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

January 2004

January 2004
13:153-235

Authors:

Anton Hausknecht

University of Vienna

Irmgard Krisai-Greilhuber

University of Vienna

Hermann Voglmayr

University of Vienna

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.

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Österr.

Z. Pilzk. 13 (2004) 153

Type studies in North American species of

Bolbitiaceae

belonging to the

genera

Conocybe

and Pholiotina

ANTON

HAUSKNECHT

Sonndorferstraße

A-3712

Maissau, Austria

IRMGARD

K.R1SAI-GRE1LHUBER

HERMANN

VOGLMAYR

Institut

für Botanik der Universität Wien

Rennweg

A-1030

Wien. Austria

Received

12.8.2004

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

Ilava

und l'holiolina plumbeilmcla werden vorgeschlagen Mikroskopische Zeichnungen zu allen unter-

suchten

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

ofthetaxa

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-

tated

Materials and methods

Microscopical investigations were performed with an Olympus BH-2 microscope, the drawings with a

drawing tube at magnifications of

1:2500

(spores) or

1:1000

(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

hours.

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

the

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-

nate,

ventncose, broad, inserted, medium distant, entire, fulvous, with white lamellar

edges,

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 =

1.4-1.7,

irregu-

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

diam.

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

pileipellis.

/-/ ('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-

leus.

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:

Pileus

1.2-3.5

cm latus, convexus, deinde planus. pallide ochraceus. nudus, camosus,

lamellae latae, subliberae, ochraceae; stipes cylindricus, basi inflatus. pallidus, carina-

tus

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 =

1.4-1.6,

el-

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

holotype).

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.

angusticeps

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.

atkinsonii

see procera

besseyi

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-

SEYand E. A. BESSEY.

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

8.5-10

x 7-8.5 urn. mean 15.0 x 94 x 7.8 um, Q =

1.5-1.7,

ellip-

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

found.

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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

diam

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

1912.

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=

3-2.1.

ellipsoidal

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)

161

Fig. 3 a-e. (jalervla canalipes (type), a Spores, b basidia. c cheilocystidia, d stipe covering, e pileipel-

lis.

1-1.

(ialemla capillanpes (type). /Spores, g basidia. h cheilocystidia ' stipe covering, j-m. Agari-

LUS

crocosporus

(type).

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

„cotype").

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-

tures,

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

8.5-10

urn.

Clamp connections: not observed

Cheilocystidia: lecythiform, 12-17 x 6-9.5 urn, with capitulum 3-4 5 urn in

diam.

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-

nous,

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

diam.

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

(NYS.

holotype).

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:

Pileus

1.5-3.5

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

diam.

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)

165

Fig. 4 a-e. (iaiera cnspa (from type locality) a Spores, h basidia. c cheilocystidia. J stipe covering

from the middle of

stipe,

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

(1998).

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).

crispella

(¥\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.

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Österr. Z. Pilzk. 13(2004)

167

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,

pileipellis.

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168

HAUSKNECHT

& 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

elements

Specimen examined: USA: Florida, Alachua County, Gainesville, on grass on open lawn, 5. 8.

1941.

leg. W. A.

MURRILL

(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

j-m)

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

10:

168

Conocybe crocospora

(BERK

& M A CURTIS) KUHNER 1935, Le genre Galera 137,

nom inval (no basionym cited)

Original

description:

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

Microscopical

characters:

Spores: 11

5-14.5

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 several sections)"

Stipe covering: composed of non-lecythiform. roundish, cylindrical to shortly

capilliform elements, no lecythiform elements found.

Pileipellis: some roundish-stipitate elements found, ca 30 x 18-26 urn in diam.,

mostly collapsed.

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Osten. Z. Pilzk. 13 (2004) 169

Specimen examined: USA: South Carolina, on moist straw mat in house. August (K 102521.

holotype)

The holotype consists of

two

molded fruitbodies in very poor condition

Agaricus crocosporus undoubtedly belongs to Conocybe, sect Pilosellae, stirps

Siliginea The large, ellipsoidal, non-lentiform spores and the pileus described as "sor-

did brown, viscid" place this taxon in the vicinity of Conocybe fuscimarginata, which

has smaller, somewhat lighter spores.

Due to the very poor preservation of the type material we refrain from combining

this species into the genus Conocybe.

cryptocystis (Fig 5 a-e)

Galerula cryptocystis G. F ATK. 1918, Proc. Amer. Phil. Soc. 57: 368

Conocybe cryptocystis (G. F. ATK.) SINGER 1954, Sydowia 8 125

Original description:

Gregaria, 6-10 cm alta; pileo ovali dein campanulato,

1,5-2,5

cm lato, ochraceo, ochra-

ceo-fulvo vel ferrugineo, demum pallidiore, sparsim villoso; lamellis stipite adnexis.

latis;

stipite aequali, pileo concolore sed pallidiore, striato, sursum pruinoso, 2-4 mm

crasso.

Microscopical characters:

Spores: 13.5-16 5 x 8 5-11 x 8-9.5 urn, mean 15 6 x 100x8.8 um, Q =

45-1.65,

ellipsoidal, conspicuously lenticular, mustard yellow in KOH with a ca 15-2 urn thick

wall and a large, truncate germ-pore.

Basidia: 2(

)-spored, 12-17 x 7-10.5 urn.

Clamp connections: not observed

Cheilocystidia: lecythiform, 14-17 x 6-8.5 urn, with capitulum 3-4.5 um in

diam

Stipe covering: composed of lecythiform caulocystidia, 9-13 x 6-9 urn, with

very small capitulum 2-2.5 urn in diam.; in-between abundant hairs up to 2 urn thick

and some non-lecythiform elements

Pi lei pel

is hymeniform, composed of roundish elements up to 33 x 30 urn, no

pileocystidia present.

Specimen examined: USA: California, Santa Clara County, near Stanford University, on de-

cayed horse manure, 30 11 1901, Pacific Slope Fungi No. 127, leg. C. F. BAKER (CUP, holotype).

The type consists of about four fruitbodies broken into many fragments in good

condition The microscopical analysis and the following comment are a translation

from HAUSKNECHT (2003).

Conocybe cryptocystis was described by ATKINSON (1918) as a "common little

fungus on decayed horse manure in old pastures", and should therefore be quite com-

mon in the western USA. The spore sizes measured by us are considerably smaller

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170

HAUSKNECHT & al.: Type studies

North American Holbinaceae

than given by ATKINSON (1918: 368) and are in-between Conocybe merdaha AR-

NOLDS & HAUSKN. (ARNOLDS & HAUSKNECHT 2003) and Conocybe macrospora

(ATK.) HAUSKN. However, the spores are distinctly lenticular and in front view wider

than those of both species, and also thicker-walled, but in K.OH paler, mustard yellow

and not reddish brown. In addition, no pileocystidia were found in the pileipellis of

Conocybe cryptocystis. and the hymeniform elements of the pileipellis are overall

wider Due to the different spore characteristics we preclude conspecificity with Cono-

cybe merdaria and C. macrospora.

curia (Fig. 5 f-j)

GalerulacurtaG. F ATK. 1918, Proc. Amer. Phil Soc 57: 1918

Conocybe curia (G. F. ATK.) WATLING 1981, Bibl Mycologica 82 103

Original description:

Gregana. 4-6 cm alta; pileo campanulato dein subexpanso, 2-4 cm lato, 1-2 cm alto,

obtuso. glabro. non stnato. ochraceo vel ochraceo-fulvo demum ochroleuco; lamellis

stipite adnexis, ellipsoideis, ochraceis; stipite aequali, glabro, striatulo, sursum prui-

noso.

cavo. 3-5 mm crasso

Microscopical characters:

Spores:

1.5-16

x 7.5-10 urn. mean 13.9-14.0 x

8.5-8.7

um, Q =

1.4-1.9,

ellipsoidal,

somewhat limoniform. not compressed, brownish orange in K.OH with up to 15 urn

thick wall and a conspicuous germ-pore

Basid ia: 2-spored, 17-22 x 11-13 urn

Clamp connections: not observed

Cheilocystidia: lecythiform, 13-19 x 5-10 urn, with capitulum 3-5 um in

diam.

Stipe covering: composed of lecythiform caulocystidia and capilliform ele-

ments, many of which ending in a conspicuous capitulum (12-22 x 3-6 urn. capitulum

2.5-3.5 um in diam), many without apical capitulum.

Pileipellis: hymeniform, composed of roundish-stipitate elements, 16-30 x 15-

21 urn; no pileocystidia present.

Specimens examined: USA:

New

York, Ithaca, Buffalo Street

grass

roadside.

20. 7. 1899,

leg.

G. F.

ATKINSON 3210 (CUP, holotype);

- -

13.

1899.

leg. G. F.

ATKINSON (CUP. paratype).

The type consists of three well preserved fruitbodies, the stipes of which are bro-

ken Also the paratype is in good condition The microscopical analysis and the fol-

lowing commentary are a translation from HAUSKNECHT (2003).

Both investigated specimens originate from the same locality, they were collected

in the interval of one week. The species shows a great similarity with a very robust,

thick-stiped Conocybe hckenii. but has stipe covering of sect. Mixtae with many lecy-

thiform elements between the stipe hairs. The spore size resembles that of Conocybe

merdaria. from which it differs in size and habitus, the very pale, not hygrophaneous,

non-stnate pileus and the rather slightly limoniform spores.

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Österr. Z. Pilzk. 13(2004)

171

Fig. 6 a-c. Oalera cyanopes (neotype). a Spores, h two cheilocystidia. c pileipellis. d-g. Oalerula cya-

mopvs. d Spores, e basidia. /cheilocystidia g pileipellis. h-k. (onocybe dennisu (type), h Spores. /

h>asidia.y cheilocystidia. k stipe covering.

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172 A. HAUSKNECHT &al.: Type studies in North American Holbiiiaceae

cyanopes (Fig 6 a-c)

Galera cyanopes K.AUFFMAN 1918, Mich. Geol. & Biol. Survey 26, Biol. Ser. 5,1: 500

Pholiotina cyanopes (K.AUFFMAN) SINGER in RUMACK & SALZMANN 1978, Mush-

room Poisoning: 203

Conocybe smithii WATLING in BENEDICT, TYLER & WATLING 1967, Lloydia 30: 152

Pholiotina smithii (WATLING) ENDERLE in ENDERLE & HIIBNER 1999, Z Mykol.

65:

Original description (from WATLING in BENEDICT & al. 1967):

Pileus 3-10(-13) mm. obtusely conic soon expanding at the margin to become dis-

tinctly umbonate, ochraceous tawny, tawny cinnamon, darker at the disc which is usu-

ally more tawny, stnate for 2/3 or more, hygrophanous, fading to pinkish buff or dingy

tan on drying, glistening Stipe 10-50(-70) x 0.75-1.0 mm (15 mm at base), slender

slightly swollen at the very base, fragile, pure white then watery white and finally

flushed with ochraceous pallid but only faintly so, base particularly tinged greenish

grey or greyish blue but sometimes tinged above as well, finally quite ochraceous, fi-

brillose to glabrescent at base, smooth or faintly pubescent at very apex. Gills adnate

soon receding, subdistant to crowded, narrow to moderately broad, ochraceous buff at

first becoming cinnamon rust, margin ± concolorous with face, even or becoming mi-

nutely crenulate or flocculose. Flesh watery white; odour none; taste not distinctive.

Habitat: scattered amongst mosses in boggy areas and wet ditches.

Microscopical characters:

Spores: 7-9x4-5 urn, mean 7.8 x 4.4 um, Q =

1.6-1.9,

slender ellipsoidal, not lenti-

form, thin-walled with conspicuous germ-pore, ochre yellow to rusty orange in K.OH.

Basidia: 4-spored

Cheilocystidia: mostly collapsed, two more or less fusiform elements with

apical capitulum found, these 30-31 x 8-10 um in size.

Stipe covering: completely collapsed; according to WATLING (in BENEDICT &

al.

1967) similar to the cheilocystidia, but more variable.

Pileipellis: hymeniform, composed of roundish-stipitate elements, 28-40 x 14-

20 urn, in-between sporadic pileocystidia similar to the cheilocystidia, but more slen-

der.

Specimens examined: USA: Michigan, Washtenaw County, Chelsea, in huckleberry bog on

mosses. 27. 7. 1909, leg. C. H. KAUPFMAN (MICH, designated as neotype). - Montana, Flathead Lake,

Yellow Bay, near Biological Station, on soil among moss, 18. 7. 1989, leg. BAILEY (ZT; see HORAK &

HAUSKNECHT 2002: 244).

The specimen designated as "part of neotype" consists of one strongly collapsed

fruitbody, which has almost no intact lamellae.

WATLING (in BENEDICT & al. 1967) has seen fresh material of this species, which

is rather common in North America, and he mentions the following differences to

Pholiotina cyanopus (G. F. ATK.) SINGER: The lamellae are more cinnamon brown in

smithii, the pileus is paler and moist more distinctly striate, the fruitbodies are more

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Osten. Z. Pilzk. 13 (2004) 173

delicate and smaller, the spores narrower, the number of pileocystidia smaller and the

cheilocystidia wider and have a less distinct capitulum Unfortunately, the differences

in spore size and form given in STAMETS (1999) are incorrect, as a comparison of both

type specimens shows (means of both specimens almost identical). Also the different

number of pileocystidia is not confirmed in our studies Therefore, the only confirmed

differences concern the somewhat wider cheilocystidia of Pholiotma smithii and some

macroscopical differences A detailed analysis of European material of sect. Cyanopo-

dinae (SlNGER) ARNOLDS is still missing, and as we know Pholiotina smithii only from

herbarium specimens, we tentatively agree with WATLING (in BENEDICT & al 1967)

and consider that it is a distinct species. In case of conspecificity, Pholiotina cyanopus

is the valid name due to priority reasons, as it has been published a few months earlier

cyanopus (Fig 6 d-g)

Galerula cyanopus G F ATK. 1918, Proc Amer Phil Soc 57: 367

Conocybe cyanopus (G. F ATK.) KÜHNER 1935, Le genre Galera: 128 (as "cyano-

poda")

Pholiotina cyanopus (G. F. ATK.) SINGER 1950, Acta Inst. Bot. Komarov Acad. Sei.

URSS,

Series 2,6: 425 (as "cyanopoda")

Original description:

Gregaria vel solitana, 2-3 cm alta; pileo ovali, campanulato-convexo, dein expanso,

obtuso. ad marginem striatulo. fragili. brunneo-ferrugineo, 5-10 mm lato; stipite 1-1,5

mm crasso, leniter bulboso, albo ad basem cyaneo, sursum pruinoso, deorsum leniter

velutino, fragili.

Microscopical characters:

Spores: 7-9 x 4-5 urn, mean 7.8 x 46 urn, Q = 1.55-1.85, ellipsoidal, slightly li-

moniform, thin-walled with a small, but distinct germ-pore, ochre yellow in K.OH.

Basidia: 4-spored, 15-20 x 7-10 5 urn.

Clamp connections: not found.

Cheilocystidia: 14-30 x 4-12 urn, very polymorphic, slightly utriform, cylin-

drical-lageniform with capitulum to almost slightly moniliform.

Pileipellis: hymeniform, composed of sphaeropedunculate elements, 21-31 x

13-18 urn, pileocystidia not observed, according to WATLING in BENEDICT & al. (1967)

similar to cheilocystidia, but with a more distinctly capitulum-like apex.

Specimen examined: USA: New York. Ithaca, among grass on campus. Aug. 1912, leg. G. F.

ATKINSON 23302 (CUP. holotype)

The holotype consists of fragments of about 8-12 fruitbodies in good condition,

and all microscopical features could be observed.

Galerula cyanopus is another member of Pholiotina, subsect. Cyanopodinae (AR-

NOLDS 2003). It has been recorded from North America and Europe. The blueing of

the stipe base is of quite variable intensity and may be easily overlooked. Then, the

species can be determined by the combination of shape and size of cheilocystidia,

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174 A. HAUSKNECHT & al.: Type studies in North American Holbiiiaceae

shape and poms features of the spores and presence/absence of pileocystidia For the

differences to Pholiotina smithii which has not yet been reliably recorded for Europe,

see discussion under cyanopes.

dennisii(\

\g 6 h-k)

Conocybe dennisii HAUSKN. in HORAK & HAUSKN. 2002, Österr Z Pilzk 11214

Original description:

Pileus 3-5 mm diam. convex, umbo absent, centre and striation light ochraceous

buff,

ochraceous

buff,

smooth, pale elsewhere, hygrophanous. dry. Lamellae rather distant,

emarginate-adnexed, narrow, pale brown. Stipe 6-10 x 0.2 mm (from dried specimen),

cylindrical, filiform, equal, white over whole length, minutely pruinose, dry Context

fragile.

