Macrofungi were sampled on three Douglas-fir dominated chronosequences in the
Coastal Western Hemlock xeric maritime (CWHxm) biogeoclimatic subzone in
southeastern Vancouver Island. Each chronosequence has four stands: a regeneration
stand (G) aged 10-12 years in 1997, an immature stand (1) aged 38-49 years, a mature
stand (M) aged 83-105 years and an old-growth stand (O) aged 251-322 years old. The
macrofungal cornmunity had a high richness - 384 species were observed from 1995 to
1997 - and evenness (J'= 0.85). Species accumulation curves indicate that the observed
richness is only a fraction of the actual richness of the macrofüngal community. The order
of importance of macrofüngal guilds in Douglas-fir dominated CWHxm forests is
mycorrhizal > litter-decay > wood decay > general saprobes > parasitic, fùngicolous and
basidiolichens. The genera with the most species were Mycena > Cortinarius >
Tricholorna > Russula > Inocybe. The dominant species or species groups were
Cortinarius sg. telamonia spp. > Guepiniopsis alpinus > Inocybe spp. > Mycena Ispp. >
Russula spp. > Galerina spp. > Lactarius rubrilacteus > Russula placita >Hemimycena
delectabilis > Marasmius salalis. There were three distinct h i t i n g seasons, spring,
surnmer, and fall, and most species of macrofungi fniited predominantly in one season.
The following components of the macrofungal community varied significantly
with stand age: community genus richness, community species richness, total fiequency,
mycorrhizal guild richness and Erequency, litter decay guild richness, general saprobes
guild richness and frequency, and wood decay guild richness. Results can be
surnmarized: genus richness G < M < O < 1, species richness G < M < O < 1, total
fiequency G < M < O < 1 ,mycorrhizal guild richness G < M < 1< O, mycorrhizal guild
fiequency G < M < 1 < 0, litter-decay guild richness G < M < O < 1, litter decay guild
frequency G < M < 1 < O, general saprobes guild richness G < M < O < 1, general
saprobes guild frequency G < M < 1 < O, wood decay guild richness O < G < M < 1 and
wood decay guild frequency O < G < M < 1. The frequencies of 18 macrofungal genera
showed significant changes with stand age: 17 genera had lower fiequencies after
clearcutting and one genus had higher fiequency after clearcutting. The frequencies of 58
s thesis is dedicated to my father who instilled in me a fascination with
1'11 always fondly remem r our nature waiks.
ryce Kendrick, who suggested this study, for his
I would like to
for this study, the Coastal Forest Chronosequences Project, and for his help with the
experimental design and analysis of this study. I thank
official faculty supervisor, and Dr. Richard Ring and Dr. Will Wintz for being on my
my entire cornmittee for reviewing my
Paul Kroeger, Christine Roberts, Adolf and Oluna Ceska,
vided valuable help
Demis, Brenda Callan, Scoît Re
identimg mushroom specirnens. 1
specimens. The fieldwork for
following volunteers: J
chad Winder, John
Ian Gibson for helping to organize herbari
s study would not
_
been possible without the
aumbrough, Adolf and Oluna Ceska,
e, Carol Harding, Stuart Lee, Sean
arker, Josh Press, C
stine Roberts, Tony
d Winder, Sergei Yazvenko and many others.
members of the South V
SVIMS and the Pacific No
any of the above are
ycological Society (SV
st Key Council for many enjoyable and educational
mushroom forays.
I th
Sciences and Engineering Research Council (NSERC) for
GS-A Fellowship, the Canadi
Forest service for providing an NSE
supplement, and the Universi@ of Victoria for a President" Sc
Teaching Research Fellowship and Teaching Assistantships. Additional
provided by an NSERC gant to Bryce Kendrick and by the Canadian Forest Service
ough Tony Tro
...................................................................................................................ii
tract
ents
les
................................................................................................
v
........................................................................................................
List of Figures
X
.......................................................................................................x i i
racterization of
...............................
1.1 . 1
Roles of macrofungi in the forest ecosystem ............................................................. 3
1.1.2
Dispersal and survival abilities of fun@
1.1.3
Difficulties in studying ecology of macm
1.2.1
Data collection methods ..........................................................................................10
1.2.2
Data analysis methods ........................................................................................... 13
1.3.1
CommuniQ Results...............................................................................................1 7
1.3.2
Guild ResuIts ...................................................................................................... 19
1.3.3
Genera Results .........................................................................................................22
....................................................................
6
......................................................... 9
viii
2
1.4.4
Species..................................................................................................................... 34
1.4.5
Review .....................................................................................................................
35
Effects of clearcu
easiern Vancouver
................................................................................
5
2.1.1
Effects of forestry on macrofungi ............................................................................59
2.1.2
Other rhreats to macrofùngi .....................................................................................65
2.1.3
Effects of clearcutting on other groups of organisms ..............................................65
2.2.1
Data analysis............................................................................................................ 67
2.2.2
Similarity of stand a g a ..........................................................................................-70
2.2.3
Cluster maSysis of species ......................................................................................
-70
2.2.4
Power Anaiysis ........................................................................................................
70
2.2.5
Evaluation of experimental design .......................................................................... 72
2.3
2
in Douglas-fr
Results
.............................................................................................................72
2.3.1
Effects of clearcutting on the rnacrofimgal cornmunity ........................................... 72
2.3.2
Similady of the macrofùngal communities on different stands ..............................73
2.3.3
Species-area curves for each age .............................................................................73
2.3.4
Effects of clearcuning on the richness and hquency of macrofungal gui1ds ..........74
2.3.5
Effects of clearcutting on the cumulative fiequency of genera ................................76
2.3.6
Effects of clearcutting on the hquencies of macrofungal species ..........................77
2.3.7
.
........................................................................... 78
Power analysis ......................... .
.
Discussion
2.4.1
.......................................................................................................
Richness and hquency of the macrofimgal community ......................................... 81
Simiiarity of the macro
al cornmuni9 on different stands .................................85
Effects of cleareutting on maetofungai guilds ......................................................... 8'7
Sensitivity of macrofungal genera to clearcutting....................................................92
Sensitivity of rnacrofungal species to elearcutli'ng...................................................93
Power Analysis ...................................................................................................... 105
.1 Richness of the macrofimgal CO
Douglas-fi dominated forests on southeastern Vancouver Island.
37
.2 Comparison of macrofungal comunity composition in
CWHxm Douglas-fir dominated forests for each month in 1997 using
37
Sorenson's coefficient of similarity.
d a t i v e frequency and evenness macro
Douglas-fir dominated forests
chness and proportion of total richness of macrofungal
guilds of Douglas-fir dominated forests in the month of October 1995,
1996, and 1997.
38
.5 Dominant rnacro
equency 0 and their relat
fir dominated forests.
38
gal genera by species
proportions (Ps,pf) in
s(S),and
m Douglas-
ost frequently encountered species in C
dominated forests in 1997.
40
.7 Presence/absence and fiequency of macro
Douglas-fir dominated forests on southeni Vancouver Island fiom 1995
to 1997.
Comparison of habitat, guild structure
species in each guild), total rie
previous studies.
Plots that had the largest ANOVA residuals.
49
to
110
110
of age efTects on macrofungal genus
2.2 Summary of AN
quency,
and
species
richness
and
ss, species richness
f?equency of macrofungal guilds with Tukey's post-hoc comparisons of
al1 pairs of stand ages.
111
Sirnilarity of species composition of macrofungal
communities in 1997 for all pairs of chronosequence plots using
Sorenson's coefficient
EIlfects of clearcutting on rnacrofungal genus frequency in
CWHxm Douglas-fir dominated stands.
e 2.6 Effects of clearcutting on macrofungal species in CWHxm
Douglas-fir forests in 1997.
e 2.7 Macrofungal species found only on old-gro
Douglas-fir stands frorn 1995 to 1997.
Power of post-hoc cornparisons of age classes for different
ANOVA tests.
ary of results of ANOVAs on genus ri
each rnonth.
xii
ap of coastal forest chronosequence sites.
were sampied on the three southeastern-most chronosequences,
Victoria Watershed North, Victoria Watershed South, and Koksilah.
.2 Layout of transect/quadrats for sampling macrofungi at
each stand in 1995,1996 and 1997.
.3 Species rank vs. log abundance.
Seasonal variation in the richness and to fiequency of
the macrofungai communiQ in 1997 as measured by sporocarp
production.
Seasonal variation in species composition of the
as measured by the
51
52
53
54
Nmber of macro
Number of macrofungd species observed versus t h e .
56
Cladogram of chronosequence plots grouped by
eies coniposition.
119
gal species area curves for dif7Ferentlyaged
xrn Douglas-fir stands.
120
Effects of clear-cutting on ric
a) mycorrhizal guild richness vs. stand
richness vs. stand age, c) wood-decay gui1
d) general saprobes guild richness vs. stand age.
Pgaires continued
.5 Effects of clear-cutting on cumulative species frequency
acrofungal guilds: a) mycorrhizal guild fiequency vs. stand age,
122
b) litter-decay guild richness vs. stand age, c) wood-decay guild
richness vs. stand age. , d) general saprobes guild richness vs. stand
age.
Changes in macrofungal guilds with stand age: a)
proportions of total richness, b) proportions of total frequency
123
ure 2.7 Power versus number of replicates for ANOVA to
detect differences in macrofungal genus richness related to stand age
at three îype-1 error rates.
124
Power versus number of replicates for ANOVA to
detect differences in macrofungal species richness related to stand
age at three type-1 error rates.
125
ower versus number of replicates for Tukey's post-hoc
cornparisons (a = 0.10) ïnvolving regeneration age.
126
Power versus number of replicates for Tukey's
cornparisons (a= 0.10) among
ages.
Effect size versus power of Tukey's post-hoc
arisons (a = 0.1 0).
128
ure
Effect size detectable with
pst-hoc cornparisons (a= 0.10) versus
129
igure 2.13 Box plots of genus ric
indicated by white bands.
130
The temperate old-growth forests of British Columbia represent an increasingiy rare,
invaluable and perhaps irreplaceable resource. They are a record of baseline information
about biological diversity and ecosystem functioning in the absence of the influence of
industrial man. Sadly, this information is not being collected as quickly as the forests
themselves are being cut down. On Vancouver Island. British Columbia, less than half
(45 %) of the mature first growth forests remained in 1991 (BCMOF 1991). These forests
are being cut down and replaced witb even-aged stands which are projected to be
harvested again when they are m
(90- 120 years for
so may never
allowed to reach the old-growth stage again. The longest stage in the life cycle of
forest is thus being eliminated, with potentially devastating effects on biodiversity. We
must strive to understand the impacts of current forestry practices, and rnodi@ those
practices radically, if necessary, before these unique reservoirs of biodiversity are lost.
The macrofungi of Vancouver Island have been little studied.
Western Hemlock (C
) zone is the largest biogeoclimatic zone in
its mycota is probably similar, as is the ora. Fewer than 1% of the maero
Columbia have been treated in studies with systematic sampling tec
e@estimated that the species richness ratio of fungi to vascular plants is
about 6 :1 and possibly as high as 9: 1 (Hawhwo
translates to about 15,000 to 22,500 species of fungi. Arnong these would be an estimated
1,500 to 2,000 species of macrofungi of which only 500 have been reported in the
literature (Ryan et al. 1993).
One ongoing study by Gamiet has published a list of some 133 fungi in C
growth on the south mainland of BC (Gamiet and Berch 1992). There is another study
under way, analyzing the effects of tree monoculture on macrofungal
Vancouver Island (Renata Outerbridge pers. comm). Roberts is engaged on a Ph.D. on the
genus Russula on Vancouver Island (Christine Roberts pers. comm.). An amateur group,
the South Vancouver Island Mycological Society, has compiled an unofficial checklist of
Vancouver Island mushrooms with over 800 species, and is just starting to document
these species with herbarium specimens (Ian Gibson pers. comm.).
Many countries have recognized the need to conserve macrofùngi and their
habitats and have compiled RED lists of rare and endangered species. These countries
include: Germany (Blab et al. 1984; Benkert et al. 1996), Austria (Krisai 1986), Finland
(Rassi and Vaisanen 1987; Rassi et al. 1992), Norway (Bendiksen and Hoiland 1992),
Slovakia (Lizon 1995),and the Netherlands (Arnolds 1989). RED is an acronym, and
organisms on RED lists are rated for three attributes: Rarity, Endangerrnent and
Distribution. Once compiled, these lists can be used to make management decisions about
forest habitats. No RED list exists for ritish Columbian macrofungi (Conservation Data
Centre, BC Ministry of Environment Lands and Parks), partially due to the absence of the
baseline information necessary to compile these lists.
The objective of this chapter is to characterize the macrofungal community in
CWHxm Douglas-fir stands. This information can be used as a starting point for
managed and maintained.
ore
Macrofungi ean be defined as non-lichenized ascomycetes and basidiomycetes
that produce fruiting bodies visible to the naked eye (Redhead and Berch 1995). The
macrofungi are an integral component of the forest ecosystem, yet they are frequently
overlooked because the above-ground fmiting bodies, or sporocarps, are often ephemeral
and are produced somewhat sporadically - in some cases only once in several years. The
main part of the organism, the perennating thallus or mycelium, is composed of a network
of hyphae and is hidden in the substrate (soil, wood, litter, etc.). Macro
major roles in the forest ecosystem as: 1) decomposers, 2) mycorrhizal symbionts, 3)
parasites or pathogens, 4) food for other organisms (Pilz and Molina 1996).
The primary and probably ancestral role of macrofungi is that of the decomposer.
Al1 fungi are heterotrophic, Le., they cannot fix carbon as plants do and so rely on the
existence of other organisms or their remnant biomass. Fungi produce a suite of enzymes
that can digest bio-polymers such as cellulose, lignin, keratin and chitin (Kendrick 1992).
Bacteria can also digest these substrates but the hyphal growth form of fungi gives them
an advantage over bacteria by enabling them to penetrate, permeate and utilize large solid
masses of these substrates, whereas bacteria can act only at surfaces (Cooke and Rayner
1984). Thus, fungi are very important for breaking down and releasing nutrients from the
large volumes of plant remains that accumulate in the forest (O'Dell et al. 1996).
1.l.
1.2 Mycorrhizal symbionts
Mycorrhizal fungi are extremely important to the plants in the forest ecosystem.
Harley and Smith (1983) recognize seven types of mycorrhizae: vesicular-arbuscular,
ectomycorrhizae, ectendomycorrhizae, arbutoid, monotropoid, ericoid and orchid.
Macrofungi may be involved in al1 of these except vesicular arbuscular mycorrhizae
which are formed by microscopic memebers of the Glomales (Karley and Smith 1983).
Ectomycorrhizae are particularly important in the coniferous forests of the Pacific
Northwest because al1 conifers, except members of the Cupressaceae, are
ectomycorrhizal. Ectomycorrhizae are dual structures comprised of short lateral plant
roots with fungal hyphae surrounding them and penetrating between their cortical cells.
Ectomycorrhizal fungi benefit their host plants by providing nihogen, phosphorus and
increased drought tolerance, and in retwn receive carbohydrates (Trappe and Fogel 1977).
These authors even state at most woody plants cannot survive in the field without
mycorrhizae. Ectomycorrhizae can provide pine trees with up to 3.2 times more P and 1.8
times more N than non-mycorrhizal roots (Bowen 1973). With their apically extending,
repeatedly branching hyphal growth form, mycorrhizal fungi act as far-reaching
extensions of the plant root systems that can explore and extract nutrients from a much
larger volume of soi1 than could be explored by root hairs.
Ectomycorrhizae have other beneficial effects as well. Some ectomycorrhizal
fungi have been s h o w to suppress occurrences of root rot in their hosts. Laccaria
laccata, Pisolithus tinctorius and Paxillus involutus inhibit the root rot-causing fungus
Fusarium oxysporum (Sinclair et al. 1982, Chakravarty and Unestam 1987, Duchesne et
al. 1989); and the presence of Thelephora terrestris was negatively correlated with root
rot in Pinus strobus seedlings (Ursic et al. 1997).
Mycorrhizae are important to the physical structure of the soil. Hyphae of
mycorrhizal fungi physically bind soil particles into macroaggregates and produce
polysaccharides that cement the soil into aggregates (Miller and Jastrow 1990).
Different mycorrhizal fungi occupy different niches. They vary in their host
specificity - some being generalists while others are limited to one host species (Molina
1979) - ecophysiological requirements, nutrient uptake and water absorption capabilities
(Mejstrick and Krause 1973, Dixon et al. 1983). Simard et al. (1997) have shown that one
ectomycorrhizal
gus may be involved in symbioses with trees of two different species
at one time and can actually transfer carbon bi-directionally between them. Conversely,
one ectomycorrhizal tree may have many different ectomycorrhizal fungal partners at one
time, and over time (Trappe 1977). Ectomycorrhizal fùngi exhibit succession through the
lifetime of a forest stand on both naturally disturbed sites (Visser 1995) and sites
disturbed by man (Dighton et al. 1986). On an old-growth site, the trees should be
associated with mycorrhizal fungi that are specifically adapted to the environmental and
nutrient conditions and tree hosts of the site. If we continue to clear-cut our old-growth
forests some late-successional or old-growth-dependent mycorrhizal fungi might become
extinct, therefore management practices that conserve fungal diversity may be important
in maintaining long-term forest productivity (Kropp and Albee 1996)
Besides being mutual symbionts of the forest trees, macrofùngi are also important
tree parasites and pathogens.
ecosystem, they actually have some beneficial effects. Trees killed by pathogenic
become snags and logs that are inhabited by wildlife (O'Dell et al, 1996). The gaps
created in the canopy by the dying trees allow shade intolerant plants to grow (Holah ef
al. 1993) and release the next generation of canopy trees.
Food sources
Macrofungi are important as food sources for wildlife such as bear, elk, deer,
small marnmals, and mollusks (Fogel and Trappe 1978, O'Dell et al. 1996). Also, many
arthropods such as Coleoptera (beetles) (Fogel and Peck 1975), Diptera (true flies)
(Okland 1994, Worthen and MacGuire 1990) and Collembola (springtails) (Klironomos
1994) depend on
gi for food. Mycophagy brings the energy and nutrients extracted
fkom dead material by decomposer
gi back into the food web, continuing the various
nutrient cycles.
1.1.2.1 Dispersal
The epigeous macrofùngi are teleomorphs (sexually reproducing stages of the life
cycles) of basidiomycetes and ascomycetes. There are a number of methods by which
they may recolonized heavily disturbed sites, such as clearcuts, and disperse to new
locations and substrates. These are: vegetative growth of mycelium, animal dispersal,
7
water dispersa1 of spores, and air dispersa1 of spores. Vegetative growth is not important
as a method of long range dispersal. Rhizomorphs of Armillaria bulbosa grow in soi1 at
0.2 rnlyear although they may grow as fast as 9.0 mmlday (3.3rnJyear) in culture (Smi
al. 1992). Vegetative growth may be limited by obstacles, zones of unfavourable
conditions and the presence of other fungal mycelium. Some fùngi produce structures that
c m survive unfavourable conditions and serve as inoculum when conditions are
favourable again. Animal dispersa1 is important for hypogeous macrofùngi, but is not
likely to be a major mode of dispersa1 for epigeous macrofungi that can expel their spores
into the air. Water dispersal is probably of secondary importance in the terrestrial
ecosystem, though spores can be carried short distances by rainsplash and longer
distances in streams and surface runoff.
The primary method of long-range dispersa1 for most epigeous macro
airborne dispersa1 of spores. A laminar boundary of still air surrounds al1 surfaces, and
fungal spores must penetrate this boundary and enter the turbulent layer of air in order to
achieve effective dispersal (Gregory 1973). This is achieved by a number of mechanisms
that forcibly discharge the spore or by passive methods that rely on kinetic energy from
wind, raindrops, or other disturbances. Once in the turbulent air, spores may travel great
distances, but over 90 % are deposited within 100 m of their source. Precipitation has a
strong effect on spore dispersal; rainfall decreases the number of spores in the air
exponentially over time, depositing more spores closer to their source (Lacey 1996)
Besides recolonization by one of the above dispersa1 methods, fungi may survive
disturbances or unfavourable conditions in a resting stage. Fungi produce three types of
persistent resting structure: 1) stromata, which are indeterrninate thick plates of hyphae;
2) sclerotia, which are discrete, solid masses of hyphae, often with a hard, pigmented
rind; and 3) pseudosclerotia, which are discrete masses of friable material bound tog
by hyphae (Cooke and Rayner 1984). Functionally, these structures are equivalent: they
lie dormant during unfavourable conditions when the rest of the myceliurn dies off, and
begin growing again when conditions are suitable. During maturation, these structures
accumulate glycogen, trehalose and other food reserves and become dehydrated (Dix and
Webster 1995). Not al1 fùngi produce sclerotia but at least some mycorrhizal macrofungi,
e.g. Paxillus involutus, and some saprobic macrofiuigi, e.g. Collybia tuberosa, produce
sclerotia which may persist in the soil for some time before germinating. However, the
longevity of sclerotia (Brundrett and Abbot 1995) and even which fungi can produce
them has not been well studied. Sclerotia of Verticillium sp., a parasitic hyphomycete,
have been reported as being viable after 14 years in soil (Sussman 1973) and it is
conceivable that some ectomycorrhizal fungi can also produce such persistent sclerotia.
1.1.2.2 Ectomycorrhizal host switc
Many ectomycorrhizal fiuigi are obligate syrnbionts and cannot survive without
their host trees, although a few are able to use ericaceous shrubs as alternate hosts (Perry
el al. 1987), and might be able to survive clearcutting. For example, Astraeus
hygrometricus, Tricholomajlavovirens, Suillus albidipes, Suillus tomentosus, Lactarius
paradoxus, and Coltriciaperennis form ectomycorrhizae with jack pine (Pinus
banksiana) and also form arbutoid mycorrhizae with bearberry (Arctostaphylos uva-ursi,
Ericaceae) (Danielson 1983). Facultative ectomycorrhizal fùngi can survive saprobically
without their host but often with reduced vigour and they may be out-competed by more
vigorous saprobes.
Despite their enormous ecological importance, the community ecology,
biogeography and conservation status of macro
gi are poorly understood (Rydin et al.
1997) partly as a result of the inherent difficulties presented by the nature of macrofùngal
communities. There is high variability £rom year to year in the abundance and species
composition of sporocarps. Redhead and Berch (1995) suggest that macrofungal
cornrnunities cannot be adequately characterized in less than 3 years and that a study
period of five or more years is preferable. Macrofwigi exhibit a species accumulation
curve over time that may not level off for more than 20 years (Tofts and Orton 1998). The
ephemeral nature of sporocarps also causes problems, because while sporocarps of some
gi such as Cantharellusformosus can persist for more than three months (Redhead et
al. 1997), others last only a few days or less and thus sarnpling must be frequent if these
ephemeral species are not to be rnissed (Redhead and Berch 1995). Another problem is
the fact there may be little correlation between above-ground (sporocarps) and belowground (mycelium) assessments of species composition and abundance (Gardes and
Bruns 1996).
This study took place within the framework of the Coastal Forest
Chronosequences study initiated by Trofjmow et al. (1997). Macro
were sampled at three chronosequence sites on southeastern Vancouver Island,
Columbia, Canada: Victoria Watershed South (VWS), Victoria Watershed North (VWN),
and Koksilah (Figure 1.1). Each chronosequence site has four stands: a regeneration stand
aged 10-12 years in 1997, an immature stand aged 38-49 years, a mature stand aged 83105 years and an old-growth stand aged 25 1-322 years old. VWS contains Douglas-fir
dominated stands in the eastern variant (C
Very Dry Maritime biogeoclimatic subzone, and VWN and Koksilah chronosequences
have Douglas-fir dominated stands in the western variant (CWHxm2) of the Coastal
Western Hemlock Very Dry
aritime biogeoclimatic subzone (Tro
Other criteria used to select chronosequence sites were that al1 stands within a
chronosequence were within a 5 km by 5 km or smaller area, had a similar slope, a midslope position, similar elevation, similar aspect, and had second growth stands of harvest
origin (Tro@mow et al. 1997). Some stands did not meet al1 these criteria and the
discrepancies are outlined below. More detailed descriptions of each plot including
environmental characteristics, selected soil chemistries and soil horizon descriptions,
stand descriptions, and indicator vegetation species, have been extracted from Trofjmi~ow
et al. (1997) and are presented in Appendix I
At VWS the regeneration and mature plots were gently sloping while the
immature and old-growth plots were moderately steeply sloping. The VWS regeneration
plot is on an upper slope, and the mature plot is on a lower slope. The VWS mature plot
has a northwest aspect while the other plots have northeast aspects. Also the VWS mature
stand was salvage-logged after wildfire (Trofymow pers. comm.) and the site series
appears to be transitional between the zonal 01 (western hemlock - Douglas-fir Kindbergia) and the moister 07 (western redcedar - Foamflower) site series while the
rest of the stands are zona1 01 site series.
At VWN, the slope varies from gently sloping (immature and mature plots), to
moderately sloping (regeneration plot) to moderately steeply sloping (old-growth plot).
The aspect varies greatly: the old-growth plot faces south-west, the regeneration plot
faces west, the i
ature plot faces north and the mature plot faces northeast. Al1 second-
growth stands are of logging origin and al1 plots are in the zonal CWHxm2IOl (western
hemlock - Douglas-fir - Kindbergia) site series.
On the Koksilah chronosequence the mature plot is moderately sloping and faces
southwest while the others are gently sloping and face south. The mature stand is of
wildfire origin. The regeneration and mature plots are in the zonal CWWxm2101 (western
hemlock - Douglas-fir - Kindbergia) site series and the mature and old-growth plots are
xm2103 (Douglas-fir - western hemlock -salal) site series.
Macrofungi were sampled from 1995 to 1997 and the sampling methods changed
over time. In the fa11 of 1995, macrofungi were sampled on two chonosequences, VWS
and VWN, by members of the Southern Vancouver Island Mycological Society (SVIMS),
an amateur mushroom-hunters' club. Sampling took place in October and November, but
the November sampling was incomplete due to snow. In each plot macrofùngi were
sampled along four transects, 40 m long and 5 m wide arranged in the shape of a square
(Figure 1.2). The abundance of each species was recorded as one of three abundance
classes:
F - few, about 1 to 10 sporocarps per transect
M - many, about 11 to 100 sporoc
A - abundant, more than 100 sporocarps per transect
The substrate of each collection was recorded as one of seven classes:
1 - organic litter over minera1 soil
2 - well-decayed wood
3 - organic litter over well-decayed wood
4 - relatively undecayed log or snag
5 - moss on rock
6 - moss on minera1 soil
7 - exposed mineral soil
In the fa11 of 1996, the author with the aid of SVIMS members, sampled
macrofùngi on three chronosequences, VWS, VWN and Koksilah. On each plot,
macrofùngi were sampled along 2 transects, 44 m long by 4 m wide (Figure 1.2).
Fungi with a cap diameter less than 2 cm were sampled on six 2 m by 4 m quadrats
located at the ends and midpoint of each transect (Fig 1.2). Relative abundance and
substrate data were recorded as in 1995.
Starting in March of 1997, macro ngi were sampled monthly at VWS, VWN,
chronosequences. Fungi with a cap diameter greater than 2 cm were
sampled on 4 m by 4 m quadrats and those with caps smaller than two cm were sampled
on 1.25 m by 1.25 m quadrats nested within the larger quadrats (Figure 1.2). There were
twenty quadrats per stand spaced 4 m apart along four 40 m transects that form the sides
of a square (Figure 1.2). Because there were so few
gi fmiting from March to August
1997, the 4m by 4m quadrats were expanded to 4 m by Sm and al1 macrofungi, including
those with a cap diameter less than 2 cm, were sampled on these quadrats (Figure 1.2).
Substrate data was recorded as in 1995 and 1996, but instead of counting the number of
sporocarps for each taxon, their presencelabsence on the quadrats was used to calculate
their frequency.
In al1 years, specimens that could not be identified to species in the field were
collected and taken to the lab where they were stored at 4 O C until
eY could be studied
more closely. Specimens were identified to genus and, if possible, to species. When
specimens could not be identified to species they were placed into distinct groups within
a genus e.g. Mycena sp. 1, and those that were not placed in distinct groups were simply
lumped under the genus narne, e.g. Mycena spp. For purposes of analyses, these groups of
related species were treated as distinct species. Voucher specimens were dried for future
reference and have been deposited in the herbarium at the Pacific Forestry Centre,
Victoria, BC (DAVFP). No voucher specimens were kept from 1995 collections.
Data fi-om al1 monthly samples were pooled to find the total number of genera and
the total number of species encountered. The percent frequency of an individual species
(f) was calculated as:
where
f is frequency
p is the number of quadrats on which the taxon was present, and
n is the total number of quadrats.
No distinctions were made between the two different levels of sarnpling, the large and
small quadrats. Total frequency was defined as the sum of the frequencies of al1 the
individual species on the quadrats and so could be greater than 100%. The cumulative
frequency of the guilds and genera was calculated as the sum of the frequencies of the
individual species belonging to each category.
1=
lron
where
F is the total fkequency or cumulative frequency
n is the total number of species, and
J; is the fkequency of the ith species.
Fungi were assigned to one of seven guilds, mycorrhizal guild, wood-decay guild,
litter-decay guild, fimgicolous guild, general saprobes guild, parasite guild and
basidiolichen guild, based on their observed substrates and the available literature. The
general saprobes guild contains species that are non-specific saprobes, for example
Xeromphalina fulvipes was observed growing on wood and litter, and multi-species
groups composed of both litter and wood-decay fungi, e.g. Galerina spp. Frequency of
the guilds and genera was measured as the surn of the fiequencies of their component
species and so can be greater than 100%.
