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Norsk tittel: Hvor ofte og hvor lenge er nok?
Langtidsstudium (2003-2022) av variasjon i
forekomst, artsmangfold og fruktifisering av
beitemarksopp i en naturbeitemark på Stord.
How long and how frequent is sufficient?
Long time study (2003-2022) of variation in
occurrence, species richness and fructification
of grassland-fungi in a semi-natural grass-
land in South-Western Norway. Agarica 2023
vol. 43: 55-83.
KEYWORDS
Grassland-fungi, biodiversity, frequency of
survey, monitoring, seasonality, longtime study,
fructification
NØKKELORD
Beitemarksopp, biomangfold, frekvensen av
besøk, overvåkning, sesongmessige forhold,
langtidsstudium, fruktifisering
SAMMENDRAG
Hovaneset på Stord har vært inventert for
beitemarksopp i 20 år, fra 2003 og frem til i
dag (2022). Dette har resultert i funn av 92
forskjellige arter etter totalt 180 besøk. Endring
i frekvensen av inventering til ca. ukentlige
besøk i soppsesongen i årene mellom 2010
og 2022 resulterte i stor økning av artsmang-
foldet også av rødlistearter. I dette tidsrommet
mer enn doblet det totale artsantallet seg, og
tallet på rødlistearter mer enn tredoblet seg, noe
som tydelig viser den store effekten i endring
av inventeringsfrekvens. Inventeringen viser
også klart at antall funn ved en inventering
varierer stort fra år til å og fra uke til uke. I et
optimalt år for beitemarksopp kan en totalt
finne i overkant av 70% av artene som vokser
der ved en ukentlig inventering hele sesongen,
mens et dårligere år gir maksimalt funn av
40% av artene som vokser der. Inventeringen
viser også at tidspunkt for besøk i et bestemt
år er avgjørende for hvor mange arter som
blir funnet. I et optimalt år kan en ved best
mulig treff med tidspunkt, finne oppunder
50% av de artene som vokser der på ett besøk,
men mest sannsynlig vil det ligge på rundt
35%. Dette viser ganske tydelig at besøk
gjennom hele sesongen er nødvendig for å få
oversikt over alle soppene som vokser her.
Langtidsserien viser også at mange beite-
marksopp har en svært uregelmessig frukti-
fisering fra år til år, noe som særlig gjelder
arter innenfor slekten Clavaria og Entoloma.
Bare ti arter ble funnet hvert år i løpet av 20
år med inventering, der de fleste var voks-
sopper (Hygrocybe s.l.), mens 14 arter bare
ble funnet i ett av årene i inventeringen.
Tidspunkt for fruktifisering for mange arter
synes også å ha endret seg i løpet av de 20
årene undersøkelsen har pågått. Uken med
flest funn i 2021/22 er ca. 3-4 uker seinere
enn den var i 2013/14. Dette er særlig tydelig
for slekten Entoloma som normalt har en
tidlig og begrenset fruktifiseringsperiode.
AGARICA vol. 43 55
How long and how frequent is sufficient? Long time study
(2003-2022) of variation in occurrence, species richness and
fructification of grassland-fungi in a semi-natural grassland
in South-Western Norway
Per Fadnes
Faculty of Education, Science and Sport, Institute for Sport, Nutrition andNatural
Sciences, Western Norway University of Applied Sciences, campus Stord
Per Fadnes
ABSTRACT
The semi natural grassland described in this
paper, Hovaneset in Stord municipality, South-
Western Norway, has been examined for grass-
land fungi for 20 years, from 2003 until today
(2022). This has resulted in the discovery of
92 different species after a total of 180 visits
to the locality. Change in the frequency of
survey to approximately a weekly visit during
the seasons from 2010 resulted in a large
increase in species diversity, also of red-listed
species. During these last 13 years of survey,
the total number of species more than doubled,
and the number of red-listed species more
than tripled, which clearly shows the huge
effect in changing the frequency of visits to a
visit once a week during the season. The
survey also clearly shows that the number of
finds during a visit varies greatly from year
to year and from week to week. In an optimal
year for grassland fungi, around 70% of the
species that grow there can be found in a
weekly survey throughout the season, while
a less good year gives a maximum of 40%
finds. The survey also shows that the time of
visit in a particular year is decisive for how
many species are found in one visit. In an
optimal year for grassland fungi, at the best
possible match with time of survey, you can
find up to 50% of the species growing there
in one visit, but more likely it will be around
35%. It therefore shows very clearly that
visits during the whole season is necessary to
get hold on all fungi growing here.
The survey also shows that many grass-
land fungi have very irregular fructification
from one year to the other, which particularly
applies to species within the genera Clavaria
and Entoloma. Only ten species were found
each year during the survey, most of which
were waxcaps (Hygrocybe s.l.), while 14
species were only found in one of the years
of survey. The time of fruiting also seems to
have changed during the 20 years the survey
has been ongoing. The week with the highest
number of species recorded in 2021/22 is
approximately 3-4 weeks later than it was in
2013/14. This is particularly clear for the
genus Entoloma, which normally has an early
and limited fruiting period.
INTRODUCTION
Grassland fungi is a concept normally restric-
ted to a few genera and families of fungi that
primarily grow in unfertilized pastures, often
described as semi-natural grasslands. This is
in accordance with several authors (Nitare
1988, Griffith et. al. 2013). Concepts like
“grassland fungi” and “waxcap-grasslands”
(seminatural grasslands rich in Hygrocybe
species) were given attention as far as back
in the 18th century (Griffith et. al. 2013).
Grassland fungi, used as described above,
consist of a taxonomical diverse group of
fungi with similar ecological preferences.
These are Hygrocybe (s. l.) (waxcaps), Ento-
loma (redspores), Camarophyllopsis (s. l.),
Clavaria, Clavulinopsis, Ramariopsis, Pseudo-
tricholoma (Porpoloma), Dermoloma and
Geoglossaceae (Geoglossum (s. l.) and Tri-
choglossum) (Boertmann 2010, Nitare 1988,
Noordeloos 1992, Jordal 1997, McHugh et.
al. 2001, Griffith et al. 2013), and the former
member of Geoglossaceae, Microglossum,
which is now placed in the Leotimycetes
(Ohenoja et al. 2010, Hustad et al. 2013). In
Norway and Sweden around 150-160 different
species of grasslandfungi have so far been
registered (Nitare 1988, Jordal 2011, 2013).
In the United Kingdom, these fungi are
also often referred to as CHEGD-fungi, which
is an acronym of most of the names listed
above (Griffith et. al. 20013). In accordance
with Evans (2003) and Griffith et. al. (2013),
the group consists of 180-200 species in the
UK.
Many grassland fungi are rare and grow
only in unfertilized pastures and meadows,
which are the main habitats for these fungi.
Termination of grazing leads to invading of
56 AGARICA vol. 43
Per Fadnes
trees, and fertilization are favoring fungi and
plants which prefer more nutrition.
