V5 Floraagaricinaneerlandicavol.5.Agaricaceae

FLORA AGARICINA NEERLANDICA
Critical monographs on families of agarics and boleti occurring

in the Netherlands
Edited by
M.E. NOORDELOOS, TH.W. KUYPER, & E.C. V E L L I N G A
With assistance of
J. V A N OS (illustrations)
VOLUME 5
A. General part
B. Taxonomic part
Agaricaceae
A.A. B A L K E M A P U B L I S H E R S LISSE / ABINGDON / EXTON (PA) / TOKYO

Library of Congress Cataloging-in-Publication Data
Applied for
Part of the work for the Flora agaricina neerlandica has been supported by research grants of the
Foundation Rijksherbariumfonds Dr. E. Kits van Waveren.
Typesetting: Red Barn Publishing, Skeagh, Skibbereen, Ireland.

Printed by: Krips, Meppel, The Netherlands.
© 2001 Swets & Zeitlinger B.V., Lisse
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form
or by any means, electronic, mechanical, by photocopying, recording or otherwise, without the prior written permission of
the publishers.
For the complete set of Flora agaricina neerlandica (cloth): ISBN 90 6191 861 8
For Volume 1 (cloth): ISBN 90 6191 859 6
For Volume 1 (paper, o.o.p.): ISBN 90 6191 758 1
For Volume 2 (paper, o.o.p.): ISBN 90 6191 972 X
For Volume 5 (paper): ISBN 90 5410 495 3
Contents
A . General part
1. Scope, methods, and presentation by C.Bas 3
2. Glossary by M . E . Noordeloos & E . C . Vellinga 6
3. Abbreviations of authors' names by E . C . Vellinga, M . M . Nauta & M . E . Noordeloos 12
4. Bibliographic abbreviations by E . C . Vellinga, M . M . Nauta & M . E . Noordeloos 15
B . Taxonomic part
Family Agaricaceae by E . C . Vellinga 21
Key to the genera 21
1. Genus Agaricus by M . M . Nauta 23
2. Genus Allopsalliota by M . M . Nauta 62
3. Genus Macrolepiota by E . C . Vellinga 64
4. Genus Chlorophyllum by E . C . Vellinga 74
5. Genus Leucocoprinus by E . C . Vellinga 76
6. Genus Leucoagaricus by E . C . Vellinga 85
7. Genus Lepiota by E . C . Vellinga 109
8. Genus Chamaemyces by E . C . Vellinga 152
9. Genus Cystolepiota by E . C . Vellinga 154
10. Genus Melanophyllum by E . C . Vellinga 161
Index 163
Index to the generic names i n volumes 1-5 169

A
General part
CHAPTER 1
Scope, methods and presentation

C. BAS
Scope and outside the Netherlands, are given but distribution maps are not
The ' F l o r a agaricina neerlandica' contains keys to, correct names of, included.
concise synonymies for, descriptions and illustrations of, and distribu- O n l y i n the cases o f very rare species (five or fewer localities
tional data o n agarics and boleti (Agaricales sensu lato and Russulales) k n o w n i n the Netherlands) are i n d i v i d u a l localities enumerated.
occurring i n the Netherlands. F o r practical reasons, the genera o f so- Frequency classes (extremely rare, very rare, rare, rather rare, m o d -
called c y p h e l l o i d fungi are excluded. erately c o m m o n , rather c o m m o n , c o m m o n , very c o m m o n ) are esti-
E x t r a l i m i t a l taxa o f w h i c h occurrence i n the Netherlands c a n be mated from the number o f collections f r o m the Netherlands studied
expected o n account o f their e c o l o g y and/or g e o g r a p h i c a l distribu- and the number o f reliable references i n literature.
tion are i n c l u d e d i n the keys whereas a concise s y n o n y m y , references A guidance for the estimation o f the frequency classes is found i n
to selected descriptions and i l l u s t r a t i o n s , and a short diagnostic A r n o l d s et a l . (Overzicht van de paddestoelen i n N e d e r l a n d . 1995,
description are g i v e n i n the textual part. F o r various reasons a n u m - reprint 1999) and is based o n numbers o f quadrants o f 5 X 5 k m o n
ber o f taxa less l i k e l y to be encountered i n the N e t h e r l a n d s are topographic maps i n w h i c h the taxa have been found.
i n c l u d e d o n l y i n the keys m e r e l y w i t h their author citation and one extremely rare: 1-2 quadrants.
reference to the literature. very rare: 3-6 quadrants.
F u n g i that spontaneously establish themselves i n the Netherlands rare: 7-17 quadrants.
after human intervention (e.g. m y c o r r h i z a l symbionts o f the introduced rather rare: 1847 quadrants.
Larix species as Suillus grevillei) are considered indigenous. Hothouse moderately common: 48-113 quadrants.
fungi are not treated except some species o f Leucocoprinus frequent- rather common: 114-246 quadrants.
l y found i n flowerpots i n houses. A strongly s i m p l i f i e d map o f the common: 247-426 quadrants.
Netherlands is given i n F i g u r e 1. very common: more than 426 quadrants.
Introductory chapters Illustrations

For chapters o n the history o f agaricology i n the Netherlands, ecology A l l species are illustrated by line-drawings o f basidiocarps, spores and
and distribution o f agarics and boleti i n the Netherlands, specific and cystidia ( i f present), i f necessary complemented by drawings o f other
infraspecific delimitation, generic concepts, orders and families o f the microscopical characters. T h e magnifications o f these drawings are:
agarics and boleti, and nomenclature the reader is referred to Chapters basidiocarps X 1 (unless indicated otherwise)
1-6 i n V o l u m e 1 o f this flora. spores X 1500
cystidia and basidia X 1000 (unless indicated otherwise)
Descriptions tissues X 500 (unless indicated otherwise).
In p r i n c i p l e a l l descriptions are based o n observations referring to W i d e l y dotted areas indicate the presence o f a gelatinous substance.
material from the Netherlands. In cases where this material is not rep- For the abbreviations used i n the illustrations, see the list o f abbrevia-
resentative, however, observations o n other collections f r o m north- tions at the end o f this chapter.
western E u r o p e are sometimes i n c l u d e d . C o l l e c t i o n s revised for this
flora have been m a r k e d w i t h special labels added to them. Formulae
If authentic observations, particularly o n m a c r o s c o p i c characters For formulae o f c h e m i c a l reagents and stains, the reader is referred to
are l a c k i n g , data are taken from literature, but this is then e x p l i c i t l y current m y c o l o g i c a l literature (e.g. K i i h n e r & R o m a g n e s i , F l . anal.
indicated w i t h reference to the sources o f information. C h a m p , sup., 1953; M o s e r , R o h r l i n g e - B l a t t e r p i l z e , 5. A u f L , 1983;
Spore print colours are preferably recorded w i t h c o l o u r code nota- Singer, A g a r i c a l e s m o d . Taxon., E d . 4, 1986).
tions. Unfortunately such precise records are still l a c k i n g for many
taxa. C o l o u r indications without quotation marks have been extracted Nomenclature
f r o m the field-notes o f the collections studied; those between quota- In nomenclatural matters, c l o s e adherence to the latest e d i t i o n o f the
tion marks are taken f r o m literature or are based o n general consensus. 'International C o d e o f B o t a n i c a l N o m e n c l a t u r e ' is a i m e d at. F o r the
Sizes o f spores a m measured i n tenths o f a |jun but rounded off to procedures f o l l o w e d i n cases i n w h i c h the C o d e is not perfectly clear,
halves o f jxm. Sizes o f basidia, cystidia, and other m i c r o s c o p i c a l ele- particularly i n connection w i t h sanctioned names, see V o l . 1, Chapter
ments are measured and given i n |xm, unless they are narrower or 6.
smaller than 10 |a,m, i n w h i c h case they are treated as the spores. S y n o n y m y is not complete, but restricted to synonyms that have
appeared i n m o d e j ^ monographs, w e l l - k n o w n floras, D u t c h m y c o l o g -
Ecological and distributional data i c a l l o g i c a l literature, and popular books w i t h a w i d e distribution. A
Extensive notes o n e c o l o g y and geographical distribution both w i t h i n similar selection is made f r o m the m i s a p p l i e d and e x c l u d e d names.
3
4 FLORA AGARICINA NEERLANDICA
F i g . 1. Topographical map o f the Netherlands(l-12: provinces; 1 = Groningen, 2 = Friesland, 3 = Drenthe, 4 = Overijssel, 5 = F l e v o l a n d , 6,=
Gelderland, 7 = Utrecht, 8 = N o o r d - H o l l a n d , 9 = Z u i d - H o l l a n d , 10 = Zeeland, 11 = Noord-Brabant, 12 = L i m b u r g ) .
SCOPE, METHODS AND PRESENTATION (Bas) 5
N a m e s o f pteridophytes and phanerogams mentioned are i n accor- D r . T h . W . K u y p e r , A g r i c u l t u r a l University, Section S o i l B i o l o g y

dance w i t h H e u k e l s / V a n der M e i d e n (1996), F l o r a van N e d e r l a n d , and Plant F e e d i n g , D r e i j e n p l e i n 10, N L 6700 E C Wageningen, T h e
22nd ed., G r o n i n g e n . Netherlands, E - m a i l : T h o m . K u y p e r @ b b . b e n p . w a u . n l .
D r s . M . M . Nauta, N a t i o n a l H e r b a r i u m o f the Netherlands, L e i d e n
New names and new taxa University B r a n c h , P . O . B o x 9514, 2300 R A Leiden, E-mail:
N e w names found to be required and new taxa discovered i n the course [email protected]
o f the investigations carried out for this flora, are not published i n the D r . M . E . N o o r d e l o o s , N a t i o n a l H e r b a r i u m o f the Netherlands,
flora itself, but i n a series o f separate notes titled 'Notulae ad F l o r a m L e i d e n U n i v e r s i t y B r a n c h , P . O . B o x 9514, 2300 R A L e i d e n , E - m a i l :
agaricinam neerlandicam' appearing i n the j o u r n a l Persoonia. In these Noordeloos @nhn.leidenuniv.nl
Notulae the more lengthy discussions on taxonomic and nomenclatur- D r s . E . C . V e l l i n g a , U n i v e r s i t y o f C a l i f o r n i a at Berkeley, Dept. o f
a l problems are also placed. Plant and M i c r o b i a l B i o l o g y , B r u n s L a b , 111 K o s h l a n d H a l l - 3 1 0 2 ,
Berkeley, C A 94720-3102, U . S . A . E - m a i l : V e l l i n g a @ u c l i n k 4 . b e r k e -
ley.edu
Addresses of the authors and editors
D r . C . B a s , N a t i o n a l H e r b a r i u m o f the Netherlands, L e i d e n U n i v e r s i t y
B r a n c h , P . O . B o x 9514, 2300 R A L e i d e n , E - m a i l : B a s @ n h n . l e i d e -
nuniv.nl
LIST OF ABBREVIATIONS IN TEXT AND DRAWINGS
ad int. - ad interim nom.conserv. - nomen conservandum

auct. - auctores ( = authors) nom.nud. - nomen n u d u m
auct. eur. - auctores europaei ( = E u r o p e a n authors) not val. p u b l . - not v a l i d l y published
auct. neerl. - auctores neerlandici ( = D u t c h authors) Nov. - November
A u g . - August Oct. - October
av. - average pc - pileus covering
cc - caulocystidia p i - pleurocystidia
cf. - confer(= compare) p i . - plate
ch - cheilocystidia pp - pileipellis or elements o f pileipellis
D e c . - December p.p. - pro parte
descr. - description p.p.maj. - pro parte majore ( = for the greater part)
diagn. - diagnosis p.p.min. - pro parte minore (for the m i n o r part)
dpt. - departement prov. - province
emend. - emendatus ( = corrected) R o m a g n . - R o m a g n e s i , L e s Russules d ' E u r o p e (colour chart)
et al. - et a l i i Q - quotient o f length and w i d t h or breadth
e x c l . - excluded Qav - average quotient
f. - forma sect. - section
Feb. - February sel. - selected
F i g . , fig. - Figure, figure sensu auct. maj. - sensu auctores majores
F i g s . , figs. - Figures, figures Sept. - September
illegit. - illegitimate s.l. - sensu lato ( = i n a w i d e sense)
Jan. - January s.str. - sensu stricto ( = i n a restricted sense)
K . & W . - K o r n e r u p , A . & Wanscher, J . H . , M e t h u e n handbook o f subgen. - subgenus
colour; Farver i farver. subsect. - subsection
1 - number o f lamellulae between two lamellae subsp. - subspecies
L - number o f lamellae subvar. - subvariety
lit. - literature trib. - tribus
loc. cit. - l o c o citato ( = i n the passage already quoted) var. - variety
misappl. - m i s a p p l i e d vern. - vernacular
M u . - M u n s e l l s o i l c o l o r charts
CHAPTER 2
Glossary
E . C . V E L L I N G A & M . E . NOORDELOOS
abrupt papilla - (on pileus) ( V o l . 1, F i g . 29.30). binding hyphae - branching, rarely septate, t h i c k - w a l l e d , narrow
abruptly bulbous - (base o f stipe) ( V o l . 1, F i g . 31.18). hyphae b i n d i n g the other elements o f a tissue together.
acanthocyte - globose, spiny element produced on the basal m y c e l i u m binucleate - w i t h two nuclei.
o f some Strophariaceae. bivelangiocarpy - a type o f angiocarpic development o f the basidio-
acrophysalidic - (tissue) consisting o f connective hyphae and abun- carp i n w h i c h partial and universal v e i l participate.
dant, large, terminal, inflated elements ('acrophysalides'). breadth - (of spore) largest distance between sides as seen from frontal
acuminate - (cystidia) tapering f r o m i n w a r d l y curved sides o f f to a view ( V o l . l . F i g . 32.2).
point ( V o l . 1, F i g . 34.39). brevipes-type - see urticiform.
acute - (cystidia) tapering off to a sharp point ( V o l . 1, F i g . 34.40); - broadly clavate - (cystidia) clavate w i t h Q < 1.5 ( V o l . 1, F i g . 34.14).
(spore apex) pointed ( V o l . 1, F i g . 33.30). broadly conical - (pileus) ( V o l . 1, F i g . 29.13), see also truncately
acute papilla - (on pileus) ( V o l . 1, F i g . 29.31). broadly c o n i c a l ; - (cystidia) c o n i c a l w i t h Q < 1.5 ( V o l . 1, F i g .
adnate - (lamellae) broadly attached to stipe ( V o l . 1, F i g . 30.21), see 34.18).
also narrowly adnate. ,- broadly cylindrical — (cystidia) c y l i n d r i c a l , w i t h Q < 2 ( V o l . 1, F i g .
adnexed- (lamellae) rounded towards stipe ( V o l . 1, F i g . 30.19). 34.11) .
aeriferous - (pileus and stipe surfaces) w i t h silvery shine because o f broadly ellipsoid -^(spores) Q = 1.15-1.3 ( V o l . 1, F i g . 33.3); (cystidia)
air present a m o n g superficial hyphae. Q = 1.15-1.3 ( V o l . l , F i g . 34.3).
allantoid - (spores) w i t h adaxial side concave and parallel to abaxial broadly fistulose - (stipe) w i t h very w i d e tube ( V o l . 1, F i g . 31.12).
side ( V o l . 1, F i g . 33.21). broadly fusiform - (spores) fusiform w i t h Q = 1.5-2.0 ( V o l . 1, F i g .
amygdaliform - (spores) w i t h adaxial side straight or less convex than 33.12) ; - ( c y s t i d i a ) fusiform w i t h Q = 1.5-2.0 (Vol. 1, F i g . 34.21),
abaxial side ( V o l . 1, F i g s . 33.18, 33.19). see also very broadly fusiform.
amyloid - (spore w a l l , spore ornamentation, h y p h a l walls), staining broadly lageniform - (cystidia) ( V o l . 1, F i g . 34.26).
greyish to b l a c k i s h blue i n M e l z e r ' s reagent. broadly utriform - (cystidia) ( V o l . 1, F i g . 34.29).
anastomosing - (lamellae) p r o v i d e d w i t h irregular transverse connec- broadly ventricose - (lamellae) ( V o l . 1, F i g . 30.14).
tions ( V o l . 1, F i g . 30.4). broom-cells - elements i n pileipellis or h y m e n i u m consisting o f a more
angiocarpy - a type o f development o f the basidiocarp i n w h i c h at or less clavate or spheropedunculate body, bearing few to numer-
some stages the developing h y m e n i u m is situated i n a closed cav- ous warts o f finger-like appendages i n upper half. T w o types are
ity; see also primary and secondary angiocarpy. distinguished: Rotalis-type w i t h rounded-warty appendages ( V o l .
annular belt - (stipe) remnants o f partial v e i l i n f o r m o f a girdle round 3, Fig. 139), and Siccus-type with elongate, finger-like
(upper half of) stipe. appendages ( V o l . 3, F i g . 154).
annulus - (stipe) ring-like structure r o u n d (apex of) stipe, formed by brosse - see en brosse.
partial v e i l . bulbangiocarpy - a type o f angiocarpic development o f the basidio-
apex - (of spores) summit ( V o l . 1, F i g . 3 2 . 4 A ) . carp i n w h i c h thevbasidiocarp is initiated i n a cavity o f the p r i -
apical - (spores) situated at the tip. m o r d i a l bulb.
appendage - (of spores) see hilar appendage. bulbous - (base o f stipe) enlarged ( V o l . 1, F i g . 31.17), see also abrupt-
appendiculate - (margin o f pileus) w i t h small appendages ( V o l . 1, F i g . l y bulbous and marginately bulbous.
29.48). campanulate - (pileus) bell-shaped ( V o l . 1, F i g . 29.12).
applanate - (pileus) flattened, flat ( V o l . 1, F i g s . 29.7, 29.19). canaliculate - (stipe) ( = channelled) w i t h a l o n g i t u d i n a l groove or
arachnoid - (pileus and stipe surfaces or velum) cobwebby. channel.
arcuate - (lamellae) w i t h concave l a m e l l a edge (Vol. 1, F i g s . 30.15, cantharelloid - (basidiocarp) w i t h decurrent obtuse r i b - or v e i n - l i k e
30.16). lamellae.
areolate-rimose - (pileus surface) m a r k e d w i t h numerous superficial capitate - (apex o f cystidia) having a distinct and abrupt k n o b ( V o l . 1,
clefts or cracks f o r m i n g angular patches. F i g . 34.44).
ascending - (hyphae) c u r v i n g upward. cartilaginous - (stipe) cartilage-like, flexible but tough.
aseptate - (hyphae) without septa. catenate - (of cheilocystidia) united as i n a chain.
auriscalpioid - (habit) shaped l i k e Auriscalpium spp.: w i t h vertical catenulate - (elements) united or l i n k e d as i n a chain.
eccentric stipe and horizontal pileus. caulocystidium - c y s t i d i u m situated o n surface o f stipe.
bacilliform - (spores) Q > 3.0 ( V o l . 1, F i g . 33.7). central - (stipe) attached to centre o f pileus ( V o l . 1, F i g . 31.1); - (germ
ballistosporic basidium - a b a s i d i u m that actively discharges its spores. pore) situated at the central tip o f the spore ( V o l . 1, F i g . 33.27).
basidiocarp - fruitbody p r o d u c i n g spores o n basidia. chambered - (stipe) w i t h several cavities ( V o l . 1, F i g . 31.13).
basidiole - immature b a s i d i u m . cheilocystidium - c y s t i d i u m situated on edge o f l a m e l l a or tube.
bilateral - see divergent. cheiloleptocystidium - t h i n - w a l l e d c y s t i d i u m situated o n edge o f
6
GLOSSARY (Vellinga & Noordeloos) 1
lamellae (only used w h e n confusion w i t h c h e i l o c h r y s o c y s t i d i a deliquescent - (lamellae and/or basidiocarp) b e c o m i n g l i q u i d after