Microscopical characters:

Spores: 4.5-5.5 x 3.5-4 um, mean 5.0 x 3.8 urn, Q = 1.3-1.55, roundish to submitri-

form in front view, subellipsoidal in lateral view, slightly lentiform. wall double,

slightly thickened, germ-pore absent, pale orange-brown in KOH.

Basid ia: 4-spored, 10-13 x 6.5-7.5 (im.

Clamp connections: present

Cheilocystidia: lecythiform, 13-19 x 6.5-8.5 urn, with capitulum 2.5-4 urn

diam

Stipe covering: consisting of lecythiform caulocystidia 10-15 x 4-6 urn,

capitulum 2.5-3.5 urn in diam . generally smaller than cheilocystidia.

Pileipellis: hymeniform. composed of clavate cells up to 20 urn in diam. Pileo-

cystidia not observed

Material examined: Trinidad and Tobago: Trinidad. St. Joseph, on soil among moss under

Hambusa

spec.

(Poaceae),

14. 10. 1949, leg. R. W. G. DENNIS

176

(K. holotype).

The holotype consists of one small fruitbody in good condition

The species belongs to the Conocybe mesospora complex, and it is clearly distinct

from all known European and North American species by the combination of small, in

front view slightly mitriform spores without a germ-pore, small fruitbodies and stipe

covering consisting of only lecythiform elements (see HORAK & HAUSKNECHT 2002)

fibrillosipes (Fig. 23 h-k)

Conocybe fibrillosipes WATLING 1971, Persoonia 6: 325

Original description:

Pileus 10-17 mm, convex, semiglobate. hardly expanding, rich dark tawny orange,

("'tawny'") tinted with „chestnut" particularly towards the disc, stnate to half-way and

covered particularly at margin with small, pale ochraceous fragments of appendiculate

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östenr. Z. Pilzk 13 (2004) 175

and marginal veil Stipe 18-32 x 3-5 mm, distinctly "yellow ochre", stout, fibrillose-

streaky to woolly floccose, particularly towards the base, veil peronate or present as

mere flecks of membranous material towards the stipe-apex, then fibrillose-woolly,

matt, giving a rough appearance below, floccules finally yellow ochraceous. Lamellae

pale ochraceous then rusty tawny, fairly close. Flesh dark brown in pileus when fresh,

slightly ochraceous in stipe-cortex, dark buff in stipe-centre, umber brown ("Saccardo's

umber") in stipe base.

Microscopical characters:

Spores: 7.5-9.5 x 4-5 5 urn, mean 8.2 x 4.6 um, Q =

1.6-2.0.

ellipsoidal, in side view

partly slightly bean-shaped, thin-walled with conspicuous, small germ-pore, pale yel-

low in K.OH

Basidia: 4-spored (no 2-spored observed), 18.5-22.5 x 7-8 urn.

Clamp connections: present at the basis of the basidia and in the trama.

Cheilocystidia: lageniform to lecythiform, usually with conspicuous capitu-

lum, 20-35 x 7-9.5 urn, capitulum 3-6 urn wide.

Stipe covering: consisting of variable elements, cylindrical-ventricose to al-

most filamentous with capitate apex, often also lecythiform similar to the cheilocys-

tidia or arranged in chains.

Pileipellis: hymeniform, composed of roundish-stipitate elements, 25-45 x 12-

24 urn, in-between rarely pileocystidia similar to the cheilocystidia, but less ventri-

cose.

Specimen examined: USA: Washington, Pend' Oreille County, Cusick, on roadside amongst

herbaceous debris in conifer woodland, 12. 10. 1966, leg. R.

WATLING

(E,

holotype).

The type material consists of about 25 well-preserved fruitbodies in various devel-

opmental stages.

WATLING (1971) based his description on a single collection, it should differ from

Pholiotina brunnea (WATLING) BON mainly in its more compact habitus and its abun-

dant, yellow velum. We had collections from Europe which are intermediate in habitus

and velum between Conocybe fibrillosipes and typical Pholiotina brunnea. Also a col-

lection from North Carolina (USA) could not be unequivocally assigned to one of

these two taxa, it had a similarly rich velum like Conocybe fibrillosipes but a slender,

thinner stipe like Pholiotina brunnea. Therefore, HORAK & HAUSKNECHT (2002) con-

sidered Conocybe fibrillosipes synonymous with Pholiotina brunnea from Europe. As

microscopically there are no differences between collections with scarce and rich ve-

lum, respectively, on the one hand, and between those with a compact and slender

stipe,

respectively, on the other hand, we fully agree with conspecificity of both taxa

after the study of the type collection.

It should be noted that Conocybe brunnea has been published in the same publica-

tion (WATLING 1971) only few pages ahead of Conocybe fibrillosipes, the former

name has therefore priority.

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176 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

filipes (Fig 7 a-e)

Galerula filipes G. F. ATK. 1918, Proc. Amer Phil. Soc. 57: 367

Conocybe filipes (G. F. ATK.) KÜHNER 1935, Le Genre Galera: 127 (invalid)

Pholiotina filipes (G. F. ATK.) SINGER 1950, Acta Inst. Bot Komarov Acad Sei.

URSS,

Ser. 2,6:435

Original description:

Gregana, 4-6 cm alta, pileo ovali dein campanulato, obtuso, ochraceo, 3-6 mm lato;

lamellis stipite late adnatis, dente decurrentibus, ochraceis vel ochraceo-tulvis, stipite

aequali, bulbilloso, sursum pruinoso, 1 mm crasso.

Microscopical characters:

Spores: 7-9.5 x4.5-5 (im, mean 8.3 x 4.8 um, Q =

1.5-1.9,

ellipsoidal, not lentiform,

with double wall and a germ-pore up to 1 urn in diam.. yellow to orange-yellow in

KOH

Basidia: 4-spored, ca 12-18 x 6-8 urn.

Clamp connections: not observed

Cheilocystidia: 20-35 x 6-11 urn, lanceolate, irregularly fusiform ventricose

base,

usually collapsed and with broken tip (certainly many longer than 35 (im)

Stipe covering: consisting of caulocystidia similar to the cheilocystidia, but

larger, up to 45 x 15 urn.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 25-35 x 14-

20 urn; pileocystidia abundant, similar to cheilocystidia in shape, but much larger (up

to 80 x 8 urn), sometimes yellowish pigmented.

Specimen examined: USA: New York, Essex County, Adirondack Mountains, North Elba, leg.

H. PECK as (ialera capillanpes (NYS. holotype).

The type collection consists of six to seven fragmented fruitbodies; especially the

pilei are heavily broken and most cystidia collapsed.

If the original description and also the microscopical analysis are compared with

Pholiotina sulcatipes (PECK) BON and Pholiotina aberrans (K.UHNER) SINGER, there is

only one major difference against the former: the stipe base of Pholiotina sulcatipes

has been described as "blue or green at the base". The spores and caulocystidia are

almost similar, and both have abundant very large caulo- and pileocystidia HAUSKNECHT

(2001) synonymized the European Pholiotina aberrans with Pholiotina sulcatipes.

This synonymy has not been accepted by some authors; they consider both taxa as dif-

ferent species (e.g., ARNOLDS 2004). In this case, the species of

ATKINSON

would have

priority and European collections would have to be named Pholiotina filipes.

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Österr. Z. Pilzk. 13(2004)

177

Fig. 7 a-e. Galerula filipes (type), a Spores, b basidia, c cheilocystidia, d caulocystidia, e pileipeilis

with numerous pileocystidia /-/. Conocybe fimicola (type)./Spores, g basidia, h cheilocystidia, / pi-

leipeilis.

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178 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

fimicola (Fig 7 f-i)

Conocybefimicola WATLING 1971, Persoonia 6: 335

Pholiotina fimicola (WATLING) ENDERLE 1997, Z Mykol 63 32

Original description:

Pileus 10-25 mm conico-convexus, expanso-convexus vel obtuso-umbonatus, glaber

humidus. striatus. jove sicco exstriatus. hygrophanus. ferrugineus jove sicco ochraceo-

fulvus. Stipes 30-40 x 2-3 mm, aequalis, farctus, cinnamomeo-bubalinus ad basim ex

luteolo-bubalinus postremo umbrinus. ad apicem fibrilloso-pruinosus ad basim ap-

presso-tlbnllosus, annulus medius, membranaceus, bubalmus, striatus, fugaceus. La-

mellae adnatae, L 20-25, I 1-3, sordide fulvae, subconfertae. Caro tenuis, fragil is, fer-

ruginea.

Microscopical characters:

Spores: 7-9.5 x 4-5 urn, mean

8.3-8.8

x 4.6-4.7 |im, Q =

1.7-2.0,

ellipsoidal, slightly

pip-shaped, not lentiform, with relatively thin wall and a germ-pore ca 1 urn in diam..

yellow in K.OH.

Basid ia: 4-spored, 16-23 x 6.5-8.5 um.

Clamp connections: present

Cheilocystidia: 22-36 x 6-8 5 (im, fusiform-ventricose with a long, pointed

beak, sometimes with an apical slime cap up to 6 urn in diam.

Pileipellis: hymeniform, consisting of pyriform to roundish-stipitate elements,

30-40 x 15-22 urn. pileocystidia absent.

Specimens examined: USA: Washington, Clallam County, Olympic NP., Lake Crescent, gre-

garious on mature pile of

dung.

29. 5. 1939, leg. A. H. SMITH 13856 (MICH, holotype): - - 9. 6. 1939,

leg A. H. SMITH 14192 (MICH).

The holotype consists of fragments of one well-preserved fruitbody, the material

from the second collection - from the type locality - is much more profuse Micro-

scopically, both agreed perfectly.

WATLING (1971) characterized his new species particularly by its habitat (dung

inhabiting) and the features of the ring, but conceded that "the cheilocystidia shape

would place it close to Pholiotina filaris (FR.) KÜHNER". The combination of

EN-

DERLE (1997) was without commentary. Actually, Pholiotina fimicola and Pholiotina

filaris cannot be distinguished macro- and microscopically, neither concerning the

cheilocystidia nor the shape and size of the spores. One collection of the third author

from Tennessee, Gatlinburgh, 1. 4. 2000, by no means dung inhabiting and determined

as Pholiotina filaris by us, has features identical to Pholiotina fimicola, but it grew on

bare soil at a roadside. On the other hand it is well-known that European collections of

Pholiotina filaris often grow on mineral rich soil, under stinging-nettle or in leaf litter

Some collections of H.-J. HÜBNER originating from southern Germany grew on fresh

compost soil, and there is not much difference to dung.

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Osten. Z. Pilzk 13(2004)

179

Fig. 8 a-e. (ialera flava (type) a Spores, b basidia c cheilocystidia d elements of

the

stipe covering, e

two elements of pileipellis. /-/. (onocybe flexipes (type). /Spores, g basidia h cheilocystidia ; pilei-

pellis.

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180 A. HAUSKNECHT &al.: Type studies in North American ßolbmaceae

Despite these arguments it is advisable to wait with a definite statement on con-

specifity of both taxa until further collections from North America admit evaluation of

macro- and microscopical variabilty

flava (Fig 8 a-e)

Galeraflava PECK 1893, Ann Rep New York State Mus Nat. Hist. 45: 79

Galerulaflava (PECK) MURRILL 1917. North American Flora 10: 166

Conocybe flava (PECK) KÜHNER 1935, Le Genre Galera. 137, nom inval (no ba-

sionym cited)

Pholiotina flava (PECK) HAUSKN., KRISAI-GREILH. & VOGLMAYR, comb, nova

Basionym: Galeraflava PECK. 1893, Ann. Rep. New York State Mus. Nat. Hist. 45: 79

Original description:

Pileus 12-24 mm broad, ovate or campanulate, obtuse, moist or subhygrophanous, yel-

low, atomate when dry, surface breaking up into squamules, finely plicate-striate to the

middle Context thin. Lamellae adnate, whitish then yellowish-cinnamon, close, nar-

row. Stipe 5-7.5 cm long, 2-4 mm thick, white or slightly tinged with yellow, with

white mealy particles, slightly stnate at top. equal or slightly tapering upward, hollow

Microscopical characters:

Spores: 9.5-11 5 x 6-8 x 5.5-7 um, mean 10.6 x 7.5 x 6.2 um, Q =

1.3-1.5,

ellipsoi-

dal to slightly angular, conspicuously lentiform. with thick wall and distinct germ-

pore,

orange-yellow to rust-yellow in K.OH

Basid ia: 4-spored, 19-22 x 10-12 urn.

Clamp connections: not observed

Cheilocystidia: one lamellar fragment with some cheilocystidia found, these

are rusiform-ventncose to slightly lagemform, 17-25 x 5-9 urn

Stipe covering: consisting of some hairs and cylindrical-twisted elements, 18-

40x2-7 urn

Pileipellis: hymeniform, consisting of pyriform elements, ca 35-47 x 30 urn.

No pileocystidia found

Specimen examined: USA: New York, Tompkins County, Freeville, on damp vegetable mould

in woods. July 1892 (NYS. holotype).

The type specimen is in very poor condition. It consists of sparse fragments of

about two fruitbodies glued to paper

HESLER (in his unpublished records) did not find cheilocystidia. The lamellar frag-

ment investigated by us had altogether 8-10 cheilocystidia. of which five were in good

condition and resembled in shape and size those of Pholiotina mairei (WATLING)

EN-

DERLE. but the spores are completely different. It may be related to Pholiotina cop-

rophila, from which it differs by completely different spores and cystidia. It should be

mentioned here that within the genus Pholiotina lentiform, slightly angular spores are

hitherto unknown for European and North American species.

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Österr. Z. Pilzk. 13(2004) 181

According to the original description, the pileus surface breaks up into squamules.

However, according to the pileipellis structure this should only be the product of

weather conditions. Also HESLER (unpubl.) did not find squamules in the exsiccate.

Although the type specimen is in very poor condition, we decided to combine

Galera flava into Pholiotina due to the unique combination of distinctive microscopi-

cal features, the yellow pileus and the white stipe.

flexipes (Fig. 8 f-i)

Conocybe flexipes WATLING 1971, Persoonia 6: 336

Pholiotina flexipes (WATLING) ENDERLE 1997, Z Mykol. 63: 32

Original description:

Pileus 5-15 mm, conicus, convexo-expansus vel campanulatus vix plano-umbonatus,

glaber, humidus, hygrophanus, striatus, pallide ochraceo-fulvus vel incarnato-buba-

linus.

Stipes 50-70 x 1-1,5 mm, aequalis, saepe flexuosus, ad apicem pallido-bubali-

nus.

ad basim pallide ochraceo-fulvus ad apicem pruinosus, ad basim appresso-fibril-

losus vel glaber annulus apicalis, membranaceus. pallidus vel pallido-bubalinus ad

marginem crassus et plumosus. Lamellae confertae, adnatae, pallido-bubalinae pos-

tremo ochraceo-fulvae vel fulvae. Caro delicatula, pallido-ochracea.

Microscopical characters:

Spores:

8.5-11

x 4.5-6 urn, mean 9.7 x 5.3 um, Q =

1.7-2.0,

ellipsoidal to slightly

drop-shaped, with slightly double wall and a germ-pore 0.5-0.8 urn in diam, yellow in

KOH

Basidia 4-spored, 19-24 x 8-10 5 urn

Clamp connections: common at the base of the basidia and in the trama

Cheilocystidia: 30-40x8.5-14 urn, usually utriform, lageniform-capitate with

up to 13 urn wide, capitate apex and a thick neck, rarely only lageniform, sporadically

roundish up to 25 urn wide elements in-between.

Stipe covering: not observed.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 20-45 x 15-

25 urn; pileocystidia absent.