Overgrowing can, to some extent, be rever-
sed by reintroducing grazing animals, since
the grassland fungi might have survived and
start fructification again, while fertilization
seems to be an irreversible process wiping out
most of the species (Arnolds 1989, Vesterholt
and Knudsen 1990). Some species prefer
calcareous soil, but we can also find many
rare and red-listed species growing on more
poor and acidic soil, especially if the locality
has long continuity. Many of these species
are therefore good indicators of valuable
grasslands, probably better indicators than
vascular plants (Rald 1985, Öster 2008).
Different species of grassland fungi often
occur in clusters on the same locality.
The ecology of grassland fungi is to date
not clearly understood. Are they saprotrophes,
decaying dead materials, or do they have
biotrophic relationships with herbs or gras-
ses? New results indicate that hyphae from
several waxcaps do colonize plant roots,
indicating a biotrophic lifestyle (Seitzman et
al. 2011, Halbwachs et al. 2013).
Very few longtime studies of fungal fructi-
fication have been performed, and those that
exist are relatively old. The existing surveys
of fungi have been criticized for being too
short and random not hitting the right time
for the best results (Halme & Kotiaho 2012).
Generally, there is a clear correlation
between the number of species found at one
location and the intensity of visits (Parker-
Rhodes 1955, Watling 1995). According to
Parker-Rhodes, you will seldom find more
than 25% of the species that grow at a speci-
fic site in one visit. Orton (1986) concluded
that the data from at least 10 years of investi-
gation is needed to get an accurate picture of
the species growing in a locality. Newton et
al. (2003) visited five localities with old past-
ures between 10 and 16 times over a period
of three years (1999-2001). Species diversity
increased with each visit, and they concluded
that at least 16 visits to a certain site is neces-
sary to get a good picture of fungal diversity
at the locality. Straatsma et al. (2001) surveyed
a forest area for 21 years from May to Sep-
tember. New species appeared every year,
but only eight species occurred every year.
The most species rich localities in Norway
until now had 71 different species recorded,
including 32 waxcaps (Fadnes 2014), in
Sweden one locality had 76 species of which
33 were waxcaps (Phil 1992), and in UK the
most species rich locality had 78 species, of
which 34 were waxcaps (Griffith et. al. 2013).
Increased survey of grasslands in SW-
Norway the last decade has led to a great
increase in the number of grassland fungi
known, and today between 60 and 130 species,
including many rare and red-listed species
are found in the municipalities situated here
(Brandrud et al. 2021). It has previously been
shown that SW-Norway has a particularly
rich and varied occurrence of earthtongues
(Fadnes 2011), and for most of the other genera
the numbers are high (Artsdatabanken 2022).
SW-Norway therefore appears to be a “hot
spot” for localities with rare and red-listed
grassland fungi. After around 20 years of
survey of old grasslands in SW-Norway, it is
apparent that there are many factors in addi-
tion to fertilizing, continuity, geology and
soil, that have an influence on which species
we actually find on a particular locality at a
certain time (Fadnes 2014).
Results from the first 11 years of survey
have previously been published (Fadnes
2014). The number of species presented in
this study differs to a degree from those
presented here. This is due to several new
finds the last nine years, some changes in red
listed species (Brandrud et.al. 2021), and that
some species, especially species of Entoloma,
have been subjected to molecular studies.
In 2003, when this study started, no grass-
landfungi were known from the location
AGARICA vol. 43 57
Per Fadnes
studied. However, every year of survey grad-
ually increased species diversity. Some new
fungi appeared every year, and some found
the last year did not reappear. Such a long-
term study of a semi-natural grassland has
probably not been carried out before.
The aim of this study will focus on the
variations in diversity and fruiting of grass-
land fungi through a 20-year survey with
almost weekly visits from July to November
of this particular grassland in SW-Norway,
described in detail below.
MATERIALS AND METHODS
Sampling procedure
The locality that is the main basis for this
article, Hovaneset (a more detailed description
follows under), was investigated from 2003
to 2022. From 2003 to 2009 the frequency of
visits was random during the season.
The intensity ranged from two to five visits
every year. In 2010, the number of visits in-
creased to ten, and the last 12 years (2011-22)
the locality was visited almost every week
from late July to November, totaling 10 to 15
times a season. The total number of visits to
the locality accounts to around 180 performed
during the last 20 years. A path of about 2 km
has been followed, and in this way, the whole
area has been systematically surveyed on each
visit. Every find was recorded at each visit,
and for rare and red-listed species coordinates
were noted. Dried specimens of red-listed
species were sent to the fungaria of Natural
History Museum (O), (University of Oslo
2023), all other finds were registered at the
Norwegian Species Observation Service
(Artsdatabanken 2022). Both datasets are
available for downloading at GBIF-org (2023).
Fungi mainly growing in nutritious soil
and manure were noted, but not included in
this survey. Some species had to be examined
microscopically. Fresh material of the fungus
was studied after soaking in water using a
LEICA DM750 microscope.
Based on the number of visits to the locat-
ion, the survey can be divided into two periods:
Period 1. The first seven years (2003-2009)
with a more random number of visits, ranging
from two to five.
Period 2. The last 13 years (2010-2022)
with a more systematic survey with a visit
almost every week during the season (10-15
visits/year).
Description of the investigated locality
Hovaneset, the location studied here, is a
peninsula in the Hardangerfjord just north of
the village Leirvik on the east side of the
island Stord and about 70 km south of Bergen
defined by the coordinates 32VLM0534
(Figure 1). It is a large natural grassland of
approximately 5 ha with some small marshy
areas and is grazed by sheep (Figure 2).
Probably some fertilizer has been provided
earlier on the upper flattest part of the area.
Therefore, the prevalence of grassland fungi
is mainly highest in the rugged peripheral
zones of the peninsula (Figure 2).
Calcareous bedrocks surround the pasture
where you can find plants like Lysimachia
minima (EN), Sagina nodosa and Erophila
verna in addition to typical mountain plants
like Saxifraga oppositifolia (NT) and S.
aizoides. In the pasture we find a variety of
demanding grass and herbs like Cynosurus
cristatus, Briza media (NT), Linum cathari-
ticum and Orchis mascula. The soil quality
varies from thin soil with Pimpinella saxi-
fraga and Aira praecox to more humid areas
with Carex hostiana and C. flacca (NT).
Here we also find the rare and red-listed
species Isolepis setacea (EN).
Seen from archaeological traces, the area
has a continuous use by man from the Bronze
Age up to present (Figure 2c) At the end of
the last ice age (10000 years ago) the area
was mainly under the sea level because the
land had been pressed down by the large
icecap. Later, in the Bronze Age, Hovaneset
58 AGARICA vol. 43
Per Fadnes
AGARICA vol. 43 59
Figure 1. Map showing the location of the locality Hovaneset in Western Norway
Figure 2. a. Arial photo of the locality Hovaneset (Norgeskart), b. The peninsula Hovaneset seen
from the south, c. Archaeological traces (burial mound) going back to the Bronze age, d. Part of the
grassland from Hovaneset. Photo P. Fadnes.