1.2.2.1 Evenness methods
Species were ranked according to their abundance, i.e. the total number of
quadrats on which they were present in 1997. The ranks were then plotted against the
loglo of their abundance giving a graphical representation of the distribution of abundance
among species. Two indices of equitability were calculated, the Shannon-Weaver
equitability (J'):
where p, is the proportion of individuals of the ith species and N is the total
number of species; and the erger-Parker dominance index (d):
d = Am,/AT
where A,,
is the abundance of the most common species and ATis the abundance
of al1 species.
1.2.2.2 Similarity of species composition for each month
Similarity of macrofungal species composition between al1 months was calculated using
Sorenson's coefficient of similarity (S,) (Krebs 1983).
where :
S, = Sorenson's coefficient of similarity
j
= the number of
species present in both months
a = the number of species present in the first month
b = the number of species present in the second month
The resulting similarity matrix was converted to a dissimilarity matrix
(dissimilarity = 1- S,) and agglomerative hierarchical cluster analysis with average
linkage was perforrned using the "hclust"
etion in S-PLUS.
1.2.2.3 Species-area curves methods
Data from al1 months in 1997 were pooled and species presencelabsence was
summarized by quadrat. Data from al1 years could not be pooled because of the different
sampling regimes that were used. The 1997 data set was chosen because it was the most
complete, spanning 9 months of the year rather than just one, and has a large number of
smaller sampling areas. Species accumulation curves were developed by rando
sampling the data without replacement, also known as jack-knifing or rarefaction, and
keeping a tally of the number of species present versus area sampled or nurnber of
quadrats sampled. No distinctions were made between the two different levels of
sampling, the large and small quadrats. This random sampling was repeated 10 times
using the S-plus program "s.a.curve" (Appendix III). The resultant cloud of points was
plotted and a regression line fitted.
esul
The results are presented in a hierarchical manner, community-level results are
presented first followed by guild-level results, genus level results, and species level
results. Within this hierarchy, data are summarized by three different timeframes. Data
from 1995 to 1997 were pooled to give the most complete view of the macrofungal
community; data from 1997 were extracted from the complete data to examine
frequencies, seasonal trends and allow comparison with data presented in the following
chapter; and data from October of 1995, 1996, and 1997 were extracted to examine
annual trends.
1.3.1.1 Diversity
From 1995 to 1997, a total of 385 species or groups of macrofùngi were observed,
representing 128 genera (Table 1.l) and seven fungal guilds. Two hundred species from
92 genera were found in 1995, 144 species from 65 genera were found in 1996, and 302
species from 108 genera were found in 1997.
The total frequency of al1 species and species groups in 1997 was 1388 %, Le. the
average 16m2 quadrat had almost 14 different rnacrofungal species fruit on it during the
course of a year.
The slope of the species rank vs. log abundance curve for 1997 (Figure 1.3) is
steep at the start and then levels out, showing that abundance is less evenly distributed
among the most common species than it is among the uncommon and rare
Shannon Weaver equitability index was 0.85, and the Berger-Parker dominance index
was 0.04, both of which indicate that abundance is fairly evenly distributed among
species.
18
1.3.1.2 Annual variation
October was the only month in which complete data was collected in al1 three
years. In October 1995 140 species from 71 genera were observed, 120 species from 59
genera were observed in October 1996,212 species from 79 genera were observed in
October 1997 (Table 1.1). In al1 three Octobers, a total of 296 species from 103 genera
were observed.
The species composition of the macrofungal cornrnunity in October changed
considerably from year to year. Species composition was most similar between October
1996 and October 1997 (S, = 0.5 l), intermediate between October 1995 and October
1997 (S, = 0.46), and least similar between October 1995 and 1996 (S, = 0.42).
1.3.1.3 Phenology
The peak macrofungal fruiting season in richness and frequency is in the fall,
starting in September and peaking in October and November (Figure 1.4). Two smaller
peaks occur in the spring and early summer (May and July). The maximum nurnber of
species that fruited in the fa11 was 21 1 species, triple the summer maximum of 70 species.
The maximum total frequency in the fa11 was 758%, more than 6.5 times the total
frequency in the summer. Very few species fruited in the months of March, April and
August.
Figure 1.5. is a cladogram based on the dissimilarity of the macro
community in each month. Months linked at higher values are less similar in species
composition and months linked at low values are more similar in species composition.
Four clusters are apparent, a spring cluster with the months of March, April and May, a
sumrner cluster with June and July, a fa11 cluster with the months of September, October
and November and, lastly, the month of August. The complete similarity matrix is
presented in Table 1.2
Species accumualtion curves
Figure 1.6 shows the relationship between the number of species observed and the
area sampled in 1997 fitted with a regression line and 95% confidence intervals. The
equation of the regression line is a second order polynomial with logarithmic terms.
The total nurnber of species on the species list increased with time (Figure 1.7).
The average increase was 93 species per year for al1 data and 78 species per year for
October data.
The richness and frequency results compiled by fùngal guilds are listed in Table
1.3. The dominant guild in terms of species ric ess was the myconhizal guild, 41.7% of
al1 species recorded in 1997 and 38.4% of al1 species from 1995 to 1997 were
mycorrhizal. This was the largest change of al1 guild proportions when 1997 data were
compared to 1995 to 1997 data. The next richest guild was the litter-decay guild with
about 30% of al1 species, the wood decay guild with almost 20% of al1 species, and the
general saprobes guild with just under 10% of al1 species. The fùngicolous guild,
basidiolichen guild and parasite guild each had fewer than 1% of al1 species.
20
When the guilds are compared by their total frequency (Table 1.3,1997 data only),
the trends are similar to the trends shown by richness. The mycorrhizal guild is still the
dominant guild but it is even more dominant, accounting for 48.5% of the total fiequency.
The litter-decay guild and wood- decay guild are less dominant by frequency (25.6% and
11.6 of the total frequency, respectively) than by richness. The litter-decay guild
maintains its position as the second most fiequent guild but the wood decay guild slips to
fourth position. The general saprobes guild is more dominant by frequency (12.9% of the
total fiequency) and is the third most fiequent guild. The fimgicolous guild, basidiolichen
guild and parasite guild each accounted for fewer than 1% of the total fiequency
The evenness of the mycorrhizal guild and the litter decay guild (Table 1.3) were
similar to the evenness of the whole macrofungal community, although they had slightly
higher Berger-Parker dominance indices. The evenness of the wood-decay guild and the
general saprobes guild was lower than the evenness of the whole community and the
mycorrhizal and litter-decay guilds (Table 1.3).
1.3.2.2 Annual variation
The mycorrhizal guild, litter-decay guild, general saprobes guild, and wood-decay
guild showed the same pattern when richness data fiom October 1995,1996 and 1997
were compared (Table 1.4). For the mycorrhizal, litter decay, and general saprobes guilds
richness was highest in 1997, intermediate in 1995 and lowest in 1996. The mycorrhizal
guild showed particularly high richness in October of 1997 with 102 species, more than
double the number seen in 1995. The wood decay guild showed the lowest annual
variation in richness with 29 species in both 1997 and 1995, and 24 species in 1996.
The proportion of richness that belonged to each guild varied f?om year to year
(Table 1.4). The mycorrhizal guild showed the largest change, accounting for a minimum
of 33.6% in 1995 and a maximum of 48.3% of al1 species in 1997. The litter-decay guild
decreased by about 3% each year from 1995 to 1997; the general saprobes guild showed
little variation; and the wood-decay guild was at about 21% in 1995 and 1996, and only
1.3.2.3Phenology
The seasonal variation in richness and cumulative frequency of the macrofungal
guilds (Figure 1.8) corresponds to the phenology of the macrofungal community as a
whole with a few exceptions. The mycorrhizal guild did not show a peak fniiting season
in the spring and showed only a minor peak in the early summer; only 3 % (4 species) of
al1 mycorrhizal species observed in 1997 fruited in May and 16 % (20 species) fi-uited in
July. The mycorrhizal guild had a very strong fi-uiting season in the fall, almost 82 % (103
species) of al1 mycorrhizal species observed in 1997 fi-uited in October. The litter-decay
guild had three peak fniiting seasons. The spring and early summer peaks were not as
strong as that of the wood-decay guild or general saprobes guild, 22 % (19 species) of al1
litter-decay species observed in 1997 fruited in May and 24 % (2 1 species)
The fa11 peak was strong with a maximum of 65% (58 species) of litter-decay fungi
fruiting in October. The general saprobes guild showed two peak seasons, the proportion
of species fniiting increased steadily fiom May to July with a peak of 39% (1 1 species) in
July, and the fa11 season peaked in October with 71% (20 species) of al1 general saprobes
fniiting. The wood-decay guild had three peak fniiting seasons but showed the least
difference between the spring/early sumrner and fa11 seasons, 34% ( 1 9 species) of wood
decay fùngi fruited in May, 30% (17 species) fiuited in July and 52% (29 species) fruited
in October.
Ge
esul
The dominant genera by species richness are listed in Table 1.5; complete data are
presented in Appendix II. The dominant genus by richness was Mycena, with 11.3% (34
species) of al1 species observed in 1997 and 12.7% (49 species) of al1 species from 1995-
1997 were members of this genus. The second richest genus was Cortinarius with 7.6%
(23 species) of al1 species in 1997 and 8.3% (32 species) fiom 1995-1997. The next
richest genera, in bo
1997 and 1995 to 1997, in order were Tricholoma, Russula,
Inocybe, Clitocybe, Collybia and Lactarius, decreasing gradually from about 4% of total
richness for Tricholoma down to about 2% for Lactarius. Combined, these eight richest
genera accounted for 36.7% of al1 species in 1997 and 37.3 % of al1 species fiom 1995 e remaining genera, 23 each had between 1% and 2% of the species richness
in 1997, and together accounted for 3 1.4% of the species richness. From 1995 - 1997, 17
genera each had between 1% and 2% of the species richness, and together accounted for
24.3% of the species richness. Sixty-seven genera each accounted for less than 1%, and
combined accounted for 3 1.9% of the species richness in 1997. From 1995 to 1997, 1 O3
genera each accounted for less than 1%, and combined accounted for 39.4% of species
richness.
The most frequent genera are listed in Table 1.5, complete data are presented in
Appendix III. The most frequent macrofungal genus in 1997 was Mycena ( 1 77 %) which
had 12.8% of the total frequency, followed by Cortinarius (158%) with 11.4% of the total
frequency. Ten more genera each accounted for more than 2% of the total frequency in
1997, in order they are Russula (8.5%), Inocybe (5.3%), Galerina (4.5%), Lactarius
(4.0%), and Tricholoma, Marasmius, Guepiniopsis, Nolanea, Hemimycena and Laccaria,
decreasing from 3.3 % to 2.2 %. Together al1 these dominant genera ( > 2% of total
frequency each) accounted for 63.8% of the total frequency. Nine genera each accounted
for 1% to 2% of the total frequency, and combined accounted for 12.5% of the total
frequency. Eighty-seven genera each accounted for less than 1% of the total frequency in
1997 and combined accounted for 23.7% of the total frequency.
The most common macrofungal species and species groups are listed in Table 1.6.
One species group had a frequency greater than 50%, 10 species or species groups had
frequencies fiom greater than 25% to 50%, 22 species or species groups had frequencies
from greater than 10% to 25%, 33 species or species groups had frequencies from greater
than 5% to 1 O%, 61 species or species groups had frequencies fiom greater than 2% to
5%, 4 1 species or species groups had frequencies from greater than 1% to 2%, and 134
species or species groups had frequencies of 1% or lower.
Four of the top five are species groups rather than individual species. The most
comrnon species group in 1997 was Cortinarius sg. telamonia spp. which had a frequency
of 57.9% and accounted for 4.2% of the total frequency. Guepiniopsis alpinus was the
most common individual species with a frequency of 43.8% and 3.2% of the total
frequency. Inocybe spp., Mycena spp., Russula spp., Lactarius rubrilacteus, Russula
placita, Hemimycena delectabilis, Marasmius salalis, and Cortinarius acutus vel afl, al1
had greater than 25% frequency. Cortinarius obtusus vel a f ,Hebeloma crustuliniforme,
Russula brevipes, Mycena pura, Cortinariusfulvescens vel afl, Laccaria laccata,
Pseudohydnum gelatinosum, Nolanea holoconiota vel afl, Galerina emetensis vel afl,
Lyophyllum semitale, Nolanea spp., Russula xerampalina, Mycena alcalina, Mycena
rorida, Dermocybe phoenicea, Ramaria spp., Clavulinopsis fusiformis, Helvella
lacunosa, Tricholoma spp., Omphalina ericetorum, Mycena sp. 1, and Xeromphalina
campanella al1 had greater than 10% frequency. Table 1.7 lists al1 species observed from
1995 to 1997, their guilds, presencelabsence in 1995 and 1996, and fiequency in 1997.
Al1 cornmon species listed for the mycorrhizal, litter-decay and general saprobes
guilds had 10 % or higher fiequency, species listed for the wood decay guild had 5% or
higher fiequency. The most common mycorrhizal fungi were: Cortinarius sg. telamonia
spp., Inocybe spp., Russula spp., Lactarius rubrilacteus, Russulaplacita, Cortinarius
acutus vel a#, Cortinarius obtusus vel afl, Hebeloma crustuliniforme,Russula brevipes,
Cortinariusfulvescens vel a#, Laccaria laccata, Russula xerampalina, Dermocybe
phoenicea, Ramaria sp., Tricholoma spp., and Inocybe ovatocystis. The most common
litter-decay fungi were Hemimycena delectabilis, Marasmius salalis, Mycena pura,
Galerina emetensis vel afl, Lyophyllum semitale, Mycena rorida, Clavulinopsis
fusiformis, Helvella lacunosa, Mycena sp. 1, Mycena oregonensis, and Otidia alutacea
vel afl The most common general saprobes were Mycena spp., Galerina spp., Nolanea
holoconiota vel afl, Nolanea spp., and Mycena alcalina. The most common wood-decay
fungi were Guepiniopsis alpina, Pseudohydnum gelatinosum,Xeromphalina campanella,
Mycena elegantula, Nidula niveotomentosa, Trichaptum abietinum, and Hypholoma
capnoides.
.3.4.2 Phenology
Most macrofungal species fruited in one season only; 17 species fniited only in
the spring, 13 species fruited only in the summer, and 187 species fniited only in the fall.
Other species had high frequencies in one season but fruited in other seasons as well. The
most frequent spring-specific species were Ciboria rufofusca, Discinaperlata,
Pseudoplectania melaena, Agrocybe sp., and Omphalina chlorocyanea. Nolanea
holoconioia vel a$ had a high fiequency in the spring and also fniited less frequently in
the summer. Omphalina ericitorum had a high fiequency in the spring and the fall. The
most frequent summer-specifie species were Nolanea cetrata vel afl, Kuehneromyces
vernalis, and Peziza spp. Plectania melastoma and Dasyscyphus bicolor had high
frequencies in the sumrner and lower frequencies in the spring. Coltriciaperennis and
Inocybe mixtilis were frequent in the summer but also fmited in the fall. The most
comrnon fall-specific species were Cortinarius obtusus vel a$, Cortinariusfulvescens vel
a f ,Lyophyllum semitale, Russula xerampalina, Dermocybe phoenicea, Clavulinopsis
fus formis, Helvella lacunosa, Mycena sp. 1, Mycena oregonensis, and Otidia alutacea
vel a 8
Discussion
.l.
1 Diversity of the macrofungal CO
m has very high macrofungal richness, in fact it shows the highest
richness of any study done with a similar size of sampling area in North America, and
shows similar richness to a European study (Salo 1993) that covered an area 15 times
larger (Table 1.8). The Douglas-fir forests of the CWHxm biogeoclimatic zone on
southern Vancouver Island are a hotspot of macro
gal diversity that has been very little
studied and it should be ensured that sufficient areas of this habitat are protected. Further
research is required to determine if other forest ecosystems on Vancouver Island or
ritish Columbia are equally diverse, and why macrofungal diversity is so high in
s habitat or region.
The total frequency is very high (1388 %), showing that macrofungi fmit very
abundantly in the C
zone. Villeneuve et al. (1988) reported a maximum ftequency
of 185% on balsam fir-paper birch forests and Brunner et al. (1991) reported a maximum
of 350% on Alnus crispa forest in Alaska. However, both these studies used quaclrats one
quarter of the area used in this study, which could reduce their fiequencies by a
theoretical maximum of a factor of four but more likely by a much smaller factor. Also,
Villeneuve et al. (1988) and Brunner et al. (1991) did not sarnple through the entire
fmiting season and so missed fungi that were present on their sites but that fi-uit at
different times of year. Additionally, the Douglas-fir forests in the present study had
higher species richness, which contributes to higher total ftequency. The frequency
recorded in this study seems more reasonable when these factors are taken into
consideration.
The rank abundance curve shows that there are some dominant species in the
macrofungal community. The steeper slope at the start of the cuve may be an artifact
because many of the most common "species" are actually species clusters containing
multiple species, e.g. Cortinarius spp. subgenus Telamonia, which artificially inflates
their abundance. The Shannon Weaver equitability index and the Berger-Parker
dominance index both indicate that equitability is high in the macrofbngal community.
This may seem contradictory to the rank-abundance curve results, but this can be
reconciled when one considers that high numbers of rare
that the most dominant species accounts for only 4% of the total abundance. The many
rare species contribute more to the equitability index. The equitability reported here
closely agrees with that of Villeneuve et al. (1989) who examined macro
communities in sugar maple - yellow birch forests, balsarn-fir - paper birch forests, closed
canopy black spruce forests and open canopy black spruce forests in Quebec and found
equitabilities of 0.87, 0.82,0.80 and 0.87, respectively. However, the results from both
these studies are biased by the method of measuring abundance. Measuring abundance as
the frequency of occurrence on quadrats is not an ideal method because it tends to
underestimate the abundance of common species because it sets upper and lower limits
on abundance according to the nurnber of quadrats sampled. Also it does not consider
biomass; the relative importance of macrofbngi that produce very large sporocarps, like
Russula brevipes, is down played while the importance of macro
miniscule sporoc
s, such as Hemimycena delectabilis, is enhanced. If biomass had been
used as measure of abundance in this study, then the equitability might have been much
lower.
1.4.1.2 Annual variation in the macrofungal communi
The observed richness of the macrofungal community varied greatly from year to
year as expected. The macrofungal CO
and root tips for mycorrhizal fùngi, are always becoming available for colonization,
allowing new species to become established. Although part of the variation in richness
may be due to the loss or addition of species in the community, it is most likely that the
actual richness of the community changed little and the observed variation is due to the
sporadic nature of sporocarp production. The predominant factors that affect sporoc
production are temperature and rainfall (Widden 1981). Many species were probably
present in al1 three Octobers, but only fruited when the environmental conditions were
right. Also, because of the changes in sampling technique from year to year, it is not clear
how closely the observed variation in sporoc
production relates to the actual variation
in sporocarp production.
The change in species composition from year to year can also be attributed to the
sporadic nature of sporocarp production rather than the loss or addition of species. In
order to differentiate between variable sporocarp production and actual compositional
changes in the community it is necessary to combine sporoc
SurveYs with molecular
techniques for identiQing mycelia as Gardes and Bruns (1996) did for
ectomycorrhizal guild in Pinus muricata forests. Additionally, the fact that different areas
were sampled each year adds to the variability in species composition. If the exact same
quadrats had been sampled each year, then it is expected that the variability in species
composition would have been lower.
henology of the macrofungal community
The monthly changes in species composition were much greater than the annual
species that are fmiting represent
just the sarne species fruiting at different times. This indicates that different species are
induced to fmit by different environmental conditions and may have specific fmiting
seasons. The species composition in the month of August was very dissimilar from al1
other months not because of unique species that fmited but because very few macrofungi
fniited in August.
The fa11 season is the time of maximum diversity of macrofungal sporocarps and
if the objective of some research is to answer questions that do not require complete
characterization of the macrofungal community, then sarnpling effort should be
concentrated in this time. Maximum knowledge for minimum effort could be gained by a
single visit in October to CWHxm sites on Southern Vancouver Island. However, if more
complete knowledge of the macrofungal community is required then year round sampling
is necessary.
Species accumulation curves for the rnacrofungal communi
The equation of the species area curve can be used to extrapolate, with great
uncertainty however, the number of species expected if a larger area were sampled. If the
area sampled in 1997 had been doubled, the equation predicts a 26% increase in the
3O
number of species to 382. If one km2 was sampled, 260 times the area sampled in 1997,
then 1034 species of macrofùngi are expected. This shows that sampling effort must
increase exponentially for linear increases in the number of species observed and that a
complete list of al1 macrofungi in one habitat is not achievable through systematic
sampling. It should be noted that the true equation of the line must be asyrnptotic because
the nurnber of species cannot increase indefinitely and that the logari
no upper bound, so these nurnbers are likely overestimates.
The number of species observed also increased over time and might continue to
increase linearly for 20 years or more as has been reported for Caledonian pine forests
(Tofts and Orton 1998). If the number of species continued to increase at 93 species per
year for 17 more years it would bring the total number of species to 1966. It was an
is rate. Match Maker (Gibson 2000), a computer key to gilled macrofimgi, lists 2093
species expected to occur in the Pacific Northwest based on literature and anecdotal
evidence, and this does not include non-gilled macrofùngi which may represent another
1000 species.
The discussion of guild results is limited primarily to the mycorrhizal guild, litterdecay guild, and wood-decay guild. Results from the general saprobes guild are difficult
to interpret beeause it contains many groups of unidentified species that belong to
different guilds, and the parasite, fimgicolous and basidiolichen guilds each contain very
few species.
1.4.2.1 Diversity of macrofungal gui1
The CWHxm zone has a rich mycorrhizal guild due to the fact that the major tree
species (Douglas-fir, western hemlock, red alder), with the exception of western red
cedar, are ectomycorrhizal. Other studies of ectomycorrhizal host forests showed similar
percentages of mycorrhizal species (Table 1.8). Alder stands showed much lower
ectomycorrhizal richness because they tend to have alder-specific fungal symbionts and to
grow in pure stands. Percentages of other guilds also appear similar to those in other
studies in coniferous forests. Gamiet and Berch (1992) reported a lower percentage of
mycorrhizal species in the community, but also reported that that in the year of their study
the summer was unusually wet and the fa11 unusually dry. This may have reduced the
nurnber of mycorrhizal fungi fmiting since they primarily b i t in the fall. The lower
percentage of wood-decay
gi recorded in this study than in that by Gamiet and
(1992) can be explained by the fact that coarse woody debris was not systematically, or
selectively, sampled in this study. Until fùrther studies are conducted, it cannot be
concluded whether the reported relative proportions of guild richness are typical of this
forest type.
The mycorrhizal guild is the most frequent, accounting for 48.5% of the total
frequency while its share of species richness is 41%.
s indicates that it contains some
dominant species and indeed 10 of the 15 most frequent species are mycorrhizal. The
wood-decay fungi, on the other hand, account for a lower share of the total frequency than
they do of species richness, indicating that wood-decay species are not dominant. This
could be partly because coarse woody debris was not studied systematically. Litter-decay
fùngi also accounted for a much lower percentage of the total frequency than they did of
the total species richness. This is an artifact due to the sampling method used because
macrofùngi smaller than 2 cm cap diameter (mostly litter-decay fungi of the genera
Mycena and Marasmius) were sampled from nested smaller quadrats in the fa11 of 1997,
the main fruiting season. Thus, less area was sampled and their overall frequency was
reduced. If al1 fungi had been sampled on the larger quadrats, some species of Mycena
would undoubtedly have been among the most frequent fungi. However, this partitioning
in sarnpling effort may have resulted in a more accurate view of the dominance of species
in macrofungal community by correcting for differences in biomass. If 1000 sporocarps
of Hemimycena delectabilis, one of the most abundant litter-decay fungi sampled on the
smaller quadrats, were collected, their biomass would be less than that of one sporocarp
of Russula brevipes.
The high evenness in the mycorrhizal and litter-decay guilds might be due to the
dynamic state of the resources they use. Mycorrhizal fungi must compete for space and
root tips as the root tips of their host grow. Replacement of one mycorrhizal species by
another often occurs when root growth resurnes following periods of cold or drought
(Fleming 1985, Fleming et al. 1984) This turnover and competition could prevent species
from becoming dominant. A similar process occurs in the litter decay guild where the
influx of new substrates from the canopy presents the opportunity for competition. In both
situations, random factors - e.g. being the lucky spore in the right place at the right time may play an important role in deciding the outcome of the competition. In the wooddecay guild, the colonization of new substrates occurs less frequently and the substrates
are much larger discrete units that exist for much longer periods and may allow more
cornpetitive species to become dominant.
1.4.2.2 Annual variation in rnacrofun
The general trends in annual variation in the richness were similar for each guild,
showing that sporocarp production of al1 guilds is affected similarly by the changing
environmental conditions. The fact that the relative proportions of the number of species
in each guild changes Erom year to year shows either that the magnitude of the response
varies with each guild or that some other factors besides temperature and rainfall, perhaps
guild specific, are affecting sporocarp production. For example, if it was a particularly
good year for tree growth, the increased photosynthate production would be passed on to
the mycorrhizal guild immediately, the increased needle production might affect the litterdecay guild a few years later when the needles drop, and the wood-decay guild would be
unaffected.
henology macrofungal guilds
The phenology of fmiting for the different macrofungal guilds may be related to
their strategies for colonizing new substrates. For litter decay fungi, fa11 fniiting puts
their spores in place to compete for the flush of litter that accumulates over the winter and
becomes available when the snow melts. For litter fungi, the fa11 season is less dominant
over the spring and surnmer seasons than it is for mycorrhizal fungi, possibly because
litter falls year round, and litter-decay hngi must fiuit throughout the year to colonize
new substrates and new areas. The opportunity to colonize new substrates is not strongly
tied to season for wood-decay fungi and this may be why the wood-decay guild showed
the least variation with season. In addition, many wood-decay fùngi produce perennial
sporocarps, stabilizing the observed richness and frequency of the guild through the year.
ener
The dominant genera in terms of frequency are similar to the dominant genera in
terms of richness with the large (species-rich) genera being the most frequent. One
notable change is that Tricholoma is ranked third for ric
ss but seventh for frequenc~
indicating that its species are not cornmon fungi
ecies
1.4.4.1 Frequencies of individual species
Five of the six most frequent species are actually groups of species not identified
further and should not be considered as dominant. Some of
to the west, e.g. Guepiniopsis alpinus (Arora 1986), Lactarius rubrilacteus, which grows
with Douglas-fir (Arora 1986), Marasmius salalis which grows with sala1 (Gaultheria
shallon), and Strobilurus trullisatus which grows only on Douglas-fir cones. Others are
more cosmopolitan, such as Hebeloma crustuliniforme,Russula brevipes and Mycena
pura (Arora 1986).
1.4.4.2 Species
The cluster analysis of the monthly species composition identified that there were
three distinct fmiting seasons, each with a distinct assemblage of species, and
examination of the monthly fkequencies allowed the identification of species that h i t e d
in each seasan.
evie
Normally, ecological studies like this one are carried out in areas that have been
thoroughly mycologically described, since this makes studies faster and more efficient.
But when the forests are being cut down as fast as they are being cut today it does not
leave time to do things in the normal way. Thus, out of necessity, this study has gone
somewhat against the nom. Because of this, it was somewhat ambitious and precocious. I
did not know how many species would be encountered and certainly was no expert in
fungal identification. It was a learning process and the primary goal was to determine the
effects of clearcutting on macro
gi, not to carefully catalogue the macrofungi present.
This meant studying as many sites as possible. In the field there was not always time to
find specimens suitable for herbaria; some specimens were in poor condition and had to
be discarded after identification. Also, some easily identified
e field and never collected because it was assumed that voucher specimens already
existed. In the lab there were simply too many specimens to process before some of them
rotted. Some specimens were identifiable but not suitable for herbarium specimens and
were discarded with the assumption that more would be collected at a later date. The
species list compiled during this study is by no means complete, many uncertain species
were lumped under their genus name, and many species have no voucher specimens.
There are also undoubtedly some misidentifications, but uncertain species have been
noted with veE a 8
The changes in sampling methodology in each of the years were unavoidable. The
1995 data were collected by the South Vancouver Island Mycological Society and the
author had no input into the design of that study. The number of
onosequences was
36
increased to three to give more statistical power and the sampling area at each
chronosequence was reduced to what was considered manageable, al1 macrofungi
sampled on two transects per stand. On the first day in the field in 1996, it was discovered
that it was not possible to sample al1 macrofungi on the transects because of the
ovenvhelming abundance and diversity of the small litter decay fungi. A decision was
made in the field to subsample al1
gi with a cap diarneter of less than 2 cm. Sporocarp
counts collected in 1995 and 1997 proved difficult to analyse because they were
somewhat subjective, and were biased towards the litter decay fùngi that produce
abundant small sporocarps. In 1997,20 quadrats were sampled in each stand to allow
frequency to be used as a measure of abundance. Because of these changes in sampling
methods, al1 year to year comparisons must be interpreted with caution.
Douglas-fir dominated
e 1.1 Richness of the macrofungal community in C
forests on southeastern Vancouver Island.
