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Anamorphs in the Strophariaceae (Basidiomycota,
Agaricales)
Grit Walther and Michael Weiß
Abstract: We describe and illustrate conidiogenesis in 21 species of Strophariaceae sensu Singer in culture, including first
reports of conidiogenesis from nine species. Two modes of thallic conidiogenesis were revealed. The first mode, present in
Hypholoma capnoides (Fr.) P. Kumm., Hypholoma fasciculare (Fr.) P. Kumm., Hypholoma marginatum (Pers.) J. Schro
¨t.,
Hypholoma subericaeum (Fr.) Ku
¨hner, Hypholoma sublateritium (Schaeff.) Que
´l., Kuehneromyces mutabilis (Schaeff.)
Singer & A.H. Sm., Pholiota lenta (Pers.) Singer, Pholiota lucifera (Lasch) Que
´l., Pholiota mixta (Fr.) Kuyper &
Tjall.-Beuk., Pholiota spumosa (Fr.) Singer, Psilocybe cf. coprophila (Bull.) P. Kumm., Psilocybe inquilina (Fr.) Bres.,
Psilocybe semilanceata (Fr.) P. Kumm., Psilocybe sp., Stropharia rugosoannulata Farl. ex Murrill, and
Stropharia semiglobata (Batsch) Que
´l., is characterized by straight or often coiled conidiogenous hyphae, not or only
slightly sympodially proliferating conidiophores lacking geniculate parts, and hyaline, rod-shaped conidia. Conidiogene-
sis in Pholiota aurivella (Batsch) P. Kumm., Pholiota gummosa (Lasch) Singer, and Pholiota squarrosa (Batsch)
P. Kumm. followed a second mode, distinguished by straight conidiogenous hyphae, sympodially proliferating genicu-
late conidiophores, and coloured, swollen conidia formed on older mycelia. These two modes also differed concerning
the number of nuclei in the conidia. Pholiota alnicola (Fr.) Singer and Pholiota tuberculosa (Schaeff.) P. Kumm. pos-
sessed specific modes of conidiogenesis that deviated from the two prevalent modes described above. Careful analysis
of anamorph descriptions compiled from literature supports these modes and allows conidia to be distinguished from
nondetaching hyphal swellings that have also been designated as conidia in the past.
Key words: Agaricales, anamorphs, conidia, conidiogenesis, Strophariaceae.
Re
´sume
´:Les auteurs de
´crivent et illustrent la conidioge
´ne
`se en culture, chez 21 espe
`ces de Strophariaceae sensu Singer,
incluant les premiers rapports sur la conidioge
´ne
`se de neuf espe
`ces. On de
´couvre deux modes de conidioge
´ne
`se thallique.
On retrouve un premier mode chez les Hypholoma capnoides (Fr.) P. Kumm., Hypholoma fasciculare (Fr.) P. Kumm., Hy-
pholoma marginatum (Pers.) J. Schro
¨t., Hypholoma subericaeum (Fr.) Ku
¨hner, Hypholoma sublateritium (Schaeff.) Que
´l.,
Kuehneromyces mutabilis (Schaeff.) Singer & A.H. Sm., Pholiota lenta (Pers.) Singer, Pholiota lucifera (Lasch) Que
´l.,
Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk., Pholiota spumosa (Fr.) Singer, Psilocybe cf. coprophila (Bull.) P. Kumm.,
Psilocybe inquilina (Fr.) Bres., Psilocybe semilanceata (Fr.) P. Kumm., Psilocybe sp., Stropharia rugosoannulata Farl. ex
Murrill et Stropharia semiglobata (Batsch) Que
´l.; ce premier mode se caracte
´rise par des hyphes conidioge
´nes droits ou
souvent enroule
´s montrant une prolife
´ration de conidiophores peu ou pas faiblement sympodiaux sans parties ge
´nicule
´es,
et des conidies hyalines en ba
ˆtonnet. Chez les Pholiota aurivella (Batsch) P. Kumm., Pholiota gummosa (Lasch) Singer et
Pholiota squarrosa (Batsch) P. Kumm. on observe le second mode qui se distingue par des hyphes conidioge
`nes droits
avec des conidiophores a
`prolife
´ration sympodiale et ge
´nicule
´s, ainsi que des conidies colore
´es et renfle
´es sur le vieux my-
ce
´lium. Ces deux modes diffe
´rent e
´galement par le nombre de noyaux dans les conidies. Les Pholiota alnicola (Fr.) Singer
et Pholiota tuberculosa (Schaeff.) P. Kumm. posse
`dent des modes spe
´cifiques de conidioge
´ne
`se qui de
´vient par rapport
aux deux modes de
´crits plus haut. Une analyse soigne
´e des descriptions d’anamorphes compile
´es a
`partir de la litte
´rature
supporte ces modes et permet de distinguer les conidies des renflements non-de
´tachables venant sur les hyphes et qu’on a
souvent appele
´conidies dans le passe
´.
Mots-cle
´s:Agaricales, anamorphes, conidie, conidioge
´ne
`se, Styrophariaceae.
Introduction
The Strophariaceae sensu Singer (1986) are characterized
by cinnamon-coloured to blackish-lilac spore prints, smooth
basidiospores that usually possess a germ pore, a pileipellis
in the form of a cutis or a trichodermium, absence of truly
free lamellae, and saprotrophy.
Since mycologists started to study mycelia in culture, nu-
merous anamorphs of Strophariaceae have been reported.
Several comprehensive descriptions of conidiogenesis were
Received 27 November 2007. Published on the NRC Research Press Web site at botany.nrc.ca on 21 May 2008.
G. Walther.1CBS Fungal Biodiversity Centre, P.O. Box 85167, NL-3508 AD Utrecht, the Netherlands.
M. Weiß. Eberhard-Karls-Universita
¨t Tu
¨bingen, Botanisches Institut, Lehrstuhl fu
¨r Spezielle Botanik und Mykologie, Auf der
Morgenstelle 1, D-72076 Tu
¨bingen, Germany.
1Corresponding author (e-mail: grit_walther44@yahoo.de).
551
Botany 86: 551–566 (2008) doi:10.1139/B08-036 #2008 NRC Canada
published for single species, such as Pholiota aurivella
(Batsch) P. Kumm. (Martens and Vandendries 1933), Pho-
liota gummosa (Lasch) Singer (Ku
¨hner 1946a, 1946b,
1946c, as Flammula gummosa,Hypholoma fasciculare (Fr.)
P. Kumm. (Garibova et al. 1986, as Naematoloma fascicu-
lare), and Psilocybe merdaria (Fr.) Ricken (Valenzuela and
Garnica 2000). In addition, various studies addressing a
wider spectrum of homobasidiomycetous anamorphs in-
cluded members of these families (e.g., Yen 1950; Oddoux
1955; Ka
¨a
¨rik 1970; and Res
ˇetnikov 1991). Except for Res
ˇet-
nikov (1991), who intensively examined the ontogeny of nu-
merous anamorphs in Agaricales, these studies lack details
concerning conidiogenesis and only the contribution of Yen
(1950) includes some general comparative remarks concern-
ing conidiogenesis in light-spored and dark-spored agarics.
Detailed comparative studies on ontogeny and morphol-
ogy of anamorphs have been published for only two genera
of the Strophariaceae, Pholiota (Hu
¨bsch 1978; Jacobsson
1989) and Psilocybe (Heim and Wasson 1958). The objec-
tive of the present study was a comparative analysis of ana-
morphs of the Strophariaceae as observed in pure culture.
To assess the systematic value of the observed characters,
we also made a comprehensive compilation and evaluation
of published reports on anamorphs.
This study is part of a project focusing on anamorphs of a
broader range of Agaricales. A survey of the detected types
of conidiogenesis, together with a molecular phylogenetic
analysis of the studied species and a general discussion of
the systematic relevance of anamorphs in Agaricales, has
been presented elsewhere (Walther et al. 2005).
