Corresponding author: firstname.lastname@example.org
Norsk tittel: Noen lite kjente eller nye rød-
sporer (Entoloma; Tricholomatinae, Basidio-
mycota) for Norge
Brandrud TE, Bendiksen E, Jordal JB, Weholt
Ø, Dima B, Morozova O, Noordeloos ME,
2019. On some Entoloma species (Tricholoma-
tinae, Basidiomycota) little known or new to
Norway. Agarica 2019, vol. 39: 31-52.
rDNA ITS-sequencing, morphology, ecology,
rDNA ITS-sekvensering, morfologi, økologi,
Dette er den andre artikkelen med resultater
fra det norske Entoloma-prosjektet 2015-2017.
I denne artikkelen presenterer vi arter som er
nye for Norge eller lite kjent, fra følgende
mindre grupper/klader; Caeruleopolitum,
Claudopus, Entocybe, Leptonia, Prunuloides
og Sphagneti. I tillegg tar vi med enkelte arter
som plasserer seg noe isolert i de foreløpige
fylogenetiske analysene basert på ITS.
Følgende arter rapporteres her nye for Norge:
E. cuboidoalbum, E. jahnii, E. percoelestinum,
E. pseudoconferendum, E. ritae og E. venustum.
Den svært lite kjente E. juniperinum og
synonymi med E. mirum blir også diskutert.
Entoloma porphyrocephalum er tidligere
beskrevet som en varietet, men blir her endret
til artsnivå basert på DNA-analyser.
This is the second paper reporting major results
from the Norwegian Entoloma project 2015-
2017. Here we present species that are new to
Norway or little known, from the following
smaller clades: Caeruleopolitum, Claudopus,
Entocybe, Leptonia, Prunuloides and Sphagneti.
In addition, a few species taking a somewhat
isolated position in our unpublished preliminary
ITS tree are presented. The following species
are here reported new to Norway: E. cuboido-
album, E. jahnii, E. percoelestinum, E. pseudo-
conferendum, E. ritae, and E. venustum. The
synonymy of E. mirum with the older E. juni-
perinum, is also discussed. Entoloma por-
AGARICA vol. 39 31
On some Entoloma species (Tricholomatinae, Basidiomycota)
little known or new to Norway
Tor Erik Brandrud1, Egil Bendiksen1, John Bjarne Jordal2, Øyvind Weholt3, Bálint
Dima4, Olga Morozova5, Machiel E. Noordeloos6
1Norwegian Institute for Nature Research, Gaustadalléen 21, NO-0349 Oslo,
2Biolog J.B. Jordal, Skrøovegen 21, NO-6610 Øksendal, Norway
3Nord University, Nesna, NO-8700 Nesna, Norway
4Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University,
Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary
5Komarov Botanical Institute of the Russian Academy of Sciences, 197376, 2
Prof. Popov Str., Saint Petersburg, Russia
6Naturalis Biodiversity Center, section Botany, P.O. Box 9517, NL-2300 RA
Leiden, The Netherlands
Brandrud et al.
phyrocephalum, formerly described as a
variety, is now raised to species level based
on its phylogenetic position.
The present paper is the second in a series of
papers in Agarica reporting results from the
Norwegian Entoloma project 2015–2017. The
Norwegian Entoloma project has been a part
of The Norwegian Taxonomy Initiative, funded
by The Norwegian Biodiversity Information
Centre (NBIC). In the first paper, results on
the Rhodopolia clade were reported (Brand-
rud et al. 2018). In this second paper, some
small, basal clades in Entoloma are emphasized,
including the clade Entocybe, often disting-
uished as a separate subgenus, by some even
as a separate genus (Baroni et al. 2011).
Furthermore, the present paper also treats
some taxa from the Claudopus and Prunu-
loides groups/clades (in the sense of e.g.
Noordeloos 1992) and Leptonia s. str. (in the
sense of Morozova et al. 2014), as well as
some taxa with uncertain phylogenetic affinity,
which are new to Norway or at least little
known. A third paper, on the large clade
Cyanula (= Leptonia s. lat.) is in preparation
for a later issue of Agarica. All results pre-
sented here are based on samples being
verified by rDNA ITS sequencing, mainly
through NorBOL (Norwegian Barcode of
Life Network; see Ratnasingham and Hebert
Entoloma is one of the most species-rich
genera within Agaricales, well characterized
by many-angled spores that leave a pinkish
brown spore print. As commented more deeply
in the first paper (Brandrud et al. 2018), we
consider that Entoloma should still be kept as
one single, very large genus and not split into
several smaller genera, due the fact that varia-
bility in Entoloma is complex and still not
fully understood. This is mainly due to incom-
plete sampling and the fact that large regions
are still underexplored (e.g. Africa, S. America,
Asia) and the fact that many clades in recent
phylogenetic analyses show low support with
the genetic markers applied so far (see e.g.
Morgado et al. 2013).
MATERIAL AND METHODS
Altogether approx. 700 samples of Entoloma
species have been collected during the Nor-
wegian 2015–2017 Entoloma project, and
1080 samples were verified by ITS-sequencing.
The latter number also includes many herbar-
ium collections. In addition, sequences were
obtained of numerous type specimens and
some important reference material from out-
side Norway studied in connection with the
present project. All sequenced material from
Norway are listed under material sequenced.
Collections labelled NOBAS or CAFUN are
sequenced through NorBOL, those labelled
ALV are sequenced by Pablo Alvarado
(ALVALAB, Spain), and those with no
sequence label are sequenced by us (BD, in
Eötvös Loránd University, Budapest). All
sequences are analysed by us (see below).
All material is deposited or will be deposited
in herb. Oslo (the Oslo fungarium).
Morphology and molecular study
The macromorphological observations are
based on the field notes and photographs of
the available material. Microscopical obser-
vations were made using standard methods
(see e.g. Morozova et al. 2014), and the mole-
cular study was likewise performed according
to standard methods, described in detail in
the first paper in this series (Brandrud et al.
RESULTS AND DISCUSSION
During our NBIC Entoloma project 2015-2017,
we could distinguish 194 well-supported,
phylogenetic species (Operational Taxonomic
Units, OTU’s). Of these, 87 species were new
to Norway. In the following, we will present
32 AGARICA vol. 39
Brandrud et al.
in more detail some of our results on a number
of small clades.
1. The basal Entoloma clades
The basal Entoloma lineages (the basal Ento-
loma grade as defined by Baroni et al. (2011)
and Co-David et al. (2009)), includes several
distantly related clades, among others the
apparently polyphyletic (sub)genus Entocybe.
Many species in this basal group are charac-
terized by a peculiar spore morphology, being
rather small, thin-walled, often rounded and
weakly angular, almost nodulose-warted,
resembling a type of spores that is also found
in the Rhodocybe clade. There are, however,
exceptions, like in the case of E. sphagneti,
which has more elongate-heterodiametrical
spores, which at the same time also have a
wavy-nodulose outline instead of being
sharply angled as in most Entoloma species.
A macromorphological feature found in this
basal Entoloma clades is the occurrence of
blue and/or violaceous pigments in many
species, a character shared with the Cyanula
and Leptonia clades, but otherwise a rare
feature in the genus Entoloma. Many species
in these clades also have a characteristic shape
of the stipe, which is distinctly swollen just
above the often rather distinctly tapering base.
Members of the basal clades occur widespread
all over the world, with several species recently
described from Australasia, and America
(Horak 2008, Largent 1994, Noordeloos and
Gates 2012). Many European species prefer
oligotrophic habitats, such as Sphagnum peat
bogs and heathland on poor, acidic soil.
1.1. Clade Caeruleopolitum
This small clade consists of two, morphologi-
cally rather similar species, characterized by
having relatively large spores, up to 10 (11)
µm long, and with normally thick walls and
well-developed angles. Both species may have
blue-violaceous tinges in the basidiocarp or
not. The more common of the two, E. caeruleo-
politum occurs widespread in Norway, while
E. juniperinum is more rarely recorded, but
seems locally not infrequent.
