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Type studies in North American species of Bolbitiaceae belonging to the genera Conocybe and Pholiotina



56 taxa described in the genera Agaricus, Conocybe, Galera, Galerella, Galerula, Gastrocybe or Pholiolina, which were considered to have affinities to Bolbitiaceae, are investigated and their taxonomic status is discussed. The new combinations Agrocybe ludoviciana, Conocybe humicola, Conocybe pulchra, Pholiolina flava and Pholiolina plumbeitincta are proposed Microscopical drawings of all investigated type specimens are given.
Z. Pilzk. 13 (2004) 153
Type studies in North American species of
belonging to the
and Pholiotina
Maissau, Austria
r Botanik der Universität Wien
Wien. Austria
Key words: Fungi, Agaricales, Holbiliaceae. Agrocybe. Conocybe. Cialerina. l'holiolina. - Taxon-
omy, type studies, new combinations. Mycoflora of North America.
Abstract: 56 taxa described in the genera Agancus, Conocybe, (ialera, Galerella, (jalerula. (.iastro-
cvhe or l'holiolina. which were considered to have affinities to Holbmaceae, are investigated and their
taxonomic status is discussed. The new combinations Agrocybe ludoviciana. ( 'onocybe hunucola,
( onocybe pulchra, l'holiolina /lava and l'holiolina plunihemncla are proposed Microscopical draw-
ings of all investigated type specimens are given.
Zusammenfassung: 56 aus Nord- und Mittelamerika in den Gattungen Agaricus. Conocybe. (la/era.
Cialerella, (ialerula. Crasirocvbe oder l'holiolina beschriebene Taxa, die mit den Holbitiaceae in
Zusammenhang gebracht worden sind, werden behandelt und ihre taxonomische Zuordnung diskutiert
Die Neukombinationen Agrocybe ludoviciana. ('onocybe humicola, ('onocybe pulchra. l'holiolina
und l'holiolina plumbeilmcla werden vorgeschlagen Mikroskopische Zeichnungen zu allen unter-
Typusbelegen werden angefertigt.
The present study comprises taxa described from material originating from the USA.
Mexico and Greenland; as many North American floras also include specimens from
Cuba or Puerto Rico, types from the Caribbean were also included.
The descriptions of the present study are not always identical with the original di-
agnoses, as all microscopical measurements were excluded If the diagnoses are short
or uninformative, they were supplemented with details from the English descriptions
or even replaced with the latter. The descriptions are therefore not identical with the
original protologue but include all available information on the microscopical features
For a better orientation, the taxon name supposed to be actually valid according to
our investigations is given in bold.
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154 A. HAUSKNECHT &al.: Type studies in North American Holbitiaceae
The microscopical measurements are exclusively from investigations of the type
specimens by the first author Supplements from other sources are especially anno-
Materials and methods
Microscopical investigations were performed with an Olympus BH-2 microscope, the drawings with a
drawing tube at magnifications of
(spores) or
(other elements), and scaled down ac-
cordingly for the publication The microscopic slides were prepared in L4 Congo red, NH
OH or
KOH. Spore colour was recorded from slides in KOH (sometimes in NH4OH when no differences
were observed between the latter and the former), with a fully open aperture diaphragm and using a
blue filter. Ammonia reaction was performed using a commercial or slightly diluted solution; the
slides were investigated for the formation of spicular crystals after 15-20 minutes and after ca 12
Spore measurements are recorded from ten spores: usually eight spores were selected by
chance, and the upper and lower limits were estimated by choosing one particularly large and one
particularly small spore. Means of spore sizes were calculated from at least ten different spores. In
lenticular spores, five were measured each in side and front view, respectively, but care was taken that
spore sizes did not differ strongly In lenticular spores, the length-breadth ratio was calculated from
measurements front view and not from side view.
Enlargement of
microscopical figures: Spores x 2000, all others x 800.
