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The macrofungal community and fire in a Mountain Ash forest in southern Australia

Authors:

Abstract

Changes in the occurrence of macrofungi with time following forestry activities and fire were studied at 14 sites in Mountain Ash (Eucalyptus regnans) dominated forests, in the Eastern Central Highlands, Victoria, Australia. Forests of 0-57 years after fire were used to compare macrofungal communities. Pattern analysis through classification and ordination showed that there was a distinct change in the macrofungal community over time since disturbance. Three phases were apparent in the process of recolonisation after fire: (1) immediate post-fire (0-year), (2) an intermediate phase (2- and 4-year-old), and (3) a mature phase (7-year-old and older). The macrofungi evident in the Mountain Ash forest during the first year after fire were the most distinctive. The change in the suite of macrofungi closely reflected the changes in macrofungal substrates in the forests of different ages. Macrofungi found to be specific to certain stages of regeneration after fire will provide a subset of indicator taxa suitable for use in further surveys.
57
The macrofungal community and fire in a Mountain Ash
forest in southern Australia
Sapphire J.M. McMullan-Fisher1,2*, Tom W. May3 and Phil J. Keane1
1School of Botany, La Trobe University, Bundoora, Victoria 3084, Australia
2Current address: University of Tasmania, GPO Box 252-78, Hobart, Tasmania 7001,
Australia
3Royal Botanic Gardens Melbourne, Birdwood Ave, South Yarra, Victoria 3141, Australia
McMullan-Fisher, S.J.M., May, T.W. and Keane, P.J. (2002). The macrofungal
community and fire in a Mountain Ash forest in southern Australia. In: Fungal Succession
(eds. K.D. Hyde and E.B.G. Jones). Fungal Diversity 10: 57-76.
Changes in the occurrence of macrofungi with time following forestry activities and fire
were studied at 14 sites in Mountain Ash (Eucalyptus regnans) dominated forests, in the
Eastern Central Highlands, Victoria, Australia. Forests of 0-57 years after fire were used to
compare macrofungal communities. Pattern analysis through classification and ordination
showed that there was a distinct change in the macrofungal community over time since
disturbance. Three phases were apparent in the process of recolonisation after fire: (1)
immediate post-fire (0-year), (2) an intermediate phase (2- and 4-year-old), and (3) a
mature phase (7-year-old and older). The macrofungi evident in the Mountain Ash forest
during the first year after fire were the most distinctive. The change in the suite of
macrofungi closely reflected the changes in macrofungal substrates in the forests of
different ages. Macrofungi found to be specific to certain stages of regeneration after fire
will provide a subset of indicator taxa suitable for use in further surveys.
Key words: classification, Eucalyptus regnans, forestry, indicator taxa, macrofungi,
ordination, recolonisation, Victoria.
Introduction
Macrofungi, such as agarics, coral fungi, polypores and cup- and
disc-fungi, play important roles in the ecology of Eucalyptus forest
communities, particularly as saprotrophs and through mycorrhizal
partnerships (May and Simpson, 1997). Fire is one of the driving forces
shaping the Australian landscape, particularly eucalypt forests. It is well-
known that some macrofungi are stimulated to produce fruit bodies after
fire (Petersen, 1970; Cribb and Cribb, 1971; Carpenter and Trappe, 1985;
Carpenter et al., 1987; Watling, 1988; Duchesne and Weber, 1993), and
some studies have been carried out in Northern Hemisphere forests to
investigate the effect of fire on the macrofungal community (Watling 1988;
Visser 1995; Visser and Parkinson, 1999). There has however, been little
research on the effect of fire on the macrofungal community in Australian
forests. Gill and Ashton (1971) listed macrofungi as part of a study of
* Corresponding author: Sapphire J.M. McMullan-Fisher; email: smcmulla@postoffice.utas.edu.au
Appendix 12
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58
eucalypt forests on different soils; fire was part of the site history but its
direct effects were not considered. Warcup (1990) demonstrated a
succession of epigeal and hypogeal larger ascomycetes in the first three
years after fire in plantations of Eucalyptus maculata in South Australia.
The effect of fire on the production of fruit bodies of hypogeal fungi, which
provide food for mycophagous Australian mammals, has been investigated
by a number of authors (Taylor, 1991; Claridge, 1992; Johnson, 1997).
The analysis of macrofungal communities in different continents has
mainly been carried out using visual examination of tables of species of
fungi from different sites (Winterhoff, 1992). Multivariate analyses are
often used for plant community data (Whittaker, 1978), but Arnolds (1992)
highlights the lack of application of such techniques to the study of
macrofungal communities. Villeneuve et al. (1991a,b) used pattern analysis
to successfully elucidate fungal communities within Canadian forests. For
Tasmanian wet forests, multivariate analysis of replicated plot-based data
was used by Packham et al. (2002) in their study of the macrofungi of old
growth forests in comparison to silvicultural regrowth. For saprotrophic
fungi on logs of Mountain Beech (Nothofagus solandri var. cliffortioides)
in a New Zealand temperate forest, Allen et al. (2000) used multivariate
analysis to compare the community of fungi on logs differing in decay
stage and other characteristics.
The present study uses multivariate analysis of plot-based samples to
investigate the effects of disturbance by fire on the macrofungi in forest
dominated by Mountain Ash (Eucalyptus regnans) in the Eastern Central
Highlands, Victoria. The study focuses on the relationships between the
diversity of macrofungi, plants and substrates in forests of different age
since fire.
Materials and methods
Study area
The 14 study sites were located in the Eastern Central Highlands,
Victoria, Australia. All sites were within 30 km of each other and were
located in the Loch Valley, Latrobe River and Powelltown areas. Sites were
all on well-drained granitic soils as part of the Kirchubel land system. The
forests were subject to management and logging operations under the
control of the Victorian Department of Natural Resources and
Environment. Sites were located in E. regnans forest of different ages since
disturbance (Table 1).
Appendix 12
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59
Table 1. Site and disturbance details.
Year of fire
Years since fire
Geographic coordinates
Type of disturbance*
1996
0
37°52’45”S, 145°53’23”E
RB
37°53’09”S, 145°57’55”E
RB
37°47’37”S, 146°00’53”E
RB
1994
2
37°52’46”S, 145°52’03”E
RB
37°46’42”S, 145°56’40”E
RB
1992
4
37°47’47”S, 145°58’00”E
RB
37°45’47”S, 145°56’29”E
RB
1989
7
37°53’50”S, 145°57’45”E
RB
1983
13
37°45’31”S, 145°59’09”E
RB
37°52’57”S, 145°48’07”E
WS
37°52’45”S, 145°48’16”E
WS
1939
57
37°52’45”S, 145°53’09”E
W
37°53’25”S, 145°57’41”E
W
37°53’47”S, 145°58’47”E
W
* RB = regeneration burnt after logging; W = burnt by wildfire; WS = burnt by wildfire
and salvage logged.