Number of
species
Number of
genera
200
144
302
385
140
120
212
296
92
65
108
128
71
59
69
103
1995
1996
1997
1995-1997
Oct. 1995
Oct. 1996
Oct. 1997
Oct. 1995,1996 & 1997
le 1.2 Comparison of macrofungal community composition in CWHxm Douglas-fir
dominated forests for each month in 1997 using Sorenson's coefficient of similarity.
0.48
ay
June
July
August
Se tember
0.36
0.43
0.37
0.31
0.40
0.28
0.23
0.39
0.59
0.00
0.00
0.03
0.10
Q.22
0.20
0.13
0.26
0.33
0.47
0.25
June
July
August
September
October
October November
0.10
0.08
0.18
0.23
0.36
0.15
0.5 1
0.13
0.10
0.23
0.24
0.38
0.17
0.47
0.71
e 1.3 Richness, cumulative frequency and evenness of macrofungal guilds of in
C W x m Douglas-fir dominated forests.(s - the number of species, p, - percentage of the
total richness,f - cumulative frequency, pf - percentage of total frequency, J' - ShannonWeaver equitabiliy index, d - Berger-Parker dominance index)
1995-1997
Guild
mycorrhizal
litter-decay
general saprobes
wood-decay
fungicolous
basidiolichen
parasite
S
P,
("w
148 38.4
118 30.6
36 9.4
76 19.7
3 0.8
2 0.5
2 0.5
1997
s
P, f
Pf
("A) (%) (%)
126 41.7 673.3 48.5
88 29.1 355.0 25.6
28 9.3 178.8 12.9
56 18.5 160.8 11.6
2 0.7
2.5
0.2
1 0.3 10.8 0.8
1 0.3 7.1 0.5
J'
0.82
0.85
0.72
0.75
d
0.09
0.07
0.24
0.27
Richness and proportion of total richness of macrofungal guilds of
dominated forests in the month of October 1995, 1996, and 1997. (s - number of species,
p - percentage of the total richness)
Oct. 1995
Guild
s
mycorrhizal
litter-decay
general saprobes
wood-decay
fùngicolous
basidiolichen
parasitic
47
47
14
29
2
1
0
p (%)
33.6
33.6
1O
20.7
1.4
0.7
O
Oct. 1996
S
43
37
12
25
2
1
O
p (%)
35.8
30.8
10
20.8
1.7
0.8
0
Oct. 1997
S
102
58
20
29
1
O
1
p(%)
48.3
27.5
9.5
13.7
0.5
0
0.5
Oct. 1995,1996
& 1997
S
p(%)
123
41.6
90
30.4
28
9.5
51
17.2
2
0.7
1
0.3
1
0.3
al genera by species richness (S), and frequency V) and
their relative proportions ( p , p f) in CWHxm Douglas-fir dominated forests.
.5 Dominant macro
Genus
1995 - 1997
Genus
1997
Genus
1997
(%) p (%
Mycena
Cortinarius
Tricholoma
Russula
Inocybe
Clitocybe
Collybia
Lactarius
Gymnopilus
Hydne llum
Hydnum
Hygrocybe
Marasmius
Galerina
Hygrophorus
Suillus
Camarophyllus
Crepidotus
Dermocybe
Lycoperdon
Pholiota
49
32
14
13
10
9
9
8
7
7
7
7
7
6
6
6
5
5
5
5
5
12.7
8.3
3.6
3.4
2.6
2.3
2.3
2.1
1.8
1.8
1.8
1.8
1.8
1.6
1.6
1.6
1.3
1.3
1.3
1.3
1.3
Mycena
Cortinarius
Tricholoma
Russula
Inocybe
Clitocybe
Collybia
Lactarius
Galerina
Gymnopilus
Suillus
Dermocybe
Hydnellum
Hydnum
Hygrophorus
Lycoperdon
Marasmius
Camarophyllus
Gomphidius
Hygrocybe
Nolanea
34
23
13
11
9
7
7
7
6
6
6
5
5
5
5
5
5
4
4
4
4
11.3
7.6
4.3
3.6
3.0
2.3
2.3
2.3
2.0
2.0
2.0
1.7
1.7
1.7
1.7
1.7
1.7
1.3
1.3
1.3
1.3
Mycena
Cortinarius
Russula
Inocybe
Galerina
Lactarius
Tricholoma
Marasmius
Guepiniopsis
Nolanea
Hemimycena
Laccaria
Hebeloma
Dermocybe
Hygrophorus
Collybia
Pseudohydnum
Lyophyllum
Clitocybe
Xeromphalina
Hydnellum
176.7
157.5
117.9
73.8
61.7
55.8
46.3
44.2
43.8
37.5
36.7
30.4
28.3
27.5
18.3
17.9
17.9
17.1
16.3
15.8
15.0
12.8
Most frequently encountered species in C W x m Douglas-fir dominated forests
in 1997. Guilds: a - basidiolichen guild, f - fungicolous, 1 - litter-decay, m - mycorrhizal,
p- parasite, s - general saprobe, w - wood-decay
Species
Guild
Frequency
Proportion by fre
(%)
Cortinarius sg. telamonia sp.
Guepiniopsis alpinus
Inocybe spp.
Mycena spp.
Russula spp.
Galerina spp.
Lactarius rubrilacteus
Russula placita
Hemimycena delectabilis
Marasmius salalis
Cortinarius acutus vel af
Cortinarius obtusus vel a 8
Hebeloma crustulinforme
Russula brevipes
Mycena pura
Cortinariusfulvescens vel a f
Laccaria laccata
Pseudohydnum gelatinosum
Nolanea holoconiota vel a f
Galerina emetensis vel a f f
Lyophyoyllum semitale
Nolanea spp.
Russula xerampafina
Mycena alcalina
Mycena rorida
Dermocybe phoenicea
Ramaria sp.
Clavulinopsisfusiformis
Helvella lacunosa
Tricholoma spp.
Omphalina ericetorum
Mycena sp. 1
Xeromphalina campanella
Inocybe ovatocystis
Mycena oregonensis
Otidea alutacea vel af
m
w
m
s
m
s
ln
m
1
1
m
m
m
m
1
m
m
w
s
1
1
s
m
s
1
m
m
1
1
m
a
1
w
m
1
1
57.9
43.8
43.3
43.3
40.4
38.3
38.3
30.4
26.3
25.8
25.4
25.0
24.6
22.1
21.3
18.8
18.8
17.9
17.5
17.1
15.8
15.8
13.8
12.9
12.9
12.5
12.5
12.1
11.7
11.3
10.8
10.4
10.4
10.0
10.0
10.0
4.2
3.2
3.1
3.1
2.9
2.8
2.8
2.2
1.9
1.9
1.8
1.8
1.8
1.6
1.5
1.4
1.4
1.3
1.3
1.2
1.1
1.1
1
0.9
0.9
0.9
0.9
0.9
0.8
0.8
0.8
0.8
0.8
0.7
0.7
0.7
m Douglas-fir
.7 Presencelabsence and fiequency of macroîungi in C
dominated forests on southern Vancouver Island fiom 1995 to 1997.
G = guild. Guilds are: a - lichen, f - fungicolous, 1-1itter decay, m - mycorrhizal, p - parasitic, s - general
saprobes, and w - wood-decay; + = present, numbers are % frequency.
1997
ay Jun Sul Aug Se
Species
Agaricus a m i s
Agaricus semotus vel afl
Agaricus sp.
Agrocybe sp.
Albatrellus sp.
Amanita porphyria
Amanita silvicola
Amanita vaginata
Armillaria ostoyae
Auricularia sp.
Auriscalpium vulgare
Baeospora myosura vel af
Bisporella citrina
Boletopsis subsquamosa
Boletus chrysenteron vel a#
Bondarzewia montana
Bovista plumbea vel afi
Callistosporium luteo-olivaceum vel
a4
Calocera viscosa
Camarophflus borealis
Camarophyllus niveus
Camarophylluspaupertinus vel afi
Camarophyllus recuwatus vel a#
Camarophyllus sp.
Cantharellula umbonara
Cantharellusformosus
Cantharellus infundibuliformis
Cantharellus subalbidus
Chlorociboria aeruginascens
Chroogomphus tomentosus
Ciboria rufofùsca
Clavaria vermicularis
Clavariadelphusligula
Clavariadelphustruncatus
Clavicorona taxophila
Ctavulina cinerea
Clavulina cristata
Clavulinopsis corniculata
ClavuIinopsisfus formis
Clitocybe avellaneialba
1
1 +
1
1
m
0.4
0.4
0.8
1.3
0.4
0.4
0.8
0.4
m
m
m
P
0.4
0.4
0.4
0.4
0.4
0.8
0.4
0.4
0.8
0.4
7.1
2.5
+
+
0.4
0.4
+ +
1.3
4.6
w +
1
1
w
m
m
s
s
w
1.7
+
1.3
1.3
1.3
1.3
+
+
+
w +
m
1.3
+
+
4.2 0.4
2.1
s +
s +
0.8
s +
0.4
s
1
m
1.7
+
1.3
0.8
3.8
0.4
0.8
m + +
1.7
1
3.3
m
m
+ +
w
m +
m
m
+
5.0
1
m
+
0.4
0.4
1.3
m +
+
0.8
3.3
+
0.8
12.1
s
1 +
s +
2.1
0.8
0.4
0.4
1.7
0.8
0.4
1.7
0.4
0.8
0.8
0.4
0.4
0.4
0.4
0.4
0.8
0.8
0.4
1.7
0.4
0.8
3.3
0.8
3.3
1.7
2.5 2.1
0.4
0.4
0.4
0.4
0.8
2.5
0.8
0.4
0.4
0.8
7.9 6.7
1.3 0.8
Species
Clitocybe conferophila vel aff
Clitocybe dealbata vel afl
Clitocybe decepliva
Clitocybe harperi
Clitocybe inversa vel aff
Clitocybe nebularis
Clitocybe sp. 1
Clitocybe spp.
Collybia acemata
Collybia butyracea
Collybia cirrhata
Collybia confluens
Collybia dryophila
Collybia maculata vel aff
Collybia racemosa
Collybia spp.
Collybia tuberosa
Coltriciaperennis
Conocybe tenera vel af
Cordyceps sp.
Coriolellus sepium vel a f f
Cortinarius aculus vel a f f
Cortinarius alboviolaceus
Cortinarius anomalus
Cortinarius boulderensis vel aff
Cortinarius brunneus vel a 8
Cortinarius callisteus
Cortinarius caninus vel aiff
Cortinarius casfaneus vel afl
Cortinarius clandestinus
Cortinarius claricolor
Cortinarius dilutus vel afl
Cortinariusjülvescens vel a@
Cortinarius gentilis vel afl
Cortinariusjubarinus vel af
Cortinarius nigrocuspidatus vel aff
Cortinarius obtusus vel aff
Cortinariusplumiger vel aff
Cortinarius pyriodorus
Cortinarius rapaceus vel aff
Cortinarius scaurus vel afS
Cortinarius sg. leprocybe spp.
Cortinarius sg. myxacium spp.
Cortinarius sg. phlegmacium spp.
Cortinarius sg. serioqbe spp.
1997
G 9 5 '96 '97 Mar A r May Jun Jul Aug Se
s
3.8
1
1
1
+
+ +
+
1
1 + +
1
1 +
+
w
1
f
1
1
s
f
s
f
m
1
P
w
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
m
1.3 2.9
+
1.7 0.8
0.4
0.4
O .4
0.4
0.8
0.4
0.8
5.0
0.4
2.9
1.7 1.3
6.3
0.8
0.8
+
+
+
+
2.5
0.8
3.8
0.4
0.4
0.8
0.4
1.7
0.4
0.4
+
0.8
+
+ +
+ +
+ +
7.5
1.7
6.3
5.4
0.8
2.1 2.1
2.1 2.1 2.5
1.3 2.9
2.1
1.3
0.8
1.3 0.8
0.8 0.4
0.8
0.4
+
+
+ + 25.4
0.4
1.3
+
+
+
0.4
15.8 13.3
0.8 0.4
0.4
+
0.4
0.4
1.3
0.8
+
+ +
+
0.8
+
6.3
6.3
+
18.8
0.8
0.8 18.3
0.8
0.4 2.1
2.5
0.4
+
+ +
+
25.0
0.8 21.7
0.4
0.4
0.4
0.4
0.4
0.8
0.4
0.4
2.5
2.1
0.4
0.4
5.4
+
0.4
0.4
Cortinarius sg. telarnonia spp.
Cortinarius sp. 1
Cortinarius sp. 2
Cortinarius squamuloszrs vel aff
Cortinarius superbus vel aff
Cortinarius vanduzerensis
Cortinarius vibratilis
Cortinarius violaceus
Crepidotus applanatus
Crepidotusfusisporus
Crepidotus herbarum
Crepidotus mollis vel aff
Crepidotus spp.
Cryptoporus volvatus
Cudonia circinans
Cudonia monticola
Cystoderma amianthinum
Cystodermafallax
Cystoderma granulosum
Dacrymyces palmatus
Dasyscyphus bicolor
Dentinum umbilicatum
Dermocybe californica
Dermocybe cinnamomes vel aff
Dermocybe malicoria
Dermocybe phoenicea
Dermocybe sernisanguinea
Discina perlata
Entoloma sp.
Fomitopsis cajenderi
Fomitopsis oflcinalis
Fomitopsis pinicola
Galerina autumnalis
Galerina badipes vel a#
Galerina emetensis vel a#
Galerina mammilata
Galerina spp.
Galerina vexans vel aff
Geoglossum sp.
GloeophylIum saepiarium
Gomphidius oregonensis
Gomphidius smithii
Gomphidius spp.
Gomphidius subroseus
Guepiniopsis alpina
m +
+ 57.9 0.4 0.8
ay Jun Jul Aug Sep Oct Nov
0.8 1.3 5.8
2.1 51.3 24.6
43
Species
Gymnopilus bellulus vel a@
Gymnopilus croceoluteus vel a8
Gymnopilus picreus
Gymnopilus sapineus vel afl
Gymnopilus spectabilis
Gymnopilus spp.
Gymnopilus terrestris
Gyromitra esculenta
Hebeloma crustuliniforme
Hebeloma mesophaeum vel a 8
Hebeloma sacchariolens
Helvella lacunosa
Helvella queletii vel aff
Hemimycena delectabilis
Hemimycena sp. 1
Hemimycena spp.
Hydnellum aurantiacum
Hydnellum caeruleum
Hydnellum conigenum
Hydnellum peckii
Hydnellum scrobiculatum vel ag
Hydnellum spp.
Hydnellum suaveolens
Hydnum calvatum
HydnumJennicum
Hydnum fuscoindicum
Hydnum scabrosum vel aff
Hydnum sp. 1
Hydnum sp. 2
Hydnum subrosum vel aff
Hygroqbe conica
Hygrocybe flavescens
Hygrocybe miniata
Hygroqbe sp. 1
Hyk7ocrbe SPP.
Hygrocybe unguinosus vel a f
Hygrophoropsis aurantiaca
Hygrophoropsis olida
Hygrophorus agathosmus
Hygrophorus bakerensis
Hygrophorus camarophyllus
Hygrophorus eburneus vel a f
Hygrophorus picea
Hygrophorus sp. 1
Hygrophorus spp.
1997
ay Jun Jul Aug Se
w
w
+
w
w
w
+
w +
+
+
m +
m +
m +
1 +
+
+
+
+
1
1 +
1
+
1
+
+
+
2.9
0.4
0.8
24.6
2.9
0.8
11.7
0.4
26.3
8.3
2.1
6.7
2.1
0.8
1.7 0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
20.0 7.5
2.9
0.8
0.8 6.7 9.2
22.5 14.6
6.3 4.2
1.3 0.4
1.3 2.5 1.7 3.3 1.3
0.4 0.4 0.4 1.7
+
+
+
+
+
+
m
m +
m +
1 +
1 +
1
1
1
1
+
+
s
+
1 +
m
2.1
0.4 1.7 0.4
3.8
2.9
0.4
1.7
0.4
0.4
7.1
1.3 5.4 0.4
1.3 1.7 1.3
0.4 2.1 2.1 0.8
0.4
0.8 0.8
0.4
0.4
0.4
+
+
m
m
m
m
m
m
m
0.8
0.4
0.4
1.7 2.1
+
s
1
m
m
m
m
m
m
m
m
m
m
0.8
0.4
0.4
3.8
+
8.8
+
3.3
0.4
+
+
0.4
1.3
0.4 7.5 2.1
2.1
0.4
2.5
4.6
0.8
0.4
2.9
0.4
0.4
1.7
0.8
1.3 4.2
0.4 0.4
0.4
+
+
+ +
7.9
0.4
4.6
1.7
0.8
1.3 5.0
0.4
2.5 2.1
le 1.7 continued
Species
Hypholoma capnoides
Hypholoma fusciculare
Inocybe calamistrala
Inocybe geophylla
Inocybe lacera vel a#
Inocybe lilacina
Inocybe maculata vel a#.'
Inocybe mixtilis
Inocybe napipes vel afl
Inocybe ovatoqstis
inocybe sororia
Inocybe spp.
Kuehneromyces vernalis
Laccaria altaica vel a#
Laccaria amethysteo-occidentalis
Laccaria bicolor
Laccaria laccata
Lactarius luculentis
Lactarius mucidus ve2 a$
Lactarius pallescens
Lactarius psezrdomucidus
Lactarius rubrilacteus
Lactarius rufus vel aff
Lactarius scrobiculatus
Lactarius spp.
Leccinum manzanitae
Lentaria spp.
Lepiota clypeolaria
Lepiota seminuda vel aff
Lepiota sp.
Leptonia usprella vel a 3
Leptonia carnea
Leptonia sp.
Leptonia undulatella
Lycoperdonfoetidum
Lycoperdon molle
Lycoperdonperlatum
Lycoperdon pyriforme
Lycoperdon sp.
Lyophyllum decastes
Lyophyllum semitale
Macrotyphula~stulosavel afl
Marasmiellus candidus vel f l
Marasmius androsaceus
Marasmius epiphyllus vel afl
ar Apr May Jun Jul Aug Se
w
5.4
1.3 4.2
Species
Marasmius limosus vel a f f
Marasmius salalis
Marasmius sp. 1
Marasmius spp.
Marasmius umbilicatus
Marcelleina sp.
Melanoleucu graminicola
Melanotus textilis vel aff
Merulius tremellosus
Micromphale foetidum vel a f f
Micromphale perforans vel aff
Mitrufa abietis vel a 8
Mollisia sp.
Multiclavula mucida vel afl
Mycena fusco-ocula vel a f f
Mjcena subsanguinolenta vel afl
Mycena acicula
Mycena adonis
Mycena alcaiina
Mycena alnicola
Mycena amabilissima
Mycena amicla
ficena aurantiidisca
Mycena brownii vel aff
A@cena capillaripes
Mycena capillaris
Mycena citrinomarginata
Mycena clavularis vel ag
Mycena concolor vel aff
Mycena elegantula
Mycena epipterygia
Mycena epipterygioides
Mycena jlavoalba
Mycena galopus
Mycena gaultherii
Mycena haematopus
Mvcena inclinata
Mycena iodolens vel aff
Mycena leptocephala vel aff
Mycena Iitoralis vel aff
Mycena m a c h t a
Mycena metata
Mycena oregonensis
Mycena paucilamellata
Mvcena pura
1
1 +
1
s
+
l +
w
1
s +
w
1
1 +
1 +
w
a +
1 +
1
0.4
+ 25.8
2.9
+ 8.3
+
0.8
1.7
+
1997
ar Apr May Jun Jul Aug Sep
0.4
0.8
0.4
8.8
1.7
0.8 2.1 2.5
2.1
Oct Nov
16.7 5.4
1.3 0.8
1.3 0.4
0.8
0.4
0.4
1.3
0.4
0.4
+
0.8
0.4 0.8
+
1
s
+
+
s
w
1
w
1
w
1
1
1
w
1
w
s
1
1
1
1
w
w
1
s
1
w
+ + 12.9
0.4
+ +
+
+ +
+
+
+
0.8 0.8
1.3
0.8
5.4
0.4
2.5
0.4
7.1
0.4
1.7 5.4 4.6
0.8
0.4 4.2 1.7
0.4
1.7 0.4
4.2
2.1
+
0.8
0.4
0.4
+
+
+ +
+
0.4
1.7 5.8 2.1
5.0 1.3
1.3
1.3
0.8 0.4
0.8 0.4
+
+
+
0.4
9.2
5.4
+
+
0.8
0.8
0.4
0.4
0.8
+
+
+
+ +
1
1 +
1
1 +
1.3
0.4
+ 10.0
O .4
+ 21.3
1.3
0.4
0.8 8.8
0.4
1.7
0.8
4.6 16.3 2.5
Speeies
Mycena purpureofusca
Mycena rorida
Mycena rubromarginala
Mycena sanguinolenla
Mycena sp. 1
Mycena sp. 2
Mycena sp. 3
Mycena sp. 4
Mycena sp. 5
Mycena sp. 6
Mycena sp. 7
Mycena sp. 8
Mycena spp.
Mycena strobilinoides
@cena subcana vel a f f
Mycena tenerrima
Mycena vitilis vel a f f
Mycena vulgaris vel aff
Neournula pouchetii
Nidula niveotomentma
Nolanea cetrata vel a&?
Nolanea holoconiota vel afl
Nolanea sericea vel afl
Nolanea spp.
Oligoporus caesius
Oligoporusfragilis
Oligoporus tephroleucus vef afl
Omphalina chloroqanea
Omphalina chrysophylla
Omphalina ericetorum
Osteina obducta
Otidia alutacea vel aff
Panellus serotinus vel aff
Panellus stipticus
Paraeccilia sericeonitida var.
ligniphila
Paxillus atrotomentosus
Paxillus involutus vel afl
Peziza spp.
Phaeolus schweinitzii
Phellinus sp.
Phellodon atratus
Phellodon melaleucus
Phellodon tomentosus
Pholiota decorata
w +
1
+
+ +
1997
ay Jun Jul Aug Sep Oet Nov
0.4
0.4 0.4 0.8
1.3 0.8 2.1
6.7 1.7 1.3
2.1
12.9
W + +
1
+ 0.4
1
+ 10.4
1
+ 9.6
1
4.2
1
0.4
1
7.9
1
0.4
1
w
s
1
10.4
1.7 6.7
2.5 0.8 1.3
0.4
7.5 0.4
1.3
0.4
+
0.4
43.3 2.5 2.5
+ 1.3
0.4
0.4
+ +
w +
1
1 +
1
+
1
w
s
+ +
s
s
s +
w
w
w +
1
w
a +
w
1 +
w +
w
w
m +
m +
0.8
7.9
+ 3.8
17.5
0.4
+ 15.8
0.4
1.3
+
0.4
0.4
4.6
0.4
0.8
1.3 2.1 2.5
1.3 2.9
12.1 2.1 6.3
+
m +
m
m +
+
+
+
7.1
1.3
2.5 3.8
0.4
0.4
5.4
1.7
0.4
1.3
1.3
5.8 0.8
0.4
2.9
0.8
0.8
1.7 9.2 3.3
0.4
0.4
0.4
0.4
0.4
0.4
2.1
1.7
2.1
1.3
5.8
0.8
0.4
0.4
0.4
1.3
0.4
+
2.5 32.5 5.8
0.4 0.8
1.7 4.2
3.3
+ 10.8 0.4 0.4
0.4 0.4
+ 10.0
1
w +
w
w
0.4
1.7 0.8
0.4
0.4
0.8
1.7
0.4
0.8
0.8
0.4
2.5 3.8
0.8
0.4
e 1.7 continued
Species
Pholiota highlandensis vel a 8
Pholiota malicola vel a f f
Pholiota sp.
Pholiota terrestris
Phylloporus rhodoxanthus
Plectania melastorna
Pleurocybella porrigens vel a#
Pleurotus ostreatus vel aff
Pluteus cewinus
Pluteus flavofulgineus
Polyporus elegans
Polyporus hirtus
Psathyrella gracilis vel a$
Psathyrella longistriata
Psathyrella spp.
Pseudocoprinus disseminatus vel a 8
Pseudohydnum gelatinosum
Pseudoplectania melaena
Pseudoplectania nigrella
Psilocybe corneipes vel a$
Psilocybe rnontana
Ramaria sp.
Rhodocybe hirneola vel aff
RickenellaJibula
Russula JLagilis vel afl
Russula adusta vel a f
Russula bicolor
Russula brevipes
Russula cascadensis vel aff
Russula emetica vel a$
Russula marei
Russula occidentalis vel afjr
Russula placita
Russula sp. 1
Russula spp.
Russula variata vel a f
Russula xerampalina
Sarcosphaera crassa
SpathulariaJavida
Stereum hirsutum vel aff
Stereurn sanguinolentum
Strobilurus albipilatus
Strobilurus trullisatus
Stropharia ambigua
Suillus borealis vel af
G
w
w
w
w
m
1
w
w
w
w
w
'95 '96 '97 Mar Apr May Jun Jul Aug Se
+
0.8
0.4
3.8
0.8
+
+
+ +
+ +
1
w +
1
1
1 +
1
m +
1 +
1
1.7 0.4
0.4
1.7
0.4
0.4
5.4 0.4
s +
1 +
1 +
1
m
m
m
m
m
m
m
m
m
m
m
m
m
1
1
w
w
s
1
1
m
+
+
+
2.5
0.4
2.5 0.4
1.3
2.1
0.8
0.4
0.4
0.4
0.4
0.4 0.4
0.4
0.8
0.8 0.4 0.8
+ 17.9 1.3
1.3 0.8
0.8 0.8
0.8 0.4 0.4
0.8
5.4 11.3 10.8
1.3 1.3 0.4
0.8
0.4
8.8 1.3 0.4
+ 12.5
0.8
2.1
0.4
1.3 0.4
9.2 3.8
2.1
0.8
1.3
+ 3.8
+ 22.1
1.3
0.4
0.4 0.4
2.9
1.7 6.3
0.8 0.4
0.4
0.4
0.8
17.1 7.5
0.8
2.5
1.3
+ 30.4
0.8
+ 40.4
0.4
6.3 0.4 1.3
0.8 0.4
+
+
0.8 0.4
+
+
0.4
2.5
1.3
0.4 7.1 24.6 4.6
0.8
2.5 2.1 11.3 30.4 1.3
+
+ 13.8
+
+ +
+
+ +
0.4
0.4
1.7
0.4
1.3
9.6
0.8
0.8
9.6
5.8 0.4
0.4
0.4
0.4
0.8
0.4
0.4
1.3
5.4 3.8 0.4
0.4 0.4
0.8
Species
Suillus granulatus
Sztillus lakei
Suillus luteza
Sztillus spp.
Suillus subolivaceous
Thelephora terrestris
Trametes hirsuta
Trametes versicolor
Tremella mesenterica vel a 8
Trichaptum abietinum
Tricholoma atroviolaceum
Tricholoma~avovirens
Tricholoma inamoenum
Tricholoma magnivelare
Tricholoma pessundatum
Tricholomaportentosum
Tricholoma saponaceum
Tricholoma sejunctum
Tricholoma sp. 1
Tricholoma spp.
Tricholoma sulphureum
Tricholoma lerreum vel a$"
Tricholoma virgatum
Tricholoma zelleri
Trichopilus plebiodes
Tubaria sp.
Tyromyces chioneus
Xeromphalina campanella
Xeromphalinafilvipes
Xeromphalina spp.
Xylaria hypoxyIon
m +
1.3
ay Jun Jul Aug Sep Oct Nov
1.3
Cornparison of habitat, guild structure (percentage of al1 species in each guild),
total richness, and area sampled of this study to previous studies. (M - mycorrhizal guild,
S - soi1 fungi guild, L - litter-decay guild, W - wood-decay guild, P - parasitic guild)
P
CWH old-growth
alder
sugar maple yellow
birch
balsam-fir paper
birch
black spruce
feather moss
taiga
Pinus sylvestris
hemiboreal
coniferous and
deciduous forests
boreal forests and
mixed forests and
peatlands
hornbeam oak
woods
beech woods
acid oak woods
evergreen oak
woods
No. of
Area
Country
Study
B.C. Canada
south coast, B.C.
Canada
4000 US(A1aska)
2000 Quebec
Gamiet and Berch
( 1 992)
Bntnner et al. (1 992)
Villeneuve et al. 1988
2000 Quebec
Villeneuve et al. (1988)
2000 Quebec
Villeneuve et al. (1988)
2000 Quebec
1O00 Nonvay
Estonia
Villeneuve et al. (1 988)
Sastad (1995)
Kalamees (1 980)
59600 Finland
Salo (1993)
Belgium
Darimont (1 973)
Europe
Netherlands
Italy
Lisiewski (1 974)
Jansen (1 98 1)
De Dominics and
Barluzzi (1 983)
.1 Map of coastal forest chronosequence sites (TroQxnow et al. 1997).
Macrofungi were sarnpled on the three southeastern-most chronosequences, Victoria
Watershed North, Victoria Watershed South, and Koksil
.2 Layout of transect/quadrats for sampling macrofimgi at each stand in 1995,
1996 and 1997. 1995: al1 macrofungi were sampled on four 40 m by 5 m transects. 1996:
macrofungi with a cap diameter greater than 2 cm were sarnpled on two 40 m by 4 m
transects and those with caps smaller than 2 cm were sampled on six 2 m by 4 m
quadrats. March - August 1997: al1 macrofùngi were sarnpled on 20 4 m by 8 m quadrats.
September - November 1997: macrofimgi with a cap diameter greater than 2 cm were
sarnpled on 20 4 m by 4 m quadrats and those with caps smaller than 2 cm were sampled
on 20 1.25m by 1.25m quadrats nested within the larger quadrats.