List of treated taxa
alachuana (¥\g I a-e)
Galerula alachuana MURRILL 1940, Bull. Torrey Bot. Club 67: 229
Galera alachuana (MURRILL) MURRILL 1940. Bull. Torrey Bot Club 67: 235
Conocybe alachuana (MURRILL) HESLFR in WATLING & GREGORY 1981, Bibl
Mycologica 82: 89
Original description:
Pileus conic to convex, gregarious, 5-7 mm broad; surface dry, glabrous, striate, avel-
laneous-isabelline. isabelline on the small umbo, margin straight, entire; lamellae ad-
ventncose, broad, inserted, medium distant, entire, fulvous, with white lamellar
stipe tapering upwards, smooth, glabrous, stramineous, about 6 cm long and
1 mm or less thick.
Microscopical characters:
Spores: 11-13.5 x 7.5-9 x 6-7.5 urn, mean 11.9 x 8.0 x 7.0 um, Q =
larly ellipsoidal to ellipsoidal-subcylindrical in front view, slightly lentiform and
somewhat angular, with a thick wall and a large germ-pore, red brown in KOH.
Basidia: 4-spored, 17-21 x 10-12.5 urn.
Clamp connections: not observed.
Cheilocystidia: lecythiform, 15-21 x 6-8 urn, with a capitulum 2.5-4 um in
Stipe covering: roundish to capilliform, rather collapsed; no lecythiform cau-
locystidia present.
Pileipellis: hymeniform. consisting of roundish-stipitate elements up to 50 x 28
jim; pileocystidia not observed.
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Fig. I a-e. Clalenila alachuana (type), a Spores, h basidia c cheilocystidia. d stipe covering, e ele-
ments of
/-/ ('onocvhe anmiophila (type). /Spores, g basidia h cheilocystidia / stipe cov-
ering. / elements of pileipellis.
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156 A. HAUSKNECHT & al.: Type studies in North American Holbitiaceae
Specimen examined: USA: Florida. Alachua County. Gainesville, Planera Hammock, under
deciduous trees. 21. 10. 1938. leg. W. A. MURRILL (FLAS, holotype).
The type consists of fragments of two fruitbodies and is moderately preserved; es-
pecially the stipe coverings are rather collapsed.
Conocybe alachuana is a member of sect Pilosellae and is close to Conocybe ve-
lutipes (VELEN ) HAUSK.N & SVRCEK However, the latter usually has much larger,
robust fruitbodies with a darker pileus and is especially different by the shape of the
spores. In both species the spores are lentiform. but the spores of Conocybe velutipes
are regularly ellipsoidal in side and front view, whereas those of Conocybe alachuana
are often subcylindric in front view and in side view flattened to slightly lemon-shaped. As
such a spore shape was never observed in the numerous European collections of Co-
nocybe velutipes, we consider Conocybe alachuana as a well-distinct species.
Conocybe fuscimarginata (MURRILL) SINGER has spores of similar size, which are
never lentiform, and the fruitbodies are usually larger and have a paler, non-striate pi-
The spores of Conocybe lenticulospora WATL1NG are lentiform. but slightly
smaller and paler in K.OH; in addition, they are never subcylindrical in side view. In
addition, Conocybe fuscimarginata and C. lenticulospora are usually found on dung or
heavily manured soil.
ammophila (F
ig. 1 f-j)
Conocybe ammophila M. LANGE 1957, Medd Grönland 148 9
Original description:
cm latus, convexus, deinde planus. pallide ochraceus. nudus, camosus,
lamellae latae, subliberae, ochraceae; stipes cylindricus, basi inflatus. pallidus, carina-
Microscopical characters:
Spores: 9.5-11.5 x 6.5-7.5 x 6.0-7.0 um, mean 10.7x6.9x6.3 um, Q =
lipsoidal, often slightly lentiform and rarely angular with ca 0.5 urn thick wall and a
germ-pore up to 1.5 urn in diam.. reddish brown in KOH.
Basid la: 4-spored, 22-26 x 8 5-11 urn.
Ammonia reaction: negative even after 12 hours.
Clamp connections: not observed.