may occur). maturing.
cheilochrysocystidium - c h r y s o c y s t i d i u m situated o n edge o f lamellae. depressed - (pileus) w i t h central depression/sinking ( V o l . 1, F i g .
chrysocystidium - c y s t i d i u m w i t h y e l l o w amorphous body or bodies i n 29.25) , see also slightly depressed.
contents after treatment w i t h a m m o n i a or K O H . depression - (of pileus), see depressed; - (of spores), see suprahilar
circular- (pileus) round ( V o l . 1, F i g . 29.1). depression.
clavate - (stipe) club-shaped ( V o l . 1, F i g . 31.8); - (cystidia, basidia), derm - a pileipellis consisting o f erect elements or o f ascending ele-
Q = 1.5-4 ( V o l . 1, F i g . 34.13), see also narrowly, and very nar- ments, see trichoderm and h y m e n i d e r m .
r o w l y clavate, and broadly clavate. dextrinoid - (spore w a l l , h y p h a l walls) staining red to reddish b r o w n
clitocyboid - see o m p h a l i o i d . i n M e l z e r ' s reagent.
collarium - a tube around, but free from, the apex o f the stipe to w h i c h dimitic - (tissues) consisting o f generative hyphae and skeletal hyphae
the lamellae are attached ( V o l . 1, F i g . 30.5). or b i n d i n g hyphae.
colliculose - (pileus surface) covered w i t h h i l l o c k - l i k e elevations. divergent - ( = bilateral) (hymenophoral trama) having downward hyphae
collybioid - (habit) characterized b y pileus neither umbilicate, nor con- turning outward from a median line (Vol. 1, Figs. 35.5, 35.6).
i c a l ; lamellae free or adnate; context tough; context o f pileus c o n - diverticulate - (cystidia) w i t h short finger-like excrescences ( V o l . 1,
tinuous w i t h context o f stipe ( V o l . 1, F i g s . 28.2, 28.3). F i g . 34.45).
concave - (pileus) ( V o l . 1. F i g . 29.21). discontinuous - context o f pileus and stipe not f o r m i n g one entity;
congophilous - (spore w a l l ) accumulating C o n g o red. pileus easily separating f r o m stipe, see also pluteoid.
conical - (pileus) cone-shaped ( V o l . 1, F i g . 29.14), see also broadly duplex - (context) consisting o f two structurally differing layers.
c o n i c a l and n a r r o w l y c o n i c a l , truncately broadly c o n i c a l , trun- eccentric - (stipe) not attached to centre o f pileus ( V o l . 1, F i g . 31.2);
cately c o n i c a l , and obtusely c o n i c a l ; - (cystidia) Q = 1.5-4 (Vol. - (germ pore) situated at the abaxial side o f the spore ( V o l . 1, F i g .
1, F i g . 34.17), see also narrowly c o n i c a l and broadly conical. 33.26) .
conidium - a non-motile asexual spore. echinate - (spores) w i t h spines.
connate - (stipes) g r o w n together at their bases. ectosporium - the very thin outer layer o f the basidiospore w a l l .
connective hyphae - usually narrow undifferentiated hyphae o f the ellipsoid - (spores) Q = 1.3-1.6 ( V o l . 1, F i g . 33.4); - (cystidia) Q =
context o f a basidiocarp connecting a l l other elements (used i n 1.3-1.6 ( V o l . 1, F i g . 34.2), see also broadly e l l i p s o i d .
opposite o f fundamental hyphae). emarginate - (lamellae) notched near the stipe ( V o l . 1, F i g . 30.23).
constriction - see m e d i a n constriction. en brosse - (of cystidia) with excrescences, diverticulate ( V o l . 1, F i g .
continuous - context o f pileus and stipe f o r m i n g one entity; pileus and 34.45).
stipe not easily separating. encrusted, encrusting - see incrusting.
convex - (pileus) ( V o l . 1, F i g . 29.9). endosporium - the electron transparent inner layer o f the basidiospore
coralloid - irregularly branching, l i k e i n c o r a l . w a l l at the inside o f the episporium, but l a c k i n g i n many white-
corneous - (stipe) horny, horn-like i n texture. and pale-spored taxa.
cortina - a w e b - l i k e v e i l connecting pileus m a r g i n and stipe. entire - ( l a m e l l a edge) straight, smooth, and glabrous ( V o l . 1, F i g s
costate - (pileus and stipe surfaces) w i t h ridges. 30.30, 30.38).
crenate - (margin o f pileus) w i t h rounded teeth ( V o l . 1, F i g . 29.45); - episporium - the electron opaque fundamental layer present i n a l l
(lamella edge) ( V o l . 1, F i g . 30.33). basidiospores o f the H y m e n o m y c e t e s ; it is the innermost layer o f
crenulate - (margin o f pileus) m i n u t e l y crenate ( V o l . 1, F i g . 29.46); - the spore w a l l w h e n the endosporium is l a c k i n g .
(lamella edge) ( V o l . 1, F i g . 30.34). epithelioid hymeniderm - a h y m e n i d e r m made up o f elements w i t h Q
crepidotoid - see pleurotoid. = 1.0-1.15 ( V o l . 1, F i g . 3 6 . 3 C ) .
curved - (stipe) bent f r o m substrate upwards as to adjust to the gravi- epithelium - a pileipellis made up o f globose to broadly e l l i p s o i d ele-
tation. ments i n more than one layer deep; see regular e p i t h e l i u m and
cutis - a pileipellis consisting o f repent non-gelatinizing hyphae ( V o l . irregular epithelium.
l . F i g . 36.1 A ) . equal - (stipe) o f equal diameter f r o m apex to base.
cyanophilous - (spore w a l l ) accumulating C o t t o n blue. erect - (hyphae or projections o f hyphae) perpendicular to surface o f
cylindrical - (stipe) c i r c u l a r i n cross-section and o f equal diameter pileus.
from apex to base ( V o l , 1, F i g . 31.4); - (spores) Q = 2.0-3.0, i n eroded - (margin o f pileus) irregularly toothed ( V o l . 1, F i g . 29.47); -
outline w i t h p a r a l l e l sides; - (cystidia) Q = 2-4 ( V o l . 1, F i g . (lamella edge) ( V o l . 1, F i g . 30.37).
34.10), see also narrowly c y l i n d r i c a l and broadly c y l i n d r i c a l . euhymeniderm - a h y m e n i d e r m made up o f elements w i t h Q = 1.15-
cyphelloid - (basidiocarp) more or less cup-shaped w i t h smooth 6 ( V o l . 1, F i g . 3 6 . 3 B ) .
hymenophore. eusporium - the inner set o f firm and resistant layers o f the basidiospore
cystidioid - cystidium-like. w a l l , consisting o f the episporium and the endosporium.
cystidium - sterile, differentiated, terminal element i n the h y m e n i u m or even - ( l a m e l l a edge) straight, smooth and glabrous, entire ( V o l . 1,
on the surfaces o f the basidiocarp. F i g s . 30.30, 30.38).
decurrent - (lamellae) descending d o w n the stipe (angle lamellae-stipe exceeding - (margin o f pileus w i t h regard to lamellae) ( V o l . 1, F i g .
4 0 - 6 0 ° ) ( V o l . 1, F i g . 30.28), see also deeply decurrent. 29.42).
decurrent tooth - (of lamellae) ( V o l . 1, F i g s . 30.25. 30.26). exosporium - a layer o f the basidiospore w a l l between perisporium and
deeply decurrent - (lamellae) (angle lamellae-stipe < 4 0 ° ) ( V o l . 1, F i g . episporium, frequently responsible for the ornamentation o f
30.29). spores.
deeply infundibuliform - (pileus) ( V o l . 1, F i g . 29.23). exscissa-type - see urticiform.
deeply umbilicate - (pileus) w i t h deep abrupt depression ( V o l . 1, F i g . fasciculate - (basidiocarps) g r o w i n g i n a bundle.
29.28). felted - (pileus and stipe surfaces or volva) c o m p o s e d of, or covered
deflexed- (margin o f pileus) bent downwards ( V o l . 1, F i g . 29.37). with, densely compressed, matted hairs or fibrils.
fertile - ( l a m e l l a edge) c o m p o s e d o f basidia only. homoiomerous - (tissue) consisting o f hyphae and without nests o f
fibrillose - (pileus and stipe surfaces) covered w i t h thin, thread-like sphaerocysts.
fibres. hymeniderm - a derm made up o f non-septate elements originating a
filiform - (cystidia, stipe), very l o n g and narrowly c y l i n d r i c a l . the same level, see also epithelioid h y m e n i d e r m , and transition
fimbriate - ( l a m e l l a edge) w i t h regular h a i r - l i k e projections ( V o l . 1, between h y m e n i d e r m and epithelium.
F i g . 30.39). hymenocarpy - a type o f development o f the basidiocarp i n w h i c h the
fissurate - (pileus and stipe surfaces) w i t h deep and/or distinct clefts. differentiation o f the hymenophore starts as a circular zone i n the
fistulose - (stipe) h o l l o w ( V o l . 1, F i g . 31.11), see also broadly fistu- otherwise still undifferentiated p r i m o r d i u m .
lose. hymenopodium - a special (usually large-celled) layer between subhy-
flabelliform - (pileus) fan-shaped ( V o l . 1, F i g . 29.2), see also rounded m e n i u m and hymenophoral trama.
flabelliform. hyphal peg - fascicle o f unbranched hyphae projecting beyond basidia.
flattened - (spores) spores adaxially flattened, resulting i n a different imbricate - (basidiocarps) g r o w i n g immediately above each other.
measure for breadth and w i d t h (this volume, F i g . 1.1). incrusted - see incrusting.
flexuous - (stipe) full o f bends; - (cystidia) c y l i n d r i c a l but w i t h bends incrusting - (pigment) situated o n the outer side o f the w a l l , and v i s i -
( V o l . 1, F i g . 34.36). ble as bands, granules or patches.
flocci - s m a l l flocks or tufts. inflated- (hyphae) consisting o f swollen elements, constricted at septa,
floccose - (pileus and stipe surfaces) covered w i t h tufts o f soft hairs. not c y l i n d r i c a l .
flocculose - (pileus and stipe surfaces) minutely floccose. inflexed - (margin o f pileus) bent inwards ( V o l . 1, F i g . 29.38).
free - (lamellae) not attached to stipe ( V o l . 1, F i g . 30.18). infundibuliform - (pileus) funnel-shaped ( V o l . 1, F i g . 29.22), see also
fringed - ( l a m e l l a edge) w i t h irregular appendages. deeply infundibuliform.
frontal view -(of spores) ( V o l . 1, F i g . 32.2). insititious - (stipe) grafted o n substratum; base o f stipe seems insert-
fugacious - (veil) evanescent, early disappearing. ed i n substrate because basal hairs or tomentum are l a c k i n g .
fundamental hyphae - the inflated hyphae g i v i n g the fleshy basidio- intercalary element - between two other elements, not terminal.
carp its firmness. intercellular - (pigment) situated between the elements.
furcate - (lamellae) forked ( V o l . 1, F i g . 30.2). intervenose - (lamellae) p r o v i d e d w i t h veins between the lamellae
fusiform - (spores) spindle shaped, tapering at both ends, w i t h Q = ( V o l . 1, F i g . 30.3).
2.0-4.0 ( V o l . 1, F i g . 33.13); - (cystidia) w i t h Q = 2.0-4.0 ( V o l . 1, intracellular - (pigment) situated inside the elements.
F i g . 34.20), see also narrowly, broadly, and very broadly fusiform. intricate trichoderm - a trichoderm" made up o f interwoven elements
horsehair-like - (stipe) very l o n g and thin, stiff, like the hair o f a horse. ( V o l . 1, F i g . 3 6 . 2 B ) .
gelatinous — j e l l y - l i k e i n consistence. inverse - (hymenophoral trama) having d o w n w a r d convergent hyphae,
generative hyphae - the basic type o f septate, thin- to t h i c k - w a l l e d , i.e. turning i n w a r d to a median line ( V o l . 1, F i g . 35.4).
branching hyphae, present i n a l l (young) basidiocarps, from involute - (margin o f pileus) r o l l e d i n ( V o l . 1, F i g . 29.39).
w h i c h a l l differentiated hyphae arise (used i n opposite o f b i n d i n g irregular - (hymenophoral trama) having interwoven hyphae ( V o l . 1,
and skeletal hyphae). F i g . 35.3).
germ pore - (of spores) apical thin-walled spot i n spore w a l l (Vol. 1, irregular epithelium - an epithelium made up o f irregularly disposed
F i g . 3 2 . 6 E ) , see also central and eccentric germ pore. elements ( V o l . 1, F i g . 3 6 . 4 B ) .
glabrous - (pileus and stipe surfaces) bald. irregular trichoderm - ( V o l . 1, F i g . 3 6 . 2 C ) .
globose - spherical, (spores) Q = 0.95-1.05 ( V o l . 1, F i g . 33.1); - (cys- isocarpy - a type o f development o f the basidiocarp i n w h i c h the dif-
tidia) Q = 1.0-1.05 ( V o l . 1, F i g . 34.5). ferentiation o f pileus, stipe, and hymenophore starts at the same
glutinous - slimy, very v i s c i d . time.
granulose - (pileus and stipe surfaces) covered w i t h (or composed of) ixocutis - a cutis made up o f gelatinizing hyphae ( V o l . 1, F i g . 3 6 . 1 B ) .
minute grains. ixohymeniderm - a h y m e n i d e r m made up o f gelatinizing elements.
gregarious - (basidiocarps) g r o w i n g i n a group. ixohyphidium - a gelatinizing, more or less differentiated, hypha-like
guttate - (pileus and stipe surfaces) w i t h roundish darker spots. terminal element i n the pileipellis.
guttulate - (contents o f spores and other elements) w i t h (oil) droplets. ixotrichoderm - a trichoderm, made up o f gelatinizing elements ( V o l .
guttule - droplet ( V o l . 1, F i g . 3 2 . 6 G ) . 1, F i g . 3 6 . 2 D ) .
gymnocarpy - a type o f development o f the basidiocarp i n w h i c h the laccate - (pileus surface) as though varnished.
h y m e n i u m is exposed f r o m initiation t i l l maturity. lacrymoid- (spores) w i t h confluent hilar appendage; tear-shaped ( V o l .
hemispherical - (pileus) w i t h shape o f a h a l f sphere ( V o l . 1, F i g . l , F i g s . 33.16, 33.17).
29.10). lageniform - (cystidia) characterized by neck narrower than half w i d t h
heterogeneous - (lamella edge) composed o f cystidia and basidia. of c e l l body ( V o l . 1, F i g . 34.25), see also narrowly lageniform and
heteromerous - (tissue) consisting o f hyphae and 'nests' o f sphaero- broadly lageniform.
cysts. lamellate - (hymenophore) consisting o f lamellae ( = gills).
hexagonal - (spores) six-angled ( V o l . 1, F i g s . 33.22, 33.23). lanate - ( = w o o l l y ) (pileus and stipe surfaces) covered w i t h long, soft,
hilar appendage - (of spores) short process at basal end o f spore by matted hairs.
w h i c h it was attached to sterigma ( V o l . 1, F i g . 3 2 . 4 C ) . lateral - (stipe) attached to one side o f pileus ( V o l . 1, F i g . 31.3).
hilum - scar left o n the hilar appendage o f the basidiospore after its lecythiform - (cystidia) lageniform and abruptly capitate ( V o l . 1, F i g .
discharge f r o m the sterigma. 34.33).
hirsute - (pileus and stipe surfaces) covered w i t h rather l o n g , rather length - (of spores) distance from apex to bottom as seen i n side v i e w
coarse hairs (coarser than i n pubescent, less coarse than i n hispid). ( V o l . 1, F i g . 32. 1).
hispid - (pileus and stipe surfaces) covered w i t h l o n g or short, erect leptocystidium - thin-walled c y s t i d i u m . O n l y used i n the combination
stiff hairs or bristles. cheiloleptocystidium, w h e n confusion w i t h cheilochrysocystidia
hispidulous - (pileus and stipe surfaces) minutely hispid. may occur.
lichenized - l i v i n g i n s y m b i o t i c association w i t h an alga, f o r m i n g a widest part above ( V o l . 1, F i g . 34.32).

