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MYCOTAXON
Volume 92, pp. 361–369 April–June 2005
Taxonomy and nomenclature of Gymnopilus bellulus and
G. microsporus (Agaricales, Cortinariaceae)
JAN HOLEC
jan.holec@nm.cz
National Museum, Mycological Department, Václavské nám. 68
115 79 Praha 1, Czech Republic
Abstract—Original descriptions, types and other material of Gymnopilus bellulus
described from North America and Gymnopilus microsporus from Europe were
studied to determine whether the European G. bellulus differs sufficiently from the
American concept to be named G. microsporus. It was found that the G. bellulus
type matches the original description and recent material from both continents and
that American and European collections of G. bellulus are conspecific. Singer based
Gymnopilus microsporus on a collection from the Caucasus validated by Bresadolaʻs
Latin description of Flammula liquiritiae. F. liquiritiae sensu Bresadola is difficult
to interpret due to discrepancies in its description. The only available herbarium
specimens represent Pholiota mixta, whereas the Singerʼs material of G. microsporus
is identical with Gymnopilus bellulus. This paper selects a lectotype for Gymnopilus
bellulus and epitype for G. microsporus to support the fact that G. microsporus sensu
Singer is a synonym of G. bellulus.
Key words—fungi, type study
Introduction
Gymnopilus bellulus (Peck) Murrill is a well known fungus included in most
mycological floras and monographs of Gymnopilus in Europe and North America (e.g.
Kühner & Romagnesi 1953, Hesler 1969, Moser 1983, Singer 1986, Høiland 1990,
Orton 1993, Breitenbach & Kränzlin 2000, Keller & Moser 2001, Ludwig 2000, 2001,
Bon & Roux 2002). However, some problems concerning its original versus recent
concepts and interpretation of type specimen are mentioned by Hesler (1969: 38) and
Singer (in Moreno & Esteve-Raventós 1990: 288). They indicate that the recent concept
(species with spores measuring about 4.5–6.5 x 3–4 µm) is not in accordance with the
type specimen, which is reputedly a fungus with much larger spores (8–9.5 x 5–5.5 µm,
see Hesler 1969: 38).
Gymnopilus microsporus (Singer) ex Singer is a similar but less well-known fungus
omitted in most recent taxonomical works on Gymnopilus. It is only included in the
key by Moser (1983) and discussed by Moreno & Esteve-Raventós (1990). Works only
mentioning the name are not included here. Moreno & Esteve-Raventós (1990) consider
G. microsporus a correct name for Gymnopilus bellulus sensu European authors (non
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Peck), but with a question mark. To clear up these discrepancies, I tried to revise original
descriptions, types and material collected by Peck and Singer. The aim was to evaluate
the taxonomic concept of both taxa. The study was carried out within the framework of
my monographic work on the genus Gymnopilus in the Czech Republic and adjacent
regions of Central Europe (Holec 2001b, 2002, Holec et al. 2003). Type studies in
Gymnopilus are recently carried out also by Guzmán-Dávalos (2003).
Material and Methods
Microcharacters were studied on material mounted in a 5 % KOH solution. Pigmentation
of hyphae of the pileus and stipe cuticle was studied in pure water and their arrangement
was observed on cap surface. Iodine reaction was studied in Melzerʼs reagent prepared
according to the formula given in Moser (1983), cyanophilous reaction in cotton blue
(according to Kotlaba & Pouzar 1964, Singer 1986) after short boiling. Examinations
were made using an Olympus BH-2 microscope. For spore size measurements, 20
randomly selected mature spores were used per collection. Immature spores (extremely
small ones or those having a thin wall and hyaline content) and aberrant spores (1.5-
2 times longer than the normal ones) were not measured. Marginal values of spore
size are given in brackets and represent at most 10 % of all spores measured in each
species. Illustrations of microcharacters were drawn at a magnification of 1250x using
the drawing tube.
Acronyms of herbaria are cited according to Holmgren et al. (1990). Abbreviations: E:
length/width ratio of the spores. Q: mean value of E for all spores studied.