Macrofungal identification
Macrofungi were those species with readily visible fruit bodies
(generally larger than 1 mm). Where possible macrofungi were identified to
species, either to named species, or sometimes using informal names for
distinct but undescribed species as applied at the National Herbarium of
Victoria (MEL). Most identification was carried out in the field, but in
some cases specimens were examined microscopically to confirm
identification, using standard methods (Singer, 1986). Fungal names follow
the Catalogue and Bibliography of Australian Macrofungi (May and Wood,
1997; May et al., 2002). Macrofungi that were unable to be identified to
species were grouped into “field taxa”, which were macroscopically similar
fungi that would mostly have belonged to a single genus. Some field taxa,
which varied very little in morphology, may represent a single undescribed
species. Other field taxa, however, had variable morphology within a single
genus and so probably represent a group of species. If material suitable for
a herbarium specimen was present, a voucher was collected and lodged at
MEL. Some field taxa observed were not collected due to time and
processing constraints and a lack of suitable specimens.
Macrofungal survey
At each site, ten 10 m2 (1 × 10 m) strip plots were established at
random positions and orientation within a 2 ha area. A priori decisions were
made not to place plots on tracks or log landings (which are often
compacted) or along creek lines (where different vegetation may
Appendix 12
201
60
Table 2. Substrate groups.
Substrate group
Description
Burnt soil
Burnt soil red and black coloured depending on the temperature of the fire,
black burnt soil contained charcoal pieces.
Burnt wood
Wood, including stumps which has been partially burnt or entirely charred.
Bare soil
Soil, which has not obviously been burnt, and is not covered by moss or
litter.
Litter
Wood that is smaller than 1 cm in diam. and dead leaves covering the
ground.
Bark
Eucalyptus regnans bark which has fallen off the trees.
Small wood
Dead trees and branches from 1-5 cm in diam.
Large wood
Dead wood, including stumps and moss-covered wood, which was greater
than 5 cm in diam.
Wire grass
Dead and living Tetrarrhena juncea.
Live regnans
Live Eucalyptus regnans buttresses, surveyed to a height of 2 m.
Moss
The total covering of moss on plants, wood and the ground.
Dung
Herbivore dung.
be present). Strip plots were used to prevent disturbance due to the
surveying (e.g. trampling) and to ensure that fruit bodies were not
overlooked in the dense undergrowth. Every visible surface in the strip
plots to a height of 2 m was surveyed for the presence of fruit bodies.
Vegetation and fallen timber was disturbed as little as possible during each
survey. Presence of each macrofungal taxon on each strip plot was
recorded, along with the range of its substrates. Because of the irregular
distribution of fruit bodies, the macrofungi found outside the plots but
within the 2 ha area (within the same habitat) were also recorded, along
with their substrates.
Surveys of the strip-plots and the 2 ha sites were undertaken in April,
June, July and October 1996 and in January 1997. In addition, short (15
minute) surveys at all sites were carried out monthly from April 1996 to
January 1997 inclusive, by walking around the same 2 ha sites in which the
plots were located (excluding the plots themselves) searching for
macrofungi. The 2- and 4- year-old sites were surveyed from July 1996
onwards.
Plant and substrate surveys
In January 1997, at each site, each of the ten strip plots used for the
macrofungal survey were extended to 300 m2 (3 × 10 m) and the cover of
vascular plants, bryophytes and substrates was recorded.
The percentage cover of each plant species on each strip plot was
estimated using the following percentage cover classes: < 5% rare, < 5%
common, 5-10%, 10-25%, 25-50%, 50-75% and 75-100%. Bryophytes
were not identified but were grouped as “Liverwort species” or “Moss
species”. The daisies Olearia lirata and O. phlogopappa were not
Appendix 12
202
61
differentiated and neither were tree ferns (Cyathea australis and Dicksonia
antarctica). Vascular plant names follow Ross (2000).
The percentage cover of each substrate type (Table 2) was estimated
on each strip plot using the same percentage cover classes as for the plant
survey.
Numerical analysis
Multivariate analyses were carried out using the PATN: Pattern
Analysis Package (Belbin, 1993). Data matrices subject to ordination and
classification were: (1) macrofungi presence / absence for each site pooled
across all visits, (2) plant average cover across strip plots for each site, and
(3) substrate average cover across strip plots for each site. For (2) and (3)
the average cover was calculated from the midpoint of each cover class.
Dissimilarity matrices were constructed using the Bray-Curtis association
measure. Ordinations were produced using SSH (“Semi-Strong Hybrid”
Multidimensional Scaling) with default options except that random starts
were set at 25. Dendrograms were produced using flexible UPGMA
(Unweighted Pair-Group Method using Arithmetic averages) with default
options, because UPGMA gives equal weight to the objects, not the groups.
The Bray-Curtis association measure and hybrid multidimensional scaling
were used for analysis, as these are robust ecological measures (Faith et al.,
1987).
A series of Mantel Tests were carried out between all combinations of
the three dissimilarity matrices produced from the macrofungi data (data
matrix 1 above), plant data (matrix 2) and substrate data (matrix 3). The
Mantel tests were run for 10,000 iterations.
Results
Plant species richness
A total of 67 plants were recorded from the sites. Floristic data from
the 7-, 13- and 57-year-old sites was consistent with the Eucalyptus
regnans dominated “Wet Sclerophyll Forest” community of Mueck (1990),
although site PT39 had some Nothofagus cunninghamii in the understorey,
which is more typical of “Cool Temperate Rainforest”.
Plant community
A classification of the plant data produced two main groups. These
groups are superimposed on the ordination (Fig. 1). One group included all
0-year-old sites and one of the 4-year-old sites (LO92). The other group
Appendix 12
203
62
Fig. 1. Ordination of sites based on the average percentage cover of plants. Age since fire
(years): = 0, = 2, = 4, = 7, = 13, = 57.
contained the other 4-year-old site (BO92) and all of the 2-, 7-, 13- and 57-
year-old sites. The two-dimensional ordination had a relatively high stress
(0.22), which suggests that whilst the plot shows the major groupings it
does not depict the full range of relationships between sites.
The separation of the two groups is mainly due to the greater cover of
the shrubs Coprosma reptans, Correa lawrenciana, Persoonia arborea,
Pimelea axiflora, Prostanthera lasianthos, and especially of Eucalyptus
regnans on the older sites. The recently disturbed sites are separated out by
the greater cover of herbs and grasses such as Acaena novae-zelandiae,
Dryopoa dives, Euchiton sphaericus, Geranium potentilloides, Hydrocotyle
hirta, Mentha laxiflora, Pelargonium inodorum, Senecio velleioides, S.
linearifolius, Taraxacum sp., and liverworts.
Substrates
The spectrum of substrates varied considerably across sites of
different age since fire (Fig. 2). Burnt wood and burnt soil covered more
than 70% of the recently burnt sites (0-year-old sites), and was present on
the 2- and 4-year-old sites; but on the 7-year-old (and older) sites evidence
of burnt wood and burnt soil had disappeared. Small wood and litter were
not found on the recently burnt sites as such substrates were consumed by
fire. Small wood has
-2
-1
0
1
2
-2
-1
0
1
2
SSH Axis 1
SSH Axis 2
Appendix 12
204
Fig. 2. Average cover (%) of substrates on sites of different age since fire (bars are standard error).