Species Rank
igure 1.3 Species rank vs. log abundance. Abundance is measured as the number of
quadrats on which the species was present.
1
=--O-
-..&..
richness
frequency
..
6
Mar Apr May Jun Jul Aug Sep Oct
Month
Seasonal variation in the richness and total frequency of the macro
cornrnunity in 1997 as measured by sporocarp production.
.5 Seasonal variation in species composition of the macrofungal eornmunity in
1997 as measured by the presencelabsence of sporocarps.
1O00
2000
Area sampled (m2)
3000
Number of macrofimgal species vs. area sampled in 1997
- - A &
- - October data
A
A
h'
'
year
.7 Number of macrofungal species observed versus time.
mycorrhizal
- -n. litter-decay
.ageneral saprobes
-o-
/
.P . . .
ar Apr May Jun Jul Aug Sep Oct
Month
Mar Apr May Jun Jul Aug Sep Oct
Month
Seasonal variation in the richness and cumulative fkequency of the four largest
macrofungal guilds in 1997 as measured by sporocarp production.
Considering that macrofüngi are known to be important to the
forest ecosystem (Trappe and Fogel1977, Fogel and Trappe 1978, O'Dell et al. 1996,
Pilz and Molina 1996), it is surprising that little research has been done on the long-term
effects of clearcutting on these organisms. Many studies have addressed the issue
indirectly, looking at things such as comparisons of different ages of second-growth
stands but not at the comparison of old-growth and rotation-age second growth. The
comparison of mature second growth to old-growth is essential, because if an old-growth
gus is absent or less abundant in mature second-growth then it will surely rernain
absent or become even less abundant in third-growth and later rotations. We have not
been clearcutting forests in B.C. for even one fifth the potential life-span of a Douglas-fir
tree. How can we conduct experiments testing the effects of forestry if we do not use oldgrowth as a control?
The objectives of this section are to address this
owledge gap, to determine if
the macrofungal community is affected by stand age, to identify components of the
community that may be at risk, and to suggest ways to reduce this risk. A part of this
chapter is devoted to power analysis. This is because being unable to prove that a given
component of the macrofungal community is affected by stand age does not conclusively
59
mean that the component is not affected; it is important to know how large an effect could
be detected and how likely it was that biologically significant effects could be detected.
The important îunctions of macrofungi in the forest ecosystem make
understanding the effects of forestry on macro
gi im~erative.Of p r i m interest
~
are
the effects of clear-cutting old-growth forest on macrofungal diversity and cornmunity
structure. There have been a number of studies on the effects of harvesting on different
aspects of the fùngal comrnunity such as the occurrence and diversity of ectomycorrhizal
sporocarps (Kropp and Albee 1996), formation of ectomycorrhizae on seedlings (Pilz and
Perry 1983), hypogeous sporoc
abundance (Clarkson and Mills 1994), and epigeous
and hypogeous sporocarp biomass (North et al. 1997). Others studies have investigated
the effects of other aspects of forestry on fungi such as the effects of burning on
macrofungi (Peterson 1971), the effects of logging waste removal or retention on
macroîungal communities (Wasterlund and Ingelgog 198l), the effects of soi1
compaction and organic matter removal on ectomycorrhizal root-tip abundance and
diversity (Amaranthus et al. 1996), the effects of nitrogen fertilization (Brandrud 1995)
and nitrogen-free fertilizer (Karen and Nylund 1996) on ectomycorrhizal fùngi. Despite
al1 of this attention to forest h g i , few studies have directly compared fungal diversity
and community structure in old-growth and clear-cut stands and those that did so focused
on mycorrhizal fungi.
2.1.1.1 Effects of clearcutting on fungal diversity and community structure
Kropp and Albee (1996) examined the effects of clearcutting and thinning on the
occurence of mycorrhizal sporocarps in lodgepole pine stands. The clearcut was 16 years
old, the thinned stand and undisturbed stands were 80 to 100 years old, and the thinned
stand had the number of stems per hectare reduced by about 50% 10 years prior to the
study. Clear-cutting reduced the species richness, total number of mycorrhizal sporocarps
and the fi-equency of fruiting of al1 species. Thinning reduced the total number of
mycorrhizal species and the fkequencies of some species but did not affect the total
nwnber of sporocarps. Different species and groups responded differently to the
treatments. Suillus brevipes becarne less abundant with increased levels of disturbance
but the genus Suillus as a whole had a higher relative abundance on clear-cuts than on
undisturbed plots. Fungi in the family Hygrophoraceae were most aflected by thinning.
Garbaye and Le Tacon (1982) also reported that some fungi responded differently to
different levels of thinning while others were unaffected. In another study of variable
retention forestry practises, Krannabetter and Kroeger (2001) found that reduction of
basal area of western hemlock could lead to increase, unchanged, reduced
ectomycorrhizal sporocarp ric ess and thus demonstrated that partial cutting systems can
be superior to clearcuts for maintaining mycorrhizal diversity.
O'Dell et al. (1992) conducted a preliminary study comparing mycorrhizal
communities on old-growth (400 years old), rotation age (50-60) years old, and closedcanopy young (25-30 years old) Douglas-fir stands. The two younger age classes were
plantations. Only 15 mycorrhizal mus oom taxa and 10 truffle taxa were observed.
number of mushroom species increased with stand age but the number of truffle species
showed little variation with stand age.
Kranabetter and Wylie (1998) examined changes in mycorrhizal root-tip diversity
on naturally regenerated western hemlock seedlings across 4-year-old 50 to 75m patch
cuts in old-growth western hemlock forests. Mycorrhizal richness was highest on
seedlings under the old-growth canopy. Seedlings at the forest edge showed 27%
reduction in mycorrhizal richness and seedlings in the forest opening showed a 40%
reduction in mycorrhizal richness. Pilz and Perry (1983) studied the formation of
ectomycorrhizae on Douglas-fir seedlings planted in clear-cuts and undisturbed Douglasfir dominated forests. Two of three undisturbed sites were old-growth. A greater
abundance of mycorrhizal root-tips formed on seedlings planted in clear-cuts but
diversity of mycorrhizal mo
otypes was higher on seedlings planted in undisturbed
forest.
North et al. (1997) examined the epigeous and hypogeous sporocarp biomass on
young, mature, and old-growth western hemlock (Tsuga heterophylla) stands. Epigeous
sporocarp biomass did not change with stand age. Hypogeous sporocarp biomass was
five times higher in mature and old-growth stands than in managed young stands.
Clarkson and Mills (1994) examined the differences in ectomycorrhizal hypogeous
sporocarp abundance (truffle abundance) between clear-cuts and old-growth Douglas-fir
forest remnants. Forest remnants had 20 to 40 times more truffles than adjacent clearcuts. Truffles were also found more often under coarse woody debris, and red-backed
voles were associated with areas with more truffles. Red-backed voles and other
mammals serve as vectors for truffle dispersal, and it was hypothesized that low ispersal
abilities might limit the recolonization of tniffles afier clearcutting. Late-sera1 forest
remnants were identified as vital refuges for hypogeous fungi and animal mycophagists
such as the red-backed vole.
Goodman and Troflmow (1998) compared ectomycorrhizal morphotype diversity
in old-growth and mature (fire origin, not logging) Douglas-fir stands on southeastern
Vancouver. However, their mature stands were of fire rather than harvest origin. They
found no differences in mycorrhizal diversity between these two forest ages.
2.1.1.2 Effects of logging waste on sporocarp biomass, numbers an
guilds
Wasterlund and Ingelgog (198 1) examined the effects of logging waste removal or
retention on macrofungi. No significant differences in the biomass or numbers of
sporocarps were seen between sites with and without logging waste. The ratio of
mycorrhizal uild fungi to decomposer uild fungi was apparently unaffected by the
removal of logging waste. However, sites that had logging waste had higher diversity
indices than those that had no logging waste.
2.1.1.3 Soil Compaction
Amaranthus et al. (1996) examined the effects of soi1 compaction and organic
matter removal on ectomycorrhizal root-tip abundance and diversity of Douglas-fir and
western white pine seedlings. Three levels of compaction (none, moderate, and heavy)
were combined with three levels of organic matter removal (bole removal, bole and
crown removal, and bole, crown and forest floor removal). Seedlings were planted in the
resultant clear-cuts and mycorrhizal root-tip abundance and diversity were examined one
year later. Western white pine was unaffected by any treatment, but Douglas-fir seedlings
showed significantly lower root-tip abundance on moderately and severely compacted
sites that had boles and crowns or boles, crowns and forest floor removed, and lower
mycorrhizal diversity on al1 severely compacted sites. Soil compaction has been shown to
decrease tree growth in Pinus contorta plantations (Clayton et al. 1987) and this may be
partly due to reduced mycorrhizal root-tip diversity and abundance.
2.1.1.4 Soil nitrogen and nitrogen fertilization
Brandrud (1995) examined the effects of nitrogen fertilizers on ectomycorrhizal
fungi. Addition of nitrogen caused a reduction in both the number of mycorrhizal species
present and the total nwnber of fruit-bodies formed. Different genera and species
responded differently to the addition of nitrogen. The genus Lactarius seemed to be
unaffected by nitrogen addition, while Cortinarius and Russula showed the largest
decreases in fmit-body production. By contrast, Paxillus involutus and Lactarius rufus
exhibited increased sporoc
production after nitrogen was added. Menge and Grand
(1978) also reported a decrease in overall numbers of sporocarps and number of species
of mycorrhizal fungi after nitrogen fertilization, with a variable response from species to
species.
Karen and Nylund (1996) examined the effects of nitrogen-fiee fertilizer on
underground ectomycorrhizal fungi communities. This fertilizer had no apparent effect on
fungal biomass, as measured by ergosterol analysis, or on the number of ectomycorrhizal
root-tip morphotypes, but a 50% reduction in the production of epigeous sporocarps was
noted.
De Vries et al. (1995) noted that increased atmospheric nitrogen deposition in the
Netherlands has led to a build up of nitrogen-rich litter and humus and a decrease in
ectomycorrhizal fungal richness. Removal of this layer led to increased species richness
and a change in community composition to include many earlier succession species.
2.1-1.5 Soil acidity and organic matter content
Soil pH is a major factor determining the distribution of many macrofungi (Tyler
1985, Hansen and Tyler 1992). In one study, macrofungi were studied in 300 plots in
Swedish beech forests, and metal ion saturation - a direct measure of acid-base condition
of the soil - and organic matter content were measured. It was found that 18 species were
associated with less acid soils, 24 species were evenly distributed across the edaphic
range and 21 species were associated with the more acid and hi
organic soils (Tyler
1985). Thus it was demonstrated that soil acidity and organic matter content are important
factors defining the niches of some macrofùngi, with some fungi being specialists and
others being generalists.
2.1.1.6 Soil temperature and moisture
Peredo et al. (1983) studied the factors affecting the occurrence and spatial
distribution of sporocarps of the ectomycorrhizal fungus Suillus luteus. They found that
average soil temperature and moisture were the most important factors and that solar
radiation was an important factor affecting temperature and moisture. Suillus luteus
sporocarps were confined to a narrow average soil temperature range of 13 - 15 OC and a
soil moisture range of 25-38% with an optimum of 32-33%. Although the fungus can
fruit only in a narrow range of conditions this does not necessarily correlate to the
conditions necessary for growth and survival of the mycelium. However, these results do
show that clear-cutting, which greatly alters solar radiation, soil temperature and moisture
regimes, may have a significant effect on fungi.
Theodorou (1978) showed that different ectomycorrhizal fungi grow optimally at
different soil water potentials and have differing abilities to survive drought. Different
ectomycorrhizal fungi also respond differently to temperature, showing varying growth
rates and abilities to form ectomycorrhizae (Theodorou and Bowen 1971). When clearcutting alters soil temperature and moisture regimes, it may also alter the fungal
community by affecting competition between fùngi.
Collecting and trampling due to collecting presumably have only local effects
(Rydin et al. 1997) unless collecting is carried out repeatedly and systematically, as with
Tricholoma magnivelare. However, the only detailed study of the effects of collecting,
that on Cantharellusforrnosus by Norvell et al. (1996), showed no effects.
Clearcutting is known to have deleterious effects on other groups of organisms, and
results from some studies on plants, insects and birds are given as examples. Herbaceous
understory plant communities in Appalachian mixed forests have lower richness and
cover after clearcutting, and do not recover within 87 years (Du@ and Meier 1992).
Zobel(1993) found that plant communities in pine forests (Pinus sylvestris) responded
differently after clearcutting according to edaphic factors. In Douglas-fir forests and
western hemlock forest on Vancouver Island, regeneration sites had higher species
richness than old-growth forest sites because most forest species survive, though in
reduced numbers, and invasive herbs appear (Ryan et al. 1994). However, in the western
hemlock forests there were a few les@ liverworts that were restricted to mature andor
old-growth stands and in the Douglas-fir forests seven species were restricted to mature
andor old-growth stands.
Okland (1994) found that clearcutting has a long-lasting effect on the
mycetophilid (fungus gnat) fauna in Nonvegian spruce (Picea abies) forests. Even after
120 years of regrowth, former clear-cuts had lower richness, and contained fewer rare
species than semi-natural (selectively cut) forests. The larvae of most species of
Mycetophilidae develop in fungal microhabitats (Hackman et al. 1988). OMand (1994)
also showed that the number of mycetophilid species was positively correlated wi
richness and abundance of polypores (wood-decay fungi), and suggested both that
diversity directly influences the diversity of Mycetophilidae, and that
maintained by a high degree of continuity, i.e., the absence of major disturbances such as
clearcutting or fire.
The diversity of other groups that feed on fungi, such as Collembola, may also be
directly influenced by changes in fungal diversity. Setala and Marshall (1994) exarnined
collembolan cornmunities in stumps on three Douglas-fir chronosequences (the sarne
ones used in this study). Old-growth and mature stands had the highest species richness
and regeneration stands had the lowest ric ess. Stand age was shown to be the most
important factor affecting species composition of the collembolan community. Al1 sera1
stages shared 63% of the collembolan species and the similarity was highest between
mature and old-growth and lowest between immature and regeneration.
Silvicultural treatments that retain structural and compositional vegetation
complexity or develop late old-growth characteristics quickly are used by more bird
species than stands that do not retain or develop these characteristics (Chambers and
McComb 1997). Selective or partial cutting exemplifj the former, and clearcutting the
latter.
Data from al1 monthly samples in collected in 1997 as described in Chapter 1
were pooled to find the total number of genera and the total number of species on each
plot. Data fi-om 1995 and 1996 are not presented because the different sampling methods
used in these years do not allow analysis of frequency data. No distinctions were made
between the two different levels of sampling, the large and small quadrats. Fungi were
assigned to one of seven guilds as defined in Chapter 1. Total fi-equency, cumulative
fi-equenciesof guilds, cumulative fi-equenciesof genera, and species frequencies were
calculated for each stand as in Chapter 1.
Genus richness, species richness, species frequency, guild richness and
cumulative frequency, cumulative frequency of individual genera and frequencies of
individual species were analyzed as unreplicated Model-III ANOVAs with three randomeffect levels (sites) and four fixed-effect treatment levels (stand age). Frequency data was
analysed as raw counts of the nurnber of occurrences, i.e. without being converted to
percentages. Moderate deviations fiom normality were seen in the residuals of most
variables but these were considered unimportant and ANOVA is robust enough to
accommodate these deviations from normality. Plots of residuals showed no strong
evidence of heteroscedasticity except for genus and species frequencies. The variances of
individual genera and species frequencies increased with increasing mean values so these
data were transformed, base 2, and reanalyzed. Stands that showed the largest residual
values are listed in Table 2.1.
The interaction mean squares in an unreplicated model-III ANOVA is used as the
denominator in F-tests. Since the site effect in this ANOVA is a random effect, its
expected variance is composed of the site variance plus
interaction variance, and the
site effect can be inte reted only if one assumes that an interaction between age and site
is either not possible or is unlikely (Sokal and Rohlf 1995). Interaction plots show that
there is some interaction between site and age (Figures 2.1,2.3,2.4) so the site effect was
not tested. Only age effects were tested because there was no replication within sites, site
effects could not be tested without assuming that there is no interaction between age and
site. Post-hoc comparisons of treatment means were made using Tukey's simultaneous
confidence intervals. A type-1 error rate of 0.10 was used as the cutoff for significance.
Genera and species were rated for their sensitivity to clearcutting in the following
categories: 1) highly sensitive, 2) sensitive 3) moderately sensitive, 4) non-sensitive nonspecific, 5) non-sensitive early succession, and 6) non-sensitive intermediate succession.
A t a o n was considered sensitive to clear-cutting if its frequency was significantly lower
on second growth stands than on old growth stands, and considered non-sensitive if its
frequency was unchanged or increased after clearcutting. Taxa that had significantly
lower frequency on the mature stands than on the old-growth stands were considered
highly sensitive. Taxa that had significantly lower frequency on the immature stands than
on the old-growth stands, and had no significant difference between the old growth and
mature stands were considered sensitive. Taxa that were significantly less frequent on the
regenerations stands than on the old growth stands and showed no significant difference
between the immature and old-growth stands were considered moderately sensitive. Taxa
that were not significantly less frequent on the regenerations stands than on the old
growth stands but were significantly less frequent on the regeneration stands than either
the immature or mature stands, and showed no significant difference between
immature and old-growth stands were also considered moderately sensitive. Taxa that
were more fiequent on the regeneration stands than on the old-growth stands were rated
non-sensitive early-successional. Taxa that were more frequent on the immature or
mature stands than on the old-growth stands were rated non-sensitive intermediatesuccessional. Taxa that did not change in frequency with stand age and had total
.
frequencies higher than 10% were considered non-sensitive non-age-selective. Taxa that
varied significantly in frequency with age were rated as potentially belonging to one of
the above categories according to trends at a=0.20 or complete absence from differently
aged stands.
Relationships between coarse woody debris (C
) descriptors (volume by size
class, volume by decay class) from Wells and Trofymow (1998) and wood-decay guild
richness were examined with scatter plots. The relationship between volume of CWD in
decay class 4 and number of wood-decay species was further characterized by linear
regression.
Similarity of macrofungal genus composition between al1 pairs of stand ages was
calculated using Jaccard's and Sorenson's coefficients of sirnilarity (
where :
Sj = Jaccard's coefficient of sirnilarity
S, = Sorenson's coefficient of sirnilarity
j
= the number of
genera present in both ages
a = the number of genera present in the first age
b = the number of genera present in the second age
cies
Sorenson's coefficient of similarity was calculated for al1 pairs of plots.
Hierarchical cluster analysis was performed on the resulting similarity matrix using the
"hclust" function in S-PLUS with "method = average".
Power analyses of both the ANOVA and post-hoc cornparisons were carried out
via simulation using programs written in SPLUS 4.01. Genus and species richness
ANOVA power was estimated using the SPLUS program "super.powwow" (Appendix
III) with 1000 iterations. This program repeatedly simulated data from a normally
distributed theoretical population, with a variance equal to the variance from actual data,
and treatment means equal to actual treatment means. One-way ANOVA was performed
for each simulated data set and power was calculated as the proportion of data sets in
which the nul1 hypothesis was rejected. Curves of ANOVA power versus number of
replicates were generated for three type-1 error rates, 0.05,O. 10, and 0.15 The curves were
smoothed using the smoothing spline option in SPLUS.
The power of the multiple comparisons was estimated using the SPLUS script file
"multi.powwow" (Appendix III) with 100 or 1000 iterations. This program is an
extension of the program "super.powwow" that conducts multiple comparisons on each
simulated data set. Computer memory limitations prevented me from combining these
two programs into one.
Curves of power of Tukey's post-hoc comparisons versus number of replicates
were generated for each comparison by running 100 iterations of "multi.powwow" and
changing the number of replicates in the simulated data sets. The curves were smoothed
using the smoothing spline option in SPLUS.
The change of power of the post-hoc comparisons of genus richness group means
with effect size was estimated using the SPLUS program "effect.size" (Appendix III) with
100 iterations of different effect sizes (variables e l , e2 and e3). This program is similar to
"multi.powwow" except that the group means of the simulated data are the three effect
sizes, el, e2, and e3, and zero. Two type-I error rates were used, 0.05 and 0.10. Data from
al1 runs of the program were pooled and curves of effect size versus power were plotted
and smoothed using the smoothing spline option in SPLUS.
The plot of detectable genus richness effect size versus number of replicates was
generated from the curves of the power of multiple comparisons versus number of
replicates (Figures 2.7 and 2.8) by interpolating (or extrapolating in one case) the number
of replicates required to achieve 90% power for each comparison. One point in the middle
was estimated using the program "effect.sizemto generate a power versus effects size
curve for 20 replicates and then interpolating the effect size with 90% power. The curve
was fitted with a line using the power function fit option in SPLUS.
To determine the usefulness of repeated monthly sampling on the same sites,
ess data from each month was also analyzed by ANOVA in the same manner as the
pooled data.
ni
The number of genera, number of species and total frequency of the macrofungal
community varied significantly with stand age (Table 2.2). The overall pattern was the
same regardless of whether nwnber of genera, number of species or total species
frequency vs. age was examined (Figure 2.1). Also, the patterns of genus richness, species
ess and total species Erequency for each chronosequence were very similar. The
regeneration age had significantly lower genus richness, species richness, and Erequency
than the three closed-canopy ages. No significant differences in macrofungal genus
richness and species richness and frequency existed amongst the immature, mature and
old-growth ages but the immature stands had the highest average richness and frequency,
closely followed by the old-growth and mature stands. The crossing of lines for each
chronosequence from one age to the next suggests that there is some interaction between
site and age. It is impossible to test for this interaction with an unreplicated design and
the apparent interaction may simply be due to normal variation. The old-growth age is
also more variable in genus richness and species richness than other ages.
Stands closer in age were more similar in macro
gal genus composition
(presence/absence of genera) than stands further apart in age (Table 2.3). There was a
trend of increasing similarity with increasing age. The regeneration age is most dissimilar
from the three older ages, which show a high similarity to each other (Sj > 0.62). The
immature stage shows the highest similarity to the regeneration stage and the old-growth
stage was the least similar to the regeneration stage. The mature and old-growth stands
show the highest similarity to each other.
Cluster analysis of the similarity of the species composition individual stands
shows that the regeneration stands are more similar to each other than to any of the
forested stands, and that the forested stands are more similar to each other than to the
regeneration stands with no definite patterns of similarity by site or age (Figure 2.2, Table
The area sampled for each age was adequate. For small (Le. inadequate) areas
sampled, a two-fold increase in sampling area means a greater than two-fold increase in
number of species found, based on interpolation of species area curves. A two-fold
increase in area sampled for each age (960 m2) in this study would mean only a 25%
increase in the nurnber of species for the immature and old-growth ages and even less for
mature and regeneration (Figure 2.3).
acrofun
Mycorrhizal guild richness and frequency varied significantly with stand age
(Table 2.2). The regeneration stands had significantly lower richness and frequency than
e older ages. The difference in mycorrhizal guild frequency between the
regeneration and mature ages was significant only at ~ 0 . 1 0No
. significant differences
existed among the closed-canopy stands. The mean mycorrhizal richness and fiequency
were highest on the old-growth stands (50.3 species, 970%) followed by the immature
stands (47 species, 945%) with a noticeable drop to the mature stands (37.7 species,
703%) (Figure 2.4a, 2.5a and Table 2.2).
Litter-decay guild richness varied significantlywith stand age but litter-decay
guild frequency did not (Table 2.2). The difference in richness between the regeneration
and immature ages was significant at a=0.10. The litter-decay fùngi do not show as
strong a difference in mean richness and frequency as the mycorrhizal fimgi between the
regeneration stands (15.7 species, 200%) and the closed-canopy stands (29.6 species,
407%). Mean litter-decay richness and frequency in the closed-canopy stands was highest
on the immature stands followed by the old-growth and mature stands (Figure 2.4b, 2.5b
and Table 2.2).
Richness and frequency of the general saprobes guild varied significantly with
stand age (Table 2.2). Regeneration stands had significantly fewer species than the
immature stands and old-growth. The frequency of the regeneration stands was
significantly lower than the fiequency in al1 three closed-canopy ages. General saprobes
showed the same general pattern as litter-decay fïmgi. Mean richness and fiequency were
highest on the immature stands ,followed by the old-growth, mature and regeneration
stands (Figure 2.4d, 2.5d Table 2.2). The regeneration stands have about one quarter the
frequency and one half the richness of the closed-canopy ages.
Wood-decay guild richness varied significantly with stand age but wood-decay
guild fiequency did not (Table 2.2). The old-growth and regeneration stands had
significantly fewer wood-decay species than the immature stands. No other significant
differences were found. The wood-decay guild richness and fiequency showed a different
pattern from the other guilds (Figures 2 . 4 ~and 2%). The immature stands had the
highest mean richness and frequency followed by the mature stands and the lowest mean
richness and fiequency were on the old-growth stands and regeneration stands. Richness
was not correlated to any of the following variables: total amount of coarse woody debris
(CWD), arnount of CWD in different size classes, amount CWD in decay class 1,2, or 5.
Richness showed a significant relationship with the amount of CWD in decay class 4
(linear regression, p=0.005), 56% of the variability in wood-decay guild richness was
explained by the amount of CWD in decay class four.
The overall patterns of mycorrhizal richness and total fiequency, and litter-decay
ess and frequency (Figures 2.4,2.5) were very similar to the patterns of total
richness and frequency (Figure 2.1), i.e., the regeneration stands had the lowest ric
or frequency and the closed-canopy stands had higher values that were similar to each
other.
The relative proportions by richness and frequency of the macrofùngal guilds
changed with stand age (Figure 2.6). The mean proportion of mycorrhizal species
increased with stand age. The mean proportion of wood-decay fùngi decreased with stand
age. The mean proportion of litter-decay fùngi was highest on the regeneration stands and
lowest on the closed-canopy stands. Mean proportions of general saprobes did not
change. Guild proportion data were not tested for statistical significance.
Eighteen genera had frequencies that varied significantly with age (Table 2.5).
Camarophyllus and Pholiota had significantly lower frequency on the mature stands than
the old-growth stands and were classified as highly sensitive although their frequencies
were not significantly lower than old growth levels in some of the younger stands.
Clavariadelphus was present only on old growth stands but was too infrequent to show
significant differences in the multiple comparisons and was classified as potentially
highly sensitive. Cortinarius, Hemimycena, Inocybe, Laccaria, Mycena, Nolanea,
Russula, and Tricholoma had significantly lower frequency on the regeneration stands
than al1 the closed canopy stands and were classified as moderately sensitive. Dermocybe,
Gomphidius, Hebeloma, Lactarius, and Strobilurus were al1 classified as moderately
sensitive, al1 had significantly lower frequencies on the regeneration stand than some of
the forested stands and had no significant differences in frequency between the immature
stands. Hydnellum was absent from the regeneration stands and present in
the forested stands but showed no significant differences between ages, and was classified
as potentially moderately sensitive. Psilocybe was non-sensitive early successional,
having a significantly higher frequency on the immature stands than the old-growth
stands. Clavulinopsis, Gymnopilus, Lycoperdon, Omphalina, and Suillus were very
frequent genera that showed no significant changes in frequency with stand age and were
classified as non-sensitive, non-age-selective. Many other genera showed non-significant
changes in frequency with stand age that might have been revealed as significant if more
chronosequences had been sarnpled (Appendix II).
Of the 302 species observed in 1997,64 species met the criteria necessary to
classify them into sensitivity categories (Table 2.6). The frequency of 58 species varied
significantly with stand age. More than half (37 species, 58%) of these 64 species were
sensitive to clearcutting, the remainder were not sensitive. Of the sensitive species, 11
species were highly or potentially sensitive, two species were sensitive or potentially
sensitive, and 24 species were moderately or potentially moderately sensitive. Of the nonsensitive species, five species were early-succession h g i , sixteen were intermediate
succession fmgi, and six species were non-sensitive non-age-selective. Mycorrhizal h g i
accounted for 70% of the sensitive species, 16% were litter-decomposers, 8% were
general saprobes and 5% were wood decomposers. For the non-sensitive species, only
22% were mycorrhizal, 37% were litter decomposers, 15% were general saprobes, 22%
were wood decomposers and one species (4%) was a basidiolichen.
78
Many other species showed non-significant trends which might have been
detected as significant if a larger number of chronosequences had been sampled. A total
of 48 species or species groups were found only in old-growth stands fiom 1995 to 1997
(Table 2.7). Appendix IV lists the complete species frequency ANOVA results and
fiequencies of al1 species on al1 stands in 1997.
The power of the ANOVA to detect the effect of stand age on the number of
genera of macrofungi present given the observed variance and effect size was about 65
percent (Figure 2.7). If the type-1 error rate is relaxed to 0.10 then the power increases to
about 80%. Power at a = 0.15 is 88%. If the nurnber of c onosequences is increased,
power increases and 90% power can be achieved at a
= 0.05 by
sampling five
chronosequences instead of three.
Power of the ANOVA for species richness (Figure 2.8.),given the observed
variance and effect size, was about 45 percent. Even if the type-1 error rate is relaxed to
0.15, power is still below 80%. To achieve 90% power at a = 0.05, six chronosequences
would have to be sampled.