Specimen examined: USA: Washington, Pierce County, Mt. Rainier NP, in moss, decaying

wood and detritus from herbaceous plants. 19. 10. 1952. leg. A. H. SMITH 41179 (MICH, holotype).

The type specimen consists of two well-preserved fruitbodies with conspicuously

yellow colour in dried condition

ENDERLE & HUBNER (1999) in their publication of Pholiotina utricystidiata EN-

DERLE & HUBNER shortly listed the differences ofthat species to Pholiotina flexipes

Except for the size of the fruitbodies and the in Pholiotina flexipes more distintly

flexuous stipe, which we do not consider taxonomically significant, they regard the

somewhat smaller spores and the cheilocystidia as distinguishing features.

Actually, both taxa are very close, and we would not consider the mentioned fea-

tures sufficient enough to separate both taxa at the species level. However, after com-

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182 A. HAUSKNECHT & al.: Type studies in North American Bolbiriaceae

parison of both type specimens we found another distinguishing feature: not only are

the spores of Pholiotina utricystidiata about 1-2 urn longer, but these are also thicker-

walled and in KOH not yellow but yellowish-brown to rust-brown. We therefore do

not consider both taxa conspecific

floridana (Fig 9 a-e)

Galerulafloridana MURRILL 1941. Mycologia33: 445

Conocybe antipus (LASCH)

FAYOD

i. floridana (MURRILL) SINGER 1950, Sydowia 4:

131

Original description:

Pileus conic to subexpanded, solitary, 2-5 cm broad; surface dry. smooth, glabrous,

uniformly isabelline. margin entire, even, upturned with age; context thin, pallid,

odorless, lamellae adnate or adnexed, narrow, close, unequal, entire, soon becoming

pale fulvous; stipe long, slender, equal, smooth, glabrous, long-radicate, pallid. 7-10 x

0.1-0.3 cm.

Microscopical characters:

Spores: 8-9.5x5.5-7x5-5.5 urn, mean

8.3-8.6

x 6.2-6.4 x 5.3 urn, Q= 1.25-1.45, in

front view angular-submitriform, in side view ellipsoidal, conspicuously lentiform,

with double wall and large germ-pore, rust-orange to orange-brown in KOH.

Basidia: 4-spored, 16-19x7-10 um.

Clamp connections: not observed.

Cheilocystidia: lecythiform, 14-18 x 6-9 urn, with capitulum 3-5 5 urn wide.

Stipe covering, consisting exclusively of lecythiform, often collapsed ele-

ments similar ot cheilocystidia, but often thinner, partly almost cylindrical with capi-

tulum up to 4 urn wide.

Pileipellis: hymeniform. consisting of sphaeropedunculate elements, 22-36 x

19-24 urn. Pileocystidia not observed.

Specimens examined: USA: Florida, Gainesville. Sugarfool Hammock, in sandy soil under trees,

6. 1938. leg. W. A. MURRILL (FLAS. holotype): - Florida Gainesville, in oak woods. 3. 6. 1938,

leg.

W. A. MURRILL (FLAS. designated as "cotype").

The type specimen consists of fragments of one moderately preserved fruitbody,

particularly the stipe covering is heavily collapsed. The second specimen is somewhat

better preserved and has a conspicuous, long root.

SINGER (1950) downgraded Galera floridana as a form of Conocybe antipus "with

somewhat smaller spores". The spore size of both specimens fully corresponds with

that of many collections from Europe; therefore, we consider Galera floridana to be a

Synonym of Conocybe antipus

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Osten- Z Pilzk 13(2004) 183

Fig. 9 a-e. Cialerulaßoridana (type), a Spores, b basidia. c cheilocystidia, d stipe covering, e pileipet-

\is.f-j. GaleraJragilis (type)./ Spores, g basidia, h cheilocystidia. / stipe covenng. / pileipellis.

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184 A. HAUSKNECHT & al.: Type studies in North American fiolbmaceae

fragilis (Fig 9 f-j)

Galerafragilis PECK 1897, Bull. Torrey Bot. Club 24: 144

Galerulafragilis (PECK) MURRILL 1917. North American Flora 10: 164

Conocybe fragilis (PECK) SINGER 1950, Acta Bot Inst Komarov Acad Sei URSS,

Series 2, 6: 438

Original description:

Pileus submembranous, very fragile, broadly campanulate, glabrous, dull flesh colour;

lamellae ascending, adnate, subdistant, dark yellow or subochraceous, becoming fer-

ruginous; stem slender, flexuous, hollow. Pileus 3-5 lines broad; stem 10-15 lines long, 0.5

line thick.

A very small and fragile plant.

Microscopical characters:

Spores:

8.5-10

x 5-6 urn, mean 9.1 x 5.4 urn, Q = 1.55-1.8, ellipsoidal, lentiform,

with slightly double wall and a germ-pore up to 1.5 urn in diam., bright yellow to light

orange-yellow in K.OH

Basidia: 4-spored, 18-22x9-11 urn.

Clamp connections: not observed.

Cheilocystidia: lecythiform, 14-21 x 6.5-9.5 urn, with capitulum 3.5-4.7 urn

wide

Stipe covering: consisting mainly of hairs and non-lecythiform elements; in-

beetween up to 20 % lecythiform caulocystidia similar to cheilocystidia in shape and

size.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 22-40 x 15-

25 urn; no pileocystidia observed

Specimen examined:

I ISA:

Kansas, Rooks County, among short grass in pastures, leg. E.

BARTHOLOMEW

2313 (NYS, holotype).

The type material consists of hundreds of very small fragments of several well-

preserved fruitbodies The description given above is a translation from HAUSKNECHT

(2003).

Conocybe fragilis belongs to sect. Mixtae due to its stipe covering This taxon has

long been misinterpreted in Europe; recently, ARNOLDS & HAUSKNECHT (2003) dem-

onstrated that two different species are involved which are not even closely related

with each other All specimens from Europe identified as Conocybe fragilis are likely

to be incorrectly determined; whether specimens from overseas (NATRAJAN & RAMAN

1983:

India, WATLING 1992: Brazil, WATLING 1994: Malaysia) are conspecific with

the type is questionable, as all authors assumed a stipe covering of

sect.

Pilosellae.

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Österr. Z. Pilzk. 13(2004)

185

Fig. 10 a-d. Galerula juscimarginata (type), a Spores, b basidia, c cheilocystidia, d stipe covering, e-i.

('onocybe

aniipus var. hunucola (type), e Spores. / basidia. g cheilocystidia. h stipe covering, /

pileipellis.

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186 A. HAUSKNECHT & al.: Type studies in North American Bolbiiiaceae

fuscimarginata (Fig lOa-d)

Galerulafusamarginata MURRILL 1942, Lloydia 5: 148

Conocybe fuscimarginata (MlRRILL) SINGER 1969, Beih Nova Hedwigia 29 210

Original description:

Pileus conic, not expanding, gregarious, about

cm, surface dry, smooth, glabrous,

uniformely isabelline. margin even, entire, becoming brownish with age. context

membranous; lamellae adnate, inserted, narrow, crowded, entire, stipe stnate. pallid,

enlarged and pubescent above, clavate at the base. 4-5 cm long, 1 mm or less thick.

Microscopical characters:

Spores: 10-12.5 x 6-8 urn, mean 11.2 x 6.8 (im, Q =

1.5-1.9

um. ellipsoidal, not

lentiform. with 0.3-0.5 um thick wall and a germ-pore up to 1.5 urn in diam . rust-

yellowish brown in KOH.

Basidia: 4-spored. 16-21 x 9-11 urn.

Clamp connections: not observed

Cheilocystidia: lecythitbrm, 14-17 x 6-8 urn, with capitulum 3-4 urn wide.

Stipe covering: cosisting of hairs, roundish to fusiform-lagemform elements,

the latter up to 55 x 11 urn, lecythiform caulocystidia absent.

Pileipellis: hymeniform. consisting of roundish-stipitate elements

Specimen examined: I'SA: Florida. Alachua County. Gainesville, in partly shaded cultivated

field. 12. 5. 1939, leg W. A. MURRILL F 15977 (FLAS, holotype).

The type consists of fragments of altogether about five fruitbodies, the pilei almost

not fragmented

Conocybe fuscimarginata is widely distributed especially also in Europe. Its size

can be much larger than the type specimen, with pilei up to 5 cm wide. There are no

microscopical differences between the numerous European collections investigated by

the first author and the type.

glabra (Fig 11 a-i)

Galerula glabra MURRILL 1917. North American Flora 10: 163

Conocybe glabra (MlRRlLl.) WATLING 1976, Kew Bull 31 593

Original description:

Pileus conic to campanulate. not expanding, solitary, reaching 8 mm broad; surface

moist, not stnate, entirely smooth and glabrous, uniformly dull-isabelline with a ful-

vous tint; lamellae adnate or adnexed, attenuate behind, very regular, not crowded,

somewhat ventricose. dull-isabelline to fulvous, whitish-pruinose on the edges, stipe

slender, equal, smooth, glabrous, concolorous. paler and pruinose at the apex. 2.5 cm

long, 0.5 mm thick.

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Österr. Z. Pilzk. 13(2004)

187

Fig. 11 a-e. (ialerula glabra (holotype, small specimen), a Spores, b basidia, c cheilocysridia, d stipe

covering, e pileipellis. /-/. (ialerula glabra (holotype, larger specimen). /Spores, g basidia. h cheilo-

cystidia, i pileipellis.

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188 A. HAUSKNECHT & al.: Type studies in North American Bolbiliaceae

Microscopical characters:

As already observed by HESLER (in herb), the type consists of fragments of two fruit-

bodies, which clearly belong to two different taxa:

1) Small specimen (Fig 11 a-e):

Spores: 12-14 x 7-8 x 6.5-7(-7.5) urn, mean 12.7x7.6x6.9 um, Q =

1.6-1.8,

ellip-

soidal in front view, oblong-ellipsoidal to slightly bean-shaped in side view, thick-

walled with germ-pore up to 1.5 urn in diam., rust- to orange-yellow in KOH.

Basidia: 4-spored, 18-21 x 10.5-13 (im.

Ammonia reaction: with regard to the very sparse type material not observed

Clamp connections: rare.

Cheilocystidia: lecythiform, 155-20 x 6-9 urn, with capitulum 3.5-5 (im

wide.

Stipe covering: consisting exclusively of lecythiform caulocystidia similar to

cheilocystidia, but often with a longer, more slender neck.

Pileipellis: hymeniform, consisting of roundish-stipitate to broadly clavate

elements, 18-35 x 10-18 urn, in-between numerous lecythiform pileocystidia (similar

to cheilocystidia).

2) Larger specimen (Fig 11 f-i):

Spores: 7.5-9.5 x 4.5-5.5 um, mean 8.7 x 4.8 um, Q =

1.7-2,

ellipsoidal, not lenti-

form, slightly thick-walled with a germ-pore ca

(im in diam., ochre yellow in KOH.

Basidia: 4-spored, 19-21 x 7.5-9.5 (im.

Clamp connections: not observed

Cheilocystidia: 28-56 x 11-16 urn, lageniform. utnform, with long, often re-

peatedly constricted neck and conspicuously capitate apex 5-8 urn wide

Pileipellis: hymeniform, consisting of sphaeropedunculate elements, 20-50 x

15-22 urn, without pileocystidia

Specimen examined: ISA: New York State, Lake Placid, Adirondack Mountains, in soil on the

side of

bank in the open, 17-29. Juli 1912, leg W. A. & EDNA L. MURR1LL (NY)

The collection date indicates that both fruitbodies have been collected on two days

almost two weeks apart. The microscopical features given in the original description

are from the smaller fruitbody, whether the macroscopical description is also from this

is uncertain The smaller fruitbody is a member of sect. Conocybe. closely related with

the European taxa Conocybe semiglobata KUHNER & WATL1NG and Conocybe subxe-

rophytica SINGER & HAUSKN , but with different colours. In addition, the spores of the

American taxon are slightly bean-shaped to irregularly impressed which we have

never observed from European material.

The larger fruitbody is obviously a Pholiotina without velum Pholiotina flexipes

has similar microscopical features but is a member of the species with ring The chei-

locystidia of the European Pholiotina utricystidiata are also very similar, but it also

has an annulate stipe and additionally larger spores.

HESLER (in his unpublished notes) asks the question whether this taxon should be

"excluded" Since it cannot be ruled out that especially colour notes of the larger

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Österr Z. Pilzk 13(2004) 189

specimen have been included into the original diagnosis, we agree with HESLER and

consider Conocybe glabra to be a nomen dubium.

humicola (¥\g lOe-i)

Conocybe antipus (LASCH) FAYOD var. humicola THIERS 1959, Mycologia

51:

529

Conocybe humicola (THIERS) HAUSKN., KRISAI-GREILH. & YOGLMAYR, comb &

stat. nov

Basionym: Conocybe antipus var. humicola THIERS 1959, Mycologia

51:

529

Original description:

Pileus 0.9-1.3 cm broad, conic to subconic, unchanging or becoming convex with age,

margin slightly incurved when young becoming straight when older, entire to eroded;

surface moist, atomate, glabrous, striate to the disk when moist, evenly dark brown to

umber to occasionally brown during all stages of development Flesh thin (c. 2 mm),

brittle, concolorous with the surface: taste and odor not distinctive Lamellae close to

subdistant, ascending-adnate to adnexed, becoming free with age, narrow, thin, light

brown to pale fulvous at maturity; margin entire, concolorous with the gill faces; la-

mellulae of 1-2 lengths Stipe 4-6.5 cm long, 0.5-1 mm broad at the apex, tapering

slightly toward the apex, solid, translucent; surface glabrous except for white fibrils at

the base, moist, olivaceous to yellow during all stages of development.

Microscopical characters:

Spores: 7.5-10 x 6-6.5 x 4.5-5 urn, mean 8.3 x 6.3 x 4.9 um, Q =

1.2-1.5,

in front

view angular-mitriform, in side view ellipsoidal, conspicuously lenticular, with double

wall and a large germ-pore, ochre-yellow in K.OH

Basidia: 2(-4)-spored. 14-18 x 7.5-9.5 um.

Clamp connections: not observed

Cheilocystidia: lecythiform, 13-17 x 6-10.5 urn, with capitulum 3-4.5 urn

wide.

Stipe covering: consisting only of lecythiform caulocystidia similar to the

cheilocystidia but with a longer, more slender neck and a capitulum 2-4 urn wide

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 22-35 x 14-

19 urn; no pileocystidia observed

Specimen examined: USA: Texas, Brazos County, near Wellbourne. in plant debris under oak,

18.

1953.

leg H D THIERS 1883 (MICH, holotype).

The specimen consists of fragments of three to four fruitbodies in rather good con-

dition; the stipe base is apparently not radicating.

Concerning shape and size, the spores of this taxon are within the range of varia-

tion of European collections of Conocybe antipus Besides the differences listed in

THIERS (1959), which concern the colour of pileus and stipe and the missing root,

there is another difference which we consider even more important: the mostly two-

spored basidia which bear spores of the same size As already WATLING & GREGORY

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190 A. HAUSKNECHT & al.: Type studies in North American Bolbiliaceae

(1981:

93) note "probably worthy of specific rank", we propose a new combination at

the species level.

intermedia (Fig 12 a-d)

Pholiota intermedia A H.

SM.

1934, Ann. Mycol. 32: 479

Conocybe intermedia KÜHNER 1935, Le genre Galera: 143

Pholiotina intermedia (A. H. SM.) SINGER 1936, Beih Bot Centralblatt B, 56 170

Pholiota septentnonalis A. H.

SM.

1935, Mycologia 27: 227

Pholiotina septentnonalis (A. H. SM.) SINGER 1945. Notul. Syst. Sect. Crypt. Inst.

Komarov Acad. Sei 5: 98

Original description:

Pileus

-2 cm latus, obtuse conicus demum expansus, glaber, viscidus, hygrophanus,

rufo-fulvus, striatus; lamellae confertae, angustae, adnatae, fulvae, stipes 1-2 cm lon-

gus.

2-3 mm crassus, ochraceus demum fulvus, fibrillosus, annulatus; annulus distans,

reflexus. albus, striatus.