lichen. obtuse - (apex o f cystidia) rounded ( V o l . 1, F i g . 34.41).
lignicolous - growing on wood. obtusely conical - (pileus) c o n i c a l w i t h rounded apex ( V o l . 1, F i g .
linear - (lamellae) w i t h straight l a m e l l a edge and parallel upper side 29.18).
( V o l . 1, F i g . 30.6). omphalinoid - see o m p h a l i o i d .
luminescent - (basidiocarp) fluorescent; g i v i n g light i n darkness. omphalioid (including o m p h a l i n o i d or c l i t o c y b o i d ) (habit) character-
marasmioid - (stipe) tough, slender, and y e l l o w - b r o w n , red-brown, or i z e d by pileus plano-convex to deeply infundibuliform; lamellae
grey-brown at least at base; - (habit) habit w i t h plicate pileus and decurrent ( V o l . 1, F i g s 28.4, 28.5, 28.6).
horsehair-like stipe, revivescent. opaque - (pileus) not translucent.
marginately bulbous - (base o f stipe) p r o v i d e d w i t h a b u l b w i t h a open pore type ofhilum - h i l u m w i t h a depression or perforation at one
raised border ( V o l . 1, F i g . 31.19). side and sometimes a perforation or tear at the other side o f the
marmorate - ( = marbled) pileus and stipe surfaces, and context) look- hilar appendage often connected by a slit (frequent i n thick-
i n g like marble: faintly and irregularly striped or innately veined. walled spores).
medallion clamp - a c l a m p connection w i t h an opening between the ovoid - egg-shaped (spores) ( V o l . 1, F i g . 33.8); - (cystidia) ( V o l . 1,
c l a m p connection itself and the elements connecting b y it. F i g . 34.7).
median constriction - (of spores) transverse contraction i n the middle, palisade - (pileipellis) special type o f h y m e n i d e r m , i n w h i c h the ele-
( V o l . 1, F i g s . 33.24, 33.25); - (of cystidia) ( V o l . 1, F i g . 34.30). ments are very l o n g and slenderly c y l i n d r i c a l .
mediostratum - m i d d l e layer. papilla - s m a l l n i p p l e - l i k e protuberance (on pileus) ( V o l . 1, F i g .
metachromatic - (spore wall) turning reddish to violet i n solution o f 29.29); - (on spores) ( V o l . 1, F i g . 33.28), see also abrupt and
C r e s y l blue i n H 0 .2
acute papilla.
metuloid - deep-rooting c y s t i d i u m b e c o m i n g d i s t i n c t l y t h i c k - w a l l e d papillate - (pileus surface) covered w i t h papillae.
and often incrusted w i t h age. paraboloid - (pileus) ( V o l . 1, F i g . 29.11).
micaceous - (pileus surface) w i t h glistening particles or spots. paravelangiocarpy - a type o f angiocarpic development o f the basid-
moniliform - (cystidia) c y l i n d r i c a l but contracted at regular intervals, iocarp i n w h i c h o n l y a partial v e i l participates.
l i k e a string o f beads ( V o l . 1, F i g . 34.35). parietal - (pigment) situated i n the h y p h a l w a l l .
monomitic - (tissue) built up o f one type o f hyphae. partial veil - see v e l u m partiale.
monovelangiocarpy - a type o f angiocarpic development o f the basid- patent - (hyphae or projections o f hyphae) perpendicular to surface o f
iocarp i n w h i c h o n l y a universal v e i l participates. stipe.
mitriform - (spores) shaped like a mitre (this volume, F i g . 2.3). pedicillate - (cystidia) p r o v i d e d w i t h a stalk ( V o l . 1, F i g . 34.48).
mucilaginous - consisting o f mucilage ( = viscous substance). pedunculate - (cystidia) provided w i t h a stalk ( V o l . 1, F i g . 34.48).
mucous - (pileus and stipe surfaces) slimy. pellicle - an easily peeling ixocutis.
mucronate - (cystidia) w i t h s m a l l abrupt, acute or blunt protuberance perisporium - the often m u c i l a g i n o u s layer o f the basidiospore w a l l
at apex ( V o l . l , F i g . 34.37). just inside the ectosporium; sometimes early disappearing, some-
mycenoid - (habit) characterized by pileus c o n i c a l to paraboloid; times f i l l i n g the spaces between exosporial ornamentation.
lamellae free to adnate; stipe u s u a l l y l o n g and slender, context phaseoliform - (spores) w i t h concave adaxial side, not parallel to abax-
usually brittle ( V o l . 1, F i g s . 28.7, 28.8). ial side ( V o l . 1, F i g . 33.20).
myxosporium - the set o f often mucilaginous layers on the outside o f pilangiocarpy - a type o f secondary angiocarpy i n w h i c h the original-
the basidiospore w a l l enveloping the eusporium; its components l y exposed hymenophore i n later stages is enclosed by (an out-
are ectosporium, p e r i s p o r i u m and e x o s p o r i u m . growth of) margin o f pileus.
narrowly adnate - (lamellae) ( V o l . 1, F i g . 30.20). pileipellis - cortical layer(s) o f pileus.
narrowly clavate - (cystidia, basidia) clavate w i t h Q = 4.0-8.0 ( V o l . 1, pileocarpy - a type o f development o f the basidiocarp i n w h i c h the
F i g . 34.12), see also very narrowly clavate. pileus is the first part initiated i n the p r i m o r d i a l bulb.
narrowly conical - (pileus) ( V o l . 1, F i g . 29.15); - (cystidia) conical pileocystidium - c y s t i d i u m situated o n surface o f pileus.
with Q > 4 ( V o l . 1, F i g . 34.15). pileostipitocarpy - a type o f development o f the basidiocarp i n w h i c h
narrowly cylindrical - (cystidia) c y l i n d r i c a l w i t h Q > 4 ( V o l . 1, F i g . the differentiation o f pileus and stipe starts at the same time
34.9). before that o f the hymenophore.
narrowly fusiform - (spores) fusiform w i t h Q > 4.0 ( V o l . 1, F i g . plage - (of spores) see suprahilar plage.
33.14); - (cystidia) fusiform w i t h Q > 4 ( V o l . 1, F i g . 34.19). plano-concave - (pileus) slightly concave ( V o l . 1, F i g . 29.20).
narrowly lageniform - (cystidia) ( V o l . 1, F i g . 34.24). piano-conical (pileus) slightly c o n i c a l , almost flat.
narrowly utriform — (cystidia) ( V o l . 1, F i g . 34.27). plano-convex - (pileus) slightly convex ( V o l . 1, F i g . 29.8).
necropigment - dark b r o w n i s h b l a c k pigment, usually i n f o r m o f intra- pleurocystidium - (lamella or tube) c y s t i d i u m situated o n sides o f the
cellular pigment-clots. hymenophore.
nettle-hair shaped - (cystidia) lageniform w i t h l o n g , narrow, slender pleurotoid- (including crepidotoid), (habit) characterized b y absent or
neck ( V o l . 1, F i g . 34.23). lateral stipe ( V o l . 1, F i g s . 28.12, 28.13).
nodulose type ofhilum - h i l u m an approximately circular area covered plicate - (pileus) folded radially, like a fan.
w i t h protuberances (frequent i n thin-walled spores). pluteoid - (habit) characterized by lamellae free; context o f pileus dis-
non-amyloid - (spore w a l l , spore ornamentation, h y p h a l w a l l s ) not continuous w i t h context o f stipe; stipe usually longer than d i a m -
changing colour or o n l y b e c o m i n g y e l l o w i s h i n M e l z e r ' s reagent. eter o f pileus ( V o l . 1, F i g . 28.1).
oblong - (spores) Q = 1.6-2.0 ( V o l . 1, F i g . 33.5); - (cystidia) Q > 1.6 polar view - (of spores) ( V o l . 1, F i g . 32.3).
( V o l . l , F i g . 34.1). primary angiocarpy - a type o f angiocarpic development i n w h i c h the
obovoid - (spores) reversely o v o i d , w i t h the broadest and widest part primordial h y m e n i u m is initiated i n a c l o s e d cavity.
uppermost, ( V o l . 1, F i g . 33.9); - (cystidia) ( V o l . 1, F i g . 34.8). primary mycelium - the uninucleate m y c e l i u m produced by a germi-
obpyriform - (cystidia) reversely p y r i f o r m , w i t h the broadest and nating basidiospore.
primordium - a very young, not fully differentiated basidiocarp. Schaeffer-reaction - (pileus surface and surface o f base o f stipe) cross-
protocarpic tuber - a non-persisting fleshy tuber o n w h i c h one or more reaction o f aniline and concentrated nitric acid.
basidiocarps may develop. sclerotium - a (long) persisting compact m y c e l i a l body.
pruinose - (pileus, l a m e l l a and stipe surfaces) covered w i t h a (often seceding - (lamellae) at first attached to stipe, but later separating from
white or whitish) powdery ' b l o o m ' . it ( V o l . 1, F i g . 30.22).
pseudoangiocarpy - secondary angiocarpy (see there). secondary angiocarpy - a type o f angiocarpic development i n w h i c h i n
pseudocystidium - differentiated prolongation o f vascular hypha into its later stages the p r i m o r d i a l h y m e n i u m is enveloped by hyphae
the h y m e n i u m . originating f r o m the stipe and/or the pileus.
pseudoinsititious - (stipe) at first sight appearing truly insititious, but o n segmentiform - (lamellae) w i t h straight l a m e l l a edges and convex upper
closer inspection (lens) w i t h very poorly developed basal m y c e l i - side ( V o l . 1, F i g s . 30.7, 30.8, 30.9).
um. separable pellicle - (pileus) pileipellis a (thick) ixocutis, easily peeling
pseudoparaphyses - short cells i n the h y m e n i u m , surrounding the off w i t h help o f a needle.
basidia. septate - (hyphae) w i t h septa.
pseudorhiza - a root-like extension o f the stipe (Vol. 1, F i g . 31.15). sericeous - ( = s i l k y ) (pileus and stipe surfaces) covered w i t h fine,
pubescent - (pileus and stipe surfaces) ( = downy) covered w i t h soft, straight, appressed, glossy hairs or fibrils.
fine hairs. serrate - (lamella edge) toothed like a saw ( V o l . 1, F i g . 30.35).
pulverulent - (pileus and stipe surfaces) covered w i t h powder. serrulate - (lamella edge) minutely serrate (Vol. 1, F i g . 30.36).
pustulate - (stipe and pileus surfaces) w i t h small rounded warts. sessile - (cystidia) without a stalk ( V o l . 1, F i g . 34.47).
pyriform - (cystidia) pear shaped (Vol. 1, F i g . 34.31). setiform - (cystidia) narrowly conical w i t h thickened w a l l (Vol. 1, F i g .
quadrangular - (spores) w i t h shape o f rectangle or square; Q equal or 34.16).
less than 1.15 (Vol. 1, F i g . 33.10). Siccus-type - see broom-cells.
radially rimose - (pileus surface) marked w i t h numerous, superficial, siderophilous - (particles i n basidia) turning b l a c k i s h purple or black-
radial clefts or cracks. ish violet i n acetocarmine i n presence o f metal ions.
radiate growth - (hymenophoral trama) formed by hyphae g r o w i n g . side-view - (of spores) (Vol. 1, F i g . 32.1).
radially at the underside o f the pileus from centre towards margin sinuate - (lamellae) having a concave indentation near the stipe (Vol. 1,
o f pileus. F i g . 30.24).
Ramealis structure - (of pileipellis) w i t h irregularly shaped and skeletal hyphae - aseptate, t h i c k - w a l l e d , straight o f slightly flexuous
arranged, nodose or en brosse or diverticulate elements ( V o l . 3, hyphae.
F i g . 122). slightly depressed - (pileus) w i t h shallow central depression ( V o l . 1,
reduced - (of stipe) very short ( V o l . 1, F i g . 31.3). F i g . 29.24).
reflexed - (margin o f pileus) bent upwards (Vol. 1, F i g . 29.40). smooth - (pileus and stipe surfaces) without elevations, ridges, grooves,
regular - (hymenophoral trama) having parallel hyphae ( V o l . 1, F i g . veins, etc.
35.1). solid - (stipe) made up o f homogeneous tissue. (Vol. 1, F i g . 31.9).
regular epithelium - an epithelium made up of elements i n erect rows solitary - (basidiocarp) single.
( V o l . 1, F i g . 3 6 . 4 A ) . spathuliform - (pileus) elliptic or oblong tapering gradually towards
reniform - (pileus) kidney-shaped ( V o l . 1, F i g . 29.6). eccentric or lateral stipe (Vol. 1, F i g . 29.3).
repent - (hyphae) creeping, not ascending. spheropedunculate - (cystidia) globose or subglobose w i t h long stalk
resinous — consisting o f or covered w i t h resin or resin-like substance. (Vol. 1, F i g . 34.6).
reticularly venose - (pileus surface) marked with anastomosing veins spinulose - (spores) covered w i t h s m a l l spines.
forming angular patches. spiny - covered w i t h spines, see also spinulose.
revivescent - (basidiocarp) r e v i v i n g after desiccation w h e n remoist- squame - small scale.
ened, restoring the fresh habit and producing spores again. squamose - (pileus and stipe surfaces) covered w i t h coarse (appressed)
revolute - (margin o f pileus) r o l l e d back (Vol. 1, F i g . 29.41). scales.
rhizomorph - a visible root-like m y c e l i a l strand (Vol. 1, F i g . 31.14). squamule - scale.
rhomboid - (spores) more or less quadrangular spore w i t h concave squamulose - (pileus and stipe surfaces) covered w i t h minute scales.
sides (this volume, F i g . 2.2). squarrose - (pileus and stipe surfaces) covered w i t h projecting, coarse
rimose - see radially rimose; areolate-rimose. scales.
rimulose - (pileus surface) minutely rimose. squarrulose - (pileus and stipe surfaces) covered w i t h small projecting,
rostrate - (cystidia) provided w i t h a beak-like extension at apex (Vol. coarse scales.
1, F i g . 34.38). statismosporic basidium - a basidium that does not discharge its spores;
rostrum - beak-like extension at the top o f a cystidium. they simply break off from the basidium.
Rotalis-type - see broom-cells. sterile - (lamella edge) composed o f cystidia only.
rounded flabelliform - (pileus) ( V o l . 1, Figs. 29.4, 29.5). stipitipellis - cortical layer o f stipe.
rounded triangular - (spores) rounded three-angled (Vol. 1, F i g . 33.15). stipitocarpy - a type o f development o f the basidiocarp i n w h i c h the
rugose - (pileus surface) irregularly w r i n k l e d . first differentiating hyphae o f the p r i m o r d i u m are those o f the
rugulose - (pileus surface) w i t h minute irregular wrinkles. stipe.
sarcodimitic - (tissue) consisting o f generative hyphae and chains o f straight - (margin o f pileus) not bent upwards or inwards (Vol. 1, Figs.
very long, thin- to t h i c k - w a l l e d elements ('sarcoskeletals'). 29.34, 29.35, 29.36).
sarcoskeletals - see sarcodimitic. striate - (pileus and stipe surfaces) marked w i t h regular lines.
sarcotrimitic - (tissue) consisting o f (1) generative hyphae, (2) chains strigose - (pileus and stipe surfaces) covered w i t h long, coarse or thick,
o f very l o n g thin- to thick-walled elements ('sarcoskeletals') and rather stiff hairs.
(3) t h i c k - w a l l e d b i n d i n g hyphae. stuffed - (stipe) having central part filled w i t h tissue different from tis-
scabrous - (stipe surface) scurfy. sue i n outer part ( V o l . 1, F i g . 31.10).
sub- - slightly or almost, under. twisted - (stipe) fibrils ranged spirally round axis because o f the base
subbulbous - (base o f stipe) slightly bulbous ( V o l . 1, F i g . 31.16). o f stipe being rotated w i t h regard to apex.
subcapitate - (apex o f cystidia) h a v i n g a k n o b ( V o l . 1, F i g . 34.43). umbilicate - (pileus) h a v i n g a n a v e l - l i k e depression ( V o l . 1, F i g .
subclavate - (stipe) slightly club-shaped ( V o l . 1, F i g . 31.7). 29.27), see also deeply umbilicate.
subcylindrical - (spores) Q = 2.0-3.0 ( V o l . 1, F i g . 33.6). umbo - (on pileus) broad rounded k n o b ( V o l . 1, F i g . 29.33).
subdecurrent - (lamellae) slightly decurrent, angle lamella-stipe 60- umbonate - (pileus) with broad rounded k n o b ( V o l . 1, F i g . 29.33).
8 0 ° . ( V o l . 1, F i g . 30.27). undate - (margin o f pileus) w a v y ( V o l . 1, F i g . 29.43); - (lamellae)
subglobose - (spores) nearly spherical Q = 1.05-1.15 ( V o l . 1, F i g . undulating l a m e l l a edge ( V o l . 1, F i g . 30.31).
33.2); - (cystidia) Q = 1.05-1.15 ( V o l . 1, F i g . 34.4). undulate - minutely undate (margin o f pileus) ( V o l . 1, F i g . 29.44); -
subgregarious - (basidiocarps) g r o w i n g i n a s m a l l group or g r o w i n g i n (lamella edge) ( V o l . 1, F i g . 30.32).
a group o f widespread specimens. uniguttulate - (spores) w i t h one droplet ( V o l . 1, F i g . 32.6).
subpellis - lower layer o f (pilei-)pellis. uninucleate - with one nucleus.
subregular - (hymenophoral trama) h a v i n g slightly flexuous, nearly universal veil - see v e l u m universale.
parallel hyphae ( V o l . 1, F i g . 35.2). urticiform — (cystidia) lageniform w i t h a long, tapering neck, bearing
subumbilicate - (pileus) having a s m a l l navel-like depression ( V o l . 1, needle-shaped crystals, reminiscent o f the nettle-cells o f Urtica
F i g . 29.26). species. T w o types are distinguished i n the genus Melanoleuca:
subumbonate - (pileus) w i t h low, broad u m b o ( V o l . 1, F i g . 29.32). brevipes-type i n w h i c h the upper c e l l is slender, c y l i n d r i c a l , and
subventricose - (lamellae) w i t h slightly convex l a m e l l a edge ( V o l . 1, distinctly separated from the lower c e l l , and the septum without
F i g . 30.10). median spot (this volume, F i g . 2.4, and exscissa-type with upper
sulcate - (pileus and stipe surfaces) w i t h grooves. c e l l gradually attenuated towards apex, and base o f upper c e l l
suprahilar depression - (of spores) s i n k i n g just above the hilar rather w i d e (this volume, F i g . 2.5).
appendage ( V o l . 1, F i g . 3 2 . 4 B ) . utriform - (cystidia) characterized b y neck broader than h a l f w i d t h of
suprahilar plage - (of spores) rounded, smooth area just above the c e l l body ( V o l . 1, F i g . 34.28), see also narrowly utriform and
hilar appendage ( V o l . 1, F i g . 3 2 . 5 D ) . broadly utriform.
tapering downwards - (stipe) b e c o m i n g narrower f r o m apex to base vascular hyphae - usually aseptate, often irregular, flexuous hyphae
( V o l . 1, F i g . 31.5). w i t h refractive contents.
tapering upwards - (stipe) b e c o m i n g narrower from base to apex ( V o l . velar sock - (stipe) covering o f v e i l on lower part o f stipe, f o r m i n g a
l . F i g . 31.6). sock.
thick-walled - (spores) ( V o l . 1, F i g . 32.6F). velum partiale - (= partial veil) p r i m o r d i a l tissue between the
tibiiform - (cystidia) lageniform w i t h a l o n g neck and capitate ( V o l . 1, hymenophore and the stipe and tissues differentiating from this
F i g . 34.34). p r i m o r d i a l tissues.
tomentose - (pileus and stipe surfaces) densely covered w i t h matted velum universale - (=universal veil) outer differentiated layer envelop-
(more or less appressed), soft hairs. i n g the p r i m o r d i a l basidiocarp and tissues differentiating from
tooth - (of lamellae) see decurrent tooth. this layer.
tortuous - twisted o f bent i n different directions. velutinous - ( = v e l v e t y ) (pileus and stipe surfaces) densely covered
trabecular - (hymenophoral trama) h a v i n g transversely oriented ele- w i t h fine, short, erect hairs.
ments. venose - (pileus surface) w i t h vein-like wrinkles, see also reticularly
tract - (spores) channel i n the spore w a l l leading to the germ pore. venose.
transition between hymeniderm and epithelium - ( V o l . 1, F i g . 3 6 . 3 D ) . ventricose - (lamellae) w i t h convex l a m e l l a edge ( V o l . 1, F i g s . 30.11,
transvenose - (lamellae) p r o v i d e d w i t h veins o n the surface ( V o l . 1, 30.12, 30.13), see also broadly ventricose; - (stipe) considerably
F i g . 31.1). broader i n the m i d d l e than at base and apex.
triangular - (lamellae) ( V o l . 1, F i g . 30.17); - (spores), see rounded tri- verrucose (pileus and stipe surfaces, spores) covered w i t h wart-like
angular. elevations; - (cystidia) w i t h small h o l l o w or solid protuberances
trichoderm - a p i l e i p e l l i s made up o f erect, straight elements, septate, (Vol. 1, F i g . 34.46).
and/or not originating at the same level ( V o l . 1, F i g . 3 6 . 2 A ) , see verruculose - (pileus and stipe surfaces, spores) covered w i t h fine,
also intricate and irregular trichoderm. wart-like elevations, see also verrucose.
trichohymeniderm - a h y m e n i d e r m made up o f elements w i t h Q > 6 very broadly fusiform - (spores) fusiform w i t h Q = 1.15-1.5 ( V o l . 1,
( V o l . 1, F i g . 3 6 . 3 A ) . F i g . 33.11); - (cystidia) fusiform w i t h Q < 1.5 ( V o l . 1, F i g .
tricholomatoid - (habit) characterized b y lamellae neither free, nor 34.22).
decurrent; stipe about the same length as pileus diameter or some- very narrowly clavate - (basidia) clavate w i t h Q > 8.0.
what longer; context fleshy; context o f pileus continuous w i t h villose - (pileus and stipe surfaces) covered w i t h fairly l o n g , soft, more
context o f stipe ( V o l . 1, F i g s . 28.9, 28.10, 28.11). or less straight, not interwoven hairs.
truncate - (apex o f spores) ending abruptly as i f cut off, ( V o l . 1, F i g . virgate - (pileus and stipe surfaces) streaked.
33.29); - (apex o f cystidia) ( V o l . 1, F i g . 34.42). viscid - (pileus and stipe surfaces) sticky.
truncately broadly conical - (pileus) broadly c o n i c a l w i t h as i f cut off water-spots - (surface o f pileus and stipe) w i t h roundish, darker spots,
apex ( V o l . l . F i g . 29.16). as i f stained b y drops o f water.
truncately conical - (pileus) c o n i c a l w i t h as i f cut off apex ( V o l . 1, F i g . width - (of spores) largest distance between sides as seen i n side-view
29.17). ( V o l . l , F i g . 32.1).
tubular - (hymenophore) consisting o f tubes. zonate - (pileus surface) w i t h concentric paler and darker alternating
tubuliform - (trama elements) tube-shaped. coloured zones or bands.
CHAPTER 3
Abbreviations of authors' names in this volume

E . C . V E L L I N G A , M . M . N A U T A & M . E . NOORDELOOS
A . & S. - J . B . v o n A l b e r t i n i & L . D . von S c h w e i n i z Bres. - G . Bresadola

Albert - L . Albert Bresinsky - A . Bresinsky
Alessio - C . L . Alessio B r e s i n s k y & Haas - A . B r e s i n s k y & H . Haas
A n g e l - F. A n g e l Britz. - M . Britzelmayer
Anon. - Anonymus R. Brown - R. Brown
Arnolds - E . J . M . Arnolds B r u n e l l i - F. B r u n e l l i
A r n o l d s & Veerkamp - E . J . M . A r n o l d s & M . T . Veerkamp B u l l . - J.B . F B u l l i a r d
Arora - D . Arora Candusso - M . Candusso
Atk. - G.F. Atkinson Cappelli - A . Cappelli
B . & Br. - M . J . Berkeley & C . E . Broome Cejp - K . Cejp
B . & C . - M . J . B e r k e l e y & M . A . Curtis Cetto - B . Cetto
Babos - M . B a b o s Charbonnel - J. Charbonnel
B a g l . - F. Baglietto Chatin & B o u d . - G . A . Chatin & E . Boudier
Ballero - M . Ballero Cherubini - A . Cherubim
Barbier - M . Barbier Chev. - F.F. C h e v a l l i e r
B a r l a - J.H.J.B. Barla Chiari - M . Chiari
Bas - C . Bas Chiusa - L . L a Chiusa
Batt. - G . A . Battara C h o d . & M a r t . - R . H . Chodat & C . E . M a r t i n
Beeli - M . Beeli Chrispijn - R. Chrispijn
B e l l u - F. B e l l u Clel.-J.B.Cleland
Bender - H . Bender Clemencon - H . Clemencon
A . v.d. B e r g - A . P . van der B e r g Clericuzio - M . Clericuzio
F A . v.d. B e r g h - F . A . van den B e r g h Coccia - M . Coccia
Berk. - M . J . Berkeley C o h n - F. C o h n
Besl - H . Besl C o l l i n - P. C o l l i n
W. Beyer - W. Beyer Contu - M . Contu
Big. & Guill. - R. Bigeard & H . Guillemin Contu & Currelli - M . Contu & L . Currelli
Bizio - E. Bizio C o n t u & Serra - M . C o n t u & D . Serra
Bizzi - A . Bizzi Cooke - M . C . Cooke
B o d i n & P r i o u - M . B o d i n & J.P. P r i o u Cool - C. Cool
Bohus - G . Bohus C o r d a - A . K . J . Corda
Boisselet & M i g l . - P. Boisselet & V . M i g l i o z z i Courtecuisse - R . Courtecuisse
B o l t . - J. B o l t o n Courtec. & D u h e m - R . Courtecuisse & B . D u h e m
M . Bon - M . Bon C z e r n . - V . M . Czernajew
B o n & B e l l u - M . B o n & F. B e l l u Dahncke - R . M . D a h n c k e
B o n & Boiff. - M . B o n & J. Boiffard Dahncke & D a h n c k e - R . M . D a h n c k e & S . M . D a h n c k e
B o n & Candusso - M . B o n & M . Candusso D a l S a v i o - F. d a l S a v i o
B o n & Cappelli - M . B o n & A . Cappelli D a m b l o n & L a m b i n o n - J. D a m b l o n & J. L a m b i n o n
B o n & Chevassut - M . B o n & G . Chevassut D a n i e l - A r r a n z - J. D a n i e l - A r r a n z
B o n & C o l l i n - M . B o n & P. C o l l i n Demoulin - V. D e m o u l i n
B o n & Courtec. - M . B o n & R . Courtecuisse D e r b s c h & Schmitt - H . D e r b s c h & J . A . Schmitt
B o n & G u i n b . - M . B o n & J. Guinberteau Donk - M . A . Donk
B o n & H a l u w i j n - M . B o n & C h . van H a l u w i j n Dorfelt - H . Dorfelt
B o n & M i g l . - M . B o n & V. Migliozzi D0ssing - L . D0ssing
B o n & Orton - M . B o n & P . D . Orton Earle - F . S . E a r l e
B o n & G . Riousset - M . B o n & G . Riousset Einh. - A . Einhellinger
Boud. - J.L.E. Boudier Enderle - M . Enderle
Bousset & Joss. - M . Bousset & M . Josserand Enderle & L a u x - M . Enderle & H . E . L a u x
B r e b i n a u d - P. B r e b i n a u d Essette - H . Essette
Breitenb. & K r a n z l . - J. Breitenbach & F. K r a n z l i n Esteve-Rav. & A l t e s - F. Esteve-Raventos & A . A l t e s
12
ABBREVIATIONS OF AUTHORS' NAMES (Vellinga, Nauta & Noordeloos)
Eyre - W.L.W. Eyre C . H . K a u f f m . - C . H . Kauffman

J . Favre - J . Favre Kelderman - P H . Kelderman
Fay. - V . F a y o d K e l l e r - J . Keller
Fillion - R. Fillion Kerrigan - R . W . K e r r i g a n
G . Fischer - G . F i s c h e r Knudsen - H . Rnudsen
F o r t i - P. Forti K o b l e r - B . Kobler
Fourre - G . Fourre R. d e K o k - R . P J . d e K o k
Fr. - E . M . Fries K o k & V e l l i n g a - R.P.J, de K o k & E . C . V e l l i n g a
A . E . Freeman - A . E . H . Freeman Konr. - P. K o n r a d
G a l l i n a r i & T o m a s i - A . G a l l i n a r i & R . Tomasi Konr. & M . - P. K o n r a d & A . M a u b l a n c
Gea & Honrubia - F J . Gea & M . Honrubia Korhonen - M . K o r h o n e n
Genev. - L . G . Genevier Kreisel - H . Kreisel
Gennari - A . Gennari Krieglst. - G . J . Krieglsteiner
Gerault & G i r r e - A . Gerault & L . G i r r e L . Krieglst. - L . Krieglsteiner
Gerhardt - E . Gerhardt K r o m b h . - J.V. v o n K r o m b h o l z
Gidholm - L . Gidholm Kuhner - R. Kiihner
E.J. Gilb. - E . J . Gilbert Kiihner & Romagn. - R. Kuhner & H . C . L . Romagnesi
Gillet - C . C . Gillet K u m m . - P. K u m m e r
Gminder - A . Gminder O. Kuntze - C . E . O . Kuntze
Godey - Godey K u y p . & Schreurs - T h . W . K u y p e r & J . Schreurs
Godfrin - J. Godfrin L . - C . Linnaeus
S.F. G r a y - S.F. G r a y Laber - D . Laber
GxilU - E . G r i l l i L£ess0e - T. La;ss0e
Groves - J.W. Groves Chr. L a n g e - Chr. L a n g e
Gueguen - F. Gueguen J. L a n g e - J . E . L a n g e
G u i n b . - J . Guinberteau Lanzoni - G . Lanzoni
G u i n b . & B o d i n - J . Guinberteau & M . B o d i n Lasch - W.G. Lasch
H a l l e r - R . Haller Lavorato - C . Lavorato
B . Hanff - B . Hanff L a z z a r i & B e l l u - G . L a z z a r i & F. B e l l u
Hansen - L . Hansen Lincoff - G . Lincoff
Hardtke & R o d e l - H . - J . Hardtke & T. R o d e l Locq. - M . V Locquin
A . H a u s k n . - A . Hausknecht Lohmeyer - T.R. Lohmeyer
Heilmann-Clausen - J. Heilmann-Clausen Lonati - G . Lonati
R. Heim - R. Heim L u n d . & Nannf. - S. L u n d e l l & J . A . Nannfeldt
H e i m & Becker - R. H e i m & G . Becker Liitjeharms - W . J . Liitjeharms
H e i m & Romagn. - R. H e i m & H . C . L . Romagnesi M a k e - R.C.J.E. Maire
H e i n e m . - P. H e i n e m a n n M a i . & Bert. - J . L . G . M a l e n c o n & R . Bertault
P. H e n n . - P. Hennings Malenc. - J . L . G . Malencon
Herink - J. Herink Marchand - A . Marchand
Hlavacek - J. Hlavacek M a s s . - G . E . Massee
Hohn. - F . X . R . von Hdhnel Massart - F. Massart
H0iland - K . H0iland Mattirolo - O. Mattirolo
H o f f m . - G . F . Hoffmann Mattuschka - H . G . von M a t t u s c h k a
H o n g o - T. H o n g o Melzer - V. Melzer
Hora - K B . Hora Menier - C h . Menier
Horak - E . Horak M . Meusers - M . Meusers
H o r n s c h . - Chr.F. H o r n s c h u c h G . Meyer - G.F.W. Meyer
H o t s o n & Stuntz - J . W . H o t s o n & D . E . Stuntz Michael - E . Michael
Hotz - O. Hotz Michael & Hennig - E . Michael & B . Hennig
Huds. - W . Hudson M i g l . - V. M i g l i o z z i
Huijser - H . A . Huijser M i g l . & Rava - V. M i g l i o z z i & M . Rava
Huijsman - H . S . C . Huijsman O . K . M i l l e r - O . K . M i l l e r , Jr.
Imaz. - R . I m a z e k i Mitchell & Bresinsky - A . D . Mitchell & A . Bresinsky
Imbach - E . J . Imbach M o h r - P. M o h r
Imler - L . Imler F . M 0 l l e r - F . H . M0ller
Johnson & V i l g a l y s - J . Johnson & R . V i l g a l y s M 0 l l e r & J. Lange - F . H . M 0 l l e r & J.E. Lange
Joss. - M . Josserand M 0 l l e r & Schaeff. - F . H . M 0 l l e r & J . Schaeffer
Joss. & L . Riousset - M . Josserand & L . Riousset M o n t . - P. M o n t a g n e
J u l i c h - W . F . B . Jttlich Moreno - G . Moreno
K a l c h b r . - K . Kalchbrenner Morg. - A.P. Morgan
P. Karst. - R A . Karsten Mornand - J. Mornand
Kasparek - F. Kasparek Mos. - M . M . Moser
M o s . & Jul. - M . M . Moser & W . R B . Jiilich J. Schaeff. - J . Schaeffer

