A confusing duo: Calocybe cerina and Callistosporium pinicola 
(Agaricales)

EEF ARNOLDS

Holthe 21, NL 9411 TN Beilen, eefarnolds@hetnet.nl

A r n o l d s  E.: A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales). 
Acta Mycol. 41 (1): 29 40, 2006.

The name Calocybe cerina has recently been applied for two different species of agarics. 
After studying collections from different European countries it is concluded that the true 
Calocybe cerina is very close to C. chrysenteron and best regarded as a variety. The new 
combination Calocybe chrysenteron var. cerina is proposed and a full description of that taxon 
is given. The majority of collections, identified as C. cerina, appear to belong to a fungus, 
provisionally described by Bon as Callistosporium luteoolivaceum fo. minor. It is described here 
as a new species: Callistosporium pinicola. Calocybe juncicola is another species that might be 
confused with the two species mentioned before and therefore a full description is provided. 
The status of the genera Rugosomyces and Calocybe is discussed. The new combinations 
Calocybe obscurata and Calocybe pudica are made.

Key words: Callistosporium, Calocybe, Rugosomyces

INTRODUCTION

In The Netherlands in recent years several collections were made of a fairly small, 
white-spored agaric with an orange-brown pileus and crowded, orange-yellow lamel-
lae, growing on stumps of coniferous trees. It was initially identified as Calocybe 
cerina (Pers.: Fr.) Donk (A r n o l d s , B e c k e r  1993). An important indication was 
the presence of granules in the basidia that were apparently siderophilous, staining 
dark violet in acetocarmine with addition of iron. However, some characters were 
deviating from most descriptions in literature, in particular the striking red to violet 
staining of all tissues in KOH and ammonia, the complete absence of clamp-connec-
tions and the lignicolous habitat, suggesting that it could be an undescribed taxon. 

Recently I came across descriptions of Callistosporium luteoolivaceum var. minor 
M. Bon ined. (L u d w i g  2001; W i l h e l m  2003) that showed striking resemblance 
with our material, e.g. the red staining in ammonia and KOH. In this paper I shall try 
to unravel the identity of the collected material and I will discuss the taxonomic posi-
tion of Calocybe cerina (Pers.: Fr.) Donk and the related C. juncicola (Heim) Sing.

 ACTA MYCOLOGICA
 Vol. 41 (1): 29-40
 2006

Dedicated to Professor Alina Skirgiełło
on the occasion of her ninety fifth birthday



30 E. Arnolds 

WHAT IS CALOCYBE CERINA?

Agaricus cerinus was initially described by P e r s o o n  (1801: 321) as a fairly small 
agaric, growing in pine forests (substrate not explicitly mentioned) with dark wax-
yellow (‘flavo-cerinus opacus’), fleshy, flattened to depressed pileus; crowded, nar-
row lamellae and a taste becoming bitter after a while. In his sanctioning description 
F r i e s  (1821:89) mainly copied Persoon’s description, adding that the context and 
spore print are white. Most authors have interpreted the name as a species close to 
or identical with Calocybe chrysenteron (Bull.: Fr.) Sing. P e r s o o n  (1801) treated 
Agaricus chrysenterus Bull. immediately after his A. cerinus. The two taxa are micro-
scopically characterized by e.g very small, ellipsoid spores, a pileipellis in the form of 
a cutis with transitions to a trichodermium, and presence of clamp-connections.

A different interpretation of Agaricus cerinus was published by A r n o l d s  & 
B e c k e r  (1993) and accepted by e.g. K a l a m e e s  (1995, 2004). The basidiocarps of 
that fungus occur on dead wood, are slightly smaller and thin-fleshed, their tissues 
turn reddish in alkaline solutions and clamp-connections are absent.

After comparing collections and modern descriptions of these taxa I believe that 
the first-mentioned interpretation is in better agreement with the original diagno-
sis, mainly in view of the larger size of the basidiocarps, the thicker context and the 
substrate on soil and litter. Therefore I present first a description of this little-known 
fungus. 

Calocybe chrysenteron (Bull.:Fr.) Sing. var. cerina (Pers.: Fr.) Arnolds nov. comb. 
– Fig. 1.

Basionym: Agaricus cerinus Pers.: Fr., Syst. mycol. 1: 89. 1821.
Agaricus cerinus Pers., Syn. meth. Fung.: 321. 1801; Agaricus cerinus Pers.: Fr., 

Syst. mycol. 1: 89. 1821; Tricholoma cerinum (Pers.:Fr.) Quél., Champ. Jura Vosges 
1; 81. 187; Tricholoma chrysenteron subsp. cerinum (Pers.: Fr.) Konr. & M., Icon. sel. 
Fung. 3: pl. 267, fig. 2. 1937; Calocybe cerina (Pers.: Fr.) Donk in Beih. Nova Hedwi-
gia 5: 43. 1962; Rugosomyces cerinus M. Bon in Doc. mycol. 21 (82): 66. 1991. – Tri-
choloma pseudoflammula J. Lange in Dansk bot. Ark. 8(3): 24. 1933; Spec. Pl.: 1173. 
1753; Calocybe pseudoflammula (J. Lange) Sing. in Sydowia 15: 47. (‘1961’) 1962; 
Rugosomyces pseudoflammula (J. Lange) M. Bon in Doc. mycol. 21(82): 65. 1991.