Microscopical characters:

Spores: 6-7.5 x 3.5-4.5 urn, mean 6.8 x 4.0 um, Q =

1.5-1.9,

ellipsoidal, not lenti-

form. partly slightly bean-shaped, thin-walled with a small germ-pore, pale yellow to

yellow in KOH.

Basidia: 4-spored, 14-20x6-8 urn.

Clamp connections: not observed.

Cheilocystidia: "nine pin" shaped, 12-22(-30) x 5-7 urn, with capitulum 2.5-4

urn wide

Pileipellis: hymeniform, consisting of clavate elements, 28-35 x 10-16 urn; no

pileocystidia observed

Specimen examined: USA: Michigan. Emmet County. Harbor Springs. Blisswood. on rotting

maple wood. 19. 8. 1933. leg. A. H. SMITH (MICH, part of holotype).

The part of the type specimen made available to us consists of two complete and a

half fruitbody which are well-preserved. The younger fruitbody has a broad, membra-

naceous ring spanning between pileus margin and stipe

Pholiotina intermedia is very closely related with the European Pholiotina brun-

nea (WAITING) BON. It is widely distributed in North America (especially in Michi-

gan) (WATLING 1971). WATLING (1971) has intensively studied this group and differ-

entiates two taxa, which differ in velum features as well as colour and density of la-

mellae, and we fully agree with him.

intrusus (Fig. 12 e-h)

Cortmarius intrusus PECK 1896, Bull Torrey Bot Club 23: 416

Conocybe intrusa (PECK) SINGER 1950, Sydowia 4 133

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Österr. Z. Pilzk. 13(2004) 191

Original description:

Pileus fleshy, rather thin, convex, then expanded, glabrous, somewhat viscid when

moist, even or radiately wrinkled on the margin, yellowish or

buff,

sometimes with a

reddish tint, flesh white; lamellae thin, close, rounded behind, at first whitish or creamy

white, then cinnamon, often uneven on the edge; stem equal or slightly tapering either

upward or downward, stuffed or hollow, sometimes beautifully striate at the top only

or nearly to the base, minutely floccose when young, soon glabrous, white; pileus 1-2.5 in.

broad; stem 1-3 in long, 3-6 lines thick.

Microscopical characters:

Spores: 5-7x4-5 urn, mean 6.0 x 4.7 urn, shortly ellipsoidal, Q = 1.1-1.35, not len-

tiform. with double wall, completely without germ-pore or callus, orange-yellow in

K.OH

Basidia: 4-spored, 13-17 x 6-8 urn

Clamp connections: present

Cheilocystidia: 14-21 x 6.5-11.5 urn, with capitulum 3-4.5 urn wide.

Stipe covering consisting only of lecythiform elements similar to cheilocys-

tidia

Pileipellis: hymeniform, consisting of roundish-stipitate, almost hyaline to

yellowish elements. 30-40 x 20-28 urn; pileocystidia absent.

Specimen examined: USA: New Jersey. Haddonfield. in mushroom beds, manured soil in con-

servatories or in plant-pots, leg. MC1LVAINE (NYS. holotype).

The holotype consists of ca four fruitbodies in rather good condition.

An extensive documentation of this species which has been recorded from green-

houses, flower cases or pots kept indoor and exceptionally also from gardens outdoor

is given by WATLING (1977). It is the only member of sect. Giganteae SINGER and is

widely distributed but rare at similar habitats also in Europe.

lateritia (Fig.

j-m)

Gastrocybe lateritia WATLING 1968, The Michigan Botanist: 19-24

Original description:

Pileus 5-25 (30) mm high x 2-10 (15) mm broad, ellipsoid-campanulate to conic, long,

subacute, not expanding or hardly significantly so, greasy, tacky or viscid, membra-

nous,

pellucid-striate to disc, especially in immature specimens, then rapidly reduced

to a gelatinous mass which adheres to the apex of the stipe, either retaining former shape or

becoming amorphous, uniform or slightly darker at disk, light chocolate brown, dark-

ening at maturity and in dried material more dark chestnut, even bay, polished. Stipe

50-100 (130) mm x

1-1.5(3)

mm. pure white, shining, hardly discolouring, equal,

tlexuous or straight, slightly attenuated upwards, dry, smooth or sparsely pruinose

(more so towards base) obscurely striate. fragile, easily collapsing, hollow. Gills rich

rust-brown, well-formed with acute margin, branched, forked and interveined, the

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192 A. HAUSKNECHT & al.: Type studies in North American Botbitiaceae

branches fusing with adjacent gill plates, thus forming a fairly regular honeycomb

structure with smaller "locules" nearer the margin of the pileus. Flesh white in stipe,

hardly discernible in pileus; smell and taste not noted.

In grass lawns, gregarious to solitary.

Microscopical characters:

Spores: 10.5-13 x 6-8 urn. mean 11.5 x 7.2 urn, Q = 1.5-1.75, ellipsoidal, not lenti-

form with thin, simple wall and a germ-pore 1-1.5 urn in diam., mature spores yellow

in KOH.

Basidia: 4-spored, 19-28 x 11-14 urn.

Clamp connections: not observed

Cheilocystidia: lecythiform, almost all collapsed, one intact cystidium meas-

uring 14x9 um, with capitulum 4.5 urn wide.

Stipe covering: presumably collapsed, no true caulocystidia observed, only

some capilliform, slightly twisted elements.

Pileipellis: hymeniform, consisting of roundish cells covered by a thick, not

stainable amorphous mucilage. Pileocystidia abundant, lecythiform, 25-30 x 4-6 |im,

with capitulum 4-5,5 urn wide, yellow.

Specimen examined: USA: Michigan, Gratiot County, Ithaca, gregarious on grassy ground in a

cemetery. 31 8. 1947, leg V. POTTER (MICH, holotype).

The part of the type specimen made available to us consists of fragments of one

pileus and two stipes

Gastrocybe lateritia is widely distributed but rare in North America, northern Af-

rica and southern to southeastern Europe.

As demonstrated by HALLEN & al. (2003), the slimy pileus is caused by infection

with bacteria from the Chryseobactenum gleumlmdologenes group. DNA-investiga-

tions unambiguously placed Gastrocybe lateritia into sect Candidae of Conocybe As

Gastrocybe lateritia may be a malformed, already described species of Conocybe, they

did not propose a new combination within Conocybe. In case it proves to be a distinct

species the epithet "lateritia" will have to be changed as it is a later homonym.

liratus (Fig 13 a-d)

Agaricus liratus BERK. & M. A. CURTIS 1860, Proc Amer Acad Arts & Sei. 4: 116

Galera lirata (BERK. & M A. CURTIS) SACC. 1887, Sylloge Fungorum 5: 865

Conocybe lirata (BERK. & M. A, CURTIS) MURR1LL 1912, Mycologia 4: 248

Original description:

{Galera) liratus.

BERK

CURT

: pusillus; pileo umbilicato sulcato atomato rufo,

stipite brevi; lamellis paucis latis adnatis. On the bark of Oak-trees. Mare Island. San

Francisco Bay, California. - Resembles a Marasmius, with the habit of/), corticola.

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Osten Z. Pilzk. 13 (2004)

193

Fig. 12 a-d. I'holioia intermedia (part of holotype). a Spores, b basidia. c cheilocystidia, d pileipellis.

e-h. ( oriinanus mirusus (type), e Spores, /basidia. g cheilocystidia. h stipe covering, ; pileipellis j-m.

Ciasirocybe

laieruia (type).7 Spores, k basidia, / cheilocystidiuin. m pileocystidia.

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194 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

Microscopical characters:

Spores: 12-15 x 7-9 urn, mean 13.2 x 7.8 urn, Q = 1.45-2.1, ellipsoidal, thin-walled,

without germ-pore, pale yellow to yellow in K.OH

s l

a: 4-spored, ca 30 x 10 |im, with sterigmata up to 5 um long.

Clamp connections: present at the base of the basid ia and in the trama.

Cheilocystidia: cylindrical-clavate to roundish, also elongated vesicular. 23-

37x8-14 urn.

Stipe covering: not investigated.

Pileipellis: consisting of horizontal hyphae with erected end cells, 3-4 um

wide; pigment roughly encrusted.

Specimen examined: I'SA: California, San Francisco Bay, Mare Island, on bark of oak, ex Herb.

HOOKER (WRIGHT). 16. 1. 1856 (K, holotype).

The type specimen consists of several fruitbodies of which two are comparatively

well-preserved.

Due to the structure of the pileipellis, Agaricus liratus is not a member of the Bol-

bitiaceae, but belongs to the genus Galehna In the monograph of SMITH & SINGER

(1964) Galera lirata has not been listed; we could also find no species with completely

smooth spores and similarly shaped cheilocystidia. However, we refrain from a com-

bination in Galehna as important macroscopical features are unknown.

locellina (Fig. 13 e-j)

Galerula locellina MURRILL 1946, Lloydia 9: 327

Conocybe locellina (Ml'RRlLL) WATLING 1976, Kew Bull 31: 593

Original description:

Pileus conic, gregarious, about 4 cm broad; surface dry, isabelline, disk darker, margin

closely stnate. upturned at times; context thin, dull-isabelline. mild, odor slightly un-

pleasant; lamellae adnate, close, narrow, pallid, entire; stipe tapering upward from a

clavate base, hollow, white, atomate and pubescent to mostly glabrous, 7-9 x 0.5-1 cm;

volva limb entire, small, white, sometimes persisting.

Microscopical characters:

Spores: 10.5-16 x 8-9.5 x 6.5-8 urn, mean 12.4-12.9 x 8 3-8 5 x 7 2-7.4 um. Q =

1.4-1.6.

ellipsoidal to slightly angular, in front view also slightly navicular. lentiform.

with thick wall and a large germ-pore, rust-orange yellow in K.OH.

Basid ia: 4-spored. 19-26 x 10-13 urn.

Clamp connections: not observed.

Cheilocystidia: lecythiform. 17-25 x 7-10 (am. with capitulum 2 5-4 urn wide

Stipe covering: consisting only of hairs and non-lecythiform elements, hairs

3.5-5 urn wide, lecythiform elements absent.

Pileipellis: hymeniform, consisting of roundish-stipitate elements. 22-40 x 15-

22 urn, in-between commonly capilliform pileocystidia

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Osten. Z. Pilzk. 13 (2004)

19S

Fig. 13 a-d. Agaricus liratus (type), a Spores, b basidium. c cheilocystidia, d pileipellis e-j. (ralerula

locellma (type), e young fhiitbody redrawn from a drawing of MURRILL (x 0.8),/ spores, g basidia, h

cheilocystidia, / stipe covering, j pileipellis.

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196 A. HAUSKNECHT & al.: Type studies in North American Bolbmaceae

Specimens examined: USA: Florida. Alachua County. Gainesville, on exposed sawdust from a

stable, 19. 2. 1945, leg. W. A. MURRILL (FLAS, holotype); - - on manure pile. 19. 9. 1939. leg. W. A.

MURR1LL (FLAS).

The type material is fragmented into numerous pieces, but all microscopical structures

are well-preserved The drawing of a young fruitbody enclosed by MURRILL shows a

wide, membranaceous volva at the stipe base, but no velar remnants at the pileus mar-

gin or at the stipe above the volva.

Conocybe locellina is a member of subgenus Singerella and is closely related to

the European Conocybe hornana SINGER & HAUSKN. Striking macroscopical differ-

ences to the latter are the somewhat more dull pileus colour, the dense striation, the

smooth and never wrinkled pileus surface, no velum residues at the pileus margin and

especially the membranaceous, wide volva at the stipe base. The spores of Conocybe

hornana are approximately of the same size but less lentiform and in front view hardly

navicular.

ludovicianus (Fig. 14a-g)

Gymnopilus ludovicianus MURRILL 1917, North American Flora 10 204

Flammula ludoviciana (MURRILL) KAUFFMAN 1926, Amer J. Bot. 13: 19

Conocybe ludoviciana (MURRILL) WATLING 1976, Kew Bull.

31:

593

Agrocybe ludoviciana (Ml RRILL) HALSKN., KRISAI-GREILH. & VOGLMAYR,

comb,

nova

Basionym: Gymnopilus ludovicianus MURRILL 1917, North Amer. Flora 10: 204

Original description:

Pileus 6-10 cm broad, depressed, pale-yellowish-tan, moist, not viscid, glabrous, mar-

gin non striate. Context white, mild. Lamellae adnate, pale cinnamon, narrow, crow-

ded. Stipe 10-12 cm long, 10 mm thick, white, glabrous, cylindric, often curved, firm

(veil not mentioned)

Microscopical characters:

Spores:

8.5-11

5.5-6.5

x 5-6 urn, mean 9.4 x 6.0 x 5.3 urn, ellipsoidal, slightly

lentiform, in side view also bean-shaped, with double wall and a conspicuous germ-

pore ca

urn in diam.

Basidia: 4-spored, 25-30 x 7-9 urn.

Clamp connections: common in the context.

Cheilocystidia: fusiform, cylindrical with slightly swollen venter to slightly

utriform, 18-40 x 6-11.5 urn.

Pleurocystidia: similar to cheilocystidia, but not that polymorphic, more con-

spicuously fusiform with very acute tips, 22-45 x 8-10 urn

Stipe covering: consisting of short, roundish to irregularly ellipsoidal ele-

ments.

Pileipellis: four-layered, consisting of roundish, oval, fusiform-ventricose to

slightly capitate elements, pigment parietal and slightly encrusting.

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Österr. Z. Pilzk. 13 (2004)

197

Fig. 14 a-g. Gymnopilus ludovicianus (type), a Spores, b basidia. c cheilocystidia, d pleurocystidia, e

stipe covering, /'section of gill trama, g pileipellis.

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198 A. HAUSKNECHT & al.: Type studies in North American Holbiiiaceae

Specimen examined: USA: Lousiana. New Orleans. City Park, on the base of living maple tree

(Acer),

leg. F. S. KARLE. 3. 9. 1908 (NY, holotype).

The type specimen consists of two fragmented fruitbodies in good condition.

The structure of the pileipellis, presence of pleurocystidia and growth at the basis

of a living tree indicate that Gymnopilus ludovicianus is a member of the genus Agro-

cybe. Therefore, a new combination is proposed in Agrocybe.

Galerula magnispora MlJRRlLL 1943, Mycologia 35: 530

Conocybe magnispora (Ml'RRlLL) SINGER 1950, Sydowia

135

Original description:

Pileus convex, not expanding, gregarious, reaching 1.2 cm broad, surface moist, smooth,

glabrous, isabelline. estriate, margin even, entire; context thin, odor not characteristic;

lamellae adnate, broad, ventricose. medium distant, inserted, entire, soon becoming

fulvous; stipe slightly enlarged below, hollow, smooth, glabrous, white, shining. 3-4.5

x 0.12-0.2 cm.

Microscopical characters:

Spores: 15-16.5 x

8.5-11

urn, mean 15.6 x 9.6 um, Q =

1.5-1.7,

ellipsoidal, not len-

tiform, thick-walled with a germ-pore up to

5 urn in diam . orange-brown in K.OH.

Basidia: 4-spored, 25-30 x 12-16 urn

Clamp connections: not observed.

Cheilocystid ia: rather collapsed, ca 12-16 x 6 5-8 urn. with capitulum 2.5-4

urn wide

Stipe covering: consisting only of non-lecythiform elements and hairs, hairs

up to 60 urn long

Pileipellis: hymeniform, consisting of sphaeropedunculate elements, 32-45 x

20-32 urn. not interspersed with pileocystidia.

Specimen examined: USA: Florida Levy County, Gulf Hammock, on old cow dung. 8 3. 1942.

leg.

W. A. MURRILL(FLAS, holotype).

The type material consists of numerous fragments, especially the lamellar edge

with the cheilocystidia heavily collapsed

MURRILL (1943) reports much larger spore sizes than measured by us (15-20 x 10-

12 urn). Concerning the spore characteristics and stipe covering. Conocybe magni-

spora is similar to the European and African Conocybe singeriana, but there are some

important differences. Conocybe singeriana is much larger, more robust, with a strongly

bulbous stipe base, the pileus is darker coloured in fresh condition, moist always stri-

ate and strongly pubescent, the stipe never white Microscopically, the many capilli-

form pileocystidia are conspicuous in Conocybe hornana, in the type of Conocybe

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Österr. Z. Pilzk. 13(2004)

199

Fig 15 a-e. Cialerula magmspora (type), a Spores, b basidia. c cheilocystidia, d stipe covering, e

pileipellis./-/. Pholioiina maireiaffinis (type)./Spores, g basidium. h cheilocystidia, / stipe covering.