Material and methods
Dikaryotic mycelia were isolated from basidiospore
deposits of freshly collected basidiocarps. To obtain single-
spore cultures, basidiospores were suspended in sterile dis-
tilled water using an inoculation loop. Several dilutions of
this suspension were plated on malt extract agar (2.5%, malt
extract produced by Difco, Augsburg, Germany) in Petri
dishes. The outgrowing mycelia were isolated and cultivated
on malt extract agar (2.5%) in Petri dishes at room temper-
ature.
At least eight single-spore mycelia and one dikaryotic
mycelium isolated from the same basidiocarp were studied
macroscopically and microscopically when the cultures had
reached about 3–4 cm in diameter. Dikaryotic mycelia were
identified by the presence of clamp connections. In cases
where the first examination did not reveal conidia, three of
these nine strains were examined a second time after 2–
3 months. Mounts for microscopic examinations were made
in two different ways for each species, (i) by pressing a
small piece of adhesive tape onto the colony, and (ii) by
squashing a small portion of the colony. The fungal material
was mounted in lactic acid with cotton blue (2 mg cotton
blue!ml–1 lactic acid) and examined using a microscope of
type JENAVERT (Zeiss, Jena, Germany). Drawings were
prepared at 100"magnification with the aid of a camera lu-
cida. The terminology concerning mycelia, conidia, and con-
idiogenous structures and processes is explained in Walther
et al. (2005).
Herbarium vouchers of the specimens from which the
strains were isolated are preserved either at the Staatliches
Museum fu
¨r Naturkunde Go
¨rlitz (GLM; Go
¨rlitz, Germany)
or at the Centraalbureau voor Schimmelcultures (CBS;
Utrecht, the Netherlands). Isolates are either kept at Bayer
Health Care (BAYER; Wuppertal, Germany), the Deutsche
Sammlung von Mikroorganismen und Zellkulturen (DSMZ;
Braunschweig, Germany), the Hans-Kno
¨ll-Institut (HKI) in
Jena (Germany), or the Centraalbureau voor Schimmelcul-
tures (CBS; Utrecht, the Netherlands). Herbarium and strain
numbers are given below where the respective specimens
are discussed. Strains lack an accession number in cases
where the respective strain died before deposition in a cul-
ture collection. The majority of the strains described in the
present paper have been included in the molecular phyloge-
netic analysis in Walther et al. (2005).
To test the germination potential of the conidia, a conidial
suspension was obtained by rinsing a richly sporulating
mycelium with sterile distilled water. The suspension was
mixed 1:1 (v/v) with a malt extract solution (2.5%) and stored
at room temperature. Every day, a drop from this suspension
was examined microscopically for germinating conidia.
To stain the nuclei of the conidia, conidiogenous parts of
the mycelium were spread out in a water drop as loosely as
possible on a slide previously coated with a mixture of glyc-
erol and egg white 1:1 (v/v). After drying at room tempera-
ture, the material was fixed in 3:1 absolute ethanol – pure
acetic acid (v/v) for 30 min. The slides were then briefly
rinsed in 70% ethanol and distilled water. Preparations were
hydrolyzed for 7.5 min in 1N HCl at 60 8C, and rinsed in
distilled water and subsequently in phosphate buffer
(0.1 mol!L–1 KH2PO4–KOH, pH = 7.0). The washed mate-
rial was stained in 1:10 Giemsa stock solution (Merck,
Darmstadt, Germany) – phosphate buffer (v/v) for 24 h. Sur-
plus stain was removed by washing in distilled water, fol-
lowed by rinsing with 70% ethanol and subsequently with
distilled water for a few seconds, respectively. Dried prepa-
rations were mounted in Euparal (Roth, Karlsruhe, Ger-
many) and examined microscopically.
Results
Thalloconidia
Thallic conidiogenesis was the dominating anamorph type
in the species of the Strophariaceae studied in culture. Two
characteristic modes of thallic conidiogenesis were revealed.
In mode I (Figs. 1–6, 12, 13, and 14–18), conidia were
formed by fragmentation of straight to strongly coiled coni-
diogenous hyphae arising singly as lateral or terminal
branches (undifferentiated conidiophores) or slightly sympo-
dially (conidiophores slightly differentiated in a sterile stipe
and fertile conidiogenous apical part). In mode II, conidia
developed singly or in straight chains on sympodially prolif-
erating geniculate conidiophores (Figs. 19–21). While the
conidia in mode I remained rod-shaped and hyaline, those
in mode II became swollen and coloured in older cultures.
The majority of the 21 species studied belonged to mode I,
while mode II was observed in Ph. aurivella,Ph. gummosa,
and Pholiota squarrosa (Batsch) P. Kumm. Pholiota alni-
cola (Fr.) Singer and Pholiota tuberculosa (Schaeff.)
P. Kumm. possessed specific modes deviating from the pre-
vailing modes I and II.
552 Botany Vol. 86, 2008
#2008 NRC Canada
In mode I as well as in mode II, conidial delimitation
started apically and continued either basipetally or ran-
domly. Most frequently this process started by an initial de-
limitation of larger hyphal segments by septa. Subsequently
these larger segments disarticulated into two or more coni-
dia. The conidia seceded rhexolytically. Compared with
other agaricalean families, the protoplasmic contraction,
which is part of the rhexolytic secession, was distinct in
most cases.
The conidia formed in culture were either dry or mucoid.
Mucoid conidia building slimy drops after release mostly
developed from coiled conidiogenous hyphae (mode I) or
from conidiogenous hyphae of geniculate conidiophores
(mode II). Conidia were produced on haploid, as well as on
dikaryotic mycelia.
In the following we give details of conidiogenesis as
studied in cultures. The species are grouped according to
their mode of conidiogenesis. In Agaricales, differentiated
conidiophores are characterized by a sterile stipe and an
apical part consisting of densely packed (condensed) sympo-
dially branching conidiogenous hyphae (Walther et al.
2005). The number of conidiogenous hyphae condensed by
Figs. 1–6. Conidiogenesis of mode I in the Strophariaceae. Fig. 1. Hypholoma capnoides (from Walther et al. 2005, reproduced with the
permission of Mycol. Res., Vol. 109, pp. 525–544. #2005 The British Mycological Society.) Fig. 2. H. marginatum. Fig. 3.
H. subericaeum. Fig. 4. Pholiota lenta. Fig. 5. Ph. mixta. Fig. 6. Ph. spumosa. Scale bar = 10 mm.
Walther and Weiß 553
#2008 NRC Canada
sympodial branching may serve as a measure for the differ-
entiation of the conidiophores. ‘‘Clearly mucoid’’ conidia
adhere in compact masses after release. ‘‘Slightly mucoid’’
conidia form only loose accumulations.
Mode I: conidiogenous hyphae straight or coiled,
geniculate conidiophores absent, conidia hyaline, rod-
shaped to slightly rounded in older cultures
The conidiogenous hyphae were mostly long and often
coiled. They arose singly as lateral or terminal hyphae, or
sympodially on slightly differentiated conidiophores. Geni-
culate conidiophoral structures were absent. Conidiogenous
hyphae of differentiated conidiophores were occasionally
branched. In cases where sporulation was particularly in-
tense, conidiophoral stipes eventually disintegrated. Sporula-
tion of dikaryotic mycelia was often less intense. If clamps
were involved in the fragmentation, they usually were trans-
formed into small, beak-like appendages of the conidia.
Conidia were one-celled, hyaline, thin-walled, dry or mu-
coid, and cylindrical, with the exceptions of Pholiota mixta
(Fr.) Kuyper & Tjall.-Beuk. (Fig. 5) and Stropharia rugo-
soannulata Farl. ex Murrill (Fig. 12), where slightly rounded
conidia occurred. Their size was not species-specific and
was in the range of 2–10 mm"1–2.5 mm.
Two different types of mycelial cystidia were found in
this group. We most frequently observed diverticulate cysti-
dia (hereinafter, cyanophilous mycelial cystidia; Figs. 7–8
and 10), which were incrusted by a strongly cyanophilous
substance in the central part. In Stropharia, additionally spi-
nulose mycelial cystidia occurred that we designate as acan-
thocytes, according to Farr (1980) (Figs. 9 and 11).