Results
Gymnopilus bellulus (Peck) Murrill
Bas.: Agaricus bellulus Peck, Bull. Buffalo Soc. Nat. Sci. 1: 51, 1873.
≡ Gymnopilus bellulus (Peck) Murrill, North American Flora, vol. 10, part 3: 200, 1917. ≡
Naucoria bellula (Peck) Sacc., Syll. fung. 5: 841, 1887. ≡ Flammula bellula (Peck) Pilát, Klíč k
určování našich hub hřibovitých a bedlovitých (Agaricalium europaeorum clavis dichotomica):
351, 1951.
Lectotype (designated here): USA, New York, herb. NYS: mycology type database
450; data from old label and original newspaper envelope: Agaricus bellulus, Lowville,
Lewis Co., Sept., leg. C.H. Peck 68; the specimen is labelled as “type” (on box) or
“syntype” (on computer-written label). The collection consists of about 20 well-
preserved but slightly overdried mature and young fruitbodies.
Macrocharacters: see the original description (Peck 1873: 51) and later descriptions
by Peck (1874: 58), Hesler (1969), etc.
Lectotype study: Spores small, (5.2–)5.6–6.4 x 3.2–3.6 µm, E = 1.56–2.00, Q = 1.72,
rusty yellow in KOH, ellipsoid to ellipsoid-amygdaliform, with distinct suprahilar
depression in side view, wall rusty, surface moderately to coarsely verrucose, without
plage, slightly dextrinoid (with pale reddish-brown tinge in Melzerʼs reagent), mature
spores acyanophilous or only slightly cyanophilous, the immature ones and those with
363
a broken wall distinctly cyanophilous. Basidia 4-spored, 19–22 x 5–6 µm, cylindrical
to narrowly clavate with a median constriction. Cheilocystidia numerous but poorly
preserved, mostly collapsed, 18–25 x 3–6 µm, cylindrical to narrowly lageniform with
obtuse to capitate apex, neck 1.5–2 µm, head 3–4 µm, thin-walled. Pleurocystidia not
observed. Lamellar trama regular, made up of cylindrical hyphae 2–8(–10) µm wide,
wall yellow-rusty incrusted. Pileus cuticle studied on surface only in order not to damage
the material, consisting of cylindrical to slightly inflated cells 4–13(–16) µm wide,
terminal elements slightly clavate, cells incrusted in a “tiger” pattern, pileocystidia not
observed. Stipe cuticle a cutis of densely arranged parallel hyphae 1.5–3 µm broad, with
yellow-rusty incrustations, at places with nests of interwoven hyphae, caulocystidia not
observed. Clamp connections present.
Fig. 1. Collections identified as Gymnopilus bellulus by C.H. Peck (as Agaricus bellulus or
Naucoria bellulus on original labels). 1: Lowville, lectotype, NYS. 2: Forestburgh, NYS. For
details see collections studied. Note the clear difference in spore size and shape of cheilocystidia.
B: basidium, CH: cheilocystidia, S: spores. Scale bar = 10 µm.
Conclusion: The lectotype agrees not only with the original description of Gymnopilus
bellulus by Peck but also with material from the USA and Europe commonly identified
as G. bellulus (see the collections studied). This means that the American and
European collections can be considered conspecific, at least according to the classical
morphological and anatomical characters.
Discussion: Peck (1873, 1874) characterised his Agaricus bellulus as a smal fungus
with bright watery-cinnamon pileus, yellow lamellae, reddish-brown stipe and spores
measuring 5 x 3.5 µm. At the subgeneric level, he placed it into Naucoria. Some
authors (e.g. Moreno & Esteve-Raventós 1990) are convinced that G. bellulus in the
sense of recent (mostly European) authors is different from the type of G. bellulus.
Moreno & Esteve-Raventós (1990) suggested to use the name Gymnopilus microsporus
for European G. bellulus. Similarly, Hesler (1969: 38) writes that „Peckʼs type is
confusing“. However, his description (Hesler 1969: 37) well agrees both with the
original description by Peck (1873) and recent concept of G. bellulus.