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200 A. HAUSKNECHT &al.: Type studies in North American Bolbiuaceae

magnispora and also in two other collections from the USA investigated by the first

author, pileocystidia are always missing.

Conocybe pseudopubescens THOMAS, HAUSKN & MANIMOHAN (THOMAS & al.

2001),

described from India from elephant dung, has similarly delicate fruitbodies like

Conocybe magnispora. but is different in brighter orange to orange white, conspicu-

ously stnate pilei, a similarly coloured, never white stipe and somewhat smaller, much

thicker-walled, in K.OH red-brown spores

Conocybe magnispora was recently reported also from Europe (ARNOLDS & HAUS-

KNECHT2003).

maireiaffinis (Fig 15f-i)

Pholiotina maireiaffinis SINGER 1989, Fieldiana n s 21 106

Original description:

Pileo flavido-ochraceo, hygrophano, siccando pallidiore, haud viscoso, in siccis bre-

viter et debiliter sulcato, glabro, convexo, obtuso, centro interdum depresso, 10-15 mm

lato Lamellis ochraceobrunneis, confertis vel mediocnter confertis, ventncosis, latis,

adnexis Stipite albo, ad apicem minute pruinato, aequali. 8-21 x 1,3-2 mm Velo nullo.

Came alba, interdum in basi stipitis fuscescente, inodora.

Microscopical characters:

Spores: 6 5-8.3 x 4-5 um, mean 7 5 x 4.4 urn, Q =

6-2 0, ellipsoidal, not lentiform,

thin-walled, with small, often hardly visible germ-pore, pale yellow in KOH

Basidia: 4-spored, 18-20 x 6.5-9 urn.

Clamp connections present.

Cheilocystidia: 20-40 x 6-8 urn, cylindrical, slightly lageniform, with rather

obtuse apex, also slightly forked.

Stipe covering: consisting of capilliform. fusiform to clavate elements

Pileipellis: due to the scarce type material not investigated According to SINGER

(1989) the pileipellis is hymeniform, consisting of roundish-stipitate elements, in-

between few pileocystidia similar to cheilocystidia

Specimen examined: I'SA: Illinois. Ogle County. White Pine State Park, on the ground of a for-

est track {(Juenus. Ira.xinus). 9 10 1976. leg. R. SINGER N 7591 (F).

The type consists of

single fragmented fruitbody

According to SINGER (1989) it is close to Pholiotina mairei (WATLING) ENDERLE,

but has larger fruitbodies, not lacrimating lamellae, somewhat broader spores and

other cystidia. The most important difference should be the shape of the cheilocystidia,

which in Pholiotina mairei are always more delicate with long, beak-shaped apex

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Österr. Z. Pilzk. 13(2004)

201

Fig. 16 a-c. Agaricus martianus (type), a Spores, b collapsed cheilocystidia (?), c pileipellis. d-h.

(ialerula mexicana (type), d Spores, e basidia, /cheilocystidia, g stipe covenng, h pileipellis.

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202 A. HAUSKNECHT & al.: Type studies

North American Bolbiliaceae

murtianus (Fig 16a-c)

Agahcus martianus

BERK

& M A

CURTIS 1869,

Linn. Soc. 10:

291

Galera martiana

(BERK

& M A. CURTIS)

SACC.

1887. Sylloge Fungorum 5: 864

Conocybe martiana (BERK

CURTIS) SINGER 1955, Sydovvia 9:

402

Original description:

Helvolus; pileo tenui umbonato piano glabro, stipite gracili leviter sursum deorsumque

mcrassato; lamellis latis ventricosis laete ferrugineis liberis.

Pileus

inch across; stem

inch high, capillary Spores bright ochraceous.

Microscopical characters:

Spores:

7-8.5 x 5-5.5 urn,

mean

7.7 x 5.2 urn,

ellipsoidal with prolonged apiculus,

without germ-pore, sporadically with weak callus, with thick, very wrinkled-uneven,

distinctly calyptrate wall, strongly yellow-brown

K.OH

Basidia:

not

observed.

Cheilocystidia: sparingly very collapsed, cylindrical-clavate

lageniform;

22-26

x 7-8 urn

large elements present.

Stipe covering: entirely collapsed.

Pileus covering:

cutis

thin, lying hyphae with weak incrusting pigment,

no pileocystidia present

Material investigated: Cuba:

rotten wood, June, leg.

WRIGHT no. 70 (K, holotypej.

The type material consists

two separated pilei

and two

stipe fragments

very

poor condition

and is

heavily molded

by a

contaminating fungus.

The

microscopical

analysis given above

and

the following commentary

are

from HAUSKNECHT & KRISAI-

GREILHUBER(1998).

Because

the structure

the pileus covering

and

also

the spore surface this

not

member of the Bolbitiaceae,

but a

representative

the genus Galenna This

was

already stated

DENNIS

(1953:

192),

who

placed

the

species

in the

vicinity

Gale-

rina thscopa (FR.) KUHNER

Due

to the

poor condition

the type material

and the

missing information

cys-

tidial features

consider

combination into Galenna

not to be

meaningful

mexicana (Fig 16d-h)

Ga/erula mexicana MURRILL 1917, North Amer Flora

10: 169

Galera mexicana (MURRILL)

SACC.

TROTT. 1925, Sylloge Fungorum 23:

278

Conocybe mexicana (MURRILL) W'ATLING 1981, Bibl Mycologica

82 120

Original description:

Pileus

3 cm

broad, subhemisphenc,

not

umbonate, moist, uniformely ochroleucous

(yellowish buffish), glabrous,

not

striate. Lamellae adnexed, isabelline. darker

rnatu-

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ÖsterT. Z. Pilzk 13(2004) 203

rity, subcrowded, broad, edges whitish and slightly crenulate Stipe 11 cm long. 3 mm

thick, stramineous, erect, equal, fistulöse

Microscopical characters:

Spores: 12.5-15.5 x 8-9.5 um, mean 14.3 x 8.7 urn, Q = 1.55-1.75, ellipsoidal to

broadly ellipsoidal, sporadically subcylindrical, not lentiform. thick-walled with con-

spicuous germ-pore, orange-brown in KOH.

Basidia: 2-(

1-)spored,

no 4-spored observed, 18-23 x 11-13 um.

Clamp connections: present at the base of the basidia and the trama.

-reaction: negative even after 12 hours.

Cheilocystidia: lecythiform. 12-22 x 6-9 um. with capitulum 2.5-5 urn in

diam.

Stipe covering: consisting mainly of lecythiform caulocystidia, these usually

smaller than the cheilocystidia. 11-19 x 5-9.6 urn. with capitulum 2-4 um in diam :

additionally some short-cylindrical elements present.

Pileipellis: hymeniform, consisting of roundish-stipitate. almost colourless

elements, 30-43 x 19-24 (im; pileocystidia not observed.

Specimen examined: Mexico: Jalapa, in grass by roadside, 12-20. 12. 1909. leg. W. A & E. L.

MURRILL 139 (NY, holotype).

The type specimen consists of one pileus and a stipe fragmented into two pieces in

good condition. The microscopical analysis and the majority of the following commentary

are a translation from HAUSKNECHT (2002 a).

This is the only two-spored member of the Conocybe tenera-group. DENNIS

(1953:

192) doubted that Conocybe mexicana is a separate species and compared it with C.

semiglobata The microscopical description of MURRILL (1917) is too short to infer

additional differences, and microscopically only the two-spored and the more broadly-

ellipsoidal, sometimes subcylindrical spores are evident. Additional, well documented

collections are necessary before a definitive statement about the status can be given.

michiganense (Fig 17 a-e)

Psathyre/la michiganense A. H. SM. 1941, Cont. Univ. Mich. 5: 35

Conocybe michiganensis (A. H. SM.) WATLING 1975, Notes Roy Bot Gard. Edin-

burgh 34:248

Description (from WATLING 1975: 248):

Pileus 10-25 mm broad, obtusely conic and with the margin depressed against the stipe

when young, broadly conic or in age the margin flaring somewhat, moist, when young

minutely pubescent from projecting hairs, soon glabrescent, opaque when young and

moist, only very faintly striate at maturity, varying from sordid "tawny olive" to bistre

or nearly black when the spores mature, sometimes the disk becoming "wood-brown'

(Ridgway 1912) and the margin "avellaneous", hygrophanous atomate when faded,

fading to sordid ashy brownish grey or greyish white, margin regular and non-striate or

folded in age. Stipe 20-50 mm x 1-1.5 mm, equal, strictly rigid, tubular, densely white

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204 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

pubescent (under a lens) at first, soon glabrous or with scattered fibrillose flecks, whi-

tish above, base sordid brown, in a yellowish or sordid brown over all except apex,

base tinged reddish at times, occasionally longitudinally striate over the lower

half.

Flesh very thin and fragile; smell none and taste faintly of radish Gills ascending ad-

nate,

not readily seceding, moderately close (23-37 reach the stipe), moderately broad,

pale avellaneous when young slowly becoming fuscous brown, edges white.

Microscopical characters:

Spores: 9-10 x 6-7 urn, mean 9.5 x 6.5 urn, Q =

1.4-1.6,

ellipsoidal, not lentiform,

thick-walled and with conspicuous germ-pore, violaceous-brown, chocolate-brown in

KOH

Basidia: 4-spored, 19-21 x 9-10 urn.

Clamp connections: not observed.

Cheilocystidia: lecythiform, 15-22 x 7-10 mm, with capitulum 3-4 5 urn in

diam.

Stipe covering: consisting of a mixture of hairs, clavate, cylindrical-ventri-

cose and capilliform-capitate elements.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 32-43 x 16-

30 urn; intermingled with many capilliform pileocystidia.

Specimen examined: ISA: Michigan, Milford, on sawdust, 15. 9. 1939, leg. A. H. SMITH 10920

(MICH, part of holotype).

The part of the holotype accessible to us consists of three well-preserved fruitbod-

ies.

Conocybe michiganensis has been originally placed into Psathyrella (SMITH 1941)

due to the "fuscous" spore colour, which is highly exceptional for a member of Bolbi-

tiaceae WATLING (1975) investigated several collections of this species comparatively

common in Michigan and combined it into Conocybe. He placed it into a new section

Conocybella (SlNGHR) WATUNU, with Conocybe michiganensis as type species. The

species has up to date only been recorded from North America

microspora (Fig 17f-i)

Pholiotina microspora SINGER 1989, Fieldiana n s 21: 107

Original description:

Pileo pallide brunneo, in umbone brunneo, hygrophano, leniter pellucide striato in hu-

midis,

glabro, nudo, convexo, umbonato, ± 13 mm lato. Lamellis ochraceis, confertis,

mediocnter latis, rotundato-adnatis. Stipite albo, ad basin mox brunneo, ad apicem

fannoso, subaequali, haud bulboso, 20 x 1,5 mm. Velo haud appendiculato in margine

pilei sed annuliformi; annulo membranaceo, fugaci. Carne inodora. in basi stipitis

brunnea.

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Osten. Z. Pilzk. 13(2004)

205

Fig. 17 a-e. I'salhvrella michiganense (type) a Spores, h basidia. c cheilocystidia. J stipe covering, e

pileipellis./-/. l'holiolina microspora (type)./ Spores, g basidia. h cheilocystidia. / pileipellis.

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206 A. HAUSKNECHT & al.: Type studies in North American Holhiiiaceae

Microscopical characters:

Spores: 6-8 x 3.5-4 um, mean 7.0 x 3.9 um, Q =

1.5-1.9,

ellipsoidal, not lentitbrm,

thin-walled, without germ-pore, at most with callus, pale yellow in KOH.

a s

a: 4-spored, 15-21 x 5-7 urn.

Clamp connections: present

Cheilocystidia: 21-55 x 4-8 (im, cylindrical-lageniform with a long neck and

an obtuse, rarely slightly capitate apex, partly with a slime cap

Pileipellis: hymenitbrm. consisting of roundish-stipitate elements, 19-28 x 12-

20 urn. no pileocystidia observed; pigment roughly encrusted.

Specimen examined: I'SA: Illinois. Cook County. Harms Woods, on dead stump of deciduous

tree

(Acer

Ouenus),

16. 10.

1983.

leg. R. SINGER

5147 (F, holotype).

The type specimen consists of two well-preserved specimens, the ring-shaped ve-

lum is not present any more.

SINGER (1989) described the spore features as "poro germinativo truncato instruc-

tis'\

"dilute ferruginascente-ochraceo-brunneis". This disagrees with our observations

(see above).

Pholiotina microspora has been considered by SINGER (1989) to be closely related

with Pholiotina intermedia (presumably because both are lignicolous), but the cheilo-

cystidia have a completely different shape. We consider it to be similar to a delicate

Pholiotina arrhenn with a short stipe, but it has lighter coloured spores without a

germ-pore (in P. arrhenii a germ-pore is always present, although it may be small),

and also somewhat differently shaped cheilocystidia Further collections from North

America are necessary whether these differences are sufficient to separate these two

taxa

neoantipus (Fig 18 a-j)

Galerula neoantipus G F ATK 1918. Proc Amer Phil Soc 57 371

Conocybe neoantipus (G F. ATK.) SINGER 1936. Ann Mycol 34 433

Original description:

Gregaria 3-7 cm alta; radix 2-5 cm; pileo campanulato-convexo,

1,5-2,5

cm lato; la-

mellis cum in Galerula antipus

Cystidiis solum in acie lamellarum. numerosis, lecythiformibus, 15-30 x 6-9 u;

sporis quaternis. ellipsoideis, 12-17 x 6-9 u

Microscopical characters:

The type of

neoantipus consists of three partly heavily fragmented fruitbodies (WAT-

LING 1986: 92 recorded "two fragmentary basidiomes and several pieces of stipe, un-

doubtedly previously rooting"). These parts can with some troubles (usually with the

differently sized spores which are on the stipe surface) be attributed to two if not three

different taxa

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österr. Z. Pilzk. 13(2004)

207

Fig. 18 (ialemla neoanlipus (labelled with red ink as "(ialenila neoantipus ATKINSON n. sp type").

a-c large pileus a Spores, b basidia. c cheilocystidia cl-g. medium-sized pileus with stipe fragment.

d Spores, i' basidia. /cheilocystidia. # elements of stipe covering /;.;. small pileus with stipe fragment.

h Spores. ; elements of stipe covering,

elements of stipe covering from light-coloured stipe fragment

(presumably belonging to large pileus).

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208 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

1) large pileus (specimen very pale, stipe fragments rooting) (Fig 18 a-c,j)

Spores: 11-13(-14.5) x 7-9 x 6.5-7.5 urn, mean 12.3 x 8.0 x 7 3 urn, Q =

1.5-1.8,

ellipsoidal to inconspicuously angular, lenticular, with wall 1 urn thick and a germ-

pore 1.5-2 urn in diam , yellowish brown in K.OH.

Basidia: 4(2)-spored, 16-23 x 11-13 um.

Clamp connections: not observed

Cheilocystidia: lecythiform, 20-25 x 7.5-11 urn. with capitulum 3.7-6 urn in

diam.

Stipe covering: strongly collapsed, but evidently consisting of lecythiform

caulocystidia with a thin venter and a captitulum 4.5-5.5 um wide, and with hairs up to

55 x 3 urn.

Pileipellis: hymeniform, consisting of roundish-stipitate elements.

2) medium-sized pileus (root not traceable, specimen much darker than large pi-

leus) (Fig 18 d-g)

Spores: 11-14.5 x 7.5-8.5 x 7-8 um, mean 12.8 x 7.8 x 7.6, Q =

1.5-1.8,

distinctly

ellipsoidal and not lentiform, with wall 0.5-1 urn thick and a germ-pore

1.5-1.8

um in

diam., distinctly yellow-brown in KOH.

Basidia: 4-spored, 18-28 x 12-14 urn

Clamp connections: not observed.

Cheilocystidia: lecythiform, 17-23 x 6.5-9 urn, with a capitulum 3.7-5 urn in

diam.