0
10
20
30
40
50
60
70
0 2 4 7 13 57
Years since Fire
Average Cover (%)
Burnt soil
Burnt wood
Bare soil
Litter
BARK
Small wood
Large wood
Wire grass
Appendix 12
205
64
Fig. 3. Ordination of sites based on the average substrate abundance cover. Age since fire
(years): = 0, = 2, = 4, = 7, = 13, = 57.
a maximum of about 8% cover in the 13-year-old sites, probably as a result
of death of many pole stage trees in this age class. Litter reached more than
50% cover by the 7-year-old age class and remained at this level on the
older sites. Large wood was present on the recently burnt sites, but wood of
this size was nearly always burnt and so it was included in the “burnt
wood” class. On 7-year-old sites, burnt logs were observed to have become
moss covered while wood that had not been burnt was more often bare. The
2-year-old sites and LO92 were relatively open and consequently the soil
and wood present was comparatively dry, especially the large logs.
Note that, because of the layering of substrates, the total area may add
up to more than 100%; and even where layering did not occur, use of the
midpoint of each cover class means that the total substrate cover may add
up to less than 100%.
Substrate groupings
Classification of the substrate cover data produced a dendrogram with
four main groups. These groups have been superimposed on the ordination
space (Fig. 3). The first cluster contained the 0-year-old sites in a tight
cluster, well separated from the other sites. The second cluster contained
one of the 2-year-old sites (LO94) and the third cluster contained one each
of the 2-year-old (AD94) and 4-year-old (LO92) sites. The final cluster
contained all the 7-, 13-, and 57-year-old sites and the other 4-year-old site
(BO92), which was a less open site and had wetter substrates, more similar
-2
-1
0
1
2
-2
-1
0
1
2
SSH Axis 1
SSH Axis 2
Appendix 12
206
65
to the situation on the older sites. The two-dimensional ordination of the
data has a low stress value (0.09) indicating a good fit of the ordination
against the original data.
Macrofungal taxon richness
A total of 116 taxa (including individual species and field taxa) were
recorded from the sites, both on and off the subplots. Of these taxa, 59
(51%) were previously described species, 45 (39%) represent a single
undescribed species and the remaining 12 (10%) are “field taxa”, which are
usually a group of species in one genus. For the 95 taxa found on more than
one site, a presence absence table has been arranged to indicate the change
in taxa over time (Table 3). There were 21 taxa that were found on only one
site (Table 3).
Of the total number of taxa, about a quarter (and up to 81% for
AD94) were not recorded from the strip plots. All analyses presented below
are based on presence / absence of macrofungal taxa from all visits to the
sites (including taxa recorded on and off the strip plots). For the strip plots
the first sampling in autumn yielded 30 to 68% of the total number of taxa
eventually found, and further samplings yielded further taxa on all sites,
although few additional taxa were recorded from the summer sampling.
Taxon richness was low for the first four years after fire (Table 1).
The 2- and 4-year-old sites had slightly lower numbers of taxa than the 0-
year-old sites. The peak of macrofungal taxon richness was in the 13-year-
old class that, however, had a similar level of taxa to the 7- and 57-year-old
sites. The highest number of taxa on any one site was 59 on LO83, and the
minimum was 11 on AD94. Sites within each age class were reasonably
similar in their taxon richness, except that there were very few taxa on
AD94.
Macrofungal community
Preliminary multivariate analyses of data based on the abundance of
taxa present on strip plots showed similar patterns to analyses of presence /
absence data for the whole site (including taxa found off the strip plots).
Therefore we present analyses based on presence / absence data for the
whole site. Three main groups were produced in the classification of the
macrofungal data; these are superimposed on the two-dimensional
ordination plot (Fig. 4). The stress of this ordination was 0.14, which
indicates a good fit with the original data. The most dissimilar group is
made up of the 0-year-old age class, the second group is made up of the 2-
and 4-year-old age classes, and the final group contains the mature sites (7-,
13- and 57-year-old).
Appendix 12
207
66
Table 3. Macrofungi present on sites of different age since fire. Species found on only one
site are listed at the foot of the table (and included in totals).
Macrofungal taxa
Site (Years since fire)
AD96
(0)
LO96
(0)
CC96
(0)
LO94
(2)
AD94
(2)
LO92
(4)
BO92
(4)
CC89
(7)
PT83
(13)
BU83
(13)
LO83
(13)
PT39
(57)
AD39
(57)
CC39
(57)
Discomycete sp. B
+
+
+
Mycelium sp. A
+
+
+
Mycena sp. B
+
+
+
Neolentinus
dactyloides
+
+
+
Peziza echinospora
+
+
+
Laccocephalum
sclerotinum
+
+
+
Laccocephalum
tumulosum
+
+
+
Tubaria sp. A
+
+
+
Fuligo septica
+
+
Tricholomataceae sp. A
+
+
Laccocephalum mylittae
+
+
Coprinus sp. A
+
+
+
+
+
Schizophyllum commune
+
+
+
+
Hygrocybe sp.
+
+
+
Physalacria inflata
+
+
+
Hypocrea sp.
+
+
+
+
+
Discinella terrestris
+
+
+
+
+
+
+
+
+
+
Psathyrella sp.
+
+
+
+
+
+
+
+
+
Gymnopilus/Hypholoma
+
+
+
+
+
+
+
+
+
+
+
+
+
Stereum illudens
+
+
+
+
+
+
+
+
+
+
+
Galerina patagonica
+
+
+
+
Ramaria sp. A
+
+
+
+
Heterotextus
peziziformis
+
+
+
+
+
+
+
+
+
+
Psathyrella echinata
+
+
+
+
+
+
+
+
+
+
Laccaria sp.
+
+
+
+
+
+
+
+
+
Marasmius sp. A
+
+
+
Cortinarius sp. A
+
+
+
+
+
+
+
+
+
+
+
+
+
Mycena sp. C
+
+
+
+
+
+
+
Agaricus sp.
+
+
+
+
Polypore sp. B
+
+
+
+
+
+
Dictyopanus pusillus
+
+
+
+
Clavaria miniata
+
+
Pycnoporus coccineus
+
+
+
+
Panellus stipticus
+
+
+
+
Scutellinia sp. A
+
+
Trametes versicolor
+
+
+
+
Trametes hirsuta
+
+
+
+
+
Appendix 12
208
67
Table 3 continued.
Site (Years since fire)
Macrofungal taxa
AD96
(0)
LO96
(0)
CC96
(0)
LO94
(2)
AD94
(2)
LO92
(4)
BO92
(4)
CC89
(7)
PT83
(13)
BU83
(13)
LO83
(13)
PT39
(57)
AD39
(57)
CC39
(57)
Meruliopsis corium
+
+
+
+
+
+
+
Stereum sp. A
+
+
+
+
+
+
Clavaria amoena
+
+
+
+
+
+
Crepidotus variabilis
+
+
+
+
+
+
+
Stereales sp. A
+
+
+
+
+
+
+
Stereales sp. D
+
+
+
+
+
+
+
+
+
Stropharia semiglobata
+
+
+
+
+
Marasmius sp. B
+
+
+
+
+
Mycena pura
+
+
+
+
Discomycete sp. G
+
+
+
+
+
+
+
Mycena sp. D
+
+
+
+
+
+
+
+
Amauroderma rude
+
+
+
Mycelium sp. C
+
+
+
Lepiota sp. A
+
+
+
Xylaria apiculata
+
+
+
+
Discomycete sp. F
+
+
Stereales sp. C
+
+
Calyptella sp.