Per Fadnes
was an island, and as the mainland gradually
raised from the pressure of the ice, it became
a peninsula, which is how it looks today.
After 20 years of investigation, a total of
92 different species of grassland fungi have
been discovered on Hovaneset, where 44 are
on the Norwegian red list and 21 of them are
vulnerable (VU) or endangered (EN) while
five are defined as data deficient (DD) (Table 1).
In addition, more than 15 species not belong-
ing to the grasslandfungi are found in the area
(Arrhenia spp., Agaricus sp., Mycena spp,
Rickenella sp., Panaeolus spp., Cystoderma sp.,
Coprinopsis sp., and Stropharia sp.), making
the total number of fungal species more than
100. Some of them are also relatively rare in
Norway like Arrhenia rickenii, and Agaricus
phorphyrocephalus).
With a totality of 180 visits during the last
20 years, the locality is probably the best
investigated semi natural grassland in Norway
and the most species rich known today (Fadnes
2014, 2021). Most of the species from Hov-
neset are well documented on the authors
webpage “Grasslandfungi.no” (Fadnes 2023).
Molecular studies
Molecular studies were performed with
standard procedures. DNA extraction, PCR
amplification and sequencing were done in
accordance with the Norwegian Barcode of
Life Network (NorBOL) as part of BOLD by
the University of Oslo or following Alvalab
(Alvarado et al. 2012) or Dima et al. (2016).
For BOLD methods see Ratnasingham &
Hebert (2007, 2013).
For the DNA extraction and PCR reactions
the Phire® Plant Direct PCR Kit (Thermo
Scientific, USA) was used following the
recommendations of the manufacturer, but in
some cases the NucleoSpin Plant II Kit (Mac-
herey-Nagel, Düren, Germany) was also used.
The ITS region of the nrDNA was amplified
with the primer pairs ITS1F/ITS4 (White et
al. 1990, Gardes & Bruns 1993). PCR proto-
cols were followed Papp & Dima (2018). The
amplicons were sequenced at LGC Genomics
60 AGARICA vol. 43
Fgure 3. Changing in diversity of grassland fungi during 20 years of survey.
Per Fadnes
(Berlin, Germany) with the same primers used
in PCR reactions. The electropherograms
were checked, assembled, and edited with
the CodonCodeAligner package (CodonCode
Corporation, Centerville, MA, U.S.A).
RESULTS
Change in number, diversity and fructi-
fying of grassland fungi during 20 years of
survey.
The number of visits to the locality was
sparse from 2003 to 2009 (period 1), during
the season from July until November, varying
between two and five random visits per year.
The yearly records of different species varied
from 20 to 26, even if the accumulated num-
ber of different species found during the first
seven years increased from 20 to 41 (Figure 3).
The same effect was seen among the fungi
listed in the Norwegian red list (Brandrud et.
al. 2021). The number of red-listed species
found in Period 1 varied between four and
seven each year, and the accumulated number
found during Period 1 increased to 13 in
2009 (Figure 3).
In period 2, from 2010 until 2022, the
investigation of the locality increased to a
visit almost once a week during the season
from end of July to medio November. This
change in method had a profound impact on
the number of fungi recorded. It resulted in a
doubling in number of species found in one
year from 2009 to 2010, from 23 to 45 species.
This also increased the total (accumulated)
number of species found at the locality to 54
in 2010. A total of 21 of 24 red-listed species
were found that year, showing the profound
effect of the increased survey activity from
one year to the next, especially on red-listed
species which normally are rarer than the
others.
The next 12 years with intensive survey
increased the total number of species almost
every year and reached the substantial number
of 92 species accumulated in 2021 (Figure 3).
The number of red-listed species also
increased gradually during these years, with
the numbers starting to level out in 2015,
ending up at 44 species in 2021, which
constitute almost half of the total number of
AGARICA vol. 43 61
Figure 4. Occurrence of different groups of grassland fungi the 13 last years of survey (Period 2).
Per Fadnes
species. The rise in number of species in
2021 is partly due to the molecular study of
several Entoloma species which could have
occurred earlier but have not been registered
before (Table 3). Moreover, some species of
waxcaps and Clavariaceae were identified by
molecular studies in 2021 and contributed to
the rise in number of species (Table 3). Since
the change to a more intensive survey in
2010, the number of red-listed species more
than tripled, and the total number of species
has more than doubled at the end of the
survey (Figure 3).
Figure 4 shows the changing in number of
the different groups of grassland fungi in the
years from 2010 until 2022. The waxcaps
show a stable number of species from one
year to the other. Species of Entoloma varies
a lot from one year to the other and had two
years with relatively high numbers, 2014 and
2021. The number of earthtongues and mem-
bers of the Clavariaceae show like the waxcaps
a relatively stable number from year to year.
The exception was 2018 which had an extra-
ordinary dry summer (NMI 2022), resulting
in a dramatic drop in the number of finds of
all groups compared with all years back to
2010, especially for Clavariaceae which had
only one find in 2018. This shows the clear
effect of weather on the number of records in
a given year. However, much unexpected was
the occurrence of tree new species that year,
all on the Norwegian red list. The most sur-
prising species was Hygrocybe spadicea,
occurring for the first time, together with
Entoloma neglectum and E. ameides. H.
spadicea is a rare, southerly species which
probably prefers warm summers. E. ameides
is a species also occurring on many other
locations this year. The year 2014 also had a
relatively warm summer in Western Norway
but not as dry as in 2018 (NMI 2022), and
was as shown, one of the most species rich
years in the survey (Figure 3) together with
2021.
62 AGARICA vol. 43
Figure 5. Accumulated finds of different groups of grassland fungi from Period 1 to Period 2.
Per Fadnes
Figure 5 shows the accumulated number of
the different groups of grassland fungi from
2009 to 2022. The year 2009 represents the
accumulated number for the first seven years
of survey (Period 1). This shows that waxcaps
constituted the most abundant group in Period
1 with as many as 21 species where six are
redlisted. This number raised significantly
already the first year of Period 2, and later to
34 species in 2021, which is the final number
of waxcaps. Among these are 13 on the Nor-
wegian red list (Brandrud et al. 2021) (Table 1).
The genus Entoloma has a slower develop-
ment in the number of species found, starting
with only seven species from Period 1, but
starting to increase already the first years of
Period 2 and ending at 29 species in 2021
(Figure 5), with as many as 13 species on the
red list (Brandrud et al. 2021).
Most of the earthtongues (s.l.) were already
found in Period 1, with eight different species,
and increasing to 11 as late as 2021. The
earthtongues from the South-Western Norway
are presented in an earlier article (Fadnes
2011).