2.3.7.2 Post-hoc comparisons
Power of post-hoc comparisons to detect the observed differences in macrofungal
genus richness between pairs of stand ages for three different alpha levels is shown in
Table 2.8. Power at al1 three alpha levels is very low for detecting differences among the
immature, mature and old-growth ages. Power for the regeneration stands versus
immature stands and the regeneration stands versus old-growth comparisons is below
0.80 at a = 0.05, above 0.80 at a = 0.10 and above 0.90 at a = 0.15. Power of the
regeneration stands vs. mature stands comparison is 0.521 at a = 0.05, and still below
0.80 at a = 0.15.
Power c w e s for the multiple comparisons ( a = 0.10) of genus richness involving
the regeneration stands are shown in Figure 2.9. Power increases with sarnple size. Power
greater than 0.90 can be achieved with four replicates for the regeneration versus
immature comparison, and with five replicates for both the regeneration versus mature
and the regeneration versus immature comparisons. This power analysis was done as a
paper for a statistics course at a time when the data had been analysed only to the genus
level. 1 decided that repeating this analysis with species level data was not necessary to
meet the requirements of my Master's degree.
Power of the comparison between immature and mature (Figure 2.10) is
extremely low with three replicates but increases to 0.80 with about 65 replicates and
0.90 with about 85 replicates. Power of the comparison between mature and old-growth is
almost zero with three replicates but increases to 80 percent with 90 replicates and, by
extrapolation, 90 percent power could be attained with 110 replicates - completely
impracticable, of course. Power of multiple comparisons for al1 variables d o m to guild
level is present in Table 2.8 but not discussed.
The curves of power versus effeet size (Figure 2.1 1) have a sigrnoidal shape and
power increases as the effect size (nurnber of genera) becomes larger. Power at a = 0.10
is 0.15 higher over much of the curve than that at a = 0.05 or altematively the lower a
can detect an effeet 3 to 3.5 genera smaller than the higher a.
The minimum detectable effect size is large with small numbers of replicates and
decreases quickly with increases in the number of replicates, but starts to plateau after
about twenty replicates (Figure 2.12). The formula of the fitted curve is
y = 50.91 * x(-0.557').
2.3.7.3 Evaluation of experimental design
en data Erom each month are analyzed separately for effects on genus richness,
September shows a significant age effect. October and November almost showed
significant age effects at a = 0.10. The number of genera peaks twice, once in the late
spring and again in the fall, with the fa11 season showing the highest richness for al1 sites
and ages (Figure 2.13). The variance in richness each month increases with increasing
richness. In al1 months, the regeneration age shows the lowest genus richness.
Discussion
Although 1 attempted to identifi al1 specirnens to species, this was possible for
only two-thirds of the fungi encountered, and the remaining third was identified to genus.
Thus, although the species results are more detailed and meaningfùl, they are potentially
less accurate. The sarnple size was small, with only three replicates of each age, so the
effect must be very large to be detected. The small sample size led to difficulties in
assessing whether the data met the assumptions of ANOVA, normality of residuals and
homogeneity of variances. There is no obvious biological reason why the residuals should
be non-normal, so the data was analyzed without a normalizing transformation. The nonnormality was caused by outliers. Victoria Watershed South mature stand and Koksilah
old-growth were the most consistent outliers.
Koksilah old-growth had lower richness and frequency than the other old-growth
stands. This plot always seemed drier and more open than al1 the other closed-canopy
stands. This plot had a southern exposure unlike the other old-growth stands, and there
was a nearby clearing for a logging road to the north of the plot and an approximately
1000 m2 clearing on a rocky outcropping that abutted ont0 the west side of the plot. These
factors may have combined to dry out the LFH layer that would othenvise have been
protected by the canopy. Fungi require moistwe in order to fniit, and the dryness could
explain the lower observed diversity. However, soi1 moisture data did not show that this
site was drier than others (Trofjmow pers. comm.).
82
The Koksilah old-growth also has a much higher cover of salal (Gaultheria
shallon) than the other stands in this chronosequence (Trofymow et al. 1997). Salal is an
extremely dense, broad-leaved, evergreen, ericaceous understory s
that may reduce
the measured macrofungal richness, as opposed to the actual richness. It does this in two
ways: 1) Salal makes it very difficult to spot macrofungi that fruit beneath it, leading to
lower measured richness; and 2) many macrofungi require light to fruit and the shading
by the salal may be enough to prevent them from fruiting. Salal litter is high in tannins
which may reduce litter decay guild richness as well.
VWS mature plot had a different site series than the other plots. It was a rich site
(site series 07), whereas al1 the other sites were poor sites (site series 01 and
02)(Tro@mow et al. 1997). The minera1 soil has the highest total N value, 2146 kgha,
almost double the value of the second highest plot, 1310 kg Nha. Nitrogen fertilization
has been s h o w to reduce ectomycorrhizal diversity (Menge and Grand 1978, Brandrud
1995). The VWS mature plot also has the lowest amount of nitrogen in the LFH.
Nitrogen in the LFH is in complex organic forms that ectomycorrhizal fùngi can make
available to trees, so trees in the other plots, with lower minera1 soil N and higher LFH
nitrogen, may rely more heavily on ectomycorrhizal fungi. Also, pools of water formed in
some parts of the plot. No macrofungi were observed fruiting undenvater so these pools
may have reduced fùngal diversity.
Regardless of the outliers, there was a very large and significant general decrease
in macrofungal diversity and frequency after clearcutting. The number of macro
dropped from a mean of 50 genera and 103 species found in the old-growth stands to a
mean of 27 genera and 41 species on ten-year-old regeneration stands. This is a decrease
of about 46% in the genus richness and 60% in species richness, and it is likely that an
even larger decrease occurred immediately after clearcutting and the regeneration stands
have partially recovered due to spore dispersa1 from nearby stands. The return to oldgrowth levels of macrofungal richness on the immature stands suggests that there are
factors that change drastically in the 10 to 20 years between the regeneration and
immature ages.
The decrease in richness immediately following clear-cutting can be attributed to
a number of causes (not necessarily in order of importance): disturbance and compaction
of the soil; the change in environmental characteristics of the site; the loss of input of
substrates required by saprobic fungi; and the removal of ectomycorrhizal trees. The first
two points are discussed below, and the last two points are discussed under the
appropriate guilds.
Disturbance and compaction of the soil caused by harvesting may kill fungal
colonies directly, or the changed soil conditions may no longer be optimal for the fungi
growing there. Amaranthus et al. (1996) showed that soil compaction resulted in
decreased mycorrhizal diversity associated with planted seedlings.
Clearcutting changes environmental conditions such as moisture, temperature and
illumination. The regeneration plots had a higher average daily temperature than the
forested sites in the summer but during the rest of the year al1 ages had similar
temperature regimes (TroQmow pers. comm.). Surface litter in the regeneration plots also
dried out more rapidly at the start of the summer (Trofjmow pers. comm.) Clear-cut sites
lose moisture from the surface litter more readily because there is no canopy to provide
shade fiom the drying effects of the Sun or shelter from the drying effects of the wind.
The canopy also moderates temperature; so clear-cut sites are more subject to extremes of
temperature due to increased solar radiation during the day and higher radiative cooling at
night. In the summer, this means not only a higher average temperature but also a more
exfieme temperature range - hotter in the day, colder at night. During the rest of the year,
cloudy weather reduces this effect. Temperature and moisture are important factors
controlling the growth of fungi, and different fungi have different minima, maxima and
optima (Cooke and Rayner 1984). Temperature and moisture were the dominant factors
affecting the distribution of Suillus sporocarps (Peredo et al. 1983). Fungi whose optima
were exceeded by the summer temperatures would be out-competed by hardier species in
unity, the net result being lower richness. y the immature age, the trees have
grown enough that the canopy has closed, allowing the less-hardy species to return.
It appears from the present study that overall ric
ss and frequency of the
macrofungal community does in fact return to old-growth levels by the immature age (38
to 49 years). The mature age has a non-significant, slightly lower richness than the
immature and old-growth ages. The return in fungal richness appears to be linked with the
re-establishment of a fünctioning forest ecosystem and the substrates and microhabitats it
provides. The immature and mature ages both had closed canopies and more resembled
forests than clear-cuts. This is good news, but it does not mean that macrofüngi are
invulnerable to clearcutting. The effects on the different macrofüngal guilds, individual
species and comunity composition must be considered.
ichness by itself is not a complete measure of the impacts of clearcutting on
macrofungal communities; the community composition must also be taken into
consideration because a community could change completely in its composition, but still
have the same richness.
The immature, mature and old-growth stages al1 showed a moderate similarity in
macrofùngal genus composition. Fifty-four genera were common to al1 three ages of the
closed-canopy stands. The regeneration stand was most dissimilar because it had lower
richness than the other stands as well as some unique genera. Even if al1 the genera in the
regeneration had been present in the immature stage, Jaccard's coefficiant of similarity
would still be only 0.62 . The immature stands are most similar to the regeneration stands.
e reason that the immature has the same richness as the old-growth is that its
fimgal community is in the process of changing over fiom one
ical of the clear-cut to
one typical of a forest, so it has higher richness because representatives of both clear-cut
fùngi and forest fungi are present. The drop in richness from the immature to the mature
age could be because some early succession fungi still present in the immature age had
disappeared by the mature age which does not yet, however, have the full complement of
late succession fùngi.
The macrofwigal community is characterized by having a few common taxa and
many rare taxa (Figure 2.3). These rare taxa explain why the similarity coefficients are
low. Rarity introduces observational error, since rare taxa recorded for one stand or age
may be absent on another because of their rarity or sporadic occurrence rather than their
inability to grow there. Sorenson's coefficient is not as sensitive to these rare
because it weights matches more heavily than mismatches and thus gives a higher
similarity than Jaccard's coefficient.
Presence or absence of a canopy appears to be a major factor in determining the
species composition of the macrofungal community. In the cluster analysis al1 the
regeneration stands, which lack a canopy, were grouped together (clade A), and al1 the
stands that have a canopy were grouped together (clade B) (Figure 2.2). In the closedcanopy stands, there was no major pattern of age or site relatedness, i.e., stands of
same age were not always the most similar, nor were stands in the same chronosequence
most similar. Other factors may explain the patterns of distribution of macrofùngi in the
closed-canopy sites but they do not appear to be age related. This is not to Say that there
are no age-related effects in the closed-canopy sites: they could be present, but masked by
the effects of other factors. Studying the relationships between the species composition
and other site factors with multivariate techniques would be useful and could increase our
understanding of the effects of forestry on macro
gi but this is beyond the scope of
present study.
The dissimilarity of the regeneration stands to the closed-canopy stands can be
attributed partly to their lower richness, Le., they lack forest mushroom species, and
partly to the presence of species characteristic of the clear-cuts such as Psilocybe
montana and Rickenellafibula. The Watershed South regeneration stand is less similar to
the other regeneration stands because it has a higher richness, meaning it has more
species not found in the other two sites. This higher richness is due to the presence of
some forest-associated fùngi such as Trichaptum abietinum and Mycenapura. In the
closed-canopy stands, Koksilah old-gro
and Watershed South mature plots are seen to
have species compositions less similar to the rest of the closed-canopy plots. These two
plots were also consistently identified as outliers in terms of their overall richness and
fi-equency, and the richness and frequency of the major fimgal guilds. These plots have
some edaphic characteristics that differentiate them from the other plots and each other.
Koksilah old-growth is more open, and drier than the other closed-canopy sites.
Watershed South mature plot is a much more nutrient rich site, with a flatter slope and
less well drained. These characteristics were discussed in detail under the section on
outliers.
.3.1 Mycorrhizal guild
The mycorrhizal guild was the guild most severely affected by clearcutting in
terms of nchness and frequency. The regeneration stands have only one-eighth of the
species richness of the old-growth. The immature age has 88% of the nurnber of
mycorrhizal species found in the old-growth, while the mature age has only 70% of that
number, although the last two differences were not signifiant.
Goodman (1995) studied ectomycorrhizal root-tips on two of the plots used in this
study, Koksilah mature and old-growth plots. During two years, he took 36 soi1 cores
fi-om each site. a total area of 0.0706 m2, and found 35 ectomycorrhizal morphotypes on
the mature stand and 36 morphotypes on the old-growth. By contrast, in this study 320 m2
per stand was sampled in 1997 and sporocarps of 50 ectomycorrhizal species were found
in the mature stand versus 42 species in the old-growth stand. Goodman (1995) studied
only these two plots plus two others for 2 years while 1 studied these two plots plus 10
others for 1 year and looked at litter- and wood-decay fungi as well as mycorrhizal
Therefore, a much smaller above-ground sampling effort detected more species than
below-ground sampling, thus, although production of sporocarps can be sporadic and may
not always be related to below-ground biomass (Gardes and Brun 1996), sporocarp
surveys are still an efficient way to examine species richness and community
composition.
Many ectomycorrhizal fungi are obligate symbionts and cannot survive without
their host trees, although a few may find alternate hosts in ericaceous shnibs (Perry et al.
1987). Facultative ectomycorrhizal fungi can survive without their host but often with
reduced vigour and they may be out-competed by more vigorous saprobes. Harvey et al.
(1980) found that almost al1 ectomycorrhizae are dead one year after clearcutting without
replanting. However, spores of ectomycorrhizal fungi may persist in the soil for at least
two years (Miller et al. 1994). Some mycorrhizal fmgi produce sclerotia which may
persist in the soil for some time before germinating, but the longevity of sclerotia
(Bmdrett and Abbot 1999, and even which fungi can produce them, have not been well
studied. The major ectomycorrhizal genera, Cortinarius,Lactarius, and Russula, were
completely absent from the regeneration stands and it is highly unlikely, though not
impossible, that their occurrence in the immature stands is due to the presence of sclerotia
capable of surviving for 30 to 40 years. The re
of ectomycorrhizal diversity, and
macrofungal diversity in general, by the immature age must rather be considered as being
due primarily to re-colonization by airborne spores, with other dispersa1 methods playing
a secondary role.
It has been shown that ectomycorrhizal fungi exhibit succession (Dighton et al.
1986, Visser 1995) so the drop in richness after clearcutting is due to the lack of latesuccession genera. Although late-succession species are probably physically able to return
by spore dispersa1 they are unable to establish themselves as syrnbionts of the saplings on
the regeneration stands since they are adapted for symbiosis with mature trees.
Late-succession ectomycorrhizal fungi must re-colonize when the new stand is
mature because they are probably eradicated by clearcutting. Re-colonization by late
succession ectomycorrhizal fungi is almost certainly dependent on a source of airborne
spores from an old-growth forest nearby, rather than the persistence of propagules in the
soil. If there is no old-growth nearby, re-colonization by airborne spores will probably be
much less effective - the log of number of fùngal infections by airborne spores decreases
with the log of distance from the spore source (Lacey 1996) - and late successional stage
species may not retum. Of course, this would be a moot point if e second-growth stands
are clear-cut before they reach the old-growth stage. Aerial dispersa1 of spores may not be
as effective in the rainy Pacific northwest as it is in other areas because spores are washed
out of the air during rainfall at an exponential rate (Lacey 1996). Thus, proximity of oldgrowth refugia is probably of even greater importance than it would be in a drier climate.
Further research is needed in this area and could have important implications at the
landscape level for forest management.
2.4.3.2 Litter-decay guild
Many saprobic fungi live in the forest floor litter. While loss of e trees does not
create an immediate loss of habitat, the litter layer is no longer being replenished by
90
continued litter fàll from the forest canopy, so species that are primary colonizers of the
litter disappear. Although there is fiesh litter fiom understory plants such as salal, many
fùngi are substrate-specific and need conifer needles or cones. Fungi that specialize in
larger litter, such as twigs and branches, do not seem to suffer from the loss of the input
of fi-esh litter because much slash is generated from clearcutting and the coarser material
takes longer to break d o m . Fungi that break down coarse woody debris may also initially
benefit from the slash created by clearcutting.
Saprobic fùngi in general do not seem to be as adversely affected as mycorrhizal
fungi by clearcutting. The number of non-wood-decay saprobic species on the
regeneration age is 68% of that found in old-growth, while the immature and mature
show an increase over old-growth levels at 129% and 104% of old-growth numbers. The
litter-decay guild proper was less sensitive than the "other saprobes" guild. 1 would have
suspected the reverse since the other saprobes use a wider range of substrates. Perhaps the
generalist fungi are out-competed by the litter specialists for the more limited resources
available on the regeneration stands.
The arnount of litterfall on these sites was greatest in the old-growth, while the
immature and mature stands had significantly lower litterfall, roughly 70-80% of the oldgrowth level (Trofjmow 1998). The higher litterfall in old-growth stands did not
correspond to higher
gal richness or frequency. Litterfall on the regeneration age was
not measured because it lacks a canopy, but litterfall increases with stand age and levels
off upon canopy closure (Trofymow et al. 1991). Litterfall on the regeneration stage may
have been less than one-tenth of old-growth levels.
ood-decay guild
The frequency of wood-decay fungi was not significantly affected by stand age but
their richness was affected, showing that the fewer species on the stands with lower
richness had higher individual frequencies. It was unexpected that the highest richness of
wood-decay fungi would be found in the immature age. The expected relationship would
be highest in the old-growth and regeneration stands and lowest in the immature and
mature stands if the volume of coarse woody debris (CWD) followed the theoretical Ushaped curve of Spies and Franklin (1988). However, CWD volumes were low on these
sites and no overall trends were found (Wells and Trofjmow 1998). It was suggested by
these authors that CWD was low because the sites were low-productivity
area, but these sites had to be chosen because of a paucity of productive old-growth sites.
Also, al1 the second growth sites have been subject to burning, either post-harvest
broadcast burning, piling and bwning, or salvage logging after wildfire (Trofj-mow pers.
comrn.). Levels of C
in the old-growth could also be low because of non-stand-
destroying fire.
No relationship was found between total volume of CWD and wood-decay guild
richness, but a strong positive correlation was seen between volume of CWD decay class
4 and wood-decay guild richness. This agrees with Renvall(1995) who found that the
richness of wood-decay fungi was highest on decay class 4 and second highest on decay
class 3.
Wood-decay fungi may not have been adequately sampled. Unlike litter-decay and
mycorrhizal species whose substrate is more or less uniforrnly distributed at the scale of
this study, CWD is very patchily distributed and systematic sampling using CWD as
sampling units would have obtained better results. Only three of the 40 most frequent
fungi were wood-decay fungi, suggesting that this group was not adequately sampled.
2.4.3.4 Guild proportions and stand age
The relative proportions of the guilds change with stand age because the different
guilds are affected differently or to different extents by stand age. The
ages show nearly identical proportions of the guilds, indicating that the macrofungal
communities of ecosystems in these ages are functioning similarly. The macrofungal
community in the forested stands is roughly 50% decomposers and 50% mycorrhizal
though the mycorrhizal guild is slightly higher in the old-growth stands.
sarne 5050 ratio was reported for balsam-firlpaper birch and black spruce forest in
Quebec (Villeneuve et al. 1988) and Pinus sylvestris forests in Nonvay (Sastad 1995) and
may be typical of forests dominated by ectomycorrhizal trees. In contrast, in the
regeneration stands the macrofungal community is weighted predominantly toward
decomposers, less than 20% of the macrofungi were mycorrhizal.
Three genera were found to be highly sensitive to clearcutting: Camarophyllus,
Clavariadelphus and Pholiota. Clavariadelphus was found only in old-growth but the
differences between stand ages were not significant because of its very low frequency.
Four different species of Camarophyllus were seen in 1997. The frequency of al1 species
was very low and al1 were seen on either the regeneration or immature age so it is hard to
Say whether this genus really is highly sensitive or not. Pholiota also had very low
frequencies. Two species of Pholiota were found in 1997, Pholiota terrestris was found
only on the regeneration age so this species cannot be considered sensitive at al1 and
Pholiota
was primarily on the old-growth stands. This illustrates the inaccuracy
looking for patterns in whole genera which are composed of many species that may be
affected differently by clearcutting.
Only four of the 14 genera that were moderately sensitive were saprobic, the rest
were mycorrhizal, whereas of the 1 1 non-sensitive genera (ne, ni and nn) only Suillus was
mycorrhizal. Suillus species may be facultative saprobes (Bonello et al. 1998) and this
may be why Suillus is one of the few mycorrhizal genera on the regeneration stands,
possibly having survived clearcutting.
ighly sensitive species
The majority of the highiy sensitive species were mycorrhizal, showing that this
guild is the most sensitive to clearcutting. Russula spp. and Russula xerampelina were
significantly less frequent in the mature stands than the old-growth, unlike the
ectomycorrhizal guild as whole or the genus Russula as a whole, both of which were
moderately sensitive. This shows the importance of species-level studies in revealing
details that would be missed in a more general analysis. Russula spp. is a lurnped group
of many species. Its odd pattern of attaining a level of diversity equivalent to old-growth
levels by the immature age and then dropping significantly in diversity by the mature age
may be due to two component species or groups, one that is non-sensitive intermediate
successional and one that is highly sensitive. It could also be argued that this group is
moderately sensitive because its fiequency returns to old gro
levels by the immature
age. The value of this result and the results of other multi-taxa groups is limited except to
Say there are some unidentified species that are affected by clearcutting. Russula
xerampelina is unequivocally highly sensitive. Its frequency was higher in old-growth
stands than on al1 younger stands, although is was not significant on the immature
stands. Because of the reduced populations on immature and mature stands, this species
will not be extirpated fiom this habitat imrnediately, but may be extirpated with
subsequent rotations if no adjacent old-growth stands are available for recolonization.
These Russula species/groups recover to old growth levels by the immature age
and then decline again in the mature which could indicate that there are some effects of
clearcutting that take 80 - 90 years to be manifested. The litter decay fungus Clitocybe
deceptiva follows this same trend in fiequency with age. The richness and fiequency of
macrofungal community, mycorrhizal guild, litter-decay guild and general saprobes guild
also followed this trend, although the drop in values in the mature stands was not
significant. One possible explanation for this trend (if it is real) is the intermediate
disturbance hypothesis (Conne1 1978). This would imply that it takes somewhere between
50 and 80 years after the disturbance of clearcutting for the macrofungal community to
reach an equilibriurn state when competitive exclusion reduces diversity. The high
diversity in old-growth stands may be maintained by their higher structural diversity and
by periodic disturbances such as ground fire or wind-throw.
Agaricus sp. 1, Gomphidius smithii, Inocybe lacera vel afl, Russula sp. 1,
Russulafragilis vel a#, Tricholoma atroviolaceum vel a 8 and Tricholoma sulphureum
al1 had low frequencies, less significant age effects, and differences in their frequency
between the old growth and other stands were not significant. They were classified as
potentially highly sensitive because they showed trends at p = 0.2. These species warrant
her investigation and may turn out to be old growth indicators.
were highly sensitive.
2.4.5.2 Sensitive species
Russulu brevipes was one of the most severely affected common species (rated
sensitive). Its frequency was highest on the old-growth stands, half of old-growth levels
on the mature stand, one-tenth of old-growth levels on the immature stand, and it was
absent fkom the regeneration stands. Only the mature s
different from the old-growth.
There are many possible reasons why R. brevipes failed to reach old-growth
levels in the second-growth stands. Spore dispersa1 could be a limiting factor. Sporocarps
of R. brevipes start growth beneath the duff and push up a large patch of it as they mature.
Sometimes they fail to completely emerge from the duff, or they emerge long after they
have started dropping spores. Because of this, a large proportion of their spores may fa11
to the substrate immediately beneath them and fail to reach the air colurnn, where they
can be effectively dispersed. Many members of Family Russulaceae have become
sequestrate, i.e. their spore bearing layer or hymenium is never openly exposed or even
remains subterranean (Kendrick 1994). It is possible that Russula brevipes is just starting
d o m the evolutionary path to becoming sequestrate.
Russulu brevipes produces some of the most massive sporocarps of al1 Pacifie
st mycorrhizal fùngi and may place a carbohydrate demand on its hosts that only
an old-growth forest can sustain. Fungi can be very long-lived and their mycelia can be
massive; clones of Armillaria bulbosa can live for 1,500 years, occupy 15 hectares, and
weigh 10,000 kg (Smith et al. 1992).Russula brevipes could have a long-lived mycelium
that relies on somatic growth to slowly out-compete other fungi. Somatic growth is
probably an important way in which mycorrhizal fungi colonize new root-tips. Ninety
years may simply not be enough time for Russula brevipes to establish itself as a
dominant mycorrhizal fungus and attain a colony size large enough to produce its massive
sporocarps. This would be a good organism to study in more detail.
Because of the difficulty of identifying Ramaria, al1 species were lumped under
the genus narne. The genus was considered sensitive to clearcutting because it tended to
have lower frequency on the immature and regeneration stands than the old growth
stands. Ramaria spp. were absent from the regeneration and immature stands in 1997, but
were found a few times on the immature stands in 1995 and 1996. Further investigation
may reveal that this genus contains some highly sensitive species. Some species of
Ramaria, like Russula brevipes, produce very large sporocarps, and they may take a long
time to build up sufficient biomass to fmit.
2.4.5.3 Moderately sensitive species
Many mycorrhizal fungi were moderately sensitive to clearcutting. Cortinarius
acutus vel a ! , Cortinarius obtusus vel a!,
Cortinarius sg. telarnonia spp., Inocybe
ovatocystis, Inocybe spp., Laccaria bicolor, Laccaria laccata, Russula placita, Lactarius
rubrilacteus, and Tricholorna spp. were significantly less frequent on the regeneration
stands than the old growth and had no significant differences in frequency between the
old growth and immature stands. Cortinariusfulvescens vel a#, Hebeloma
97
crustuliniforme, and Inocybe geophylla were classified as moderately sensitive, they had
significantly higher frequency in the immature or mature stands than the regeneration
stands and showed trends of lower fi-equencyin regeneration than old-growth stands.
Tricholoma sejunctum and TricholomaJlavovirens were classified as moderately
sensitive because they showed no significant difference in frequency between the mature
and old-growth stands and were significantly less frequent on the regeneration stands than
the mature stands. Cortinarius vibratilis was classified as potentially moderately sensitive
because it tended to have lower frequency on the regeneration stands than the forested
stands.
Mycorrhizal fungi have been shown to exhibit succession with the advancing
maturity of their host tree (Dighton et al. 1986, Visser 1995) and this succession occurs
much more rapidly in young ages. Cortinarius spp., Lactarius spp., Russula spp., and
Tricholoma spp., have been identified as late stage fungi in pine forests (Visser 1995).
Mycorrhizal species that were moderately sensitive can be viewed as late-stage fungi
because they were present on the old-growth. Although they first return at the immature
age, they persist into the mature age, and can be expected on the old second-growth. The
dominant factor affecting the return of moderately sensitive, sensitive and highly sensitive
mycorrhizal species may be the maturity of the host trees, although effects of canopy
closure on temperature and moisture regimes may also be important.
The litter-decay
i Galerina emetensis vel afi, Hemimycena delectabilis, and
Otidia alutacea vel afl were moderately sensitive to clearcutting. These species depend
on the input of fresh litter fiom the canopy. Strobilurus trullisatus grows exclusively on
Douglas-fir cones and its return is limited by the return of cone production. The other
species grow in conifer needle litter.
en this input is eliminated by clearcutting, these
species disappear either as the litter decays or when they are out-competed by other litterdecomposing fungi that are better adapted to the moisture and temperature fluctuations
experienced in the clearcut. These fungi may
when the regenerating second-growth
stands start producing enough litter or they may not be able to return until the canopy
closes and begins to moderate soi1 temperature and moisture fluctuations.
Three general saprobes were classified as moderately sensitive. One of these
is a species group, Mycena spp., which was classified as a general saprobe because it
contains litter-decay and wood-decay species. Nolanea holoconiota typically grows on
litter but may also be found on well-decayed wood, and Xeromphalina fulvipes grows
predominantly on fine woody debris but is also found on coarse woody debris and litter.
of these species to older stands can be attributed to increased litter-fa11 and the
moderation of temperature and moisture by a closed canopy.
Only one wood-decay fungus was found to be moderately sensitive to clearcutting.
Pseudohydnurn gelatinosum grows on a wide range of sizes and decay classes of woody
substrates, often partially or completely buried. It was not found on the clear-cut,
probably because it could not survive the extreme temperature and moisture fluctuations.
The increased input of fallen branches from the canopy was probably also important for
its return.
The mature stands in this study developed in a landscape with a higher proportion
forests than exists today, and thus a larger source of late successional
fungal spores. The immature stands have developed in a landscape with a much smaller
proportion of old-growth, and the regeneration s
ds are developing in a landscape with
minimal old growth. By the time their canopies close there may be no old-growth left to
serve as sources of spores of highly-sensitive species. Species that recover from decreased
frequency after clear-cutting in this study may not have done so in a landscape with less
old-growth. Also, as the proportion of old-growth in a landscape decreases, the average
distance from a regenerating stand to the nearest old-growth stand increases. About 90%
of aerial spores are deposited within the first 100 m from their source (Lacey 1996), so
not only is there a smaller source of late-successional specific fungal spores but the
source is farther away and much less effective in providing inoculum. One study found
that ectomycorrhizal diversity on naturally regenerated hemlock seedlings decreased with
increasing distance from the old-growth forest edge in 50 to 75m openings and that the
decrease may be partly due to unsuccessful spore dispersa1 (Kranabetter and Wylie 1998).
on-sensitive early-successional species
One litter-decay fungus, Psilocybe montana, had a significantly higher frequency
in regeneration stands than old-growth stands. Rickenellafibula and Omphalina
chlorocyanea showed the same trend but the difference was not significant. R. Jibula and
P. montana were ofien found growing together, likely occupying the same niche, and are
good examples of early successional species adapted to the conditions on a clearcut.