+
+
Discomycete sp. A
+
+
+
+
Discomycete sp. C
+
+
+
+
Discomycete sp. E
+
+
+
Marasmius elegans
+
+
+
Trogia sp.
+
+
+
Collybia sp. B
+
+
+
+
+
+
Melanophyllum
haematospermum
+
+
+
+
Amanita sp.
+
+
+
Campanella olivaceonigra
+
+
+
+
+
+
+
Mycena hispida
+
+
+
+
+
+
+
Polypore sp. D
+
+
+
+
+
Russula sp.
+
+
+
+
Marasmius sp. D
+
+
+
+
Clavicorona pyxidata
+
+
+
Mycena austrororida
+
+
+
+
Entoloma lampropus
+
+
+
+
Hydnum repandum
+
+
+
+
Calocera sp.
+
+
+
+
Marasmiellus affixus
+
+
+
+
Mycena austrofilopes
+
+
+
+
+
Mycena interrupta
+
+
+
+
+
Appendix 12
209
68
Table 3 continued.
Site (Years since fire)
Macrofungal taxa
AD96
(0)
LO96
(0)
CC96
(0)
LO94
(2)
AD94
(2)
LO92
(4)
BO92
(4)
CC89
(7)
PT83
(13)
BU83
(13)
LO83
(13)
PT39
(57)
AD39
(57)
CC39
(57)
Marasmius sp. C
+
+
+
+
+
Stereum sp. B
+
+
+
+
+
Zelleromyces sp.
+
+
+
+
+
Crepidotus eucalyptorum
+
+
+
+
Lactarius eucalypti
+
+
+
Steccherinum ochraceum
+
+
+
Chlorociboria
aeruginosa
+
+
+
Melanotus hepatoch rous
+
+
+
Mycena sp. A
+
+
+
Discomycete sp. D
+
+
Gloiocephala sp. A
+
+
Galerina aff. hypnorum
+
+
+
Mycena subgalericulata
+
+
Mycena epipterygia
+
+
+
Clavulina rugosa
+
+
Cortinarius sp. B
+
+
+
Collybia sp. A
+
+
Agaricus xanthodermus
+
+
Cyptotrama aspratum
+
+
Total fungi on plot
13
11
9
13
2
10
10
29
21
35
41
36
26
27
Total fungi off plot
13
8
10
7
9
8
6
15
16
11
17
12
11
20
Total fungi
26
19
19
20
11
18
16
44
37
46
58
48
37
47
Total vascular plants
29
28
32
36
19
27
26
17
29
32
30
26
25
20
Species found only on one site. LO96: Mycena sanguinolenta; CC96: Ascomycete sp. A;
LO94: Polypore sp. A; BO92: Polypore sp. C; CC89: Geastrum triplex, Macrolepiota sp.
and Stereales sp. B; PT83: Mycena viscidocruenta; BU83: Ascocoryne sarcoides and
Mycena sp. E; LO83: Amanita sp. A, Antrodiella zonata, Discomycete sp. H, Entoloma
aromaticum and Tremella fuciformis; PT39: Clavaria sp. A, Collybia eucalyptorum,
Dermocybe austroveneta and Omphalotus nidiformis; AD39: Vibrissea melanochlora;
CC39: Leotia lubrica.
The distinctiveness of the 0-year-old sites is due to the presence of
thirteen taxa which were restricted to this age class. Eight of these were
present on all three sites of this class, including Laccocephalum
sclerotinum, L. tumulosum, Neolentinus dactyloides and Peziza
echinospora. The intermediate group containing the 2- and 4-year-old sites
was characterised by the occurrence of Panellus stipticus, Pycnoporus
coccineus and Trametes versicolor, which were found on three out of the
four sites, and on one other site outside the 2- and 4-year-old sites. Clavaria
miniata was found only on the intermediate group sites (on two of the four
sites). A third group, comprising
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69
Fig. 4. Ordination of sites based on the presence / absence of macrofungi for all surveys.
Age since fire (years): = 0, = 2, = 4, = 7, = 13, = 57.
the mature sites (7-, 13- and 57-year-old) was characterized by the
occurrence of 64 taxa which were restricted to this group, including:
Amauroderma rude (3 of 7 sites), Campanella olivaceonigra (7),
Chlorociboria aeruginosa (2), Clavicorona pyxidata (3), Crepidotus
eucalyptorum (4), Entoloma lampropus (4), Hydnum repandum (4),
Marasmiellus affixus (4), Melanophyllum haematospermum (4), Melanotus
hepatochrous (3), Mycena austrofilopes (5), M. austrororida (4), M.
hispida (7), M. interrupta (5) and Xylaria apiculata (4). Most species found
on more than one of the mature sites, were found on both the 13- and 57-
year-old sites, although nine taxa were found only on the 57-year-old sites,
including Agaricus xanthodermus (2), Chlorovibrissea melanochlora (1),
Cyptotrama aspratum (2), Dermocybe austroveneta (1), Leotia lubrica (1)
and Omphalotus nidiformis (1). No taxa were found on all sites, and few
(11) taxa were found on sites of all three ordination groups, examples are
Dictyopanus pusillus (4), Discinella terrestris (9), Galerina patagonica (4),
Heterotextus peziziformis (10), Psathyrella echinata (10) and Stereum
illudens (11). There were 15 species found on both the intermediate and
mature sites, including Clavaria amoena (6), Crepidotus variabilis (7),
-2
-1
0
1
2
-2
-1
0
1
2
SSH Axis 1
SSH Axis 2
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70
Table 4. Results of Mantel Tests based on comparisons among Bray-Curtis association
matrices for macrofungi, plant and substrate data.
Comparison
R
SD
R2
Significance
Macrofungi-Substrate
0.81
0.1
0.6561
<0.0001
Macrofungi-Plant
0.56
0.1
0.3136
<0.0001
Plant-Substrate
0.50
0.1
0.2500
<0.0001
Meruliopsis corium (7), Mycena pura (4), Trametes hirsuta (4) and T.
versicolor (4).
Species found only on the 0-year-old site were mainly found on burnt
soil or burnt wood (the major substrates present) and occasionally on the
few large logs that remained unburnt. Neolentinus dactyloides and the three
species of Laccocephalum all arose from sclerotia or pseudosclerotia,
sometimes attached to buried wood. In the intermediate sites, Pycnoporus
coccineus occurred on larger wood (burnt or unburnt) on three of the four
sites, but not on BO92, where the wood was wetter, but it also appeared on
a log on AD96. Chlorovibrissea melanochlora and Cyptotrama aspratum
were restricted to 57-year-old sites and grew only on large moss-covered
logs. The latter species was also occasionally observed on moss-covered
branches. Species found on the mature sites did fall into different groups
based on substrate preference: some species were found only or
predominantly on one type of substrate, and others were found on a wider
range of substrates. However, apart from the two species mentioned above,
there was no strong relationship between the presence of fungi on sites of
different ages and the spectrum of substrates on the mature sites. The only
effect of the latter was to alter the abundance of particular taxa.