Excl. – Calocybe cerina sensu Arnolds & Becker in Coolia 36: 70-73, fig. 1. 1993 
(  Callistosporium pinicola); sensu Dermek in Fung. rar. Ic. col. 17: 10. 1987 (  C. 
fallax); sensu Kalamees in Scripta mycol. 18: 84. 2004 (  Callistosporium pinicola); 
Rugosomyces pseudoflammula sensu M. Bon, Fl. Mycol. Eur. 5, Collybio-Marasmïoï-
des: 110. 1999 (  C. chrysenteron).

Sel. icon. – Cetto, Funghi Vero 3: pl. 1024. 1979; Flora Batava 25 : pl. 1994a. 
1920; Konr. & M., Icon. sel. Fung. 3: pl. 267, fig. 2. 1927; J. Lange, Fl. Agar. Dan. 1: 
pl. 24B. 1935 (as T. pseudoflammula); Schweizer Pilzt. 5: pl. 42. 1972.

Sel. descr. & figs. – M. Bon, Fl. Mycol. Eur. 5, Collybio-Marasmïoïdes: 111. 1999; 
Konr. & M., Icon. sel. Fung. 3: pl. 267 II. 1927; J. Lange, Fl. Agar. Dan. 1: 57. 1935 
(as T. pseudoflammula).

Pileus 12-40(-50) mm, convex at first, with thin, involute margin, then plano-
convex to flattened or slightly depressed with straight, often undulating margin, with 
or without weak umbo, not hygrophanous, brownish yellow, brownish orange to or-
ange-brown from the beginning (e.g. K. & W. 5B7, 5C8, 6C7, 6C8), dull and dry, 



 A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales) 31

smooth to slightly tomentose, especially at centre, not striate. Lamellae, L  38-60, 
l  1-7, adnexed, emarginate to adnate, very crowded, segmentiform, up to 5 mm 
broad, thin, lemon- to golden-yellow or orangeish yellow, with concolorous, entire 
edge. Stipe 20-50 x 2-6(-10) mm, cylindrical or compressed, often tapering to base, 
stuffed to fistulose, concolorous with pileus, sometimes weakly yellow striate length-
wise, at apex pruinose, at base white tomentose or strigose-hairy. Context whitish, 
pale yellow to lemon-yellow. Smell farinaceous when cut; taste first farinaceous, 
then often more or less bitter. Spore print white.

Spores 2.5-4.0(-4.5) x 2.0-3.0 μm, av. 3.1-3.5 x 2.3-2.5 μm, Q  1.2-1.5, Qav.  1.3-
1.4, broadly ellipsoid to ellipsoid. Basidia 17-23 x 4.0-7.0 μm, clavate, 4-spored, with 
numerous small granules. Lamella edge fertile. Hymenophoral trama subregular, 
made up of hyphae with cylindrical or slightly inflated elements, 3.0-12 μm broad. 
Pileipellis a cutis of repent hyphae, 2.0-6.0 μm broad, often with some broader, el-
lipsoid to subglobose cells, often at centre with trichodermial tufts of erect and as-
cending hyphae, with yellowish intracellular pigment. Stipitipellis a cutis, made up of 
repent hyphae, 2.0-4.0 μm wide, at stipe apex with clusters of undifferentiated, erect 
hyphal tips up to 40 μm long, 2.0-5.0 μm wide. Clamp-connections present. 

Chemical reactions: No part of basidiocarp reddening with KOH or ammonia. 
Granules in basidia staining dark purple in acetocarmine with iron (siderophilous).

Habitat and distribution: Mainly on needle litter in coniferous forests (Picea, 
Abies), also reported under broad-leaved-trees (Quercus); on mesic to dry, calcare-
ous soil. Probably widespread in Europe but much rarer than var. chrysenteron. Ex-
act distribution unknown in view of taxonomic confusion (see notes). With certainty 
recorded from Denmark, Germany, Switzerland (K o n r a d & M a u b l a n c 1937) 
and The Netherlands (Flora Batava, 1920).

Collections examined. – DENMARK: Rubjerg, Knude Plantage, 26 Sept. 1990, 
J. Vesterholt 90-505 (C); Same loc., 21 Oct. 1991, J. Vesterholt 91-801 (C) – GER-
MANY: Bavaria, Frankische Schweiz, Ebermannstadt, Aufsesstal, 19 Sept. 2005, E. 
Arnolds 05-87 (L).

Fig. 1. Calocybe chrysenteron var. cerina. A. Basidiocarps x 1; B. Spores x 2000; C. Basidia x 
1000; D. Pileipellis x 1000 (all from Arnolds 05 87).