Entoloma caeruleopolitum Noordel. &
Brandt-Ped. Fig. 1 A, B.
Characteristics: Pileus up to 30 mm broad,
conical at first, soon convex or applanate,
umbonate or slightly depressed at centre,
hygrophanous, when moist deeply translucently
striate, reddish or porphyry brown, as young
often tinged blue or more greyish brown,
darker at centre, pallescent on drying, glabrous.
Lamellae rather distant, broadly adnate or
emarginate, sometimes with decurrent tooth,
segmentiform to ventricose, pallid creamy-
beige or tinged brownish, then with pink
tinge. Stipe 20–60 × 2–5 mm, cylindrical or
compressed with longitudinal groove, some-
times tapering towards base, blue, violaceous-
AGARICA vol. 39 33
Figure 1. A, B) E. caeruleopolitum. A) KB&EB107-16;
B) JBJ0251. Photos: A) E. Bendiksen; B) J.B. Jordal.
Brandrud et al.
brown or greyish brown, smooth, polished.
Context dark grey in pileus, in cortex of stipe
with blue tinge. Smell and taste indistinct.
Spores 8.0–10.0(–11.0) × 6.5–8.0 µm, Q =
1.1–1.3, subisodiametrical, 5–6-angled in
sideview. Basidia 4-, and occasionally also 2-
spored, clamped. Cystidia absent. Pileipellis
an ixocutis of narrow, cylindrical, 3–8(–11)
µm wide hyphae, subpellis usually well
differentiated, made up of inflated elements,
up to 25 µm wide. Pigment brown, intracel-
lular in pileipellis. Clamps abundant in all
Habitat & distribution: In SE and E Norway
the species has been found in poor grasslands,
mainly abandoned but pastured summer
farms on oligotrophic bedrock. Along the W
Norwegian coast it is found in similar habi-
tats, but there is also a few finds in calcareous
soils. It is recorded up to 965 m a.s.l. Oppdal,
South Trøndelag. The species is found mainly
in sites poor in Entoloma species (only with
E. conferendum, E. clandestinum and E.
sericeum) and it also occurs much later in the
season (together with a.o. Hygrocybe spp.)
than most Entoloma species.
Collections sequenced: NORWAY. Hedmark:
Rendalen, Grøndalen, old summer farm pasture,
3 Sept 2016, K. Bendiksen, E. Bendiksen KB
EB 107/16 (O-254028). Hordaland: Bømlo,
Holme, in semi-natural pasture, 9 Oct 2008,
leg. A. Knutsen, J. B. Jordal (O-287891).
Bømlo, Tverborgvika ved Lykling, in semi-
natural pasture, 10 Oct 2008, leg. A. Knutsen,
J. B. Jordal (O-287897). Møre og Romsdal:
Averøy, Vågsholmen by Kårvåg, in semi-
natural pasture, 30 Sep 1994, leg. G. Gaarder,
J. B. Jordal (O-178137). Oppland: Sel, Otta-
dalen, Rustmo, leg. H. Schwencke, pasture,
22 Sep 2016, HS-E1-15 (ALV7294). Sør-
Trøndelag: Oppdal, Losfjellet, Medlisætrene,
in semi-natural pasture, 30 Aug 2016, leg. J.
B. Jordal, JB16994 (O-304665).
DENMARK. Jutland: Silkeborg, Høvild forest,
18 Oct 1982, leg. Brandt-Pedersen 82.268 (C
67093 holotype E. caeruleopolitum , ITS2
T. G. Frøslev, pers. comm.).
Comments: Entoloma caeruleopolitum, origi-
nally described from Jutland, Denmark, has a
wide distribution in Northwestern Europe, but
it is often overlooked because of its small,
inconspicuous basidiocarps. Distinctive
characters are the glabrous, translucently
striate pileus, polished stipe, bluish colours,
(sub-)isodiametrical spores, clamped hyphae
and simple ixocutis. Like in other species in
this clade, the colour of the basidiocarps varies
a great deal. Typically, the pileus is bluish
tinged as young, soon turning moderately dark
reddish or porphyry brown; the stipe is blue
or violaceous-blue, but in some collections,
the stipe lacks blue tinges. Entoloma reae,
also with blue pileus and stipe, may appear
to be conspecific with E. caeruleopolitum as
discussed by Noordeloos (1992). A type
sequence is needed to prove the synonymy.
Entoloma juniperinum is closely related, but
it differs by having a darker, inconspicuously
translucently striate pileus and normally less
Entoloma juniperinum Barkman &
Syn.: Entoloma mirum Kokkonen
Emended description (incl. features of E.
mirum): Pileus 5–20 mm broad, convex or
applanate, with weak central depression or
with small papilla, with deflexed then straight
margin, hygrophanous, when moist translu-
cently striate at margin up to 2/3 of radius,
dark brown to grey-brown, especially at centre,
sometimes when young with blue tinges near
margin, occasionally pinkish with darker
centre, pallescent on drying, opaque, almost
glabrous or covered with fine whitish fibrils,
at centre sometimes minutely squamulose.
Lamellae, L = 10–25, l = 1–5(–7), (moderately)
distant, adnate sometimes emarginate with or
without decurrent tooth, segmentiform, rarely
34 AGARICA vol. 39
Brandrud et al.
ventricose, up to 6 mm broad, sometimes
transvenose, white, or pale grey-brown then
with pink tinge. Stipe 15–45 × 1–5(–8) mm,
cylindrical, sometimes slightly to distinctly
swollen at base, brown or pink, more or less
concolorous with pileus or with blue or steel-
grey tinge, fading to brown with age, smooth,
polished or covered with fine fibrils length-
wise. Smell indistinct to subfarinaceous. Taste
often distinctly farinaceous, sometimes absent.
Spores 8–10(–11) × 6–8 µm, O = 1.1–1.2
(–1.25), subisodiametrical, 5–7-angled in
side-view. Basidia 4-spored. Cheilocystidia
absent. Pileipellis a cutis with transitions to a
trichoderm, made up of cylindrical to slightly
inflated hyphae, 5–20 µm wide, subpellis well-
differentiated, made up of inflated elements,
20–45 × 15–35(–45) µm. Pigment brown,
intracellular in pileipellis, especially in
subpellis. Caulocystidia scattered, simply
cylindrical to subclavate. Clamps abundant
in all tissues.
Habitat & distribution. The Norwegian collec-
tions are from eutrophic Rubus idaeus thickets
on clear-cuttings, where it seems locally not
infrequent, as well as from a semi-natural
pasture and Alnus incana floodplain (delta)
Collections sequenced: NORWAY. Oppland:
Lunner, S. Oppdalen, Mørkomdalen, 27 Aug
1979, E. Bendiksen, EB 266/79 (O; as E. juni-
perinum, conf. M.E. Noordeloos), 9 Sept 1990,
E. Bendiksen (O). Lunner, Brovoll, semi-
natural forest pasture, 6 Oct 2000, E. Bendik-
sen (O). Nordre Land, Dokka, Dokkadeltaet
nature reserve, Bergsrønningen, floodplain
Alnus incana–Prunus padus woodland, 15
Sept 1989, K. Bendiksen, E. Bendiksen, KB
EB 269/89 (O).
Comments: This species, normally being less
bluish tinged than its sister species E. caeruleo-
politum, is very little known and seldom col-
lected. Nevertheless, it seems in Norway to
be locally rather frequent in eutrophic Rubus
idaeus shrublands after clear-cutting of low-
herb spruce forest. Unfortunately, the nomen-
clature of this taxon is still somewhat unsettled.
Kokkonen (2015) published E. mirum as a
new species, with normally pink or sometimes
rather dark brown basidiocarps and small,
more or less isodiametrical or slightly hetero-
diametrical spores. All three known collections
were found in the type locality. She noted the
remarkable differences in colour between the
specimens, some being pink, others dark brown.