Mode Ia: conidiogenous hyphae straight or sporadically
coiled
The species showing mode I conidiogenesis differed con-
cerning constancy and extent of coiling. Numerous species
belonging to Hypholoma,Stropharia,Pholiota, and Kueh-
neromyces formed conidiogenous hyphae that were only
sporadically coiled. The presence or the extent of coiling de-
pended on the age of the culture and the intensity of sporu-
lation. We could not derive clear rules, but intensive
sporulation was frequently coupled with distinct coiling,
whereas in sparsely sporulating cultures, coiling was absent
in most cases. Sometimes it was necessary to observe the
cultures over 2–3 months to observe coiling. In some cul-
tures, conidiogenous hyphae condensed by sympodial
branching were rare, and fragmentation of normally
branched hyphal systems or of simple or little branched
short lateral hyphae predominated.
Hypholoma capnoides (Fr.) P. Kumm.
MATERIAL STUDIED: GLM 45937.
Figs. 7–13. Mycelial cystidia, conidiogenesis of mode I in the Strophariaceae, and conidiogenesis of Pholiota alnicola. Figs. 7–8. Cyano-
philous mycelial cystidia of Stropharia coronilla. Fig. 9. Acanthocytes of S. coronilla. Fig. 10. Cyanophilous mycelial cystidia of
S. rugosoannulata. Fig. 11. Acanthocytes of S. rugosoannulata. Fig. 12. Conidiogenesis in S. rugosoannulata. Fig. 13. Conidiogenesis in
S. semiglobata. Scale bar = 10 mm.
554 Botany Vol. 86, 2008
#2008 NRC Canada
DESCRIPTION: Fig. 1. Macroscopic, mat white to cream, cott-
ony. Microscopic, up to 6 conidiogenous hyphae (ch) con-
densing, nearly straight in some mycelia, but distinctly
coiled in most mycelia; conidia 3.5–10.3 mm"0.9–3.0 mm,
dry or slightly mucoid.
Hypholoma fasciculare (Fr.) P. Kumm.
MATERIAL STUDIED: GLM 45938, HKI ST 27356.
DESCRIPTION: Macroscopic, mat white to cream, cottony to
floccose. Microscopic, ch not or slightly (up to 3) condens-
ing, straight to strongly coiled depending on colony age;
conidia straight or curved, 2.5–9.5 mm"1.0–2.6 mm,
mucoid, with or without minute intracellular droplets.
Hypholoma marginatum (Pers.) J. Schro
¨t.
MATERIAL STUDIED: GLM 45939; GLM 46073.
DESCRIPTION: (Fig. 2). Macroscopic, mat is whitish to ochre,
plain, cottony. Microscopic, haploid mycelia sparsely sporu-
lating; up to 4 ch condensing, ch coiled; conidia straight or
Figs. 14–18. Conidiogenesis of mode I in the Strophariaceae. Fig. 14. Pholiota lucifera. Fig. 15. Psilocybe cf. coprophila. Fig. 16. Ps. in-
quilina. Fig. 17. Ps. semilanceata. Fig. 18. Psilocybe sp. Scale bar = 10 mm.
Walther and Weiß 555
#2008 NRC Canada
curved, 2.3–6.7 mm"1.6–1.8 mm (conidia measured in the
chain, because no free conidia were present at the time of
measurement).
Hypholoma subericaeum (Fr.) Ku
¨hner
MATERIAL STUDIED: GLM 45940, BAYER G524.
DESCRIPTION: (Fig. 3). Macroscopic, mat is whitish, plain, cott-
ony. Microscopic, only haploid mycelia sparsely sporulating;
ch not condensing by sympodial branching, straight or only
slightly curved; conidia 3.5–11.5 mm"1.1–1.7 mm (conidia
partly measured in the chain because only few free conidia
were present at the time of measurement), dry, with one or
two small intracellular droplets or without intracellular drop-
lets.
Hypholoma sublateritium (Schaeff.) Que
´l.
MATERIAL STUDIED: GLM 45941, BAYER G269.
DESCRIPTION: Macroscopic, mat white, cottony to floccose.
Microscopic, ch not or slightly (up to 3) condensing,
straight; conidia 2.0–7.5 mm"1.0–2.5 mm, dry; cyanophi-
lous mycelial cystidia present.
Figs. 19–21. Conidiogenesis of mode II in the Strophariaceae. Fig. 19. Pholiota aurivella. Fig. 20. Ph. squarrosa. Fig. 21. Ph. gummosa.
Scale bar = 10 mm.
556 Botany Vol. 86, 2008
#2008 NRC Canada
Kuehneromyces mutabilis (Schaeff.) Singer & A.H. Sm.
MATERIAL STUDIED: GLM 45999, DSMZ G288; GLM 46077,
HKI ST 21599.
DESCRIPTION: Macroscopic, mat white, dikaryotic mycelia or-
ange-brownish in age, cottony. Microscopic, ch not or
slightly (up to 3) condensing, straight, curved or slightly
coiled depending on colony age and strain, arising mainly
as short lateral branches; the location of clamps within ch
of dikaryotic mycelia depending on the differentiation of
the conidiphore (cph); ch of undifferentiated cph regularly
clamped; ch of differentiated cph only basally clamped (as
shown in Fig. 4 for Pholiota lenta (Pers.) Singer); conidia
straight or curved, 2.0–9.5(–14) mm"0.8–1.8(–4) mm, dry
or mucoid, with minute to small intracellular droplets or
without intracellular droplets, apically formed conidia some-
times slightly capitate (capitate apices were also observed at
somatic hyphae).
Pholiota lenta (Pers.) Singer
MATERIAL STUDIED: GLM 45997, HKI ST 27314.
DESCRIPTION: (Fig. 4). Macroscopic, young mat white, subse-
quently orange-brown, plain, cottony. Microscopic, up to
4 ch condensing, ch straight or coiled; cph of dikaryotic my-
celia often arising from clamps, within the cph clamps only
present at the base of the ch; conidia straight or curved, 2.8–
9.0 mm"1.1–2.5 mm, mucoid, containing one or more
small intracellular droplets.
Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk.
MATERIAL STUDIED: GLM 45998, BAYER G509.
DESCRIPTION: (Fig. 5). Macroscopic, mat white to cream-
coloured, plain, cottony to floccose. Microscopic, only hap-
loid mycelia sporulating; up to 5 ch condensing, ch
straight, arising mainly as short lateral hyphae; conidia
3.6–8.5 mm"1.0–1.6 mm, slightly rounded, dry, contain-
ing one or several more or less large intracellular droplets
(also ch containing intracellular droplets).
Pholiota spumosa (Fr.) Singer
MATERIAL STUDIED: GLM 46000, DSMZ G508; GLM 46076,
BAYER G351.
DESCRIPTION: (Fig. 6). Macroscopic,matwhitetocream,plain,
cottony to floccose. Microscopic, ch rarely condensing by
sympodial branching (condensation of 4 ch observed in one
case), but arising mainly as short lateral branches (monopo-
dial type of hyphal branching); these ch often condensing,
ch straight, shortened, especially in dikaryotic mycelia; ch
of dikaryotic mycelia mostly forming only 2–3 conidia; ch
of haploid mycelia forming 4–6(–10) conidia; conidia 2.5–
8.5(–11) mm"1.0–2.2 mm, slightly rounded, not or slightly
mucoid, with one to several minute intracellular droplets or
without intracellular droplets; numerous intercalary, terminal
or lateral thick-walled swellings present, 4.5–8.5 mm in di-
ameter.
Stropharia coronilla (Bull.) Que
´l.
MATERIAL STUDIED: GLM 46028, DSMZ G606; GLM 46074,
BAYER G294.
DESCRIPTION: (Figs. 7–9). Macroscopic, mat white, cottony.
Microscopic, no anamorph found, but numerous cyanophi-
lous mycelial cystidia and acanthocytes present.
Stropharia rugosoannulata Farl. ex Murrill
MATERIAL STUDIED: GLM 46075.
DESCRIPTION: (Figs. 10–12). Macroscopic, mat white, cottony.
Microscopic, only haploid mycelia sporulating; ch not con-
densing by sympodial branching, straight; conidia 3.4–
7.1 mm"1.0–2.0 mm, slightly rounded, mucoid with one or
two small intracellular droplets, or without intracellular
droplets; acanthocytes present in haploid mycelia; cyanophi-
lous mycelial cystidia present in both haploid and dikaryotic
mycelia.