32 E. Arnolds 

The studied collections differ from typical Calocybe chrysenteron morphologically 
only in the duller, more brownish colours of pileus and stipe and the paler context. 
In var. chrysenteron all parts of the basidiocarps are bright yellow to orange-yellow 
(e.g. L u d w i g  2000). In addition var. cerina seems to be almost confined to conifer-
ous forests, whereas var. chrysenteron is mostly (but not always) found in deciduous 
forests. Both taxa are calciphilous. Some modern authors synonymize C. chrysenter-
on and C. cerina (e.g. L u d w i g  2001), others treat them as separate species (e.g. 
B o n  1999; H o r a k  2005). The differences are small but seem to be rather constant 
and warrant in my opinion a distinction in the rank of variety.

Many modern authors regard Tricholoma pseudoflammula J. Lange as a synonym 
of Calocybe chrysenteron (e.g. L u d w i g  2001; K a l a m e e s  2004), but Lange’s plate 
(24B) is typical of var. cerina in view of the orange-brown pileus, already in young 
basidiocarps, and the pale context. 

For differences with C. cerina sensu Arnolds and Becker (  Callistosporium pini-
cola) see notes on that species.

A DOUBLE OF CALOCYBE CERINA: CALLISTOSPORIUM PINICOLA

Callistosporium pinicola Arnolds nov. spec. – Fig. 2.
Pileus (5-)10-30 mm convexus margine involutus, dein planiusculus vel depres-

sus, haud vel leviter hygrophanus, flavo-brunneus, aurantio-brunneus, rufo-brun-
neus, numquam striatus, siccus, centro tomentosus. Lamellae adnatae vel emargina-
tae, confertae, tenuae, flavae vel aurantio-flavae. Stipes (12-)14-32 x (1,5-)2-3,5 mm, 
cylindricus vel compressus, pileo subconcolorus. Caro tenuis, fragilis, pallide flava. 
Odor nullus vel farinaceus. Sapor nullus vel amarescens. 

Fig. 2. Callistosporium pinicola. A, E. Basidiocarps x 1; B, F. Spores x 2000; C, G. Basidia x 1000; 
D, H. Pileipellis x 1000 (A D from Arnolds 6100 (holotype), E H form A. Becker, 6 Oct. 1990).



 A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales) 33

Sporae (2,5-)2,8-4,7 x 2,0-3,5 μm, ellipsoideae vel late ellipsoideae, laeves, hya-
linae, saepe corpusculis xanthis vel brunneis. Basidia 12-25 x 3,5-6,0 μm, clavatae, 
tetraspora. Lamellarum trama regularis. Cystidia nulla. Pileipellis cutis vel trichode-
rmium, hyphis 2,0-5,5 μm latis, interdum ex cellulis inflatae ad 8,0 μm. Fibulae ab-
sentes. Basidia, trama et pileipellis granulis numerosis brunneis in aqua, rubescentis 
vel violascentis in KOH et ammonia.

Habitatio ad truncos putridos coniferarum. Aestate- automno. 
Holotypus: ‘The Netherlands, Dwingeloo, Lheederzand, 4 Oct. 1990, E. Arnolds 

6100’ (L).
Synon.: Callistosporium xanthophyllum (Malenc. & Bert) M. Bon fo. minor M. 

Bon ad int., in Flore mycol. Europe 2: 83. 1991 (invalid, without latin diagnosis and 
without designation of type).

Misapplied names: Calocybe cerina sensu Arnolds & Becker in Coolia 36: 70. 
1993; sensu Kalamees in Scripta mycol. 18: 84. 2004.

Sel. plates. – Ludwig, Pilzkompendium 1: pl. 15, fig. 6.2 A, C. 2000 (as Callist-
osporium luteoolivaceum var. minor M. Bon ined.); Wilhelm in Schweiz. Z. Pilzk. 81: 
64. 2003 (as Callistosporium luteoolivaceum var. minor M. Bon).

Sel. descr. & figs. – Arnolds & Becker in Coolia 36: 70-73, fig. 1. 1993 (as Calo-
cybe cerina); Ludwig, Pilzkompendium 1: 37-38. 2001 (as Callistosporium luteooli-
vaceum var. minor M. Bon ined.); Wilhelm in Schweiz. Z. Pilzk. 81: 62-66. 2003 (as 
Callistosporium luteoolivaceum var. minor M. Bon) .