The type sequence appeared to be similar to
some earlier collections from Norway identi-
fied by one of us (MEN) as E. juniperinum,
as well as another one from the Netherlands,
identified as such (J. Schreurs 892, L).
Especially the latter should be regarded as
a representative reference collection for the
original concept of E. juniperinum. Entoloma
juniperinum, also known from juniper heaths
in Denmark, Germany and the Netherlands,
has similar morphological features as those
described by Kokkonnen for E. mirum, but a
still larger range of colours, including forms
with distinct blue tinges in the stipe. This
phenomenon occurs also frequently in the
related taxa E. vinaceum, E. caeruleopolitum,
and the North American E. trachyosporum.
Unfortunately, the holotype of E. juniperinum
did not produce usable DNA sequence for
analysis. We are convinced, however, based
on the morphological similarities, and match
with reference material, that E. mirum is a
later synonym of E. juniperinum. More
material will be sequenced and studied
morphologically to support the supposed
1.2. Clade Sphagneti
So far, three members of this clade are known
from Europe, all of which occur in Norway,
and seem preferably to be associated with
Sphagnum in peaty areas in more or less
AGARICA vol. 39 35
Brandrud et al.
acidic environments. The spores are thin-
walled, and more or less nodulose-bumpy in
outline, heterodiametrical in E. sphagneti
and E. pseudoconferendum, isodiametrical in
E. chamaemori. Entoloma sphagneti is a rela-
tively large, stout fungus, almost tricholomatoid,
the other two are more slender, mycenoid or
collybioid. At present, these are verified from
only two collections each. Whether these are
truly rare in a frequent habitat, or just over-
looked, remains to be seen.
Entoloma sphagneti Naveau. Fig. 2 A, B.
Characteristics: A fairly stout, tricholomatoid
(-mycenoid) fungus. Pileus up to 120 mm
broad, conical, expanding to convex with
broad umbo, dark red-brown, slightly trans-
lucently striate, glabrous, but with fibrillose
patches along the margin. Lamellae deeply
adnate-emarginate, ventricose, reddish pink.
Stipe 35–150 × 3–17 mm, from, brown, paler
than pileus, strongly fibrillose striate length-
wise. Smell indistinct or farinaceous. Spores
9.5–12.5 × 6.5–9.0 µm, distinctly hetero-
diametrical with a wavy-nodulose, many-
angled outline. Basidia 4-spored. Cystidia
absent. Pileipellis an ixocutis or cutis of
narrow, cylindrical hyphae, 2.5–9 µm wide.
Pigment very abundant, brown, intracellular,
in some collections also encrusting. Clamps
present in all tissues.
Habitat & distribution: In Norway the species
is verified only from two collections, one
(ALV5672) in a muddy dried-out “pond”,
with needle litter in an oligotrophic spruce
forest, the other one (ALV14353) in a
recently (1 year) dug muddy ditch on peaty
Collections sequenced: NORWAY. Østfold:
Fredrikstad, Hystad, near Prestmyra, 11 Aug
2014, Ø. Weholt, OW E2 14, ALV5672 (O).
Fredrikstad, Blåkollen, 13 Oct 2017, M.
Pettersen, MP-3.131017, ALV14353 (O; as
E. aff. turbidum).
Comments: Entoloma sphagneti is a rare spe-
cies, mainly confined to (Atlantic) Sphagnum
bogs in Northwestern Europe. The species was
originally described from Belgium (Naveau
1923). The type is not sequenced, but fits in
all other respects, and since this is morpho-
logically and ecologically a quite well defined
species, we choose to apply this name. Most
records of E. sphagneti are from the Nether-
00041140) where it is usually found at the
margin of peat bogs, often in dry summers in
places where the Sphagnum is dying, some-
times it grows on dried out peat, and occasio-
nally it is also found in swamp forest with
Salix aurita. The two Norwegian sequence-
verified collections are from more or less
dried-out ditches and pond-like depressions,
probably with Sphagnum peat. It is also
36 AGARICA vol. 39
Figure 2. A, B) E. sphagneti. A) JV02-695; B) Ø.
Weholt E2 14. Photos: A) J. Vesterholt. B) Ø. Weholt.
Brandrud et al.
recorded from Britain (Wickens and Legon
2005) and was published from Norway and
Denmark by Noordeloos in Knudsen and
Entoloma pseudoconferendum Noordel. &
Wölfel Fig. 3 A-E.
Syn. Entoloma dolosum Noordel. & Wölfel,
non Entoloma dolosum Corner & Horak.
Characteristics: Habit slender, mycenoid.
Pileus up to 3 cm broad, convex with umbo
soon applanate, dark reddish-greyish brown,
deeply translucently striate, hygrophanous,
pallescent on drying, glabrous. Lamellae
moderately distant, deeply adnate-emarginate,
brownish pink. Stipe much paler than pileus,
silvery-fibrillose striate, base with white
tomentum. Smell slightly farinaceous. Spores
10–14 × 7–10 µm, heterodiametrical, weakly
angular-nodulose in outline. Basidia 4-spored.
Cystidia absent. Pileipellis a cutis of cylindrical
hyphae with intracellular pigment. Clamps
Habitat & distribution: Found in (very) moist
oligotrophic bogs and ditches, apparently
fruiting mainly in very dry summers with a
lowered water table.
Collections sequenced: NORWAY. Aust-Agder:
Risør, Opstadtjern, A. Omberg, AO 3-18,
ALV18069. Froland, Øynaheia, I.-L.- Fonne-
land & D. Pettersen, ILF 2018-001 (O-76265).
THE NETHERLANDS. Prov. Gelderland:
Bronckhorst, Kruisbergsebossen, 25 Sept
2016, G.M. Jasen C1594394 (L).
Comments: Entoloma pseudoconferendum
was described in 1995 based on a single
collection from Germany, and have never been
recorded afterwards until 2016, when it was
found in the Netherlands in September and in
Eastern Germany in October, in both occasions
in the margin of peat bogs where the Sphag-
num had been dried out. The species was
found in a number of dried-out bogs, ditches
and exposed lakeshores during the very dry
summer of 2018 in southernmost Norway
AGARICA vol. 39 37
Figure 3. A-E) E. pseudoconferendum. A) (habitat), B), C) ILF 2018-001; D) AO 3-18; E) G.M. Jansen C1594394.
Photos: A)-C) I.-L. Fonneland; D) A. Omberg; E) G.M. Jansen & M. Vegte.
Brandrud et al.
(Agder; I.-L. Fonneland, pers. comm.), and
these represents the first finds in Norway.
Entoloma pseudoconferendum has strong
similarities with E. conferendum, which is
not infrequent in bog habitats during summer
in Norway. The latter has, however, completely
different spore morphology. Microscopically,
this species is similar to E. sphagneti, but it
can readily be distinguished on the slender,
mycenoid habit, and the glabrous, deeply
translucently striate pileus, and possibly also
on a more extremely seasonally inundated
Entoloma chamaemori Noordel., Weholt,
Eidissen & Lorås. Fig. 4 B.
Characteristics: Habit mycenoid, with 25–40
mm wide, conico-convex, expanding pileus,
(reddish) brown with darker centre, glabrous.
Lamellae distant to moderately crowded, nar-
rowly adnate-emarginate to (almost) free,
greyish white, becoming pinkish tinged, with
concolorous edge. Stipe 50–70 × 3–5 mm,
greyish white, apex pruinose, silvery striate
lengthwise. Smell farinaceous. Spores 6.0–8.0
(–8.5) × 5.5–7.0(–7.5) μm, isodiametrical to
subisodiametrical, with 6–8 rather weak angles,
thin-walled. Basidia 4-spored. Cystidia absent.
Pileipellis a differentiated (ixo-) cutis of thin
hyphae, with scattered clavate to cystidiform
terminal elements, 20–35 × 4–12 μm and
brown intracellular pigment. Clamps present.