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Singer (in Moreno & Esteve-Raventós 1990: 288) concluded that the type of Gymnopilus
bellulus represents a species with large spores, whereas G. bellulus of recent authors is
a species with small dextrinoid spores. According to him, G. microsporus sensu Singer
(having non-dextrinoid spores) is not identical with G. bellulus of recent authors but
could be identical with G. bellulus sensu Kühner & Romagnesi (1953). In my opinion,
Singerʼs conclusions based on the reaction of spores with Melzerʼs reagent are of
limited value. In the lectotype of G. bellulus and epitype of G. microsporus I studied,
the reaction is quite identical – the spores of both species are slightly dextrinoid.
To solve all these discrepancies, I tried to obtain the original material of G. bellulus
including the type on loan and compare it with collections from Europe. I obtained 8
specimens of Agaricus bellulus (= Gymnopilus bellulus) identified (and mostly also
collected) by C.H. Peck from NYS herbarium. There were no collections of G. bellulus
by Peck in BUF and NY, the other herbaria housing his material. One of the specimens
from NYS is labelled as type or syntype (Lewis Co., Lowville). Together with the
specimen from Canada collected by J.M. Macoun (see Tab. 1 and the collections
studied) it represents quite another species (having small spores) than the remaining
ones (having much larger spores). The latter collections probably represent a species
close to Gymnopilus picreus (which should be judged by an American monographer of
Gymnopilus).
Tab. 1. Comparison of the spore size in collections of Gymnopilus bellulus at NYS
identified by C.H. Peck
NYS specimens Peck identified as
NAUCORIA BELLULA
(for details see Collections studied)
SPORE SIZE IN µM
(specimens with small spores)
SPORE SIZE IN µM
(specimens with large spores)
Lowville (lectotype) (5.2–)5.6–6.4 x 3.2–3.6
Canada 4.8–5.6 x 3.2–3.6
Adirondacks Mts. 8.4–9.6 x 6.0–6.4
Forestburgh (8.8–)9.2–9.6 x 6.0–6.4(–6.8)
Warrensburg (8.0–)8.8–9.2 x 5.6–6.8
Catskill Mts. 8.8–9.6 x 6.0–6.4
Lewis Co., Greig 9.2–10.0 x 6.0–6.4
North Elba (8.4–)8.8–9.6 x 6.0–6.8
As all collections were identified by Peck, it seems that he did not have a clear concept
of his new species. Similarly, L.R. Hesler, a monographer of Gymnopilus, writes
(Hesler 1969: 38) that „Peckʼs type is confusing, it appears that the labels of the type
and of a collection of another species have been interchanged. In the collection marked
type and which we examined, the spores are 8–9.5 x 5–5.5 µm – much too large for G.
bellulus“.
All collections I had on loan except for the syntype from Lowville and the Canadian
collection (see above) really have the large spores as indicated by Hesler. However, the
syntype from Lowville (for type study see above) well agrees with both the original
description by Peck and the current concept of Gymnopilus bellulus, which is a species
365
with spores measuring about 4.5–6.5 x 3–4 µm. It is hard to explain why Hesler
found much larger spores in the type specimen. However, he does not indicate exactly
which type collection he studied (he only writes “collection marked type which we
examined“). It could be a collection from the second locality given by Peck (1873) in
his original description (Sandlake).
I asked current NYS, curator John Haines whether such a collection is available at NYS.
He wrote me that „we do not have the second presumed Sandlake syntype (presumed
because it was mentioned in the protologue)“ and that the specimen „has not been seen
since I came to the museum in 1969“.
Fortunately, the syntype from Lowville is available and agrees well with the current
concept of Gymnopilus bellulus. Consequently, it is logical to designate it as the
lectotype of Gymnopilus bellulus. In this way, the current concept of G. bellulus will be
fixed in spite of the discrepancies in the material collected by Peck.
Another bit of helpful information is a painting made when the Lowville specimen was
fresh. John Haines scanned and sent it to me via Internet. According to him, „it bears the
same collection number as the Lowville specimen and was removed from that box some
years ago“. The depicted fungus looks exactly like Gymnopilus bellulus, which I know
well from the field in Europe (in paragraph „Collections studied“ I cite only a limited
selection of representative specimens from Europe which I revised for comparison with
the type).