Cheilocystidia: 17-27 x 7-11 urn. with capitulum 5-7 urn in diam
Stipe covering: consisting almost exclusively of lecythiform elements, which
are similar in shape to cheilocystidia. but up to 30 x 13 um. with capitulum 4-8 ^m in
diam.; in-between some sparse non-lecythiform caulocystidia and hairs, especially in
the middle of the stipe.
Pileipellis: hymeniform, consisting of round ish-stipitate elements up to 37 x 17
iim in diam., which are almost hyaline, at the base encrusted yellowish.
Specimen examined: Greenland: Sondre Stromfjorden. on sand bank along creek close to river,
8. 1946, leg. M. LANGE (C. holotype)
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Österr Z Pilzk. 13(2004) 157
Fig. 2 a-d (ialera hulbifera (part of
a Spores, h basidia. c isolated capitulum of cheilocys-
ndium. d stipe covering, e-i. (ialemla caespuosa (holotype). e Exsiccate (x 0.8), / spores, g basidia. h
cheilocystidia. ; stipe covering.
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158 A. HAUSKNECHT &al.: Type studies in North American Holbihaceae
The specimen consists of numerous well-preserved, partly broken fruitbodies. the
investigation of which is only hampered by the attached sand. WATLING annotated in
his previous investigation of the specimen "sect. Conocybe with capitate caulocystidia;
good species in stirps Leucopoda which should be looked out for".
The combination fruitbodies Agrocybe-Wke in habitus, lecythiform caulocystidia
with large capitulum and slightly lentiform spores does not fit any other described spe-
cies The spore dimensions and spore colour in K.OH as well as the cystidia with large
capitulum are somewhat reminiscent of Conocybejuniana (VELEN.) HAUSKN & SVRCEK,
but clearly differ by the stout, compact habitus, the whitish stipe and the lentiform,
partly slightly angular spores.
Galera angusticeps PECK 1897, Bull. Torrey Club 24: 143
Galerula angusticeps (PECK) MURRILL 1917. North Amer. Flora 10: 168
Galeropsis angusticeps (PECK) SINGER 1962. Sydowia 15 83
Agrocybe angusticeps (PECK) WATLING 1981, Bibl Mycologica 82: 26
Original description:
Pileus thin, narrowly and irregularly conical or subcylindrical, obtuse acute or abruptly
acuminate at the apex, even, glabrous, viscid and dark ochraceous when young and
moist, nearly white when old and dry, the margin somewhat incurved and appressed to
the stem; lamellae close, narrow, adnate, somewhat white-margined, more or less anasto-
mosing, brownish-ferruginous when mature, stem slender, glabrous, hollow, equal or
slightly thickened at the base, whitish or tinged with yellow, shining when dry. Pileus
8-15 lines long, 4-6 lines wide, stem 1.5-3 in long, 1-15 lines thick.
WATLING (in WATLING & GREGORY 1981) combined the species into Agrocybe.
The original description and the combination of SINGER (1962) into Galeropsis show
that it is a gasteroid species which has nothing to do with Conocybe or Pholiotina.
Therefore, we have not examined the type once again.
see procera
Galera besseyi PECK 1909, Bull. New York State Mus. Nat Hist 131: 35
Galerula besseyi (PECK) MURRJLL 1917. North American Flora 10: 163
Conocybe besseyi (PECK) R HEIM 1931, Le genre lnocybe: 65
Cyttarophyllum besseyi (PECK) SINGER 1936, Ann Mycol 34 344
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Öaerr. Z. Pilzk. 13(2004) 159
Original description:
Pileus thin, ovate or oval, rarely subglobose, obtuse, glabrous, never expanding,
isabelline or pale dingy ochraceous. the margin abruptly contracted and closely em-
bracing the stem; lamellae thin, close, ascending, adnate, ferruginous brown, stem
slender, slightly flexuous. hollow, glabrous, even or slightly stnate. colored like the
pileus. Pileus 2.5-6 lines long, 2-5 lines broad; stem 1-2 inches long. 5-1 line thick.