Pileus (5-)10-30 mm, first convex with narrow, involute margin, then plano-con-
vex to flattened with straight margin, often with slightly depressed centre, not or 
weakly hygrophanous, usually vividly yellow-brown, orange-brown, rusty-brown to 
red-brown, occasionally with olivaceous hue near centre (e.g. K. & W. 5D7, 6D7, 
7E7; Mu. 5 YR 6/6, 10 YR 4/6, 5/8), rarely entirely butter-yellow (K. & W. 4A5), not 
striate, dry, smooth or slightly radially fibrillose, often at centre tomentose. Lamel-
lae, L  28-36, l  (1-)3-7, adnate to strongly emarginate, crowded to very crowded, 
segmentiform, up to 4 mm broad, thin, butter-yellow, golden-yellow to yellowish or-
ange (Mu. 2.5 Y 6/8; K. & W. 4A5, 5C6, 4C6), with concolorous, entire edge, becom-
ing dark red-brown to almost black in exsiccata. Stipe (12-)14-32 x (1.5-)2-3.5 mm, 
cylindrical or sometimes compressed, solid or narrowly fistulose, concolorous with 
pileus or paler yellow-brown, slightly aerenchymatic at first, then smooth or white 
striate lengthwise, at apex slightly pruinose, at base often white tomentose. Context 
in pileus thin, up to 2 mm thick, fragile, concolorous with surface, inside stipe pale 
yellow. Smell almost absent or weakly farinaceous; taste mild to slightly bitter. Spore 
print pale cream-coloured.

Spores (2.5-)2.8-4.5 x 2.0-3.5 μm, av. 3.0-4.2 x 2.3-3.4 μm, Q  1.1-1.5(-1.8), 
Qav.  1.2-1.4, in majority broadly ellipsoid or ellipsoid, some subglobose or oblong, 
smooth, not amyloid, in water colourless or with yellow, refractive body, in part also 
with brown granules (‘necropigment’), staining reddish in ammonia and KOH, dark 
red to violet-brown in congored and acetocarmine. Basidia 12-25 x 3.5-6.0 μm, cla-
vate, 4-spored, in water with numerous brown granules and larger clots, staining like 
spores. Lamella edge fertile. Hymenophoral trama subregular, made up of rather 
narrow hyphae with cylindrical or slightly inflated elements, 13-90 x 3.0-14 μm, with 
numerous brown granules and clots. Pileipellis mainly a poorly differentiated cutis, 
made up of repent, interwoven hyphae, 2.0-5.5 μm broad, with yellow intracellu-



34 E. Arnolds 

lar pigment, towards the centre often with trichodermial fascicles of ascending and 
erect hyphae, often with ellipsoid to subglobose terminal cells up to 8.0 μm wide, 
with brown granules and clots. Stipitipellis a cutis, made up of repent hyphae, 2.0-4.0 
μm wide. Clamp-connections absent. 

Chemical reactions: all parts of the fresh basidiocarp turning immediately dark 
red-brown to violet-brown with KOH 5% and 10% ammonia (macroscopically); 
preparations of all tissues turning immediately violet in KOH 5% and reddish in 
10% ammonia, also staining surrounding liquid; granules and clots becoming vina-
ceous red or dark violet. Granules staining dark purple in acetocarmine with iron 
(seemingly siderophilous). 

Habitat and distribution: Saprotrophic, solitary or in small groups, on strongly 
decayed stumps or dead trunks of coniferous trees, mainly recorded from Pinus syl-
vestris, also on Pinus pinea and Picea abies, in coniferous and mixed stands, mainly on 
acidic, sandy and loamy soils. July-Nov. Apparently widespread but rare in Central-
Europe; recorded from The Netherlands, southern and eastern Germany (L u d w i g 
2001; W i l h e l m  2003), Switzerland (K a l a m e e s  2004, as C. cerina), Austria (K a -
l a m e e s  2004, as C. cerina), France (W i l h e l m  2003; B o n  1995) and Italy.

Collections examined. AUSTRIA: Steiermark, Gleichenberg, Trautmannsdorf, 
21 Aug. 2003, W. Wofar & A. Hausknecht (WU, as Calocybe cerina). – FRANCE: 
Elsass, Hardt, 16 Sept. 1998, M. Wilhelm 596 (herb. Wilhelm); Sondersdorf, 31 July 
2000, M. Wilhelm 709 (herb. Wilhelm); Oltingue, 27 Aug. 2002, M. Wilhelm 793 
(herb. Wilhelm). – ITALY: Tuscany, Orbetello, Bosco di Patanella, 27 Oct. 1987, 
Th. W. Kuyper 2804 (L, as Calocybe cerina). –– THE NETHERLANDS: Dwingeloo, 
Lheederzand, 4 Oct. 1990, E. Arnolds 6100 (L, as Calocybe cerina); Smilde, Hard-
ersbosch, 13 Nov. 1991, E. Arnolds 6232 (L, as Calocybe cerina); Diever, State Forest 
Smilde, 3 Nov. 1986, B. de Vries 5193 (L, as Calocybe spec.); Diever, State For-
est Smilde, 31 Aug. 1992, J. Baar 92/11 (L, as Calocybe chrysenteron); Ede, estate 
Roekel, 21 Sept. 2002, P.J. Keizer s.n. (L, as Calocybe cerina); Bergen-Schoorl, State 
Forest, A.G. Becker, 6 Oct. 1990 (L, as Calocybe cerina).