Fig. 22. Conidiogenesis in Pholiota tuberculosa. Scale bar =
10 mm.
Walther and Weiß 557
#2008 NRC Canada
Stropharia semiglobata (Batsch) Que
´l.
MATERIAL STUDIED: GLM 46030, BAYER G601.
DESCRIPTION: (Fig. 13). Macroscopic,matwhitetopalebrown,
cottony. Microscopic, only haploid mycelia sporulating; up
to 3 ch condensing, ch straight or strongly coiled depending
on colony age; conidia straight to curved, 2.8–6.0 mm"
1.1–1.9 mm, mucoid, with one to several, but mostly two
small intracellular droplets, or without intracellular droplets;
cyanophilous mycelial cystidia present.
Mode Ib: conidiogenous hyphae constantly coiled
Members of Psilocybe and also Pholiota lucifera (Lasch)
Que
´l. constantly formed coiled conidiogenous hyphae. Com-
Fig. 23 .Conidiogenesis in Pholiota alnicola. Scale bar = 10 mm.
Table 1. Number of nuclei in the conidia formed by haploid (H) and dikaryotic (D) mycelia observed
in the present study.
Teleomorph
Nuclear number in the conidia formed
by haploid (H) or dikaryotic (D) mycelia
HD
Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk. 1 (2; 3)
Ph. spumosa (Fr.) Singer 1
Psilocybe inquilina (Fr.) Bres. 1
Psilocybe sp. 1 (2)
Stropharia semiglobata (Batsch) Que
´l. 1
Hypholoma capnoides (Fr.) P. Kumm. 1 1 (2; 3)
H. fasciculare (Huds.) P. Kumm. 1 (2) 1?*
H. sublateritium (Schaeff.) Que
´l. 1 1?*
Kuehneromyces mutabilis (Schaeff.) Singer & A.H. Sm. 1 (2) 1 (2)
Pholiota lenta (Pers.) Singer 1 (2) 1 (2)
Ph. lucifera (Lasch) Que
´l. 1 (2; 3) ?
Ph. aurivella (Batsch) P. Kumm. 1–6 2–4 (1)
Ph. gummosa (Lasch) Singer 1–8 2 (1; 3)
Ph. squarrosa (Batsch) P. Kumm. 1–4 2–6 (1)
Ph. alnicola (Fr.) Singer 1–4 2
Ph. tuberculosa (Schaeff.) P. Kumm. 1 2 (1; 3)
Note: *, Conidia are supposed to be uninuclear because conidia-forming dikaryotic mycelia were transformed
into haploid mycelia (see text); rarely occurring numbers are given in parentheses.
558 Botany Vol. 86, 2008
#2008 NRC Canada
pared with the species described above, the coiling of the
conidiogenous hyphae was more intense. Coiling in these
species was evident in every studied strain, at any mycelial
age. All species of this group produced mucoid conidia. The
resulting mucoid masses of conidia reached diameters of up
to 100 mm. In some cases, the conidiogenous hyphae were
coiled to such an extent that their exact number could only
be inferred for young conidiophores.
Pholiota lucifera (Lasch) Que
´l.
MATERIAL STUDIED: GLM 46002, DSMZ G829; GLM 46080,
HKI ST 20126.
DESCRIPTION: (Fig. 14). Macroscopic, mat white, orange-brown
with age (especially dikaryotic mycelia), cottony. Micro-
scopic, haploid and dikaryotic mycelia sporulating; up to
5 ch condensing; cph of the dikaryotic mycelium often aris-
ing from clamps; conidia in old mycelia sometimes swelling
(hyphae in old mycelia also swelling), but normally straight
to strongly curved, 2.3–7.0(–10.5) mm"1.0–2.2(–3.5) mm,
with small intracellular droplets, or without intracellular
droplets.
Psilocybe cf. coprophila (Bull.) P. Kumm.
MATERIAL STUDIED: CBS-H 19953; CBS 121966.
DESCRIPTION: (Fig. 15). Macroscopic, mat white to pale yellow,
plain, cottony to slightly floccose, slightly zonate. Micro-
scopic, haploid and dikaryotic mycelia sporulating; dikary-
otic mycelium forming clamps; cph of the dikaryotic
mycelium arising clampless hyphae; up to 4 ch condensing
(number inferred for young conidiophores); conidiophoral
stipe often slightly widened; conidia straight to curved, 2.0–
5.0 mm"1.5–2.2 mm, with one to several minute to small
intracellular droplets.
Psilocybe inquilina (Fr.) Bres.
MATERIAL STUDIED: GLM 46021, HKI ST 27345; GLM 46078,
BAYER G255.
DESCRIPTION: (Fig. 16). Macroscopic, aerial mycelium whitish,
sparse, floccose to cottony, plain; substrate mycelium orange-
ochre. Microscopic, only haploid mycelia sporulating; up to
4 ch condensing (number inferred for young conidiophores);
conidia straight to curved, (1.8-)3.0–8.0(–11.3) mm"1.2–
2.6 mm, containing one or several small to medium-sized
intracellular droplets.
Psilocybe semilanceata (Fr.) P. Kumm.
MATERIAL STUDIED: GLM 46022, BAYER G844.
DESCRIPTION: (Fig. 17). Macroscopic, mat white to pale orange-
ochre, cottony to felted. Microscopic, only haploid mycelia
investigated, haploid mycelia displacing dikaryotic mycelia
in polysporic cultures; up to 4 ch condensing (number in-
ferred for young conidiophores); conidia straight to curved,
2.0–8.0 mm"1.1–2.0 mm, with one to several, minute to
medium-sized intracellular droplets, or without intracellular
droplets.
Psilocybe sp.
MATERIAL STUDIED: GLM 46023, BAYER G523.
DESCRIPTION: (Fig. 18). Macroscopic, mat white, plain, cottony.
Microscopic, only haploid mycelia sporulating; up to 4 ch
condensing, ch slightly wider than the vegetative hyphae;
conidia straight to curved, 2.6–6.0 mm"1.1–1.7 mm, with
one to several small intracellular droplets, or without intra-
cellular droplets; thicker-walled swellings with intracellular
droplets present.
Mode II: conidiogenous hyphae straight, geniculate
conidiophores present, conidia coloured and swollen in
older cultures
Pholiota aurivella, Ph. gummosa, and Ph. squarrosa
formed straight conidiogenous hyphae and sympodially pro-
liferating geniculate conidiophores (Figs. 19–21). Fre-
quently, the directions of the sympodial outgrowths
alternated. In some cases, the conidiogenous hyphae were
reduced to a single cell. In addition to the production of
conidia on differentiated conidiophores, conidia were
formed by simple lateral branches that could also be reduced
to a single cell.
Conidia of older mycelia swelled. When they developed
singly or in chains of two, they mostly showed a more dis-
tinct swelling. Generally, the simpler the conidiophore archi-
tecture, the larger and more thick-walled were the conidia.
While in conidiogenesis of mode I the conidia rarely ex-
ceeded 2.5 mm in width, widths of 3–4 mm were often found
in mode II. Occasionally, swelling started before the septum
appeared, so that the conidiogenesis might be considered as
blastic in these cases. The conidia of young mycelia were
hyaline, those of older mycelia became yellowish to yellow-
brown. They were smooth-walled and contained one to sev-
eral medium-sized to large intracellular droplets.
Both haploid and dikaryotic mycelia formed thalloconidia.
Haploid mycelia showed a higher variation in the number of
conidia in a chain and in the differentiation of the conidio-
phores.
Pholiota aurivella (Batsch) P. Kumm.
MATERIAL STUDIED: GLM 45995, HKI ST 27321.
DESCRIPTION: (Fig. 19). Macroscopic, young mat white, golden
yellow in age, cottony. Microscopic: up to 10 ch per cluster;
ch of dikaryotic mycelia forming (1–)2(–4) conidia, ch of
haploid mycelia forming 1–6 conidia; conidia cylindrical to
nearly globose, 4.0–11.5 mm"1.5–5.5 mm, thin- or thick-
walled, mucoid, hyaline or yellowish, up to yellow brown
in older cultures.