M o s . & S i n g . - M . M . M o s e r & R . Singer J. Schaeff. & M 0 l l e r - J . Schaeffer & F . H . M 0 l l e r
O.F. M u l l . - O . F . M i i l l e r J. Schaeff. & Steer - J . Schaeffer & Steer
Murrill - W.A. Murrill Schatzle & Ottmann - E . Schatzle & G . Ottmann
N a r d u c c i & C a r o t i - R . N a r d u c c i & V . Caroti Schnizl. - A . C . F . H . C . Schnizlein
Nauta - M . M . Nauta Schrank - F. v o n Paula von Schrank
P.D. Orton - P.D. Orton Schreurs - J . Schreurs
Over. - C . van O v e r e e m Schroet. - J . Schroeter
Pacioni - G . Pacioni S. S c h u l z . - S. Schulzer v o n M u g g e n b u r g
P a n i z z i - F. P a n i z z i Schulz-Weddigen - J. Schulz-Weddigen
Papeti - C . Papeti S c h u m . - H . C . F . Schumacher
Parra - L . A . Parra Schwobel - H . Schwobel
Partacini - G . Partacini Scop. - G . A . S c o p o l i
Pat. - N . T . Patouillard Seer. - L . Secretan
Patane - F. Patane Serafin - Serafin
P a u l . - J.J. Paulet Signorello - P. Signorello
Pazmany - D . Pazmany S i n g . - R . Singer
A . Pears. - A . A . Pearson Sing. & C l e m . - R . Singer & H . C l e m e n c o n
Peck - C . H . Peck Sing. & D i g i l i o - R . Singer & A . P . L . D i g i l i o
P e g l . - D . N . Pegler A . L . Sm. & Rea - A . L . Smith & C . Rea
P e g l . & B r a n d - D . N . Pegler & A . W . B r a n d A . H . Smith - A . H . Smith
Pegl. & C a l o n g e - D . N . Pegler & F . D . Calonge H.V. Smith - H . V Smith
P e g l . & L e g o n - D . N . Pegler & N . W . L e g o n W . G . S m . - W . G . Smith
Perrone - L . Perrone Sow. - J . Sowerby
Pers. - C . H . Persoon Speg. - C . L . Spegazzini
J. Petersen - J . H . Petersen Stridvall - L . S t r i d v a l l
Petch - T. Petch Stridvall & A . Stridv. - L . S t r i d v a l l & A . S t r i d v a l l
R. Phillips - R. Phillips Sturm - J . Sturm
P i l a t - A . Pilat Sundb. - W . J . Sundberg
Pilat & U s a k - A . P i l a t & O . U s a k Svrcek - M . Svrcek
Pilat & P o u z . - A . P i l a t & Z . Pouzar Szujko-Lacza - J. Szujko-Lacza
Potz - H . Potz Tabares - M . Tabares
Q u e l . - L . Quelet T r i m b . - J. T r i m b a c h
Q u e l . & B e r n . - L . Quelet & G . Bernard Trimb. & Augias - J. Trimbach & Augias
Rab. - G . L . Rabenhorst Ulje - C . B . Ulje
Raithelhuber - J . Raithelhuber Vasas - G . Vasas
Raid - E . Raid Velen. - J. Velenovsky
Rauschert - S. Rauschert Vellinga - E . C . Vellinga
Rea - C. Rea Vesterholt - J . Vesterholt
Redhead - S . A . Redhead V i l a - J. V i l a
D. Reid - D . A . Reid Vitt. - C . V i t t a d i n i
Reid & Eicker - D . A . Reid & A . Eicker V o g l . - P. V o g l i n o
Reijnders - A . F . M . Reijnders G . A . de V r i e s - G . A . de V r i e s
Relh. - R. Relhan W a k . & Pears. - E . Wakefield & A . A . Pearson
Richon & Roze - C . E . Richon & E . Roze Wallr. - C . F . W . W a l l r o t h
J. R i c k - J . R i c k Wasser - S.P. Wasser
G . Riousset - G . Riousset W e h o l t - 0 . Weholt
L . Riousset - L . Riousset Weinm. - J . A . Weinmann
Rocabruna - A . Rocabruna Wichansky - E . Wichansky
Rodriguez Armas - L . Rodriguez Armas Wilhelm - M .Wilhelm
Romagn. - H . C . L . Romagnesi Winterhoff & B o n - W . Winterhoff & M . B o n
R y m a n & H o l m a s e n - S. R y m a n & I. H o l m a s e n W u i l b a u t - J.J. W u i l b a u t
Sacc. - P . A . Saccardo Zalin & M i g l . - G . Zalin & V. Migliozzi
Sacc. & S y d . - P . A . Saccardo & P. S y d o w Zecchin - G . Zecchin
Sacc. & Trott. - P . A . Saccardo & A . Trotter Zeller - S . M . Zeller
Saut. - A . E . Sauter Zschieschang - G . Zschieschang
Schaeff. - J . C . Schaeffer Zuccherelli - A . Zuccherelli
CHAPTER 4
Bibliographic abbreviations
E . C . V E L L I N G A , M . M . NAUTA & M . E . NOORDELOOS
a. Books tif des champignons superieurs de l a zone subalpine du Pare

N a t i o n a l Suisse. In Ergebn. wiss. Unters. schweiz. NatnParks, n . F
A r n o l d s & Veerkamp, G i d s Paddest. Meetnet - G i d s voor de pad- V I , 42.
destoelen i n het meetnet. J. Favre, C h a m p , sup. Z o n e alpine - L e s champignons superieurs de l a
A r n o l d s et a l . , O v e r z . Paddest. N e d e r l a n d - O v e r z i c h t van de pad- zone alpine d u Pare N a t i o n a l Suisse. In E r g e b n . wiss. Unters.
destoelen i n N e d e r l a n d . schweiz. NatnParks, n.F. V , 33.
A r o r a , M u s h r . demyst. - M u s h r o o m s demystified. Fr., Epicrisis - E p i c r i s i s systematis m y c o l o g i .
B . & B r . , N o t i c . B r . F u n g i - Notices o f B r i t i s h fungi. In various jour- Fr., H y m e n o m y c . eur. - H y m e n o m y c e t e s europaei sive Epicriseos sys-
nals. tematis m y c o l o g i c i .
B a r l a , F l . m y c o l . i l l . - F l o r e m y c o l o g i q u e illustree. L e s champignons Fr., Ic. sel. H y m e n o m y c . - Icones selectae h y m e n o m y c e t u m selectum
des A l p e s - M a r i t i m e s . nondum delineatorum.
B a s et a l . , F l . agar, neerl. - F l o r a agaricina neerlandica. Fr., M o n o g r . H y m e n o m y c . Sueciae - M o n o g r a p h i a H y m e n o m y c e t u m
B i g . & G u i l l . , F l . C h a m p , sup. France - F l o r e des champignons Sueciae.
superieurs de France. Fr., Syst. m y c o l . - Systema m y c o l o g i c u m .
B o l t . , H i s t . F u n g . H a l i f a x - A history o f fungusses g r o w i n g i n Halifax. Fr., Syst. m y c o l . , Ind. g e n . - Systema m y c o l o g i c u m , Index generalis.
M . B o n , C h a m p . Eur. o c c i d . - C h a m p i g n o n s d ' E u r o p e occidentale. Fr., Syst. Orb. veg. - Systema orbis vegetabilis.
M . B o n , F l . m y c o l . Eur. 3, Lepiotes - F l o r e m y c o l o g i q u e d ' E u r o p e 3. Gerhardt, G r . Pilzf. - GroBe B L V Pilzfuhrer.
L e s Lepiotes. D o c . m y c o l . M e m o i r s hors serie no. 3. Lepiotaceae E . J . G i l b . , Genre A m a n i t a - L e genre A m a n i t a Persoon.
Roze. Gillet, C h a m p . France - L e s champignons (fungi, hymenomycetes) q u i
B o u d . , Ic. m y c o l . - Icones m y c o l o g i c a e . croissent en France et description et iconographie, proprietes utiles
Breitenb. & K r a n z l . , P i l z e S c h w e i z - P i l z e der S c h w e i z . ou veneneuses.
Bres., F u n g i trident. - F u n g i tridentini. G i l l e t , Hymenomycetes - L e s H y m e n o m y c e t e s .
Bres., Iconogr. m y c o l . - Iconographia m y c o l o g i c a . S.F. Gray, Nat. A r r . B r . PI. - N a t u r a l Arrangement o f B r i t i s h Plants.
B u l l . , Herb. France - Herbier de l a France. Groves, E d i b l e pois. M u s h r . C a n a d a - E d i b l e and poisonous mush-
Candusso & L a n z o n i , L e p i o t a - L e p i o t a s.l. F u n g i europaei 4. rooms o f C a n a d a E d . 2. In Pubis D e p . A g r i c . C a n . 1112.
C a p p e l l i , A g a r i c u s - A g a r i c u s . F u n g i europaei 1. Hansen & K n u d s e n , N o r d i c M a c r o m y c . - N o r d i c M a c r o m y c e t e s .
Cetto, F u n g h i V e r o - F u n g h i dal vero. Hoffm., A b b . S c h w a m m e - A b b i l d u n g e n der Schwartime.
Cetto, G r . P i l z f . - D e r groBe Pilzfuhrer. H o r a k , S y n . G e n . A g a r . - Synopsis G e n e r u m A g a r i c a l i u m . In Beitr.
Chev., F l . generale Env. Paris - F l o r e generale des environs de Paris,... K r y p t o g . F l . S c h w e i z 13.
C h r i s p i j n , C h a m p . Jordaan - C h a m p i g n o n s i n de Jordaan. D e pad- Hornsch., S y l l . PI. nov. Ratisb. - S y l l o g e plantarum novarum itemque
destoelen van A m s t e r d a m . minus cognitarum a praesantissimis botanicis adhuc viventibus c o l -
C l e l . , Toadst. M u s h r . South A u s t r a l i a - Toadstools and mushrooms and lecta et a societate regia botanica Ratisbonensi edita.
other larger fungi o f South A u s t r a l i a . Imaz. & H o n g o , C o l . 111. M u s h r . Japan - C o l o r e d illustrations o f mush-
C l e m e n c o n , A n a t . H y m e n o m y c . - A n a t o m i e der H y m e n o m y c e t e n . rooms o f Japan.
C o h n , K r y p t . - F l . Schlesien - K r y p t o g a m e n f l o r a v o n Schlesien. Imaz. et al., F u n g i Japan - F u n g i o f Japan.
C o o k e , Handb. B r i t . F u n g i - H a n d b o o k o f B r i t i s h fungi. Jiilich, H i g h e r T a x a B a s i d i o m y c . - H i g h e r taxa i n Basidiomycetes. In
C o o k e , 111. B r i t . F u n g i - Illustrations o f B r i t i s h fungi. B i b l t h e c a m y c o l . 85.
C o o k e & Q u e l . , C l a v . syn. H y m e n o m y c . eur. - C l a v i s synoptica Kalchbr., Ic. sel. Hymenomyc. Hung. - Icones selectae
h y m e n o m y c e t u m europaeorum. H y m e n o m y c e t u m Hungariae.
C o r d a , Icon. F u n g . - Icones fugnorum hucusque cognitorum... P. Karst., R y s s l . , F i n l . S k a n d . H a l f 0 n s Hattsvamp. - Rysslands,
Courtec. & D u h e m , G u i d e C h a m p . France E u r o p e - G u i d e des Finlands o c h den S k a n d i n a v i s k a half0ns hattsvampar. In B i d r .
champignons de France et d ' E u r o p e . K a n n . F i n l . Nat. F o l k 32.
Diihncke, 1200 P i l z e - 1200 P i l z e . K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g - P a r a s o l z w a m m e n van Z u i d -
D i i h n c k e & D i i h n c k e , 700 P i l z e - 700 P i l z e i n Farbfotos. L i m b u r g . L e p i o t a s.l. e x c l . M a c r o l e p i o t a .
D e r b s c h & Schmitt, A t l . P i l z e Saarlandes - A t l a s der P i l z e des Keller, A t l . B a s i d i o m y c . - A t l a s des B a s i d i o m y c e t e s vus aux m i c r o -
Saarlandes. Teil 2: Nachweise, Okologie, Vorkommen und scopies electroniques.
Beschreibungen. Aus Natur und Landschaft im Saarland. Knudsen & Vesterholt, Truede Storsvampe Danmark - Truede
Sonderband 3. storsvampe i D a n m a r k - en rodliste.
Essette, Psalliotes - L e s Psalliotes. A t l a s m y c o l o g i q u e s 1. Konr. & M . , Ic. sel. F u n g . - Icones selectae fungorum.
J. Favre, Cat. descr. C h a m p , sup. Z o n e subalpine - Catalogue descrip- K r o m b h . , Naturg. A b b . Beschr. essb. schadl. verd. S c h w a m m e -
15
Naturgetreue A b b i l d u n g e n und Beschreibungen der essbaren, R y m a n & H o l m a s e n , Svampar - Svampar.

schadlichen und verdachtigen Schwamme. S a c c , S y l l . F u n g . - S y l l o g e fungorum.
K i i h n e r & R o m a g n . , F l . anal. C h a m p , sup. - F l o r e analytique des Schaeff., F u n g . Bavariae - F u n g o r u m q u i i n B a v a r i a et Palatinatu c i r c a
champignons superieurs. Ratisbonam nascuntur icones.
K i i h n e r & R o m a g n . , C o m p l . F l . anal. - Complements a la "Flore analy- Scop., F l . earn. - F l o r a c a m i o l i c a .
tique'. In various journals. B i b l t h e c a m y c o l . 56. Sing., A g a r . m o d . T a x o n . - T h e Agaricales i n modern taxonomy.
K u m m . , Ftihr. P i l z k . - D e r Fiihrer i n die Pilzkunde. A . H . S m i t h , M u s h r . nat. Hab. - M u s h r o o m s i n their natural habitats.
O . Kuntze, Rev. G e n . P L - R e v i s i o generum plantarum. W . G . S m . , Synopsis B r i t . B a s i d i o m y c . - Synopsis o f the B r i t i s h basid-
L . , Spec. PI. - Species plantarum. iomycetes.
J. Lange, F l . agar. dan. - F l o r a agaricina danica. Sow., C o l . F i g s E n g l . F u n g i - C o l o u r e d figures o f E n g l i s h fungi or
L a z z a r i & B e l l u , A t l . iconogr. - Atlante iconografico. Suppl. B o l l . G r . mushrooms.
m i c o l . G . Bres. 24. Sturm, Deutschl. F l . - Deutschlands F l o r a .
Lincoff, F i e l d G u i d e N . A m e r . M u s h r . - T h e A u d u b o n Society F i e l d S z u j k o - L a c z a , F l . Hortobagy natn. P a r k - F l o r a o f the Hortobagy
G u i d e to N o r t h A m e r i c a n Mushrooms. N a t i o n a l Park (Budapest).
M a i . & Bert., F l . C h a m p , sup. M a r o c - F l o r e des champignons Urbonas et a l . , K o n s p . agar. G r i b . L S S R , L S S R , E S S R - K o n s p e k t
superieurs du M a r o c . i z u c h e n i v a a g a r i k o v y k h gribov L i t o v s k e i S S R , L a t v i s o i i S S R ,
M a n z , Unters. K u l t u r Lebensz. Leucoagaricus naucinus, Macrolepiota E s t o n s k o i S S R (Materialy 1778-1973).
procera - Untersuchungen iiber die Kultur und den Lebenszyklus von Velen., C e s k e H o u b y - Ceske houby.
Leucoagaricus naucinus (Fr.) Sing, und Macrolepiota procera (Scop. Velen., N o v i t . m y c o l . - N o v i t a t e s m y c o l o g i c a e .
ex Fr.) Sing. Velen., N o v i t . m y c o l . nov. - Novitates m y c o l o g i c a e novissimae.
M a r c h a n d , C h a m p . N o r d M i d i - Champignons du nord et du m i d i . Vitt., Descr. F u n g h i mang. Italia - D e s c r i z i o n e dei F u n g h i mangerec-
Mass., B r i t . Fungus F l . - B r i t i s h fungus flora. c i p i u c o m u n i d e l l ' Italia.
Massart, Champignons - L e s champignons. O u et quand les trouver. Wasser, Tr. A g a r i c e a e Soviet U n i o n - T r i b e A g a r i c e a e Pat. o f the
G . M e y e r , Pr. F l . essequ. - Primitiae florae essequeboensis. Soviet U n i o n .
M i c h a e l , Fiihr. Pilzfr. - Fiihrer fur Pilzfreunde. < Wasser, A g a r i k o v y e G r i b y S . S . S . R . - A g a r i k o v y e G r i b y S . S . S . R .
M i c h a e l & H e n n i g , Handb. Pilzfr. - Handbuch fur Pilzfreunde. Wasser, F l . F u n g . R . S . S . U c r a i n i c a e , A g a r i c a c e a e - F l o r a fungorum
M i c h a e l et al., Handb. Pilzfr. - Handbuch fiir Pilzfreunde. R S S U c r a i n i c a e . Basidiomycetes, Agaricaceae C o h n .
M o s . , Blatter-, B a u c h p i l z e - D i e Blatter- und Bauchpilze. Z u c c h e r e l l i , F u n g h i Pinete Z o n e medit. - I funghi delle pinete delle
M o s . , R o h r l i n g e , Blatter-, B a u c h p i l z e - D i e Rohrlinge, Blatter- und zone mediterranee.
Bauchpilze.
M o s . , Rohrlinge Blatterpilze - Rohrlinge und Blatterpilze.
b. Journals and series
M o s . & Jiil., Farbatl. B a s i d i o m y c . - Farbatlas der Basidiomyceten.
Nauta & vellinga, A t l . N e d e r l . Paddest. - A t l a s van Nederlandse pad- A c t a M u s . nat. Prag. - A c t a musei nationalis Pragae (Sbormk narodnflio
destoelen. musea v Praze - Sb. nar. M u s . Praze).
Pat., Essai tax. H y m . - E s s a i taxonomique sur les families et les genres A c t a phytotax. bare. - A c t a phytotaxonomica barcinonesia.
des Hymenomycetes. Agarica - Agarica.
Paul., Tr. C h a m p . - Traite des champignons. A n . M u s . argent. C i e n c . nat. - A n a l e s del M u s e o argentino de Ciencias
P e g l , A g a r i c F l . Lesser A n t i l l e s - A g a r i c F l o r a o f the Lesser Antilles. naturales 'Bernardino Ravadavia' Buenos A i r e s .
K e w B u l l . add. Series 9. A n . M u s . nac. Hist. nat. B . A i r e s - A n a l e s del M u s e o national de histo-
Pegl., A g a r i c F l . S r i L a n k a - A g a r i c F l o r a o f S r i L a n k a . K e w B u l l . add. ria natural de Buenos Aires.
Series 12. A n . Soc. cient. arg. - Anales de la Sociedad cientifica argentina.
P e g l , Prel. A g a r i c F l . East A f r i c a - A preliminary A g a r i c flora of East A n n . M a g . nat. Hist. - A n n a l s and M a g a z i n o f Natural History.
A f r i c a . K e w B u l l . add. Series 6. A n n l s hist.-nat. M u s . natn. hung. - Annates historico-naturales M u s e i
Pers., Observ. m y c o l . - Observationes mycologicae. nationalis hungarici.
Pers., S y n . meth. F u n g . - Synopsis methodica fungorum. A n n l s m y c o l . - Annates m y c o l o g i c i .
Pers., Tent. D i s p . meth. F u n g . - T e n t a m e n dispositionis mehtodicae fun- A n n . R e p . N . Y . State B o t . - A n n u a l Report o f the N e w Y o r k State
gorum... Botanist.
R . Phillips, Paddest. S c h i m m . - Paddestoelen en schimmels van West- A n n l s Sci. nat., Bot. - Annales des sciences naturelles, ser. II, Botanique.
Europa. A t t i A c c a d . naz. L i n c e i M e m o r i e - A t t i d e l l ' A c c a d e m i a nazionale dei
Pilat, K l i c urc. H u b hfib. bedl. - K l i c k urcovani nasich hub hfibovitych Lincei. Memorie.
a bedlovitych. A t t i X I X C o m . scient. naz. Serina (Bergamo) - A t t i d i X I X Comitate
Pilat & Usak, Nase H o u b y - Nase Houby. Scientifico Nazionale, Serina (Bergamo).
Quel., C h a m p . Jura Vosges - L e s Champignons du Jura et des Vosges. B e i h . N o v a H e d w i g i a - Beihefte zur N o v a H e d w i g i a .
In various journals. B e i h . S y d o w i a - Beihefte zur S y d o w i a .
Quel., Enchir. F u n g . - E n c h i r i d i o n fungorum q u i i n E u r o p a media et B e i h . Z . P i l z k . - Beihefte zur Zeitschrift fur M y k o l o g i e .
praesertim i n G a l l i a vigentium. Beitr. K e n n t n . P i l z e Mitteleur. - Beitrage zur Kenntnis der P i l z e
Quel., F l . m y c o l . France - Flore mycologique de l a France et des pays Mitteleuropas.
limitrophes. B e l g . J . Bot. - B e l g i a n Journal o f Botany.
Rab., Deutschl. K r y p t . F l . - Deutschlands Kryptogamen-Flora. Ber. bayer. bot. Ges. - Bericht der Bayerischen botanischen Gesellschaft
Rea, Brit. B a s i d i o m y c . - B r i t i s h Basidiomycetae. zur Erforschung der heimischen F l o r a .
R i c h o n & R o z e , A t l . Champ, comest. veneneux - Atlas des champignons Ber. naturh. Ver. A u g s b u r g - Bericht des naturhistorischen Vereins i n
comestibles et veneneux de l a France et des pays circonvoisins. Augsburg.
BIBLIOGRAPHIC ABBREVIATIONS (Vellinga, Nauta & Noordeloos) 17
Ber. naturf. Ges. Augsburg - Bericht der naturforschenden Cryptog. M y c o l . - Cryptogamie, M y c o l o g i e .