Callistosporium pinicola does not belong to the genus Calocybe in view of the 
presence of small granules and larger clots of brown necropigment in the basidia, 
spores trama and pileipellis, as well as by the complete absence of clamp-connec-
tions. However, the necropigment can only be recognized as such in preparations in 
water (also of exsiccata), because it is immediately turning red to violet in ammonia 
and KOH. Since exsiccata are usually revived in these solutions the brown pigment is 
easily overlooked in dried specimens. Moreover the necropigment is strongly stain-
ing purple with acetocarmine, thus mimicking the siderophilous granules of Calo-
cybe species. Therefore it is not surprising that the majority of herbarium collections, 
identified as Calocybe cerina, appears to belong to Callistosporium pinicola. In this 
context it is interesting to note that M a i r e  (1937) described Callistosporium luteoo-
livaceum under the name Tricholoma chrysenteron (  Calocybe chrysenteron) var. 
olivascens Maire (R e d h e a d  1982). Apparently confusion between the two genera 
has occurred before.

The presence of necropigment, the red staining of tissues in KOH and ammonia 
and the absence of clamp-connections are all characters of the genus Callistosporium 
Sing. The present species belongs to subgenus Callistosporium in view of the crowd-
ed lamellae, small, ellipsoid spores, short basidia and the pigment not turning blue 



 A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales) 35

in ammonia (Bon, 1991). Taxonomy and nomenclature of this group are still rather 
controversial. Bon (1991) distinguished two species, C. xanthophyllum (Malenc. & 
Bert.) M. Bon and C. elaeodes (Romagn. ex) M. Bon, but R e d h e a d  (1982) and 
N o o r d e l o o s  (1995) united these taxa in one species and synonymised it with the 
North-American C. luteoolivaceum (Berk. & Curtis) Sing, which name has priority in 
that case. L u d w i g  (2001) described C. foetens E. Ludwig as additional new species 
in this subgenus, characterized by a strong unpleasant smell.

Callistosporium pinicola differs from all species mentioned above in the first place 
by the very small spores. Ludwig (2001) described the spores of C. luteoolivaceum 
as 5-5.5(-6.5) x 3.5-4 μm and B o n  (1991, as C. xanthophyllum) as (5.0-)5.5-6.5(-7) 
x (3-)4-4.5(-5) μm. Spores in the type collection of C. elaeodes measured 6.5-7.5 x 
3.5-4.4(-5.0) μm (B o n  1976). In addition it is striking that the basidiocarps of C. 
pinicola are generally lacking green colours, so characteristic for other species of 
Callistosporium. Only in one of the studied collections a weak olivaceous tone was 
observed in the centre of the pileus. The habitat on stumps and trunks of conifer-
ous trees is also characteristic for C. pinicola. Other species are usually growing on 
deciduous wood or litter. In view of these characteristics the identity of the collec-
tion, depicted by L u d w i g  (2000) in Pl. 6.2 B as C. luteoolivaceum var. minor, is in 
my opinion doubtful since the pileus is distinctly green and the spores are described 
as larger: 4-5 x 3-4 μm.

In the field Callistosporium pinicola may also be mistaken for a species of Simocybe 
or Gymnopilus.

THE TAXONOMIC POSITION OF CALOCYBE JUNCICOLA

Next to Calocybe cerina sensu Arnolds & Becker, there is a second European 
species of Calocybe in which red staining of tissues in alkaline solutions has been re-
ported, viz. Calocybe juncicola (R. Heim) Sing. (M o r e n o  1995). Since this reaction 
is considered a characteristic feature of the genus Callistosporium, I have examined 
two collections of that species in order to re-evaluate its taxonomic position.

Calocybe juncicola (R. Heim) Sing. in Sydowia 15: 47. (‘1961’) 1962. – Fig. 3.
Tricholoma chrysenteron var. juncicola R. Heim in Treb. Mus. Cienc. nat. Bar-

celona 15: 101. 1934; Calocybe chrysenteron var. juncicola (R. Heim) G. Moreno in 
Cryptog. Mycol. 15: 240. 1995; Rugosomyces chrysenteron var. juncicolus (R. Heim) 
M. Bon in Doc. Mycol. 21 (82): 65. 1991.

Sel. icon. – Cetto, Funghi Vero 6: pl. 2414. 1989; Ludwig, Pilzkompendium 1: pl. 
19, fig. 7.9. 2000. 

Sel. descr. & figs. – Hauskn. & Zuccherelli in Boll. Gruppo micol. G. Bres. 1994: 
68-69, fig. 1 a-c. 1994; Ludwig, Pilzkompendium 1: 48. 2000; G. Moreno in Cryptog. 
Mycol. 15: 240-241, Fig. 1. 1995.