Few species (5) from the family Clavaria-
ceae were found during Period 1 but showed
a significant rise in number in 2013/14 adding
seldom species like Clavaria tenuipes and C.
flavipes to the genus. Especially many of the
club fungi in the genus Clacvaria have a very
irregular fructification from year to year which
can explain this development.
Figure 6 shows the appearance of different
species during 20 years of survey. As many as
14 species have only appeared once during
the survey period: Pseudotricholoma meta-
podium, Clavaria tenuipes, Clavulinopsis
umbrinella, Entoloma ameides, E. neglectum,
E. indutoides, Hygrocybe spadicea, H. auran-
tiosplendens, H. coccineocrenata, Geoglos-
sum cookeanum. This also includes some
AGARICA vol. 43 63
Figure 6. Number of years with number of different species.
Per Fadnes
new species identified by molecular study in
2021/22: Entoloma violaceoviride, E. allos-
permum, E. carneogriseum, Hygrocybe flavi-
pesoides and Lamelloclavaria petersenii.
Some Entoloma species could of course have
occurred earlier without being discovered,
since some of them have just been described.
Only 10 species were registered every single
year of the survey. They were mostly rela-
tively common species and mainly waxcaps:
Geoglossum fallax, Hygrocybe chlorophana,
H. coccinea, H. conica, Cuphophyllus praten-
sis, C. virgineus, C. colemannianus, C. russio-
coriaceus, Gliophorus psittacinus and G.
laetus.
Figure 6 also shows that 56 different species
are only found between one and ten years,
while 36 species were found between 11 and
20 years of the survey (se also Table 1).
All species in the location but one (91)
were found during Period 2, while only 41
species were found in Period 1. This shows
that a more intensive survey with almost a
visit every week in the season results in more
finds, especially of more rare species.
If we only look at period 2, a total of 22
different species were found every year, and
13 species were found only once during this
period, again indicating that these fungi are
rare and/or have a very irregular fructification.
Table 2 shows the number of different
species found each year the last ten years. It
varies from 39 to 67 species and constitute
from 42 to 73% of the total number (92).
These high numbers presuppose an intense
survey of the location during the whole season.
Seasonal variation of grasslandfungi
Based on own experiences grasslandfungi
fructify at different times during the season
depending on latitude. In the end of July and
beginning of August the fructification in the
north of Norway is optimal especially among
Entoloma species. At that time there is almost
nothing in the south-western part of Norway.
In the UK the best time to look for grassland-
64 AGARICA vol. 43
Figure 7. Weekly finds of grassland fungi in some selected years in Period 2. (July: 29-30,
August: 31-34, September: 35-39, October: 40-43, November: 44-46.)
Per Fadnes
fungi is in October and November, showing
the effect of latitude. The seasonal variation
among the different genera and species of
grasslandfungi is also great. Different species
are fructifying to different time of the season
resulting in different numbers from one week
to the other.
Figure 7 shows the variation of finds of
different species from week to week for six
different years. The season in SW-Norway
normally starts in the beginning of September
with relative few numbers of finds, but the
number increases as September passes and
October begins. However, it seems that the
highest peak of different fungi has changed
from 2013 to 2022. The last years the main
peak has been in the beginning to middle of
October, which is a change in several weeks
from 2013 (Figure 7). This is particular
evident for the genus Entoloma which is
normally an early fructifying genus, but the
last years have fructified three to four weeks
later compared to 2013-2014 (Figure 8). This
figure also shows the great variation of finds
of different Entoloma species between years.
The total number is kept relatively high
over a longer period mostly due to the waxcaps
which fructify over a longer period of time
and where there also are some early and late
fructifying species (Figure 10). Many waxcaps
were found during the whole season like the
more common species Hygrocybe chloro-
phana, H. conica and the more seldom H.
quieta. The early fructifying species are species
like Hygrocybe helobia, H. acutoconica, H.
intermedia and Neohygrocybe ingrata. None
of those were found in the late visits to the
locality. The late fructifying species like
Hygrocybe splendidissima, H. punicea and
Cuphophyllus russocoriaceus were not found
in the start of the season but started to show
up in the middle of September or later. The
AGARICA vol. 43 65
Figure 8. Weekly finds of Entoloma species for four selected years in Period 2.
Per Fadnes
latest fungi to show up in the location was
always Cuphophyllus fornicatus. As shown
in figure 7 the start of the season has shifted
the last years, which will also be the case for
many of the species shown in Figure 10.
The high number of fungi found in a
particular year is dependent on several visits
which is the case in this study. However,
figure 7 shows a great variation in finds
depending on the time of visit. Normally you
only visit a locality a few times during the
season, and the figure shows that you should
be lucky to hit the most perfect week con-
cerning the number of different species you
can expect to find.
Table 2 gives you a picture of this. The
maximal number of different species you
will find, provided you choose the optimal
week, varies between 18 and 47 in the years
2013 to 2022. The most probable number is
between 30 and 40 of a total number of 92
species. This makes the maximal percentage
of different species found in one visit in this
survey around 50% (2021), but the most
probable number is between 34% and 38%.
Figure 7 also shows the dramatic effect of
high and stabile summer temperatures and
little precipitation on the number of collected
fungi, which was the case in 2018.
Survey statistic
Figure 9 shows the connection between the
accumulating number of fungi and the number
of visits during 20 years of survey. It seems
like the accumulated number of species found
correlate very well with the number of visits
until year 2009 (Figure 9). Then with an
increased number of visits the number of
species still increases until 2014 and around
90 visits and 78 different species, when the
number seems to even out. A further increase
of visits from 2014 still adds some new
species and ended in 2021 on 92 different
species. So even if the number of new species
doesn’t follow the number of visits from
2015, the intense survey still added 12 new
species to the total number during these last
eight years, which constitute 13% of the total
number of species, and the last 90 visits were
therefore not in vain even if it can seem
overwhelming.
Some comments on the survey of different
groups of grassland fungi
Waxcaps (Hygrocybe s.l.)
Waxcaps growing in semi-natural grasslands
were until recently placed in the genus Hygro-
cybe, distinguished from the ectomycorrhizal
species associated with trees belonging to the
genus Hygrophorus. During the last years,
there has been a revision of both genera
resulting in splitting the genus Hygrocybe into
several genera based on molecular studies
(Lodge et al. 2014). In the survey during 20
years on this locality, 34 different waxcaps
were found. It raised from 21 species to 28
from 2009 to 2010 after a more intensive
survey started (Period 2) (Figure 5). As late as
2018 the very rare waxcap Hygrocybe spadi-
cea was found for the first time (Figure 11a).
The summer of 2018 was unusual warm and
dry (NMI 2022), and this seems to be advanta-
geous to this fungus. The same year it was
found on several other localities in Norway
(Artsdatabanken 2022) indicating the same.
In addition, the species H. coccineocrenata
and H. aurantiosplendens were also only
found one year. Other waxcaps with relatively
few yearly finds are Neohygrocybe nitrata
(4) and H. subpappilata (3). Cuphophyllus
flavipes has with molecular studies been
shown to be two species (Jordal & Larsson
2021). In 2021/22 material of C. flavipes
coll. were subjected to molecular studies to
see which of the two species were present.