Psilocybe montana and RickenellaJibula are found on mossy outcrops in forest clearings
and are probably saprobic on moss, so they do not require litter-fa11 from the canopy.
Clearcutting increases the habitat available to them. By comparison, it should be noted
that no mycorrhizal species had significantly higher frequency on regeneration stands
than old-growth stands.
1O0
The trends in frequency of the wood-decay fungus Hypholoma fasciculare
indicate that it is potentially an early successional species. Hypholoma fasciculare is
especially combative and excellent at secondary resource capture, Le. actively attacking
and replacing other
gi in woody substrates (Cooke and Rayner 1984). The sub-
optimal growth conditions on the clearcuts may cause fungi that would normally be able
to defend their resources to succumb to its attacks.
on-sensitive intermediate-successional species
Three mycorrhizal species, Dermocybe croceifolia and Hebelorna rnesophaeum
vel afl, and Coltriciaperennis were significantly more frequent on immature stands than
on old growth stands and classified as non-sensitive intermediate-succesional species.
The mycorrhizal species Dermocybe semisanguinea, Hygrophorus bakerensis, and
Phellodon melaleucas were potentially non-sensitive early successional species. These six
fungi may be ruderal species that require older trees (not saplings), are good at dispersing
to new areas but are not very competitive. Their preference for the immature stands can
be attributed to the phenornenon of mycorrhizal succession. Coltricia perennis can form
arbutoid mycorrhizae with ericaceous plants as well as ectomycorrhizae with conifers
(Danielson 1983) and may have been able to persist after clearcutting with an ericaceous
host.
Two litter-decay species groups, Mycena sp. 1 and Mycena sp. S., were more
frequent on the immature stands than the old-growth stands. This difference was
significant for Mycena sp. 2 but not Mycena sp. 1. Both of these taxa are species groups
or aggregates, Mycena sp. 1 is possibly two different species that both smell like raw
cucumber, and Mycena sp. 2 is a group of species that macroscopically fit Smith's (1947)
concept of the infrageneric section diminutivae. Both these groups grow on needle beds
so their return is probably due to the increased litter-fall. Their return by the immature age
and subsequent decline may be because these groups are able to colonize stands after
clearcutting but are less competitive than other litter decay fungi and are slowly
eliminated.
Two general saprobes, Galerina badipes vel a f and Marasmius spp., were nonsensitive intermediate-succession fungi. Galerina badipes vel afJ: was a group of
Galerina species that had black or dark-brown stipe bases, resembling the true G.
badipes. As suggested above, these species may have good dispersa1 and survival
characteristics but may be less competitive than other fungi.
on-sensitive non-age-specific s
Six species with a total fiequency greater an 10% showed an even distribution
arnong the different stand ages, and were classified as non-sensitive non-age-specific.
These were the litter-decay fun@Clavulinopsisfusiformis, Marasmius salalis, and
Mycena pura, the general saprobe Mycena alcalina, the wood-decay fungus
Guepiniopsis alpina and the basidiolichen Omphalina ericetorum. Marasmius salalis
grows almost exclusively on the leaves of sala1 (Gaultheria shallon) and Oregon grape
(Mahonia nervosa). These plants are a ubiquitous ground cover in stands of al1 ages, so it
is not surprising that Marasmius salalis is likewise found in al1 ages and is not sensitive
to clearcutting. The fact that Clavulinopsisfusiformis, Mycenapura, M. rorida and M.
alcalina are not affected by clear-cutting indicates that they âre vigorous competitors,
able to withstand the conditions on the regeneration stands and out-compete other fungi
for the available resources. Guepiniopsis aZpinus was the most Kequently encountered
single species on these chronosequences. 1have seen it fruiting on woody debris of
almost al1 size and decay classes. It is not s
rising that this versatile fungus is
unaffected by clearcutting. The lichenized Omphalina ericetorum obtains its energy from
the photosythates of its algal partner so it is energetically independent from the trees and
may benefit fiom the temperature and moisture changes on the regeneration plots.
2.4.5.7 Implications of sensitivity ratings
Species that were highly sensitive to clearcutting were old-growth selective
species that did not re-colonize to old-growth levels by the mature age (83 -105 years).
The re-colonization of highly sensitive species may be controlled by factors other than
those brought about by canopy closure with its resultant temperature and moisture
moderation and increased litterfall input. There could be limitations to dispersal, or other
site conditions may be unsuitable for their growth, e.g. it is possible that later-succession
fungi cannot establish mycorrhizae with younger trees (Kranabetter and Wylie 1998).
Reductions or eliminations of older sera1 stages or old-growth forests could mean that
highly sensitive species may be extirpated or have their frequencies significantly reduced.
Sensitive species were found only in, or are most abundant in, old-growth or
mature stands. These species take more than 40 years, and up to 90 years, to re-colonize
after clearcutting and could be lost with successive rotations or shorter rotation ages. Like
highly sensitive species, their return is likely controlled by factors beyond canopy
closure, soi1 temperature and moisture, and increased litterfall.
Moderately sensitive species were found on old-growth, mature and immature
stands but were absent or rare on regeneration stands. These species take 25 to 40 years to
re-colonize after clearcutting. Their return coincides with, and is probably linked to,
canopy closure and its resultant changes in soi1 moisture, temperature and litterfall,
although other factors may also be important.
The three classes of sensitive fungi can be considered late succession fùngi
because they do not grow in the early stages of forest succession and are present in oldgrowth. These sensitive fungi may require special management if their populations are to
be maintained. Kropp and Albee (1996) found that a selective cut which reduced an evenage lodgepole pine stand from 840 to 395 stemsfha, maintained 73% of the richness of the
undisturbed stands, while clear-cuts retained only 15%. Further research is needed to
examine the effects of different variable retention silviculture practices on
macrofungi in CWHxm Douglas-fir forests.
Non-sensitive early-succession fungi could be said to benefit fiom clearcutting
because their fiequency in the regeneration age has increased over old-growth levels.
These species are either better competitors in the more extreme temperature and moisture
regime or simply have better dispersa1 abilities and were the first to re-colonize. Also,
some of these species may have been present in the old-growth forest and survived after
clearcutting.
Non-sensitive intermediate-succession species also could be said to benefit from
clearcutting, since their fiequencies are higher in the immature andlor mature stands than
they are in the old-growth stands and
ey may be present in the regeneration stands. The
fact that there are intermediate-successionfungi shows that although overall species
richness and frequency of the immature and mature stands are equivalent to old-growth
values, al1 of their site characteristics are not.
Non-sensitive non-specific species were equally present on al1 age classes, Le.
clearcutting did not affect their overall frequency. Fungi that belong to one of the above
three non-sensitive classes do not require any special management to maintain their
populations.
2.4.5.8 Species not rated for sensitivity to clearcutting
Many of the species that did not show significant effects still warrant discussion.
Tricholoma magnivelare, the pine mushroom, and Cantharellusformosus, the yellow
chanterelle, are two of the most economically important wild mushrooms harvested
primarily for export to foreign markets (Re ead et al. 1997). Neither of these species
was found in sufficient quantity to determine their response to clear-cutting. Both these
species are mycorrhizal and probably have to re-establish themselves in second-growth
stands via aerial spores. Cantharellusformosus was reported only on the old-growth
stands but 1have also found it on irnrnatue second growth Douglas-fir stands outside of
the study plots, so it is probably moderately sensitive to clearcutting. Tricholoma
magnivelare was found only on the mature stands but mushroom stwnps that smelled like
T. magnivelare and mushroom pickers with guns were found on the old-growth stands.
The armed mushroom pickers, who were quite friendly when they found out 1 wasn't a
rival picker, informed me that they each had picked $1000 worth of pine mushrooms
(about 45 kg) the day before in a nearby old-growth Douglas-fir stand. This anecdotal
evidence indicates that old-growth Douglas-fir forests may be important pine mushroom
habitat.
The large proportion of wood-decay fùngi that are not affected by clearcutting
argues that there is some mechanism helping them survive. It is possible that wood-decay
fungi are simply more robust than other fungi, that their substrate moderates the
temperature and moisture fluctuations of fresh clearcuts, or simply that wood takes a long
time to decay so wood-decay fungi do not run out of resources.
Power of the ANOVA testing the null hypothesis that stand age has no effect on
genus richness was low at 66 percent as estimated using the program "super.powwow"
in SPLUS (Appendix A) with a = 0.05. If the type-1 error rate is relaxed to 0.10 then the
power increases to a more acceptable 82%. Power at a = 0.15 is 93%. A type4 error in
this hypothesis test would lead one to falsely assume that clearcutting has a significant
effect on macrofùngal richness. A type-II error, failing to reject a false null hypothesis,
would lead one to erroneously conclude that clearcutting has no effect on macrofùngal
diversity. From a perspective of minimizing the potential loss of macrofungal diversity, a
type-II error is more dangerous than a type-1 error. From the results of the power analysis
it is clear that the less stringent type-1 error rate of 0.10 should be used for this study to
reduce the chance of a type-II error. In order to increase the power of the ANOVA to
90%, the number of chronosequences observed must be increased to four with a =O. 10 or
six with a = 0.05. It may even be preferable, given the consequences, to set type-1 error
rate so that it produces an equal type-II error rate.
ower of multiple comparisons
The power of the multiple comparisons was low at a = 0.05 . Even
comparisons between the old-growth and regeneration, and between immature and
regeneration, which had differences between their means of 23 and 24 genera
respectively, were below 80% power with a = 0.05. As expected, the comparisons with
the largest differences had the greatest power. Power can be increased by relaxing the
type-1 error rate to 0.10. The power for the comparisons of old-growth and regeneration,
and immature and regeneration exceeds 80% at a = 0.10, but is still less than the
preferred 90%. A larger number of replicates is required. Power of
regeneration comparison is increased from 52% to 66% when the alpha level is reIaxed to
0.10. There is no significant difference between the mature and the regeneration age at a
= 0.05 but
at a = 0.10 the mature age is significantly richer. There is insufficient power to
detect the difference between the mature and old-growth, and mature and immature ages.
Very large sample sizes of 80 to 110 chronosequences would be required to detect this
effect. Such intensive sampling would simply be too time consurning with whole
chronosequences. It may be possible if only mature vs. old growth comparisons were
made and more personnel were involved.
The minimum detectable effect size with three chronosequences is quite large
(Figure 2.12). If 90% power is desired then only a difference of 29 genera (a = 0.05) or
26 genera (a = 0.20) can be reliably detected. This is larger than the difference in genus
richness between the most diverse and the least diverse ages in the present study. Even if
power of 50% can be accepted then the minimum detectable effect size is still large, 18
genera (a = 0.05) or 15 genera (a = 0.10). A decrease in richness of 18 genera is
unquestionably highly biologically significant, and if fùrther studies were to be carried
out they should be redesigned to increase the power and decrease the minimum detectable
effect size.
Detectable effect size can be minimized by increasing the nurnber of
chronosequences sarnpled. Detectable effect size starts off large and decreases quickly at
first with small increases in numbers of replicates, but then levels off. The addition of a
few more chronosequences would have a large effect on the minimum detectable effect
size. There are five chronosequences in the CWHxrn biogeoclimatic subzone and if al1
five were sampled an effect of about 20 genera, as opposed to 30 with three replicates,
could be detected. If ten chronosequences were sampled
n an effect of 15 genera could
be detected.
The detectable effect size and the power of the analyses are limited by the high
variability of the macrofungal genus richness fiom site to site. A difference of 15 genera
is very biologically significant and future studies should be designed to recognize the
significance of smaller effects.
These statistical problems do not mean that the effects observed during the present
study are not real, but a more powerfùl experimental design is necessary to prove this. If
the difference in number of mycorrhizal species between the mature and old-growth
stands is real, then mycorrhizal diversity will decline with subsequent rotations. Thus it is
important that a more in-depth study be conducted.
sefulness of repeated monthly sam
ANOVAs of the effect of stand age on genus richness for each month from March
to November show that only one month, September, shows a significant effect. October
and November also show probabilities that are close to significant at a = 0.10 . The others
are al1 type-II errors due to low power and sampling at the wrong time of year.
oughout the year the same pattern of richness was observed, the regeneration
stands were consistently poorer and the closed-canopy stands consistently richer (Figure
2.13). Of the three closed canopy stand ages, the immature stands had the highest mean
richness for six months out of nine, were tied for the second lowest richness for two
months and had the lowest richness in one month. The old-gro
had the highest
richness one month, the second highest richness for seven months out of nine and the
ess in one month. The mature stands had the highest ric ess in two months,
e second lowest rie ess in one month and the lowest richness for five months. The
immature age had the highest richness in the sumrner and fall, but the lowest in the
spring, suggesting that spring-fhiting and fall-fniiting fùngi may be affected differently
by forestry practices.
A more efficient use of sampling effort would have been to have sampled
different sites each month rather than visiting the same sites repeatedly.
data on one site does give a stronger signal for that site, the signal of the effect of
clearcutting can be seen with a month's data and although
s gives a weaker signal on
each site, the increased sample size should more than compensate for this.
The drawback of not visiting the same sites throughout the year is that the
knowledge of community composition for a site would be incomplete. If only three
chronosequences could be sarnpled each month, a compromise would be to visit one
chronosequence repeatedly and two new ones each month.
110
le 2.1 Plots that had the largest ANOVA residuals. g = regeneration, i= immature,
m= mature O=old-growth, s=watershed south, n =watershed north, k= Koksilah, + or indicate whether the value was higher or lower than expected.
ANOVA test
genus
richness
species richness
species frequency
mycorrhizal richness
wood-decay richness
litter-decay richness
other saprobes richness
non-wood-decay richness
mycorrhizal frequency
wood-decay frequency
litter-decay fiequency
other saprobes frequency
non-wood-decay frequency
ms-, os+, okik+, ms-, okms-, okgs+, ms-, okin-, ik+, mkms-, os+, okin-, ik+, okik+, ms-, okgs+, ms-, mk
ik+, okms-, mk+
ik+, okms-, ok-
le 2.2 Summary of ANOVA of age effects on macrofungal genus richness, species
richness and frequency, and species richness and frequency of macrofungal guilds with
Tukey's post-hoc comparisons of al1 pairs of stand ages. Boldface indicates significance
at a=0.05 level, underlining indicates significance at a=0.10 level. g = regeneration, i=
immature, m= mature O=old-growth
Comparisons
Test
ANOVA g-i
P
genus richness
total species richness
total fiequency
mycorrhizal richness
mycorrhizal frequency
wood-decay richness
wood-decay frequency
litter-decay richness
litter-decay fiequency
general saprobes
richness
general saprobes
frequency
0.025
0.014
0.018
0.009
0.009
0.038
g-m
g-o
0.05 0.10 0.05
0.05 0.10 0.05
0.05 0.10 0.05
0.05 0.05 0.05
0.05 0.10 0.05
0.05
58.0
0.05
0.10
0.182
0.05
0.005
0.05
0.05
i
0.iO
0.10
-
0.016
m-O g
27.3 51.7
41.3 114.7
161
638
7.3 47.0
75.0
945
11.3 18.0
137
213
15.7 33.3
200
480
6.3 14.0
0.138
0.083
i-m i-o
Means
m
242
o
47.0 51.0
91.3 102.7
483
569
37.7 50.3
703
970
14.0 10.7
162
132
27.0 28.3
358
382
10.7 11.7
207
208
e 2.3 Similarity of the generic composition of macrofungal communities in different
stand ages as measured by Jaccard's coefficient of similarity. A coefficient of 1.O is exact
similarity and a coefficient of zero is no similarity. g = regeneration, i= immature, m=
mature O=old-growth
6-M
6-0
1-M
1-0
M-O
34
34
58
59
59
genera in
genera in
Coefficient
of
49
49
79
79
72
72
73
72
73
73
0.390805
0.386364
0.623656
0.634409
0.686047
Coe
Sorenson (S,)
0.561983
0.557377
0.768212
0.7763 16
0.813793
Similarity of species composition of macrofungal CO unities in 1997 for al1
pairs of chronosequence plots using Sorenson's coefficient. g - regeneration, i - immature,
m - mature, O - old-growth, s - Watershed South, n - Watershed North, k - Koksilah.
gs
gn
gk
is
in
ik
ms
mn
mk
OS
on
gn
gk
is
in
ik
ms
mn
mk
os
on
ok
0.444
0.273
0.508
0.385
0.344
0.233
0.288
0.281
0.226
0.523
0.314
0.265
0.206
0.546
0.590
0.350
0.356
0.262
0.517
0.458
0.448
0.302
0.269
0.211
0.581
0.605
0.575
0.506
0.273
0.309
0.256
0.539
0.609
0.596
0.489
0.622
0.310
0.272
0.225
0.563
0.475
0.520
0.534
0.559
0.545
0.302
0.245
0.206
0.530
0.529
0.603
0.503
0.657
0.612
0.587
0.191
0.234
0.300
0.443
0.477
0.463
0.439
0.468
0.555
0.452
0.587
able 2.5 Effects of clearcutting on macrofungal genus frequency in CWHxm Douglas
fir dominated stands. S is the number of species. Sensitivity ratings: hs = highly sensitive,
s = sensitive, ms = moderately sensitive, ne = non-sensitive, early succession fungus, ni
= non-sensitive, intermediate succession, nn = non-sensitive non-age-selective, phs =
potentially highly sensitive. The p-value reported is the probability that the variation in
frequency with age is due to chance. Comparisons - numbers indicate a at which test is
significant, comparisons that were not significant at a = 0.10 are lefi blank. Frequency
data reported are from 1997 Stand ages: g = regeneration, i = immature, m = mature O =
old-growth.
Comparison
AOV
Genus
Camarophyllus
Clavariadelphus
Clavulinopsis
Cortinarius
Dermocybe
Gomphidius
Gymnopilus
Hebeloma
Hemimycena
Hydnellum
Inocybe
Laccaria
Lactarius
Lycoperdon
Mycena
Nolanea
Omphalina
Pholiota
Psilocybe
Russula
Stro bilurus
Suillus
Tricholoma
S Sens. p
4 hs
2 phs
2 nn
23 ms
5 ms
4 ms
5 nn
3 ms
3 ms
5 pms
9 ms
3 ms
7 ms
5 nn
34 ms
4 ms
2 nn
2 hs
2 ne
I l ms
2 ms
6 nn
13 ms
0.04
0.07
0.98
0.00
0.04
0.09
0.78
0.03
0.01
0.08
0.00
0.00
0.02
0.93
0.01
0.00
0.68
0.01
0.05
0.00
0.03
0.91
0.00
g-i
g-m g-O i-m i-O m-O
O. 1
0.05
0.05 0.05 0.05
0.05
O. 1
0.05
0.05 0.05 0.05
0.05 0.05 0.05
0.05 0.05 0.05
0.05
0.05
0.05 0.05 0.05
0.05 0.05 0.05
0.05 0.05
O. 1
0.05 0.05 0.05
0.1
0.05
0.05 0.05 0.05
Frequency
g
i
m
O
1.7
3.3
0.0 15.0
0.0
0.0
0.0
3.3
16.7 13.3
8.3 13.3
0.0 238.3 168.3 223.3
1.7 66.7 15.0 26.7
0.0 11.7
6.7 16.7
13.3 10.0
5.0
5.0
1.7 55.0 33.3 23.3
0.0 38.3 70.0 38.3
0.0 26.7 28.3
5.0
8.3 85.0 96.7 105.0
0.0 38.3 30.0 53.3
1.7 90.0 41.7 90.0
18.3 10.0 13.3 10.0
66.7 251.7 196.7 191.7
3.3 56.7 43.3 46.7
18.3 10.0 10.0 10.0
3.3
0.0
0.0
5.0
23.3
8.3
3.3
1.7
0.0 151.7 96.7 223.3
0.0 15.0
8.3 20.0
15.0
8.3 13.3 15.0
0.0 38.3 81.7 65.0
Table 2.6 Effects of clearcutting on macrofkngal species in CWHxrn Douglas-fir forests in 1997. Guilds: a = basidiolichen, f =
fùngicolous, 1 = Iitter-decay, m = mycorrhizal, p = parasitic, s = general saprobes, w = wood-decay. Sensitivity ratings: hs = highly sensitive, s = sensitive,
ms = moderately sensitive, ne = non-sensitive, early succession fungus, ni = non-sensitive, intermediate succession, nn = non-sensitive non-age-selective,
p denotes potential sensitivity rating. Frequency data reported (nurnbers) are from 1997.Stand ages: G = regeneration, 1 = immature, M = mature O =
old-growth. The p-value reported is the probability that the variation in frequency with age is due to chance. This table shows only species with significant
age effects and those with greater than or equal tolO% total frequency that showed no change in frequency with age.
Species
Agaricus sp.
Clavicorona taophila
Clavulinopsisfusiformis
Clitocybe deceptiva
Coltricia perennis
Cortinarius acutus vel aff:
Cortinariusfulvescens vel a f
Cortinarius obtusus vel a f
Cortinarius sg. telamonia spp.
Cortinarius vibratilis
Dacymyces palmatus
Demzocybe malicoria
Dermocybe semisanguinea
Fomitopsis pinicola
Galerina badipes vel a f
Galerina emetensis vel afl
Gomphidius smithii
Guepiniopsis alpina
Hebeloma cmstuliniforme
Hebeloma mesophaeum vel aff
Hemimycena delectabilis
Hygrophorus bakerensis
Guild Sensitivity p(age)
1
phs
0.07
1
pni
0.09
1
0.98
1
hs
0.01
m
ni
0.08
m
ms
0.00
m
ms
0.07
m
ms
0.00
m
ms
0.00
m
pms
0.09
w
pne
0.07
m
ni
0.04
m
pni
0.07
w
pni
0.09
s
ni
0.02
1
ms
0.06
m
phs
0.09
w
0.47
m
ms
0.04
m
ni
0.00
1
ms
0.05
m
pni
0.07
G-1
G-M
GO
-0.2
-0.2
0.2
-0.2
-0.05
-0.1
-0.05
-0.05
-0.2
0.2
-0.1
-0.2
-0.05
-0.2
-0.05
-0.05
-0.2
-0.2
1-M
-0.05
-0.05
-0.2
-0.05
-0.05
-0.2
0.2
-0.2
-0.2
-0.2
0.2
0.1
1-0
-0.2
0.2
M-O
-0.2
-0.05
0.1
-0.05
-0.2
-0.05
-0.05
-0.2
0.2
0.2
0.2
-0.2
0.1
0.2
0.2
O. 1
-0.1
-0.2
-0.2
-0.2
-0.1
-0.1
-0.2
0.05
0.05
0.2
0.2
-0.2
G
O
O
17
O
5
O
O
O
O
O
3.3
O
O
O
O
1.7
O
53
1.7
O
O
O
I
O
5
12
3
20
33
28
42
80
12
O
10
7
O
10
18
O
52
42
10
22
3
M
O
O
8
O
O
32
25
22
70
12
O
2
O
12
5
23
O
30
32
2
47
O
O
3
O
12
7
O
37
22
37
82
15
O
O
O
O
2
25
5
40
23
O
37
O
w
F
W
Hypholomafasciculure
Inocybe geophylla
lnocybe lacera vel a$
Inocybe ovatocystis
Inocybe spp.
Laccaria bicolor
Laccaria laccata
Lactarius rubri/acteus
Marasmius salalis
Murasmius spp.
Mollisia sp.
Mycena alcalina
Mycena flavoalba
Mycena para
Mycena sp. 1
Mycena sp. 2
Mycena spp.
Nolanea holoconiota vel afl
Omphalina chlorocyanea
Omphalina ericetorum
Otidia alutacea vel aff
Phellinus sp.
Phellodon melaleucz4s
Pholiota sp.
Polyporus hirtus
Pseudohydnum gelatinosurn
Psilocybe montana
Ramariu spp.
w
m
m
m
m
m
m
m
pne
ms
phs
ms
ms
ms
ms
ms
1
s
w
ni
pni
S
1
pni
1
1
1
S
S
1
pni
ni
ms
ms
pne
a
1
w
m
w
s
W
1
*
ms
pni
pni
phs
pni
ms
ne
Ps
Species
Rickenella fibulu
Russula iagilis vel iiff
Russula brevipes
Russula placita
Russula sp. 1
Russula spp.
Russula xerampalina
Strobilurus îrullisatus
Tricholoma atroviolaceum
Tricholomaflavovirens
Tricholoma sejunctum
Tricholoma spp.
Tricholoma sulphureum
Xeromphalina fulvipes
Guild Sensitivity
p(age)
G-1
C-
6-0
0.2
1-M
1-0
G
8.3
1
O
e 2.7 Macrofungal species found only on old-growth CWHxm Douglas-fir stands
from 1995 to 1997.
Species
Agaricus sp.
Agrocybe sp.
Amanita porphyria
Amanita vaginata
Auricularia sp.
Baeospora myosura vel aff
Boletus chrysenteron vel aff
Cantharellusjormosus
Clavariadelphus ligula
Clavariadelphus truncatus
Cortinarius brunneus vel a$
Cortinarius claricolor
Cortinarius nigrocuspidatus vel a$
Cortinarius rapaceus vel aff
Cortinarius scaurus vel a f
Cortinarius sg. leprocybe spp.
Cortinarius squamulosus vel a f
Cortinarius vanduzerensis
Cortinarius violaceus
Cryptoporus volvatus
Geoglossum sp.
Gomphidius oregonensis
Gomphidius smithii
Helvella queletii vel a f
Species
Hydnellum scrobiculatum vel aff
Hygrocybe SPP.
Lactarius mucidus vel afi
Leccinum mamanitae
Lepiota sp.
Macrotyphulafistulosa vel a#
Marasmius limosus vel a f f
Micromphale foetidum vel a f
Mycena adonis
Mycena brownii vel afl
Mycena iodolens vel aff
Mycena metata
Mycena mida vel a f
Mycena sp. 6
Paxillus involutus vel a f f
Peziza spp.
Pholiota malicola vel afl
Russula adusta vel a f f
Russula cascadensis vel a f f
Russula sp. 1
Sarcosphaera crassa
Tricholoma atroviolaceus
Tricholoma sulphureum
Tricholoma terreum vel aff
Power of post-hoc comparisons of age classes for different ANOVA tests.
regeneration, i= immature, m= mature O=old-growth
Comparison
Test
genus richness (~0.05)
genus richness (a=0.10)
genus richness (a=O.
15)
species richness (a=0.05)
species richness (~0.10)
species richness(a=O.15)
total frequency (~0.05)
total frequency (a=O.
1 O)
mycorrhizal rich. (a=0.05)
mycorrhizal rich. (~0.05)
mycorrhizal freq. (a=0.05)
mycorrhizal freq. (a=O.
10)
wood-decay rich. (a=0.05)
wood-decay rich. (a=O.
1 O)
wood-decay freq. (a=0.05)
wood-decay freq. (w0.10)
litter-decay rich. (a=0.05)
litter-decay rich. (a=O.
10)
litter-decay freq. (~0.05)
litter-decay freq. (a=0.10)
other saprobes rich. (a=0.05)
other saprobes rich. (a=O.
10)
other saprobes freq. (~~0.05)
other sa probes freq. (a=O.
1 0)
non-wooddecay rich. (~0.05)
non-wood-decay rich. (a=O.
10)
non-wood-decay freq. (a=0.05)
non-wood-decay freq. (a=O.
10)
. Summary results of ANOVAs on genus richness data from each month.
Significant probabilities are in bold.
ont
March
April
May
June
July
August
September
October
November
square
2.75
2.64
6.75
16.64
20.08
3.31
14.92
59.81
35.13
0.191
0.627
0.155
0.775
0.222
0.464
0.101
0.113
Effects of clear-cutting on macrofungi: a) genus richness vs. stand age, b)
species richness vs. stand age, c) total species frequency vs. stand age. (G - regeneration, 1
- immature, M - mature, O - old-growth)
Cladogratn of chronosequence plots grouped by similarity of species
composition. g = regeneration, i = immature, m = mature O = old-growth, s = Watershed South, n =
Watershed North, k = Koksilah.
1 00
300
500
700
900
Area sampled (m2)
ure 2.3 Macrofùngal species area cwves for differently aged CWWxm Douglas-f~
stands.
igure 2.5 Effecü of clear-cutting on cumulative species fkequency of macrofungal
guilds: a) mycorrhizal guild frequency vs. stand age, b) litter-d
stand age, c) wood-decay guild richness vs. stand age. ,d) ge
vs. stand age. (g - regeneration, i - immature, m - mature, o - old-growth)
Eother saprobes
Ed litterdecay
O paraslic
Owooddecay
C3 other saprobes
ed litterdecay
parasitic
a wooddecay
Changes in macr
) proportions of to
old-growth)
guilds with stand age: a) proportions of total
ueney. (g - regeneration, i - immature, m - mature, o -
7.5
10.0
12.5
umber of replicates
igure 2.7 Power versus number of replicates for ANOVA to detect differences in
macrofungal genus richness related to stand age at three type-1 error rates.
alpha = 0.05
....&... alpha = 0.10
.-.a-.alpha = 0.15
Power versus number of replicates for ANOVA to detect differences in
macrofungal species ric ess related to stand age at three type-1 error rates.
1
Regen. vs. Mature
Regen. vs. Old-growt
I
2
3
4
5
6
7
8
9
10
Number of replicates
Power versus nurnber of replicates for Tukey's post-hoc cornparisons (a=
0.10) involving regeneration age.
-
Immature vs. Mature
Immature vs. Old-growtt
--o- Mature vs. Oid-growth
l
I
I
I
I
I
I
I
l
70
90
1O
30
50
Number of replicates
Power versus number of replicates for Tukey's post-hoc cornparisons (aa =
ature, mature and old-growth ages.