Pholiota gummosa (Lasch) Singer
MATERIAL STUDIED: GLM 45996, HKI ST 27313.
DECRIPTION: (Fig. 21). Macroscopic, mat white to orange-
brown, cottony. Microscopic, considerable differences be-
tween haploid and dikaryotic mycelia. Haploid mycelia:
conidiogenesis by fragmentation of simple lateral branches
or of ch sympodially proliferating from the apex of cph
stipes; transitions between both modes present; up to 11 ch
per cluster; ch forming 1–8 conidia; conidia 1.9–10.0 mm"
1.5–4.0 mm. Dikaryotic mycelia: conidiogenesis by fragmen-
tation of ch sympodially proliferating from the apex of cph
stipes; up to 5 ch per cluster; ch with a basal clamp, forming
(1–)2(–3) conidia; conidia 4.8–10.8 mm"2.0–4.0 mm; con-
idia of both haploid and dikaryotic mycelia truncate at one
or both ends, mucoid, hyaline or yellow in older cultures;
striking refractive swellings (‘‘allocysts’’ of Ku
¨hner 1946a)
present, mainly terminal, but occasionally formed laterally,
Walther and Weiß 559
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singly or in chains of two or three, hyaline, ellipsoid or pyri-
form, thick-walled, smooth-walled, 3.5–31.5 mm"3.5–
13.0 mm.
Pholiota squarrosa (Batsch) P. Kumm.
Material studied: GLM 46001, HKI ST 22286; GLM
46082, BAYER R170.
DESCRIPTION: (Fig. 20). Macroscopic, young mat white, later
orange-brown, cottony. Microscopic, sporulation sparse;
sympodially proliferating cph rare; up to 6 ch per cluster,
but conidia mainly formed by short lateral hyphae; ch form-
ing 1–2(–3) conidia; conidia truncate, 6.0–11.5 mm"2.8–
7.1 mm, thick-walled, hyaline or yellowish. The formation
of strongly swollen conidia was often associated with the
absence of a distinct protoplasmic contraction.
Thalloconidia of Pholiota tuberculosa (Schaeff.)
P. Kumm.
Pholiota tuberculosa also produced permanently coiled
conidiogenous hyphae, however, its conidiogenesis differed
from mode Ibmainly by the formation of swollen and
strongly curved conidia and of sporodochia, and by acrope-
tal conidial maturation (Fig. 22).
MATERIAL STUDIED: GLM 46026, HKI ST 27336.
DESCRIPTION: Macroscopic, aerial mycelium sparse, whitish,
with small sporodochia; substrate mycelium pale orange-
ochre, tuberculate. Microscopic, haploid and dikaryotic my-
celia sporulating; ch disarticulating into chains of 2 or 3
conidia, strongly coiled, branched; up to 3 ch condensing
by sympodial branching, up to 5 if monopodial branching
was also present; conidia swelling during maturation; coni-
dial maturation acropetal (it is unclear whether apical
growth of ch stops before maturation starts basally); ch of
dikaryotic mycelia regularly clamped; clamps mostly open-
ing and becoming beak-like appendages of the conidia,
rarely transformed into small conidia (3.0–5.5 mm"1.5–
2.0 mm); strongly curved conidia occasionally disarticulating
into two truncate conidia; cph single or concentrated in
sporodochia; sporodochia completely disarticulating into
conidia; vegetative hyphae and ch of ageing cultures swelling
irregularly and thus forming irregularly shaped conidia;
conidia hyaline, but pale orange-ochre in mass, swollen,
strongly curved or irregularly shaped, (4.0-)6.0–
11.5(–4.0) mm"2.0–3.5(–5.2) mm, with large intracellular
droplets, mucoid; conidial masses forming slimy drops.
Thalloconidia of Pholiota alnicola (Fr.) Singer
In contrast to other members of the family sensu Singer,
its conidiogenesis lacked a protoplasmic contraction. The
conidia were distinctly larger and their genesis was often un-
ambiguously blastic (Fig. 23).
MATERIAL STUDIED: GLM 45994, BAYER G515.
DESCRIPTION: Macroscopic, mat white to yellowish or beige,
cottony. Microscopic, haploid and dikaryotic mycelia sporu-
lating; conidiogenesis blastic to thallic with continuous tran-
sitions; conidia formed singly, in chains of two, or
sympodially, terminally on main hyphae or mostly on short
lateral supporting hyphae or laterally without supporting hy-
phae, mostly swelling; supporting hyphae in mycelia with
intense sporulation sometimes finally disarticulating into
thalloconidia that subsequently swell; secession of conidia
delayed after conidial maturation; conidia truncate, 6.0–
17.5 mm"5.2–13.5 mm, smooth-walled with walls up to
1mm thick, hyaline or pale yellow with large intracellular
droplets, dry.
Germination of the conidia
All germination tests were successful. Thus germination
of the conidia was observed for the following species:
Hypholoma capnoides,H. fasciculare,Kuehneromyces mu-
tabilis,Pholiota alnicola,Ph. aurivella,Ph. gummosa,Ph.
lenta,Ph. mixta,Ph. spumosa,Ph. squarrosa,Ph. tubercu-
losa,Psilocybe sp., and Stropharia semiglobata.
Numbers of nuclei in the conidia
Species with rod-shaped conidia (conidiogenesis mode I)
produced uninucleate conidia on haploid mycelia (Table 1).
Bi- or tri-nucleate conidia occurred rarely. Hypholoma cap-
noides,Kuehneromyces mutabilis, and Pholiota lenta pro-
duced uninucleate conidia on dikaryotic mycelia. In
Hypholoma fasciculare,H. sublateritium, and Pholiota lenta
the conidia-forming clamped dikaryotic mycelia transformed
into clampless conidia-forming mycelia after longer cultiva-
tion. Therefore, we assume that dikaryotic mycelia of
H. fasciculare and H. sublateritium also produced uninu-
cleate conidia, and that the germinating haploid mycelia re-
placed the dikaryotic culture.
In contrast to conidiogenesis mode I, species with mode II
produced conidia with one to several nuclei on haploid my-
celia. Conidia of dikaryotic mycelia predominantly con-
tained two nuclei and germinated by producing clamped
mycelia. The swollen conidia of Pholiota aurivella and
Ph. squarrosa contained up to four or six nuclei, respec-
tively. In rare cases, the conidia possessed only one nucleus,
probably as a result of irregular cell divisions. This assump-
tion is based on the observation that in the accumulations of
conidia found in the preparations a nearly equal number of
free uninucleate and trinucleate conidia occurred. In two
cases a uninucleate conidium was still connected in chain to
a trinucleate conidium.
Hyphal swellings
We often observed spore-like hyphal swellings in cultures
of the Strophariaceae. Because of their unspecific ontogeny
and morphology they probably possess no systematic rele-
vance. In many ageing or desiccating mycelia, cytoplasm
was retracted into swollen hyphal parts. Submerged mycelia
often contained irregularly swollen hyphae. Swellings devel-
oped singly or in chains, terminal or intercalary on undiffer-
entiated hyphae. Generally, the swellings varied
considerably in size, shape, and thickness of the cell walls,
and reached a distinctly larger size than true conidia. Their
cell walls were often refractive.
Discussion
The dominance of the thallic mode of conidiogenesis in
the Strophariaceae is supported by numerous literature re-
ports (Table 1 and Table 2) predominantly referring to cul-
ture studies. However, Garibova et al. (1986) detected
560 Botany Vol. 86, 2008
#2008 NRC Canada
thalloconidia in vivo on rhizomorphs of H. fasciculare.
Blastic conidiogenesis was described only by Garibova et
al. (1986), who observed budding cells and chains of blasto-
conidia in cultures and on the surface of rhizomorphs of
H. fasciculare.However,thesemighthavebeencausedby
a contaminant, since these types of cells were not mentioned
by any other author, nor did we observe such cells in our
culture studies.