The study showed that both species were
growing in the location. Most molecular
studies of this complex in the region have
showed that the most common species of the
two is C. flavipesoides, and that C. flavipes
66 AGARICA vol. 43
Per Fadnes
probably is a southernly species reaching as
far north as SW-Norway (Jordal & Larsson
2021). At the moment it is not possible to
separate the two species by morphological
differences, only by sequencing, making it
difficult determining the right species in the
field. Among the other waxcaps, nine species
were found every year, and these were rela-
tively abundant species mentioned earlier. In
addition to the 34 different waxcaps species,
two subspecies have also been found in the
locality. These are Cuphophyllus pratensis
subsp. pallidus and C. virgineus subsp. ochra-
ceopallidus, the last one is a very common
species occurring almost every year.
All together a number of 41 different
species of waxcaps have until now been
recorded in SW-Norway, which means that a
substantial number (34) of these fungi grow
in this particular grassland studied here
(Artsdatabanken 2022).
Earth tongues (Geoglossaceae s. l.)
The taxonomy of Geoglossaceae has histori-
cally been difficult due to lack of good morph-
ological characters to distinguish the different
species. Therefore, they have until lately
been placed in the same family Geoglossaceae.
However, after several revisions of the family
there have been great changes. The genus
Geoglossum has recently been revised by
many authors. Schoch et. al. (2009) reduced
the family only to include the genera Sarco-
leotia, Geoglossum and Trichoglossum. Micro-
glossum, which has earlier been included in
Geoglossaceae, is now excluded, and shown
to be very distant related based on molecular
methods (Schoch et al. 2009, Sandnes 2006).
This genus is now placed in the Leotiomycetes
(Schoch et al. 2009). Hustad et al. (2011 and
2013) also included the genus Nothomitra in
Geoglossaceae and created a new genus
Sabuloglossum (including the former Geo-
glossum arenarium) and Glutinoglossum
AGARICA vol. 43 67
Fgure 9. Survey statistic: Number of visits vs. number of accumulated species and yearly finds of
grassland fungi during 20 years of survey.
Per Fadnes
(including the former Geoglossum glutinosum).
Studies of Fedosova et al. (2018) have later
proposed 13 species in the genus Glutino-
glossum by molecular studies, where seven
species are known from Europe.
The studies of Arauzo & Iglesias (2014)
confirmed the genus Leucoglossum proposed
by Imai (1942). They also proposed a new
genus Hemileucoglossum, which includes
species with hyaline spores and setae on the
stipe showing resemblance with those of the
genus Trichoglossum.
Eleven different earthtongues are found in
the locality, and they are often appearing in
great numbers. This is true for Geoglossum
fallax which is also a very macroscopically
variable species and could very well be a
complex of different species. In the wet part of
the locality, we find Geoglossum uliginosum
(Figure 11f), G. simile and Trichoglossum
hirsutum. The latter also grows in completely
dry areas where we also find Geoglossum
umbratile, G. starbaeckii, Glutinoglossum
glutinosum, Trichoglossum walteri, Microglos-
sum atropurpureum and Hemileucoglossum
pusillum. The last one was found for the first
time in 2010 but was confirmed by molecular
studies in 2020 (Fadnes et al. 2021). This is
the only known record of this species in Nor-
way and is otherwise only known from a few
localities in Slovakia and Spain. Surprisingly
a record of a new earthtongue was made in
2021, Geoglossum cookeanum. It could
probably have been recorded earlier, but since
the occurrence of earthtongues are high, it
can occasionally have been overlooked.
Redspores (Entoloma)
The genus Entoloma consist of many different
species where many are restricted to semi-
natural grasslands, especially in the subgenus
Cyanula with their characteristic scaly cap
and blue colors on stem and/or hat. However,
also species from the subgenus Nolanea and
Trichopilus are common in this habitat.
Around 30 different species of Entoloma
are found at Hovaneset until now, based on
the keys developed by Noordeloos (1992 and
2004) and Vesterholt (2002). However, in the
Norwegian Entoloma project (Brandrud et al.
2020, Noordeloos et al. 2020) where many
collections have been subjected to molecular
study, the present keys show substantial
failures in determination for a great number of
species. This is especially the case for many
of the Cyanula species with blue colors. In
Noordeloos et al. (2022b) they are also con-
firming that species of the genus Entoloma
very frequently have been misidentified in
literature resulting in high level of incorrect
names in gene databases as Gen Bank and
UNITE. This study has also shown that a
species like Entoloma corvinum, which was
known as a common species in Norway, now
seems not to grow here. In many cases in this
survey, it has been shown by molecular studies
to be E. atrocoeruleum (Table 3). The number
of Entoloma species in the locality is probably
higher than the number reported here. This is
due to the rejection of several species because
of unsecure determination. In 2021, which
was an optimal year with many Entoloma
collections, many species were subjected to
molecular studies (Table 3). This resulted in
seven new species of Entoloma to the locality:
Entoloma allospermum, E. glaucobasis, E.
carneogriseum, E. cyaneoliliacinum, E. soro-
pratulense, E. violaceoviride and E. maju-
sculum, explaining the rise in numbers of
Entoloma recorded in 2021 (Figure 4). Some
of these species have just recently been
described (Noordeloos et al. 2021, Dima et
al. 2023). The same is for E. pentagonale
which is a member of the Rhombisporum
clade (Noordeloos et al. 2022a). All the
species are also described in Fungi Europaei:
Entoloma s.l. (Noordeloos et al. 2022b).
Some of the Entoloma species described here
are shown in Figure 11.
68 AGARICA vol. 43
Per Fadnes
Most Entoloma species normally appears
early in the season, and the season for most
of them is relatively short. The slope of the
locality facing north has the richest occur-
rence of Entoloma species, and the ground
here probably also contains calcareous sedi-
ments that many Entoloma species seems to
prefer (Figure 2).
In the years before 2010 when the survey
was conducted in a smaller scale (Period 1),
only seven different species of Entoloma
were found (Figure 5). This could partly be
due to visits not conducted on the optimal
time for Entoloma species, or to difficulties
in determining the right species. However,
the number generally differs much from one
year to the other. The years 2014 and 2021,
which had warm summers in the western part
of Norway and enough precipitation later in
the autumn (NMI 2022), provided a high
number of species of Entoloma, with 20 and
25 species found, respectively. However, in
the years in between and especially after the
dry summer in 2018, the occurrences have
been sparser (Figure 4). This change in
occurrence from year to year is difficult to
explain, but difference in weather from one
year to the other and irregular fructification
of many species is worth mentioning.