Pileus 10-30 mm, truncate campanulate to hemispherical at first, then convex 
to flattened, with or without umbo, often irregular, not hygrophanous, not striate, 
bright orange to golden yellow or brownish orange (e.g. K. & W. 4A8, 6A6, 6A7, 
6B8), uniformly coloured or with slightly paler margin, slightly aerenchymatic in 
places when young, glabrous and dry. Lamellae, L  30-40, l  3-7, emarginate to 
adnexed, moderately crowded to crowded, segmentiform, up to 4 mm broad, thin, 
vividly yellow, butter-yellow or ochre-yellow (K. & W. 4A5/6, 4B6), with concolor-



36 E. Arnolds 

ous, entire or eroded edge. Stipe 30-60 x 2-10 mm, subcylindrical or tapering towards 
base, often compressed or twisted, often slightly broader towards apex or base, solid, 
paler than the pileus, pale yellow, butter-yellow to brownish yellow (e.g. K. & W. 
4A4, 5, 6, 4B6), fibrillose striate lengthwise, appearing pruinose by aerenchymatic 
tissue. Context in pileus up to 4 mm thick, pale yellow. Smell and taste strongly fari-
naceous, taste slightly adstringent. Spore print unknown.

Spores (3.5-)4.5-6.5 x (2.5-)3.0-3.7(-4.0) μm, av. 5.0-5.4 x 3.2-3.3 μm, Q  1.3-1.9, 
Qav.  1.5-1.65, ellipsoid or ellipsoid-oblong, smooth, not amyloid, in water and am-
monia colourless. Basidia 25-36 x 5.0-8.0 μm, clavate, 4-spored or a few 2-spored, in 
part with few to fairly numerous small granules. Lamella edge fertile, but sometimes 
with scattered cylindrical basidiola, 18-23 x 4.0-5.0 μm. Pleurocystidia absent. Hy-
menophoral trama subregular, made up of rather narrow hyphae with cylindrical or 
slightly inflated elements, 2.0-14 μm broad. Pileipellis a poorly differentiated cutis, 
made up of repent, interwoven hyphae, 1.5-5.0 μm broad, in places with ascending 
to erect hyphae, with pale yellow intracellular pigment. Stipitipellis a cutis, made up 
of repent hyphae, 2.0-4.0 μm wide. Clamp-connections abundant in hymenium and 
trama.

Chemical reactions: Preparations of all tissues turning immediately reddish to 
violet in KOH 5% and ammonia 10%; macroscopic reactions on fresh basidiocarps 
unknown. Granules in basidia staining dark purple in acetocarmine with iron (si-
derophilous).

Habitat and distribution: Saprotrophic, usually fasciculate, on roots and litter in 
dense tussocks of Juncus spp. (e.g. Juncus maritimus, Juncus acutus) in open forests 
and dune slacks. Nov. Apparently very rare in southern Europe, known from Italy 
and Spain.

Fig. 3. Calocybe juncicola. A. Basidiocarps x 1; B. Spores x 2000; C. Basidia x 1000 (all from 
Noordeloos 20266).



 A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales) 37

Collections examined. – ITALY: Ravenna, Pineta San Vitale, Bardello, 6 Nov. 
2000, A. Hausknecht et al. (WU no. 20752); same dat. and loc., M.E. Noordeloos 
20266 (L). 

The reddish staining of tissues in alkaline solutions was only reported by M o r e n o 
et al. (1995), who mentioned a violet discoloration of the solution when making a 
preparation of the lamellae in KOH. The reaction was not mentioned in other des-
criptions (H a u s k n e c h t ,  Z u c c h e r e l l i  1994; C e t t o  1989; L u d w i g  2001) or in 
the descriptive notes accompanying the collections. The reddening in ammonia and 
KOH is very similar to that of Callistosporium pinicola, although slightly weaker. 
Nevertheless, Calocybe juncicola seems not to be related to the latter species in view 
of the absence of brown necropigment, the abundance of clamp-connections and 
the presence of truly siderophilous granules in the basidia, although these granules 
seem to be less numerous and striking than in other Calocybe species. Otherwise, 
C. juncicola differs from C. chrysenteron (incl. var. cerina) mainly in duller colours, 
striate stipe, larger and more elongate spores and specific habitat.

A second European species of Calocybe with violaceous red discoloration of tis-
sues in KOH is Rugososmyces pudicus M. Bon & Contu, collected in moist forest of 
Populus on Sardegna (C o n t u ,  B o n  2000). It has small basidiocarps with a pinkish 
brown to yellow-brown pileus up to 20 mm, differing mainly from C juncicola in im-
mediate reddening of the context on exposure to the air and broader spores (5.0-6.0 
x 3.5-4.5 μm) that are slightly rugulose, as was also demonstrated with SEM micros-
copy (C o n t u ,  O r t e g a  2001).

CALOCYBE OR RUGOSOMYCES?

The genus Calocybe was erected (but invalidly published) by K ü h n e r  (1938) 
in order to accommodate tricholomoid, white-spored agarics with basidia contain-
ing siderophilous granules in combination with vividly coloured basidiocarps. Donk 
(1962) validated this name and selected Calocybe georgii (L.) Kühner (  C. gambosa 
(Fr.: Fr.) Donk) as type species. Several authors have discussed the possible hetero-
geneity of this genus since. K a l a m e e s  (1992) has proposed the genus Tricholomella 
for Calocybe constricta (Fr.: Fr.) Sing., deviating from other species in presence of a 
partial veil, verrucose spores, large basidia and ecological preference for substrates 
with high ammonia content. This distinction was later confirmed by phylogenetic 
studies using molecular characters (H o f f s t e t t e r  et al. 2002; M o n c a l v o  et al. 
2002).

