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Type study of
Mycena phaeophylla
reveals its conspecificity with
M. clavata
Anna Ronikier
1
Institute of Botany, Polish Academy of Sciences, Lubicz
46, 31-512 Krako´w, Poland
Arne Aronsen
Torødveien 54, N-3135 Torød, Norway
Abstract
: An extensive taxonomic study based on the
type collection of
Mycena phaeophylla
revealed its
conspecificity with
M. clavata
. Redescription of
Ku¨hner’s original collections of
M. phaeophylla
is
provided as well as a detailed description of
M.
clavata
based on European material. A lectotype for
M. phaeophylla
is designated. Additional study on
collections of
M. speirea
confirmed that this closely
related species can be separated easily from
M.
clavata
. New key to subsection
Omphaliariae
of
section
Hiemales
is proposed.
Key words:
Basidiomycota, Europe, North
America, subsection
Omphaliariae
, taxonomy,
Tricholomatales
INTRODUCTION
Section
Hiemales
Konrad & Maubl., subsection
Omphaliariae
Ku¨hner ex Maas Geest. comprises
species of
Mycena
characterized by small to medium
size basidiomes, white to gray-brown, absence of odor,
arcuate lamellae and nonamyloid spores (Maas
Geesteranus 1991a). Apart from the widely distribut-
ed and common species
M. speirea
(Fr.) Gillet the few
others belonging to this subsection are rare or
insufficiently known. This is why there are many
difficulties in species delimitation in this group. Two
problematic species where identity and affinities to
one another have not been demonstrated clearly are
Mycena phaeophylla
Ku¨hner and
M. clavata
(Peck)
Redhead. The recent proposal to treat
M. phaeophylla
synonymous with
M. speirea
(Horak 2005) brought
additional complications to the taxonomy of the
subsection.
M. clavata
was described from North America in
the genus
Omphalia
by Peck (1898). Redhead (1986),
based on the type study of
Omphalia clavata
Peck,
proposed the new combination
Mycena clavata
(Peck) Redhead. He also synonymized another
American species,
Mycena thujina
A.H. Sm., with
M.
clavata
. Smith (1947) synonimized
M. phaeophylla
with
M. pallida
(Murrill) A.H. Sm., but this was
rejected by Redhead (1986) and Maas Geesteranus
(1991a) who, after re-examination of the type of this
taxon,
Omphalopsis pallida
Murrill, concluded that it
does not belong to genus
Mycena
.
Mycena phaeophylla
was described by Ku¨hner (1938), and since then it has
been reported from several countries although it
seems to be rare in Europe.
Difficulties in the precise identification of speci-
mens belonging to
M. phaeophylla
and
M. clavata
collected or examined recently by us encouraged us
to study the type collection of
M. phaeophylla
as well
as to re-examine other collections of
M. phaeophylla
and
M. clavata
deposited in some European herbaria
to find the answer to the question about their
conspecificity. Comparative analysis of
M. speirea
also
was included.
MATERIALS AND METHODS
The revision is based on the study of 35 European
collections labeled ‘
Mycena phaeophylla
’or‘
M. clavata
’:
seven from C, five from S, 13 from G (including the type),
six from KRAM, one from O and three other Norwegian
collections (herbarium Aronsen). Some collections were
misidentified therefore they were excluded from the
present study. All descriptions of both species available in
the literature (Peck 1898; Ku¨hner 1938; Smith 1947; Favre
1948; Redhead 1986; Maas Geesteranus 1991a, b) also were
taken into consideration. The type collection of
M. clavata
was not examined because it is scanty; Maas Geesteranus
(1991a) was allowed to study only the pileipellis and stipe
cortex. We recognize Peck’s type description and especially
the notes from Redhead’s (1986) and Maas Geesteranus’
(1991a) re-examinations as sufficient for our analysis. In
addition five collections of the most closely related species,
M. speirea
from O and KRAM, were examined to check its
affinity to
M. clavata
.
The material was examined following the standard
methods used in the taxonomy of Basidiomycetes. The
spores were measured in 5% KOH. Whenever possible 30
spores of each collection were measured (apart from a few
collections where not enough mature spores could be
found). For each collection spore length and width ratio
(q) were calculated and the average quotient value (q
av
)is
given in the description. Melzer’s reagent was used to check
amyloidity of spore walls. Spores to be measured were taken
when possible from natural spore deposit present at stipe
apex or pileus surface. Drawings of microcharacters were
made with a drawing tube (Nikon Y-IDT). All measurements
were taken directly through the microscope (under oil
immersion objective), not from drawings. The description
of morphological characters is based on our own observa-
Accepted for publication 13 August 2007.
1
Corresponding author. E-mail: a.ronikier@ib-pan.krakow.pl
Mycologia,
99(6), 2007, pp. 924–935.
#
2007 by The Mycological Society of America, Lawrence, KS 66044-8897
924
tions as well as notes of collectors included in herbarium
material and descriptions available from literature. SEM
pictures were taken at the Laboratory of Field Emission
Scanning Electron Microscopy and Microanalysis, Institute
of Geological Sciences, Jagiellonian University, Krako´w, with
a Hitachi S-4700 microscope at 20 kV and a working
distance of about 12 mm. Air dried spores were coated
with gold. Chart presenting spore size was prepared with
Statistica 6 software.