Gesellschaft A u g s b u r g . D a n s k bot. A r k . - D a n s k botanisk A r k i v .
B i d r . K a n n . F i n l . Nat. F o l k - B i d r a g t i l k a n n e d o m o m Finlands natur Denkschr. bayer. bot. Ges. Regensburg - Denkschriften der
och folk. K b n i g l i c h e n B a y e r i s c h e n botanischen Gesellschaft z u Regensburg.
Blyttia - Blyttia. D o c . m y c o l . - Documents mycologiques.
B o l . S o c . B r o t . - B o l e t i m da Sociedade Broteriana. Ergebn. wiss. Unters. schweiz. NatnParks - Ergebnisse der w i s -
B o l . S o c . m i c o l . M a d r i d - B o l e t m de l a S o c i e d a d M i c o l o g i c a de senschaftlichen Untersuchungen des schweizerischen
Madrid. Nationalparks.
Bolets Catalunya - Bolets de Catalunya. F l . batava - F l o r a batava.
Boletus - Boletus. Fl. mycol. Eur. - Flore mycologique d'Europe. Documents
B o l l . A s s . micol. ecol. Romana - Bollettino dell'Associazione mico- mycologiques M e m o i r e hors Serie.
l o g i c a ed ecologica romana. F l . i c o n . C h a m p . C o n g o - F l o r e iconographique des champignons du
B o l l . Circ. m i c o l . G . Carini - Bollettino del Circolo micologico Congo.
«Giovanni Carini». F o o d C h e m . - F o o d Chemistry.
B o l l . G r u p p o m i c o l . G . Bres. - B o l l e t t i n o del G r u p p o m i c o l o g i c o ' G . Friesia - F r i e s i a .
Bresadola'. F u n g . rar. Ic. c o l . - F u n g o r u m rariorum icones coloratae.
B o l n A c a d . nac. C i e n c . C o r d o b a - B o l e t i n de l a A c a d e m i a n a t i o n a l de F u n g h i A m b . - F u n g h i e ambiente.
ciencias en C o r d o b a . F u n g i exs. suec. - F u n g i exsiccati sueci.
B o r b a s i a - Borbasia, Dissertationes botanicae. Fungus - Fungus.
Bot. B u l l . A c a d , sinica - B o t a n i c a l B u l l e t i n o f A c a d e m i a S i n i c a . Goteborgs S v a m p e k l . A r s s k r . - Goteborgs Svampeklubbs Arsskrift.
Bot. G a z . - B o t a n i c a l Gazette. Grevillea - Grevillea.
Bot. Tidsskr. - B o t a n i s k Tidsskrift. Hedwigia - Hedwigia.
B u l l . F e d . m y c o l . D a u p h . Savoie - B u l l e t i n trimestriel de l a Federation Iheringia, B o t . - Iheringia, B o t a n i c a .
M y c o l o g i q u e Dauphine-Savoie. J. Bot., Paris - Journal de Botanique, Paris.
B u l l . Inst, agron. Stns R e c h . G e m b l o u x - B u l l e t i n de l'lnstitut J. Jap. B o t . - Journal o f Japanese botany.
agronomique de l'etat et des stations de recherches de G e m b l o u x . J. M y c o l . - Journal o f M y c o l o g y .
Bull. Jard. bot. Buitenzorg - Bulletin du Jardin botanique de J. T o x i c o l . C l i n . T o x i c o l . - Journal o f T o x i c o l o g y . C l i n i c a l T o x i c o l o g y .
Buitenzorg. Jordstjarnan - Jordstjarnan.
B u l l . Jard. bot. Etat - B u l l e t i n du Jardin B o t a n i q u e de l ' E t a t a Kew Bull. - K e w Bulletin.
Bruxelles. K e w B u l l . add. Series - K e w B u l l e t i n A d d i t i o n a l Series.
B u l l . Jard. bot. natn. B e l g . - B u l l e t i n de Jardin botanique national de Lejeunia - Lejeunia.
Belgique. L i b r i bot. - L i b r i botanici.
B u l l . mens. Soc. l i n n . L y o n - B u l l e t i n mensuel de l a Societe Linneenne Lilloa - Lilloa.
de L y o n . Linnaea - Linnaea.
B u l l . N . Y . bot. G d n - B u l l e t i n o f the N e w Y o r k B o t a n i c a l Garden. Lloydia - Lloydia.
B u l l , semest. S o c . m y c o l . N o r d - B u l l e t i n semestriel de l a Societe Madrono - Madrono.
M y c o l o g i q u e du N o r d . Magy tudom. Akad. Ertek. termeszettud. Koreb. - Magyar
B u l l . S o c . bot. Fr. - B u l l e t i n Societe botanique. France. Tudomanyos A k a d e m i a . Ertekezesek a termeszettudomanyok
B u l l . Soc. bot. Geneve - B u l l e t i n de l a Societe botanique de Geneve. korebol.
B u l l . S o c . H i s t . nat. A f r . N . - B u l l e t i n de l a Societe d'histoire naturelle M e d e d . N e d . m y c o l . Vereen. - M e d e d e e l i n g e n van de Nederlandsche
de 1'Afrique d u N o r d . M y c o l o g i s c h e Vereeniging.
B u l l . S o c . m y c o l . F r . - B u l l e t i n de l a Societe m y c o l o g i q u e de France. M e m . N . Y bot. G d n - M e m o i r s o f the N e w Y o r k B o t a n i c a l Garden.
B u l l . S o c . nat. O y o n n a x - B u l l e t i n de l a Societe des naturalistes M e m . Fac. E d u c . S h i g a U n i v . N a t . S c i . - M e m o i r s o f the Faculty o f
d ' O y o n n a x pour F etude et l a diffusion des sciences naturelles dans Education, S h i g a University. N a t u r a l Science.
la region. M e m . Soc. E m u l . M o n t b e l i a r d - M e m o i r e s de l a Societe d'emulation
B u l l . S o c . roy. bot. B e l g . - B u l l e t i n de l a Societe R o y a l e de Botanique de M o n t b e l i a r d .
de B e l g i q u e . M e m o r i e S o c . tosc. S c i . nat. - M e m o r i e d e l l a Societa Toscana d i
B u l l . Torrey bot. C I . - B u l l e t i n o f the Torrey B o t a n i c a l C l u b . Scienze N a t u r a l i .
B u l l , trimest. S o c . m y c o l . Fr. - B u l l e t i n trimestriel de l a Societe M e t r o d i a n a - Metrodiana.
m y c o l o g i q u e de France. M i c o l . ital. - M i c o l o g i a italiana.
C . r. hebd. Seanc. A c a d . S c i . , Paris - C o m p t e rendu hebdomadaire des M i c o l . Veget. medit. - M i c o l o g i a e vegetazione mediterranea.
seances de l ' A c a d e m i e des sciences. Paris. M i c o l . veneta - M i c o l o g i a veneta.
C . r. A s s . franc. A v . S c i . - C o m p t e rendu de l ' A s s o c i a t i o n francaise M i k o l . Kozlem. - Mikologiai Kozlemenyek
pour l'avancement des sciences. M i t t , aargau. naturf. G e s . - M i t t e i l u n g e n der A a r g a u i s c h e n natur-
Carinthia - Carinthia. forschenden Gesellschaft.
Carolinea - Carolinea. Mitt, naturf. G e s . L u z e r n - M i t t e i l u n g e n der naturforschenden
Ceska M y k o l . - Ceska Mykologie. Gesellschaft i n L u z e r n .
Comm. Soc. critt. ital. - Societa crittogamologica italiana. Mittbl. Arbeitsgem. Pilzk. Niederrhein - Mitteilungsblatt der
Commentario. 'Arbeitsgemeinschaft P i l z k u n d e N i e d e r r h e i n ' .
Contr. U n i v . M i c h . Herb. - Contributions f r o m the U n i v e r s i t y o f M o l . P h y l o g e n . E v o l . - M o l e c u l a r Phylogenetics and E v o l u t i o n .
M i c h i g a n Herbarium. M y c o l . Res. - M y c o l o g i c a l Research.
Coolia - Coolia. Mycologia - Mycologia.
Mycologist - The Mycologist; Mycologist. Rev. catal. M i c o l . - R e v i s t a Catalana d i M i c o l o g i a .

M y c o p a t h . M y c o l . appl. - M y c o p a t h o l o g i a et m y c o l o g i a applicata. Revue gen. B o t . - R e v u e generate de Botanique.
Mycotaxon - Mycotaxon. Riv. M i c o l . - Rivista di Micologia.
M y k o l . M i t t b l . - M y k o l o g i s c h e s Mitteilungssblatt. S. A f r . J . B o t . - South A f r i c a n Journal o f Botany.
M y k o l . Sb., Praha - M y k o l o g i c k y sbornik, Praha. S c h w e i z . Z . P i l z k . - Schweizerische Zeitschrift fur P i l z k u n d e .
Mykologia - Mykologia. Science - Science.
N . Amer. F l . - North American Flora. Sc. tot. Env. - Science o f the Total Environment.
N . Z . J . B o t . - N e w Z e a l a n d Journal o f Botany. Seem. J . B o t . - Seemann's Journal o f Botany.
N o r d i c J . B o t . - N o r d i c Journal o f Botany. Sieni Lehti - Sieni Lehti.
Not. bot. H o r t i agrobot. C l u j - N a p o c a - Notulae botanicae H o r t i agro- Svampe - Svampe.
bo tanici C l u j - N a p o c a . Siidwestd. Pilzrundschau - Sudwestdeutsche Pilzrundschau.
Notes R . bot. G d n E d i n b . - N o t e s f r o m the R o y a l B o t a n i c G a r d e n , Sydowia - Sydowia.
Edinburgh. Trans. B r . m y c o l . S o c . - Transactions o f the B r i t i s h M y c o l o g i c a l
Nova Hedwigia - Nova Hedwigia. Society.
N o v . Sist. vyssh. n i z s h . Rast. - N o v o s t i Sistematiki v y s s h i k h i n i z s h i k h U k r . bot. Z h . - U k r a y i n s ' k y i botanichnyi zhurnal.
Rastenii. Ulmer Pilzfl. - U l m e r Pilzflora.
Papers M i c h . A c a d . S c i . , A r t s Letters - Papers from the M i c h i g a n Verh. bot. Ver. Prov. Brandenb. - Verhandlungen des B o t a n i s c h e n
A c a d e m y o f Science, A r t s and Letters. Vereins fur die P r o v i n z Brandenburg.
Persoonia - Persoonia. Verh. z o o l . bot. Ges. W i e n - Verhandlungen der Zoologisch-botanis-
P i l z f l . Nordwestoberfrankens - P i l z f l o r a Nordwestoberfrankens. chen Gesellschaft i n W i e n .
P u b i s D e p . A g r i c . C a n . - Publications. Department o f A g r i c u l t u r e . W i s s . Z . Ernst M o r i t z A r n d t - U n i v . G r e i f s w a l d - Wissenschaftliche
Canada. Zeitschrift der Ernst M o r i t z Arndt-Universitat G r e i f s w a l d .
Rep. N . Y . St. M u s . nat. Hist. - Report on the N e w Y o r k State M u s e u m Z . M y k o l . - Zeitschrift fur M y k o l o g i e .
of natural History. Z . P i l z k . - Zeitschrift fur P i l z k u n d e .
R e p r i u m nov. Spec. R e g n i veg. - R e p e r t o r i u m specierum novarum
regni vegetabilis.
B
Taxonomic part
Agaricaceae (Fr.) Cohn
ELSE C . VELLINGA
Agaricus 'subordo' Agaricini Fr., Syst. Orb. veg.: 65. 1825; Agaricus 'ordo' Agariceae Chev., Fl. generale Env. Paris
1: 121. 1826; Agaricaceae (Fr.) Cohn in Hedwigia 11: 17. 1872. - Lepiotaceae Over, in Bull. Jard. bot. Buitenzorg 9:
19. 1927. - Leucocoprinaceae Sing, in Annls mycol. 34: 323. 1936 (not valid); Leucocoprinaceae Sing, ex Juhch,
Higher Taxa Basidiomyc: 376. 1981.
Basidiocarp pluteoid or, more rarely, collybioid or mycenoid; lamellae free, irregular in secotioid species; universal and
partial veil in most species present; annulus or annular zone often present; spore print white, cream, pale lilac, pale
pink, brown, (pale) grey-brown, dark reddish brown or reddish brown, green, in one species discolouring on drying to
blackish, even blue in one tropical genus.
Spores smooth, rarely ornamented (in most cases small warts, in one tropical genus an intricate reticulate pattern),
uninucleate or binucleate, with or without germ pore, without or with apical endosporal thickening, thick-walled, dex-
trinoid, rarely not, and often metachromatic in Cresyl Blue, but those reactions may be obscured by pigmentation;
cheilocystidia almost always present; pleurocystidia almost always absent; hymenophoral trama regular or trabecular;
clamp-connections absent or present. Development bivelangiocarpic in most species, rarely monovelangiocarpic, and
pileocarpic, pileostipitocarpic, hymenocarpic, stipitocarpic, or isocarpic. - Type genus Agaricus L.: Fr.
HABITAT & DISTRIBUTION. - Solitary to gregarious or in fairy rings, saprotrophic and terrestrial, rarely on decaying wood,
in woods and in grasslands, often on soil relatively rich in nutrients (esp. nitrogen), widespread, and cosmopolitan; with
many diverse representatives in the tropics.
Singer (Agar. mod. Taxon., Ed. 4: 465-467. 1986) had a wider concept of the family than is practiced here; he included
tribus Cystodermateae Sing, and the genus Pseudobaeospora Sing, within the Agaricaceae. As argued by Bas (in Bas et
al., Fl. agar, neerl. 1: 44. 1989) and confirmed by molecular evidence (Johnson & Vilgalys in Mycologia 90: 976. 1998;
Vellinga, unpublished results) neither of these taxa belong to the Agaricaceae.
Singer (Agar. mod. Taxon., Ed. 4: 465-467. 1986) recognized three tribus (excluding the Cystodermateae), viz.
Leucocoprineae, Agariceae, and Lepioteae. This subdivision of the family is not supported by molecular evidence
(Johnson & Vilgalys in Mycologia 90: 971-979. 1998; Johnson in Mycologia 91: 443-458. 1999). Further molecular
research by the present author on this group shows that a division into two groups is more plausible, combining the
former tribus Agariceae and Leucocoprineae, while transferring Melanophyllum Velen. to tribus Lepioteae.
Coprinus comatus (O.F. Mull.: Fr.) Pers. and C. sterquilinus Fr. are treated under Coprinus (Pers.: Fr.) S.F. Gray,
though they are considered more closely related to members of the Agaricaceae than to the rest of the genus Coprinus
(Hopple & Vilgalys in M o l . Phylogen. Evol. 13: 1-19. 1999).
Some secotioid genera are closely related to members of the Agaricaceae. A n example is Endoptychum Czern.,
which is represented with one species in Europe, viz. E. agaricoides Czern. Singer (Agar. mod. Taxon., Ed. 4. 1986)
did not mention this genus in his treatment of the Agaricales.
Several species are considered excellent edibles and are widely cultivated or gathered, but the family also harbours
deadly toxic members.
K E Y TO THE G E N E R A
1. Spore print brown, (dark) reddish brown, or reddish turning blackish on drying; lamellae turning brown with age,
or pinkish red
2. Pileus covering granular, made up of globose elements only 10 (1) Melanophyllum haematospermum
2. Pileus covering smooth to radially fibrillose or squamose, made up of radial hyphae, rarely with clavate terminal
elements
21
3. Basidiocarps discolouring yellow or red when scratched or cut, or not at all; spores without apical endosporal
thickening; cheilocystidia clavate to globose, sometimes utriform or cylindrical, but without apical capitulum,
or cheilocystidia absent 1- Agaricus
3. Basidiocarps when cut or scratched first discolouring yellow, then after c. 15 minutes vinaceous red to red;
spores with apical endosporal thickening; cheilocystidia narrowly lageniform to almost cylindrical with api-
cal capitulum 2. Allopsalliota
1. Spore print white or very pale (cream, light lilacinous, pink etc.), or green
4. Annulus complicated, with a double, fringed edge (see Fig. 44), movable with age
5. Spores and spore print white, cream or pale pink 3. Macrolepiota p.p.
5. Spores and spore print green 4. Cnlorophyllum
4. Annulus simple, without fringes etc., only present as a thickened band on the stipe, or absent
6. Lamellae turning green or red in ammonia vapour 6. Leucoagaricus p.p.
6. Lamellae not turning green or red in ammonia vapour
7. Pileus radially sulcate; context in pileus thin and pliable; basidia surrounded by pseudoparaphyses;
clamp-connections absent 5. Leucocoprinus p.p.
7. Pileus not radially sulcate, though radially fibrillose or squamose, smooth, or floccose; context thin to
thick; pseudoparaphyses absent; clamp-connections present or absent
8. Pileus covering granular, floccose, warty, made up of globose to inflated, often loosely arranged ele-
ments
9. Pileus covering made up of globose elements; clamp-connections absent or present
10. Clamp-connections absent 5. Leucocoprinus p.p.
1
10. Clamp-connections present j
11. Spore print pale green; lamellae pale blue-green 10 (2) Melanophyllum eyrei
11. Spore print white, cream; lamellae white, cream y
12. Pileus covering made up of agglutinated chains of elements; terminal elements ellipsoid
to globose 7. Lepiota p.p.
12. Pileus covering made up loosely arranged globose elements 9. Cystolepiota p.p.
9. Pileus covering made up of inflated elements; clamp-connections absent
13. Spores (3.5-)4.0-5.5(-6.0) 3 (2.0-)2.5-3.5 um, thin-walled, without germ pore; pileus not radi-
ally sulcate 9 (8) Cystolepiota pulverulenta
13. Spores 8.0-12 3 5.5-7.5 um, thick-walled, with germ pore; pileus radially sulcate in marginal
zone 5 (2) Leucocoprinus cretaceus
8. Pileus covering smooth, squamose, with pointed warts or squamules, fibrillose, made up of narrowly
clavate, or elongated, sometimes cystidioid elements, or both rounded elements in agglutinated chains
and elongated connecting elements
14. Spores with germ pore
15. Spores 11.0-18.5(-20) 3 (6.5-)7.5-12 um with hyaline cap over the pore 3. Macrolepiota p.p.
15. Spores 7.0-11.5 3 5.0-8.0 um, without or with hyaline cap over pore
16. Spores 9.5-11.5 3 6.5-8.0 um with truncate apex with germ pore; pileus covering with tight-
ly packed narrowly clavate terminal elements 3. Macrolepiota p.p.
16. Spores 7.0-10.0(-l 1) 3 5.0-7.0 um with rounded apex; pileus covering with erect short to long
cylindrical elements 6. Leucoagaricus p.p.
14. Spores without germ pore
17. Clamp-connections present, at least in pileus covering
18. Pileus covering smooth, but not gelatinized, with squamules made up of erect long ele-
ments, with pointed warts, or radially fibrillose etc.; pleurocystidia absent 7. Lepiota p.p.
18. Pileus covering smooth, a hymeniderm, gelatinized; pleurocystidia present, large, with
refractive contents 8. Chamaemyces
17. Clamp-connections absent
19. Pileus covering innately fibrillose, fibrillose-squamulose, sometimes slightly gelatinized at
centre, not lanate, nor with pointed acute squamules, ranging in colour from white to black,
pink, red, brown, or purplish lilac; basidiocarps pliable with soft tissues, and often fragile
6. Leucoagaricus p.p.
19. Pileus covering either totally fibrillose-lanate, dull purplish, or with pointed acute squa-
mules and brown; basidiocarps not pliable and fragile 7. Lepiota p.p.
AGARICUS (Nauta) 23
1. Agaricus L.
MARIJKE M . NAUTA
Agaricus L., Spec. PI. 2: 1171. 1753; Agaricus L.: Fr., Syst. mycol. 1: lvi, 8. 1821. -Agaricus tribus Psalliota Fr., Syst.
mycol. 1: 280. 1821; Psalliota (Fr.) Kumm., Fiihr. Pilzk.: 23. 1871. - Agaricus # Pratella Pers., Syn. meth. Fung.:
X V I . 1801; Pratella (Pers.) S.F. Gray, Nat. Arr. Br. PI. 1: 626. 1821.
SELECTED LITERATURE - Bohus in Mikol. Kdzlem. 34:5-36.1995; M . Bon in Doc. mycol. 15(60): 1-37.1985; Cappelli,
Agaricus. 1984; Heinem. in Sydowia 30: 6-37. ('1977') 1978; M . Meusers in Beitr. Kenntn. Pilze Mitteleur. 2: 27-56.
1985; F. M0ller in Friesia 4: 1-60. 1950; 135-220. 1952; Nauta in Persoonia 17: 221-233. 1999; Pilat in Acta Mus. nat.
Prag. 7B (1): 1-142. 1951; Wasser, Tr. Agariceae Soviet Union: 15-120. 1989.
Basidiocarps pluteoid or collybioid, sometimes mycenoid; pileus smooth or fibrillose, or squam(ul)ose, dry, white to
yellow or reddish to purplish brown, not discolouring to yellowing or reddening when touched or damaged; lamellae
free, at first (pale) pinkish grey to (pale) pinkish brown, finally dark brown, sometimes brown; stipe central, often annu-
late; veil usually well-developed, white, present in form of usually appendiculate flocks at margin, sometimes on sur-
face of pileus near margin, and as annulus attached at upper half of stipe; context not discolouring or (sometimes
faintly) yellowish or reddish on exposure or when damaged; spore print dark reddish brown (Mu. 5 Y R 3/2-3).
Spores smooth, also when observed with Scanning Electron Microscope, often relatively thick-walled, greyish brown
or brown in water, K O H or ammonia when observed with light microscope, occasionally with germ pore or apically with
thin spot; lamella edge usually with a sterile layer consisting of often catenate cheilocystidia; cheilocystidia cylindrical,
clavate to subglobose; pleurocystidia absent; hymenophoral trama regular to irregular; pileipellis a cutis; pigment usual-
ly pale, parietal or intracellular; veil usually consisting of narrow hyphae; clamp-connections lacking in all tissues.
Development bivelangiocarpic, isocarpic or hymenocarpic. - Typus conservandus: Agaricus campestris L.: Fr.
HABITAT & DISTRIBUTION - Saprotrophic, terrestrial on roadsides, in woods, city parks etc., also in grasslands, lawns,
open dunes, often on nutrient-rich places. Widespread, often cosmopolitan, relatively more common in temperate zones.
The taxonomy of Agaricus remains far from clear, despite the contributions of many authors in recent times. It has not
been possible in this study to unravel all the species-complexes to full satisfaction. A practical approach had to be cho-
sen, but more research is needed, preferably of morphological as well as molecular characters. It is to be hoped that molec-
ular studies will give a better insight in the infrageneric taxonomy of this genus.
In general the morphological features of the basidiocarps in this genus are rather variable, and the microscopical
characters do not show much differentiation. In the work of M0ller (in Friesia 4: 1-60. 1950; 135-220. 1952) the given
spore measures are usually too small, in the work of Pilat (in Acta Mus. nat. Prag. 7B (1): 1-142. 1951) the given spore
sizes are mostly larger than in reality. This combined with the scarcely preserved type material of M0ller makes it
sometimes difficult to ascertain the identity of species described by M0ller or Pilat.
Average spore size and spore form (Qav) proved to be a useful character in Agaricus, but accuracy and the use of
an oil-immersion lens is required.
The position of the annulus is given with regard to the base of the stipe.
K E Y S TO T H E SPECIES
1. Surface of basidiocarps and/or context yellow or yellowing (sometimes pale, but without brownish tinges) on expo-
sure or when handled, damaged or cut, sometimes only in base of stipe or only rhizomorphs yellowing
2. Context at first entirely and quickly bright yellow, then slowly vinaceous red, in the end surface of pileus and con-
text entirely red; lamellae at first yellowish brown; spores with apically thickened endospore
Allopsalliota geesterani
2. Context not discolouring as described above; lamellae at first greyish, greyish brown or pinkish brown; spores
not with apically thickened endospore
3. Spores with germ pore or with thin-walled spot at apex; lamella edge fertile, cheilocystidia absent; annulus
fragile, evanescent (species of sect. Agaricus with yellowing pileus) K E Y ONE
3. Spores without germ pore or thin-walled spot at apex; lamella edge sterile or partly fertile, usually with con-
spicuous cheilocystidia; annulus fragile to relatively persistent, fibrillose or squamose at underside
4. Annulus ascending or velar sock present (species of sect. Bitorques with sometimes yellowing pileus)
K E Y TWO
4. Annulus descending; no velar sock present
5. Rhizomorphs or pseudorhiza conspicuous and yellowing; Schaeffer-reaction orange on base of stipe, neg-
ative on surface of pileus
6. Pileus entirely reddish brown squamulose; stipe below annulus pruinose; veil composed of hyphae
with inflated elements, intermixed with subglobose elements 41. A . rufotegulis
6. Pileus whitish to dark brown, stipe below annulus fibrillose or with brown girdles, not pruinose; veil
composed of hyphae (sect. Spissicaules) K E Y THREE
5. Rhizomorphs or pseudorhiza if present white to brownish and not discolouring; Schaeffer-reaction either
negative on base of stipe, or if positive (orange) also orange on pileus surface
7. Pileus not discolouring or sometimes yellow when damaged, yellow discoloration usually quickly dis-
appearing, not visible on drying; stipe usually strongly yellow at base (surface and sometimes also con-
text) when damaged or on oxidation, elsewhere sometimes faintly yellowing; smell unpleasant;
Schaeffer-reaction on pileus negative (sect. Xanthodermi) K E Y FOUR
7. Pileus usually yellowing on handling, yellow discoloration not quickly disappearing, also visible or
intensifying on drying; surface of stipe not discolouring to entirely yellowing on handling, not only at
base; smell fresh pleasant, sometimes later unpleasant; Schaeffer-reaction on pileus positive (orange),
rarely negative
8. Annulus broad, 9-35 mm wide, thick, underside floccose to squamose; stipe yellowish or not on
handling; average spore length . 6.0 |xm, often > 6.5 u.m; lamella edge usually with broad layer of
catenate sterile elements (sect. Arvenses) K E Y FIVE
8. Annulus narrow, up to 1 0 mm wide, thin, underside fibrillose; stipe usually strongly yellowing on
handling; average spore length < 6.5 (xm; sterile elements on lamella edge catenate or not, not form-
ing broad sterile layer (sect. Minores) ' K E Y SIX
1. Neither surface nor context yellowing, sometimes yellowish brown tinges occur; rhizomorphs, if present, not
yellowing
9. Spores with germ pore or with thin-walled spot at apex; lamella edge fertile: cheilocystidia absent; annulus thin,
usually soon disappearing (sect. Agaricus) K E Y ONE
9. Spores without germ pore or thin-walled spot at apex; lamella edge sterile, with cheilocystidia; annulus thin or
thick or with thickened margin, rarely only velar sock present
10. Context not discolouring on oxidation
11. Spores broad, average spore width > 4.6 jxm; Qav 1.10-1.35(-1.45); terminal elements of cheilocystidia
up to 1 3 ( - 1 5 ) mm wide (species of sect. Bitorques with not discolouring context) K E Y TWO
11. Spores narrow, average spore width < 4 . 0 u.m, Qav 1.40-1.70; terminal elements of cheilocystidia up to
2 0 ( - 2 4 ) mm wide
12. Surface of stipe below annulus fibrillose-squamulose to flocculose; K O H on context or stipe yel-
lowish brown; average spore size ( 5 . 1 - ) 5 . 7 - 6 . 2 3 3.5-3.9 mm 16. A . impudicus
12. Surface of stipe below annulus fibrillose to satiny striate; K O H on context or stipe yellow to orange;
average spore size 5.3-5.7 X 3.7-4.0 u,m 37. A . phaeolepidotus
10. Context reddening or brownish on exposure or when damaged
13. Spores narrow, Qav > 1.40; cheilocystidia clavate to globose, up to 2 0 ( - 2 5 ) |Jim wide
14. Surface of stipe below annulus fibrillose to satiny striate; K O H on context or stipe yellow to orange
37. A . phaeolepidotus
14. Surface of stipe below annulus fibrillose-squamulose to lanate; K O H on context or stipe negative or
1
' yellowish brown (sect. Sanguinolenti) K E Y SEVEN
13. Spores relatively broad, Qav < 1.40; cheilocystidia clavate to cylindrical, up to 1 5 ( - 1 7 ) p,m wide
15. Annulus at underside at margin with radially arranged squames; average spore length 6.9-8.4 u,m
20. A . depauperatus
15. Annulus not with radially arranged squames at underside; average spore length 5.8-7.3 u.m, if up to
7.7 u,m only velar sock present (sect. Bitorques) K E Y TWO
K E Y ONE
Pileus not discolouring, sometimes yellowing; veil usually present as thin, evanescent annulus; context not discolouring
or discolouring faintly reddish; spores with germ pore or with thin-walled spot at apex; lamella edge fertile, cheilo-
cystidia absent (sect. Agaricus)
AGARICUS (Nauta) 25
1. Pileus covered with dark (grey-)brown to purple-brown fibrils or squamules