Habitat and distribution: Collected in an oligo-
trophic bog, among Sphagnum papillosum
and S. tenellum in open, sub-exposed, site with
predominantly Rubus chamaemorus. So far
only known from two collections from one
locality in Holmvassdalen, Northern Norway.
Collections sequenced: NORWAY. Nordland:
Grane, Holmvassdalen Nature Reserve, 10
Aug 2014, S. E. Eidissen & J. Lorås, JL69-
14 (holotype, O). GenBank: KX928955; 10
Aug 2014, S. E. Eidissen & J. Lorås, JL70-
14 (O). GenBank: KX928956.
Comments: The species was described quite
recently, based on Norwegian material from a
poor Sphagnum bog in Holmvassdalen, Nord-
land (Noordeloos et al. 2017). The small,
nodulose weak-angled spores of E. chamaemori
resemble those of E. vinaceum and E. turbidum,
but the angles are slightly more pronounced
and the walls slightly thicker, whereas those
of the other two resemble Rhodocybe spores.
In the phylogeny, E. chamaemori is rather
isolated and clusters basally with E. sphagneti
and E. pseudoconferendum (Noordeloos et
al. 2017), both of which have more elongate,
1.3. Clade Vinaceum/Entocybe
This clade more or less fits with the concept
of Entocybe, and contains species with small,
thin-walled, often rounded-nodulose or bumpy
spores, resembling those found in the Rhodo-
cybe clade. In Norway, it is represented with
38 AGARICA vol. 39
Figure 4. A) E. turbidum. J. Vesterholt (not sequenced).
B) E. chamaemori, JL69-14 (holotype). Photos: A) J.
Vesterholt, B) J. Lorås.
Brandrud et al.
four species: Entoloma nitidum, well known
by its splendid blue basidiocarps, E. vinaceum,
E. turbidum, and a fourth taxon, probably
referable to E. turbidum var. pachylamellatum.
Due to uncertainties on nomenclature, morpho-
logical variation and circumscription, the
latter is not treated here.
Entoloma vinaceum (Scop.) Arnolds &
Syn.: Entoloma vinaceum var. fumosipes
Arnolds & Noordel.; Entoloma vinaceum
var. violeipes Arnolds & Noordel.
Characteristics: Pileus 20–40 mm broad,
convex, often with slightly depressed centre,
sometimes vaguely umbonate with deflexed
margin, hygrophanous, deeply translucently
striate when moist, moderately dark horn
brown with darker centre, of grey- to reddish
brown, paler towards margin, pallescent on
drying, glabrous. Lamellae moderately distant,
adnate-emarginate, segmentiform, greyish
then with pink tinge, with entire, concolorous
edge. Stipe 20–60 × 1–3 mm, cylindrical,
often flexuous, and distinctly tapering towards
base, sometimes rooting, variably coloured,
typically yellowish to yellow brown, but
frequently also grey or with violaceous tinges,
glabrous, polished, base white tomentose.
Smell and taste indistinct. Spores 6.0–8.0 ×
5.5–7.5 µm, isodiametrical, very thin-walled
and weakly angled, sometimes appearing
almost nodulose, reminiscent those of Rhodo-
cybe species. Basidia 4-spored. Cystidia none.
Pileipellis a simple, thin (ixo)cutis of narrow,
cylindrical hyphae. Pigment brown, intracel-
lular. Clamps present in the entire basidiocarp.
Habitat & distribution: The species is in Europe
mostly confined to nutrient poor, acidic habi-
tats, like dry, poor Pinus forests of Vaccinium
vitis-idea-Calluna type, Picea forests of
Vaccinium myrtillus type, heathlands, peat
bogs and very poor grasslands. In Norway,
the verified finds so far are from oligotrophic
Pinus-Picea forests in the Oslo-Hadeland
region. Probably widespread, but little known.
Collections sequenced: NORWAY. Oppland:
Lunner, Øståsen, Morstadhaugen (study plot
15-19), oligotrophic, xeric (lichens, Calluna)
pine forest, 4 Oct 2001, E. Bendiksen EB
204/01 (O-169164). Lunner, Øståsen, Morstad-
haugen, oligotrophic, subxeric pine(-spruce)
forest, 29 Sept 2004, E. Bendiksen 246/04
(O-165807). Lunner, Øståsen, S. Korsvatn-
haugen, old Vaccinium myrtillus–Picea
forest, 9 Oct 2015, E. Bendiksen EB 279/15
(O-252032). Oslo: Grorud, Frankrig N, mixed
forest, 26 Sept 1984, E. Bendiksen EB 362/84
Comments: Entoloma vinaceum is morpho-
logically rather variable species, with a
yellow, grey or vinaceous stipe. Accordingly,
varieties have been published with greyish
stipe (var. fumosipes) and violaceous-blue
stipe (var. violeipes). Our molecular studies
indicate, however, that these varieties should
be considered phenotypic variation rather than
worthy of taxonomic rank, like in species with
similar colour variation discussed above.
Sometimes the species can be confused with
E. turbidum, which is also widespread in
oligotrophic-acid habitats. However, E. tur-
bidum is usually a stouter fungus found in
somewhat moister spruce forests, whereas E.
vinaceum is more frequent in drier pine
forests and heathlands.
Entoloma nitidum Quél.
Characteristics: Habit mycenoid or slenderly
tricholomatoid. Pileus 20–40(–50) mm broad,
conical or campanulate then expanding to
convex with broad, low umbo, not hygrophan-
ous, not translucently striate or at margin
only, bright blue, sometimes blackish blue at
centre, silky shining, glabrous or radially
fibrillose. Lamellae moderately distant,
adnate-emarginate, white then pink. Stipe
AGARICA vol. 39 39
Brandrud et al.
30–100 × 2–5(–7) mm, cylindrical with
attenuated, sometimes even rooting base,
blue, concolorous with pileus or paler, with
distinct yellow tinge near base, innately
fibrillose striate. Spores 7–9 × 6–7 µm, iso-
diametrical, thin-walled, poorly angled. Basidia
4-spored. Cystidia absent. Pileipellis an
ixocutis of 2–6 µm wide, hyphae with blue,
intracellular pigment. Clamps abundant.
Habitat & distribution: The species is in
Norway rare but widely distributed in mossy,
oligotrophic(-mesotrophic) Picea abies forests
of Vaccinum myrtillus type. Sometimes it is
recorded also in oligotrophic Pinus or Quer-
Collections sequenced: NORWAY. Akershus:
Ski, Svartoren, under spruce and pine, 12
Sept 2014, S. Hansen (O-75603). Hordaland:
Bømlo, Totland, in margin of pasture, among
moss and needles under Pinus sylvestris,
moist, 29 Sept 2009, A. Knutsen, J. B. Jordal
(O-291246).Oppland: Lunner, S. Oppdalen,
Mørkomdalen, Solbakken; open spot in low
herb Picea forest, Vaccinium myrtillus, grass,
19 Aug 2014, A. Molia (O-75140).
Comments: Entoloma nitidum is one of the
species in the genus that can easily be identi-
fied in the field by its mycenoid to slenderly
tricholomatoid, bright blue basidiocarps and
the ecology in oligotrophic, acidic coniferous
and deciduous forests. Although these habi-
tats are among our most frequent forest types,
this beautiful species is everywhere rare, at
least based on basidiocarps, which are not seen
every year. Entoloma bloxamii/madidum
differs in a more stout tricholomatoid stature,
strong farinaceous smell and habitat in cal-
careous grasslands and forests.
Entoloma turbidum (Fr.) Quél. Fig 4 A.