RESULTS
When describing the new species
M. phaeophylla
Ku¨hner (1938) gave descriptions of two collections
representing two forms: ‘‘te´trasporique’’ and ‘‘bispor-
ique’’. We studied both collections and provide their
detailed descriptions based on our own observations.
Mycena phaeophylla
Ku¨hner, Le Genre Mycena: 590,
687, F
IG. 270. 1938.
Diagnosis
(Ku¨ hner 1938)
. «Omphalia, Mycenarii, Cam-
panellae.» Sporis hypisque non amyloideis. Pileo (5–
10 mm) semiglobato, magis minusve striato, fusco vel e
fusco griseo. Lamellis distantibus, fuscidulis vel e fusco
murinis, latissime adnatis atque uncinato-decurrentibus.
Stipite (2–4 cm 3 0.7–1 mm) deorsum hirto, e fusco pallide
murino, sub lente pruinoso. Sporis 6.2–9.5 3 4.5–7.2
mm.
Cystidiis solum in marginibus lamellarum praesentibus,
filiformibus vel fusiformibus. Pigmento fusco pilei in tunica
hypharum concreto. Ad stipes vel cortices muscis oblitas et
humi jacentes.
‘‘La forme te´trasporique’’.—
Collection: FRANCE.
ISE
`
RE: Grande Chartreuse, sur souche pourrie
moussue des bois meˆle´s,
leg. R. Ku¨ hner
, 10 aouˆt
1934. ‘‘GC 64’’, No. G53734. The collection is pre-
served in a paper envelope containing: one-half of
a carpophore (a cap with a short fragment of stipe)
0.4 cm diam, brown, with half a stipe (whole length),
orange brown, pruinose at upper part, with pale
rhizomorphs at base, lamellae decurrent (at least
seven reaching the stipe); one-half of a cap (probably
the half of the previous one), the same color, at least
eight lamellae reaching the stipe, edge of lamellae
slightly ciliate; two lower halves of stipe, less than
0.5 mm diam; two small pieces of a cap.
Spores 7–9 3 6–7
mm, av. 7.65 3 6.08 m m(q5
1.08–1.45; q
av
5 1.26), broadly elliptic, broadly
amygdaliform to subglobose with rounded or more
often slightly conical apex (F
IG. 1A). Basidia 28–37 3
6.5–8.5
mm, narrowly clavate, with four sterigmata and
a basal clamp (F
IG. 1B). Cheilocystidia abundant, 44–
88 3 6.5–13
mm, narrowly lageniform to fusiform,
usually with long, often flexuous neck, 2–5
mm wide at
apex, with a basal clamp (F
IG. 1C) and sometimes
capped with a large drop of gelatinous matter. Near
the margin of a cap another type of cheilocystidia was
observed; they were scarce, narrowly clavate with
irregular outgrowths (F
IG. 1D). Pleurocystidia not
seen, but some cystidia lying on the lamella surface
close to the lamellar edge were observed (F
IG. 1E).
Lamellar trama not staining with Melzer’s reagent.
Caulocystidia 42–62 3 6–7
mm, cylindrical and flexu-
FIG.1. A–G.
Mycena phaeophylla
(coll. G 53734 –
LECTOTYPE). A. Spores. B. Basidia. C. Cheilocystidia. D.
Second type of cheilocystidia. E. Cross section of lamella
edge with cheilocystidia laying at edge and lamella surface
close to the edge. F. Caulocystidia. G. Elements of pileipellis.
Bar 5 10
mm.
RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
925
ous, numerous especially at stipe apex (FIG.1F),in
tufts or more sparsely distributed down the stipe, at
the stipe apex more similar to the cheilocystidia,
slightly broadened at lower part. Hyphae of the
cortical layer of the stipe smooth. Pileipellis com-
posed of clamped hyphae 2–3
mm wide, covered with
branched, densely packed excrescences (F
IG. 1G),
some diverticulae are elongated and protruding
above the pileipellis surface forming pileocystidia-like
hairs. Lower layers (hypoderm and context) com-
posed of wider hyphae 2.5–7
mm diam, distinctly
encrusted with brownish pigment. Clamp connec-
tions present.
‘‘La forme bisporique’’.—
Collection: FRANCE. ISE
`
RE:
Grande Chartreuse, 4 ex. sur e´corces tombe´es
moussues dans les bois meˆle´s,
leg. R. Ku¨ hner
, 15 aouˆt
1934. ‘‘GC 101’’, No. G53735. The collection is
preserved in a paper envelope containing: one
carpophore, cap brown, 60.5 cm diam; one-half of
a cap (lamellae with edges cut out); five fragments of
stipes (1–2 cm long, ,0.5 mm diam), stipe orange-
brown, upper parts of stipe pruinose, lower part
covered with pale, long rhizomorphs; four small
fragments of a cap, lamellae mostly with damaged
edge.