Sandy soil. Garden of the Gods, F.I Paso Co., Colorado. August 1908. C. E. BES-
The description and especially the attached colour plate clearly show that Galera
besseyi is not a member of Conocybe or Pholiotma SINGER (1936) placed the species
into Cyttarophyllum, which was later placed by him (SINGER 1986: 844) into the se-
cotioid family Galeropsidaceae. which, however, has close affinities to the Bolbilia-
ceae Therefore, we have not examined the type material.
bulbifera (Fig 2 a-d)
Galera bulbifera K.AUFFMAN 1918. Agahcaceae of Michigan: 496
Conocybe bulbifera (KAUFFMAN) ROMAGN. 1942, Bull Soc Mycol France 58 147
Original description:
Pileus 0.5-2.5 cm broad, oval-campanulate, obtuse, ferruginous-cinnamon when moist,
hygrophanous. ochraceous and atomate when dry. rivulose-reticulate. Gills ascending-
adnate. narrow, sublinear, close to crowded, ferruginous-cinnamon, sprinkled by fer-
ruginous spores. Stem 6-15 cm long, 1.5-3 mm thick, strict when moist, equal above
the bulbous base, pale ferruginous, hollow, glabrous-shining when dry, sometimes
faintly striatulate Odor none.
On horse dung, dung-hills in mixed woods.
Variable in size; solitary specimens attain the large size, while a patch of them is
apt to be composed of smaller sizes It has the appearance, in the large condition, of G.
ovalis. but differs by the narrow gills etc It is well marked by the gills, the bulblet at
base of stem, and the spores. The whole plant is ferruginous-cinnamon when moist,
and in large plants the pileus is finely rugose-reticulate.
Microscopical characters:
Spores: 13.5-16 x
x 7-8.5 urn. mean 15.0 x 94 x 7.8 um, Q =
soidal in front view, slenderly ellipsoidal to faintly bean-shaped in side view, distinctly
lentiform, with a wall up to
(im thick and a broad germ-pore up to 2 urn in diam., rusty-
brown in K.OH
Basidia: 4-spored. 22-26 x 11-13.5 urn.
Clamp connections: not observed.
Cheilocystid ia: entirely collapsed, only one capitulum with a diam. of 4 urn
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160 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae
Stipe covering: collapsed; besides hairs and non-lecythiform elements dis-
tinctly lecythiform caulocystidia present. 12-19 x 7-8 urn, with capitulum 3-4 (im in
Pileipellis: collapsed, but evidently a hymeniderm composed of roundish cells.
Specimen examined: USA: Michigan. New Richmond, on horse dung. 23. 9. 1912. leg. C. H.
KAUFFMAN (MICH, portion of holotype).
The specimen examined consisted of two stipe fragments and of approximately
one fifth of a pileus of reddish-brown colour The date given on the type specimen
does not correspond to that in WATLING& GREGORY (1981), which is given as 19 9
The results of our microscopical investigations confirm WATLING (1983) who con-
sidered the type of Galera bulbifera to be a member of sect Mixtae. and who therefore
described the new species Conocybe elegans for European collections previously de-
termined as Conocybe bulbifera, as these clearly belong to sect. Pilosellae.
The dung-inhabiting Conocybe bulbifera is closely related with C. pubescens (GlL-
LET) KÜHNER, but has smaller, more thick-walled and conspicuously lentiform spores
(see HAUSKNECHT 2003). Conocybe smgehana HAUSKN. with a large, tuberous base
of the stipe and similarly sized spores as C. bulbifera has no lecythiform elements on
the stipe, and its spores are not lentiform and somewhat thinner-walled Consequently
we consider Conocybe bulbifera as a distinct species well separated from European
members of sect Mixtae.
caespitosa (Fig 2 e-i)
Galerula caespitosa MURRILL 1941, Mycologia 33: 445
Galera caespitosa (Ml RRH.l.) Ml'RRlLl. 1941, Mycologia 33 448
Conocybe caespitosa (MlRRlLL) WATLING 1977, Kew Bull 31 593
Original description:
Pileus hemispheric, not fully expanding, strictly caespitose, 1.5 cm Broad, surface
dry, pulverulent, glistening like mica, long-striate. uniformly pale rosy-isabelline or
avellaneous, margin entire, even, fibrillose. incurved; context white, unchanging; la-
mellae adnatae, rather broad behind, medium distant, inserted, fimbriate on the edges,
white, whitish when dry.