Mantel tests
All three of the Mantel tests were significant at the <0.0001 level
(Table 4). The highest correlation was between fungi and substrate with an
R2 value explaining about 66% of the variation. There are also good
correlations between the macrofungi and plants (R2 = 31%) and the plants
and substrate (R2 = 25%).
Discussion
This study was based only on the presence of fruit bodies, and it is
likely that some fungi occurred solely in the vegetative state during some
sampling occasions, or did not produce fruit bodies at all during the study.
The pooling of data over several sampling occasions overcame this problem
to some extent, although there is evidence from studies of Northern
Hemisphere ecosystems that regular visits over many years are required to
locate all the species that may be present (Richardson, 1970; Rayner, 1979;
Arnolds, 1992; Watling, 1995). However, we consider that there were a
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71
sufficient number of visits and of species recorded to ensure that the
patterns observed are biologically meaningful.
The broad pattern of change in the macrofungal community after fire
in E. regnans forest is a dramatic decrease in taxon richness immediately
after disturbance, with re-establishment of the pre-fire taxon richness by
seven years after the fire. Neumann (1991) reported a similar pattern of
change for the richness of litter arthropods after fire in E. regnans forest.
The slightly lower fungal taxon richness for the 2- and 4-year-old sites
could well be due to the fewer sampling events for these sites, and
especially the lack of an autumn sampling, which was the time of greatest
taxon richness on the other sites.
In terms of the macrofungal community, three phases were apparent
in the process of recolonisation after fire: (1) immediate post-fire (0-year),
(2) an intermediate phase (2- and 4-year-old), and (3) a mature phase (7-
year-old and older). Immediately after fire the plant community began to
re-establish, but after the first year there was no clear differentiation of the
remaining age classes based on plant floristics. However, even though a
similar range of plants was present in 4-year-old and older sites the
structure of the forest did alter markedly with age. This difference in
structure was expressed through the range of substrates present, with less
leaf litter and small wood present on the 2- and 4-year-old sites, where
plants were mostly immature, and the sites generally more open and drier in
consequence. The different phases of the macrofungal community
correspond better with the range of substrates present than with the plant
community, as shown by both the ordinations and the Mantel tests. For
saprotrophs the loss of substrate appears to be responsible for the low
species diversity in the first years after fire, as was found by Visser and
Parkinson (1999) in North American forests.
The lack of samplings in April and June for the intermediate phase
(2- and 4- year old) sites was unfortunate, but caused by difficulties in
locating sites of this age that matched the vegetation and geology of the
other sites. Even with reduced sampling, the intermediate phase is
considered to be well differentiated from the other phases for two reasons.
Firstly, one of the species characteristic of the intermediate phase
macrofungal community was Pycnoporus coccineus, a large and brightly
coloured, persistent wood rotting bracket fungus. This species was absent
from all mature forest sites and from all but one of the immediate post-fire
sites, suggesting that even though the intermediate sites were likely to be
under-sampled for macrofungi, distinctive species characteristic of this
phase were present. Secondly, when the April and June data were excluded
from tabulations, inspection of a table of species by sites still showed good
separation of the three phases, with at least some species characteristic of
(and mostly restricted to) the relevant phase present at each site; although
the overall number of species was less than for the full data set.
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Laccocephalum and Neolentinus species produced fruit bodies from
underground sclerotia or pseudosclerotia in response to fire, as has been
reported previously (Cribb and Cribb, 1971; Fuhrer, 1985). Australian fungi
that have been observed on other sites to produce fruit bodies prolifically
after fire, especially Laccaria species, were rarely observed on the recently
burnt sites. In North America, Visser (1995) also noted the absence of
Laccaria on post-fire sites in her study of Jack Pine forests, whereas
Laccaria was common on other sites. Warcup (1990) found a post-fire
flush of diverse ascomycete species in spring in South Australian eucalypt
woodland, but such a flush was not observed in this study. The weather
conditions prevalent when different studies are carried out may be
responsible for some of these differences. There was certainly below
average rainfall for the spring and summer during our survey.
The few mycorrhizal taxa found during the study were mostly present
in the more mature regeneration sites, although Peziza echinospora was
present only on the immediate post-fire sites. Observations of forest
surrounding the sites, both during the study and at other times, showed that
there were other mycorrhizal macrofungi present that were not recorded
from our study sites. These mycorrhizal macrofungi either require suitable
weather conditions to produce fruit bodies (which did not occur during the
study), or else their distribution is patchy with respect to the size of the sites
that were sampled. The production of fruit bodies of saprotrophic fungi
appeared to be somewhat more reliable, in relation to varying weather
conditions, and also less patchy at the scale of the survey sites.
A succession of decomposer fungi on particular substrates such as
logs or sawdust has been suggested (Wasterlund and Ingelog, 1981;
Hintikka, 1993; Wardle et al., 1995). On litter and small diam. wood, such
a succession would be faster than on the very large fallen Eucalyptus logs
which are up to 1.5 m in diam. and which decompose over many decades.
Immediately after fire the dead plant matter that was present consisted of
large branches, trunks and stumps, most of which had been burnt, and in the
first nine months there were few signs of these being colonised. These
burnt logs were still present in the older forest and from 7-years post fire
became covered in a thick layer of moss. Various fungi were recorded from
these logs but it is of interest that on the 57-year-old sites Vibrissea
melanochlora was found only on such logs which were waterlogged at
times, and that Cyptotrama aspratum was found only on moss-covered
large fallen logs and branches. Such species might be restricted to the later
stages of saprotrophic succession on large logs, which could be occurring
over very long time periods. Berg et al. (1994) suggested that with the loss
of old growth forests in Sweden there has also been a loss of fungi that
depend on old growth habitats, in particular fungi dependent on large logs.
Management of Australian forests where timber harvesting is being carried
out will need to consider the length of forest rotation and the removal of
Appendix 12
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73
substrates such as large logs, in relation to the potential effects on fungi.
Woodchipping, now a common practice, utilises many logs that were
formerly left behind in the forest because they were not suitable for milling.
This activity might be removing the types of substrate that become
important on older sites, such as the 57-year old sites of our study.
Collection of fallen timber for firewood is another activity that might be
removing potential substrates of saprotrophic fungi.
The taxon richness of macrofungi was greater across all the sites in
comparison with vascular plants by a factor of two. On individual sites,
there were more plants than fungi on the 0-, 2- and 4-year-old sites and
more fungi than plants on the rest of the sites. An excess of macrofungi
over plants has been found in previous studies (Packham et al., 2002) and
the number of fungi would no doubt increase with further sampling.