AGARICA vol. 43 69
July August September October November
Week nr. 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
Hygrocybe
helobia
Hygrocybe
acutoconica
Neohygrocybe
ingrata
Hygrocybe
intermedia
Hygrocybe
splendidissima
Hygrocybe
punicea
Cuphophyllus
russocoriaceus
Cuphophyllus
fornicatus
Cuphophyllus
colemanianus
Cuphophyllus
pratensis
Hygrocybe
chlorophana
Hygrocybe
conica
Hygrocybe
quieta
Figure 10. Time of fructifying for some selected waxcaps based on 20 years of survey.
Per Fadnes
One species of Entoloma, which has been
subjected to molecular studies, came out as an
unknown species by two occasions (Table 3)
(Figure 11f). This species will be described
and published in a coming paper.
Clavariaceae
A total of 16 different species in the family
Clavariaceae are registered in the locality,
representing the genera Clavaria (7), Clavu-
linopsis (5), Ramariopsis (1), Camarophyl-
lopsis (1), Hodophilus (1) and Lamelloclavaria
(1) (Figure 4). Most of these can be character-
ized as club-fungi, while the genera Hodo-
philus and Camarophyllopsis (which were
earlier in the same genera) have many morpho-
logical similarities with waxcaps. The same
applies to Lamelloclavaria petersonii which
was found once in 2017 and confirmed with
molecular studies in 2022 (Table 3) (Figure
12d). This fungus was first described in 2016
(Birkebak et. al. 2016) and this is the second
find in Norway and the first in Western Nor-
way (Brandrud et al. 2021). Two additional
finds are made in Finland (type sp.) and Ger-
many (Brandrud et al. 2021), which indicate
that this must be a very rare species. Because
of lack of data, it was listed as data deficient
(DD) in the Norwegian red-list (Brandrud et
al. 2021).
Experience with the family Clavariaceae
from different locations is a very irregular
fructification from year to year for some of the
members. Two species have only appeared
once at Hovaneset, namely Clavaria tenuipes
and Clavulinopsis umbrinella. Clavaria
flavipes appeared twice, Clavaria fumosa
appeared three times while the rare species
Clavaria pullei was found in six years since
2013 (Figure 12c). In the genus Clavulinopsis
most of the species were found almost every
year, Ramariopsis subtilis was found six
times from 2010 and Hodophilus foetens was
found eight times since 2004 (Figure 12b).
However, in the dry year 2018, only one
species in the family was found, namely
Clavaria spagnophila, which grows in the
wet part of the locality. Several of the species
have been confirmed by molecular studies
(Table 3).
Other grasslandfungi
Only two different species of typical grass-
landfungi belonging to other genera are found
in the locality. These are Dermoloma cunei-
folium and Pseudotricholoma metapodium
(Porpoloma) (Figure 12e). The last one has
only been found once, in 2014, an optimal
year for grasslandfungi, and has been con-
firmed by molecular studies (Table 3). D.
cuneifolium is found almost every year since
2010 at least on three different spots.
DISCUSSION
The most interesting aspect with the survey is
the 20-year long-time study of fungal sporo-
carps in a selected semi natural grassland in
SW-Norway with a total of 180 visits during
the years, which makes it probably the most
intensive survey ever performed in a semi
natural grassland.
No records of grassland fungi were made
in the locality prior to the start of this survey
back in 2003. The first seven years (2003-
2009) differs from the rest of the survey with
more random visits throughout the season
compared with visits almost every week of
the season in the last 13 years.
The results from the first seven years are
therefore difficult to compare with the last 13
years, but it is interesting to see how the change
in sampling method affects the number of
fungi recorded during the first and second
period (Period 1 and 2).
No year give anywhere near the full infor-
mation about the fungi on the location even
with a visit once a week in the season. Com-
paring Period 1 and 2 it’s obvious that the
low score in period 1 is due to random and
low number of visits these years. The 180
70 AGARICA vol. 43
Per Fadnes
visits to the locality for 20 years resulted in
92 different species of grasslandfungi (Figure
9). Since nearly all species (91) were found
in period 2, this result (92 species) probably
could have been obtained with fewer visits
during fewer years given a systematic survey
as in Period 2. With this high number of
different grassland fungi, Hovaneset is today
the most species rich grassland in Norway,
and it probably could compete with the
richest grassland in Sweden and UK (Phil
1992, Griffith et. al. 2013). The survey indi-
cates how many fungi you can find in one
visit. If you are choosing the optimal year
and the optimal week of that year, you will
only find maximum 50% of the fungi growing
there (Table 2). This is more than stated by
Parker-Rhodes (1955), who tells that not
more than 25% of the fungi in a location will
be found in one visit even with many visits.
Orton (1986) confirms that several visits are
necessary to get the right picture of the species
growing on a location, but the timespan he
refers to is much less than what is the case in
this study both in number of years and number
of visits.
Newton et al. (2003) concluded that at least
16 visits to a certain pasture were necessary
to get a good understanding of the diversity
of grassland fungi on the spot. This is not
consistent with my survey. If you picked 16
random visits from the survey described in
this paper during three years, you would
probably find only a fraction of the fungi
growing there. This is at least true if the
pasture has a great potential for grassland-
fungi with many rare species with an irre-
gular fructification from one year to the other.
The survey of Newton et al. (2003) is almost
consistent with the sampling method of Halme
& Kotiaho (2012), who are criticizing the
more random sampling of fungi shown in
literature. In the article they are reporting an
intensive repeated study of wood-inhabiting
fungi with the aim to determine the optimal
timing and number of the surveys for reliable
estimation of the diversity of this group of
fungi. In this study they surveyed a forest once
a month in the snowless season throughout
four years, resulting in a total of 24 visits.
They conclude that this method will give the
optimal number of fungi in the location.
Even if the survey presented in this paper is
about grasslandfungi, it could be interesting
to challenge the statement made by Halme &
Kotiaho (2012) using data from this survey.
Based on data from Period 2 in this survey,
results from one visit in the middle of each
month (August -November) during the season
in four following years has been tested. This
results in ten different studies (2010-13,
2011-14 etc.), which can be compared with
the total survey described above with 180
visits for 20 years. In none of these ten studies
the number of fungi registered exceeded 67%
of the number of fungi found in the total
survey. The results ranged from 60 to 67%
with an average of 63% for all the ten studies.
Even if we included all years in Period 2
with one visit every month, it ended with a
find percentage of 82, and many of the most
seldom species from the survey would not
have been found.
As a conclusion, the survey of Hovaneset
shows that between 30 and 40% of the fungi
from the location would have been excluded
from the results using this method.
It not realistic that every survey of fungi
should last for 20 years and almost 200 visits,
but it shows clearly that it is not easy to get
the full overview of the biodiversity of fungi
in a location. One visit is normally only “a
blink of the eye” of the total number of fungi
on the spot, and the intense survey of this
grassland has shown that several visits during
the whole season for several years are neces-
sary to get a hold on the diversity of fungi in
the area. This is clearly shown in figure 9,
indicating that even if the last 100 visits did
not add many new species, it still count for
AGARICA vol. 43 71
Per Fadnes
13% of the species found in the location. This
indicates with all clarity that fructification
for many rare and redlisted grassland fungi is
a seldom and often irregular event.