Spores 7–9 3 5.5–7.5
mm, av.: 8.11 3 6.6 mm(q5
0.93–1.42; q
av
5 1.24), broadly elliptic, broadly
amygdaliform to subglobose with rounded, or more
often slightly conical apex (F
IG. 2A). Basidia 22–27 3
7–9
mm, clavate, with two sterigmata (FIG. 2B). Chei-
locystidia abundant, 35–50 3 8–11
mm, narrowly
lageniform to fusiform, usually with long, often
flexuous neck 2–3.5
mm wide at apex, sometimes
capped with a large drop of gelatinous matter
(F
IG. 2C). Pleurocystidia not seen. Lamellar trama
not staining with Melzer’s reagent. Caulocystidia 35–
78 3 5–7
mm, cylindrical and flexuous, often with
irregular outgrowths (F
IG. 2D), numerous especially
at stipe apex, in tufts or more sparsely distributed
down the stipe. Hyphae of the cortical layer of the
stipe smooth or occasionally with sparse small
excrescences. Pileipellis composed of hyphae 2–
3.5
mm wide, covered with branched, densely packed
excrescences (F
IG. 2E); some diverticulae are elon-
gated and protruding above the pileipellis surface
forming pileocystidia-like hairs. Lower layers (hypo-
derm and context) composed of wider hyphae 3–
10
mm diam, distinctly encrusted with brownish pig-
ment. Clamp connections absent.
Ku¨hner (1938) described these two forms, but he
did not recognize them as separate taxa. The two
collections were found in the same locality, but one
was collected 5 d later than the other. The original
Latin diagnosis includes the variability of the two
forms described by this author. Because the two
collections should be considered as syntypes of
Mycena phaeophylla
, according to the International
Code of Botanical Nomenclature (Art. 9.4), a lecto-
type must be chosen.
LECTOTYPUS. Sur souche pourrie et moussue, dans
les bois meˆle´s, non loin du monaste`re de la Grande
Chartreuse (Ise`re), 10 aouˆt 1934, ‘‘GC 64’’, (G53734), here
designated.
The study of the collections of
M. phaeophylla
described above as well as the other collections
deposited in some European herbaria together with
careful analysis of the descriptions of the type
collection of
M. clavata
(Peck 1898, Redhead 1986,
Maas Geesteranus 1991a) let us conclude that the two
species are conspecific. The description provided
below is based on all the material we studied.
Mycena clavata (Peck) Redhead, Mycologia
78(4):523, F
IGS. 1–4. 1986.
Omphalia clavata
Peck,
FIG.2. A–E.
Mycena phaeophylla
(coll. G 53735).
A. Spores. B. Basidia. C. Cheilocystidia. D. Caulocystidia.
E. Elements of pileipellis. Bar 5 10
mm.
926 MYCOLOGIA
Rep. NY St Mus Nat Hist 51:285. 1898. TYPE: USA,
North Elba, on trunks of arbour-vitae, Chas H.
Peck, Aug 1997 (holotype: NYS).
;
Omphalia clavata
Peck, Ann Rep NY St Mus 51:285.
1898.
;
Omphalopsis clavata
(Peck) Murrill, North Am Flora
9(5):313. 1916.
5
Mycena thujina
A.H. Sm., North American Species of
Mycena: 361, pl. 86D, F
IG. 45(9, 10). 1947. TYPE:
CANADA, Ontario, near Lake Timagami, on log of
Thuja occidentalis
, 2 Sep 1936,
A.H. Smith
, 4444
(holotype: MICH).
5
Mycena phaeophylla
Ku¨hner, Le Genre Mycena: 590,
687, F
IG. 207. 1938, TYPE: FRANCE, non loin du
monaste`re de la Grande Chartreuse (Ise`re), sur souche
pourrie et mousse, dans les bois meˆle´s, 10 aouˆt 1934,
R. Ku¨ hner
, ‘‘GC 64’’, G53734 (lectotype: G).
5
Marasmiellus phaeophyllus
(Ku¨hner) Singer, Lilloa
22:302. 1951.
FIGS. 3–9
Pileus 2–12.5 mm across, at first hemispherical,
expanding to parabolic or campanulate when young,
then convex, flattening with age, and often somewhat
depressed at the center, translucently striate, pale
ochraceous brown, pale gray brown (milky coffee),
yellow brown or grayish olivaceous brown, fairly dark
when moist and young, pallescent with age, dark sepia
at the center, paler at margin, surface minutely
pruinose-pubescent, later glabrescent (F
IG. 3A). Flesh
thin, pale beige; odor absent, flavor mild. Lamellae 8–
20 reaching the stipe, strongly arcuate, long de-
current, slightly paler than cap, pale brown, grayish
sepia to beige brown with concave edge, paler than
the sides. Stipe 8–80 3 0.2–1 mm, hollow, firm, equal,
straight to curved, terete, pruinose all over, glabres-
cent, paler than the pileus, beige brown, pale
yellowish brown, watery yellowish, watery brownish
or grayish, the base covered with coarse, whitish
fibrils.