Stipe covering: consisting only of lecythiform caulocystidia and some round-

ish to cylindrical elements, capitulum of caulocystidia 3 5-5 urn in diam

Pileipellis: hymeniform, consisting of roundish-stipitate elements.

3) small pileus (darker coloured than 2, root not traceable) (Fig 18 h, i)

Spores: 10.5-11.5 x 7-7.5 x 6.5-7 urn, mean 10.9 x 7

x 6.7 um, Q =

1.4-1.7,

only

slightly lenticular and indistinctly angular, with wall up to

urn thick and a germ-pore

1.5-2 (im in diam , distinctly yellow-brown in KOH.

Basidia: strongly collapsed, ca 17 x 13 urn.

Clamp connections: not observed

Cheilocystidia: lecythiform, ca 15-18 x 6-8 urn, with capitulum 3.5-4 urn in

diam

Stipe covering consisting almost only of lecythiform elements with capitu-

lum 3.5-4.5 urn in diam., capilliform elements not traceable

Pileipellis: hymeniform, consisting of roundish-stipitate elements.

Specimen examined: USA: Vermont, Middleburg, on newly seeded lawn, leg. E. A.

BURT

(CUP,

holotype).

The microscopical analysis given above and most of the following commentary are

a translation from HAUSK.NECHT (19%). From the above data it is apparent that the type

consists of two or even three different taxa. This is not amazing as the habitat is char-

acterized as "newly seeded lawn", as we know that in such habitats numerous species

of Conoybe can grow simultaneously and intermingled Comparing the results given

above with the original diagnoses it is evident that the spore size strongly diverges

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Österr.

Z. Pilzk. 13 (2004) 209

from that given in ATKINSON (1918) Therefore, Conocybe neoantipus is considered as

not interpretable and rejected as a nomen dubium

pinguis (Fig. 19a-d)

Conocybe pinguis WATLING 1971, Persoonia6: 338

Pholiotina pinguis (WATLING) ENDERLE 1997, Z Mykol 63 32

Original description:

Pileus 15-35 mm. convexus postremo planus, primo viscidus, striatus glaber, castaneus

postremo ochraceo-fulvus vel sordido-fulvus. Stipes 70-90 x 3-4 mm. aequalis ad ba-

sim leviter incrassatus, farctus, ad apicem albidus, ad basim obscuriore ochraceo-

brunneus ad apicem fibrillosus vel fibrilloso-squamulosus; annulus crassus, membra-

naceus, apicalis, supra striatus. Lamellae confertae L 30

1-3, adnatae. bubalinae vel

ochraceo-fulvae. Caro tenuis, concolora.

Microscopical characters:

Spores: 7-8.5 x 4-5.5 urn, mean 76 x 4.7 um, Q =

1.4-1.9,

ellipsoidal to slightly

pip-shaped, thin-walled, with very inconspicuous to missing germ-pore, often only

slight callus present, yellow in KOH.

Basidia: 4-spored. 18-22 x 8-11 urn.

Clamp connections: present

Cheilocystidia: 20-45 x 6-12 urn, very polymorphic, cylindrical-ventricose,

rarely cylindrical or slightly clavate, also twisted, very often with capitate apex (6-10

urn).

Pileipellis: hymeniform, cosisting of pynform elements with a long pedicel,

often also irregular or unilaterally flattened (like a golf-club), 30-48 x 7.5-17 urn. in-

between cylindrical-ventricose pileocystidia.

Specimen examined: I'SA: Washington, Jefferson County, Olympic NP, Clearwater River, un-

der Alnus

debris,

9. 5. 1939, leg. A H.

SMITH

13260

(MICH,

holotype).

The type specimen consists of one fragmented fruitbody, the ring is not visible any

more.

Conocybe pinguis has spores almost identical with Pholiotina arrhenii (FR) SINGER,

but cheilocystidia of a completely different shape Also the rather small and often pe-

culiarly shaped elements of the pileipellis are conspicuous. WATLING (1971) also

gives the glutinous pileus and the very conspicuous ring as species-specitlc characters.

In Europe there are no similar species, therefore Conocybe pinguis is a separate, well-

defined species.

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210 A. HAUSKNECHT & al: Type studies in North American Bolbiliaceae

plicatellus (Fig. 19 e-h)

Agaricus coprinoides PECK 1873, Bull. Buffalo Soc Nat. Sei. 1: 52, non Agaricus co-

prmoides

CORDA

1831

Agaricus plicatellus PECK 1878, Ann Rep. New York State Mus. Nat. Hist. 29: 66

Galera plicatella (PECK) EARLE 1903. Torreya 3: 136

Galerulaplicate/la (PECK) MURRILL 1917, North Amer Flora 10: 164

Conocybeplicatella (PECK) KÜHNER 1935. Le genre Galera: 137

Galerellaplicatella (PECK) SINGER 1951 „1949", Lilloa 22: 490

Original description:

Pileus membranaceus. soon expanded, often split on the margin, plicate-sulcate to the

small even disk, yellowish, inclining ochre; lamellae close, slightly rounded behind,

concolorous, stem equal, hollow, minutely hairy-pruinose, white; plant 1' high, pileus

6' broad, stem ,5" thick.

Microscopical characters:

Spores: 7-8.5 x 5-6 x 4 5-5 urn, mean 7 4 x 5 5 x46 um, Q= I 3-1 45, in front view

ventricose-ellipsoidal. rarely scarcely angular, in side view oblong ellipsoidal, with re-

latively thin wall and conspicuous germ-pore, yellow-brown in KOH.

Basidia: 4-spored. 18-20x8.5-10 5 urn

Clamp connections: common in the trama.

Cheilocystidia: cylindrical, often with a ventricose basis and a long, slender

neck, with blunt, rarely slightly capitate apex. 20-50 x 6 5-10 urn

Stipe covering: consisting of some elements similar to the cheilocystidia.

Pileipellis: hymeniform, consisting of pynform to sphaeropedunculate ele-

ments. 24-43 x 11-21 urn; no pileocystidia observed

Specimen examined: I'SA: New York, Sterling, on grassy ground, without date (NYS, holo-

type)

The type consists of five fruitbodies partly glued on paper in well-preserved con-

dition.

Galerella plicatella is the type species of the genus Galerella (HORAK 1968. SINGER

1986).

This genus is considered to be a subgenus of Conocybe by some authors, e.g..

WATLING & GREGORY (1981) or WAITING (1982) Recent investigations (ARNOLDS

& HAUSKNECHT 2003) have shown that this is based on a misinterpretation of Euro-

pean collections, which have recently been described as Pholiolina sulcata ARNOLDS

& HAUSKNECHT Therefore, the genus Galerella with its type species Galerella pli-

catella is considered and retained as an independent genus (HORAK & HAUSKNECHT

2002,

HAUSKNECHT & CONTU 2003).

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Osten.

Z. Püzk 13(2004)

211

Fig. 19

a-d.

('onocyhe

pmguis (type), a Spores, b basidia. c cheilocystidia. d pileipellis. e-h. Agaricus

plicaiellus (type), e Spores, /basidia, g cheilocystidia. /; pileipellis.

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212 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

plumbeitincta (Fig. 20 a-e)

GalerulaplumbeitmctaG. F ATK 1918, Proc Amer. Phil. Soc. 57: 372

Conocybeplumbeitincta (G F ATK.) SINGER 1950, Sydowia 4: 137

Pholiotina plumbeitincta (G. F. ATK.) HAUSKN., KRISAI-GREILH. & VOGLMAYR,

comb,

nova

Basionym: Galerula plumbeitincta G F. ATK. 1918, Proc. Amer. Phil. Soc. 57: 372

Original description:

Gregaria, 3-5 cm alta; pileo convexo, dein campanulato, 1-1,5 cm lato, adolescente

lubnco, substriato, plumbeitincto:, lamellis stipite late adnatis, subdistantibus, ventrico-

sis,

ochraceo-cinnamomei, stipite albo dein plumbeitincto, sursum pruinoso, deorsum

substriato, cavo, 2-3 mm crasso.

Microscopical characters:

Spores: 12-15 x 7-9 urn, mean 13.4 x 7.9 um, Q =

1.6-1.8,

distinctly ellipsoidal, not

lentiform, thick-walled with a large germ-pore, orange-brown in KOH.

Basidia: 4-spored, 20-24 x 11-13 urn.

Clamp connections: not observed.

Cheilocystidia: predominantly lagemform with broad, distinct beak, some-

times subcylindrical-ventricose, some cystidia with capitulum, 17-27 x 7-10 urn.

Stipe covering consisting of cylindrical to cylindrical-ventricose elements, up

to 80 urn long and 17 urn wide.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, up to 45 x

35 urn. intermingled with many pileocystidia similar to the cheilocystidia, but larger.

Specimen examined: USA: Michigan, Ann Arbor, Cascade Glen, on dung hills, leg. C. H.

KAUFFMAN 565 (CUP, holotype).

The type specimen consists of hundreds of small fragments, nevertheless, hy-

menium, stipe and pileus structures are well preserved.

In the original diagnosis the cheilocystidia are described as "ampullaeformes, fre-

quenter subcapitatae" (ATKINSON 1918), which led SINGER (1950: 137) to combine

this species into Conocybe, presumably based on material from Europe which was

later conceded by him (SINGER 1959). He then investigated the type specimen (SINGER

1959:

396) and described the cheilocystidia as "19.3-21 5 x 5-7.5 urn, varying from

ampullaceous to capitate, many subcapitate. few truly capitate as in most species of

Conocybe and few truly ampullaceous and non-capitate without any thickening in the

apical region". However, in the following discussion he considers it arguable to retain

it as an independent species within the genus Conocybe. Therefore, for a long time all

European Conocybe collections with a more or less sordid brown to grey or blackish-

grey pileus were identified as Conocybe plumbeitincta. This name was partly used for

Conocybe mosen WATL1NG up to the 1990ties (RAITHELHUBER 1991: 219).

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Österr. Z. Pilzk. 13(2004)

213

Fig. 20 a-e. Galerula plumbeilincla (type), a Spores, b basidia, c cheilocystidia, d stipe covering, e

elements of

pileipellis.

/-;. Galerula procera (type). /Spores, g basidia, h cheilocystidia, / stipe cover-

ing.

214 A. HAUSKNECHT & al.: Type studies in North American Bolbiiiaceae

The microscopical features of the species undoubtedly place it into the genus Pho-

liotma, very close to Pholiotina coprophila (K.UHNER) SINGER. Microscopically both

taxa are scarcely discernible, but Pholiotina coprophila has completely different pileus

and stipe colours and a never striate pileus We therefore propose a combination into

Pholiotina.

procera (Fig. 20 f-i)

Galerulaprocera G F

ATK.

1918, Proc Amer Phil. Soc. 57: 372

Conocybe atkinsonii WATLING 1981, nom now, Bibliotheca Mycologica 82: 9

Original description:

Gregaria, 10-12 cm alta; pileo campanulato. obtuso, ad marginem demum leniter ex-

panso, 3-4,5 cm lato, 2-2.5 cm alto, ochraceo-fulvo. dein ochraceo vel pallide-ochro-

leuco,

glabro, non stnato, lamellis stipitis adnexis, ellipsoideis, ochraceis; stipite ae-

quali, ad basem leniter crasso, recto vel flexuoso, glabro. striato. sursum pruinoso. pi-

leo concolore sed pallidiore, 3-4 mm crasso.

Microscopical characters:

Spores: 14-18 x 9-10.5 x 8-10 urn, mean 15.9 x 9.5 x9.1 um, Q =

1.6-1.8,

ellipsoi-

dal,

slightly lentiform, not angular, with wall more than 1 urn thick and a large, trun-

cate germ-pore, reddish-brown in K.OH

Basidia: 4-spored, 20-24 x 12-14 urn.

Clamp connections: not observed.

Cheilocystidia: lecythiform, 15-21 x 6-11 urn, with capitulum 4-6.5 urn in

diam.

Stipe covering: consisting of

hairs,

cylindrical, cylindrical-ventricose or roundish

elements, no lecythiform caulocystidia present.

Pileipellis: hymeniform, consisting of roundish-stipitate elements.

Specimen examined: I'SA: New York. Ithaca. Buttermilk Gorge, on ground, in humus among

leaves in moist woods. 23 7. 1902. leg. C. H. KAUFFMAN 9910 (CUP. holotype)

The holotype consists of three fruitbodies in excellent condition.

WATLING (in WATLING & GREGORY 1981) proposed a new name for that species

as the epithet "procera" was already assigned in Conocybe to another species [Cono-

cybe procera (SINGER) WATLING, see WATLING & GREGORY 1981: 127]

The spore and stipe covering features of Conocybe atkinsonii are almost identical

to Conocybe singeriana. The latter has more robust fruitbodies with a thicker stipe and

a large bulbous stipe base; its pileus is always distinctly striate in moist condition, the

cheilocystidia are somewhat smaller with a distinctly smaller capitulum, and it grows

predominantly on dung and not on leaf litter in moist forests Conspecificity cannot be

ruled out at present, however, additional we 11-documented American collections from

similar habitats are necessary before variability of the macroscopical features of Cono-

cvbe atkinsonii can be assessed.

Österr. Z. Pilzk. 13(2004)

215

Fig. 21 a-e. (ialera pulchra (type), a Spores, b basidia. c cheilocystidia, d stipe covering, e pileipellis.

J-i.

(.ialera

reiiculaia

(type),

j Spores, # basidia, h cheilocystidia.; pileipellis.

216 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

pulchra (Fig 21 a-e)

Galera pulchra CLEM. 1896, Bot. Survey Nebraska 4: 22

Conocybe pulchra (CLEM.) KÜHNER 1935, Le Genre Galera: 137 inval (no basionym

cited)

Conocybe pulchra (CLEM.) HAUSKN., KRISAI-GREILH. & VOGLMAYR, comb, nova

Basionym: Galera pulchra CLEM. 1896, Bot. Survey Nebraska 4: 22

Description (from SACCARDO & SYDOW 1899: 142):

Pileo conico, lato, membranaceo, centrum versus striato-sulcato, minute denseque

scenceo-tomentoso; umbone distincto, ochraceo, stipite elongato cartilagineo, attenu-

ato,

fistuloso, longitudinaliter lineato-stnato-pruinoso, ochroleuco; lamellis adnexis,

angustis. linearibus, lentiter curvulis, ochraceis.

Pileus 2

cm latus, 2 cm altus; stipes 7-8 cm longus, 2 mm crassus.

Hab.

ad terrain humosam, Nebraska Amer. bor

Microscopical characters:

Spores: 12.5-15 x

8.5-9.0

x 7.5-8.5 urn, mean 14.0 x 8.7 x 7.8 um, Q =

1.5-1.7.

front view ellipsoidal-submitnform, also slightly angular, in side view ellipsoidal, with

thick wall and conspicuous germ-pore, slightly to distinctly lenticular, rusty-orange in

KOH

Basidia: 4-spored, 20-30 x 12-16 um

Clamp connections: present.

Cheilocystidia: lecythiform, 18-22 x 8-10 urn, with capitulum 4-6 urn in

diam

Stipe covering: consisting only of hairs and non-lecythiform elements; also at

the apex of

the

stipe no lecythiform caulocystidia present

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 25-55 x 18-

30 urn, pedicel 10-30 urn long, with encrusted pigment; pileocystidia not observed.

Specimen examined: USA: Nebraska, Lincoln, Otowanie Woods. Aug 1890, leg. J. E. CLE-

MENTS (NEB. holotype).

The holotype consists of one fruitbody, the pileus of which is broken into two

fragments in very good condition. The specimen is of conspicuous brightly ochre-yel-

low colour

Unfortunately, we have not been able to obtain the original diagnosis, as we could

not locate the journal in European libraries.

Conocybe pulchra is a member of

sect.

Pilosellae and should be close to Conocybe

velutipes. However, it has substantially larger spores which are lenticular as in Cono-

cybe velutipes, but in front view they are never regularly ellipsoidal but submitnform

and sometimes slightly angular. In addition, the cheilocystidia, basidia and the ele-

ments of the pileipellis are larger than in the much more delicate European Conocybe

velutipes. Also the bright colour of the exsiccate, which we have never observed in

Conocybe velutipes, is conspicuous Unfortunately, the macroscopical description is

very scanty, and additional collections of Conocybe pulchra are necessary to investi-

Osten. Z. Pilzk.