Characteristics: Pileus 20–80 mm, conical or
campanulate, then convex with pronounced
umbo, not hygrophanous, not translucently
striate, grey-brown or sepia brown, glabrous,
slightly sticky in moist weather. Lamellae
deeply emarginate-adnate, crowded, white or
greyish, then pink. Stipe 30–90 × 4–10(–12)
mm, fusiform, usually broadened just above
the tapering base, greyish white, densely
covered with silvery fibrils, at base often
tinged yellow. Smell and taste indistinct to
rancid-farinaceous. Spores 6.5–8.5(–9.0) ×
6.0–7.0(–7.5) µm, Q= 1.0–1.2, thin-walled
and many-angled with weak angles, appearing
almost nodulose. Basidia 4-spored. Cystidia
absent. Pileipellis an ixocutis of 2.5–5 µm
wide, cylindrical hyphae. Pigment brown,
intracellular in pileipellis. Clamps present in
Habitat & distribution: In Norway rather
frequent in oligotrophic-mesotrophic Picea
abies forests, sometimes in Sphagnum, more
rarely in oligotrophic pine forests and grass-
lands. Distributed in all parts of Norway.
Collections sequenced: NORWAY. Møre og
Romsdal: Skodje, Skodjereitane, alt. ca. 100
m, under pine and spruce, 02 Aug 2007, P. G.
Larsen (O-75994). Oppland: Lunner, S.
Oppdalen, Hellerud NE, old Picea forest of
Vaccinium myrtillus type, 19 Aug 2014, E.
Bendiksen (O-75256). Rogaland: Rennesøy,
Bø-Asmarvik, alt. 20 m, 30 Sept 2008, J. B.
Comments: Entoloma turbidum can be distin-
guished from E. vinaceum by the more robust
stature with a conical pileus and firm stipe,
but they may occur in similar habitats.
However, the latter is more frequent in dry
pine forests and heathlands, whereas the
former prefers moister spruce forests. Ento-
loma turbidum is one of the most frequent
Entoloma species in our boreal spruce forests
(together with E. cetratum and E. conferen-
dum) and should be familiar to those looking
40 AGARICA vol. 39
Brandrud et al.
for red-spored fungi. We have nevertheless
included it for comparison with the less
frequent or new taxa within the basal clades
2. The Prunuloides clade
The Prunuloides clade/group is a small, well-
supported lineage according to Morgado et
al. (2013), including strongly farinaceous
smelling, calciphilous grassland/forest species
with small, isodiametric spores. Three taxa
are so far recognized in the Prunuloides clade
in Norway, of which E. luteobasis is presen-
ted here, and some new data on E. bloxamii
s. l. are commented. Entoloma prunuloides, a
widespread species well known from grass-
lands (Jordal et al. 2016), are not treated here.
Entoloma luteobasis Ebert & E. Ludw. Fig.
5 A, B.
Syn.: Entoloma ochreoprunuloides Morgado
& Noordel.; E. prunuloides var. obscurum
Arnolds & Noordel.
Characteristics (emended description): Pileus
20–60 mm broad, conical soon expanding to
plan-convex with low umbo, with initially
deflexed, then straight margin, expanding
with age to applanate with weak umbo and
undulating margin, hardly hygrophanous,
medium to fairly dark (grey)brown, uniformly
coloured or with dark, greyish brown centre,
possibly sometimes with a bluish grey reflex
when very young, not entirely glabrous but,
particularly at central part with an aeriferous
covering of fine, silvery hairs, sometimes also
radially wrinkled. Lamellae fairly crowded to
moderately distant, adnate-emarginate, ventri-
cose, sordid white then pink with entire,
concolorous edge. Stipe 30–50 × 5–15 mm,
cylindrical, straight or flexuous, sometimes
slightly broadened towards base, off white to
pale grey-brown, at base sometimes, but not
frequently with distinct yellow tinges, innately
fibrillose lengthwise, often with a metallic
shine. Context concolorous, rather firm. Smell
and taste farinaceous. Spores: 6.0–7.0 × 5.5–
7.0 µm, Q = 1.0–1.05, isodiametrical with
normally thick walls, with 5 to 6 angles in
side-view. Basidia 4-spored. Cystidia absent.
Pileipellis a thin ixocutis of narrow, cylindri-
cal up to 6.5 µm wide hyphae. Pigment brown
intracellular. Clamps abundant.
Habitat & distribution. In Norway, most
records are from calcareous Tilia-Corylus
forests, mainly in the inner Oslofjord (Oslo,
Bærum, Asker, Røyken), but also in the outer
Oslofjord (Porsgrunn-Bamble), a few finds
also from calcareous Pinus forests including
transitions to open shrubland and from rich
Quercus-Tilia forests. Altogether more than
20 localities are known so far (including non-
sequenced material too). Outside Norway,
this species has been recorded only a few times
from North-western Europe (Netherlands,
Germany, Great Britain), from calcareous
AGARICA vol. 39 41
Figure 5. A, B) E. luteobasis. A). TEB363-14; B)
TEB375-14. Photos: B. Dima.
Brandrud et al.
Quercus-Carpinus and Fagus forests, and
from calcareous grasslands.
Collections sequenced: NORWAY. Akershus:
Asker, Vettre, 8 Sept 2011, E. Bendiksen, EB
713-11 (O-253930). Bærum, Dælivann,
Langenga east, 6 Sept 2014, T.E. Brandrud,
B. Dima, TEB 363-14 (O); Laenga west, 6
Sept 2014, T.E. Brandrud, B. Dima, TEB
375-14 (O). Buskerud: Røyken, Bøsnipa, 13
Sept 2011, K. & E. Bendiksen, T. E. Brandrud
& I. Kytövuori, TEB 819-11 (O-248443); 11
Sept 2014, T. E. Brandrud, B. Dima, Høstsopp-
treff (foray) no. 1462 (O-F-75440). Oslo:
Bygdøy, Reinsdyrlia, 13 Sept 2014, T.E.
Brandrud, B. Dima, Høstsopptreff (foray) no.
14780 (O-75787). Oslo, Gressholmen/Ramber-
gøya, 24 Sept. 2008, E. Bendiksen, EB 259-08
Comments: We have for some time called this
E. ochreoprunuloides (see e.g. Brandrud et
al. 2016), according to the phylogeny of
Morgado et al. (2013). However, our mole-
cular study revealed that the type of E. luteo-
basis is identical with that of E. ochreoprunulo-
ides. This throws a new light on the morpho-
logical variability of this species. The type of
E. luteobasis showed a very striking yellow
stipe base, but apparently. this character is
rather variable, and may be present or absent.
However, the dark coloured pileus separates
it clearly from the related E. prunuloides, as
well as the lack of blue/violaceous tinges
differentiate this species from E. bloxamii s. l.
The species was first reported from Norway by
Gulden (2011) under the name E. luteobasis,
found in a calcareous, mixed forest along the
Oslofjord. Then the species was found several
times in connection with our Entoloma project
and a monitoring program for calcareous
Tilia forests in the Oslofjord region (see e.g.
Brandrud et al. 2016; as E. ochreoprunuloides).
It is not infrequent in this forest type, and
seems in Norway to be a more or less habitat-
specific calcareous lime forest species.
However, a few finds are documented also
from margins of calcareous pine forest-
shrublands, and from rich (but not calcareous)
oak-lime forests. Possibly it can occur also in
calcareous grasslands in Norway, as do the
related E. prunuloides and E. madidum (E.
bloxamii coll.). Also outside Norway, this
species seems to be bound to calcareous
habitats, including loamy soils, and prefe-
rentially grows in deciduous forests (Quercus
robur, Q. cerris, Fagus, Carpinus, Populus),
but is also recorded in calcareous grasslands.
So far, it has been recorded from Norway,
Germany, Great Britain and Italy (Noorde-
loos 2004). Morgado et al. (2013) described
a variety with violaceous tinges in the pileus
as E. ochreoprunuloides var. hyacinthinum
from a calcareous grassland in Wales. We
have never seen such distinctly bluish
variants in Norway.
Entoloma bloxamii s. l. in Norway
Entoloma bloxamii in its traditional morpho-
logical concept is an iconic species with its
blue, tricholomatoid basidiocarps. This morpho-
species is fairly easy to recognize in the field.