The genus Gerhardtia was created by B o n  (1994) for Calocybe borealis A. Riva 
(  Lyophyllum incarnatobrunneum Ew. Gerhardt), a species deviating in minutely 
verruculose spores and absence of clamp-connections. Since verruculose spores have 
also been observed in Rugosomyces (Calocybe) pudicus (C o n t u  and O r t e g a  2001), 
the status of this genus seems questionable. The phylogenetic relationships with the 
rest of Calocybe (and Lyophyllum) has not yet been studied.

Earlier R a i t h e l h u b e r  (1979) erected the genus Rugosomyces for the two spe-
cies of Calocybe with a cellular pileipellis (hymeniderm) instead of a pileipellis of 
slender hyphae (cutis): the type species Rugosomyces onychinus (Fr.) Raithelh. and 
R. fallax (Sacc.) M. Bon. This concept was adopted by e.g. H o r a k  (2005), but Bo n 
(1991) extended Rugosomyces to all small species with collybioid habit, a pileipellis 



38 E. Arnolds 

with more or less free ending hyphal tips, often with inflated cells, and mixed pari-
etal and intracellular pigments. Calocybe was restricted by B o n  (l.c.) to a few large, 
tricholomoid species with intracellular pigments. This concept was accepted by e.g. 
K a l a m e e s  (2004). However, molecular studies of the Lyophylleae have demon-
strated that Calocybe (with the exception of C. constricta, possibly also C. borealis; 
see above) forms a single clade and is probably of monophyletic origin (H o f f s t e t -
t e r  et al. 2002; M o n c a l v o  et al. 2002). Therefore there seems to be no good rea-
son to recognise Rugosomyces as a separate genus. 

In this connection I propose two new combinations:
Calocybe pudica (M. Bon & Contu) Arnolds comb. nov. 
Basionym: Rugososmyces pudicus M. Bon & Contu in Doc. Mycol. 29 (116): 35. 

2000.
Calocybe obscurata (P. Karst.) Arnolds comb. nov. 
Basionym: Tricholoma cerinum Pers. ssp. obscuratus P. Karst. in Meddeland. Soc. 

Fauna Fl. Fenn. 5: 43. 1879.

CALLISTOSPORIUM AND CALOCYBE

The genus Callistosporium is assigned to the Tricholomatoideae and not to the 
Lyophylloideae in view of the absence of siderophilous granules in the basidia. Also 
in phylogenetic analysis the genera seem not to be closely related to each other 
(M o n c a l v o  et al. 2002). However, it might be interesting to study the relationships 
between these groups more closely in view of the remarkable convergence between 
some species. Also the chemical composition of the necropigment in Callistosporium 
deserves attention.

Acknowledgements. This paper is dedicated with great pleasure to Professor Alina Skirgiełło, a living 
monument of vitality and a source of inspiration for all of us!
Thanks are due to Anton Hausknecht (Maissau, Austria), Markus Wilhelm (Allschwil, Switzerland) and 
the curators of the herbaria of the Botanical Museum in Copenhagen (C) and the National Herbarium in 
Leiden (L) for sending me valuable collections on loan. I am also indebted to Thomas Kuyper (Wagenin
gen, The Netherlands) for some critical remarks on the manuscript of this paper.

REFERENCES

A r n o l d s  E., B e c k e r  A. 1993. Over Calocybe cerina en enkele verwanten. Coolia 36: 69 78.
B o n  M. 1976. Le genre Callistosporium Singer. Doc. mycol. 6 (22/23): 279 286.
B o n  M. 1991. Les genres Echinoderma (Locq. ex Bon) st. nov. et Rugososmyces Raithelhuber ss. lato. 

Doc. mycol. 21 (82): 61 66.
B o n  M. 1991. Flore mycologique d’ Europe 2. Tricholomataceae 1. Doc. mycol., Mém. hors Sér. 2: 

1 149.
B o n  M. 1994. Deux Lyophylloideae interessantes et le genre Gerhardtia st. et nom. nov. Doc. mycol. 

24(93): 65 68.
B o n  M. 1999. Flore mycologique d’ Europe 5. Les Collybio Marasmïoïdes et ressemblants. Doc. mycol., 

Mém. hors Sér. 5: 1 171.
C e t t o  B. 1989. Funghi Vero 6. Trento.
C o n t u  M., B o n  M. 2000. Une nouvelle espèce de Rugosomyces ‘rougissant’. Doc. Mycol. 29(116) : 

35 36.
C o n t u  M., O r t e g a  A. 2001. Studi sulle Lyophyllaceae della Sardegna.V. Morfologia sporale di Ru

gososmyces pudicus Bon & Contu ed implicazioni sulla sua posizione sistematica. Bol. Soc. micol. 
Madrid 26 : 173 176.