Basidia 21–39 3 5–9
mm, slender-clavate or clavate,
2-spored, clampless, with sterigmata 6–7
mm long or 4-
spored, clamped (F
IG. 4). Spores 7–11 3 5.5–8.5
(–9.5)
mm, q
av
, 1.25 (2-spored basidia), or 6.5–10.5
3 5–7.5
mm, q
av
,1.28 (4-spored basidia), broadly
ellipsoid, broadly amygdaliform to subglobose, with
rounded to slightly conical apex, smooth, nonamyloid
(F
IG. 5). Cheilocystidia 20–88 3 4–16 mm, forming
a sterile band, subcylindrical, fusiform to lageniform,
clamped or clampless, apically passing into a slender,
straight to curved or somewhat flexuous, simple,
forked or somewhat branched neck 2–5
mm wide,
often capped with a large drop of gelatinous matter
(F
IG. 6). More rarely and mostly situated near the
FIG.3. A,B.
Mycena clavata
. A. Carpophores (coll. KRAM F-54292). B. encrusting pigment in hypoderm (coll. G53738).
C.
Mycena speirea
, intracellular pigment in hypoderm (KRAM F-11290). Bars: A 5 1 cm; B, C 5 10 mm.
RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
927
pileus margin another type of cheilocystidia appears
that is shorter, 17–35 3 6–15
mm, narrowly clavate to
clavate and with several simple to branched, cylindri-
cal excrescences (F
IG. 7). Pleurocystidia apparently
absent, but cystidia may be present near the pileus
margin (F
IG. 1E). Lamellar trama not staining with
Melzer’s reagent. Hyphae of the pileipellis 2.5–6.5
mm
wide, clamped or clampless, covered with simple to
much branched excrescences up to 22.5 3 1–2
mm,
which may develop into dense masses, sometimes
becoming somewhat gelatinized, with some of the
diverticulae elongated into projecting, pileocystidia-
like hairs up to 32
mm long (FIG. 8). Lower layers
(hypoderm and context) composed of wider hyphae,
3–10
mm, distinctly encrusted with brownish pigment
(F
IG. 3B). Hyphae of the cortical layer of the stipe
1.5–3.5
mm wide, clamped or clampless, smooth for
the greater part but covered with side branches and
caulocystidia 35–78 3 3–11
mm; the caulocystidia
abundant, especially at the stipe apex, in tufts or
more sparsely distributed down the stipe, cylindrical
and curved or flexuous, often with irregular out-
growths, simple to forked, sometimes capped with
a large drop of gelatinous matter (F
IG. 9).
Collections examined
: DENMARK. NORDJYLLAND:
Rubjerg Knude Plantage, on
Pinus
, 12 Apr 1992,
leg. J.
Vesterholt
(C21363); the same place, on
Pinus
and on
Picea
,
28 May 2003,
leg. J. Vesterholt
(C43330); Rubjerg Knude,
Hjørring Kommunes Klitplantage, in litter, 17 Oct 1988,
leg.
Th. Læssøe
(C32402); the same place, on
Pinus
, 17 Oct
1988,
leg. Th. Læssøe & J.H. Petersen
(C32403); Livø, on
Abies
twigs, ass.
M. speirea
, 29 Sep 1995,
leg. Th. Læssøe
(C32056); Tole Bakker, ‘Pa˚ kvasdynge i grøft’, 2 Oct 1981,
leg. Th. Læssøe & S.A. Elborne
(C without number);
SJÆLLAND: Sorø Sønderskov, on twigs in dense heap in
fern/
Urtica
stand (
Pinus
?), ass.
Mycena speirea
,
hiemalis
,
acicula
, 18 Oct 1982,
leg. T. Læssøe & S.A. Elborne
(C
without number, TL0502); FRANCE. ISE
`
RE: Grande
Chartreuse, mixed forest, on decayed, mossy trunk, 10
Aug 1934,
leg. R. Ku¨ hner
(G53734, LECTOTYPE); the same
place, mixed forest, on fallen, mossy pieces of bark, 15 Aug
1934,
leg. R. Ku¨ hner
(G53735); Grande Chartreuse, near St
Laurent du Pont, edge of a forest, among mosses, at the
base of
Abies pectinata
trunk, 1 Nov 1933,
leg. Josserand
(G53737); AIN: vicinity of Lyon, Hauteville, 26 Oct 1956,
leg. R. Ku¨ hner
(G53733); Hauteville, 26 Oct 1953,
leg. ?
(G53738). NORWAY. VESTFOLD: Nøtterøy, Torød, one
specimen among needles under
Juniperus
, 10 Aug 1993,
leg.
A. Aronsen
(Aronsen 34/93); Tjøme: Hvasser, Sønstega˚rd,
on branches of
Juniperus communis
, 3 Dec 2006,
leg. A.
FIG. 4. A–K.
Mycena clavata
basidia. A. coll. G53729. B.
coll. G53730(401d). C. coll. G53730(401b). D. coll.
G53730(401a). E. coll. KRAM F-54292. F. coll. G53728. G.
coll. G53731. H. coll. C 32056. I. coll. C43330. J. coll.
C32403. K. coll. C without number,
leg. Th. Læssøe & S.A.
Elborne
. Bar 5 10 mm.
FIG. 5. A–N.
Mycena clavata
spores. A. coll. G53729. B.
coll. G53738. C. coll. G53730(401d). D. coll. G53730(401b).