Most descriptions in the literature relate to the modes I
(Table 2) and II (Table 3). In addition, a few reports refer
to the specific modes of Pholiota tuberculosa and Ph. alni-
cola. The anamorph description for Psilocybe semilanceata
given by Brefeld (1889) is probably based on a member of
the Psathyrellaceae, because the drawings clearly show a
Hormographiella stage, the characteristic anamorph of the
Psathyrellaceae (Walther et al. 2005). On the other hand,
this author illustrated the anamorph of Psathyrella nolitan-
gere (Fr.) Pears. & Dennis (as Psathyra n.) with long,
straight to curved conidiogenous hyphae. Hence, he might
have confused Psilocybe semilanceata with Psathyrella noli-
tangere. The same author delineated thalloconidia formed by
the fragmentation of usually branched hyphae for a species
he designated as ‘‘Pholiota squarrosa Mu
¨ller’’, which we
were unable to relate to current nomenclature. Since the de-
scriptions of Hu
¨bsch (1978) and Jacobsson (1989) for Pho-
liota squarrosa (Batsch) P. Kumm. are consistent with our
observations, we suppose that Brefeld’s ‘‘Ph. squarrosa
Mu
¨ller’’ is a different species.
For Hypholoma marginatum (Watling in Kendrick and
Watling 1979, p. 515), Kuehneromyces lignicola (Peck)
Redhead (Watling in Kendrick and Watling 1979, p. 515, as
K. myriadophyllus), Pholiota conissans (Fr.) M.M. Moser
(Yen 1950, as Flammula lutaria), Pholiota highlandensis
(Peck) A.H. Sm. & Hesler (Galland 1968, as Flammula car-
bonaria), Pholiota nameko (T. Ito
ˆ) S. Ito & S. Imai (Buhalo
1988), Psilocybe caerulescens Murrill (Singer 1958), and
Ps. semilanceata (Yen 1950) thalloconidia were reported
without descriptions of their genesis.
In the phylogenetic trees of Moncalvo et al. (2002) and
Walther et al. (2005), species showing conidiogenesis mode
Ia, except for Kuehneromyces, belong to one clade that in-
cludes the small clade of species with mode II. Species
showing mode Ibthat are included in the trees do not cluster
together. The conidiogenesis type of Pholiota lucifera sup-
ports its inclusion in the Strophariaceae; this species was
(although without significant statistical support) placed dis-
tantly from the Strophariaceae cluster in the phylogenetic
tree of Walther et al. (2005). However, better resolved mo-
lecular phylogenetic trees are needed to substantially test the
systematic relevance of the conidiogenesis types discussed
here.
The rhexolytic mode of secession is typical for the dark-
spored families of the gilled Agaricales (Walther et al.
2005). Contraction of the protoplasma, which is part of the
rhexolytic secession, has been described by other authors in
certain species such as H. fasciculare (Garibova et al. 1986,
in dikaryotic mycelia only), Pholiota spp. (Jacobsson 1989),
Ph. gummosa (Ku
¨hner 1946a, as Flammula g.), Ph. auri-
vella,Psilocybe coprophila, and Stropharia semiglobata
(Res
ˇetnikov 1991). Consistently, drawings of anamorphs
(Pholiota flammans (Batsch) P. Kumm. in Oddoux (1955);
Psilocybe mexicana R. Heim, Psilocybe hoogshagenii
R. Heim in Heim and Wasson (1958)) suggest distinct pro-
toplasmic contraction. Conflicting observations concerning
conidial secession have been reported from H. fasciculare.
Vandendries (1924) and Garibova et al. (1986) did not de-
tect protoplasmic contractions in haploid mycelia of this
species, and Vandendries (1924) supposed that the conidia
were released by dissolution of hyphal septa.
The accumulation of mucoid conidia in adhesive drops
was also reported by Petersen (1992c) for Melanotus eccen-
tricus (Murrill) Singer, which possesses coiled conidioge-
nous hyphae as well. Walther et al. (2005) interpreted this
fact as an adaptation to dispersal by soil invertebrates.
Mode I
In Table 2, the species showing mode I of conidiogenesis
are arranged according to the degree of coiling of their con-
idiogenous hyphae. Consistent with our findings, the major-
ity of literature reports mention coiled conidia-forming
hyphae. Probably due to the inconstancy of this character,
our observations and those of other authors appear to be
conflicting in some cases: in Hypholoma capnoides and
Pholiota lenta, we, but none of the authors listed in Table 2,
observed coiled conidiogenous hyphae, whereas in Hypho-
loma sublateritium only Vandendries (1934) and Brefeld
(1889) noticed this character. In Kuehneromyces mutabilis,
we found slightly coiled hyphae. However, Falck (1902, as
Pholiota m.) reported conidiogenous hyphae arising in fas-
cicles, which might indicate strongly coiled conidiogenous
hyphae.
The anamorph genus Pseudohelicomyces Garnica &
E. Valenzuela, originally described for the anamorph of Psi-
locybe merdaria (Valenzuela and Garnica 2000), fits all spe-
cies related to mode 1b.
Mode II
Literature reports referring to mode II of conidiogenesis
are listed in Table 3. Consistent with our observations,
some measurements given by other authors (Table 3) also
suggest swollen conidia. Martens and Vandendries (1933)
treated them as ‘‘chlamydospores’’.
The morphological plasticity in conidial separation makes
it difficult to apply the terms ‘‘thallic’’ or ‘‘blastic’’ to the
mode of conidiogenesis in this group. This indicates, as al-
ready stated for the Onygenales by Sigler (1989), that a ba-
sic distinction between the thallic and the blastic mode,
which is widely being used in taxonomic descriptions and
keys, does not exist in some groups of fungi.
Consistent with our observations, Ku
¨hner (1946a) re-
ported fewer conidia per chain in dikaryotic than in haploid
mycelia of Pholiota gummosa. The drawings of Ph. auri-
vella by Martens and Vandendries (1933) also show a higher
variability of anamorphs formed by haploid mycelia. Ac-
cording to the extent of conidial swelling, Martens and Van-
dendries (1933) distinguished between different spore types
in Ph. aurivella: cylindrical, thin-walled ‘‘oı
¨dies’’ formed in
long chains; slightly swollen, fusiform ‘‘conidies’’ produced
in chains of two; and strongly swollen, thick-walled ‘‘chla-
mydospores’’ that developed singly. These authors reported
that ‘‘oı
¨dies’’ and ‘‘chlamydospores’’ germinated to produce
dikaryotic hyphae, while the ‘‘conidies’’ grew into haploid
Walther and Weiß 561
#2008 NRC Canada
Table 2. Reports on conidiogenesis mode I in species of the Strophariaceae, arranged according to the degree of coiling of their conidiogenous hyphae.
Teleomorph Conidial measurements (mm)
Conidia formed on haploid (H)
or dikaryotic (D) mycelia References
Conidiogenous hyphae straight
Hypholoma brunneum (Massee) D.A. Reid NA H / D Petersen 1992b
H. capnoides (Fr.) P. Kumm. [Naematoloma c.] up to 25"1.5–3.52H1/D
2Yen 19501; Zycha and Knopf 19662; Ka
¨a
¨rik 1970;
Siepmann 1969
Kuehneromyces mutabilis (Schaeff.) Singer & A.H. Sm.
[Pholiota m.]