 A confusing duo: Calocybe cerina and Callistosporium pinicola (Agaricales) 39

D o n k  M.A. 1962. The generic names proposed for Agaricaceae. Beih. Nova Hedwigia 5 : 1 320.
F r i e s  E.M. 1821. Systema mycologicum 1. Gryphiswaldiae.
H a u s k n e c h t  A., Z u c c h e r e l l i  A. 1994. Ritrovamenti interessanti dal Ravennate. 3e Parte. Boll. 

Gruppo micol. G. Bresadola 1994: 67 95.
H o f s t e t t e r  V., C l é m e n c o n  H., V i l g a l y s  R., M o n c a l v o  J M. 2002. Phylogenetic analyses of 

the Lyophylleae (Agaricales, Basidiomycota) based on nuclear and mitochondrial rDNA sequences. 
Mycol. Res. 106: 1043 1059.

H o r a k  E. 2005. Röhrlinge und Blätterpilze in Europa. Elsevier, München.
K a l a m e e s  K. 1992. Tricholomella, a new genus, with the distribution data of Tricholomella constrictum 

comb. nov. in East Europe and Asia. Persoonia 14: 445 447.
K a l a m e e s  K. 1995. On Rugosomyces fallax and allied species (Tricholomatales). Doc. Mycol. 25 (98

100): 229 236.
K a l a m e e s  K. 2004. Palearctic Lyophyllaceae (Tricholomatales) in northern and eastern Europe and 

Asia. Scripta Mycologica 18. Tartu.
K o n r a d  P., M a u b l a n c  A. 1927. Icones selectae Fungorum 3. Paris.
K ü h n e r  R. 1938. Utilisation du carmin acétique dans la classification des agarics leucosporés. Bull. 

mens. Soc. Linn. Lyon 7: 204 212.
L a n g e  J.E. 1935. Flora Agaricina Danica 1. Copenhagen.
L u d w i g  E. 2000. Pilzkompendium 1, Abbildungen. IHW Verlag, Eching.
L u d w i g  E. 2001. Pilzkompendium 1, Beschreibungen. IHW Verlag, Eching.
M a i r e  R. 1937. Fungi Catalaunici. Contribution à l’ étude de la Flore Mycologique de la Catalogne. 

Publ. Inst. bot. Barcelona 3 (4): 1 128.
M o n c a l v o  J M, V i l g a l y s  R. ,  R e d h e a d  S . A . ,  J o h n s o n  J . E . ,  J a m e s  T. Y . ,  A i m e  M . C . , 

H o f s t e t t e r  V. ,  Ve r d u i n  S . J . W. ,  L a r s s o n  E . ,  B a r o n i  T. J . ,  T h o r n  R . G . ,  J a c o b s
s o n  S . ,  C l é m e n c o n  H . ,  M i l l e r  Jr., O.K. 2002. One hundred and seventeen clades of euagar
ics. Mol. Phylogenet. Evol. 23: 357 400.

M o r e n o  G., A r e n a l  F., G o n z á l e z  V. 1995. Algunos Agaricales de Espana. Crypt. Mycol. 15: 239
254.

N o o r d e l o o s  M.E. 1995. Callistosporium. In: Bas, C., Kuyper, Th.W., Noordeloos, M.E., Vellinga E.C. 
(eds). Flora agaricina neerlandica 3: 104. A.A. Balkema, Rotterdam, Brookfield.

P e r s o o n  C.H. 1801. Synopsis methodica fungorum. Gottingae.
R a i t h e l h u b e r  J. 1979. Calocybe Kühner  Ein Sammelgattung? Metrodiana 8: 9 10.
R e d h e a d  S.A. 1982. The systematics of Callistosporium luteo olivaceum. Sydowia 35: 223 235.
W i l h e l m  M. 2003. Callistosporium luteoolivaceum var. minor M. Bon, Kleinsporiger Scheinrübling. 

Schweiz. Z. Pilzk. 2003: 62 66.

Podwójna niejasność: Calocybe cerina i Callistosporium pinicola (Agaricales)

S t r e s z c z e n i e

Nazwa Calocybe cerina była ostatnio stosowana dla dwóch różnych gatunków grzybów aga
rikoidalnych. Na podstawie analizy materiałów z różnych krajów europejskich autor dochodzi 
do wniosku, że prawdziwie gatunek Calocybe cerina jest bardzo bliski C. chrysenteron i propo
nuje nową kombinację Calocybe chrysenteron var. cerina podając opis nowego taksonu. Więk
szość badanych kolekcji określonych jako C. cerina, okazała się taksonem, który Bon opisał 
prowizorycznie jako Callistosporium luteooliraceum fo. minor. W nieniejszym artykule został 
on opisany jako nowy gatunek: Callistosporium pinicola. Opis pozwala wyraźnie odróżnić inny 
gatunek Calocybe juncicola. Równocześnie dyskutowany jest status rodzajów Rugosomyces i 
Calocybe. Podane są też nowe kombinacje Calocybe obscurata i Calocybe pudica.




		2014-01-01T11:43:19+0100
	Polish Botanical Society