(2004) 217

gate whether there are also differences in colour and striation of the pileus and whether

the microscopical features given above are constantly different from Conocybe veluti-

pes.

At the moment we consider Conocybe pulchra to be a well separated species and

therefore propose a combination into Conocybe

reticulata (Fig 21 f-i)

Galera reticulata PECK 1901, Ann. Rep New York Stat. Mus. Nat Hist 54: 150

Conocybe reticulata (PECK) WATL1NG 1977, Kew Bull. 31: 593

Original description:

Pileus 1-2.5 cm broad, hemispheric or campanulate, obtuse, hygrophanous, cinnamon

color when moist, creamy yellow or buff when dry, rugosely reticulate Context white

or whitish, thin, fragile. Lamellae ascending, yellowish, becoming bright ferruginous,

close, narrow Stipe 2.5-6 cm long, 2-4 mm thick, white, pruinose, apex striate. equal,

hollow.

Microscopical characters:

Spores: 7-9 x 3.5-5 urn, mean 7.7 x 4

um, Q =

1.76-2,

ellipsoidal, slightly pip-

shaped, not lentiform, thin-walled with germ-pore ca

urn in diam., yellow in K.OH.

Basidia: 4-spored, 15-18 x 7-9 urn

Clamp connections: not observed

Cheilocystidia: fusiform, towards the tip gradually tapered, 20-40 x 3.5-8 urn.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 25-35 x 13-

19 urn, intermingled with numerous pileocystidia similar to the cheilocystidia, but up

to 50 x 11.5 urn.

Specimen examined: USA: New York, Ithaca, on mossy ground. November

(NYS,

holotype).

The type specimen consists of four fruitbodies glued on paper in good condition.

All macro- and microscopical features of Galera reticulata (probably except the

strongly rugose pileus surface) are in perfect agreement with the European Naucoha

striaepes COOKE, described 1885 from Great Britain We therefore consider this taxon

for a later synonym of Pholiotina sthipes (COOKE) M. M. MOSER

rugosa (Fig. 22 a-d)

Pholiota rugosa PECK 1897, Ann. Rep. New York State Mus. Nat. Hist. 50: 102

Pholiotina rugosa (PECK) SINGER 1946, Pap Mich Acad Sei 32 148

Pholiotina filaris (FR ) SINGER var rugosa (PECK) SINGER 1950, Acta Inst Bot. Ko-

marov Acad. Sei. URSS, Ser. 2,6: 429

Conocybe rugosa (PECK) WATLING 1981. Bibl. Mycol 82: 133

218 A. HAUSKNECHT & al.: Type studies in North American Bolbiiiaceae

Original description:

Pileus thin, broadly conical or campanulate becoming expanded and often umbonate,

hygrophanous, yellowish-red or ferruginous and striatulate on the margin when moist,

pale yellow or buff and commonly rugose when dry. lamellae close, adnexed, yel-

lowish-white or cream-colored becoming ferruginous or brownish-ferruginous with

age,

white and minutely denticulate on the edge; stem flexuose, equal or slightly thick-

ened toward the base, hollow, fibrillose or sometimes squamulose below the annulus,

pruinose or mealy above, pallid, the annulus membranous, white or whitish, radiately

striate on the upper surface.

Pileus 6 to 12 lines broad; stem

to 2 in. long,

to 2 lines thick.

Microscopical characters:

Spores: 9 5-115 x 5-6 um, mean 10.1 x 5.6 urn, Q = 1 55-2.0, ellipsoidal, not lenti-

form, with slightly double wall and a large germ-pore, ochre-yellow to ochre-brown in

KOH

Basidia: 4-spored, 19-26 x

8.5-11

Clamp connections: present at the base of

the

basidia and in the trama.

Cheilocystidia: cylindncal-lageniform with long neck and a blunt, rarely

slightly capitate apex. 20-40 x 7-12 um.

Pileipellis: hymeniform, consisting of roundish-stipitate elements. 40-55 x 22-

35 urn; pileocystidia not observed.

Specimen examined: USA: New York, Adirondack Mountains, on ground among decaying

chips,

September, leg. C. H. PECK (NYS, holotype).

The type specimen consists of six to seven partly fragmented fruitbodies in good

condition.

Pholiotina rugosa is very closely related with Pholiotina filaris, many authors

consider both taxa conspecific. Differences may concern the larger fruitbodies and

spores of Pholiotina rugosa. the wrinkled pileus surface should not be a good distin-

guishing feature However. European specimens of Pholiotina filans investigated by

the first author are in no way homogeneous, as there are clearly two different types of

cheilocystidia. one type being very similar to Pholiotina rugosa. Whether the shape of

cheilocystidia is correlated with other features like size of fruitbodies or spores is yet

unclear We therefore prefer to maintain two separate species until more data are avail-

able.

siligineoides (Fig. 22 e-i)

Conocybe siligineoides R. HEIM 1957, Rev Mycol Paris n s 22: 197

Original description:

Pileus

1,3-2.3

cm latus, 0,9-1,9 cm altus, primum subhemisphaericus, dein conico-cam-

panulatus. numquam extensus, colore pulchro fulvo aurantiaco rufo. glaber impoli-

tusque, cum disco vix obscuriore, paulo plus aurantiaco, margine regulanter leviterque

Österr. Z. Pilzk. 13(2004) 219

crenulato, arete albido. oris breviter sed clare striatis; hygrophanes. Stipes longus.

gracilis, rigidus, aequalis, haud aut vix ad basim inflatus, 2-4.5 x 0,15 cm, albus pau-

loque farinosus ad apicem aurantiacus pallidus ad culmen. clarescens per aetatem.

semper ad basim albus, stricte fistulosus; crescens etiam postquam pileus prope finem

incrementi invenit. Lamellae potius distantes spissioresque. subadnexatae, stnctae.

lamellulis inaequalibus comitatae, concolores (brunneae-fulvae-subaurantiacae). Caro

exilis,

dilucida praesertim in pileo, fere sine colore, sapore subdulci. dein vix acres-

centi

Microscopical characters:

Spores: 12-14.5 x 7.5-9 um, mean 13.8 x 8.3, Q=

5-1.85, ellipsoidal, sporadically

slightly angular to subhexagonal, not lentiforrn. with a thick wall and a germ-pore up

to 1.5 urn in diam., olive yellowish-brown in KOH (colour probably affected by stor-

age in fixation liquid).

Basidia: 2-spored, 16-20x9-11 urn.

Clamp connections: not observed

Cheilocystidia: lecythiform. 11-16 x 7-10 urn, with capitulum 3-4 urn in

diam.

Stipe covering: consisting of a mixture of capilliform, non-lecythiform and

lecythiform elements, the latter similar to cheilocystidia, but smaller, 12-16 x 5-7 urn,

with capitulum 3-4 urn in diam

Pileipellis: hymeniform, consisting of relatively small, roundish-stipitate ele-

ments, 19-23 x 9-13 urn, in-between sporadical capitate pileocystidia similar to the

cheilocystidia, but with smaller venter.

Specimen examined: Mexico: Huantla de Jimenez. Rancho de Tenango, on living stumps of

Saurama spec , end of June until 7. 7 1955. leg. R G. & V. P. WASSON (PC. holotype).

The type specimen is stored in a flask with fixation liquid and consists of two well-

preserved fruitbodies. The microscopical analysis and the following discussion are a

translation from HAUSKNECHT (2003).

The structure of the stipe covering not mentioned in the original description une-

quivocally places this species into sect. Mixtae. Within that section, Conocybe mer-

daha is quite similar in some features However, the spores of

Conocybe

siligineoides

deviate in being slightly angular in front view, and the pileipellis consists of the small-

est elements we have up to date been able to observe in Conocybe, in addition, the

habitat, a living stump, is completely different.

According to STAMETS (1999), this species has been used by the Aztecs for sha-

manistic purposes and should contain psilocybin. This could be further indication to

consider Conocybe siligineoides as a separate species. Considering the present know-

ledge conspeeifity with Conocybe merdaha appears very unlikely and is therefore re-

jected.

smithii

see under cyanopes

220

A. HAUSKNECHT & al : Type studies in North American Bolbiiiaceae

Fig. 22 a-d. Pholioia rugosa (type), a Spores, b basidia. c cheilocystidia. d pileipellis. e-i. ( onoc.be

siligineoides (type), e Spores,/basidia, g cheilocystidia, h stipe covering.; pileipellis.

Österr. Z. Pilzk. 13(2004)

221

Fig. 23 a-c.

onocybe siercoraria (part of type), a Spores, b basidia c cheilocystidia. d-g. Galerula

subcnspa (type), d Spores, e cheilocystidia (mostly collapsed), /stipe covering, g pileipellis. h-k. Co-

nocybe Jibrillosipes (type), h Spores, / basidia, / cheilocystidia, k pileipellis.

A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

stercoraria (Fig 23 a-c)

Conocybe stercoraria WATLING 1971, Persoonia 6:334

Pholiotina stercoraria (WATLING) ENDERLE 1997,7. Mykol. 63: 32

Original description:

Pileus 10-35 mm e convexo planus, humidus, hygrophanus, glabrus. fulvus vel ochra-

ceo-brunneus. jove sicco pallido-flavidus vel incarnato-ochraceus, ad marginem mter-

dum residuis veli obtectus. Stipes 50-60 x 1,5-2 mm, aequalis, cavus, fragilis, ochra-

ceo-brunneus vel umbrmus. fibrilloso-striatus vel subglaber; annulus fugaceus. mem-

branaceus. medius, albidus, instriatus infra flocculosus. Lamellae L 19-22, I 1-3 adna-

tae,

albidae postremo terrugineo-mellinae. subconfertae. Caro ochraceo-fulva vel ochra-

ceo-brunnea jove sicco ochracea ad basim umbnnus

Microscopical characters:

Spores: 9-105 x 5-5.5 um, mean 9.7 x 5.2 iim, Q = 18-2, ellipsoidal-pip-shaped,

not lentiform, with double wall and conspicuous germ-pore ca 1 um in diam., orange-

yellow in K.OH

Basidia: 4-spored, 20-24 x

8-10.5

um.

Clamp connections: present.

Cheilocystidia: ventricose, with blunt to capitate apex, also utriform, 25-32 x

8.5-10.5

urn.

Stipe covering: above the ring similar to the cheilocystidia.

Pileipellis: hymeniform. consisting of roundish-stipitate elements

Specimen examined: USA: Washington. Hok River near Spruce. Olympic NP.. scattered on

horse dung. 17. 5. 1939. leg. A. H. SMITH 13169 (MICH, holotype).

The specimen labeled "part of

type"

consists of one well-preserved fruitbody

The close relationship with Pholiotina utricystidiata is apparent. E-NDERLE & HUB-

NER (1999) list as distinguishing features for Pholiotina stercoraria a more fugaceous,

on the upside non-stnate ring, somewhat shorter and minimally lentiform spores,

smaller basidia and narrower, more elongated cheilocystidia as well as growth on

dung.

As Pholiotina utricystidiata has meanwhile been found several times in Europe

and also India (THOMAS & al. 2001) and more is known about their variability, some

of these features have lost their distinguishing character, e.g., the size and shape of

basidia and cheilocystidia, as well as the spore size. Therefore, the only features dis-

tinguishing both species are the features of the ring and the habitat. However, an addi-

tional feature concerns the spores, which in light microscopy are consistently darker in

all collections of Pholiotina utricystidiata than in P. stercoraria Therefore, also in this

case additional American specimens are necessary to clarify the variability of Pholi-

otina stercoraria, before a final judgment about eventual conspecificity of both species

can be given.

Österr Z. Pilzk. 13(2004)

223

Fig. 24 a-f. Agahcus sulcalipes (type), a Spores, b basidia. c cheilocystidia J caulocystidia. e pilei-

pellis,/pileocystidia.

224 A. HAUSKNECHT & al.: Type studies in North American Bolbiliaceae

subcrispa (Fig 23 d-h)

Galerula subchspa MURRILL 1942, Lloydia 5: 148

Conocybe subcrispa (MURRILL) SINGER 1950, Sydowia 4: 139

Original description:

Pileo conico ad expanso depressoque 7 mm lato, pallido, disco isabellino, praesulcato;

lamellis adnexis. angustatis, subdistantibus, stipes 5-6 cm longo, albo, bulboso.

Microscopical characters:

Spores: 12.5-18.5 x 8-10 urn, mean 15.3 x 8.6 urn, Q =

1.7-2.2,

ellipsoidal, often elon-

gated with almost cylindrical-parallel walls, not lenticular, with wall up to 0 8 um

thick and a germ-pore up to 2 um wide, yellow-brown in K.OH

Basidia: rather certainly 4-spored; no intact basidia but only tetrads of accumu-

lated immature basidiospores could be observed, indicating 4-spored basidia.

Clamp connections: sporadically present.

Cheilocystidia: lecythiform, some fragments (ca 18-20 x 7-8 um) with con-

spicuous capitulum 2-4 5 urn wide observed

Stipe covering: consisting only of non-lecythiform elements and hairs up to

50 x 4 urn.

Pi lei pell is: hymeniform, consisting of round ish-pyri form elements up to 31 x

21 urn, in-between abundant pileocystidia similar to cheilocystidia, but partly larger

(up to 32 x 13 urn) with a thicker neck.

Specimen examined: USA: Florida, Alachua County, Gainesville, in grassy lawn under long-

needled pine, 26. 7 1939, leg. W. A. MURRILL (FLAS, holotype).

The type specimen consists of ca four stipes and several small pileus fragments in

poor condition. The microscopical analysis and the major part of the following discus-

sion are a translation from HAUSKNECHT (1998)

Conocybe subcrispa is a member of sect Candidae Its fruitbodies are much more

delicate and have a longer stipe than Conocybe albipes var. crispa, in addition regular

lamellae and longer, differently shaped spores. Conocybe crispella has roughly similar

colours like C. subcrispa and also delicate fruitbodies, but much smaller, regularly

ellipsoidal-ovate spores with a Q of

1.35-1.7,

compared to

1.7-2.2

in Conocybe sub-

crispa Therefore, the latter is a separate species, which according to our knowledge is

up to date only known from two North American collections (HAUSKNECHT 1998).

sulcatipes (Fig 24 a-f)

Agahcus sulcatipes PECK 1884, Ann. Rep New York State Mus Nat. Hist 35: 132

Galera sulcatipes (PECK) SACC. 1887, Sylloge Fungorum 5: 866

Galerula sulcatipes

(PECK)

MURRILL 1917. North American Flora 10: 166

Pholiotina sulcatipes (PECK) BON 1991, Doc Mycol 21/83 39

Österr. Z. Pilzk. 13 (2004) 225

Original description:

Pileus thin, ovate, then conical or subcampanulate. hygrophanous, chestnut-colored

and generally striatulate on the margin when moist, becoming paler when dry; lamel-

lae ascending, subdistant, adnate, whitish, becoming ferruginous-cinnamon, stem slen-

der, straight or flexuous, equal, hollow, rather tenacious, stnate-sulcae, silky, floccose-

pruinose toward the base, white, often tinged with blue or green at the base. Plant gre-

garious, 1.5-3' high, pileus 5"-8"broad, stem 1" thick.

Microscopical characters:

Spores 7-9 x 4-5 5 urn, mean 8.0 x 4.8 urn, Q =

1.6-1.8,

ellipsoidal, not lentiform,

slightly thick-walled and with a conspicuous germ-pore ca

urn in diam , yellowish-

brown in KOH

Basidia: 4-spored, 13-24x8-12 um.

Clamp connections: abundant, everywhere in the trama.

Cheilocystidia: fusiform to cylindrical-ventncose, with acute but not con-

spicuously differentiated beak, 25-40 x 6.5-9.5 um.

Stipe covering: with numerous caulocystidia similar to cheilocystidia, but of-

ten larger and cruder.

Pileipellis: hymeniform, consisting of pyriform to roundish-stipitate elements,

26-55 x 9-25 (im, in-between numerous pileocystidia up to 95 x 16 urn.

Specimen examined: USA: New York. East Beme. on pile of buckwheat bran in woods. August

1893.

leg C H PECK (NYS. holotype).