In Europe it appears to be a signal species for
various types of ancient and mycologically
valuable, poorly managed pastures on cal-
careous soils (Noordeloos 2004), and it has a
red-list status in many countries. Entoloma
bloxamii sensu lato is also reported from North
America and Australasia, and was further-
more placed recently on the Global Fungal
Red List (http://iucn.ekoo.se/iucn/species_view/
However, E. bloxamii sensu lato consists of
several, molecularly and morphologically
well-separated species (Morgado et al. 2007,
Ainsworth et al. 2018). Occurrences outside
Europe refer invariably to a range of similar,
but different species (Morgado et al. 2013). In
42 AGARICA vol. 39
Brandrud et al.
Europe at least four blue taxa can be recog-
nized, viz. E. bloxamii s. str., E. madidum, E.
atromadidum, and E. ochreoprunuloides var.
hyacinthinum. They are morphologically
separated, and identifiable. So far, only three
collections have been sequenced from Norway,
two from forests and one from grassland, and
they all belong to E. madidum. Likewise, one
grassland collection from Sweden was sequen-
ced, and appeared to be E. madidum (Brandrud
et al. 2018). More studies are necessary to
see if we have more taxa in the Nordic coun-
tries. And furthermore, if they have different
ecological preferences, since the collective
species E. bloxamii shows a wide range of
habitats from calcareous, (semi)natural grass-
lands-shrublands to calcareous lime- and
Collections sequenced: NORWAY. Nord-
Trøndelag: Steinkjer, Litl-Gaulstad, margin of
calcareous pine forest, 09 Sept 2009, H. Holien,
U.-B. Bøe, (O-69505). Oppland: Nord-Fron,
Syltebakkane, calcareous, dry, semi-natural
meadow, 6 Sept 2005, J. B. Jordal, JBJ3025
(O-158205). Telemark: Porsgrunn, Kongkleiv-
åsen S, calcareous Tilia forest, 30 Aug 2919,
T.E. Brandrud, TEB 280-19 (O). SWEDEN.
Jämtland: Åre, Alsen NØ, Glösa, TEB 327-16
3. The Claudopus clade
The Claudopus clade, more or less referable
to subgen. Claudopus in Noordeloos (1992),
includes a number of small to very small,
pleurotoid, more or less lignicolous species
with reduced, lateral stipe (referable to sect.
Claudopus in Noordeloos 1992). In Norway,
E. byssisedum seem to be the only widespread
species within this group. However, the
species might sometimes have been treated
collectively, and all the Norwegian material
of this should have been revised, which
unfortunately was not possible within the
frames of our Entoloma project.
Entoloma jahnii Wölfel & Winterh.
Characteristics: Habit pleurotoid. Pileus 1–15
mm broad, membranaceous, hemispherical to
convex then applanate, with involute margin,
not hygrophanous, not translucently striate,
white to pale pink, densely and finely white
hairy-tomentose all over. Lamellae, adnate-
emarginate to almost free, distant, white then
purely pink with entire, concolorous edge.
Stipe well developed in young specimens only,
soon lateral and reduced to almost lacking,
white tomentose. Context very thin, smell
and taste indistinct. Spores (9–)10–14(–15) ×
7.5–11(–11.5) µm, very variably shaped, iso-
to heterodiametrical, 5–6 angular in side-view
with pronounced angles. Basidia 4-, rarely
also 2-spored. Pileipellis a cutis with transit-
ions to a trichoderm of cylindrical to inflate,
5.0–15 µm wide hyphae, with abundant
capitate terminal elements. Similar capitate
cystidia are found on stipe surface. Clamps
Habitat and distribution: Usually found on
rotten wood and bark of deciduous trees (Alnus,
Fraxinus, Betula, Quercus) in moist and dry
woods and copses. Distribution in Europe
poorly known. The Norwegian, verified find
was from swampy Carex vegetation.
Collection sequenced: NORWAY. Aust-Agder:
Arendal, Tromøy (v/Alvekilen), on the base
of Carex sp., 30 July 2003, T. H. Dahl, 87/03
Comments: Entoloma jahnii is a very small
and inconspicuous species within the group
of pleurotoid Entoloma, which to the present
day is poorly understood. The species is char-
acterized by white, hairy-tomentose pileus and
stipe with capitate cystidia, and large spores.
There are indications that more species occur
with capitate pileo- and/or cheilocystidia, but
with different spores. The Norwegian collec-
tion, however, fitted with the type-sequence,
so its occurrence is confirmed. The species
AGARICA vol. 39 43
Brandrud et al.
are here reported new to Norway. The Nor-
wegian material was originally identified to
E. albotomentosum, which might superficially
resemble our species, but E. albotomentosum is
fibrillose, not hairy-scaly, and lack the capitate
pileo- and caulocystidia of our species. Moist
habitats in ditches, rather similar to the Nor-
wegian find, are reported for E. jahnii from
Denmark (J. Heilman-Clausen, pers. comm.),
and the species are nicely illustrated in
The identity of the Danish material should
however been verified by sequencing, since
apparently more taxa are hidden in this
4. The Leptonia clade
Leptonia s. str. comprises species with often
vivid lilac-violaceous colours, a strongly
fibrillose pileus and fibrillose to squamulose
stipe, hyphae with clamp connections and
habitat on organic debris, litter-layer or on
rotten wood (Morozova et al. 2014). The
group forms a very distinct, well-supported
clade, and was by Morozova et al. (2014)
treated as an own subgenus. In the latter study,
new phylogenetic evidence was shown not to
alter much the morphology-based taxonomy of
Leptonia s. str. Formerly, however, Leptonia
was used in a wide sense, also covering many
grassland species without clamps, now in-
cluded in subgenus or clade Cyanula (see
e.g. Noordeloos 1992). Clade Cyanula will
be treated in the third, forthcoming paper in
Agarica. Two Leptonia species new to Nor-
way are presented below. Another member of
the Leptonia clade new to Norway, E. chytro-
philum, is treated in another paper in the
present volume of Agarica. In addition to
these three new ones, the following species
from Leptonia s. str. are ITS-verified from
Norway: E. dichroum, E. euchroum, E. lampro-
pus, E. sublaevisporum and E. tjallingiorum
(E. placidum probably also occurs, but is so
far not verified).
Entoloma percoelestinum O.V. Morozova,
Noordel., Vila & Bulyonk. Fig. 6 B.
Characteristics: Pileus 5–12 mm broad, conical
or hemispherical with umbo, not hygrophanous,
not translucently striate, with straight margin,
radially fibrillose, squamulose at centre, uni-
formly dark blue, blackish blue or black.
Lamellae moderately distant, adnate-emarginate,
ventricose, white, becoming pinkish, with
entire concolorous edge. Stipe 20–40 × 1–2
mm, cylindrical, longitudinally fibrillose
striate or almost smooth, concolorous with
pileus, whitely tomentose at base. Context
thin, concolorous with the surface. Smell
indistinct. Spores 6.5–8.5(9.0) × 5.0–6.5 μm,
heterodiametrical, with 7–9 blunt angles in
side-view, almost nodulose. Basidia 4-spored.
Cheilocystidia absent. Pileipellis a trichoderm
of cylindrical to slightly inflated hyphae 10–20
μm wide with blue intracellular pigment.
Habitat and distribution: The only collection
from Norway was collected in a rather exposed
coastal habitat with drifting sand, in a heath-
like grassland. The Spanish material was col-
lected in montane Quercus pubescens-Fagus
forest and a (planted) coniferous forest, the
Russian collections are from either coniferous
forest (Pinus; also planted), mixed forest or
Fraxinus copses (Morozova et al. 2014).
Collection sequenced: NORWAY. Rogaland:
Sola, Vigdelstranda, in coastal, grazed sandy
pasture/heath, 23 Sept 2010, leg. John Bjarne
Jordal, John Inge Johnsen (O-294539).