By the way, the original envelope of the syntype from Lowville (made from a newspaper
from Peckʼs time) is labelled Agaricus (Naucoria) obnoxius (original hand writing by
Peck?). It was probably a provisional name which was changed to Agaricus bellulus in
the original publication. John Haines confirmed to me that he recorded that provisional
name in Peckʼs field notes.
Collections studied (representing G. bellulus): USA, New York, Lowville, Lewis Co.,
Sept., leg. C.H. Peck 68 (NYS: mycology type database 450; lectotype of Gymnopilus
bellulus, designated here, see paragraph Lectotype). – Canada, 22 Aug 1892, leg.
J.M. Macoun, det. C.H. Peck as Naucoria bellula (NYS). – Austria, Niederösterreich,
Hohenberg, Laghsattel, Abies, 30 June 1992, leg. A. Hausknecht (WU 10824). – Austria,
Niederösterreich, Lunz/See, Rotwald: NSG Kleiner Urwald, wood of conifer (Abies?),
29 Aug 1996, leg. A. Hausknecht (WU 16262). – Czech Republic, Beskydy Mts.,
Morávka, Travný, Picea abies, on decayed trunk, 17 Sep 1987, leg. Z. Pouzar (PRM
852262). – Czech Republic, Novohradské hory Mts., Žofínský prales virgin forest,
Fagus, on trunk, 21 Sep 1991, leg. V. Antonín (BRNM 553290). – Czech Republic,
Šumava Mts., Mt. Boubín, Boubínský prales virgin forest, Aug 1936, leg. J. Herink
(PRM 27788). – Czech Republic, Šumava Mts., České Žleby, Mt. Spáleniště, Abies
alba, on decayed trunk among mosses, 26 Oct 2002, leg. J. Holec (PRM 900962). –
Czech Republic, Šumava Mts., Nová Pec, Mt. Plechý, Picea abies, on decayed trunk,
26 Aug 1996, leg. J. Holec, 317/96 (PRM 889100). – Italy, Trento, Paneveggio, wood of
a conifer, 4 Oct 1989, leg. R. Schütz (WU 8143). – Ukraine, Eastern Carpathians, near
Dilove („Trebušany“), Berlebash („Berlebaš“) stream valley, Picea abies, Aug 1937,
leg. A. Pilát (PRM 488229).
Collections representing other species (close to G. picreus): USA, New York, North
Elba, Sept., leg. C.H. Peck, det. C.H. Peck as Naucoria bellulus (NYS) – ditto, Catskil
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Mts. (NYS). – ditto, Warrensburg, Oct. 19 (NYS). – ditto, Forestburgh (NYS). – ditto,
Lewis Co., Greig (NYS). – ditto, Adirondack Mts., on mossy hemlock trunk (NYS).
Gymnopilus microsporus (Singer) ex Singer
≡ Fulvidula microspora Singer, Rev. Mycol. 2: 239, 1937 (invalid name, as the generic name
Fulvidula is invalid, see e.g. Donk 1962). Singer (1937) published the name as a „nomen novum“
for Flammula liquiritiae (Pers.) P. Kumm. sensu Bres., Icon. mycol., vol. 16, text + tab. 783, 1930.
This is, however, not a nomen novum (see ICBN Art. 33, Note 2). Fulvidula microspora must be
treated as an invalid name of a new species. The valid publication was made later:
≡ Gymnopilus microsporus (Singer) ex Singer, Lilloa 22: 561, 1951 („1949“).
Holotype: Bresadola, Icon. mycol., vol. 16: tab. 783, 1930 (illustration serving as
holotype); as the holotype is ambiguous and cannot be critically identified for purposes
of the precise application of the name, an interpretative type – epitype – is selected here
(for discussion see below).
Epitype (selected here, explanations of data from the original label are given in
brackets): Russia (former USSR), Caucasus, Kavkazskiyi Zapovednik (north-western
part of the Caucasus), ad truncos putridos coniferarum, leg. L.N. Vasilieva in 1936 (LE
17635), for discussion see below.