A plot-based sampling strategy was used within each site because it
was unclear what the scale of spatial patterning was going to be for the
fungi. Because there were many macrofungi observed outside the plots on
all sites, the data from the whole 2 ha site was used for analysis. Thus the
subplots might seem superfluous, but even when scoring presence / absence
of macrofungi for each site as a whole, the narrow plots (1 m wide) were
useful as areas in which to concentrate survey activity, and to ensure that all
microhabitats were carefully inspected. Smaller fruit bodies of agarics such
as Marasmius could easily be missed when randomly traversing the whole
site. There is much scope to further develop survey techniques in terms of
the size and number of plots and the manner in which they are searched.
There are many unanswered questions relating to succession of
macrofungi in different plant communities, and in relation to different
forms of disturbance such as wildfire and logging. The suites of macrofungi
in different forest types are also yet to be compared for Australian
ecosystems. Such research will be important because a better understanding
of the way that the fungal community varies with disturbance and with
different habitats will be vital for conservation and management of fungal
biodiversity. The difficulty of readily identifying macrofungi (and indeed
all fungi) in ecological surveys has been a major impediment to wide scale
surveys. This difficulty is caused partly by the numerous undescribed taxa
that are encountered, especially in Australia where the taxonomy of
macrofungi has been largely neglected. Difficulties also arise from the
requirement for microscopic examination of many macrofungi for accurate
identification. In studies where there are regular visits to a number of sites,
the number of samples can be considerable. The lodging of voucher
specimens means that eventually collections can be named as taxonomic
revisions become available. In the meantime it would be useful to have a
smaller set of readily recognisable taxa for use in further surveys.
Indications from this study of named taxa that are specific to certain stages
Appendix 12
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74
of regeneration after fire provide a useful subset of taxa suitable for further
surveys.
Acknowledgements
We thank D. Ashton for sharing his time and wisdom of the Mountain Ash forests;
B. Fuhrer for his support in the field; N. Klazenga for assistance with graphics software;
and the staff of the Noojee and Powelltown offices of the Department of Natural Resources
and Environment for facilitating the project.
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... Biological properties of soils are altered by changes or loss of microbial species and population dynamics, reduction or loss of invertebrates, and partial elimination of plant roots (Doerr and Cerdà, 2005;Neary et al., 1999). Changes in substrata and habitats depend on fire severity and type of vegetation present (McMullan-Fisher et al., 2002;Motiejūnaitė et al., 2014). ...
... There are studies of fire effects on various organisms: insects (Swengel, 2001), vertebrates (Fisher and Wilkinson, 2005;Fox, 1983;Kotliar et al., 2002;Morissette et al., 2002;Penn et al., 2003;Woinarski and Recher, 1997), plants (Kirkpatrick and Dickinson, 1984;Pereira et al., 2013;Weber and Flannigan, 1997), bryophytes and lichens (Boudreault et al., 2000;Esposito et al., 1999;Holt et al., 2008), fungi (Carpenter et al., 1987;Claridge et al., 2009;Dahlberg, 2002;McMullan-Fisher et al., 2002). Different groups of organisms show varying responses to fire, post-fire recovery and succession, depending on fire severity, climate and amount of precipitation, the traits of the organisms themselves, as well as the combination of all these factors. ...
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Myxomycetes are heterotrophic eukaryote organisms that have three life stages, none of which are known to be resistant to fire. The response of myxobiota to different severity of fire is not well known either. We examined myxomycetes in Pinus mugo plantations following a crown fire and in Pinus sylvestris plantations following a surface fire during the first three years after the wildfire event in forested coastal sand dunes in western Lithuania. Additionally, we investigated myxomycetes in corresponding unburned stands. All studied sites (unburned and burned) bore rather different myxomycete assemblages but the disparities of the species compositions between both burn types were more pronounced showing that fire severity had stronger impact on myxomycete species composition than the pre-fire stand type. Analysis of myxomycete assemblages (including the results from field collections, bark and litter cultures) showed that surface fire sites bore the highest number of post-fire species compared to crown fire and unburned sites. Dynamic annual changes in species composition were observed in all studied sites but only crown fire plots showed a clear chronosequence of post-fire myxomycete assemblages. Fire impact promoted establishment and/or sporulation of myxomycete species that are rare in similar unburned stands, or are usually confined to other types of forests and substrata. In addition, individual myxomycete species tended to switch substratum usage during the course of vegetation succession, with a final return to their usual substrata. This possibly signaled the end of early stage of post-fire succession.
... These pyrophilous fungi are absent or seen rarely in unburned systems (Moser 1949;Petersen 1970;Ahlgren 1974). Studies reporting pyrophilous fungi have largely focused on boreal forests (Dahlberg et al. 2001;de Groot et al. 2013;Sun et al. 2015), western North American coniferous forests (Bruns et al. 2002(Bruns et al. , 2005Westerling et al. 2006;Sun et al. 2015;Reazin et al. 2016), the Mediterranean (Buscardo et al. 2010), and Australia (McMullan-Fisher et al. 2002. Taudière et al. (2017) summarized studies involving both wildfires and prescribed burns and their effects on ectomycorrhizal (ECM) fungi worldwide, but missing from all these studies are data for pyrophilous fungi in eastern North America. ...
... Peziza echinospora has been reported as restricted to post-fire soils (Egger 1986) and was collected from Apr to Jul 2017, 5 to 8 mo after the Chimney Tops 2 fire, on well-burned ground in hardwood forests. This is putatively a cosmopolitan species with collections under this name observed as early successional from fire zones worldwide (Egger 1986;Lisiewska 1992;McMullan-Fisher et al. 2002;Ratkowsky and Gates 2009;Gierczyk et al. 2017). ...
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Following a late fall wildfire in 2016 in the Great Smoky Mountains National Park, pyrophilous fungi in burn zones were documented over a 2-y period with respect to burn severity and phenology. Nuc rDNA internal transcribed spacer (ITS1-5.8S-ITS2 = ITS) barcodes were obtained to confirm morphological evaluations. Forty-one taxa of Ascomycota and Basidiomycota were identified from burn sites and categorized as fruiting only in response to fire or fruiting enhanced by fire. Twenty-two species of Pezizales (Ascomycota) were among the earliest to form ascomata in severe burn zones, only one of which had previously been documented in the Great Smoky Mountains National Park. Nineteen species of Basidiomycota, primarily Agaricales, were also documented. Among these, only five species (Coprinellus angulatus, Gymnopilus decipiens, Lyophyllum anthracophilum, Pholiota carbonicola, and Psathyrella pennata) were considered to be obligate pyrophilous taxa, but fruiting of two additional taxa (Hygrocybe conica and Mycena galericulata) was clearly enhanced by fire. Laccaria trichodermophora was an early colonizer of severe burn sites and persisted through the winter of 2017 and into spring and summer of 2018, often appearing in close association with Pinus pungens seedlings. Fruiting of pyrophilous fungi peaked 4–6 mo post fire then diminished, but some continued to fruit up to 2.5 y after the fire. In all, a total of 27 previously unrecorded taxa were added to the All Taxa Biodiversity Inventory (ATBI) database (~0.9%). Most pyrophilous fungi identified in this study are either cosmopolitan or have a Northern Hemisphere distribution, but cryptic endemic lineages were detected in Anthracobia and Sphaerosporella. One new combination, Hygrocybe spadicea var. spadicea f. odora, is proposed.