Comments: Entoloma percoelestinum is a
species, as the name suggests, similar to E.
coelestinum, differing mainly by the almost
nodulose spores and longitudinally fibrous
stipe. It belongs to the Leptonia clade on
account of the clamped hyphae and tricho-
dermal pileipellis. The species was described
as new in Morozova et al. (2014), and in all
44 AGARICA vol. 39
Brandrud et al.
phylogenetic analyses, it comes out as a sister
species to E. coelestinum. It is so far known
from five sites, indicating a very wide distri-
bution; two in Russia, two in Spain (incl. type)
and the one here reported from SW Norway.
The ecological preferences of E. percoelestinum
are apparently also very broad, since it has
been found in habitats ranging from exposed
heath-like grasslands to damp forests. It
should be noted that both in Siberia and in
Norway, it is found in sandy soils.
AGARICA vol. 39 45
Figure 6. A) E. venustum. Holotype; B). E. percoelestinum, T. Bulyonkova (LE254327); C) E. cuboidoalbum.
Holotype; D). E. ritae. TEB 134-14. Photos: A) F. Hampe; B) T. Bulyonkova; C) A. Hausknecht; D) T. E. Brandrud.
Brandrud et al.
Entoloma venustum Wölfel & F. Hampe.
Fig. 6 A.
Syn.: Entoloma callichroum var. venustum
(Wölfel & F. Hampe) O.V. Morozova,
Noordel. & Vila.
Characteristics: Pileus up to 30 mm, conico-
convex to convex with small umbo, not hygro-
phanous, not translucently striate, brightly
pinkish-lilac, finely squamulose. Lamellae
distant, emarginate-adnate, lilacinous, or
whitish with bluish tinges towards edge, then
pinkish. Stipe 20-50 x 203, cylindrical, steel
blue, fibrillose, not polished. Spores 11.5–13.0
(–16.0) × 5.7–8.6 μm, heterodiametrical, with
6–8 moderately pronounced angles in side
view. Cheilocystidia present. Pileipellis a
trichoderm with intracellular pigment. Clamps
Habitat & distribution: We have just one con-
firmed sequenced collection from Norway,
and it was found in a pasture with Juniperus
on calcareous soil. According to Morozova et
al. (2014) the species is elsewhere recorded
in moist Alnus and Betula forests, park-like
secondary habitats and once also in grasslands.
E. venustum is so far known from Norway,
Germany, Belarus, Western Siberia, and the
Russian Far East.
Collection sequenced: NORWAY. Rogaland:
Rennesøy, Brimse, middle of the island, in
pasture, 2 Oct 2006, J. I. Johnsen, J. B. Jordal
Comments: This is a striking species with
bright lilac tinges. Morozova et al. (2014)
considered E. venustum a variety of E. calli-
chroum. Both species are indeed very similar
morphologically, and might in many cases
only be distinguishable with the help of
molecular markers. However, on average, the
spores seem narrower and more distinctly
angled in E. venustum, and the cheilocystidia
seem more pronounced. The degree of morpho-
logical differentiation needs further study,
but the differences in ITS are significant, and
justify recognition on specific level. Entoloma
callichroum is so far not ITS-verified from
With five finds from Russia (mainly
Siberia and Far East) and one from Belarus,
E. venustum seems to be a mainly eastern
species within the boreal-boreonemoral
Eurasian range. Like many of its relatives E.
venustum apparently has a wide range of
habitats, so far recorded in Alnus-Betula
forests, parks and grasslands. It is noted on
wood or probably leaf litter, - in grasslands
probably on grass litter or woody remnants.
5. Species of uncertain phylogenetic position
new to Norway
5.1. Entoloma excentricum and E. porphyro-
cephalum: an example of semicryptic
Entoloma porphyrocephalum (Noordel. &
Wölfel) Noordel., Brandrud & Dima,
comb. & stat. nov. Fig. 7 B, C and 8
Basionym: Entoloma excentricum var. por-
phyrocephalum Noordel. & Wölfel in Int. J.
Mycol. Lich. 1(1): 54 (1982)
MycoBank MB# 83376
Characteristics: Pileus up to 60 mm broad,
convex with depressed centre, not hygrophan-
ous, not translucently striate, when young
very finely fibrillose-tomentose, soon almost
glabrous; initially whitish, soon pale greyish
brown, ivory-grey, becoming more ochraceous
to pale pinkish brown or porphyraceous brown.
Lamellae broadly adnate to slightly decurrent,
purely white then pink with serrulate, con-
colorous, often developing a (spotwise) brown-
ish edge. Stipe 65 × 4–6 mm, cylindrical,
slightly twisted, slightly swollen at base,
sometimes slightly eccentrically attached to
pileus, entirely densely white pruinose, some
with small drops exudated at apex; more or
46 AGARICA vol. 39
Brandrud et al.
less concolorous with pileus, becoming leather-
brown to porphyry brown tinged, white tomen-
tose at base. Smell faint to somewhat farina-
ceous. Taste farinaceous. Spores 10–13.5 ×
7–8.5 µm, very large and heterodiametrical.
Conspicuous fusiform to lecithiform cystidia,
up to 100 µm long occur on sides and edge of
the lamellae, as well as on the stipe surface.
The basidia often develop brown pigment
with age (necropigment). Clamps present.
Habitat & distribution: In Norway ITS-verified
from three sites; two from outer Oslofjord
(Bamble) from shallow-soil limestone plateau;
in near-shore grassland-shrubland near sea-
shore and grassland-like opening (old track) in
calcareous Pinus sylvestris forest. The third
verified collection is from a coastal shell-bed,
grazed meadow in Brønnøy, Nordland.
Collections sequenced: NORWAY. Nordland:
Brønnøy: Urdstabbvika, UTM 33W UN
73818,64534, calcareous shore meadow
(shell-bed) which was partly mown, 3 Sept
2009, Geir Gaarder 5738 (O-293670).
Telemark: Bamble, Eikstrand, limestone with
shallow soil (grassland-shrubland), 20 Aug
2012, T. Læssøe, A. Molia AM-196c-2012
(O-245576). Bamble, Røsskleiva nature reserve,
calcareous, semi-open pine forest, along
track, 12 Aug 2016, T. E. Brandrud, B. Dima,
TEB 197-16 / DB6047 (O; CAFUN090-17).
Comments: Morphologically, E. excentricum
s.l. takes a rather isolated position in the genus
Entoloma with its large spores, prominent
and large cystidia, and dark granular pigment
(necropigment) in the hymenial elements.
For this reason, Noordeloos (1981) created
the subgenus Allocybe for it. Noordeloos and
Wölfel described an aberrant collection with
more porphyry brown cap and stipe with
caulocystidia, as E. excentricum var. porphyro-
cephalum (Noordeloos 1982). An interesting
study by Senn-Irlet & Woltsche (2002), how-
ever, demonstrated that the variability in E.
excentricum can be very large, particularly
with respect to pileus colour and presence of
caulocystidia. They observed a population
over several years and concluded that the
differences indicated as diagnostic/differential
between the two varieties was seen as gradual
variation within the population studied, and
therefore must be considered as phenotypic
variation rather than of taxonomic significance.
However, during our recent studies we
successfully amplified and sequenced the
ITS region of the holotype of E. excentricum
var. porphyrocephalum, which appeared to
be different from typical E. excentricum
(where no type exists). In addition, several
collections of E. excentricum from Norway
fit with this type sequence. Morphologically,
some have slightly more brown coloured
basidiocarps, others were, however, rather
similar to typical E. excentricum. So it appears
that within the complex of E. excentricum two
semi cryptic species can be distinguished,
one of them sometimes differing also by
having porphyry-brown tinges in the pileus.
This calls for a larger scale study on many
collections from whole Europe.