E. coll. KRAM F-54292. F. coll. G53730(401a). G. coll.
G53733. H. coll. G53728. I. coll. G53731. J. coll. C32403. K.
coll. C 32056. L. coll. C43330. M. coll. C21363. N. coll. C
without number,
leg. Th. Læssøe & S.A. Elborne
. A–G. 4-
spored collections. H–M. 2-spored collections. N. collection
with 2- and 4-spored basidia. Bar 5 10
mm.
928 MYCOLOGIA
Aronsen
(A21/06); AKERSHUS: Frogn, Bonn, NM 967 213,
in moss, on a fallen trunk of
Picea abies
, 2 Oct 1983,
leg. R.
Kristiansen
(O163104); NORD-TRØNDELAG: Namdalseid,
Fla˚bekka˚sen skogreservat, alt. 180–240 m., on
Picea
bark in
old
Picea
forest, 3 Oct 2001,
leg. H. Holien & S.
Sivertsen;
POLAND. THE CARPATHIANS: the Western Tatra Moun-
tains, summit area of the Sarnia Skała massif, northern
slope, N 49u159530,E19u569380, alt. 1370 m.,
Pinetum mugi
carpaticum
, on litter, 22 Jun 2001,
leg. A. Ronikier
(KRAM F-
54295); the same place, among mosses (probably on wood),
4 Jul 2001,
leg. A. Ronikier
(KRAM F-54292); the same place,
among mosses, on litter, 11 Jun 2003,
leg. A. Ronikier
(KRAM F-54294); the same place, on wood (branch of
?
Pinus mugo
), 11 Jun 2003,
leg. A. Ronikier
(KRAM F-
54293); the same place, among mosses, 10 Jun 2002,
leg. A.
Ronikier
(KRAM F-54296); the same place, on litter, 20 Jun
2000,
leg. A. Ronikier
(KRAM F-54357). SWITZERLAND.
VAUD: Tourbie`re des Piguet-Dessus, close to Sentier Valle´e
de Joux, 19 Sep 1941,
leg. ?
(G53731); JURA: Tourbie`re des
Rousses, 7 Oct 1938,
leg. J. Favre
(G53728); GRAUBU
¨
N-
DEN: Val Sesvenna, near S-carl, alder forest, alt. 1900 m., 2
Sep 1952,
leg. J. Favre
(G53729); Haute Engadine, God
Trid, Val Trupschun, alder forest, alt. 1900 m., 1 Sep 1957,
leg. J. Favre
(G53730); near S-carl, left side of Val Sesvenna,
alt. 2000 m., 14 Aug 1948,
leg. J. Favre
(G53730); near S-carl,
Val Sesvenna, alt. 1900 m., 9 Sep 1944,
leg. J. Favre
(G53730).
Habitat: Growing solitary or in clusters on bark of
coniferous wood or on coniferous litter, once found
on deciduous wood (Elborne and Læssøe 1982).
Known distribution: Canada, Denmark, France,
Norway, Poland, Switzerland, USA.
DISCUSSION
Evidence for synonymy of M. clavata and
M. phaeophylla.—
The affinity of
M. clavata
and
M. phaeophylla
has been mentioned a few times in the
literature. Redhead (1986) was the first to say that
Mycena phaeophylla
might be conspecific with
M.
clavata
and put this name among synonyms of the
FIG. 6. A–L.
Mycena clavata
cheilocystidia. A. coll. G53729. B. coll. G53738. C. coll. G53730(401d). D. coll. G53730(401b).
E. coll. G53730(401a). F. coll. G53733. G. coll. G53731. H. coll. C without number, TL 0502. I. coll. G53728. J. coll. C32402.
K. coll. C 32056. L. coll. C43330. A–F. 4-spored collections. G–L. 2-spored collections. Bar 5 10
mm.
RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
929
latter but with a question mark. He was not sure
because he did not study the type material of
Mycena
phaeophylla
but only one of J. Favre’s collections (coll.
G25729). Moreover Redhead (1986) noted a few
differences between the two species: slightly larger
spores and darker colors of carpophores in European
material. Maas Geesteranus (1991a), who studied only
parts of the type, the hyphae of the pileipellis and the
stipe cortex of
Omphalia clavata
because of scanty
material of the type collection, took up the discussion
and brought forward two other differences: ‘‘
Mycena
clavata
is known to possess pleurocystidia, whereas
they are absent in
M. phaeophylla
, and the caulocys-
tidia of
M. clavata
are strikingly shaped, tortuous,
simple to somewhat branched, while they are cylin-
drical and rather banal in
M. phaeophylla
, at least
going by Ku¨ hner’s illustration.’’ In another paper
based on examination of a Norwegian collection Maas
Geesteranus (1991b) said that
Mycena clavata
may be
just as dark as indicated in
M. phaeophylla
and that it
also may possess 2-spored basidia. Still he was
reluctant to synonymize the two species because
‘‘Ku¨hner (1938) stated that both the 4- and 2-spored
forms of
M. phaeophylla
lacked pleurocystidia; in-
dicated the spores of the 4-spored form as definitely
more pip-shaped than globose; apparently did not
observe any branched excrescences of the hyphae of
the pileipellis; and failed to give information on the
nature of the substratum (wood of coniferous or
deciduous tree).’’
