadically,
32
T. Kowalski and K. Andruch
even moderate (i.e. the less dense) or open crown density. High stand density results in increased air humidity and direct contact of branches, enabling transfer of
mycelium from diseased to healthy branches, which is considered an essential contribution to disease spread (Butin 1995, Cech 1995, Freyer 1976 a, Holdenrieder
1993/94, Mańka 2005, Stocka 1997, Talgø and Stensvand 2008).
The finding that infected fir trees were four times more frequent among those
regenerated artificially than among those regenerated naturally is surprising. The
average tree height and density were similar for both kinds of regeneration. The
only difference was associated with the composition of trees of the main stand
which formed the canopy in the stands of different regeneration types. The contribution of different canopies to the creation of the different microclimatic conditions can not be excluded. This observation, if it is recorded also in other stands,
may favour the use of natural regeneration of silver fir.
Nematostoma parasiticum produces pycnidia in vivo from late autumn and pseudothecia in spring (Freyer 1976 a, 1976 b, Freyer and van der Aa 1975). The sporadic
production or lack of fruitbodies over a longer period, even on severely infected silver firs, was observed by Freyer (1976 a). A similar situation occurred in regenerated silver fir in the Lesko Forest District where fruitbodies were recorded only on
3.2% of needles, and mostly in spring.
The aetiology of the disease requires emphasis of the presence of Rhizoctonia
species mycelium which was recorded on 92.0% of dead needles. The occurrence
of Rhizoctonia in association with silver fir needle disease, supposedly caused by N.
parasiticum, was also observed in Germany (Pehl et al. 2003, Hartmann et al. 2007)
and Switzerland (Holdenrieder, pers. com.). The present results show that Rhizoctonia sp. may contribute greatly to development of silver fir disease manifested by
needle browning and death. Locally its role may even be dominant. More methodically directed experiments are required for better understanding of the Rhizoctonia
participation and contribution to the aetiology of the disease.
Sporulation of different fungi, particularly T. penicillioides, R. oudemansii and R.
macrospora, was observed on the brown needles attached to twigs. Apart from Abies,
T. penicillioides is known to colonize also needles of Picea, Pseudotsuga and Tsuga (van
Maanen and Gourbière 1997). According to van Maanen and Gourbière (1997), T.
penicillioides grows through the stomata and produces fruiting bodies a few months
after needle fall. Sieber-Canavesi and Sieber (1993) included T. penicillioides among
those fungi that colonize only needle litter. The present studies show, however,
that this fungus could commonly colonize needles still attached to twigs, before
their fall to the litter. Rhizosphaera oudemansii and R. macrospora are included among
primary saprotrophs (Gourbière 1980). Both species can occur as endophytes in
the green, symptomless needles of A. alba. Their earlier colonization may helps
them in further colonization of dying needles (Sieber-Canavesi and Sieber 1993,
Kowalski and Ryś 1996). Fungi such as Phomopsis occulta or Toxosporium camptospermum are known colonizers of conifers (Grove 1935, Sutton 1975).
Herpotrichia needle browning on Abies alba...
33
Streszczenie
OSUTKA ZWISOWA JODŁY W DRZEWOSTANACH NADLEŚNICTWA
LESKO: NASILENIE PROCESU CHOROBOWEGO I GRZYBY
ZARODNIKUJĄCE NA OBUMARŁYCH IGŁACH
Analiza nasilenia występowania osutki zwisowej jodły została przeprowadzona
w sierpniu 2008 roku w drzewostanach Nadleśnictwa Lesko. W 15 z nich podrost
powstał drogą naturalną, w pozostałych zaś 15 drzewostanach – w wyniku odnowienia sztucznego na gruncie leśnym lub porolnym. Ogółem analizie poddano 900
drzew (30 drzew w każdym drzewostanie). Symptomy osutki zwisowej jodły
stwierdzono u 47,2% drzew. Nasilenie choroby w poszczególnych drzewostanach
było zróżnicowane. Ogółem prawie cztery razy więcej drzew wykazywało objawy
porażenia w podrostach powstałych sztucznie niż w podrostach powstałych poprzez naturalne odnowienie. U porażonych drzew występowały objawy zamierania
igieł, które nie odłączały się od pędów lub zwisały na strzępkach grzybni. Najwięcej
było drzew, u których porażonych było mniej niż 25% igieł. Najczęściej infekowane były igły w środkowej części korony, najrzadziej zaś w górnej jej części. Objawy
zamierania pędów na gałęziach z osutką zwisową stwierdzono u 20,5% drzew.
Ogółem przeanalizowano 300 igieł martwych pobranych jesienią 2008 roku i
300 takich igieł pobranych wiosną 2009 roku z gałęzi z objawami osutki zwisowej.
Identyfikacji grzybów dokonywano na podstawie wytworzonego in vivo zarodnikowania. Odnotowywano również obecność charakterystycznej białawej do jasnobrunatnej grzybni pokrywającej igły. Na martwych igłach jodły zidentyfikowano 10
gatunków grzybów. Najliczniej występował grzyb Rhizoctonia sp. (92,0% igieł).
Licznie występowały także: Thysanophora penicillioides (52,2%), Rhizosphaera oudemansii (21,0%) i R. macrospora (7,2%). Stosunkowo rzadko wytwarzane były na
igłach owocniki Nematostoma parasiticum (3,2%), przy czym stwierdzano głównie
jego stadium anamorficzne, oraz sporadycznie niedojrzałe pseudotecja. Poszczególne fragmenty igieł były przez niektóre grzyby zasiedlane z różną częstotliwością. Grzybnia Rhizoctonia sp. ponad trzykrotnie częściej występowała na nasadowych niż na szczytowych fragmentach igieł. Grzyby T. penicillioides, R. oudemansii
i R. macrospora najliczniej odnotowywano na szczytowych częściach igieł, a ich
udział malał ku nasadzie.
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Authors’ address:
Prof. Dr. hab. Tadeusz Kowalski, Katarzyna Andruch M.Sc., Department of
Forest Pathology, Hugo Kołłątaj University of Agriculture in Kraków, Al. 29
Listopada 46, 31-425 Kraków, Poland, e-mail: [email protected]
Accepted for publication: 16.02.2010