The type consists of three pressed fruitbodies glued on paper and of five to six ad-

ditional loose fruitbodies in good condition. The microscopical drawings are taken

from HAUSKNECHT (2001).

Pholiotina

sulcatipes is a well-defined species, concerning conspecificity with the

European

Pholiotina aberrans

(KUHNER) SrNGER we refer to the different concepts of

HAUSKNECHT (2001) and ARNOLDS (2004).

tenerella (Fig. 25 a-e)

Galera

tenerella

G F ATK 1909, Ann. Mycol 7: 369

Galerula

tenerella

(G F ATK ) MURRILL 1917, North Amer Flora 10: 164

Original description:

Plants growing close together, 5-7 cm high; pileus 3-4 mm high and broad; stem 1-1.5

mm thick. Entire plant ochraceous. Pileus thin, campanulate. smooth, covered with a

very thin and delicate filamentous whitish bloom. Gills subelliptical, narrow, adnexed,

edge whitish. Stem even, pruinose above, and over entire length when young and fresh.

226 A. HAUSKNECHT & al.: Type studies in North American Bolbitiaceae

Microscopical characters:

Spores: 14 5-15 5 x 9-10.5 x 8.5-9 urn, mean 14.8 x 9.5 x 8.6 um, Q = 14-16, ellip-

soidal, slightly lentiform. with wall 0.3-0.7 |im thick and a germ-pore 1.5-2 urn in

diam., rust- to orange-brown in K.OH

Basid ia: 2-spored, 19-23 x 10-13 urn

Clamp connections: present at the base of

the

basidia and in the trama.

Stipe covering: consisting of capilliform, roundish to fusiform-lageniform ele-

ments; no lecythiform caulocystidia present.

Pileipellis: hymemform. consisting of round ish-stipitate elements. 20-30 x

18 urn, in-between abundant capilliform pileocystidia up to 5 urn wide

Specimen examined: I'SA: New York. Ithaca. Cornell Conservatory, on dung in pots in palm

house. 1.4. 1906. leg. G. F. ATKINSON (CUP. holotype).

The holotype specimen consists of two rather fragmented fruitbodies in good con-

dition.

SINGER (1950) considers Galera tenerella conspecific with Conocybe ambigua

WATLING; however, the completely different shape and the much darker spores do not

favour this. The microscopical features of this taxon are almost identical with Cono-

cybe siliginea (FR.:

) KÜHNER sensu HAUSKNECHT & PASSAUER (1997). In com-

parison to the neotype selected in HAUSKNECHT & PASSAUER (1997). the spores are

slightly thinner-walled and more distinctly lentiform; however, in our opinion these

differences are not sufficient to accept Galera tenerella as a separate species or vari-

ety/form of Conocybe siliginea We therefore consider both taxa as conspecific.

teneroides (Fig. 25 f-j)

Agaricus teneroides PECK 1878, Ann. Rep. New York State Mus. 29: 39

Galera teneroides (PECK) SACC 1887, Sylloge Fungorum 5 861

Galerula teneroides (PECK) MURRILL 1917. North American Flora 10: 166

Original description:

Pileus thin, campanulate or expanded, gregarious, 1-2.5 cm broad; surface hygropha-

nous,

brownish-cinnamon when moist, paler when dry; lamellae narrow, crowded,

yellowish-cinnamon; stipe straight, slender, hollow, concolorous, 2.5-5 cm long, about

1 mm thick.

Microscopical characters:

Spores: 6-8 x 3.5-4.5 um, mean 7.4 x 4.2 um, Q =

1.6-1.9,

ellipsoidal, slightly pip-

shaped, not lentiform, thin-walled, without or with inconspicuous germ-pore, often

only with callus.

Basidia: 4-spored, 12-16 x6-9 urn.

Clamp connections: present

Cheilocystidia: lecythiform, 13-18 x 6-9.5 urn, with capitulum 4-5.5 urn in

diam.

Österr. Z. Pilzk. 13(2004) 227

Fig. 25 a-e. (ialera lenerella (type), a Spores, b basidia, c cheilocystidia, d stipe covering, e pileipellis.

jf-j. Agaricus teneroides (type)./Spores, g basidia, h cheilocystidia,; stipe covering, j pileipellis.

228 A. HAUSKNECHT & al : Type studies in North American Holbitiaceae

Stipe covering: consisting of differently shaped elements, often with capilli-

form extensions; lecythiform caulocystidia completely absent.

Pileipellis: hymeniform, consisting of roundish-stipitate elements, 22-32 x 13-

18 urn, no pileocystidia observed.

Specimen examined: USA: New York, Lewis County, Greig, on soil or manure in woods,

Sept., leg. C H. PECK (NYS, holotype).

The type specimen consists of seven to eight partly fragmented fruitbodies in ade-

quate condition.

WATLING & GREGORY (1981) consider Agaricus teneroides PECK, to be synony-

mous to Conocybe rickeniana, which is not supported by the smaller cheilocystidia

with comparatively smaller capitula and especially by the stipe covering. The micro-

scopical features observed by us place this taxon into sect. Pilosellae, it has all features

of Conocybe pallidospora KUHNER & WATLING and would therefore have priority due

to the prior date of publication. However, as the epithet "teneroides" has already been

used for another species within Conocybe [Conocybe teneroides (J. E. LANGE) KUHNER

1935,

= Pholiotma teneroides (J. E. LANGE) SINGER], it cannot be used and Conocybe

pallidospora remains the valid name

SEM-pictures of the spores of Agaricus teneroides show a very faintly rugulose

spore surface, which has not yet been observed in European collections of Conocybe

pallidospora However, this slight roughness is so faint that it is probably an artefact

of storage of the type material about 130 years old. We therefore do not ascribe im-

portance to this feature.

tetrasporoides (Fig 26 a-d)

Conocybe tetrasporoides HAUSKN.

2003,

Österr Z Pilzk 12 78

Original description (translated from HALSKNECHT 2003):

Pileus 2-10 mm wide, flat hemispheric to flat convex, hygrophanous, but not striate,

young and in fresh condition orange-brown, pale yellow, cream, later altogether cream

to yellowish-white, margin almost whitish, pileus surface smooth, somewhat mica-

ceous.

Lamellae sinuate, moderately distant to distant, ventricose; first cream, then

ochre to pale white-coffee-brown, with smooth, somewhat paler lamellar edge Stipe

15-45 mm long, ca

mm thick, base not to slightly enlarged, not rooting, first whitish,

cream-white, then above the base darker, ochre to light yellow-brown, at the apex

white-pruinose, otherwise very finely striate. Trama very thin, translucently white to

yellowish, without smell, with strong radish-like taste.

Microscopical characters:

Spores: 9-14 x

5.5-8.5

urn, mean 10.9 x 6 5 um, Q = 1.6-2.1, fusiform-ellipsoidal,

ellipsoidal-pip-shaped, lenticular, with comparatively thin wall and conspicuous germ-

pore ca

urn in diam., light orange-yellow to ochre-yellow in KOH

Basidia: 4-spored, sporadically 2-spored. clavate-stipitate. 20-25 x 8 5-11 urn

Clamp connections: present

Osten. Z. Pilzk. 13(2004)

229

Fig. 26 a-d.

('onocybe

lelrasporoides (type), a Spores, b basidia, c cheilocystidia, d stipe covering, e-

h Agancus toriipes (type), e Spores, f basidium with immature spores, g cheilocystidia. h strongly

collapsed elements on the pileipellis (velum?).

230

HAUSKNECHT

& al.: Type studies in North American Bolbiliaceae

-reaction: negative.

Chei locystidia: lecythiform, 15-23 x 6-9 urn, with capitulum 2.5-4 |im in

diam.

Stipe covering: consisting of lecythiform caulocystidia, non-lecythiform, cy-

lindrical to capilliform elements; in some regions lecythiform elements rarer, in others

in dense clusters present especially near to the stipe apex.

Pileipellis: hymeniform. consisting of roundish-stipitate elements. 35-60 x 25-

42 urn; pileocystidia not observed

Specimen

examined: USA: Tennessee.

KJIOX

County

. Knoxville, University of Tennessee, in

park

lawn.

29. 7. 19%, leg. 1.

K.RISAI-GREILHUBER

& H VOGLMAYR (WU

17104.

holotype).

The macro- and microscopical descriptions given above only refer to the type col-

lection from Tennessee and have been translated from

HAUSKNECHT

(2003). The type

consists of

six

well-preserved fruitbodies

Conocybe tetrasporoides is a member of sect. Mixtae and is close to Conocybe

ambigua WATLING. It differs from the latter in 4-spored basidia. smaller spores,

smaller fruitbodies with a lighter coloured, completely non-striate pileus.

tortipes

(Fig 26 e-h)

Agaricus tortipes

MONT.

1856, Sylloge Crypt.: 119

Galera tortipes

(MONT.)

SACC.

1887. Sylloge Fungorun 5: 867

Galerula tortipes

(MONT.)

MURRILL 1917, North American Flora 10: 167

Conocybe tortipes

(MONT.)

WATLING 1981, Bibl. Mycologica

82:

146

Original description:

Pileus submembranaceus vel carnosulus, convexo-campanulatus, helvolus, disco lae-

vis.

margine recto crenulato-fisso striatus. 3

cm altus. 4 cm latus Stipes procerus.

subcartilagineus. fistulosus, fragilis, flexuosus, tortilis, striatus, 15 cm et quod excedit

longus, 5 mm medio crassus, basi subincrassatus, pileo concolor. intus in exsiccato

specimine fuscus. Lamellae tridymae, lineares, 2 mm vix latae, postice adnexae. antice

attenuatae, tandem cinnamomeae, acie pallidae.

Obs.

Figura

tabulae

Elenchi Fungorum Batschii hanc nostram in memona re-

vocact speciem. Vi colons nonnullis forsan ad Bolbitium earn referendam censebunt,

quam opinionem. repugnante defectu tramae inter strata hymemi. defend) non posse

manifestum est. Insuper sporae Coprinorum desiderantur Si autem sporis albis usa,

Agarico collino affinis fuisset.

Microscopical characters:

Spores:

6.5-7.5

x 4-4.5 urn, mean 6.8 x 4.3 urn, Q =

1.6-1.7,

ellipsoidal, not lenti-

form, thin-walled, without germ-pore, violet-beige-grey in KOH

Basidia: 4-spored, ca 17 x 8 urn.

Clamp connections: not observed.

Cheilocystidia: clavate-vesicular, cylindncal-clavate, 20-47 x 6-12 5 urn.

Stipe covering: entirely collapsed, no conspicuous elements observed.

Österr. Z. Pilzk. 13(2004)

231

Fig. 27 a-J. Conocybe luxlaensis (type), a Spores, b basidia. c cheilocystidia. d stipe covering, e-h.

(ialera viscosa (type), e Spores,/basidia, g cheilocystidia, h pileipellis.

232 A. HAUSKNECHT & al.: Type studies in North American Holbiliaceae

Pileipellis: structure not detectable; some roundish, strongly collapsed ele-

ments up to 45 x 40 um observed, which could be parts of the velum or of the pilei-

pellis.

Specimen examined: USA: Ohio, Columbus, leg J. P. F MONTAGNE 378 (PC, holotype).

The type consists of one large fruitbody in very poor condition, most elements

collapsed and heavily molded

Due to the spore colour in KOH and the shape of the cheilocystidia, Agahcus tor-

tipes has to be excluded from Bolbitiaceae. Most likely it is a member of Stropharia-

ceae or of the genus Psathyrella, but the rather

stiff,

contorted stipe militates against

the latter. We therefore consider Conocybe tortipes as a dubious taxon.

tuxlaensis (Fig. 27 a-d)

Conocybe tuxlaensis SINGER 1989, Fieldiana n s 21 105

Original description:

Pileo gnsello-ochreo vel ochraceo-subgnseo, hygrophano, glabro, per medium radium

pellucide striato et leniter sulculato, convexo, obtuso, 6 mm lato. Lamellis subferrugi-

neo-ochraceo-brunneis, acie albis, latis, sublibens, subconfertis vel mediocnter distan-

tibus.

Stipite pallide brunneo, subglabro, subaequali, 11 x 0,5 mm. Carne tenui, mo-

dora

Microscopical characters:

Spores: 5-7 5 x 3 5-4 (im, mean 6.2 x 3 7 urn, ellipsoidal, thin-walled, with callus or

very faintly visible germ-pore, light orange in KOH.

Basidia: 4(2)-spored, 15-20x7-8 5 urn.

Clamp connections: present

Cheilocystidia: lecythiform. 15-20 x 5-9 urn, with slender neck and capitulum

3-4.5 urn in diam.

Stipe covering: consisting of a mixture of hairs, elongated-roundish and le-

cythiform elements, the latter being dominant (in the type ca 70 %). The capitate

caulocystidia similar to those of Conocybe subpallida ENDERLE and measuring 15-35

x 4.5-8 urn, with capitulum 1.5-5 urn wide.

Pileipellis: hymeniform. consisting of roundish-stipitate elements, with lecy-

thiform pileocystidia.

Specimen examined: Mexico: Veracruz, Estaciön Biolögica de los Tuxlas. singular in tropical

rain forest,

23.

6. 1969, leg. R. StNGER

8185

(F,

holotype).

The holotype consists of one singular, tiny fruitbody in good condition

The results of our microscopical analyses are not fully congruent with those of

SINGER (1989), especially the details on the spores deviate - see HAUSKNECHT (2002

b).

Although the type specimen originate from a tropical rain forest, this species has

been reported from Europe several times (HAUSKNECHT 2002 b). A microscopically

Osten.

Z. Pilzk. 13 (2004) 233

similar species which may be confused with it is Conocybe pilosella

(PERS.:

FR.)

KÜH-

NER with much larger fruitbodies and a completely different stipe covering.

viscosa (Fig 27 e-h)

Galera viscosa CLEM.

Galerula viscosa (CLEM ) G F ATK 1918, Proc. Amer. Phil Soc. 57: 368

Original description

(label of

the

type

specimen):

Galera viscosa CLEM., Cryptogamae Formationum Coloradensium, F E. & E S.

CLEMENTS. 380. Galera viscosa n. sp., saprophilus copiosus ad fimum vaccinum

udumque in piceto raro Picea-Pseudotsuga hylio. Cameron Glen 2800 m, 25. 8. 1906.

Microscopical

characters:

Spores: 10.5-13.0 x 7-8 urn, mean 11 6 x 7.4 urn, Q =

1.4-1.6,

ellipsoidal, not lenti-

form. with double wall and large germ-pore, bright yellow to orange-yellow in K.OH.

Basidia: 4-spored, 21-26 x 10-13.5 urn.

Clamp connections: present.

Cheilocystidia: 22-45 x 9-11 urn, usually lageniform with long beak, which is

sometimes slightly moniliform, rarely lageniform with ventricose base.

Pileipellis: hymeniform, consisting of roundish-stipitate elements. 33-40 x 18-

22 um, with numerous capilliform to slightly lageniform pileocystidia.

Specimen

examined: USA: Colorado. Cameron Glen. 2800 m s. m.. on cow dung, 25 8. 1906,

leg.

F. E. & E. S.

CLEMENTS

(E, holotype).

The type specimen consists of three well-preserved fruitbodies which contain rests

of dung at the stipe base.

Galerula viscosa has been published as part of distributed exsiccata, therefore the

printed label represents the original description. Unfortunately it has not been possible

for us to find out the date of distribution of

the

respective fascicle.

Already WATLING & GREGORY (1981) assume that Galerula viscosa is an older

name for the European Pholiotina coprophila (KÜHNER) SINGER and should be com-

bined into the genus Pholiotina All microscopic features, the appearance of the well-

preserved specimen and also the habitat perfectly agree with Pholiotina coprophila. As

we have not yet been able to locate the printed herbarium schedae and the date of dis-

tribution, we refrain from a combination into Pholiotina.

are very grateful to RON H. PETERSEN for providing the unpublished notes of

R. HESLER

and

additional North American literature. ROY WATLING for valuable information and literature, and

the herbarium curators of

CUP, E, F,

FLAS,

K, MSC. MICH, NEB, NY, NYS, PC, WU and ZT

for

the loan of

herbarium

specimens.

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

Abstract

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