... described an increase in saprotrophic fungal diversity immediately after fire, with saprotrophic succession in soil more rapid than in wood. In Australian Mountain Ash forests, distinctive communities of soil fungi appeared in the year after fire disturbance, followed by much longer seral phases dominated by non-pyrophilic species(McMullan-Fisher et al., 2002). ...
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Fire has been predicted to be more severe and frequent in forests of the Australian Monsoon Tropics over the coming decades. The way in which groups of ecologically important soil fungi respond to disturbance caused by fire has not been studied in tropical forest ecosystems. Ectomycorrhizal (EM) fungi are important tree symbionts and saprotrophic fungi drive soil nutrient cycles. We analysed both publicly-available environmental DNA sequence data as well as soil chemistry data to test a hypothesis that fire events (1970 - 2017) in a contiguous tropical forest have altered the composition and diversity of EM and saprotrophic soil fungi. We tested this hypothesis by measuring community-level taxonomic composition, fungal diversity, species richness and evenness. We determined whether changes in fungal communities were associated with fire-altered soil chemical/physical properties, vegetation types, or the direct effect of fire. Soil fungi differed in abundance and community phylogenetic structure between forest sites that had experienced fire, and those sites dominated by unburned forest. Communities of EM fungi were structurally altered by fire at shallow soil horizons, as well as by vegetational changes between burned and unburned sites at deeper soil horizons. In contrast, fires influenced community composition of saprotrophic fungi by changing soil nutrient levels and altering litter composition. Pyrophilic, truffle-like EM fungi that rely on mycophagous mammals for dispersal were abundant at recently burned sites. We conclude that fire impacts EM fungi primarily by changing plant communities, whereas fire impacts saprotrophic fungi by reducing soil nutrient levels and altering litter composition. Graphical abstract Credit: Sofia Houghton (2-column fitting image. Color to be used in print.)
... Seventeen percent of jarrah forest macrofungi fruit on wood . Large logs, which are long-lasting and can progress through and support a range of decay classes, sustain the succession of fungal species that contribute to macrofungal biodiversity (Dix & Webster 1995;McMullan-Fisher et al. 2002). Consequently, larger diameter logs can exhibit greater fungal species richness than smaller diameter logs (Yee et al. 2006). ...
Article
We examined the effects of timber harvesting and fire history on coarse woody debris (CWD) at 48 sites dispersed across the dry sclerophyll jarrah (Eucalyptus marginata) forest of south-western Western Australia. These sites represent a range of fire and harvesting histories. The mean total volume of CWD (119 ± 10 m³ ha⁻¹), which varied greatly across the sites, was greatest on the recent harvest category (140 m³ ha⁻¹) and least on the never harvested category (77 m³ ha⁻¹). The ‘old harvest’ category harvested more than 40 years prior had an intermediate value (89 m³ ha⁻¹). The increased volume of CWD on the recent harvest category consisted largely of small diameter (10–50 cm) and less decayed CWD which had been on the ground for approximately 10–60 years. Although the total volume of large CWD (diameter >50 cm) was similar across the recent, old and never harvested categories, harvesting skewed the distribution of large CWD toward the less decayed classes. Multiple linear regression was used to examine the effects of harvest intensity (amount of basal area removed), harvest history, fire history and site attributes on CWD volume. The strongest single predictors of CWD volume were the total reduction in basal area from recent and historical harvesting, and the number of prescribed fires since 1937; both positively correlated with CWD volume (r = 0.47 for both). The next strongest single predictor of the CWD volume was the number of wildfires (negatively correlated, r = −0.40). Unlike prescribed fires, wildfires reduced the accumulated volume of CWD. Over time, timber harvesting and prescribed burning increased CWD loads and reduced the volume of large diameter highly decayed CWD. Increased CWD loads resulting from past harvesting and prescribed burns are a substantial carbon store that may also benefit CWD dependent species. However, forest managers need to balance the potential benefits of maintaining large volumes of CWD against the risk that fires burning under dry summer conditions will consume a high proportion of CWD resulting in severe heating of soil and vegetation, and substantial emissions of carbon to the atmosphere.
... (see Table 1). McMullan-Fisher et al. (2002) recognized three phases of fungal re-colonization during postfire conditions: (1) immediate phase (0 year), (2) intermediate phase (2-4 years) and (3) mature phase (7 years). Such clear-cut phases seem to be dependent on the type of forest and intensity of fire. ...
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Fortnightly survey in control and fire-impacted regions of scrub jungle of south-west coast of India during south-west monsoon (50 m2 quadrats up to 10 weeks) yielded 34 and 25 species of macrofungi, respectively. The species as well as sporocarp richness were the highest during the fourth week, while the diversity attained the highest during the second week in control region. In fire-impacted region, the species and sporocarp richness and diversity peaked at sixth week. Seven species common to both regions were Chlorophyllum molybdites, Lepiota sp., Leucocoprinus birnbaumii, Marasmius sp. 3, Polyporus sp., Schizophyllum commune and Tetrapyrgos nigripes. The overall sporocarp richness was higher in fire-impacted than in control region. The Jaccard’s similarity between regions was 13.5%, while fortnights of regions ranged from 0% (10th week) to 11.7% (eighth week). Control region showed single-species dominance by Xylaria hypoxylon, while multispecies dominance by Cyathus striatus and Lentinus squarrosulus in fire-impacted region. Except for air temperature, nine abiotic factors significantly differed between control and fire-impacted regions. The Pearson correlation was positive between species richness and phosphorus content in fire-impacted region (r = 0.696), while sporocarp richness was negatively correlated with pH in control region (r = −0.640). Economically viable species were 12 and 10 without overlap in control and fire-impacted regions, respectively. Keywords: Mushrooms, diversity, abiotic factors, substrate, disturbance
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Promputtha I, McKenzie EHC, Tennakoon DS, Lumyong S, Hyde KD 2019-Succession and natural occurrence of saprobic fungi on leaves of Berchemia floribunda (climber) and their association with Magnolia liliifera (host). Abstract Fungal succession on various plants from different regions of the world have been well-studied, however there has been no report comparing the fungi on leaves of a climber with those of the supportive plant. Fungi on leaves of Berchemia floribunda, a climber, were studied to fungal diversity and succession over a period of leaf decomposition. These fungi were compared with those on leaves of Magnolia liliifera, the supportive plant, using data from previous studies at the same site. Leaves of B. floribunda were placed with the upper or lower leaf surface adjacent to the forest floor, hung above the ground either under the host tree or other tree species, or placed on the forest floor under the host tree or under other trees to establish the effects of these treatments. These leaf bait trials did not affect the fungal diversity on the leaves. There was very little overlap between fungi on the climber leaves and those on the support tree. Only four saprobes from B. floribunda were also found on leaves of M. liliifera. We suspect that most of the fungi degrading leaves of B. floribunda were initially endophytes and became active saprobes once leaves started to decay.