5
1O
15
20
25
Effect size (number of genera)
30
Effect size versus power of Tukey's post-hoc cornparisons (a = 0.10).
O
2O
40
60
Number of replicates
80
1O0
2 Effect size detectable with 90% power using Tukey's post-hoc cornparisons
(a = 0.10) versus nurnber of replicates.
GlMOGlMOGlMOGlMOGlMOGlMOGlMOGlMOGlMO
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Box plots of genus richness by month. Medians are indicated by white
bands. (6- regeneration, I - Immature, M - mature, O - old-growth)
The macrofungal community in Douglas-fir dominated CWHxm forests is very
diverse, and measures should be taken to protect this diversity. Further research is
required to determine if other forest ecosystems on Vancouver Island or coastal
Columbia are equally diverse, and why macrofùngal diversity is so high in this habitat or
region.
The species list compiled for Douglas-fir dominated C
m forests is far from
complete and there may be almost 2000 species of macrofùngi in these forests. It may not
be possible to compile a complete or nearly complete speeies list through systematic
sarnpling because sampling effort must increase exponentially for linear increases in
numbers of species observed.
There is high annual variation in the macrofùngal community due to the sporadic
nature of sporoc
production and multiple year studies are necessary to characterize the
community.
There were three distinct
iting seasons and most species fmited only or mainly
in one season. The seasonal variation in the macrofungal community is more pronounced
than the annual variation and e community should be sarnpled during
seasons to be properly characterized.
Wigh sala1 cover may be related to lower litter-decay guild and mycorrhizal guild
richness and this relationship needs
er investigation.
Sporocarp studies are an effective method for studying mycorrhizal diversity and
can detect a greater range of diversity than can mycorrhizal root-tips studies with the
same sampling effort provided that the weather conditions are conducive to sporoc
production.
The richness and frequency of the macrofungal community and the mycorrhizal,
litter decay, and general saprobes guilds, and many genera and species were negatively
affected by clearcutting. The richness and frequency of the macrofungal community, and
the aforementioned guild and the frequency of many species and genera return to oldgrowth levels within 38 to 49 years.
The development of a forest canopy, with its resultant change in temperature and
moisture regimes, provision of mycorrhizal hosts, and input of litter and woody debris
appears to be the major factor in the return of pre-clearcutting macrofungal communities.
The predominant method by which macro
al diversity returns is most likely
recolonization by airborne spores from nearby forests. The effect of distance to nearest
old-growth on macrofungal richness of second growth stands and the effect of the
proportion of old-growth remaining in a landscape and macrofungal richness on secondgrowth stands need to be investigated.
Some species that did not recover to their old-growth frequencies until 83 -105
years and some that did not recover within 83-105 years. These species need to studied in
more detail and possibly managed to maintain their populations.
The mature stands were, on average, lower in richness and fi-equencythan the oldgrowth. Further research is needed to determine whether or not this difference is real.
Statistical power in this study was low and species and groups that did not show
significant changes in frequency with stand age should not be considered to be unaffected
by clearcutting because there was insufficient evidence.
The less stringent type-1 error rate of 0.10 should be used for this study to reduce
the chance of a type-II error. Power analysis should be done on al1 environmental impact
studies to determine the chance of a type-II error, i.e. the chance of mistakenly saying
there are no negative effects when in fact there was not enough statisitical power to detect
the negative effects actually exist.
The number of replicate could be increased without increasing sampling effort if
new chronosequences were visited each month. The drawback of not visiting the same
sites throughout the year is that the knowledge of community composition for a site
would be incomplete. If only three chronosequences could be sampled each month, a
compromise would be to visit one chronosequence repeatedly and two new ones each
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complete descriptions see TroQmow et al. (1 997).
Attribute
Regeneration Immature
ature
Old-growt
Biogeoclimatic Zone / Subzone /
Variant / Site Series
Age in 1991
Elevation (m)
Slope Gradient (%)
Aspect (degrees)
Total N kglha
Basal area (&/ha)
Douglas-Fir
western hemlock
western redcedar
red alder
al1 trees
Sala1 cover (%)
CWHxml/O 1
Biogeoclimatic Zone / Subzone 1
variant / Site Series
Age in 1991
Elevation (m)
Slope Gradient (%)
Aspect (degrees)
Total N kgha
Basal area (&/ha)
Douglas-FU
western hemlock
westem redcedar
al1 trees
Sala1 cover (%)
oksilah
Biogeoclimatic Zone / Subzone /
Variant / Site Series
Age in 1991
Elevation (m)
Slope Gradient (%)
Aspect (degrees)
Total N kglha
Basal area (nS/ha)
Douglas-FU
western hemlock
western redcedar
al1 trees
Sala1 cover (%)
CWHxm2/0 1
CWHxm21Ol CWHxml/Ol
CWHxm2/01
CWHxm2/0 1
CWHxm2IO1 CWHxm2I03
CWHxm2/03
CWHxml/Ol CWHxml/Ol- CWHxmlIOl
07
4
32
99
245
280
305
240
390
15
40
11
40
50
20
315
30
1428
1565
2212
76 1
22.7
7.7
0.3
20
30.4
60
8 1.8
0.6
4.8
1.5
88.7
15
73.6
5.7
2.9
0.9
83.1
40
5
595
15
170
1570
43
710
15
170
1161
77
590
35
210
862
288
630
15
180
1134
63
87.2
0.5
0.1
40
41.9
2.4
3.3
47.6
30
63
35
87.7
70
1. Authorities for macrofùngi found on C
forests on southeastern Vancouver Island.
Species
Agaricus amensis
Agaricus semotus vel aff
Agaricus sp.
Agrocybe sp.
Albatrellus sp.
Amanita porphyria
Amanita silvicola
Amanita vaginata
Armillaria ostoyae
Auricularia sp.
Auriscalpium vulgare
Baeospora myosura vel afl
Bisporella citrina
Boletopsis subsquamosa
Boletus chrysenteron vel aff
Bondarzewia montana
Bovista plumbea vel a f f
Callisrosporium luteo-olivaceum vel af
Calocera viscosa
Camarophyllus borealis
Camarophyllus niveus
Camarophylluspaupertinus vel af
Camarophyllus recurvatus vel a f f
Camarophyllus sp.
Cantharellula umbonata
Cantharellusformosus
Cantharellus inJZtndibul~formis
Cantharellus subalbidus
Chlorociboria aeruginascens
Chroogomphus tomentosus
Ciboria rufofusca
Clavaria vermicularis
Clavariadelphusligula
Clavariadelphustruncatus
Clavicorona taxophila
Clavulina cinerea
Clavulina cristata
Clavulinopsis corniculata
Clavulinopsisfusiformis
Clitocybe avellaneialba
Clitocybe coniferophia vel a f
Clitocybe dealbata vel a$
Clitocybe deceptiva
Clitocybe harperi
Clitocybe inversa vel aff
AuthoriQ
Fries
Fries
Murrill
(Fr.) Secr.
Kauffman
(Fr.) Vitt.
(Romagnesi) Herink
S. F. Gray
(Fr.) Sing.
(Batsch ex Fr.) Koef & Carpenter
(Fries) Kotl. & Pouzar
Fr.
(Quel.) Singer
Pers.
(Berk. & Curt.) Singer
(Fr.) Fries
(Pk.) Murr.
(Fr.) Wunsche
Hygophorous paupertinus Sm. & Hes.
(Pk.) M m .
(Fr.) Singer
Corner
Fr.
Smith & Morse
Chlorosplenium aeruginascens (Nyl.) Karst.
(Mm.) O.K.Miller
(Weberb.) Sacc.
Miche1li:Fr.
(Fr.) Donk
(Quel.) Donk
(Thom) Doty
(Fr.) Schroet.
(Fr .) Schroet.
(Schaeff. ex Fr.) Corner
(Sow. ex Fr.) Corner
Murrill
Bigelow
(Fr.) Kummer
Bigelow
Murr.
(Fr.) Quelet
Species
Clitocybe nebularis
Clitocybe sp. 1
Clitocybe spp.
Collybia acervala
Collybia bulyracea
Collybia cirrhata
Collybia conjluens
Coliybia dryophila
Collybia maculata vel aff
Collybia racemosa
Collybia spp.
Coliybia tuberosa
Coltriciaperennis
Conocybe tenera vel a#
Cora'yceps sp.
Coriolellus sepium vel aff
Cortinarius acutus vel aff
Cortinarius alboviolaceus
Cortinarius anomalus
Cortinarius boulderensis vel afl
Cortinarius brunneus vel a#
Cortinarius callisteus
Cortinarius caninus vel a#
Cortinarius castaneus vel aff
Cortinarius clandestinus
Cortinarius claricolor
Cortinarius dilutus vel a#
Cortinariusfulvescens vel afl
Cortinarius gentilis vel afl
Cortinariusjubarinus vel aff
Cortinarius nigrocuspidatus vel a 8
Cortinarius obtusus vel aff
Cortinariuspfumiger vel aff
Cortinariuspyriodorus
Cortinarius rapaceus vel aff
Cortinarius scaurus vel afl
Cortinarius sg. leprocybe spp.
Cortinarius sg. myxacium spp.
Cortinarius sg. phlegmacium spp.
Cortinarius sg. seriocybe spp.
Cortinarius sg. telamonia spp.
Cortinarius sp. 1
Cortinarius sp. 2
Cortinarius squamulosus vel a f
Cortinarius superbus vel afl
Cortinarius vanduzerensis
Authority
(Bat. ex Fr.) Kumm
(Fr.) Kummer
(Fr:Fr) Quelet
(Schumex Fr.) Kummer
(Fr.) Kummer
(Bu1l.e~Fr.) Kummer
(A.&S.ex Fr.) Kummer
(Fr.) Quelet
(Fr.) Quelet
(L. ex Fr.) Murr.
(Schaeff.ex Fr.) Fayod
Trarnetes sepiurn Berk.
Fr.
(Pers. ex Fr.) Fries
(Pers. ex Fr.) Fries
A.H. Sm.
Fr.
(Fr.:Fr.) Fr.
(Fr.) Fr.
Fr.
Kauff.
Fr.
Fr.
Fr.
(Fr.)Fries
Fr.
Fries
(Fr.) Fr.
Cortinarius traganus Fr.
Fr.
Fries
Pk.
Sm.
Smith & Trappe
Species
Cortinarius vibratilis
Cortinarius violaceus
Crepidotus applanatus
Crepidotus&sisporus
Crepidotus herbarum
Crepidotus mollis vel aff
Crepidotus spp.
Cyptoporus volvatus
Cudonia circinans
Cudonia monticola
Cvstoderma amianthinum
Cystodermafallax
Cystoderma granulosum
Dacrymyces palmatus
Dasyscyphus bicolor
Dentinum umbilicatum
Dermocybe californica
Dermocybe cinnarnomea vel ajf
Dermocybe malicoria
Dermocybe phoenicea
Dermocybe semisanguinea
Discina perlata
Entoloma sp.
Fomitopsis cajenderi
Fomitopsis o#icinalis
Fomitopsis pinicola
Galerina autumnalis
Galerina badipes vel a#
Galerina emetensis vel aff
Galerina marnmilata
Galerina spp.
Galerina vexans vel aff
Geoglossurn sp.
Gloeophyllum saepiarium
Gomphidius oregonensis
Gomphidius smithii
Gomphidius spp.
Gomphidius subroseus
Guepiniopsis alpina
Gymnopilus bellulus vel a i
Gymnopilus croceoluteus vel aff
Gymnopiluspicreus
Gymnopilus sapineus vel aff
Gymnopilus spectabilis
Gymnopilus spp.
Gymnopilus terrestris
Authority
(Fr.) Fries
(L.:Fr.) S.F.Gray
(Pers.) Kummer
Hesler & Smith
(Pk.) Sacc.
(Fr.) Staude
(Pk.) Hubbard
Fr.
Mains
(Fr.) Fayod
Smith & Singer
(Morg.) Smith & Singer
(Schw.) Bres.
(Bull. ex Merat) Fuckel
(Pk.) Pouzar
Ammirati
(L.: Fr.) Wunsche
(Fr.) Ricken
(Bull. ex Mre.) Mos.
(Fr.) Mos.
Fr.
(Karsten) Kotl. & Pouz.
(Vil.: Fries) Bond. & Sing
(Swartz: Fries) Karsten
(Pk.) Smith & Singer
(Fr.) Kuhner
(Murrill) Smith & Singer
(Fr.) Karst
Peck
Miller
Peck
(Tracey & Earle) Bres.
(Pk.) Murr.
Hesler
(Fr.) Karst.
(Fr.) R.Maire
(Fr.) A.H.Smith
Karsten
Hesler
.continued
Gyromitra esculenta
Hebeloma crustuliniforme
Hebeloma mesophaeum vel afl
Hebeloma sacchariolens
Helvella Iacunosa
Helvella queletii vel afl
Hemimycena delectabilis
Hemimycena sp. 1
Hemimycena spp.
Hydnellum aurantiacum
Hydnellum caeruleum
Hydnellum conigenum
Hydnellum peckii .
Hydnellum scrobiculatum vel a f
Hydnellum spp.
Hydnellum suaveolens
Hydnum caivatum
Hydnumfennicum
Hydnum Jùscoindicum
Hydnum scabrosum vel afl
Hydnum sp. 1
Hydnum sp. 2
Hygrocybe conica
HygrocybeJlavescens
Hygrocybe miniata
Hygrocybe sp. 1
Hygrocybe spp.
Hygrocybe unguinosus vel af
Hygropkoropsis aurantiaca
Hygrophoropsis olida
Hygrophorus agathosmus
Hygrophorus bakerensis
Hygrophorus camaroplgdlus
Hygrophorus eburneus vel afl
Hygrophorus picea
Hygrophorus sp. 1
Hygrophorus spp.
Hypholoma capnoides
Hypholoma fusciculare
Inocybe calamistrata
Inocybe geophylla
Inocybe lacera vel afl
Inocybe Macina
Inocybe maculata vel a 8
Inocybe mixtilis
Inocybe napipes vel afi
Authority
(Pers.) Fr.
(StAmans) Quel.
(Fr.) Quel.
Quel.
Afz. ex Fr.
Bres.
(Peck sensu Smith) Sing.
(Fr.) Karsten
(Hornem. ex Pers.) Karst.
(Pk.) Banker
Banker apud Peck
(Fr. ex Secr.) Karst.
(Scop. ex Fr.) Karst.
Harrison
Karst.
Harrison
Fr.
(Fr.) Kummer
(Kauff.) Sing.
(Fr.) Kummer
(Fr.) Karst.
(Wu1f.e~Fr.) Maire
Quel.
Fr.
Sm. & Hes.
(Fr.) Dumee, Grandjean, et Marie
(Fr.) Fr.
Kuhn. & Romagn.
Fr.
Fr.
(Fr.) Kummer ex Fries
(Huds.ex Fr.) Kummer
(Fr.) Quelet
(Fr.) Kummer
(Fr.) Kummer
(Boud.) Kauff.
Boudier
(Britz.) Sacc.
Lange
Species
Inocybe ovatocystis
Inocybe sororia
Inocybe spp.
Kuehneromyces vernalis
Laccaria altaica vel af
Laccaria amethysteo-occidentalis
Laccaria bicolor
Laccaria laccata
Lactarius luculentis
Lactarius mucidm vel aff:
Lactarius pallescens
Lactarius pseudomucidus
Lactarius rubrilacteus
Lactarius rufus vel aff:
Lactarius scrobiculatus
Lactarius spp.
Leccinum manzanitae
Lentaria spp.
Lepiota clypeolaria
Lepiota seminuda vel afJ:
Lepiota sp.
Leptonia asprella vel aff
Leptonia carnea
Leptonia sp.
Lepton ia undulatella
Lycoperdon foetidum
Lycoperdon molle
Lycoperdon perlatum
Lycoperdon pyrifornze
Lycoperdon sp.
Lyophyllum decastes
Lyophyllum semitale
MacrotyphulaJistulosa vel aff
Marasmiellus candidus vel aff
Marasmius androsaceus
Marasmius epiphyllus vel aff:
Marasmius Iimosus vel aff
Marasmius salalis
Marasmius sp. 1
Marasmius spp.
Marasmius umbilicatus
Marcelleina sp.
Melanoleuca graminicola
Melanotus textilis vel a f f
Merulius tremellosus
Micromphalefoetidum vel aff
Authority
Kuehn.
Kaufhan
(Peck) Sing. & Smith
Sing.
Mueller
(R.Maire) Orton
(Fr.) Berk. & Br.
Burlingham
Burl.
Hesler & Smith
Hesler & Smith
Hesler & Smith
(Fr.) Fr.
(Fr.) Fr.
Pers. ex S. F. Gray
(Bulliard ex Fries) Kummer
(Lasch) Kummer
(Fr.:Fr.) Kummer
Largent
Pers.
(Peck) Saccardo
Bonor.
Pers.
Pers.
Pers.
(Fr.) Sing
(Fr.) Kuh.
ClavariadelphusJistulosus(Fr.) Corner
(Bolt.) Singer
(L. ex Fr.) Fr.
(Persoonex Fr.) Fr.
Boud. & Quel
Desjardin & Redhead
(Vel.) Kuehn. & Mre.
Redhead & Kroeger
Schrad. ex Fr.
(Sow. ex Fr.) Sing.
Speeies
Micromphale perforans vel a#
Mitrula abietis vel a f
Mollisia sp.
Multiclavula mucida vel a#
Mycena fusco-ocula vel a@
Mycena acicula
Mycena adonis
Mycena alcalina
Mycena alnicola
Mycena amabilissima
Mycena amicta
Mycena aurantiidisca
Mycena brownii vel afl
Mycena capillaripes
Mycena capillaris
Mycena subsanguinolenta vel a#
Mycena citrinomarginata
Mjxena clmularis vel a#
Mycena concolor vel a#
Mycena elegantula
Mycena epipterygia
Mycena epipterygioides
Ililycena flavoalba
Mycena galopus
Mycena gaultherii
Mycena haematopus
Mycena inclinala
Mycena iodolens vel afl
Mycena leptocephala vel a#
Mycena litoralis vel a$
Mycena maculata
Mycena metata
Mycena oregonensis
Mycena paucilamellata
@cena pura
Mycena purpureofusca
Mycena rorida
Mycena rubromarginata
m e n a sanguinolenta
Mycena sp. 1
Mycena sp. 2
Mycena sp. 3
Mycena sp. 4
Mycena sp. 5
Mycena sp. 6
Mycena sp. 7
AuthoriQ
(Hofm. & Fr.) Quel
Heyderia abietis (Fr.) Link
(Fr.) Peterson
A. W. Smith
(Fr.) Quelet
(Fr.) S.F.Gray
(Fr.) Quelet
A.H. Smith
(Pk.) Saccardo
(Fr.) Quelet
Murrill
A.H. Smith
Peck
(Fr.) Quelet
A. W. Smith
Gillet
(Fr.) non Lange
(Lange) Kuhner
Peck
(Fr.) S.F.Gray
Pearson
(Fr.) Quelet
(Fr.) Quelet
A. H. Smith
(Fr.) Quelet
(Fr.) Quelet
Lundell
(Fr.) Gillet
Smith
Karsten
(Fr.) Quelet
A. H. Smith
A. H. Smith
(Fr.) Quel.
(Pk.) Sacc.
(Fr.) Quelet
(Fr.) Quelet
(Fr.) Quelet
Species
Mycena sp. 8
Mycena spp.
Mycena strobilinoides
Mycena subcana vel aff
Mycena tenerrima
Mycena vitilis vel aff
Mycena vulgaris vel aff
Neournula pouchetii
Nidula niveotomentosa
Nolanea cetrata vel aff
Nolanea hoioconiota vel aff
Nolanea sericea vel aff
Nolanea spp.
Oligoporus caesius
OligoporusJYagilis
Oligoporus tephroleucus vel aff
Qmphalina chlorocyanea
Qmphalina chrysophylla
Omphalina ericetorum
Qsteina obducta
Otidia alutacea vel a 8
Panellus serotinus vel aff
Panellus stipticus
Paraeccifia sericeonitida var. ligniphila
Paxillus utrotomentosus
Paxillus involutus vel a f
Peziza spp.
Phaeolus schweinitzii
Phellinus sp.
Phellodon atratus
Phelladon melaleueus
Phellodon tomentosus
Pholiota decorata
Pholiota highlandensis vel aff
Pholiota malicola vel aff
Pholiota sp.
Pholiota terrestris
Phylloporus rhodoxanthus
Plectania melastoma
Pleurocybellaporrigens vel aff
Pleurotus ostreatus vel a f
Pluteus ceminus
Plutezcs~jlavofilgineus
Polyporus elegans
Polyporus hirtus
Psathyrella gracilis vel aff
Authority
Peck
Smith
(Berk.) Quelet
(Fr.) Quelet
(Fries) Quelet
(Bert. & Rious.) Paden
(Henn.) Lloyd
(Fr.) Kummer
Largent & Thiers
(Bull.) P.D.Orton
(Schrad.: Fr.) Gilbn. & Ryv.
(Fr.) Gilbn. & Ryv.
(Fr.) Gilbn. & Ryv.
(Pat.) Sing.
Chrysomphalina chrysophylla (Fr.) Clc.
(Fr.) M. Lange
(Berk.) Donk
(Pers.) Mass.
(Pers. ex Fr.) Kuehn.
(Bull.ex Fr.) Karst.
Largent
Fries
(Batsch) Fr.
(Fr.) Pat.
Quelet
Harrison
(Fr.) Karsten
(Fr.) Banker
(Murr.) Smith & Hesler
(Pk.) Smith & Hesler
(Kauff.) Smith
(Fr.) Kummer
Overholts
(Schw.) Bresadola
(Sow. ex Fr.) Fuckel
(Pers. ex Fr.) Singer
(Fr.) Kummer
(Fr.) Kummer
Atkinson
BuII.: Fr.
Quel.
(Fr.) Quelet
Psathyrella Iongistriata
Psathyrella spp.
Pseudocoprinus disseminatus vel afl
Pseudohydnum gelatinosum
Pseudoplectania melaena
Pseudoplectania nigrella
Psilocybe corneipes vel aff
Psiloqbe montana
Ramaria sp.
Rhodocybe hirneola vel aff
Rickenella fibula
Russula JYagilis vel a$?
Russula adusta vel a f f
Russula bicolor
Russula brevipes
Russula cascadensis vel afi
Russula emetica vel a f f
Russula marei
Russula occidenfalis vel aff
Russula placita
Russula sp. 1
Russula spp.
Russula variata vel afi
Russula xerampalina
Sarcosphaera crassa
Spathulariaflavida
Stereum hirsulum vel aff
Stereum sanguinolentum
Strobilurus alhipilatus
Strobilurus trullisatus
Stropharia ambigua
Suillus borealis vel aff
Suillus granulatus
Suillus lakei
Suillus luteus
Suillus spp.
Suillus subolivaceous
Thelephora terrestris
Trametes hirsuta
Trametes versicolor
Tremella mesenterka vel afi
Trichaptum abietinum
Tricholoma atroviolaceum
Tricholomaflavovirens
Tricholoma inamoenum
Tricholoma magnivelare
AuthoriQ
(Murrill) Smith
Coprinus disseminatus (Persoon ex Fr.)S.F.Gray
(Scop. ex Fr.) Karst.
Fr.) Boud.
(Pers. ex Fr.) Fuckel
(Fr.) Karst.
(Pers. :Fr.) Kummer
(Fr.) Orton
(Bul1.e~Fr.) Raith.
(Persoon ex Fr.) Fnes
Fr.
Burl.
Peck
R.L.Shaffer
Fries
Peck
Singer
Burl.
Ban. apud Pk.
(Secr.) Fries
(Santi) Pouzar
Fr.
(Willd. ex Fr.) S.F. Gray
(Alb. & Schw.: Fr.) Fr.
(Peck) Wells & Kempton
(Murrill) Lennox
(Peck) Zeller
Smith, Thiers & Muller
(Fr.) Kuntze
(Mum.) Smith & Thiers
(Fr.) S.F. Gray
Smith & Thiers
Persoon
(Wulf.: Fr.) Pil.
(L. ex Fr.) Pilat
Retz.
(Dicks.: Fr.) Ryv.
Smith
(Fr.) Quelet
(Fries)Quelet
(Peck)Redhead
Species
Tricholomapessundatum
Tricholomaportentosurn
Tricholoma saponaceum
Tricholoma sejunctutn
Tricholoma sp. 1
Tricholoma spp.
Tricholoma sulphureum
Tricholoma terreum vel aff
Tricholoma virgatum
Tricholomazelleri
Trichopilusplebiodes
Tubaria sp.
Tyromyces chioneus
Xeromphalina campanella
Xeromphalinajû lvipes
Xeromphalina spp.
Xylaria hypoxylon
(Fr.) Quelet
(Fr.) Quelet
(Fr.) Kummer
(Fr.) Quelet
(Bull.: Fr.) Kummer
(Schaeffer: Fries) Kummer
(Fr.) Kummer
(Stuntz & Smith) Ovrebo & Tylutk
(Schulz.) Largent
(Fr.) Karsten
(Fr.) Kuehn.&Maire
(Murr.) Smith
(L. ex Hooker) Grev.
.Nurnber of species per
frequency for each genus in 1997 in C
Genus
Agaricus
Agrocybe
Albatrellus
Amanita
Armillaria
Auricularia
Auriscalpium
Baeospora
Bisporella
Boletopsis
Boletus
Bondarzewia
Bovista
Callistosporium
Calocera
Camarophyllus
Cantharellula
Cantharellus
Chlorociboria
Chroogomphus
Ciboria
Clavaria
Clavariadelphus
Clavicorona
Clavulina
Clavulinopsis
Clitocybe
Collybia
Coltricia
Conocybe
Cordyceps
Coriolellus
Cortinarius
Crepidotus
Cryptopoms
Cudonia
Cystoderma
Dacrymyces
Dasyscyphus
Dentinum
Dermocybe
in 1995, 1996, and 1997 and cumulative
Douglas-fir stands.
Number of species
1995 1996 1997 19951997
1
'
l
1997
frequency (%)
Genus
Entoloma
Fornitopsis
Galerina
Geoglossum
Gloeophyllum
Gomphidius
Guepiniopsis
Gymnopilus
Gyromitra
Hebeloma
Helvella
Hemimycena
Hydnellum
Hydnum
Hygrocybe
Hygrophoropsis
Hygrophorus
Hypholoma
Inocybe
Kuehneromyces
Laccaria
Lactarius
Leccinum
Lentaria
Lepiota
Leptonia
Lycoperdon
Lyophyllum
Macrotyphula
Marasmiellus
Marasmius
Marcelleina
Melanoleuca
Melanotus
Meruiius
Micromphale
Mitrula
Mollisia
Multiclavula
Mycena
Neournula
Nidula
Nolanea
Oligoporus
Number of species
1995
1996
1997 1995-
1997
.continued
Genus
Omphalina
Osteina
Otidia
Panellus
Paraeccilia
Paxillus
Pezizu
Phaeolus
Phellinus
Phellodon
Pholiota
Phylloporus
Plectania
Pleurocybellu
Pleurotus
Pluteus
Polyporus
Psathyrella
Pseudocoprinus
Pseudohydnum
PseudopIectanza
Psilocybe
Ramaria
Rhodocybe
Rickenella
Russula
Sarcosphaera
Spathularia
Stereuin
Strobilurus
Stropharia
Suillus
Thelephora
Trametes
Tremella
Trichaptum
Tricholoma
Trichopilus
Tubaria
Tyromyces
Xeromphalina
Xylaria
Total
Number of speeies
1995
1996 1997 19951997
1997
frequency (%)
,-,
2
3
12.1
.
m Douglas-fir
V Effects of clearcutting on macrofungal species in C
stands. p = probability that the observed change in frequency with stand age is due to. Guilds: a =
basidiolichen, f = fungicolous, 1 = litter-decay, m = mycorrhizal, p = parasitic, s = general saprobes, w =
wood-decay. Stands: g = regeneration, i = iumrnature, m =mature, o = old-growth, s = Watershed South, n
= Watershed North, k = Koksilah. Raw counts (number of quadrats) are presented for each stand.
p
Agaricus sp.
Agrocybe sp.
Albatrellus sp.
Amanita porphyria
Amanita silvicola
Amanita vaginata
Armillaria ostoyae
Auriscalpium vulgare
Bisporella citrina
Boletopsis subsquamosa
Calocera vscosa
Camarophy1lu.s borealis
Camarophyllus niveus
Camarophyllus remmatus vel
a8
Camarophyllus sp.
Cantharellula umbonata
Caniharellusformosus
CanthareIlus
infundibuliformis
Cantharellus subalbidus
Chlorociboria aeruginascens
Chroogornphus tomentosus
Ciboria rufofùsca
Clavaria vermicularis
Clavariadelphus ligula
Clavariadelphus truncatus
Clavicorona taxophila
Clavulina cinerea
Clavulina cristata
Clavulinopsis corniculata
Clavulinopsisfusiformis
Clitoqbe avellaneialba
Clitocybe coniferophila vel
a77
Clitocybe deceptiva
Clitocybe harperi
Clitocybe inversa vel a f f
Clitocybe sp. 1
Clitocybe spp.