2. Pileus with appressed greyish brown to greyish squamules, average spore size 7.5-7.9(-8.1) X 5.0-5.6 |j.m
lc. A. campestris var. squamulosus
2. Pileus entirely fibrillose, sometimes fibrillose-squamulose, usually dark purplish brown, average spore size vary-
ing from 6.4-8.3 |jim
3. Average spore length 7.6-8.3 pm; stipe typically attenuated towards base 5. A. cupreobrunneus
3. Average spore length 6.4-6.9 mm; stipe typically ventricose 6. A. porphyrocephalus
1. Pileus white to sometimes yellowish, smooth, fibrillose or fibrillose-squamulose, fibrils only in the end brownish or
greyish
4. Stipe up to 100 mm long, longer than diameter of pileus; context in stipe usually reddish in upper part, yellow-
ish in base 4. A. altipes
4. Stipe usually up to 65 mm long, shorter than diameter of pileus; context not discolouring except for yellow-brown
discoloration in base of stipe
5. Pileus yellowing, surface nearly smooth
6. Pileus white to yellowish, discolouring yellowish on handling or damaging, finally pale yellow; surface of
stipe above annulus flocculose; average spore length 6.2-7.1 pm 2. A. moellerianus
6. Pileus white, strongly yellowing; surface of stipe above annulus smooth; average length of spores 7.9-8.2
ixm lb. A. campestris var. equestris
5. Pileus not to only very slightly yellowing; surface usually fibrillose or fibrillose-squamulose, rarely smooth
7. Average length of spores 8.7-9.4 pm 3. A. pampeanus
7. Average length of spores 7.5-8.1 pm la. A. campestris var. campestris
K E Y TWO
Pileus not discolouring or slightly yellowing, rarely slightly reddening; annulus ascending and thin to thick or descend-
ing and thick, sometimes only velar sock present; context usually discolouring reddish or brownish on oxidation, some-
times not discolouring; spores without germ pore, relatively broad, Qav < 1.35(-1.45); lamella edge sterile,
cheilocystidia conspicuous, clavate to cylindrical, up to 15 |Jim wide (sect. Bitorques)
1. Pileus whitish from the beginning, sometimes later with brownish tinges
2. Basidiocarps small, fragile; diameter pileus up to 50(-55) mm; annulus relatively thin, narrow, velar sock absent
11. A. devoniensis
2. Basidiocarps larger, robust; diameter of pileus 3= 50 mm; annulus thick and triangular or double, if narrow or
absent velar sock present
3. Annulus thick and triangular in transverse section; basidia 2-spored . . . . 10. A. bisporus (cultivated variant)
3. Annulus double, or narrow and ascending, or velar sock present; basidia 4-spored
4. Annulus double, sometimes a high and low annulus present; velar sock absent; average spore size 5.9-6.5
X 4.6-4.9 pm 7. A. bitorquis
4. Annulus, if present, narrow and ascending; velar sock present; average spore size (6.0-)6.6-7.7 X 4.9-6.2 pm
5. Annulus usually present; average spore size (6.0-)6.6-7.3 X 4.9-6.1 pm; cheilocystidia long, cylindrical
to clavate, 35-58 X 6.5-10(-12.5) pm 8. A. bernardii
5. Annulus absent; average spore size 7.7 X 5.7-6.2 pm; cheilocystidia clavate, 20-36 X 8.5-10 pm
9. A. gennadii
1. Pileus brown(ish) from the beginning
6. Pileus fibrillose, usually with whitish velar flocks near margin 12. A. subfloccosus
6. Pileus usually squamose, if only fibrillose then with broad white rim
7. Pileus brown fibrillose to squamose, with broad white rim; smell fungoid, pleasant; basidia 2-spored; average
spore size 7.1-7.3 X 5.4-5.9 pm . 10. A. bisporus
7. Pileus squamose; smell unpleasant; basidia 4-spored; average spore size 5.8-7.0 X 5.2-5.6 pm
8. Basidiocarps solitary or in small groups, stipe clavate; cheilocystidia in clusters, width (5.5-)7.5-13 mm;
average spore size 6.6-7.0 X 5.2-5.6 pm 13. A. subperonatus
8. Basidiocarps fasciculate, with fusiform stipe; cheilocystidia not in clusters, width 6.0-9.0 mm; average spore
size 5.8-6.5 X (4.8)5.3-5.5 pm 14. A. bohusii
K E Y THREE
Pileus often slightly yellowing; base of stipe with yellowing rhizomorphs; context in base of stipe usually yellow on
oxidation, usually faintly reddish elsewhere; Schaeffer-reaction positive (orange) on base of stipe, negative on surface
of pileus; spores without germ pore; cheilocystidia either globose to clavate or inconspicuous (basidioliform), some-
times scarce (sect. Spissicaules)
1 Pileus mainly brown squamose; stipe below annulus with brown girdles 39. A. lanipes
1. Pileus mainly whitish to pale brownish, usually with appressed brownish squames at centre only; stipe below annu-
lus smooth to fibrillose
2. Pileus with appendiculate veil; average spore size (7.1-)7.4-8.0 X 5.2-5.8 p,m; lamella edge usually partly fertile
38. A. litoralis
2. Pileus without appendiculate veil; average spore size 6.0-6.7 X 4.2-4.7 (jum; lamella edge fertile to almost sterile
40. A. bresadolanus
K E Y FOUR
Pileus not discolouring or sometimes yellow on scratching, disappearing on drying, rarely slightly pink; context and/or
surface in base of stipe discolouring yellow, usually brightly; KOH-reaction yellow to orange at base of stipe; Schaeffer-
reaction on surface of pileus and base of stipe negative; smell unpleasant; spores without germ pore, length < 6.5 mm
on average (sect. Xanthodermi)
1. Pileus grey or white, or greyish brown, starting smooth, sometimes later at centre breaking up into large indistinct
squames and then brownish or reddish brown at centre
2. Lamella edge sterile; cheilocystidia 10-20 mm wide, not in chains; pileus usually white, sometimes greyish, not
discolouring pinkish on handling; annulus thickened at margin 33. A. xanthodermus
2. Lamella edge in part fertile; cheilocystidia 6.5-12 mm wide, in short chains; pileus whitish or greyish brown,
sometimes on handling becoming slightly pinkish brown
3. Context strongly yellowing at base of stipe; pileus smooth to fibrillose, not breaking up into squames; annu-
lus thickened at margin and split, forming triangle in transverse section 34. A. pilatianus
3. Context slightly yellowing at base of stipe, soon turning reddish; pileus soon breaking up into large indistinct
squames at centre, sometimes remaining smooth; annulus relatively thin but persistent, not llriangular
35. A. pseudopratensis
1. Pileus entirely covered with brown or dark greyish brown fibrillose squamules even in young specimens 4
4. Pileus with brownish to dark greyish brown small fibrillose squamules; context in base of stipe usually strongly
yellowing on exposure or when damaged; cheilocystidia (terminal elements) globose to clavate, up to 15 mm
wide 36. A. moelleri
4. Pileus with light brown, rather large fibrillose squames; context in base of stipe first pale yellow, later faintly
brownish; cheilocystidia clavate, obovoid to globose, up to 23 mm wide 37. A. phaeolepidotus
K E Y FIVE
Pileus yellowish on handling, especially on drying; annulus broad, up to 35 mm wide, with remnants of universal veil
as thick flocks or warts at underside; context not discolouring or discolouring slightly yellow or orange to pinkish;
KOH-reaction on context usually positive and yellow; Schaeffer-reaction on pileus orange, rarely negative; average
spore length > 6.0 mm; without germ pore; cheilocystidia catenate (sect. Arvenses)
1. Pileus covered with yellow to brown squames on paler background; stipe often rooting; cheilocystidia in protuber-
ant clusters and frequently of irregular form 22. A. augustus
1. Pileus white to yellowish or yellowish brown, if squames present these are mostly concolorous; stipe not rooting;
cheilocystidia not in protuberant clusters
2. Average spore size 10.1-10.7 3 6.5-7.2 mm; basidiocarps robust; smell soon unpleasant . . . . 27. A. urinascens
2. Average spore size , 9.5 3 6.0 mm; basidiocarps gracile to robust; smell usually pleasant, like almonds or anise
3. Spores 9.0-9.5 X 5.0-5.5 mm; pileus with appressed coarse squames A. arvensis var. macrolepis
Pilat & Pouz. in Acta Mus. nat. Prag. 7B (1): 134. 1951 from eastern Europe, not yet found in the Netherlands
and adjacent countries
3. Spores < 9.0 u,m; pileus without coarse squames, at most squamulose
4. Average spore size 7.7-8.6 X 5.3-5.9 mm
AGARICUS (Nauta) 27
5. Basidiocarps relatively gracile; pileus usually < 105 mm in diameter; stipe smooth to fibrillose below
annulus, cylindrical to subbulbous, 55-120 X 10-20 mm-(occasionally up to 34 mm at base); mostly in
grassland 21. A. arvensis
5. Robust basidiocarps; pileus > 120 mm in diameter; stipe usually floccose-squamulose below annulus,
bulbous, 100-200 X 20-30 mm (- 45 mm at base); mostly in woods 26. A. macrocarpus
4. Average spore size 6.0-7.5 X 4.1-5.3 |xm
6. Basidiocarps gracile, quickly yellowing; annulus usually fragile; average spore size 6.0-6.7 X 4.1-4.7
(xm; cheilocystidia globose to clavate 25. A. sylvicola
6. Basidiocarps gracile or robust, slowly and often faintly yellowing; annulus thick, persistent; average spore
size 6.5-7.5 X 4.5-5.3 mm; cheilocystidia globose to clavate, in part utriform
7. Stipe cylindrical to rarely subbulbous, (65-)80-120 X 15-28 mm; pileus usually plano-convex,
sometimes irregular, 80-175 mm in diameter; average spore size 6.8-7.5 3 4.8-5.3 |xm; in grass-
lands, rarely in woods 23. A. osecanus
7. Stipe clavate to bulbous, 50-80 X 8-18 mm, at base 20-26 mm wide; pileus regularly convex, 65-95
mm in diameter; average spore size 6.5-7.2 X 4.5-5.0 pm; in woods 24. A. pseudoumbrella
K E Y SIX
Surface of pileus and stipe strongly yellowing when touched or on drying; annulus narrow, thin, up to 10 mm wide,
sometimes absent, fibrillose, without flocks or warts at underside; context discolouring slightly yellow, orange-yellow
or yellowish brown on exposure or when damaged; Schaeffer-reaction on surface of pileus strongly orange; average
spore length < 6.5 mm; cheilocystidia shortly catenate or not (sect. Minores)
1. Pileus at first white, quickly yellowing, rarely young with some purplish fibrils
2. Pileus 17-35(-40) mm in diameter, white, soon discolouring yellow, appressed fibrillose; lamella edge with minor-
ity of cystidia, sometimes cheilocystidia hardly present 30. A. comtulus
2. Pileus 40-70(-85) mm in diameter, white at first, soon yellowing, with yellowish brown fibrillose squames, later
entirely yellow-brown; lamella edge almost sterile 31. A. xantholepis
1. Pileus vinaceous brown, or greyish brown to yellowish brown with purple brown fibrillose centre
3. Pileus reddish to purplish brown with numerous purplish brown fibrils and squames, later (reddish) brown, up to
90 mm in diameter 32. A. porphyrizon
3. Pileus at centre with vinaceous fibrils, sometimes with some squames, later yellowish brown, up to 65(-80) mm
in diam.
4. Average spore length < 5.7 mm 28. A. dulcidulus
4. Average spore length > 6.0 mm 29. A. luteomaculatus
4 % . :-v< K E Y SEVEN
Pileus not discolouring or reddening or browning; annulus relatively persistent, with thickened margin; context not dis-
colouring-or reddening or browning on exposure or when damaged; K O H and Schaeffer-reaction negative; spores nar-
row, Qav*v> (1.30-) 1.40, without germ pore; lamella edge sterile; cheilocystidia clavate to (sub)globose, up to 20 pm
wide (sect. Sanguinolenti)
1. Pileus whitish to pale (greyish) brown, if squamulose squamules concolorous with pileus
2. Basidiocarps (and context) intensely reddening when handled or damaged; spores 5.0-7.0 X 3.0-5.0 |xm; on aver-
age 5.5-6.5 X 3.5-4.5 pm 19. A. benesii
2. Pileus not discolouring on handling; context slowly but not intensely bright red on exposure or when damaged;
spores 6.0-9.5 X 4.5-6.0 |Jim; on average 6.9-8.4 X 4.8-5.6 mm 20. A. depauperatus
1. Pileus with brown to reddish brown squam(ul)es, or dark red-brown fibrillose
3. Spores < 7.0 mm long; average spore length 5.1-6.5 pm
4. Context not or hardly discolouring when cut; cystidia width up to 20(-24) pm 16. A. impudicus
4. Context clearly reddening when cut; cystidia up to 12 |xm wide
5. Pileus thin-fleshed; context up to 7 mm thick in pileus; squames dark brown or reddish brown
15. A. sylvaticus
5. Pileus thick-fleshed; context up to 17 mm thick in pileus; squames pale brown, concolorous to pileus
19. A. benesii
3. Spores > 6.5 mm long; average spore length 6.9-8.3 u,m

6. Pileus densely covered with brown to reddish brown relatively broad squames; average spore size (7.6)7.8-8.3
X 4.4-5.4 u,m 17. A . langei
6. Pileus dark red-brown fibrillose; average spore size 6.9-7.7 X 4.3-4.5 u,m 18. A . fuscofibrillosus
Subgen. Agaricus
V e i l c o m p o s e d o f hyphae w i t h usually c y l i n d r i c a l , 3.0-8.0 |i.m w i d e

elements, w i t h some inflated elements up to 12(-17) |xm.
Sect. Agaricus
Pileus not discolouring, sometimes y e l l o w i n g ; v e i l usually present as