Microscopical characters:
Spores: 10.5-15.5x6.5-9 urn. mean 12.3-13 Ox 7.7-7.9 urn. Q=
to subcylindrical-ellipsoidal, not lentiform. thick-walled with large germ-pore, orange-
yellow in KOH
Basidia: 2-3-4-spored. 18-27 x 12-15 urn
Clamp connections: not observed
Cheilocystidia: lecythiform. 15-22 x 7-10 urn. with capitulum 3-4 5 urn in
diam.. often with a long neck.
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Österr. Z. Pilzk. 13(2004)
Fig. 3 a-e. (jalervla canalipes (type), a Spores, b basidia. c cheilocystidia, d stipe covering, e pileipel-
(ialemla capillanpes (type). /Spores, g basidia. h cheilocystidia ' stipe covering, j-m. Agari-
Spores, k basidia, / stipe covenng, m collapsed elements of pileipellis.
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Üffr A. HAUSKNECHT & al.: Type studies in North American Bolbiliaceae
Stipe covering: composed of non-lecythiform elements and hairs, no capitate
caulocystidia found.
Pileipellis hymeniform, composed of roundish elements (20-32 x 25-47 urn),
in-between with hairs ca 2.5-3 um thick.
Specimens examined: USA: Florida, Gainesville, under evergreen oaks, 29. 5. 1938, leg. W. A.
MURRJLL (FLAS. holotype); - - on sawdust, 6. 1 1938, leg W. A. MURRJLL (FLAS. designated as
The holotype consists of six fruitbodies, of which three were fasciculately united,
very well preserved
SINGER & HAUSKNECHT (1988: 120) supposed a close relationship with Conocybe
leucopus KÜHNER & WATLING; however, a comparison of the microscopical features
of both taxa showed a distinctly different stipe surface in Conocybe caespitosa, hence
the former is a member of sect Conocybe whereas the latter a member of sect. Pilosel-
lae Within sect. Pilosellae there are several very rare or insufficiently known taxa
with similar spore characteristics [e.g., Conocybe crocospora (BERK. & M. A. CURITS)
KUHNER inval. from North America, Conocybe peroxydata (BERK ) REID from South
Africa, and Conocybe munnacea WATLING from Europe], but the macroscopical fea-
especially the combination of striate, pale rose-isabelline pileus and the fascicu-
late growth distinguish Conocybe caespitosa from all other species Although never
collected again, we consider Conocybe caespitosa to be a well-distinct species
canalipes (Fig 3 a-e)
Galerula canalipes MURRILL 1942, Lloydia 5: 147
Original description:
Pileus narrowly to broadly conic, gregarious, 2-3 cm broad; surface smooth, glabrous,
hygrophanous, isabelline. dark-isabelline when young and remaining so on the umbo;
margin entire, even, straight when young; context very thin, white, mild, odorless; la-
mellae adnexed. ventncose, crowded, inserted, medium broad, entire, white to isabel-
line and finally fulvous, stipe tapering upward from an onion-shaped base, slightly
radicate, cartilaginous, stuffed, pallid to isabelline. pruinose to subglabrous. canalicu-
late for its entire length, 3-4 x 0.2-0.3 cm, bulb 5 mm in diameter.
Microscopical characters:
Spores: 8-9.5 x 6-6.5 x 5-5.5 urn, in front view almost mitra-shaped to almost hex-
agonal, in side view more or less ellipsoidal, distinctly lentiform. with double wall and
a germ-pore up to 1.2 urn in diam., orange-yellow in KOH.