The number of species found every single
year of the survey was only ten, which coincide
well with observation done by Staatsma et al.
(2001), who found only eight species each
72 AGARICA vol. 43
Figure 11. a. Entoloma violaceoviride*, b. E. cyaneolilliacinum*, c. E. soropratulense*, d. E.
indutoides*, e. E. neglectum, f. Entoloma sp. (new species). Photo P. Fadnes.
*Sequenced
Per Fadnes
single year during a survey lasting for 21
years. As also shown in that study, new species
appeared every year, which is almost the
same results described in this paper.
The yearly surveys with more than ten
visits to the locality gives you an impression
of how many different species you can find in
one visit, depending on the time of the visit
and the year of visit (Table 2). If you hit the
AGARICA vol. 43 73
Figure 12. a. Hygrocybe spadicea, b. Hodophilus foetens*, c. Clavaria pullei*, d. Lamelloclavaria
petersenii*, e. Pseudotricholoma metapodium*, f. Geoglossum uliginosum. Photo P. Fadnes.
*Sequenced
Per Fadnes
right week for survey you will find between
18 (2018) and 47 (2021) different species in
one visit, ranging from 20 to 51% of the total
number growing there. That means that by
“choosing” the right year and the right week
you will find not more than 50% of the species
in the location. More likely you are going to
find far less on an occasional visit. The num-
ber varies, but shows that 2013, 2014 and
2021 were the most optimal years for grass-
land fungi in SW Norway.
The most common grassland fungi on the
locality are found on many spots and could
probably represent different mycelia. Others
have only been found once at one spot and
are probably seldom registered because there
probably is only one mycelium. This is most
likely also the case for many red-listed species
found several times. As an example, Clavaria
pullei is found six times since 2013, always
on the same spot and therefore probably
consist of only one mycelium. The same is
for Hodophilus foetens which is found eight
times always on the same three spots and
could therefore probably consist of three
different mycelia.
One interesting observation made was that
in 2013 and 2014 there was many different
Entoloma species found, and they also appe-
ared in huge amounts (Figure 8). However,
the following years the occurrence of Ento-
loma species was sparse, and for some species
very often only a single basidiocarp was
found. In 2021, a new year with many Ento-
loma finds occurred, resulting in many new
species verified by molecular studies (Table 3).
This shows, at least in SW-Norway, that the
genus Entoloma has great variation in fructi-
fying and in number of basidiocarps from
one year to the other. In this study there was
six to seven years between years with high
numbers of Entoloma species (Figure 4 and 8).
The same is true for the family Clavaria-
ceae, where the irregular fructifying is parti-
cularly observed among species in the genus
Clavaria. Seven Clavaria species are found
in the location, and especially the red-listed
species C. fumosa, C. flavipes, C. pullei and
C. tenuipes have few years of finds, varying
from one to six. This observation coincides
with observations made in other semi natural
grasslands in SW-Norway.
The species Lamelloclavaria petersenii was
found on the location as the second find in
Norway and appeared only once in 2017 and
has never been found in the later years. It is
obviously a rare species with irregular fructi-
fication.
The earthtongues (s.l.) seems to be more
stable in fructifying from one year to the other,
varying from finds of seven to ten different
species each year. Even the exceptionally dry
year 2018 resulted in finds of seven different
species although the total number of species
was low (Figure 4). Some earthtongues like
Geoglossum simile, Trichoglossum hirsutum
and T. walterii seem to have wide ecological
preferences when it comes to soil consistency.
On the other hand, Geoglossum uliginosum
was always found on swampy areas. The
variation in habitat in the location probably
explains the relatively high number of earth-
tongues growing here.
Based on the results from this study it is
legitimate to ask if the season for the bloom
of grasslandfungi have changed during this
study. The maximum number of fungi found
in one visit has been delayed by several
weeks from 2014 to 2021/22 (Figure 7). In
2013/14 the maximum peak was in week 37
(medio September) and in 2021/22 it was in
week 41 (medio October). This is even more
evident if we only look at the genus Entoloma
which has a shorter fruiting period (Figure
8). This shows a delay in fruiting of around
30 days in 2021/22 compared with 2013/14.
This is also shown to a degree in Table 2. The
reason for this delay can only be speculation.
It could be due to great variation in fructi-
fying of some of the species or local vari-
74 AGARICA vol. 43
Per Fadnes
ation in weather from one year to the other.
However, the actual weather in SW-Norway
does not show that great variation between
years and can hardly count for these differen-
ces (NMI 2022). A nearby speculation is of
course climate change, which makes the
season for grassland fungi in SW-Norway
gradually more like the season in UK. The
impact of climate change on fruiting season
of fungi has been pointed out by several
authors (Kauserud et al. 2007, Andrew et al.
2018), and could therefore also be the reason
for the observation made in this study.
One interesting follow-up study of my
survey should be comparing the carpophores
found with fungi DNA from soil samples on
the location.
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ATTACHEMENTS
Tables 1, 2 and 3 on the following pages.
78 AGARICA vol. 43
Per Fadnes
AGARICA vol. 43 79
Table 1. Different species of grassland fungi found at Hovaneset 2003-2022, redlist-category (RL) and
number of yearly finds.
* One find molecular determined: Cuphophyllus flavipes
** One find molecular determined: Entoloma pentagonale
*** Entoloma sp. subjected to molecular studies: no match
Species RL Yearly
finds
Species RL Yearly
finds
Camarophyllopsis schulzeri NT 10 Entoloma soropratulense 2
Clavaria falcata 2Entoloma turci NT 2
Clavaria flavipes VU 2 Entoloma violaceoviride DD 1
Clavaria fragilis 2Entoloma sp.*** 6
Clavaria fumosa NT 2 Geoglossum cookeanum NT 1
Clavaria pullei VU 6 Geoglossum fallax 20
Clavaria sphagnicola 4Glutinoglossum glutinosum 16
Clavaria tenuipes VU 1 Geoglossum simile NT 10
Clavulinopsis corniculata 16 Geoglossum starbaeckii 7
Clavulinopsis helvola 19 Geoglossum uliginosum VU 14
Clavulinopsis laeticolor 5Geoglossum umbratile 14
Clavulinopsis luteoalba 12 Gliophorus irrigatus 19
Clavulinopsis umbrinella NT 1 Gliophorus laetus 20
Cuphophyllus colemannianus VU 20 Gliophorus psittacinus 20
Cuphophyllus flavipes (coll)* VU 10 Hemileucoglossum pusillum DD 12
Cuphophyllus flavipesoides 1Hodophilus foetens VU 8
Cuphophyllus fornicatus VU 7 Hygrocybe acutoconica 12
Cuphophyllus pratensis 20 Hygrocybe aurantiosplendens NT 1
Cuphophyllus russocoriaceus NT 20 Hygrocybe cantharellus 13
Cuphophyllus virgineus 20 Hygrocybe ceracea 18
Dermoloma cuneifolium NT 9 Hygrocybe chlorophana 20
Entoloma indutoides DD 1 Hygrocybe coccinea 20
Entoloma allospermum 1Hygrocybe coccineocrenata 1
Entoloma ameides NT 1 Hygrocybe conica 20
Entoloma atrocoeruleum NT 16 Hygrocybe glutinipes 8
Per Fadnes
AGARICA vol. 43 80
Table 1. Different species of grassland fungi found at Hovaneset 2003-2022, redlist-category (RL) and
number of yearly finds.