Our re-examination of Ku¨hner’s original collec-
tions (both 4- and 2-spored forms) of
M. phaeophylla
showed broadly elliptic, broadly amygdaliform to
subglobose spores. Indeed they are on average slightly
larger than measured by Redhead (1986) in the type
of
Omphalia clavata
, but those measurements are
within the range of
M. phaeophylla
. The type of
M.
phaeophylla
does not possess pleurocystidia, but some
cystidia were seen on the lamella face close to the
edge (F
IG. 1E) and hence did not appear as true
pleurocystidia. This feature also was observed by
Redhead (1986) in the type collection of
Omphalia
clavata
. He mentioned that ‘‘‘pleurocystidia’ occur
on the lamellae but they are confined to the area near
the lamellar edge.’’ Maas Geesteranus (1991b)
observed scarce pleurocystidia in one European
collection. Unfortunately we did not examine this
collection. We saw the specimens found by the same
collector on the same day but in another location
(coll. O163104) and we could not find true pleur-
ocystidia in this collection either. The presence of
true pleurocystidia is the only feature that we could
not confirm, either in the type collection of
M.
phaeophylla
or in any other collections examined by
us. Following Redhead’s (1986) remarks on pleur-
ocystidia of the type of
O. clavata
(lacking or present
only close to the lamellar edge) and taking into
account the scarcity of pleurocystidia in the only one
European collection (Maas Geesteranus 1991b) we
assume that either this feature is not stable or difficult
to observe in dried specimens, and therefore it should
not be considered as taxonomically important.
Ku¨hner (1938) described the excrescences of the
hyphae of the pileipellis in
M. phaeophylla
as simple
and cylindrical, different from the much branched
excrescences of
M. clavata
. Our observations however
showed branched, densely packed excrescences both
in the 4-spored and the 2-spored forms of Ku¨hner’s
collections of
M. phaeophylla
(FIGS. 1G, 2E) similar to
the observations of Maas Geesteranus (1991a) on the
type of
Omphalia clavata
. Our observations also show
that the caulocystidia in
M. phaeophylla
are shaped
much more strikingly than in Ku¨hner’s (1938)
illustration and they are similar to the caulocystidia
of
M. clavata
(FIG. 9).
Further features indicate the conspecificity of
M.
clavata
and
M. phaeophylla
. Between the much
branched excrescences of the hyphae of the pileipel-
lis some diverticulae are elongated into projecting
hairs (F
IG. 8). This was mentioned by Ku¨ hner (1938)
for the type of
M. phaeophylla
and by Redhead (1986)
for the type of
O. clavata
and was present in all
collections examined by us. These projections are
more or less cystidia-like and up to 32
mm long. They
sometimes are difficult to observe and they usually are
FIG. 7. A–E.
Mycena clavata
cheilocystidia (the second
type). A. coll. G53730(401d). B. coll. KRAM F-54292. C. coll.
G53731. D. coll. C 32056. E. coll. C without number,
leg. Th.
Læssøe & S.A. Elborne
. A–B. 4-spored collections. C–D. 2-
spored collections. E. collection with 2- and 4-spored
basidia. Bar 5 10
mm.
930 MYCOLOGIA
scarce toward the margin of the cap while they are
more abundant at the center of the cap. They vary
also in shape and size among collections. The most
clearly visible and the largest pileocystidia were
observed in collection G53733 (F
IG. 8D).
Redhead (1986) said that the cheilocystidia of
O.
clavata
vary from ‘‘narrowly ventricose with somewhat
uneven necks to occasionally branched, or with
clavate bases’’ (and with several more or less
branched excrescences, according to his illustration).
The second type of cheilocystidia with branched
excrescences was observed in the 4-spored form of
Ku¨hner’s
Mycena phaeophylla
collection as well and
also in some of the other collections that we
examined (F
IG. 7).
In the collections examined by us the necks of the
cheilocystidia and the caulocystidia were capped by
a large drop of gelatinous material (F
IGS. 2C; 6E, F, G,
K, L; 9A, F). This feature is present in
M. clavata
mentioned by Redhead (1986) as well as in
M.
phaeophylla
both in its 4- and 2-spored forms.
Ku¨hner (1938) said that the hyphae of hypoderm
of
M. phaeophylla
have brownish encrusting pigmen-
tation ‘‘a` membrane brunaˆtre, souvent ruguleuse-
ze´bre´e.’’ This feature also was observed in the type of
O. clavata
(Redhead 1986). This pigmentation is
distinct and present in all our collections (F
IG. 3B).
Encrusting pigment is rare in the genus
Mycena
.Itis
present only in two species,
M. phaeophylla
and
M.
corticola
(5
M. meliigena
[Berk. & Cooke] Sacc.),
according to Ku¨hner (1938)
Relationship with Mycena speirea.—
The species
most closely related to
M. clavata
is
M. speirea
(FIG. 10). This common and widely distributed
fungus is similar in habit; it also has nonamyloid
spores and belongs to the same group (sect.
Hiemales
,
subsect.