2.5–6"22; 4.8–8.0 long3;
5–10"24
H1/D
4Brefeld 1889; Falck 1902; Yen 19501; Oddoux 19552;
Ka
¨a
¨rik 1970; Res
ˇetnikova et al. 19883;Jacobsson 19894
Pholiota lenta (Pers.) Singer [Flammula l. ] 3–8"22; D: 5–10"23H1/D
3Lamoure 19541; Oddoux 19552; Ka
¨a
¨rik 1970; Jacobsson 19893
Ph. lubrica (Pers.) Singer 5–10"2 D Jacobsson 1989
Ph. spumosa (Fr.) Singer [Flammula s.] 5–10"22H1/D
2Yen 19501;Jacobsson 19892
Psilocybe hoogshagenii R. Heim 6–12 long NA Heim and Wasson 1958
Ps. mexicana R. Heim NA D? Heim and Wasson 1958
Conidiogenous hyphae curved
Pholiota flammans (Batsch) P. Kumm. 2–7,5"2,21D1Oddoux 19551; Ka
¨a
¨rik 1970
Ph. squarrosoides (Peck) Sacc. 3–8"1–3 D Jacobsson 1989
Conidiogenous hyphae strongly coiled
Hypholoma fasciculare (Fr.) P. Kumm. [Naematoloma f.] 5.4–10.0"2.7–4.63H1/D
2Brefeld 1889; Falck 1902; Vandendries 19241; Ka
¨a
¨rik 19702;
Garibova et al. 19863
H. perplexum (Peck) Sacc. NA H Nichols 1904
H. radicosum J.E. Lange [H. epixanthum sensu Ricken] 4"2 D Oddoux 1955
H. sublateritium (Schaeff.) Que
´l. [Naematoloma s.] NA H1/D
2Brefeld 1889; Vandendries 19341; Yen 1950; Ka
¨a
¨rik 19702
Melanotus eccentricus (Murrill) Singer NA H Petersen 1992c
M. hartii Ammirati 2.9–3.7"1.5–2.21H1/D
2Ammirati et al. 19791;Ammirati
2in Petersen 1992a, p. 339;
Petersen 1992a
M. hepatochrous (Berk.) Singer NA H / D Petersen 1992a
M. horizontalis (Bull.) P.D. Orton [M. textilis] NA H Walker et al. 1994
Pholiota jahnii Tjall.-Beuk. & Bas 3–6"1–3 D Jacobsson 1989
Ph. polychroa (Berk.) A.H. Sm. & H.J. Brodie NA H Smith and Brodie 1935
Psilocybe acutissima R. Heim NA D? Heim and Wasson 1958
Ps. coprophila (Bull.) Que
´l. 4–10"1.5 H / D Gilmore 1926
Ps. coprophila (Bull.) Que
´l. ?? [Agaricus
coprophilus Bull.]
NA H1Eidam 1875; Falck 19021
Ps. merdaria (Fr.) Ricken 2.8–8.4 (19.6)"2–2.81H1/D
1Watling 1971; Valenzuela and Garnica 20001
Ps. mixaeensis R. Heim NA NA Heim and Wasson 1958
Ps. montana (Pers.) P. Kumm. ??
[Deconica atrorufa]
NA H Lamoure 1958
Ps. strictipes Singer & A.H. Sm.
[Psilocybe callosa]
NA H Brefeld 1889
Ps. tenax (Fr.) Ku
¨hner & Romagn. non Ricken [Naucoria t.] NA H Galland 1968
Ps. yungensis Singer & A.H. Sm. NA D Heim and Wasson 1958
Ps. zapotecorum R. Heim NA NA Heim and Wasson 1958
Stropharia melasperma (Bull.) Gillet
[S. melanosperma]
NA H Brefeld 1889
S. semiglobata (Batsch) Que
´l. NA H1Brefeld 1889; Res
ˇetnikov 19911
Note: Deviating names used by at least one author are given in square brackets; the author giving the most comprehensive information is printed in bold; details are related to authors by superscript
numbers when more than one author is cited; ??, indicates that the assignment of the specific name used by the author to the current nomenclature is uncertain; D?, indicates that the nuclear status of
conidia-forming mycelium is uncertain because conidiogenous hyphae and clamped hyphae are illustrated, but not in connection; NA, not reported.
562 Botany Vol. 86, 2008
#2008 NRC Canada
mycelia. Thus three ontogenetically and morphologically
distinct types of conidia including blastoconidia have been
assigned to Ph. aurivella. However, the strain HKI ST
27321 examined for the present study developed only one,
but morphologically variable, type of conidia covering all
of these spore types and also transitional stages. We ob-
served predominantly dikaryotic conidia; monokaryotic con-
idia occasionally occurred probably as a result of irregular
nuclear divisions. However, they had no consistent shape as
stated by Martens and Vandendries (1933).
Pholiota tuberculosa
The combination of coiled conidiogenous hyphae and
swollen and strongly curved conidia is very characteristic of
the Ph. tuberculosa anamorph. These traits match the de-
scription given by Cle
´menc¸on (2000) for Simocybe sump-
tuosa (P.D. Orton) Singer. He additionally observed
strongly swollen, thick-walled ‘‘aleuria’’ or intercalary
‘‘chlamydospores’’ depending on their position in the chain,
which developed similarly and often on the same conidio-
phores as the conidia. It is unclear whether these spores and
the irregularly swollen conidia of isolate HKI ST 27336 are
homologous structures. Based on the similarity of the
anamorphs and the placement of Ph. tuberculosa in the
phylogenetic tree of Moncalvo et al. (2002), close to
Simocybe and Crepidotus, Walther et al. (2005) discussed
an exclusion of Ph. tuberculosa from the genus Pholiota.
Pholiota alnicola
According to size and shape of the described structures
several reports refer to conidia in Ph. alnicola (Ku
¨hner
1947, as Flammula a., Denyer 1960, Nobles 1965, Ka
¨a
¨rik
1970, Hu
¨bsch 1978, as Ph. flavida, Jacobsson 1989). Ku
¨hner
(1947), Denyer (1960), and Nobles (1965) designated them
as ‘‘allocysts’’, suggesting that they did not secede. However,
the conidia of strain BAYER G515 examined in the present
study detached and germinated easily. Therefore it remains
questionable whether the allocysts of Ku
¨hner (1947), Denyer
(1960) and Nobles (1965) correspond to the conidia de-
scribed in the present study. If these are identical structures,
the term ‘‘allocysts’’ would be inappropriate for them.
In the multilocus phylogenetic overview presented by
Matheny et al. (2006) the species is positioned outside the
Strophariaceae clade, which corresponds with its deviating
anamorph morphology.
Germination tests
In addition to the positive results of our own germination
tests, successful germination of conidia has been described
for H. fasciculare (Vandendries 1924),
Hypholoma perplexum (Peck) Sacc. (Nichols 1904),
Hypholoma radicosum J.E. Lange (Oddoux 1955, as
H. epixanthum), Kuehneromyces mutabilis (Falck 1902),
Melanotus eccentricus (Petersen 1992c), Pholiota aurivella
(Martens and Vandendries 1933), Ph. gummosa (Ku
¨hner
1946a), Pholiota polychroa (Berk.) A.H. Sm. & H.J. Brodie
(Smith and Brodie 1935), Psilocybe coprophila (Gilmore
1926), Ps. merdaria (Valenzuela and Garnica 2000), Psilo-
cybe strictipes Singer & A.H. Sm. (Brefeld 1889, as Ps. cal-
losa), and Stropharia semiglobata (Brefeld 1889). Negative
results of germination tests were only reported by Brefeld
(1889) for Psilocybe semilanceata and ‘‘Stropharia stercora-
Table 3. Reports on conidiogenesis mode II in Pholiota.
Teleomorph Conidial measurements in mm
Conidia formed on
haploid (H) or
dikaryotic
(D) mycelia References
Pholiota adiposa (Fr.) P. Kumm. 4–12"1.5–61; 5–10"3–82; 7.2–20* "
1.5–2.6 and 6.0–8.2"3.5–4.0 (as
chlamydospores)3
D3Batko 19461; Semerdz
ˇieva
19652; Ka
¨a
¨rik 1970;
Buhalo 1973; Hu
¨bsch 19783;
Buhalo et al. 1985; Buhalo et
al. 1989
Ph. aurivella (Batsch) P.Kumm.
[Pholiota surivella, misapplied
Pholiota adiposa]
to 9"2.52; 5–10"3–83; 8.0–13* "
1.7–2.24; 5–11"1.5–35
H1/D
4Martens and Vandendries 19331;
Oddoux 19552; Nobles 1965;
Semerdz
ˇieva 19653;
Ka
¨a
¨rik 1970; Buhalo 1973;
Hu
¨bsch 19784; Buhalo 1988;
Jacobsson 19895
Ph. gummosa (Lasch) Singer
[Flammula g.]