thin, evanescent annulus; context not d i s c o l o u r i n g o r d i s c o l o u r i n g
faintly reddish, sometimes also y e l l o w i s h b r o w n i n base o f stipe. F i g . 2. Agaricus campestris. (habit X 'A).
K O H - r e a c t i o n negative; Schaeffer-reaction negative o n a l l parts o f
basidiocarps. not to very slightly y e l l o w i n g o n handling, finally sometimes pale y e l -
Spores e l l i p s o i d , Q a v > 1.40, w i t h germ pore or w i t h t h i n - w a l l e d l o w ( M u . 5 Y 8/3), often w i t h some finally greyish or pale greyish
spot at apex; l a m e l l a edge fertile; cheilocystidia absent. b r o w n small fibrils, usually fibrillose-squamulose, sometimes only fib-
\ rillose. L a m e l l a e , L = 95-130, 1 = l - 3 ( - 5 ) , crowded, free, undate to
1. Agaricus campestris L . , Spec. PI. 2: 1173. 1753:Fr. - F i g . 2. ventricose, up to 8 m m broad, at first p i n k i s h grey to p i n k i s h b r o w n
Agaricus campestris L . : Fr., Syst. m y c o l . 1: 2 8 1 . 1821; Psalliota (7.5 Y R 7/2; 5 Y R 5/3), later b r o w n to reddish b r o w n (5 Y R 6/2, 7.5
campestris ( L . ) Q u e l , i n M e m . Soc. E m u l . M o n t b e l i a r d ser. II, 5: 140. Y R 3/2), finally dark b r o w n , w i t h concolorous to slightly paler, entire
1872 ( C h a m p . Jura Vosges 1). edge. Stipe (40-)45-60(-100) X 9-15 m m , usually annulate, c y l i n d r i -
E X C L . - Agaricus campestris sensu C o o k e , 111. B r i t . F u n g i 4: p i . cal, often tapering towards base, sometimes slightly enlarged up to 22
527. 1885; Psalliota campestris sensu Bres., Iconogr. m y c o l . 17: p i . m m at base, straight, stuffed to narrowly fistulose, white, later greyish
823. 1931. ( = A. bisporus). i n upper half or w i t h purple tinge b e l o w lamellae, y e l l o w i s h or b r o w n -
V E R N . N A M E - Weidechampignon. ish to faintly orange-yellow at base o f stipe o n handling, above annu-
lus smooth, b e l o w annulus smooth to fibrillose, sometimes w i t h
The t a x o n o m y o f the Agaricus campestris-group is rather confusing. fibrillose girdles, or flocculose. A n n u l u s at 0.50-0.70 o f height o f stipe,
Very few characters are available, w h i c h i m p l i e s a risk o f overestimat- 1-5 m m wide, descending, pending, thin, fibrillose, later disappearing,
ing the importance o f variable characters l i k e the colour and structure white, not discolouring, w i t h smooth upperside; underside fibrillose.
o f the pileus. In this study a practical approach is used. A m o n g the Context u p to 12(-18) m m thick i n pileus, white, firm, not discolour-
m a n y infraspecific taxa and related species o f Agaricus campestris i n g w h e n cut, except for a sometimes orange y e l l o w i s h b r o w n d i s c o l -
w h i c h have been described, only a few c o u l d be recognised i n the c o l - oration at the base o f the stipe. S m e l l indistinct, or pleasant, fungoid
lections from the Netherlands and adjacent regions. F o r an extensive or nut-like. Taste fungoid.
discussion see N a u t a (in Persoonia 17. 2 0 0 1 ; i n prep.). M a c r o c h e m i c a l reactions: K O H negative (surface o f pileus), orange
(base o f stipe); Schaeffer-reaction negative to faintly orange (surface o f
K E Y TO T H E VARIETIES pileus), negative (base o f stipe).
1. P i l e u s nearly smooth, o n l y at margin slightly squamulose, strongly Spores 6.5-8.5 X 4.5-6.0 m m , o n average 7.5-7.9(-8.1) X 5.0-5.6
discolouring y e l l o w m m , Q = 1.30-1.65, Qav = 1.40-1.45(-1.50), ellipsoid, w i t h thin-walled
l b . var. equestris spot at apex. B a s i d i a 21-24 X 7.0-8.5 m m , 4-spored. L a m e l l a edge fer-
1. Pileus fibrillose o r fibrillose-squamulose, not to only very slightly tile, cheilocystidia l a c k i n g , sometimes some sterile basidia present.
yellowing Pileipellis a slightly gelatinized cutis o f 4.5-7.0 m m w i d e hyphae with
2. Pileus white, i n the end w i t h some pale greyish b r o w n fibrils c y l i n d r i c a l elements, gradually passing into pileitrama, w i t h pale y e l -
l a . var. campestris l o w i s h intracellular pigment. Stipitipellis a cutis o f unbranched, 6.5-7.0(-
2. Pileus w i t h appressed greyish to greyish b r o w n squamules 8.5) m m wide hyphae, sometimes constricted at septa, with cylindrical
l c . var. squamulosus elements, with pale yellowish, intracellular pigment.
H A B I T A T & D I S T R . - S o l i t a r y o r i n s m a l l groups, saprotrophic,
l a . var. campestris terrestrial in grasslands, meadows, grassy roadside verges,
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 9. 1984; D a h n c k e , 1200 P i l z e : sometimes i n w o o d s o n various s o i l types. C o m m o n and w i d e s p r e a d
493. 1993; J . L a n g e , F l . agar. dan. 4: p i . 1 3 8 C . 1939. i n the Netherlands, A u g . - N o v . C o m m o n a n d widespread i n E u r o p e ,
S E L . D E S C R . & F I G S . - J . L a n g e , F l . agar. dan. 4 : 59-60. 1939; F . cosmopolitan.
M 0 l l e r i n F r i e s i a 4: 56, p i . 17. 1950.
Agaricus campestris var. campestris i s here considered i n the strict
Pileus 45-90(-110) m m , at first hemispherical, later convex w i t h flat- sense o f L a n g e (in D a n s k bot. A r k . 4(12): 9. 1 9 2 6 % S i n c e the o r i g i -
tened centre to plano-convex, at first w i t h inflexed margin, later w i t h nal descriptions o f L i n n a e u s a n d F r i e s are not u n a m b i g u o u s , this
margin largely exceeding lamellae, white to whitish or greyish white, species has sometimes been understood as what m o d e r n authors
AGARICUS (Nauta) 29
w o u l d c a l l A. bisporus. T h e interpretation o f L a n g e is generally f o l - Pileus 45-85(-95) m m , at first convex, expanding to plano-convex to

l o w e d i n m o d e r n literature. (irregularly) applanate, w i t h i n f l e x e d m a r g i n , exceeding lamellae;
white to y e l l o w i s h , discolouring y e l l o w i s h o n h a n d l i n g o r damaging,
l b . var. equestris (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B (1): 14. finally pale y e l l o w ( M u . 5 Y 8/5, 9/6), at centre smooth, often near
1951. margin fibrillose-squamulose, sometimes w i t h pale b r o w n i s h fibrillose
Psalliota campestris var. equestris F. M 0 l l e r i n F r i e s i a 4: 57. 1950. squamules. L a m e l l a e , L = 105-115,1 = l-3(-7), crowded, free, sub-
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 57. 1950. ventricose, up to 9 m m broad, at first bright p i n k to p i n k i s h , remaining
p i n k i s h for a long time, later p i n k i s h b r o w n , w i t h concolorous, entire
C H A R A C T E R I S T I C S - Differing f r o m var. campestris i n the nearly edge. Stipe 40-80 X 8-15.5(-22) m m , usually w i t h annular zone, some-
smooth, strongly y e l l o w i n g pileus. times w i t h well-developed annulus, c y l i n d r i c a l to tapering at base,
Spores 7.0-9.0(-10.0) X 5.0-6.0(-6.5) |xm, o n average 7.9-8.2 X sometimes w i t h slightly enlarged base, straight to s l i g h t l y curved,
5.4-5.5(-5.9) |xm, Q = 1.35-1.60, Q a v = (1.40-)1.45-1.50, germ pore stuffed to narrowly fistulose, white, not d i s c o l o u r i n g o n handling,
conspicuous, up to 1 (xm w i d e . Other m i c r o s c o p i c a l characters as i n above annulus smooth to fibrillose o r floccose, b e l o w annulus f i b r i l -
the t y p i c a l variety. lose-pruinose to fibrillose. A n n u l u s at 0.50 o f height o f stipe, narrow,
H A B I T A T & DISTR. - Occurs i n the same habitats as the type variety. up to 2 m m wide, usually present as annular zone, descending, pend-
L e s s c o m m o n than, but as widespread as the t y p i c a l variety. ing to slightly spreading, thin, soon disappearing, white, fibrillose,
with striate upperside. Context 7-11.5 m m thick i n pileus, firm, white,
T h e t a x o n o m i c a l rank o f this taxon needs further study. T h e intensity not discolouring when cut, except i n base o f stipe where it sometimes
of y e l l o w i n g o f the pileus seems to be variable. M 0 l l e r ' s statement that turns brownish yellow. S m e l l indistinct. Taste l i k e nuts.
the spores are smaller than i n the t y p i c a l variety c o u l d not be verified. M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus);
T h e germ pore, however, o f this variety seems to be more conspicuous Schaeffer-reaction negative, sometimes w e a k l y orange (surface o f
than i n the t y p i c a l variety. A c c o r d i n g to other authors ( B o n i n D o c . pileus).
m y c o l . 15(60): 6. 1985) this variety also differs i n a stronger red dis- Spores 5.5-7.5 X 4.0-5.5 m m , o n average 6.2-7.1 X 4.1-5.1 m m , Q
coloration o f the context. T h i s c o u l d not be found i n the examined = (1.15-)1.30-1.65(-1.75), Q a v = (1.30-)1.40-1.55, ellipsoid, some-
material, nor i s i t clear f r o m M 0 l l e r ' s original description. times broadly ellipsoid or oblong, without germ pore, w i t h thin-walled
spot at apex; often aberrant spores present. B a s i d i a 22-27 X 7.0-8.0 m m ,
4-spored. L a m e l l a edge fertile, cheilocystidia lacking, sometimes some
Ic. var. squamulosus (Rea) Pilat i n A c t a M u s . nat. P r a g . 7 B (1): 14.
sterile basidia present. Pileipellis a slightly irregular cutis o f 3.5-7.0 m m
1951.
wide hyphae with cylindrical, to sometimes inflated up to 11 |xm ele-
Psalliota campestris var. squamulosa R e a i n Trans. B r . m y c o l . Soc.
ments, gradually passing into pileitrama, w i t h yellowish intracellular and
17: 50. 1932.
parietal pigment. Stipitipellis a cutis o f sometimes branched, 4.5-8.5 u-m
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 10. 1984.
wide hyphae with cylindrical elements, w i t h pale y e l l o w i s h intracellular
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 5 9 , fig. 14. 1950.
pigment and cylindrical, up to 8.5 wide terminal elements.
H A B I T A T & DISTR. - In small groups or solitary, saprotrophic, ter-
C H A R A C T E R I S T I C S - D i f f e r i n g f r o m var. campestris i n the appressed
restrial i n grasslands o n dunes o r o n slopes o f dikes, along roads, or on
greyish b r o w n to greyish squamules o n the pileus.
nutrient-poor meadows o n various soils. R a r e but widespread i n the
M i c r o s c o p i c a l characters as i n the t y p i c a l variety.
Netherlands, J u l y - N o v . Rare but widespread i n Europe.
H A B I T A T & DISTR. - Same as t y p i c a l variety; this variety seems to
be less c o m m o n than the typical one.
M a c r o s c o p i c a l l y this taxon differs i n a more smooth pileus surface and
the flocculose surface o f the stipe above the annulus. T h e s m e l l o f
2. Agaricus moellerianus M . B o n i n D o c . m y c o l . 15(60): 6. 1985. -
almonds or anise w h i c h is mentioned b y various authors ( M 0 l l e r i n
F i g . 3.
F r i e s i a 4 : 57. 1950; B o h u s i n A n n l s hist.-nat. M u s . natn. hung. 70: 107.
Psalliota campestris \w. floccipes F. M 0 l l e r i n F r i e s i a 4: 57. 1950;
1978) c o u l d not be observed.
Agaricus campestris var. floccipes (F. M 0 l l e r ) Pilat i n A c t a M u s . nat.
Prag. 7 ( B ) 1 : 14. 1951; Agaricus floccipes (F. M 0 l l e r ) B o h u s i n A n n l s
3. Agaricus pampeanus Speg. i n A n . S o c . cient. arg. 9: 280. 1880. -
hist.-nat. M u s . natn. hung. 70: 107. 1978, non A. floccipes F r . 1838.
F i g . 4.
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 11. 1984 (as A. floccipes).
S E L . I C O N . - A l b e r t i n M i k o l . K o z l e m . 36(2-3): 62. 1997; Breitenb.
S E L . D E S C R . & F I G S . - B o h u s i n A n n l s hist.-nat. M u s . natn. hung.
& K r a n z l . , P i l z e S c h w e i z 4: p i . 186. 1995; Lavorato i n R i v . M i c o l . 33:
70: 107-109, fig. 3. 1978; F. M 0 l l e r i n F r i e s i a 4: 57-58, fig. 12. 1950.
252. 1991.
V E R N . N A M E - Kleinsporige weidechampignon.
S E L . D E S C R . & FIGS. - H e i n e m . i n B u l l , trimest. Soc. m y c o l . Fr. 81:
374-375. 1965.
VERN. NAME - Steppechampignon.
Pileus 4 5 - 8 5 ( - l 15) m m , at first convex to hemispherical, expanding to

convex with applanate centre o r plano-convex, w i t h slightly inflexed
margin, w h i t i s h to cream at first, faintly y e l l o w i n g sometimes, smooth
to appressed fibrillose or fibrillose-squamulose, later sometimes fis-
sured; fibrils later usually b r o w n i s h ( M u . 5 Y R 4/3, 7.5 Y R 5/4); v e i l
present w h e n y o u n g as broad appendiculate w h i t i s h cottony zone.
L a m e l l a e , L = c. 100,1 = 3-7, crowded, free, ventricose, u p to 10 m m
broad, at first p i n k i s h grey ( K . & W . 7 - 8 D 5 ) , later dark b r o w n , w i t h
F i g . 3. Agaricus moellerianus. (habit X 'A). concolorous, entire edge. Stipe 35-65 X 11-17 m m , usually annulate
F i g . 4. Agaricus pampeanus. (habit X A).
w h e n y o u n g , tapering towards base, sometimes c y l i n d r i c a l , straight,

sometimes w i t h few white, thin rhizomorphs, stuffed, w h i t i s h , some-
what b r o w n i s h near base, above annulus w i t h p i n k i s h tinge, smooth, F i g . 5. Agaricus altipes. (habit X 'A).
b e l o w annulus w i t h white, sometimes at base pale b r o w n , floccose gir-
dles. A n n u l u s at 0.50-0.75 o f height o f stipe, narrow, 1-2 m m wide, Prag. 7 B (1): 13. 1951. - Psalliota aestivalis var. veneris H e i m &
descending, sometimes patent, thin, soon disappearing, fibrillose, B e c k e r i n B u l l , trimest. S o c . m y c o l . Fr. 76: 240. 1960 (invalid);
white, not d i s c o l o u r i n g , w i t h smooth upperside. Context up to 14 m m Agaricus aestivalis var. veneris H e i m & B e c k e r i n B u l l , trimest. Soc.
thick i n pileus, white, not d i s c o l o u r i n g . S m e l l indistinct, sometimes m y c o l . Fr. 76: 240. 1960 (invalid); Agaricus veneris ( H e i m & B e c k e r )
aromatic w h e n cut. Taste m i l d , not unpleasant. M . B o n i n D o c . m y c o l . 15(60): 7. 1985 (invalid).
M a c r o c h e m i c a l reactions: K O H 4 0 % b r o w n i s h (surface o f pileus, S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 31, 46. 1984 (as A. aestivalis);
base o f stipe); Schaeffer-reaction negative to w e a k l y orange (surface p f Breitenb. & K r a n z L , P i l z e S c h w e i z 4: p i . 167. 1995 (as A. aestivalis
pileus, surface o f stipe). ^ var. veneris); G i d h o l m i n Jordstjarnan 16: opposite 80. 1995.
Spores 7.5-10.5(-11.0) X 5.0-7.0 m m , on average 8.7-9.4 X 5.6- S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 46-48, 50-51 (as A.
6.3(-6.8) m m , Q = 1.25-1.75(-1.80), Q a v = (1.40-)1.45-1.55(-1.65), aestivalis), fig. 9, p i . 4a. 1950.
e l l i p s o i d , sometimes o b l o n g or broadly e l l i p s o i d , w i t h conspicuous V E R N . N A M E - Valse anijschampignon.
g e r m pore up to 1.5 u.m w i d e . B a s i d i a 26-30 X 8.0-9.0 m m , predom-
inantly 4-spored, u s u a l l y also ( l - ) 2 - s p o r e d basidia present. L a m e l l a Pileus 50-100 m m , at first hemispherical to convex, later convex to
edge fertile, without c h e i l o c y s t i d i a . P i l e i p e l l i s a cutis o f irregularly conico-convex, often w i t h flattened centre, sometimes umbonate, w i t h
arranged, 4.5-6.5(-8.5) m m w i d e hyphae w i t h c y l i n d r i c a l elements, inflexed margin, w h i t i s h w i t h usually pale y e l l o w i s h centre ( M u . 2.5 Y
w i t h some ascending clavate terminal elements, gradually passing into 8/2, 8/4, 10 Y R 8/3), later greyish to pale b r o w n i s h , especially near
pileitrama, w i t h pale y e l l o w i s h intracellular pigment. Stipitipellis a margin, usually y e l l o w i n g o n handling especially when young, y o u n g
cutis o f unbranched, 4.5-8.0 m m w i d e hyphae w i t h c y l i n d r i c a l , some- w i t h appressed, often y e l l o w i s h s i l k y fibrils to occasionally strongly
times up to 10.5 u.m inflated elements, w i t h pale y e l l o w i s h , parietal fibrillose, later w i t h larger fibrils, often w i t h adhering soil remnants, at
and intracellular pigment. margin sometimes flocculose-squamulose; v e i l present as appendicu-
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter- late w h i t i s h remnants. L a m e l l a e , L = 110-145,1 = 1-3, crowded, free,
restrial i n grasslands i n the coastal region, sometimes w i t h a direct subventricose, up to 8 m m broad, at first bright p i n k i s h b r o w n to pale
maritime influence. Rare, o n l y k n o w n f r o m the coastal region. A u g . - reddish b r o w n (2.5 Y R 5/6, 5 Y R 7/3), later dark greyish b r o w n , w i t h
O c t . ( - D e c ) . Rare but widespread i n Europe. A l s o recorded from north- entire, concolorous edge. Stipe (65-)70-100 X ( 9 - ) l l - 2 0 m m , usually
ern A f r i c a and S o u t h A m e r i c a . longer than pileus diameter, annulate, c y l i n d r i c a l to subclavate, at base
up to 23 m m broad, sometimes tapering towards base, straight to
T h i s taxon is distinguished from A . campestris by its larger spores w i t h somewhat curved, narrowly fistulose, white, i n lower half faintly y e l -
a m o r e conspicuous g e r m pore. T h e o r i g i n a l description o f A. pam- l o w i n g o n h a n d l i n g , at base d i s c o l o u r i n g y e l l o w to b r o w n i s h y e l l o w
peanus m a y differ i n the silvery, purely white, smooth pileus and a w h e n handled, above annulus sometimes reddish; above annulus
white stipe, and m i c r o s c o p i c a l l y i n smaller spores, 6-7 X 4-5 m m . glabrous, smooth, b e l o w annulus w i t h white to faintly y e l l o w silky fib-
However, a c c o r d i n g to H e i n e m a n n (in B u l l . Inst, agron. Stns R e c h . rils or fibrillose girdles, sometimes w i t h few fibrillose squamules.
G e m b l o u x 30: 273-282. 1962), w h o studied the type, the spore meas- A n n u l u s at 0.75-0.90 o f height o f stipe, up to 10 m m wide, descend-
ures are i n fact 8.5-10 X 5.5-7 m m . W i t h i n the scope o f this study this ing, pending, thin and evanescent, white, not discolouring, w i t h grey-
c o u l d not be verified. ish, striate upperside; underside sometimes w i t h thin fibrillose
A related taxon is Psalliota flocculosa R e a (in Trans. B r . m y c o l . squamules near m a r g i n ; m a r g i n sometimes split. Context 9-11 m m
Soc. 17: 50. 1932), w i t h a densely flocculose snow-white pileus. thick i n pileus, soft, white, usually o n l y faintly discolouring, i n upper
half o f stipe w i t h reddish tinge, i n base y e l l o w i s h to b r o w n i s h y e l l o w
4. Agaricus altipes (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B (1): 24. (2.5 Y 8/4, 8/6) w h e n cut. S m e l l indistinct to slightly fruity, sometimes
1 9 5 1 . - F i g . 5. l i k e Pelargonium or w e a k l y like Lepiota cristata. Taste indistinct to
Psalliota altipes F. M 0 l l e r i n F r i e s i a 4: 46. 1950. - Psalliota aesti- faintly nut-like.
valis F. M 0 l l e r i n F r i e s i a 4: 50. 1950; Agaricus aestivalis (F. M 0 l l e r ) M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus
Pilat i n A c t a M u s . nat. Prag. 7 B (1): 2 5 . 1 9 5 1 , n o n A . aestivalis Schum. and stipe); Schaeffer-reaction negative to slightly orange (surface o f
1803; A. albosericeus Rauschert i n N o v a H e d w i g i a 54: 213, 1992. - pileus; base o f stipe).
Psalliota aestivalis var. flavotactus F. M 0 l l e r i n F r i e s i a 4: 5 1 . 1950; Spores 6.5-9.0(-9.5) X 4.5-6.0 m m , o n average 7.6-8.0 X 4.9-5.5
Agaricus aestivalis var. flavotactus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. m m , Q = 1.30-1.70(-1.75), Q a v = 1.45-1.55, e l l i p s o i d , sometimes
AGARICUS (Nauta) 31
o b l o n g , w i t h narrow, indistinct g e r m pore, or t h i n - w a l l e d spot at apex. radially fibrillose o n paler, w h i t i s h to ochraceous or b r o w n i s h back-
B a s i d i a 15-20 X 8.0-9.0 m m , 4-spored. L a m e l l a edge fertile, usually ground; at first w i t h appendiculate floccose m a r g i n , sometimes w i t h
w i t h some sterile basidia. P i l e i p e l l i s a sometimes very slightly gelat- few w h i t i s h velar flocks near margin. L a m e l l a e , L = 8 0 - 9 0 , 1 = (0-
i n i z e d cutis o f sometimes branched, 3.5-6.5 |a.m w i d e hyphae w i t h ) l - 3 , crowded, free, ventricose, up to 7(-10.5) m m broad, at first p i n k -
usually c y l i n d r i c a l , rarely inflated elements u p to 9.0(-10.0) u.m w i d e , ish white to bright p i n k or greyish p i n k (7.5 Y R 8/2, 5 Y R 7/4, 8/2),
w i t h some ascending c y l i n d r i c a l terminal elements, gradually passing later reddish b r o w n (5 Y R 5-6/3), finally dark b r o w n (5 Y R 3/3), w i t h
into pileitrama, w i t h parietal, pale y e l l o w pigment. Stipitipellis a cutis paler, entire edge. Stipe 30-55(-80) X (7-)9-18(-22) m m , annulate or
o f 4.0-6.5 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes up to 8.5 [xm w i t h annular zone, c y l i n d r i c a l to attenuated towards base, straight to
inflated, elements, a n d some ascending clavate terminal elements, slightly curved, without r h i z o m o r p h s , n a r r o w l y fistulose, white to
w i t h y e l l o w i s h , parietal or intracellular pigment. w h i t i s h , later w i t h b r o w n i s h tinge, not d i s c o l o u r i n g or b e c o m i n g
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic, terrestrial brownish y e l l o w at base o n handling, above annulus b r o w n i s h to grey-
in parks, avenues, deciduous or coniferous woods, sometimes i n dune ish, minutely flocculose, b e l o w annulus flocculose-squamulose, occa-
scrub, often o n nutrient-rich or h u m u s - r i c h soil. Rather rare and scat- sionally w h e n y o u n g w i t h squamules i n girdles turning b r o w n i s h o n
tered i n the Netherlands. June-Oct. Rather rare but widespread i n handling. A n n u l u s at 0.60-0.70 o f height o f stipe, narrow, up to 2 m m
Europe. A l s o k n o w n from N o r t h A m e r i c a and southern South A m e r i c a . w i d e , descending, pending, thin, fibrillose, soon disappearing, white,
not discolouring, often w i t h thickened margin, w i t h striate upperside;
T h e above description i s also based o n collections from D e n m a r k . underside squamulose. Context 10 m m t h i c k i n p i l e u s , white, dis-
Because o f the y e l l o w discoloration o f the pileus and the occasion- c o l o u r i n g w e a k l y p i n k i s h b r o w n i s h i n apex o f stipe, sometimes pale
a l l y split m a r g i n o f the annulus this taxon is sometimes considered to b r o w n i n cortex o f stipe, i n pileus faintly reddish w h e n cut, y e l l o w i s h
b e l o n g to sect. Arvenses. T h e presence o f a t h i n - w a l l e d apex o f the to orange-brown i n base. S m e l l indistinct to slightly fungoid-aromat-
spores and the absence o f cheilocystidia, and also the reddish d i s c o l - ic. Taste fungoid.
oration i n the upper part o f the stipe place it clearly close to A. M a c r o c h e m i c a l reactions: K O H 10% negative (pileus surface, c o n -
campestris i n sect. Agaricus. text); Schaeffer-reaction negative (surface o f pileus).
Agaricus aestivalis (F. M 0 l l e r ) Pilat i s considered conspecific o n Spores 7.0-9.5 X 5.0-6.0 m m , o n average 7.6-8.3 X 5.1-5.5 m m , Q
account o f the same discoloration o f the context i n the stipe, and the = 1.35-1.65(-1.70), Q a v = 1.49-1.52, e l l i p s o i d , sometimes o b l o n g ,
microscopy. T h e intensity o f y e l l o w i n g o f the pileus seems to be vari- with narrow germ pore, or w i t h thin-walled spot at apex. B a s i d i a 2 0 -
able. Agaricus wasseri B o n & C o u r t e c . ( i n D o c . m y c o l . 15(60): 7. 30 X 7.0-10 m m , 4-spored. L a m e l l a edge fertile or partly sterile, usu-
1985; n o m . nov. for Agaricus longicaudus Wasser) may be a s y n o n y m . ally only composed o f basidia, sometimes with few clavate
cheilocystidia, 2 2 X 8.5-10.0 | x m . P i l e i p e l l i s a slightly irregular cutis
5. Agaricus cupreobrunneus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. of 3.0-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes up to 8.5 |jim
7 B ( 1 ) : 14. 1 9 5 1 . - F i g . 6. inflated elements, w i t h c y l i n d r i c a l terminal elements, gradually pass-
Psalliota campestris var. cupreobrunnea J . Schaeff. & Steer i n ing into pileitrama, w i t h y e l l o w i s h to b r o w n i s h parietal pigment.
M i c h a e l , Fuhr. Pilzfr. I: 147. 1939 (invalid); Psalliota cupreobrunnea Stipitipellis a cutis o f 5.5-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l ele-
J. Schaeff. & Steer ex F. M 0 l l e r i n F r i e s i a 4: 54. 1950. ments, w i t h pale yellow, intracellular pigment.
S E L . I C O N . - B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: p i . 57. 1977; H A B I T A T & DISTR. - Solitary, saprotrophic, terrestrial i n relatively
C a p p e l l i , A g a r i c u s : p i . 12. 1984; F. M 0 l l e r i n F r i e s i a 4: p i . 4b. 1950. nutrient-poor grasslands and roadside verges, often o n calcareous dune
S E L . D E S C R . & F I G S . - B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: 1-3. sand. Rather rare, more c o m m o n i n dune region o f the Netherlands;
1977; F. M 0 l l e r i n F r i e s i a 4: 54-55. 1950. Sept.-Nov. Widespread but not c o m m o n i n Europe. A l s o recorded from
V E R N . N A M E - Bruine weidechampignon. A s i a and N o r t h A m e r i c a .
P i l e u s (25-)40-60(-110) m m , at first convex w i t h flattened centre, 6. Agaricus porphyrocephalus F. M l l e r i n F r i e s i a 4: 204. 1952. - F i g .