Omphaliariae
). It is also brown, has arcuate
lamellae and occurs on similar substrate. Horak
(2005) synonymized these species, but we do not
concur in his opinion, following Ku¨hner (1938),
Moser (1983) and Maas Geesteranus (1991a). Al-
though the two species are undoubtedly close,
M.
speirea
can be separated easily from
M. clavata
on
account of (i) the shape of the spores (
M. speirea
ellipsoid to amygdaliform, 8–11 3 4.5–5.8 mm, q
av
,
FIG. 8. A–L.
Mycena clavata
hyphae of pileipellis. A. coll. G53729. B. coll. G53738. C. coll. G53730(401d). D. coll. G53733.
E. coll. G53730(401a). F. coll. KRAM F-54292. G. coll. G53731. H. coll. G53728. I. coll. C32403. J. coll. C 32056. K. coll. C32402.
L. coll. C43330. A–F. 4-spored collections. G–L. 2-spored collections. Bar 5 10
mm.
RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
931
1.9;
M. clavata
broadly ellipsoid to subglobose, q
av
,
1.3); (ii) the shape of the cheilocystidia (
M. speirea
mostly subcylindrical;
M. clavata
mostly fusiform or
lageniform, apically passing into a slender neck); (iii)
the hyphae of the pileipellis (more branched in
M.
clavata
and with some of the diverticulae elongated
into projecting pileocystidia-like hairs); (iv) the
character of pigmentation in lower layers of the
pileipellis and pileitrama-hypoderm (
M. clavata
en-
crusting (F
IG. 3B);
M. speirea
intracellular (FIG. 3C).
Examined collections of
M. speirea: NORWAY. OSLO:
Oslo, Bygdøy, Dronningberget, on wooden litter in calcar-
eous
Tilia-Corylus
forest, 13 Aug 1979,
leg. K. Høiland
(O163000); VESTFOLD: Tønsberg, Gullkrona, on twigs in
a damp area under
Fraxinus
,
Alnus
, 6 Jun 1983,
leg. A.
Aronsen M 9/83
(O66619); OPPLAND: Gjøvik kommune,
Svennevollene naturreservat, NN 8760 (1816 I), 24 Aug
1985,
leg. T.E. Brandrud & J. Stordal
, (O163003); PO-
LAND. THE CARPATHIANS: the Kotlina Orawsko-Now-
otarska basin, on bank of the Białka River, alt. ca. 540 m.,
Alnetum incanae
, on wood, 6 Aug 1985,
leg. W. Wojewoda
(KRAM F-27144); vicinity of Nowy Targ, the Bory Now-
otarskie region, Ludzmierz, on bank of the Lepietnica
River,
Alno-Padion
, on a dead trunk, 31 May 1971,
leg. W.
Wojewoda
(KRAM F-11290).
FIG. 9. A–K.
Mycena clavata
caulocystidia. A. coll. G53729. B. coll. G53738. C. coll. G53730(401a). D. coll. G53730(401d).
E. coll. G53730(401b). F. coll. G53733. G. coll. KRAM F-54292. H. coll. C32403. I. coll. G53731. J. coll. C43330. K. coll. G53728.
A–G. 4-spored collections. H–K. 2-spored collections. Bar 5 10
mm.
932 MYCOLOGIA
Differences between 2- and 4-spored forms of
M. clavata.—
According to Smith (1934), ‘‘it has
long been known that within the genus
Mycena
there
exists an unusually large number of forms bearing
basidia which produce two instead of the usual four
spores.’’ Such forms are variously treated in different
species. For instance Maas Geesteranus (1977) dis-
covered that in the Dutch populations of
Mycena
galericulata
(Scop.) Gray the 2-spored form is more
common than the 4-spored form and that the latter
can be found in specimens collected later in the
season (Oct–Dec), while the 2-spored form can be
found throughout the year. He did not recognize the
two forms as taxonomically separate, although he said
they might represent genetically different taxa (Maas
Geesteranus 1977, 1985). Similarly Aronsen and Maas
Geesteranus (1989) describing a new species,
M.
ustalis
Aronsen & Maas Geest., report 2- and 4-spored
forms from the same locality, the latter collected
several days later than the previous one. They also did
not find any morphological differences between the
two forms that might justify their separation. On the
other hand another species
Mycena radicifera
J. Favre
recently was re-examined by Moreau and Courte-
cuisse (2003) who recognized two varieties, 4-spored,
clamped, typical variety
radicifera
and 2-spored,
clampless, apogamic
M. radicifera
var.
apogama
P.A.
Moreau & Courtec., which in addition differs from
the var.
radicifera
with a structure of suprapellis.
A few characters are slightly different between the
two 2- and 4-spored forms of
Mycena clavata
. They are
presence of clamp connections, number of sterigma-
ta, spore size as well as shape and size of basidia.
In general the 2-spored form is characterized by
slightly larger spores (7–11 3 5.5–8.5
mm) than the 4-
spored form (6.5–10.5 3 5–7.5
mm). According to our
measurements (7–9 3 6–7
mm) spores of the type
collection (4-spored form) are slightly larger than
stated in the original description; according to
Ku¨hner (1938) they were 6.2–8 3 4.5–5.7. Individual
measurements of spores of 2- and 4-spored collections
largely overlap (F
IG. 11), however mean spore values
show tendency for 4-spored form to be smaller. The
differences in spore size between the 2-spored and 4-
spored forms of the same species are widely known.