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Forests fires can be either wild, prescribed or intentional and they are one of the most important threats to natural forests ecosystems in Mexico. In the years 2009 and 2010 approximately 70 ha of forests were burnt at the Barranca del Cupatitzio National Park in the municipality of Uruapan in the state of Michoacán. A study to follow development of macrofungi communities in burnt plots at this national park including changes in richness and composition of the macrofungal species associated was carried out. Three plots of 300 m² each were studied throughout three years (i.e. one year before and two years after the forests fires took place); 81 taxa of fungi were registered of which 10 are considered as pyrophilous. Results showed little differences in species richness composition in each plot before the fire; however a year before the fire and two years after the fire, great differences in species composition occurred (i.e. high temporal beta diversity). Results concluded that the macrofungal species composition is a good indicator of the intensity and effect of the fi re on forests communities.
Technical Report
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This Guide has been written to provide field naturalists and other citizen-scientists with a basic understanding of fungal survey techniques, outlining the basic steps to conducting a fungal survey under Australian conditions. The protocols were developed in consultation with mycologists and environmental managers, and field naturalist groups throughout Australia also provided input and suggestions. Other survey methods do exist, and the protocols listed here will not suit every project. This Guide aims to provide the minimum requirements for conducting a safe, enjoyable, and scientifically valid fungal survey. The intention is to provide an easy-to-follow step-by-step guide for non-specialists who, through volunteering their own time to investigate their local areas, can provide data that is incredibly valuable for this under-studied Kingdom. With support and encouragement to build slowly on their skills, high-quality data can be generated by non-specialists with nothing more than time, methodical discipline, and an eye for the detail and beauty of nature.
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In December 1997, a wildfire in the Pemberton region of southwest Western Australia burnt about 1600 ha of karri (Eucalyptus diversicolor) regrowth forest. Immediately following the fire, a network of sites was selected in burnt 17–25-year-old regrowth forest and in similar aged regrowth unburnt since establishment. We present a species list of the macrofungi recorded from 1998–2002, at 5 burnt and 5 unburnt sites. A total of 322 species of macrofungi were recognised, of which 144 were identified to species or designated an affinity species. All but four species are represented by voucher collections, and most by in situ photographs, and detailed macro and micro descriptions. Voucher collections were lodged at PERTH, and descriptions are available from the first author (R.R.).
Article
This study focuses on the diversity and ecology of wood-inhabiting macrofungal species assemblages in a regenerating tall, wet, native Eucalyptus obliqua forest in southeast Tasmania, 43 years after natural and anthropogenic distutbances. Two plots subjected to "clearfell, burn and sow" silviculture were compared with two other nearby plots that had experienced wildfire. A total of 90 species was identified from 619 macrofungal records during six fortnightly visits between May and July 2010. The plots with abundant live Pomaderris apetala trees in the understorey (i.e., those at Edwards Rd) had markedly different macrofungal assemblages from those with no or with sparse Pomaderris apetala (i.e., at Hartz Rd). This study provided evidence that a 43-year-old regenerating forest maintains a core of common wood-inhabiting macrofungal species irrespective of type of disturbance. Furthermore, species most frequently observed in older forests in Tasmania can also occur in younger managed forests if biological legacies such as large diameter wood, well-decayed wood, large living trees and a diversity of tree species remain after silvicultural treatment.
Book
Readers will perhaps be surprised to find a volume about fungi within a handbook of vegetation science. Although fungi traditionally feature in textbooks on botany, at least since Whittaker (1969), they have mostly been categorised as an independent kingdom of organisms or, in contrast to the animal and plant kingdom, as probionta together with algae and protozoa. More relevant for ecology than the systematic separation of fungi from plants is the different lifestyle of fungi which, in contrast to most plants, live as parasites, saprophytes or in symbiosis. Theoretical factors aside, there are also practical methodological considerations which favour the distinction between fungal and plant communities, as has been shown for example by Dörfelt (1974). Despite their special position the coenology of fungi has been dealt with in the handbook of vegetation science. It would be wrong to conclude that we underestimate the important differences between fungal and plant communities. The reasons for including the former are that mycocoenology developed from phytocoenology, the similarity of the methods and concepts still employed today and the close correlation between fungi and plants in biocoenoses.
Book
A large part of ecological research depends on use of two ap­ proaches to synthesizing information about natural communities: classification of communities (or samples representing these) into groups, and ordination (or arrangement) of samples in relation to environmental variables. A book published in 1973, 'Ordination and Classification of Communities,' sought to provide, through contributions by an international panel of authors, a coherent treatise on these methods. The book appeared then as Volume 5 of the Handbook of Vegetation Science, for which R. TuxEN is general editor. The desire to make this work more widely available in a less expensive form is one of the reasons for this second edition separating the articles on ordinction and on classification into two volumes. The other reason is the rapid advancement of understanding in the area of indirect ordination-mathematical techniques that seek to use measurements of samples from natural communities to produce arrangements that reveal environmental relationships of these communities. Such is the rate of change in this area that the last chapter on ordination in the first edition is already, 4 or 5 years after it was written, out of date; and new techniques of indirect ordination that could only be mentioned as possibilities in the first edition are becoming prominent in the field. In preparing the second edition the chapter on evaluation of ordinations has been rewritten, a new chapter on recent developments in continuous multivariate techniques has been included, and references to recent work have been added to other chapters.
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There has been considerable interest in characterising the general properties of food-webs and trophic interactions but little is known as to how these develop along successional gradients. From a field study we determined how the decomposer food-web develops during a three-year primary succession of sawdust. Microfloral and microfaunal groups increased over the initial 1-2 years of the study and then reached a state of "dynamic equilibrium". During this period of longer-term stability there was considerable short-term instability, with severe oscillations most likely attributable to predator-prey cycles involving nematodes, bacteria and fungi. Populations of top predatory nematodes closely mirrored those of the bacterial-feeding cephalobid nematodes three months earlier, indicative of a strong trophic association. Other relatively close linkages were also detected, but the majority of links involving nematodes appeared weak, suggesting that the distribution of food web interaction strengths is strongly skewed. Food chain length and food-web complexity increased over the first three months of the study but remained invariant thereafter, demonstrating that connectedness approaches to food-webs are extremely insensitive to primary succession. While some of our findings were in broad agreement with Odum's theory of ecosystem succession (e.g. initial biomass buildup, development of connectedness food-web properties), others were not. These included: colonisation early in the succession by the largest organisms in the food-web (mites and Collembola); little difference in early colonising ability between nematodes with r-selected and K-selected traits; a short-term "instability" of microflora and microfauna later in the succession indicative of high nutrient exchange rates; and a detectable rise in the microbial respiration:biomass ratio over the final year probably due to nutrient limitation. Consideration of trophic dynamics aided the understanding of successional trends, and an appreciation of the complex patterns of (trophic) species interactions appears necessary for a more complete interpretation of the patterns and mechanisms of ecosystem succession.