Entoloma excentricum and E. porphyrocepha-
lum have very similar habitat-preferences,
based on the Norwegian, ITS-verified col-
lections of the two taxa. They both occur
mainly in dry grassland-shrublands of coastal
shallow-soil limestone plateaus, and coastal,
grazed, calcareous shell-beds and dune slacks.
So far, three samples are sequenced and veri-
fied of E. porphyrocephalum (outer Oslofjord,
Nordland) and four of E. excentricum s. str.
(inner and outer Oslofjord, and a steppe-like
site in continental Gudbrandsdal). There seems
to be a core area for both taxa at Porsgrunn-
Bamble, Telemark county, outer Oslofjord
(cf. Brandrud 2010, Brandrud and Dima 2017,
as E. excentricum). Here the two might be
co-occurring and seem to have exact the
AGARICA vol. 39 47
Brandrud et al.
same ecological requirements. In Porsgrunn-
Bamble they occur on almost barren limestone
between seashore and adjacent calcareous
pine forests (also in openings in the forests),
often together with E. fridolfingense and E.
mougeotii. Furthermore, they are collected
from calcareous, semi-natural, grazed
meadows, especially near the seashore on
48 AGARICA vol. 39
Figure 7. A) E. excentricum TEB 189-16 / DB6044; B, C). E. porphyrocephalum; B) AM-196c-2012; C) TEB 197-
16 / DB6047. Photos: A) B. Dima; B). A. Molia ; C) B. Dima.
Brandrud et al.
shell-beds and dune slacks, of coastal Nord-
land north to Bodø (the northernmost records
of E. excentricum coll. in the world). In fact,
a rather large population of E. excentricum
coll. is documented from coastal Nordland.
So far, only one of those are verified by
sequencing, appearing to be E. porphyro-
cephalum. More sequencing is needed to see
if both taxa, or only E. porphyrocephalum
are occurring in North Norway. Outside
Norway, E. porphyrocephalum is so far ITS-
verified only from Germany (Bayern, type).
5.2. Entoloma ritae Noordel. & Wölfel Fig.
Characteristics: Pileus up to 20 mm broad,
convex to applanate, slightly undulating due to
concentric depression, with blunt or slightly
depressed centre, not hygrophanous, not trans-
lucently striate, tomentose; rather uniformly
pinkish to vinaceous brown. Lamellae adnate-
emarginate, whitish, then pink. Stipe 40–60
× 2 mm, cylindrical, yellowish pink to pale
brownish yellow, paler than pileus, more or
less polished, base with some white tomentum.
Smell and taste indistinct. Spores 8.0–12 ×
6.5–9 um, (5–8 angled with pronounced
angles. Basidia 4-spored. Lamella edge
heterogeneous. Cheilocystidia pronounced,
rendering the lamellae edge more or less
sterile (Norwegian collection), or in groups
among basidia (holotype), 25–55 × 5.0–20
µm, cylindrical to clavate. Pileipellis a cutis
with transitions to a trichoderm of up to 17
µm wide hyphae with intracellular pigment.
Habitat & distribution: In calcareous, semi-
natural grasslands; recorded in middle boreal,
mown meadow (Norway) and subalpine
meadow (Italy). So far only known and
sequence-verified from two localities; in SE
Norway and N Italy (Trentino), respectively.
The Norwegian collection was from a cal-
careous district near the lake Randsfjord.
AGARICA vol. 39 49
Figure 8. Entoloma porphyrocephalum. TEB 210-19. [Røsskleiva nature reserve.] photo: B. Dima.
Brandrud et al.
Collection sequenced: NORWAY. Oppland:
Lunner, S. Oppdalen, T. E. Brandrud, TEB
134-14 (NOBAS604-15; O-247988; originally
identified as E. indutoides).
Comments: The Norwegian collection repre-
sents the first report from N Europe and the
second known collection of this species (ITS
verified with the type sequence). It was origi-
nally described from a calcareous meadow of
N Italy (Noordeloos and Wölfel 1997). The
species is a characteristic one with its vina-
ceous brown, undulate, fibrillose pileus, and
must be one of our most rare Entoloma taxa.
Entoloma ritae resembles a Cyanula species
on account of the general habit and combi-
nation of fibrillose pileus and polished stipe,
but micro morphologically it does not fit there
because of the clamped hyphae and simple,
cutis-like pileipellis structure. Noordeloos
(2004) suggested therefore that it would be
better placed in section Roseicaules of sub-
genus Alboleptonia. Our preliminary phylo-
geny analysis suggest that it is indeed nested
within the Alboleptonia clade, but apparently
with a rather isolated position, not very close
to typical representative of Alboleptonia, such as
E. sericellum. For the time being, its phylo-
genetic and taxonomic position remains
5.3 Entoloma cuboidoalbum Noordel. &
Hauskn., Fig. 6 C.
Characteristics: Basidiocarps collybioid. Pileus
about 10 mm broad, applanate, white not trans-
lucently striate, opaque, subfelted; lamellae
adnate to slightly decurrent, whitish, then
pink; Stipe cylindrical, white to pale yellow,
polished, cartilaginous, stiff, swollen at base;
smell of honey. Spores 9–12 × 7.5–11 µm,
4–5 angled in side-view. Cystidia absent.
Ecology and distribution: In Norway recorded
in (base-)rich, steep, thermophilous deciduous
forest with Quercus and Corylus (W Norway)
and in semi-open, calcareous Pinus sylvestris-
Corylus avellana forests along track (under
pipeline). The type and the Netherlands col-
lection were found in a moist, deciduous forest
(type with Alnus incana) on wet, heavy soil.
Collections sequenced: NORWAY. Rogaland:
Hjelmeland, Hetlandsbygd below Kvitefjell,
westfaced deciduous forest, on the ground,
26 Sep 2009, O. Førland, J. B. Jordal (O-
252008). Telemark: Porsgrunn, Frierflogene
northeast (under pipeline), T.E. Brandrud, B.
Dima, 21 Sept 2015, TEB 452b-15 (O).
Comments: The Rogaland collection was first
thought to be E. sericellum (noted in the field),
but the spores had 4–5 angles, and was then
labelled E. aff. rhombisporum. The Telemark
collection was in the field identified to E. aff.
neglectum due to the adnate-decurrent lamel-
lae. The ITS sequences, however, matched
100% with the holotype of E. cuboidoalbum,
a species recently described from Austria and
recently also recorded from the Netherlands.
It differs from E. sericellum and other taxa in
the Alboleptonia group mainly by the 4-5-
angled spores. Entoloma cuboidoalbum seems,
according to the available data, to be a more
strict forest species than E. sericellum, which
is more frequent in heathlands and pastures.
E. cuboidoalbum was however, co-occurring
with E. sericellum in Porsgrunn as well as with
another white taxon in the group (E. percan-
We appreciate the permission to use photo-
graphs provided by Tatiana Bulyonkova,
Inger-Lise Fonneland, Felix Hampe, Gerrit
Jansen, Anton Hausknecht, Thomas Laessøe,
Henk Pras, Marjon van der Vegte, and Jan
Vesterholt. We thank Tobias G. Frøslev for
providing information on the type sequence
of E. caeruleopolitum. The Norwegian
Entoloma project 2015–2017 was financed
by the Norwegian Taxonomy Initiative, with
funding from the Norwegian Biodiversity
50 AGARICA vol. 39
Brandrud et al.
Information Centre (NBIC). The majority of
our material was sequenced through NorBOL,
and we thank G. Marthinsen and K. Bendiksen,
NHM, University of Oslo, as well as R. Blaalid,
NINA Bergen for performing the barcoding
work. Furthermore, we thank Pablo Alvarado
(ALVALAB, Santander, Spain), for sequen-
cing another, substantial part of our material
(collections labelled ALV). Sequencing of E.
chamaemori material from Holmvassdalen
nature reserve (Nordland) was financed by
Nord University. The foundation Rijksher-
bariumfonds Dr. E. Kits van Waveren sup-
ported the sequencing of type- and other
valuable material for this study and enabled
the necessary travelling for MEN.
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