Collybia acervata
Collybia butyracea
Coliybia conJluens
1
1
m
m
gs gn gk
is in ik ms
n mk os on ok 97 fre
m
P
1
w
m
w
m
s
s
0 . 0 7 0 0 0 0 0 0 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 0
0 . 4 5 0 0 0 0 1 0 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 0
0 . 6 5 0 0 0 0 0 0 0 1 O
0 . 4 5 0 0 0 0 0 0 0 0 0
0 . 3 1 0 0 0 1 1 9 0 1 2
0 . 4 5 0 0 0 0 0 0 1 0 0
0 . 4 5 0 0 0 0 3 0 0 0 0
0.12 O
0 0 0 0 0 1 3
0 . 6 9 1 0 0 0 1 2 1 1 1
0 . 4 5 0 0 1 0 0 0 0 0 0
0.65 O O O O 1 O O O 0
0.45 O O O O O O O O 0
s
1
m
m
0
0
0
0
m
w
m
1
m
m
m
s
0 . 4 5 0 0
0 . 4 5 0 0
0 . 4 5 4 0
0 . 1 9 0 0 0
0 . 1 3 7 2 0
0 . 4 5 0 0
0 . 4 5 0 0
0 . 0 9 0 0 0
0.45 1 O 0
0 . 1 2 2 0 1
0 . 6 5 0 0
0.983 7 0
0.41 0 0 0
0.821 O 0
1
1
1
1
1
w
1
1
0 . 0 1 0 0 0 1
0 . 4 5 0 0 0 1
0 . 4 5 0 0 0 1
0.650 1 0 0
0.25 1 0 0 4
0 . 4 5 0 0 0 0
0 . 2 5 0 3 0 1
0 . 6 5 0 0 0 0
m
1
m
m
s
1
s
. 4 5 0 0 0 1 0
. 6 3 2 0 0 0 0
. 4 5 0 0 0 0
. 4 5 0 0 0 2
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
2
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
3
0
0 0 0
1 O 0
0 0 0 0
0 0 0 0
0
0
3
0
0 2
0 7
0 0 0 0 1
0 0 0 0 0
0 0 0 0 0
0 1 0 0
0 0 0 0
0 0 0 0 0
0 0 0 0 0
2 0 0 0
0 0 0 0
0 2 0 2
0 1 0 0 0
6 0 3 2
2 1 2 0
1 0 0 4
1
0
0
0
3
1
0
1
0
0
0
0
0
0
0
1
0
3
0
0
1
1
3
0
0
0
1
0
O
O
0 0
0 0
0 0
0 1
2 2
0 0
0 0
O 0
1
0
0
0
O
O
0
0
0
0
2
0
O
1
0
0
1
0
0
0
0
0
0
0
4
1
O
0.8
1.3
0.4
0.4
0.8
0.4
7.1
0.4
1.3
1.7
4.2
2.1
0.8
0.4
0 0
0 0
0 0
0 0
1.7
1.3
0.8
3.8
1
O
0
0
2
2
1
0
0
0
1
1
5
0
0
0
0
0
4
0
O
1
0
0
O
O
2
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0.4
0.8
1.7
3.3
5.0
0.4
0.4
1.3
0.8
3.3
0.8
12.1
2.1
3.8
1
0
0
O
1
0
0
0
1
0
0
0
1
0
0
0
2
0
0
0
O
0
0
0
2.5
0.4
0.4
0.8
6.3
0.4
2.9
0.8
Species
gs gn gk
Guild
Collybia dryophila
1
Collybia racemosa
f
Coliybia spp.
s
Collybia tuberosa
f
Coltriciaperennis
m
Conocybe tenera vel afi
1
Cortinarius acutus vel aSf:
m
Cortinaritls alboviolaceus
m
Cortinarius anomalus
m
Cortinarius callisteus
m
Corfinarius clandestinus
m
Cortinarius dilutus vel aff
m
Cortinariusfulvescensvelaf m
COrtinariusgentilis vel afS
m
Cortinariusjubarinusvelaff m
Cortinarius obtusus vel a 8
m
Cortinariuspyriodorus
m
Cortinariusrapaceusvelafi m
Cortinarius sg. leprocybe spp. m
Cortinarius sg. myxacium spp. m
Cortinarius sg. phlegmacium m
SPP.
Cortinariussg.serios>bespp. m
Cortinarius sg. telamonia spp. m
Cortinarius sp. 1
m
Cortinarius sp. 2
m
Cortinarius superbus vel afl m
Cortinarius vanduzerensis
m
Cortinarius vibratilis
m
Cortinarius violaceus
m
Crepidotus applanatus
w
Crepidotus herbarum
s
Crepidotus spp.
w
Cryptoporus volvatus
w
Cudonia circinans
1
Cudonia monticola
1
Cystoderma amianthinum
1
Cystoderma.fallax
1
Cystoderma granulosum
1
Dacrymyces palmatus
w
Dasyscyphus bicolor
w
Dentinum umbilicatum
m
Dermocybe califovnica
m
Dermocybe cinnamomea vel m
af
Dermocybe malicoria
m
0 .
0 .
0 .
0.61
0 .
0 .
0.00
0 .
0 .
0 .
0.63
0 .
0.07
0 .
0.12
0.00
0 .
0.45
0.45
0.65
0 .
is in ik ms mn mk os on ok 97 fre
9 3 0 1 0 3 0
4 5 0 0 0 0
1 8 1 1 0 3
0 0 0 0 0
0 8 3 0 0 1
3 8 0 0 0 1 0
O O O 313
1 9 0 0 0 0
4 5 0 0 0 0 0
4 5 0 0 0 0 0
O O O O 0
5 9 0 0 0 0
O O O 1 13
6 5 0 0 0 0
O O O O 1
O O O 415
4 5 0 0 0 0 0
O O O O O
O O O O O
O O O O 1
6 8 0 0 0 1
3
0
1
0
0
2
7
0
0
0
O
0
O
0
O
3
0
1
O
O
0
0
0
3
1
0
4
8
1
0
0
1
4
10
1
1
13
0
O
O
O
4
0
0
0
O
0
0
7
1
0
0
O
0
3
0
1
6
0
O
1
1
0
3.8
0.8
7.5
1.7
6.3
5.4
25.4
1.3
0.4
0.4
1.3
6.3
18.8
0.8
2.5
25.0
0.4
0.4
0.4
0.8
2.5
0.45 O O O 1 O O O O O O
0.00 O O O 10 20 18 5 19 18 13
0 . 4 5 0 0 0 0 0 2 0 0 0 0
0 . 4 5 0 0 0 0 2 0 0 0 0 0
0 . 4 5 0 0 0 0 0 1 O 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 0 0
0 . 0 9 0 0 0 0 4 3 0 3 4 0
0 . 4 5 0 0 0 0 0 0 0 0 0 0
0 . 4 5 0 0 0 0 0 0 1 O 0 0
0.45 1 0 0 0 0 0 0 0 0 0
0 . 3 0 2 0 0 1 2 0 0 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 0 1
0 . 6 5 0 0 0 0 0 1 0 0 0 1
0 . 4 5 0 0 0 0 0 0 1 0 0 1
0.572 1 0 2 0 1 1 O 0 1
0 . 4 5 0 0 0 0 0 1 O 0 0 0
0 . 4 5 3 0 0 0 0 0 0 0 0 0
0.07 1 1 0 0 0 0 0 0 0 0
0.200 1 0 2 1 0 0 0 0 0
0 . 1 4 0 0 0 2 3 0 1 4 2 4
0.38 O O 0 O 1 2 O 1 O 1
0 . 4 2 0 0 0 6 0 3 0 2 2 0
O
18
0
0
0
1
6
1
0
0
0
O
O
O
1
0
0
0
0
2
1
4
O
18
0
0
0
O
3
O
0
0
0
0
0
0
O
0
0
0
0
0
O
2
0.4
57.9
0.8
0.8
0.4
0.4
9.6
0.4
0.4
0.4
2.1
0.4
0.8
0.8
3.8
0.4
1.3
0.8
1.7
7.5
2.5
7.9
0
0
2.9
0 . 0 4 0 0 0 4 1
0
0
3
3
7
3
4
0
1
1
2
1
3
0
3
6
1
O
O
O
0
1
1 1 0
0 0 0 2
1 1 2 2
0 0 0
4 0 0 0
1 1 1
211 6
1 0 0 0
0 0 0
O 0 0
O O 0
6 0 4 0
O 9 6
1 0 0 0
O O O
4 5 4
O 0 0
O O O
O O O
O O O
0 0 1 0
0
0
1
O
Guild
Dermocybe phoenicea
Dermocybe semisanguinea
Discina perlata
Entoloma sp.
Fomitopsis cajenderi
Fomitopsis oflcinalis
Fomitopsis pinicola
Galerina autumnalis
Galerina badipes vel aff
Galerina emetensis vel a 8
Galerina mammilata
Galerina spp.
Calerina vexans vel a#
Geoglossurn sp.
Gloeophyllunz saepiarium
Gomphidius oregonensis
Gomphidius smithii
Gomphidius spp.
Gomphidius subroseus
Guepiniopsis alpina
Gymnopilus bellulus vel aff
Gymnopilus croceoluteus vel
afi
G~mnopilus
picreus
Gymnopilus sapineus vel aJT
Cymnopilus spp.
Gymnopilus terrestris
Gyromitra esculenta
Hebeloma crustuliniforme
Hebeloma mesophaeum vel
af3:
Hebeloma sacchariolens
Helvella lacunosa
Helvella queletii v e l d
Hemimycena delectabilis
Hemimycena sp. 1
Hemimycena spp.
Hydnellum aurantiacum
Hydnellum caeruleum
Hydnellum peckii
Hydnellum suaveolens
Hydnum calvatum
Hvdnum scabrosum vel aff
Hydnum fennicum
Hydnum fuscoindicum
m
m
1
m
w
w
w
s
s
1
1
s
s
1
w
m
m
m
m
w
w
w
w
w
w
s
1
m
m
m
1
1
1
I
1
m
m
m
m
m
m
m
m
p
gs gn gk
is in ik ms mn mk os on ok 97 fre
0.11 1 O O 5 2 11 1 1 1 O
0 . 0 7 0 0 0 2 0 2 0 0 0 O
0 . 6 5 0 0 0 0 0 2 0 0 0 3
0 . 4 5 6 0 0 0 0 0 0 0 0 0
0 . 6 5 0 0 0 0 0 1 I O 0 0
0 . 4 5 0 0 0 0 0 0 1 O 0 0
0 . 0 9 0 0 0 0 0 0 5 2 0 0
0 . 4 5 0 0 1 0 0 0 0 0 0 0
0 . 0 2 0 0 0 1 2 3 0 2 1 O
0.06 1 O O 1 8 2 3 10 1 6
0.45 1 0 0 0 0 0 0 0 0 0
0.17 6 1 1 2 14 6 10 14 11 13
0 . 4 5 0 0 0 0 0 0 0 0 1 O
0 . 4 5 0 0 0 0 0 0 0 0 0
1
0 . 4 5 2 0 0 0 0 0 0 0 0 O
0 . 4 5 0 0 0 0 0 0 0 0 0 0
0 . 0 9 0 0 0 0 0 0 0 0 0 2
0 . 6 5 0 0 0 0 1 O 0 0 0 0
0.33 O 0 0 1 0 5 0 0 4 3
0.47 4 16 12 4 18 9 3 6 9 7
0 . 4 5 0 0 0 1 0 0 1 O 0 0
0.45 1 0 0 0 0 0 0 0 0 0
0 . 4 5 0 0 0 1
0.22 3 1 O 2
0 . 9 2 3 0 0 0
0 . 4 5 0 0 0 0
0 . 6 5 0 1 O 0
0.04 O O 1 4
0 . 0 0 0 0 0
0 .
0.15
0 .
0.05
0.16
0 .
0.31
0 .
0 .
0 .
0 .
0 .
0 .
0 .
0
O
0
0
0
8
1
0 0 0 0
1 1 O O
1 O 1 0
1 O 0 0
0 0 0 1
13 O 12 7
3 2 0 1 0
4 5 0 0 0 2 0 0 0 0 0
O O O 12 2 O 2 2 1
4 5 0 0 0 0 0 0 0 0 0
O O O 2 9 2 118 9
O O O 2 6 1 0 1 1 O
3 0 0 0 0 0 0 1 2 0 1
O 0 O O O 1 O11 1
3 8 0 0 0 0 3 0 0 1 1
1 4 0 0 0 0 1 4 0 0 0
2 0 0 0 0 0 5 2 0 0 2
4 5 0 0 0 0 0 7 0 0 0
4 5 0 0 0 0 1 O 0 0 0
4 5 0 0 0 0 1 O 0 0 0
4 5 0 0 0 0 0 0 0 3 0
3
0
0
0
0
0
0
0
1
7
0
11
0
O
0
1
1
0
5
0
0
0
0
0
0
2
0
2
0
3
0
0
0
O
O
1
1
5
0
0
12.5
1.7
2.1
2.5
0.8
0.4
2.9
1.3
4.2
17,l
0.4
38.3
0.4
0.4
0.8
0.4
1.3
0.8
6.3
43.8
0.8
0.4
0
1
1
0
O
4
0
0 0
O O
1 O
0 0
0 0
6 4
0 0
0.4
3.8
2.9
0.4
0.8
24.6
2.9
0
7
1
61
O
O
O
O
0
O
0
0
0
0
0 0
1 1
0 0
5 1
O O
0 1
2 1
0 0
0 0
0 0
0 0
0 0
0 0
1 0
0.8
11.7
0.4
26.3
8.3
2.1
6.7
2.1
2.1
3.8
2.9
0.4
0.4
1.7
1
12
0
0
160
ix IV. continued
Species
Hydnunz sp. 1
Hydnum sp. 2
Hygrocybe conica
Hygrocybe miniata
Hygrocybe sp. 1
Hygrophoropsis aurantiaca
Hygrophoropsis olida
Hygrophorus agathosmus
Hygrophorus bakerensis
Hygrophorus camarophyllus
Hygrophorus picea
Hygrophorus sp. 1
Hygrophorus spp.
Hypholoma capnoides
Hypholoma fasciculare
Inocybe calamistrata
Inocybe geophylla
Inocybe lacera vel aff
Inocybe lilacina
Inocybe mixtilis
Inocybe napipes vel aff
Inocybe ovatocystis
Inocybe sororia
Inocybe spp.
Kuehneromyces vernalis
Laccaria amethysteooccidentalis
Laccaria bicolor
Laccaria laccata
Lactarius luculentis
Lactarius pallescens
Lactarius pseudomucidus
Lactarius rubrilacteus
Lactarius rufus vel aff
Lactarius scrobiculatus
Lactarius spp.
Leccinum manzanitae
Lentaria spp.
Lepiota clypeolaria
Leptonia carnea
Leptonia sp.
Lycoperdonfoetidum
Lycoperdon molle
Lycoperdon perlalum
Lycoperdon pyrijorme
gs gn gk
Guild
m
m
1
1
1
s
1
m
m
m
m
m
m
w
w
m
m
m
m
m
m
m
m
m
w
m
(agd
0.45
0.33
0.44
0.49
0.45
0.45
0.2 1
0.34
0.07
0.45
0.42
0.45
0.19
0.13
0.09
0.45
0.07
0.07
0.30
0.79
0.45
0.02
0.53
0.00
0.4 1
0.49
is in ik ms mu mk os on ok 97 fre
Species
Guild
Lycoperdon sp.
Lyophyllurn decastes
Lyophyllum semitale
Marasmius androsaceus
Marasmius limosus vel aff
Marasnz ius salalis
Marasmius sp. 1
Marasmius spp.
Marcelleina sp.
Melanoleuca graminicola
Merulius tremellosus
Mollisia sp.
Mycena adonis
Mycena alcalina
Mycena alnicola
Mycena amabilissima
Mycena amicta
Mycena aurantiidisca
Mycena capillaripes
Mycena citrinomarginata
Mycena concolor vel aj9
Mycena elegantula
Mycena epipteïygia
Mycenaflavoalba
Mycena galopus
Mycena haetnatopus
Mycena inclinata
Mycena maculata
Mycena metata
Mycena oregonensis
Mycena paucilarnellata
Mycena sp. 8
Mycena pura
Mycena purpureofusca
Mycena rorida
Mycena sanguinolenta
Mycena sp. 1
Mycena sp. 2
Mycena sp. 3
Mycena sp. 4
Mycena sp. 5
Mycena sp. 6
Mycena spp.
Mycena strobilinoides
Mycena subcana vel aff
1
p
gs gn gk is in ik ms mn mk os on ok 97 freq
(a@)
0 . 4 5 0 0 6 0 0 0 0 0 1 O 0 0
2.9
Guild
p
gs gn gk
is in ik
k os on ok 97 freq
(a@)
Mycena tenerrima
Neournula pouchetii
Nidula niveotomentosa
Nolanea cetrata vel a#
Nolanea holoconiota vel a f
Nolanea sericea vel afi
Nolanea spp.
Oligoporus caesius
Oligoporus~agilis
Omphalina chlorocyanea
Omphalina ericetorum
Osteina obducta
Otidia alutacea vel a 8
Panellus stipticus
Paraeccilia sericeonitida var.
ligniphila
Pmcillz~satrotomentosus
Peziza spp.
Phaeolus schweinitzii
Phellinus sp.
Phellodon atratus
Phellodon melaleucus
Phellodon tomentosus
Pholiota sp.
Pholiota terrestris
Phylloporus rhodoxanthus
Plectania melastoma
Pluteus cervinus
Pluteus fiavofulgineus
Polyporus elegans
Polyporus hirtus
Psathyrella longistriata
Psathyrella spp.
Pseudohydnum gelatinosum
Pseudoplectania melaena
Pseudoplectania nigrella
Psilocybe corneipes vel a f
Psilocybe montana
Ramaria sp.
RickenellaJibula
Russula fragilis vel a#
Russula adusta vel a#
Russula bicolor
Russula brevipes
Russula emetica vel a#
1
Z
w
s
s
s
s
w
w
1
a
w
1
w
w
m
1
w
w
m
m
m
w
w
m
1
w
w
w
s
1
1
w
1
1
1
1
m
1
m
m
m
m
m
0 . 4 5 0 0 0 0 0 0 0 1 0
0 . 4 5 0 0 0 0 0 0 0 2 0
0 . 7 6 0 5 1 0 0 9 0 0 3
0 . 4 0 0 0 0 1 O 1 1 3 0
0 . 0 1 0 0 0 4 7 3 1 4 6
0.45 O O O O O 1 O O O
0.28 O 1 1 6 1 10 1 5 5
0 . 4 5 0 0 0 0 0 0 1 O 0
0 . 1 9 0 0 0 0 0 0 1 1 0
0.092 1 0 0 0 0 0 0 0
1 . 0 0 0 6 2 3 1 2 3 1 2
0 . 4 5 0 0 0 0 0 0 0 0 1
0 . 0 7 0 0 0 2 0 4 1 0 3
0 . 6 3 0 0 0 0 0 0 0 1 0
0.45 1 0 0 0 0 0 0 0 0
0
0
O
1
4
O
5
0
1
0
2
O
7
2
0
0 0
0 0
0 1
2 0
4 9
O O
2 1
0 0
O 0
0 0
2 2
0 0
5 2
0 0
0 0
0 . 3 5 0 0 0 2 0 1 0 0 1 1 0
0 . 1 2 0 0 0 0 0 0 0 0 0 3 1
0 . 4 0 1 0 0 1 0 1 2 0 0 0 0
0 . 0 9 0 0 0 0 1 2 0 0 0 0 0
0 . 4 4 0 0 0 0 7 1 O 1 4 1 O
0 . 0 7 0 0 0 0 1 1 O 0 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 0 0 1
0 . 0 2 0 0 1 0 0 0 0 0 0 1 1
0.45 O 1 O O O O O O 0 O O
0 . 4 5 0 0 0 0 0 0 0 1 0 0 0
0 . 6 5 0 1 0 1 6 0 0 2 1 2 0
0 . 1 5 0 0 0 1 0 0 1 1 0 2 1
0 . 4 5 0 0 0 1 0 0 0 0 0 0 0
0 . 2 5 0 0 0 1 1 0 1 0 3 0 0
0 . 0 9 0 0 0 1 2 0 0 0 0 0 0
0 . 4 5 0 0 0 0 0 0 1 0 0 4 0
0 . 6 5 0 0 0 1 O 0 0 0 1 O 0
0.07 O O O O 3 6 4 1 3 3 4 9
0 . 6 3 0 0 0 0 0 0 0 0 1 2 0
0 . 6 5 0 0 0 1 O 0 0 I O 0 0
0 . 4 5 0 0 0 1 0 0 0 0 0 0 0
0 . 0 4 4 9 1 1 3 0 0 1 1 0 1
0.09 O O O O O O O O 13 9 4
0 . 0 8 3 2 0 0 0 0 0 0 0 0 0
0 . 0 7 0 0 0 0 0 0 0 0 0 1 O
0 . 4 5 0 0 0 0 0 0 0 0 0 3 0
0.25 O O O 1 O 2 O O O 5 1
0.01 O O O 1 1 1 O 5 11 1 0 1 0
0 . 4 5 0 0 0 0 0 1 0 0 1 O 0
0
O
0
0
0
0
O
1
O
0
0
O
0
0
0
0
0
1
0
0
0
0
4
0
1
0
O
14
0
0.4
0.8
7.9
3.8
17.5
0.4
15.8
0.4
1.3
1.3
10.8
0.4
10.0
1.3
0.4
2.1
1.7
2.1
1.3
5.8
0.8
0.4
1.7
0.4
0.4
5.4
2.5
0.4
2.5
1.3
2.1
0.8
17.9
1.3
0.8
0.4
8.8
12.5
2.1
0.8
1.3
3.8
22.1
0.8
Gui1
Russula marei
Russula occidentalis vel aff
Russula placita
Russula sp. 1
Russula spp.
Russula xerampalina
Sarcosphaera crassa
Spathularia flavida
Stereum hirsuium vel afl
Stereum sanguinolentum
Strobilurus albipilatus
Strobilurus trullisatus
Stropharia ambigua
Suillus borealis vel afl
Suillus granulatus
Suillus lakei
Suillus luteus
Suillus spp.
Suillus subolivaceous
Thelephora terrestris
Trametes versicolor
Trichaptum abietinum
Tricholoma atroviolaceum
TrichoZomaflavovirens
Tricholoma inamoenum
Trichohma magnivelare
Tricholoma pessundatum
Tricholoma portentosum
Tricholoma saponaceum
Tricholoma sejunctum
Tricholoma sp. 1
Tricholoma spp.
Tricholoma sulphureum
Tricholoma virgatum
Tricholoma zelleri
Trichopilusplebiodes
Tyromyces chioneus
Xeromphalina campanella
Xeromphalina.fulvipes
Xeromphalina spp.
Xylaria hypoxylon
m
m
m
m
m
m
1
1
w
w
s
1
1
m
m
nl
m
m
m
m
w
w
m
m
m
m
m
m
m
m
m
m
m
m
m
1
w
w
s
s
w
gs gn gk
is in ik ms mn mk os on ok 97 freq
0.32 O O O O 1 3 O O 1 O
0 . 6 3 0 0 0 2 0 0 0 0 0 0
0.00 O O O 5 3 2 0 2 4 1 1 6
0 . 0 7 0 0 0 0 0 0 0 0 0
1
0.00 O O O 12 18 12 3 10 5 14
0 . 0 2 0 0 0 3 0 5 0 0 5 7
0 . 4 5 0 0 0 0 0 0 0 0 0 1
0 . 4 5 0 0 0 0 0 0 0 1 0 0
0 . 4 5 0 0 0 1 0 0 2 0 0 1
0.45 1 0 0 0 0 0 0 0 0 0
0 . 6 3 0 0 0 0 2 0 0 0 0 1
0 . 0 2 0 0 0 2 3 2 3 0 2 5
0 . 4 5 2 0 0 0 0 0 0 0 0 0
0 . 4 5 2 0 0 0 0 0 0 0 0 0
0.80 1 O 0 0 0 1 O 0 0 1
0.891 0 0 3 0 0 0 0 4 3
0 . 4 5 2 0 0 0 0 0 0 0 0
1
0 . 9 1 2 0 0 0 1 0 0 0 4
1
0.45 1 0 0 0 0 0 0 0 0 0
0 . 3 4 0 0 0 7 0 0 3 0 1 4
0 . 4 5 0 0 0 0 0 0 1 0 0 0
0.20 1 0 0 4 0 0 4 2 0 4
0 . 0 9 0 0 0 0 0 0 0 0 0 0
0 . 0 8 0 0 0 0 2 2 0 5 3 0
0 . 3 4 0 0 0 0 0 0 0 0 1 1
0 . 4 5 0 0 0 0 0 0 0 2 0 0
0 . 4 5 0 0 0 0 0 1 O 0 0 0
0 . 6 0 0 0 0 0 0 0 0 1 0 4
0 . 1 1 0 0 0 1 0 3 0 2 3 1
0 . 0 8 0 0 0 0 0 2 1 4 4 0
0 . 4 5 0 0 0 0 0 0 1 O 0 1
0 . 0 0 0 0 0 1 1 2 4 5 7 2
0 . 0 8 0 0 0 0 0 0 0 0 0 0
0 . 4 5 0 0 0 0 0 0 0 0 1 O
0 . 4 5 0 0 0 0 0 8 0 0 5 0
0 . 4 5 0 0 0 0 0 1 O 0 0 0
0.65 1 O 0 0 1 O 0 0 0 0
0 . 1 7 0 3 1 7 3 5 2 0 1 3
0 . 0 0 0 0 0 1 O 1 3 2 2 1
0 . 4 5 0 0 0 1 0 0 0 0 0 0
0 . 4 5 0 0 0 3 0 0 0 0 0 0
1
0
12
O
11
4
0
0
0
0
O
2
0
0
O
1
O
0
0
0
0
1
2
2
1
0
0
0
4
2
0
4
3
0
0
0
0
0
1
0
0
O
1
10
1
12
9
0
0
0
0
0
4
0
0
0
0
0
2
0
1
0
O
1
1
0
0
0
0
3
0
0
1
6
0
0
0
0
0
1
0
0
2.5
1.3
30.4
0.8
40.4
13.8
0.4
0.4
1.7
0.4
1.3
9.6
0.8
0.8
1.3
5.0
1.3
4.2
0.4
6.7
0.4
6.7
1.3
6.3
1.3
0.8
0.4
2.1
7.1
5.4
0.8
11.3
3.8
0.4
5.4
0.4
0.8
10.4
5.0
0.4
1.3
ix V - SPLUS prograrns
SPLUS function "super.po~wow'~
function(loops, std, d)
# loops is the number of iterations
# std is the standard deviation to be used for data simulation
# d is the vector of data on which simulation is based (
.mns <- c(mean(d[1:3]), mean(d[4:6]), mean(d[7:9]), mean(d[l 0: 121))
sig <- array(0, c(1 0, 3))
blcks <- vector(mode = "integer7', 10)
for(i in 1:10) (
if(i < 9) {
n<-i+2
1
else n <- n -t 5
age <- c(rep(l:4, each = n))
blcks[(i)] <- n
for6 in 1:loops) (
simdat <- rnorm(n * 4, mean = rep(grp.mns,
each = n), std)
p <- summary(aov(simdat age))[l, 51
if(p < 0.05)
sig[i, 11 <- sig[i, 11 + 1
if(p < 0.1)
sig[i, 21 <- sig[i, 21 + 1
if@ < 0. 15)
sig[i, 33 <- sig[i, 31 + 1
-
1
1
power <- sig/loops
return(blcks, power)
1
V - SPLUS programs
SPLUS program "mu1ti.powwow"
# nurnber of iterations
loops<- 1O00
# standard deviation to be used for data simulation
std<-sqrt(58.4722)
# vector of data on which simulation is based
d<-richness.df$Richness
# type one error rate of post-hoc comparisons
alph<-. 10
# nurnber of replicates for simulated data
n<-4
grp.mm <- c(mean(d[1:3]),mean(d[4:6]),mean(d[7 :9]),mean(d[10: 121))
sigm <- matrix(0,6,1)
age <- rep(~("G,"I","M,~'0"), each = n)
forCj in 1:loops) (
simdat <- rnom(n * 4, mean = rep(grp.mns9 each = n), std)
simdat.df<-data.fi-ame(age,simdat)
aov.fit<-aov(simdat age,simdat.df)
mc <- multicomp(aov.fit,alpha=alph)$table[, 3 :4]
mt <- trunc( ((me[, 11 * mc[, 21) / (abs (mc[, 11 * mc[, 2])))+0.1)
sigm <- sigm + mt
-
1
Ix V - SPLUS programs
SPLUS program "effect.sizefl
# number of iterations
Ioops<- 1O0
#
standard deviation to be used for data simulation
std<-sqrt(5 8.4722)
# type one error rate of post-hoc comparisons
alph<-. 10
#
number of replicates for simulated data
n<-3
el<-30
e2<-31
e3<-32
grp.mm <- c(0,el ,e2,e3)
names(grp.mns)<-c(0,"El ","E2","E3")
sigm <- may(O,6)
age <- rep(c("0,"E 1","E2","E3"), each = n)
for (j in 1:loops) (
simdat <- rnorrn(n * 4, mean = rep(grp.mns, each = n), std)
simdat.df <- data.frame(age,simdat)
aov.fit <- aov(simdat
age, simdat.df)
mc <- multicomp(aov.fit)$table[, 3:4]
mt <- tninc((mc[, 11 * mc[, 2])/abs(mc[, 11 * mc[, 2])+0.1)
sigrn <- sigm+ mt
-
1
names(sigm) <- names(mt)
sigrn <- sigm/loops
sigm