Revision of Gymnopilus microsporus sensu Singer
Macrocharacters: Singer (1937, 1951) refers to Bresadola (Icon. mycol., vol. 16, text
+ tab. 783, 1930).
Epitype study (the material is poorly preserved, with many collapsed cells): Spores
small, 4.4–4.8 x 2.8–3.2 µm, E = 1.38–1.71, Q = 1.51, yellow in KOH, ellipsoid in
front view, ellipsoid-amygdaliform with a slight but distinct suprahilar depression in
side view, wall yellow-brown, thick, medium to coarsely verrucose, slightly dextrinoid
(with pale reddish-brown tinge in Melzerʼs reagent), mature spores acyanophilous or
only slightly cyanophilous, the immature ones or those with a broken wall distinctly
cyanophilous. Basidia mostly collapsed, 4-spored, 16–22 x 4–5 µm, cylindrical, with
long sterigmata, often filled with rusty brown pigment. Cheilocystidia difficult to find,
probably collapsed, about 20–25 x 5–6 µm, narrowly lageniform with a slightly capitate
apex, filled with homogeneous rusty brown pigment. Pleurocystidia not observed.
Pileus cuticle (observed on surface only in order not to damage the fruitbodies) a cutis
of hyphae (5–)8–12(–14) µm broad, cells cylindrical or slightly inflated, terminal
elements clavate, up to 24 µm broad, wall coarsely rusty brown incrusted in a “tiger”
pattern, pileocystidia not observed. Stipe made up of cylindrical hyphae 4–10.5 µm
broad, cuticle a cutis of cylindrical hyphae 2.5–4 µm broad, at places (rarely) with
cylindrical outgrowths up to 5 µm long, in mass rusty brown, wall with rusty brown
membranal pigment, not heavily incrusted. Clamp connections present in all tissues.
Conclusion: The specimen LE 17635 represents the species Gymnopilus bellulus
(Peck) Murrill. The same opinion was expressed by Emma L. Nezdoiminogo on a
revision label on the envelope of LE 17635 written in 1985. However, her conclusion
was not published (personal communication). If only the herbarium specimens from
then Caucasus and its microcharacters are considered, then Gymnopilus microsporus
367
sensu Singer (1937) represents a synonym of Gymnopilus bellulus.
Discussion: Although there exists a herbarium specimen (LE 17635, for details see
paragraph Epitype) on which the name G. microsporus is based (the specimen is
labelled „holotypus“ in LE), it is not the holotype. The name Gymnopilus microsporus
is validated by Bresadolaʼs Latin description as Singer (1937, 1951) did not provide
description, Latin diagnosis, or designation of type in his publications. Consequently,
elements from the context of the validating description should be used for typification
(see ICBN Art. 7.7 Ex. 4).
Flammula liquiritiae (Pers.) P. Kumm. sensu Bresadola, on which the name G.
microsporus is based, is a fungus that is hard to interpret. In its description and
illustration, Bresadola presents a fungus with a pholiotoid to gymnopiloid habit and
small spores which may be smooth or minutely verrucose (?!). The cystidia he presents
are typically pholiotoid (see the same conclusion by Singer in Moreno & Esteve-
Raventós 1990: 288). They are too large for Gymnopilus (50–60 x 10–15 µm, whereas
20–40 x 5–10 µm is typical for European species of Gymnopilus) and lageniform-
fusiform (in Gymnopilus, the cystidia are more or less capitate). I do not know such
a fungus among European species of Gymnopilus and it is neither mentioned in any
critical treatments of Gymnopilus in Europe. Such a mixture of characters really could
be a result of erroneous observations by Bresadola (see also Singer in Moreno & Esteve-
Raventós 1990: 288) or because his illustration and description of macrocharacters is
based on other material than the description of microcharacters.