Basidia: 4-spored, 18-25 x
Clamp connections: not observed
Cheilocystidia: lecythiform, 12-17 x 6-9.5 urn, with capitulum 3-4 5 urn in
Stipe covering: collapsed, evidently composed only of lecythiform elements,
which are similar to cheilocystidia, capitulum often absent.
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Österr Z Pilzk 13(2004) . 163
Pileipellis: hymeniform. composed of round ish-stipitate elements. 21-32 x 10-
22 um; pileocystidia not observed.
Specimen examined: USA: Florida. Magnesia Springs, on cow dung in low deciduous wood, 11.
1939, leg. W. A. MURRlLL (FLAS, holotype).
The type consists of fragments of presumably only one moderately preserved fruit-
body; the base of the stipe is tuberous, but shows no marked root.
The microscopical characters of Galerula canalipes perfectly agree with those of
Conocybe antipus (LASCH) FAYOD AS MURRlLL (1942) described the stipe as "slightly
radicate" and as also the habitat corresponds perfectly, there is no doubt that Galerula
canalipes is a later synonym of Conocybe antipus, which is a rare but widely distrib-
uted species in Europe and North America
capillaries (Fig 3 f-i)
Galera capillaripes PECK 1899, Bull Torrey Bot Club 26: 66
Galerula capillaripes (PECK) MURRlLL 1917, North American Flora 10: 163
Conocybe capillaripes (PECK) WATL1NG 1977, Kew Bull 31 593
Original description:
Pileus subcampanulate, obtuse, a little broader than high, even, glabrous, hygropha-
faintly striatulate and pale ferruginous when moist, paler or buff color when dry;
lamellae rather broad, distant, adnate, pale ferruginous; stem very slender, flexuous,
glabrous, colored like the pileus. Pileus 4-6 mm broad, stem 2-3 cm long, less than
1 mm thick.
Microscopical characters:
Spores: 12 5-15 x 7.5-9.5 urn, mean 14.1 x 8.6 urn, Q = 1 5-1.8, ellipsoidal, not len-
tiform. with thick wall and large, truncate germ-pore, rusty brown in K.OH (only ma-
ture spores recorded as many spores were immature).
Basidia: 2-spored. 18-22 x 10-13.5 urn.
Clamp connections: not observed.
Cheilocystidia: lecythiform, 12-18 x 6.5-8 urn, with capitulum 2.5-3.5 urn in
Stipe covering: composed of roundish, lageniform to capilliform elements, at
the stipe apex with scarce lecythiform caulocystidia similar in shape to cheilocystidia
but larger with capitulum up to 4 urn.
Pileipellis: hymeniform, composed of sphaeropedunculate to pyriform ele-
ments. Pileocystidia not observed.
Specimen examined: USA: Ohio, lawns and grassy places. May and June, leg. C. G. LLOYD
The specimen consists of two long stipes and several small pileus fragments in
poor condition. This was the reason why we examined the pileipellis only from a min-
ute fragment and why we did not search for pileocystidia further on.
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164 A. HAUSKNECHT & al.: Type studies in North American Holbitiaceae
PECK (1899) did not give a microscopical description of the stipe covering, and
also HESLER (in his unpublished records) did not see lecythiform caulocystidia. In the
original description, PECK (1899) supposed that Conocybe capillaripes could be mis-
taken for a "dwarf form" of Galera tenera, however, the structure of the stipe covering
is different Conocybe capillaripes is a member of sect. Pilosellae, with microscopical
characters similar to those of Conocybe siliginea (FR
FR.) KÜHNER or Conocybe rickenii
(JUL. SCHAFF.) KÜHNER Considering the habitus, the species could be mistaken for a
notably delicate form of Conocybe siliginea, however, the spores are smaller, the la-
mellae more distant and the pileus is slightly striate. In case additional records bring
light into the variability of macroscopical features and spore size, it would not be sur-
prising if Conocybe capillaripes were a later synonym of Conocybe siliginea.
crispa (Fig 4 a-e)
Galera crispa LONGYEAR 1899, Bot. Gazette 28: 272
Conocybe crispa (LONGYEAR) SINGER 1951. Lilloa 22: 485
Conocybe albipes (OTTH) HAUSKN. var. crispa (LONGYEAR) HAUSKN. 1998, Österr.