* One find molecular determined: Cuphophyllus flavipes
** One find molecular determined: Entoloma pentagonale
*** Entoloma sp. subjected to molecular studies: no match
Entoloma caesiocinctum 8Hygrocybe helobia 10
Entoloma carneogriseum DD 1 Hygrocybe insipida 10
Entoloma chalybaeum NT 9 Hygrocybe intermedia VU 10
Entoloma clandestinum 8Hygrocybe miniata 16
Entoloma conferendum 13 Hygrocybe mucronella NT 9
Entoloma cyaneolilacinum 2Hygrocybe phaeococcinea 9
Entoloma exile 12 Hygrocybe punicea 19
Entoloma formosum 4Hygrocybe quieta NT 15
Entoloma glaucobasis VU 1 Hygrocybe reidii 19
Entoloma griseocyaneum NT 7 Hygrocybe spadicea VU 1
Entoloma infula 8Hygrocybe splendidissima VU 18
Entoloma juncinum 2Hygrocybe subpapillata VU 3
Entoloma majusculum 5Lamelloclavaria petersenii DD 1
Entoloma mougeotii NT 9 Mikroglossum atropurpureum VU 18
Entoloma neglectum VU 1 Neohygrocybe ingrata VU 10
Entoloma rhombisporum
(coll)**
VU 5 Neohygrocybe nitrata NT 4
Entoloma poliopus 2Neohygrocybe ovina VU 15
Entoloma prunuloides VU 10 Pseudotricholoma metapodium EN 1
Entoloma sericellum 13 Ramariopsis subtilis NT 6
Entoloma sericeum 12 Trichoglossum hirsutum 16
Entoloma serrulatum 12 Trichoglossum walterii VU 15
Per Fadnes
81 AGARICA vol. 43
Table 2. Annual and weekly finds as well as percent of finds of grassland fungi the last ten years of
survey.
Weekly Finds Yearly Finds
Year Maximal
different
finds/visit
% of total (92) Week nr. Number of
different finds
% of total (92)
2013 35 38 % 37 61 66 %
2014 35 38 % 37 66 72 %
2015 32 35 % 39 59 64 %
2016 33 36 % 38 54 59 %
2017 34 37 % 37 53 58 %
2018 18 20 % 39 39 42 %
2019 31 34 % 42 57 63 %
2020 38 41 % 39 52 57 %
2021 47 51 % 41 67 73 %
2022 38 41 % 41 51 55 %
Per Fadnes
AGARICA vol. 43 82
Table 3. List of grassland fungi subjected to molecular studies. For a description of methods used
and procedures followed, with references, see “Materials and methods”.
*Noordeloos et al. 2022a
**Fadnes et al. 2021
Sequenced specimens Voucher Dato Sequencing
Agaricus phorphyrocephalus missing 20.08.2018 Alvalab
Clavaria flavipes O-F-303977 12.10.2014 NorBOL
Clavaria pullei O-F-245863 21.08.2013 NorBOL
Clavaria tenuipes O-F-303976 02.10.2014 NorBOL
Cuphophyllus flavipes O-F-204303 16.08.2021 Dima
Cuphophyllus flavipesoides O-F-259048 11.10.2021 Dima
Entoloma allospermum O-F-204173 28.09.2021 Dima
Entoloma allospermum O-F-204178 11.10.2021 Dima
Entoloma allospermum O-F-204182 11.10.2021 Dima
Entoloma atrocoeruleum (OTU 33) O-F-242498 22.08.2011 NorBOL
Entoloma atrocoeruleum (OTU 33) O-F-204175 28.09.2021 Dima
Entoloma atrocoeruleum (OTU 33) O-F-204176 22.09.2021 Dima
Entoloma atrocoeruleum (OTU 33) O-F-204183 11.10.2021 Dima
Entoloma atrocoeruleum (OTU 33) O-F-204304 22.09.2021 Dima
Entoloma carneogriseum O-F-204185 11.10.2021 Dima
Entoloma chalybeum O-F-256344 04.09.2019 NorBOL
Entoloma clandestinum O-F-204170 14.09.2020 Dima
Entoloma clandestinum O-F-204177 22.09.2121 Dima
Entoloma cyaneoliliacinum O-F-259049 28.09.2021 Dima
Entoloma cyaneoliliacinum O-F-204184 11.10.2021 Dima
Entoloma exile O-F-204174 28.09.2021 Dima
Entoloma glaucobasis missing 16.08.2021 Dima
Entoloma indutoides missing 28.08.2010 Alvalab
Entoloma majusculum O-F-204291 23.08.2021 Dima
Entoloma majusculum O-F-204306 22.09.2021 Dima
Entoloma mougeotii (OTU 31) O-F-245808 05.08.2012 NorBOL
Entoloma pentagonale* PF 1-16* 14.09.2016 Alvarado
Per Fadnes
83 AGARICA vol. 43
Table 3. List of grassland fungi subjected to molecular studies. For a description of methods used
and procedures followed, with references, see “Materials and methods”.
*Noordeloos et al. 2022a
**Fadnes et al. 2021
Entoloma poliopus O-F-304593 19.08.2016 NorBOL
Entoloma poliopus O-F-204172 28.09.2021 Dima
Entoloma poliopus O-F-204181 11.10.2021 Dima
Entoloma soropratulense O-F-259050 10.08.2014 Dima
Entoloma unknown O-F-245862 21.08.2013 NorBOL
Entoloma unknown O-F-75970 08.09.2014 NorBOL
Entoloma violaceoviride O-F-204171 28.09.2021 Dima
Entoloma violaceoviride O-F-204179 11.10.2021 Dima
Gliophorus irrigatus O-F-256325 04.09.2021 NorBOL
Gliophorus psittacinus O-F259051 28.09.2021 Dima
Hemileucoglossum pusillum** O-F-757329 10.09.2017 Alvalab
Hodophilus foetens O-F-242513 11.09.2011 NorBOL
Hygrocybe glutinipes O-F-293270 25.09.2010 NorBOL
Hygrocybe mucronella O-F-245865 13.10.2013 NorBOL
Lamelloclavaria petersenii O-F-204305 10.09.2017 Dima
Pseudotricholoma metapodium O-F-303974 12.10.2014 NorBOL
Trichoglossum hirsutum O-F-204289 16.12.2020 Dima