H: 3–15.7"1.7–3.7, D: 4.5–
13.5"2.7–4.51; 5.5–8"2.7–4.52;
10.6–16.0* "1.3–2.63; 4–10"2–44
H1/D
1Ku
¨hner 1946a1; Ku
¨hner 1946c2;
Ka
¨a
¨rik 1970; Hu
¨bsch 19783;
Jacobsson 19894
Ph. heteroclita (Fr.) Que
´l. NA D1Ka
¨a
¨rik 1970; Jacobsson 19891
Ph. limonella (Peck) Sacc. 5–11"1.5–3 D Jacobsson 1989
Ph. subsquarrosa (the name
cannot be related to current
nomenclature)
10.0–16.8* "1.5–2.0 D Hu
¨bsch 1978
Ph. squarrosa
(Batsch) P. Kumm.
6–24"5–91; 9.0–15.0* "1.7–2.22;
6–10"3–43
D2Batko 19461;Hu
¨bsch 19782;
Ka
¨a
¨rik 1970; Jacobsson 19893
Note: Deviating names used by at least one author are given in brackets, the author giving the most comprehensive information is printed in bold; details
are related to authors by superscript numbers when more than one author is cited. *, the author treated two conidia formed in a chain as one conidium, for a
comparison with other conidial lengths the given length must be halved.
Walther and Weiß 563
#2008 NRC Canada
ria’’, which is now treated as synonymous with
S. semiglobata. Hence, Brefeld may have succeeded in ger-
minating the conidia of one out of two strains of
S. semiglobata. However, as already discussed above, the
‘‘Psilocybe semilanceata’’ culture that he studied was
possibly obtained from a member of the Psathyrellaceae. In
summary, the conidia of the Strophariaceae germinate easily
and may function as diaspores.
Numbers of nuclei in the conidia
Uninucleate conidia formed by haploid mycelia of species
showing conidiogenesis mode I have been reported for H.
fasciculare (Vandendries 1924), H. perplexum (Nichols
1904), Melanotus eccentricus (Petersen 1992c), and Psilo-
cybe merdaria (Valenzuela and Garnica 2000). Uninucleate
conidia produced on dikaryotic mycelia were also observed
in species possessing mode I of conidiogenesis, such as Hy-
pholoma radicosum (Oddoux 1955, as H. epixanthum) and
Psilocybe merdaria (Valenzuela and Garnica 2000). Accord-
ing to Gilmore (1926), the conidia of Ps. coprophila that de-
veloped within dikaryotic mycelia germinated to produce
either haploid or dikaryotic hyphae. Uninucleate conidia on
dikaryotic mycelia are also known from Flammulina velu-
tipes (Curtis) Singer (Brodie 1936; Aschan 1952) and vari-
ous aphyllophoralean species (Maxwell 1954; David 1970).
The transformation of an already established dikaryotic my-
celium to the haploid state seems to occur frequently, but its
function remains unclear.
According to Martens and Vandendries (1933) dikaryotic
mycelia of Pholiota aurivella form a distinct fusiform and
uninuclear type of conidia. We did not observe this conidial
type. Only rarely we found uninucleate conidia on
dikaryotic mycelia in all species showing conidiogenesis
mode II; such uninucleate conidia may result from irregular
cell divisions. Irregularities in nuclear division were also
observed by Ku
¨hner (1946a)whofoundtrinucleateconidia
on dikaryotic mycelia of Ph. gummosa.
Hyphal swellings and acanthocytes
Ku
¨hner (1946b, p. 152) introduced the term ‘‘allocysts’’
for more or less differentiated, but nondetaching mycelial
cells that he interpreted as equivalent to hymenial cystidia.
Although he did not explicitly restrict this term to terminal
hyphal structures, the implicit equivalence with hymenial
cystidia makes this term difficult to apply, e.g., for irregular
intercalary swellings. Therefore, we prefer the designation
‘‘hyphal swelling’’ for unspecific structures.
To clearly distinguish taxonomically relevant conidia
from nondetaching spore-like structures, such as unspecific
hyphal swellings or mycelial cystidia, we compiled all re-
ports lacking clear information about the release of such
structures separately in Table 4. Some of the reported struc-
tures may actually refer to true conidia or mycelial cystidia,
e.g., the small echinulate ‘‘chlamydospores’’ of H. fascicu-
lare (Garibova et al. 1986, as Naematoloma f.), but this can-
not be decided on the basis of the available data.
Table 4. Hyphal swellings reported in the Strophariaceae.
Teleomorph Measurements in mm
Swellings formed on
haploid (H) or
dikaryotic (D)
mycelia Reference
Hypholoma capnoides (Fr.) P. Kumm.
[Naematoloma c.]
7–20 or 4–10"10–20 D Siepmann 1969
H. fasciculare (Fr.) P. Kumm.
[Naematoloma f.]
NA H Vandendries 1924, as ‘‘grosses oidies’’
H. fasciculare (Fr.) P. Kumm.
[Naematoloma f.]
3.9–4.0"4.0–4.2 NA Garibova et al. 1986
Kuehneromyces lignicola (Peck)
Redhead [Pholiota l.]
NA D Jacobsson 1989
Melanotus hartii Ammirati 10.0–13.0 D Ammirati et al. 1979
Pholiota aurivella (Batsch) P. Kumm.
[Ph. adiposa ]
8–30"5–10 D Jacobsson 1989
Ph. conissans (Fr.) M.M. Moser
[Flammula c.,F. lutaria,Ph. graminis]
12–27"7–8.51; 11.3–
20.0"6.3–8.82; 15–
20"10–154;
H2/D
3Ku
¨hner 1947, as allocysts1; Denyer
19602;Nobles 1965, as allocysts3;
Jacobsson 19894
Ph. flammans (Batsch) P. Kumm. NA D Ka
¨a
¨rik 1970
Ph. flavida (Schaeff.) Singer 15–18"8.5–10.0 D Hu
¨bsch 1978
Ph. gummosa (Lasch) Singer
[Flammula g.]
21.0–24.7"8.7–10.62;
15–30"12–203
H1/D
1Ku
¨hner 1946a, as allocysts1;
Ka
¨a
¨rik 1970; Hu
¨bsch 19782;
Jacobsson 19893
Ph. jahnii Tjall.-Beuk. & Bas 8–45"6–10 D Jacobsson 1989
Ph. limonella (Peck) Sacc. 8–30"5–10 D Jacobsson 1989
Ph. squarrosa (Batsch) P. Kumm. 12–14"5–61;
12.0"8.0–12.02; 10–
16"8–123
D2Ku
¨hner 1946b, as allocysts1;
Ka
¨a
¨rik 1970; Hu
¨bsch 19782;
Jacobsson 19893
Ph. squarrosoides (Peck) Sacc. 9–15"6–8 D Jacobsson 1989
Note: If no other term is explicitely mentioned, they were designated as ‘‘chlamydospores’’ in the cited references. Deviating species names used in the
cited references are given in brackets; the author giving the most comprehensive information is printed in bold; details are related to authors by superscript
numbers when more than one author is cited.
564 Botany Vol. 86, 2008
#2008 NRC Canada
The term ‘‘acanthocytes’’ was created by Farr (1980) for
mycelial cystidia (allocysts sensu Ku
¨hner 1946b) with a
specific shape that occurs in Stropharia spp. This author
found this kind of cystidia in cultures and (or) herbarium
specimens of Stropharia aeruginosa (Curtis) Que
´l.,
Stropharia ambigua (Peck) Zeller, S. coronilla,
Stropharia cyanea (Bolton ex Secr.) Tuomikoski,
Stropharia hardii G.F. Atk., Stropharia hornemannii (Fr.)
S. Lundell & Nannf., Stropharia kauffmanii A.H. Sm.,
S. rugosoannulata, and Stropharia subsquamulosa A.H. Sm.
& Mitchell.
This study suggests that anamorph morphology is able to
provide new systematically relevant characters. However,
data on conidiophore morphology are lacking for many taxa
and further studies are needed to assess the specificity of the
respective characters for particular groups.
Acknowledgments
We are grateful to Walter Gams for critically reading an
earlier draft of the manuscript, Be
´atrice Senn-Irlet and
Gerhard Zschieschang for reviewing the identification of a
specimen, and Bayer Health Care for financial support.
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