Pilzk. 7: 106
Original description:
cm broad, membranaceous, persistently conico-campanulate, subacute,
uneven and somewhat rivulose, ochraceous-brown on disk, lighter toward the margin
which becomes crenulate and upturned in older specimens, slightly pruinose at first,
rugulose and a little paler when dry; lamellae slightly adnexed, not crowded, rather
narrow, interspaced with anastomosing veins, much crisped, at first nearly white, then
becoming ferruginous from the spores; stem 7-10 cm long, 0 1-0.3 cm thick, tapering
from the somewhat bulbous base, yellowish wh\te, pruinose at base, hollow, fragile
Microscopical characters:
Spores: 11.5-16 x 7 5-9.5 urn, mean 13.0-13.6 urn, ellipsoidal, in front view partly
slightly irregular or angular, not or only faintly lentiform. with 0.5-0.8 urn thick wall
and a large germ-pore up to 2,5 u.m in diam., in KOH reddish brown.
Basidia: 4-spored, 21-37 x 10-17.5 um.
Clamp connections: not observed.
Cheilocystidia: lecythiform. 23-30 x 9.5-14 urn, with capitulum 4-6 urn in
Pavement cells: present
Stipe covering: composed of cylindrical, clavate or fusiform elements (up to
35 x 13 urn), at the stipe apex also some lecythiform caulocystidia similar to cheilo-
cystidia present.
Pileipellis: hymeniform, composed of roundish-stipitate elements.
Specimens examined: USA: Plants of Michigan. Michigan State College [according to BESSEY
(1944) the type locality]. 29. 6. 1948. leg. & det. E. A. BESSEY (MSC); - - near Library Building, 30.
6. 1953, leg. & det. E. A. BESSEY (MSC).
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Österr. Z. Pilzk. 13(2004)
Fig. 4 a-e. (iaiera cnspa (from type locality) a Spores, h basidia. c cheilocystidia. J stipe covering
from the middle of
e stipe covenng from the stipe apex. /-/ (ialerula cnspella (type). /Spores, g
basidia. h cheilocystidia,; stipe covering.
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166 A. HAUSKNECHT & al.: Type studies in North American Bolbmaceae
Especially the collection from 30. 6. 1953 is in good condition, at the pileus the
conspicuous wavy-curly lamellae are clearly visible The microscopical analysis and
also the commentary of the following paragraph are a translation from HAUSKNECHT
As already mentioned by WATLING (1992: 93), the type material of Galera crispa
is lost, therefore. LONGYEARs collection 119533 was selected by him as neotype We
have never seen this collection, but two other collections of the type locality (see above).
These fully agree in all essential features with the description given by WATLING
(pers comm.) for the neotype specimen, except for the basidia which carry four insted
of two (to three) sterigmata
The strong reasons given by HAUSKNECHT (1998) to propose that Galera crispa is
only a variety of Conocybe albipes have meanwhile been confirmed by the DNA analyses
of HALLEN & al (2003) As long as there are no further molecular studies on North
American and European material, it is therefore proposed that North American collec-
tions which have been determined as Conocybe lactea (LANGE) METROD should bear
the name Conocybe albipes. and North American collections of Conocybe crispa should
be placed into Conocvbe albipes var crispa However, meanwhile it has been demon-
strated that European material described by HAUSKNECHT (1998) as var. pseudochspa
merits species rank; the new combination has been recently published as Conocybe
pseudocnspa (HAUSKN.) ARNOLDS by ARNOLDS (2003).