According to Smith (1934) the spores of 2-spored
forms are usually one-quarter to one-sixth bigger than
spores of 4-spored collections, but according to
Ku¨hner (1938) the difference in size might be smaller
in some species. The differences in size of basidia and
presence or absence of clamp connections usually go
together with the differences in number of sterigma-
ta. Ku¨hner (1938) demonstrates that basidia of 2-
spored forms are constantly slightly smaller than
those of 4-spored forms. Because no other clear
characteristics (e.g. in pileipellis structure, size of
carpophores, size and shape of cystidia) differentiated
the two forms of
M. clavata
we consider these small
differences not taxonomically significant and not
sufficient for recognition of separate taxa, either on
a level of variety or a form. It is possible that the 2-
spored
M. clavata
represents an apogamic form, but
in our opinion different reproductive strategy alone
should not be considered a basis for creation of a new
variety unless it is supported by stable morphological
characters suggesting an evolutionary differentiation
toward a parthenogenetic lineage (cf. Moreau and
Courtecuisse 2003). Moreover two collections (C
without number and C32403) are characterized by
co-occurrence of 2- and 4-spored basidia. We did not
find a correlation in number of sterigmata and time
of collection, as noticed for
M. galericulata
(Maas
Geesteranus 1977) and
M. ustalis
(Aronsen and Maas
Geesteranus 1989), but we noticed that the 4-spored
form was found more frequently in central and
southern Europe while it was almost absent in
Scandinavian countries.
Elborne and Læssøe (1982) noticed that the spores
of their 2-spored collection were not smooth. They
described the surface as covered with spots where the
spore wall is thinner, giving it an appearance of being
covered by small holes. We could observe this character
in the type collection (4-spored) under SEM (F
IG. 12).
KEY TO THE SPECIES OF SUBSECTION
OMPHALIARIAE
1. Spores ellipsoid, amygdaliform (q . 1.6, q
av
, 1.9), pigment in hypoderm intracellular,
brown ............................
M. speirea
FIG. 10. A–D.
Mycena speirea
. A. hypha of pileipellis. B.
spores. C. cheilocystidia. D. caulocystidia. A–C. (coll. O
163000). D. (coll. O 163003). Bar 5 10
mm.
RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
933
1. Spores globose to broadly ellipsoid (q # 1.6, q
av
, 1.3), pigment in hypoderm encrusting or in-
tracellular . . . ............................ 2
2. Cheilocystidia clavate to subcylindrical, pileipellis
with inflated excrescences, pigment in hy-
podermintracellular,poorlyvisible.......
M. alba
2. Cheilocystidia lageniform to fusiform with
simple or branched neck, pileipellis with
cylindrical excrescences, pigment in hypoderm
encrusting.....................
M. clavata
CONCLUSIONS
A taxonomic study based on the type collection of
Mycena phaeophylla
revealed its conspecificity with
M.
clavata
. The characters considered so far differenti-
ating the two species (shape of spores, structure of the
pileipellis, shape of caulocystidia) turned out to be
identical. Ku¨hner (1938) described two forms of
M.
phaeophylla
(2-spored and 4-spored), but in our
opinion the differences between them are not
sufficient for their recognition either on a level of
variety or a form. The 2-spored form differs slightly in
spore and basidia size, which usually is observed also
in other 2-spored forms of
Agaricales
. No correlation
with other characters was observed.
Mycena clavata
can be distinguished easily from
M.
speirea
on account of several features, of which the
most eminent is encrusting pigment.
M. clavata
is a rare but widely distributed species
known from North America and Europe. In Europe it
is connected mostly to coniferous wood and litter
(
Pinus
,
Picea
,
Abies
,
Juniperus
) and it occurs in
mountainous habitats as well as in northern part of
the continent; therefore it probably can be consid-
ered a mountain-boreal species.
ACKNOWLEDGMENTS
We thank David Hawksworth (University of Madrid, Spain)
for the nomenclatural advice, Pierre-Arthur Moreau (Uni-
versity of Lille, France) for valuable discussions on the
present study, Isobel Simonsen for linguistic corrections,
Richard W. Kerrigan and two anonymous reviewers for the
valuable comments on the manuscript. We also thank
keepers of herbaria C, S, G, O for kind loan of the material.
This work is partly a result of studies carried out under grant
No. 6 P04G 083 20, financed by the Polish State Committee
of Scientific Research (KBN).
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FIG. 11. Spore size of all examined collections of
Mycena
clavata
; square 4-spored collections, cross 2-spored collec-
tions.
FIG. 12. A–B. Spores of
Mycena phaeophylla
(coll. G
53734 – LECTOTYPE). Bar 5 1
mm.
934 MYCOLOGIA
Peck CH. 1898. Report of the state botanist. Ann Rep NY
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RONIKIER AND ARONSEN:
M.
PHAEOPHYLLA
,M.
CLAVATA
935