I contacted herbaria housing the collections by Bresadola (S, BPI, M, TR) to ask if there
are any collections of Flammula liquiritiae identified by him. There are no specimens
in S, BPI, and M, but two collections are available at TR (Museo Tridentino di Scienze
Naturali, Italy). They were collected by Julius Schaeffer in Germany (Potsdam),
identified by Bresadola (see the collections studied) and most probably used for his
description of Flammula liquiritiae. At first I thought that they represent syntypes of
Gymnopilus microsporus from which the lectotype could be selected. However, some
data from their labels are not in accordance with data in Bresadolaʼs description (Tab.
2).
Tab. 2. Differences between characters given by G. Bresadola on labels of Flammula
liquiritiae in TR herbarium and in his description published in Iconographia
mycologica.
CHARACTER LABEL
(TR B/228, TR B/239)
DESCRIPTION
(Bresadola 1930: tab. 783)
Substrate in pinetis ad terram (= on soil in
Pinus forests)
ad truncos coniferarum in
silvis (= on trunks of conifers
in forests)
Size of basidia 24–26 x 6–7 µm 28–32 x 6–8 µm
Spore size 5–6 x 3.5–4 µm 4,5–6 x 3–4 µm
Distribution or location Potsdam (Germany) Europe, Asia (Siberia)
368
It seems that the description by Bresadola was based not only on the two collections
studied but also on other collections (e.g. from Asia), as the data in the description are
broader or different than those on labels.
A careful microscopic revision of the two collections from TR clearly showed that
they represent in fact Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk. The spores are
smooth, measure 6.0–6.5(–6.8) x 3.4–4 µm and numerous cheilo- and pleurocystidia
of lageniform-fusiform shape are present. I am quite certain with this identification as
I know the species well (see Holec 2001a: 105–110). The assumption on pholiotoid
cystidia in Flammula liquiritiae expressed above proved to be correct.
It is unclear why Bresadola described the spores as smooth or minutely verrucose. This
is either an observation error (which is quite understandable) or a consequence of the
fact that other collections belonging e.g. to Gymnopilus and having verrucose spores
were used for the description of his Flammula liquiritiae.
Generally, the description and illustration of Flammula liquiritiae (Pers.) P. Kumm.
sensu Bresadola (1930) is hard to interpret and the only collections available represent
Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk. The discrepancies between the description
and labels of these herbarium specimens do not enable to consider them types.
In such a situation, the holotype of Gymnopilus microsporus is represented by the
illustration by Bresadola (Icon. mycol., vol. 16: tab. 783, 1930) – the only type material
available. It is interesting that the illustration is cited by Hesler (1969: 65) as typical
of Gymnopilus liquiritiae, although he mentions much larger spores in his description
than Bresadola.
With respect to all these discrepancies, it is better to use the collection from LE identified
by Singer for interpretation of the name G. microsporus (see the microscopic study
above). Singer himself (in Moreno & Esteve-Raventós 1990: 288) confirms that he
knows the discrepancies in Bresadolaʼs publication and that he follows his own concept
of G. microsporus (based on material from the Caucasus). In this situation, I decided
to designate the collection from LE as the epitype. If such a concept is accepted, then
Gymnopilus microsporus (Singer) ex Singer is a species identical with Gymnopilus
bellulus (Peck) Murrill and the name must be considered a synonym of G. bellulus. In
my opinion, this is the best treatment of the name G. microsporus.
Collections studied: Russia, Caucasus, Kavkazskiyi Zapovednik, ad truncos putridos
coniferarum, leg. L.N. Vasilieva in 1936 (LE 17635, epitype of Gymnopilus microsporus,
selected here, see paragraph Epitype). – Germany, Potsdam, in pinetis ad terram, leg. J.
Schaeffer, det. G. Bresadola (TR: herbarium Bresadola B/239). – ditto (TR: herbarium
Bresadola B/228).
Acknowledgements
I thank Zdeněk Pouzar (Praha), Klaus Høiland (Oslo), and Vladimír Antonín (Brno) for their
valuable advice and comments on the manuscript. Curators of the following herbaria are kindly
acknowledged for arranging loans from their institutions: LE, NYS, MICH, TR or providing
information: BUF, M, NY, S. The work was financially supported by the Grant Agency of the
Czech Republic (project no. 206/01/P05) and the Ministry of Culture of the Czech Republic
(MK0CEZ99F0201).
369
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