(¥\g 4 f-i)
Galemla crispella MURRILL 1942, Lloydia5: 148
Conocybe crispella (MlRRlLL) SINGER 1950, Sydowia 4 132
Original description:
Pileus convex to subexpanded, never conic or plane, somewhat gregarious, 2-2.5 cm
broad; surface dry. pallid, finely striate to the small isabelline disk, margin entire;
context membranaceous. white, unchanging, mild, odorless; lamellae adnexed. roun-
ded behind, narrow, close, inserted, entire, pallid
Microscopical characters:
Spores: 11 5-13 x 8-9 x 7-8 urn, mean 12.4 x 8.3 x 7.5 urn. Q =
45-1 7, ellipsoidal,
slightly lentiform. with ca 0.3-0.5 urn thick wall and a germ-pore up to 2 urn in diam..
reddish brown in K.OH.
Basidia: 4-spored, 21-28 x 10-13.5 urn. According to HESLER (unpublished
notes) 24-28x9-11 urn
Clamp connections: not found.
Cheilocystid ia: mostly collapsed; some lecythiform elements with venter up
to 8 (im wide and a capitulum 2-3 7 wide. HESLER (unpubl.) found cheilocystidia with
16-22 x 5-6 urn.
©Österreichische Mykologische Gesellschaft, Austria, download unter
Österr. Z. Pilzk. 13(2004)
Fig. 5 a-e. (ialerula cryptocysiis (type), a Spores, b basidia, c cheilocystidia, d stipe covering, e ele-
ments of pileipellis. /-/. (ialerula curia (type). /Spores, g basidia h cheilocystidia. / stipe covering,
©Österreichische Mykologische Gesellschaft, Austria, download unter
& al.:
Type studies
North American Bolbiliaceae
Stipe covering: only few hairs and non-lecythiform elements present, the stipe
surface mostly collapsed. HESLER (unpubl.) recorded "stipe with some pin-shaped
cystidia immixed", we could not find such elements even at the uppermost stipe apex.
Pi lei pel
is: hymeniform, composed of roundish-pyriform. mostly collapsed
Specimen examined: USA: Florida, Alachua County, Gainesville, on grass on open lawn, 5. 8.
leg. W. A.
(holotype, FLAS).
The type consists of numerous small fragments of several pilei and the stipes in
poor condition
Conocybe chspella is a distinct member of sect. Candidae widely distributed espe-
cially in the tropics Besides collections from the Pacific region and from flower pots
and greenhouses in Europe (HAUSKNECHT 1997), the first author examined collections
from Asia (Singapur), Africa (La Reunion, Mauritius, Seychelles) und Australasia
(Fidschi. Vanuatu).
crocosporus (Fig
Agaricus crocosporus BERK & M. A CURTIS 1853, Ann Mag Nat Hist II, 12: 421
Galera crocospora (BERK & M A CURTIS) SACC , Sylloge Fungorum 5 866
Galerula crocospora (BERK. & M. A. CURTIS) MURRILL 1917, North American Flora
Conocybe crocospora
& M A CURTIS) KUHNER 1935, Le genre Galera 137,
nom inval (no basionym cited)
Pileus 2.5 cm broad or more, conic to convex, becoming depressed, sordid-brown, vi-
scid, margin sulcate. Context thin Lamellae adnate, pale-ferruginous, becoming saf-
fron-yellow, numerous. Stipe 8-10 cm long, slender, white, silky-shining, fistulöse,
base thickened
Spores: 11
x 80-10.0 urn, mean 13.0 x 8.5 urn, Q =
4-1.6, regularly ellip-
soidal, not lenticular, with thick wall and germ-pore up to 1.5 urn in diam., orange
brown to dark brown in KOH
B a s i
a: 4-spored, ca 16-19 x 12-14 urn.
Clamp connections: not found.
Cheilocystidia: completely collapsed; the specimen is heavily molded by a
contaminating fungus, the lamellar edge completely spoilt. HESLER wrote in his un-
published notes "cheilocystidia 15-23 x 5-8 urn, flask-shaped, with a short neck and
abrupt capitellum, inconspicuous and few (found only after seve