expanding to plano-convex, often w i t h depressed centre, sometimes 7.
irregularly undulated plano-convex, w i t h i n f l e x e d , fringed m a r g i n , Psalliotaporphyrea F. M 0 l l e r i n F r i e s i a 4: 53. 1950; Agaricuspor-
later exceeding lamellae, y o u n g p i n k i s h b r o w n to p i n k i s h grey, soon phyreus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B (1): 13. 1951, n o n
darker to reddish b r o w n , often i n part dark b r o w n to (dark) purple- A. porphyreus ( A . & S.:Fr.) Fr. 1821.
b r o w n ( M u . 7.5 Y R 6-4/2, 5 Y R 4-5/3, 6/4), f l o c c o s e - f i b r i l l o s e to S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 13. 1984; F. M 0 l l e r i n Friesia 4:
entirely fibrillose, at centre later often smooth, towards m a r g i n dark p i . 4c. 1950.
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 53-54. 1950.
VERN. NAME - Porfierchampignon.
F i g . 6. Agaricus cupreobrunneus. (habit X A). F i g . 7. Agaricus porphyrocephalus. (habit X 'A).

Pileus 45-65(-85) m m , at first truncately c o n i c a l , or hemispherical to Sect. Bitorques (Heinem.) B o n & Cappelli
convex w i t h flattened centre, expanding to convex to irregularly plano-
convex w i t h u s u a l l y (slightly) depressed o r flattened centre, w i t h at Pileus not discolouring or slightly y e l l o w i n g , rarely slightly reddening;
first inflexed, later deflexed margin, w h i c h exceeds lamellae for 2 m m , annulus ascending and thin to thick or descending and thick, sometimes
pale to dark (purple-)brown to reddish brown, sometimes grey-brown only velar sock present; context usually discolouring reddish or b r o w n -
fibrillose-squamulose ( M u . 2.5 Y R 2/4, 5 Y R 3-4/2, 3-6/3, 1 0 Y R 2/2- ish o n exposure or damaging, sometimes not discolouring.
3/4) o n greyish b r o w n to pale b r o w n background (2.5 Y R 5/4, 5 Y R K O H - r e a c t i o n negative; Schaeffer-reaction negative.
7/3, 7.5 Y R 6/4, 10 Y R 7/3, 6/4). L a m e l l a e , L = 85-100, 1 = 1-3, Spores (broadly) e l l i p s o i d , relatively broad, Q a v < 1.35(-1.45),
crowded, free, subventricose, 6-11 m m broad, at first pale greyish p i n k without germ pore; l a m e l l a edge sterile; c h e i l o c y s t i d i a conspicuous,
to bright p i n k i s h , later p i n k i s h b r o w n (7.5 Y R 8/4-7/4), finally dark clavate to c y l i n d r i c a l , up to 15 |a.m w i d e .
b r o w n to p u r p l i s h b r o w n (5 Y R 3/2 - 7.5 Y R 5/4), w i t h slightly paler,
entire edge. Stipe 30-45(-70) X 9-12(-14) m m , annulate o r w i t h annu- 7. Agaricus bitorquis (Quel.) S a c c , S y l l . F u n g . 5: 998. 1887. - F i g .
lar zone, c y l i n d r i c a l to ventricose, rarely tapering downwards or 8.
clavate, sometimes at base up to 19 m m broad, straight to slightly Psalliota bitorquis Q u e l , i n C . r. A s s . franc. A v . S c i . 12: 500. 1884
curved, narrowly fistulose to stuffed, w h i t i s h , often w i t h b r o w n i s h (Champ. Jura Vosges S u p p l . 12). - Agaricus campestris var. edulis
tinge, somewhat ochraceous to b r o w n i n g towards base o n handling, V i t t , Descr. F u n g h i m a n g . Italia: 4 1 . 1832; Agaricus edulis (Vitt.)
above annulus smooth to s i l k y striate, below annulus flocculose-sub- M 0 l l e r & Schaeff. i n A n n l s m y c o l . 36: 75. 1938. -Psalliota edulis var.
squamulose, sometimes w i t h several fibrillose girdles. A n n u l u s at 0.45- validus F. M 0 l l e r i n Friesia 4: 14. 1950; Agaricus bitorquis var. validus
0.60 o f height o f stipe, narrow, u p to 2.5 m m wide, descending, thin, (F. M 0 l l e r ) B o n & C a p p e l l i i n D o c . m y c o l . 13(52): 16. 1983.
fugacious, fibrillose, white, not d i s c o l o u r i n g , w i t h striate upperside; S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 1. 1984; Dahncke, 1200 P i l z e :
underside subfloccose-fibrillose; often o n l y fibrillose remnants pres- 486. 1993; R . P h i l l i p s , Paddest. S c h i m m . : 163. 1981.
ent. Context 7-9 m m t h i c k i n pileus, white to w h i t i s h , hardly dis- S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 13-14. 1950; Wasser,
c o l o u r i n g w h e n cut, after a l o n g time somewhat sordid p i n k i s h or Tr. Agariceae Soviet U n i o n : 52-54. 1989.
faintly p i n k i s h b r o w n i n pileus, i n base o f stipe somewhat p i n k i s h y e l - VERN. NAME - Straatchampignon.
l o w i s h . S m e l l faint w h e n cut, w h e n y o u n g distinctly aromatic, later
faintly fungoid. Taste fungoid. Pileus ( 4 5 - ) 5 0 - l 10(-120) m m , at first convex, later convex, usually
M a c r o c h e m i c a l reactions: Schaeffer-reaction negative o r doubtful w i t h depressed-centre, to applanate, w i t h inflexed or involute, later
(surface o f pileus). straight margin, dirty white to greyish white, w i t h y e l l o w spots o r y e l -
Spores (5.5-)6.0-7.5(-8.0) X 4.0-5.0 m m , o n average 6.4-6.9 X l o w i n g o n handling ( M u . 2.5 Y 8/2^9/4), rarely reddening o n handling
4.1-4.3 m m , Q = (1.40-)1.45-1.75(-1.80), Q a v = 1.55-1.60, e l l i p s o i d (7.5 Y R 7/6); surface smooth -rarely w i t h some appressed
7 squames,
to o b l o n g , w i t h t h i n - w a l l e d spot at apex. B a s i d i a 21-25 X 6.0-8.0 sometimes fibrillose to fibrillose-squamulose near margin, shiny, usu-
mm, 4-spored. L a m e l l a edge fertile, c o m p o s e d o f fertile b a s i d i a , ally w i t h attached sand; m a r g i n w i t h some adhering v e i l remnants
sometimes i n t e r m i x e d w i t h some sterile b a s i d i a o f the same size and when y o u n g . L a m e l l a e , L = 95-200,1 = 0-3, crowded, free, straight,
shape. P i l e i p e l l i s a regular cutis o f 3.5-5.0(-7.0) m m w i d e hyphae up to 6.5 m m broad, at first pink, later b r o w n , finally dark b r o w n , w i t h
w i t h c y l i n d r i c a l , sometimes u p to 10 u.m inflated elements, w i t h paler, denticulate edge. Stipe 35-55(-100) X 14-20(-24) m m , annulate,
some ascending, clavate terminal elements up to 8.5 |xm w i d e , grad- c y l i n d r i c a l to clavate, at base up to 29 m m broad, straight, stuffed o r
u a l l y passing into p i l e i t r a m a , w i t h pale y e l l o w to b r o w n i s h , parietal narrowly fistulose, w h i t i s h , not d i s c o l o u r i n g o n handling, above annu-
to intracellular pigment. Squames consisting o f slightly curved, inter-
w o v e n , 4.0-6.0 m m w i d e hyphae w i t h 3 0 - 4 0 m m l o n g , c y l i n d r i c a l
elements a n d ascending, clavate, u p to 10 | x m w i d e t e r m i n a l ele-
ments, w i t h w a r m b r o w n , intracellular pigment. S t i p i t i p e l l i s a some-
times irregular cutis o f 4.0-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l ,
sometimes up to 9.5 | x m inflated, elements, w i t h pale y e l l o w parietal
or intracellular pigment.
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, sometimes i n fairy
rings, saprotrophic and terrestrial i n unmanured meadows or roadside
verges o n nutrient-poor s o i l . Very rare i n the Netherlands, O c t . - N o v .
Widespread but rare i n Europe.
Agaricus lividonitidus (F. M 0 l l e r ) P i l a t i s very close, and perhaps a

s y n o n y m . It seems to differ i n a m o r e y e l l o w i n g base o f the stipe.
Agaricus calongei D a n i e l - A r r a n z (in B o l . S o c . m i c o l . M a d r i d 24: 72.
1999) is very p r o b a b l y a s y n o n y m o f A . lividonitidus. S i n c e no type-
material has been studied o f any o f the t w o species this s y n o n y m y
c o u l d not b e made certain. A l s o , Agaricus biberi Hlavacek (in
M y k o l . Sb., P r a h a 6 1 : 74. 1984; i n v a l i d : no type designated), w i t h a
greyish instead o f p u r p l i s h b r o w n pileus, i s c l o s e l y related to A. por-
phyrocephalus.
F i g . 8. Agaricus bitorquis. (habit X 'A).

AGARICUS (Nauta) 33
lus smooth, b e l o w annulus fibrillose, sometimes w i t h velar sock form-

ing narrow ascending annulus at upper side at 0.25 o f height o f stipe,
without rhizomorphs. A n n u l u s at 0.65 o f height o f stipe, up to 11 m m
w i d e , ascending, double, usually upper m a r g i n patent or upwards and
lower m a r g i n to a lesser extent recurved, occasionally w i t h a second,
narrower annulus some distance below, thick, persistent, w h i t i s h to
b r o w n i s h (10 Y R 8/4), w i t h striate to sulcate upperside; underside
smooth. Context 12-23 m m thick i n pileus, firm, white, d i s c o l o u r i n g
reddish b r o w n or orange-red w h e n cut i n pileus above lamellae and
upper half o f stipe, sometimes faintly. S m e l l unpleasant, like canned
applesauce, or l i k e gas. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus;
context); Schaeffer-reaction negative (surface o f pileus).
Spores 5.0-7.0 X 4.0-5.5 m m , o n average 5.9-6.5 X 4.6-4.9 m m , Q
= 1.10-1.45, Q a v = 1.20-1.35, broadly e l l i p s o i d to ellipsoid, without
g e r m pore; sometimes w i t h aberrant spores up to 9 X 5 |xm. B a s i d i a
20-25 X 6.5-8.5 m m , u s u a l l y 4-spored, often also 2-spored basidia
present. L a m e l l a edge w i t h a narrow, 30-50 m m broad, almost sterile
layer consisting o f cheilocystidia intermixed w i t h some basidia; cheilo-
c y s t i d i a u s u a l l y not i n chains, 16-28 X 5.5-9.0(-13) p,m, clavate to
c y l i n d r i c a l , hyaline, rarely w i t h some basal septa, f o r m i n g rectangular
elements o f 3.0-7.0 X 4.0-5.0 u.m. P i l e i p e l l i s an often slightly gelati-
n i z e d cutis o f irregularly radially arranged, 2.5-5.5 m m w i d e hyphae
w i t h c y l i n d r i c a l elements, w i t h c y l i n d r i c a l terminal elements, w i t h
very pale y e l l o w intracellular pigment, gradually passing into pileitra- F i g . 9. Agaricus bernardii. (habit X Vi).
ma. Stipitipellis a cutis o f unbranched, irregularly arranged, 2.5-5.5

m m w i d e hyphae w i t h c y l i n d r i c a l elements. or reddish o n handling, especially i n lower half, fibrillose to slightly
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter- squamulose, above annulus smooth to slightly fibrillose and sometimes
restrial i n roadside verges, gardens, c i t y parks, o n various, usually later greyish, below annulus fibrillose; sometimes o n l y w i t h thick, fib-
nutrient-rich soils. Rather c o m m o n and widespread i n the Netherlands. rillose, w h i t i s h velar sock up to 0.45-0.60 o f height o f stipe; sometimes
May, J u l y - N o v . W i d e s p r e a d and relatively c o m m o n i n Europe. y e l l o w i n g on handling. A n n u l u s at 0.35-0.60 o f height o f stipe, (2-)6-
Cosmopolitan. 10 m m wide, ascending, patent or upward, y o u n g thick, rarely double,
later thin, rarely persistent, fibrillose, w h i t i s h to pale b r o w n , rarely y e l -
8. Agaricus bernardii Quel, in Cooke & Quel., Clav. syn. l o w i s h , w i t h smooth to striate upperside; underside fibrillose. Context
H y m e n o m y c . eur.: 89. 1878. - F i g . 9. ( l l - ) 1 5 - 2 5 m m thick i n pileus, f i r m to soft, white to w h i t i s h or pale
Psalliota ingrata F. M 0 l l e r i n F r i e s i a 4: 17. 1950; Agaricus ingra- brown, d i s c o l o u r i n g s l o w l y to q u i c k l y flesh-coloured to p i n k or red-
tus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. P r a g . 7 B (1): 5. 1951, non A. dish when cut, especially i n pileus, orange to orange-brown i n base o f
ingratus S c h u m . 1803:Fr.; Agaricus maleolens F. M 0 l l e r i n F r i e s i a 4: stipe. S m e l l usually unpleasant, v a r y i n g f r o m strong to faint, carbolic,
203. 1952. metallic or with dung component. Taste fungoid.
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 2. 1984; R . P h i l l i p s , Paddest. M a c r o c h e m i c a l reactions: a m m o n i a negative (surface o f pileus and
S c h i m m . : 161. 1981; Wasser i n F u n g . rar. Ic. c o l . 10: p i . 76. 1979. stipe, context o f pileus and stipe); K O H 1 0 % negative (surface o f
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 14-17, fig. 4, p i . 7; 17- pileus); Schaeffer-reaction negative (surface o f pileus and base o f stipe).
19, fig. 5, p i . 8 (as P. ingrata). 1950; Wasser i n F u n g . rar. Ic. c o l . 10: Spores 5.5-7.5(-8.0) X 4.5-6.5 m m , o n average (6.0-)6.6-7.3 X 4.9-
9. 1979. 6 . 1 m m , Q = 1.10-1.40(-1.50), Q a v = 1.15-1.25(-1.35), broadly ellip-
VERN. NAME - Kwelderchampignon. soid to e l l i p s o i d , without g e r m pore. B a s i d i a 27-32 X 7.0-7.5 m m ,
4-spored. L a m e l l a edge w i t h a 50-80 m m broad sterile layer; c h e i l o -
Pileus 65-125 m m , at first hemispherical, later convex to plano-con- cystidia not i n chains, (27-)35-58 X (4.5-)6.5-10(-12.5) u.m, c y l i n d r i -
vex, often w i t h flattened centre, w i t h broad, white, involute, later cal to narrowly clavate, often flexuous, sometimes irregular,
inflexed m a r g i n exceeding lamellae, w h i t e or g r e y i s h white to pale occasionally w i t h a few basal septa, f o r m i n g rectangular elements o f
b r o w n ( M u . 10 Y R 8/1-2,7/2), later rarely w i t h slightly y e l l o w or pink- 8.0-15 X 3.3-5.0 m m , usually w i t h b r o w n i s h contents, often borne by
ish patches or w i t h y e l l o w i s h fibrils (10 Y R 8/4-6); surface smooth or long oleiferous hyphae. P i l e i p e l l i s a slightly gelatinized cutis o f irreg-
innately fibrillose to appressed squamose or squamulose, sometimes ularly radially arranged, (2.0-)3.5-5.5 m m w i d e hyphae w i t h c y l i n d r i -
w i t h few large, unclearly d e l i m i t e d innate squames, occasionally are- cal elements, w i t h very pale y e l l o w , intracellular pigment gradually
olate-rimose, near m a r g i n sometimes fissurate. L a m e l l a e , L = 65-180, passing into pileitrama,. Stipitipellis a cutis o f unbranched, 3.0-5.5 m m
1 = l-5(-7), crowded, free, straight to undate, usually not very broad, w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h very pale y e l l o w intra-
sometimes u p to 11 m m broad, at first p i n k i s h to pale grey-brown (10 cellular pigment.
Y R 7/3), later reddish grey, finally dark b r o w n (7.5 Y R 3/2, 4/2), w i t h H A B I T A T & DISTR. - Solitary or i n groups, saprotrophic, terrestrial,
w h i t i s h or concolorous, granulate to denticulate edge. Stipe (30-)55- on soil i n roadside verges, i n (dune) grasslands or grazed pastures or
80(-120) X (15-)20-30 m m , usually annulate, c y l i n d r i c a l or subclavate saltings, often on eutrophicated places, o n clayey or sandy soil. Rather
to fusiform, sometimes slightly enlarged at base, straight, stuffed, rare, but widespread i n the Netherlands. June, Sept.-Nov. Widespread
w h i t i s h to pale b r o w n , sometimes later d i s c o l o u r i n g to orange-brown i n Europe. C o s m o p o l i t a n .
Agaricus maleolens F. M 0 l l e r , as described and depicted b y M 0 l l e r (in absent. Context 15 m m thick i n pileus, firm, white, not or very faintly
Friesia 4: 17-19, p i . 8, fig. 5; as P. ingrata), is considered conspecific b r o w n w h e n cut, i n base o f stipe orange-yellow (7.5 Y R 8/5). S m e l l
with A. bemardii. E x c e p t for the areolate-rimose surface o f the pileus i n weak to strong, fungoid or unpleasant. Taste not k n o w n .
specimens collected near the sea, no other possible distinguishing char- M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus);
acters c o u l d be found. Some authors (e.g. B o n i n D o c . m y c o l . 15(60): Schaeffer-reaction negative (surface o f pileus, base o f stipe).
13. 1985) interpret A. maleolens as a species w i t h smaller spores than A, Spores 6.5-8.5 X 5.5-6.5(-7.0) m m , o n average 7.7 X 5.7-6.2 m m ,
bemardii, w h i c h is not i n accordance w i t h its original sense. Q = 1.15-1.45(-1.60),Qav = 1.25-1.35, broadly e l l i p s o i d to e l l i p s o i d ,
A n o t h e r variant without an annulus but w i t h a conspicuous velar without germ pore. B a s i d i a 26-34 X 8.5-9.5 m m , 4-spored. L a m e l l a
sock differed i n no other aspect from the typical form, and is included edge w i t h a narrow sterile layer; c h e i l o c y s t i d i a 20-36 X 8.5-10 (xm,
i n the above description. c y l i n d r i c a l to narrowly clavate, w i t h h y a l i n e to pale b r o w n contents.
C l o s e l y related species are Agaricus bernardiiformis Bohus (in P i l e i p e l l i s a slightly gelatinized cutis o f 4.0-5.5 m m w i d e hyphae w i t h
A n n l s hist.-nat. M u s . natn. hung. 67: 37. 1975), differing i n the not dis- c y l i n d r i c a l elements, w i t h pale y e l l o w intracellular pigment, gradual-
c o l o u r i n g context a n d faint and not unpleasant s m e l l , and Agaricus ly passing into pileitrama. Stipitipellis a cutis o f unbranched, 3.0-7.0
robynsianus H e i n e m . (in B u l l . Jard. bot. Etat 27: 449. 1957), differing m m w i d e hyphae w i t h c y l i n d r i c a l elements w i t h pale y e l l o w intracel-
in a less reddening context and a s m e l l like Lepiota cristata. lular pigment. V e l a r sock consisting o f interwoven, slightly aggluti-
nated, branched, 3.0-5.5 m m w i d e hyphae w i t h c y l i n d r i c a l elements,
9. Agaricus gennadii ( C h a t i n & B o u d . ) P . D . O r t o n i n Trans. B r . w i t h pale y e l l o w intracellular pigment.
m y c o l . S o c . 4 3 : 174. 1960. - F i g . 10. H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter-
Chitonia gennadii C h a t i n & B o u d . i n J . B o t . , Paris 12: 66. 1898. restrial i n roadside verge o n slush o f ditch, i n Picea-wood o n calcare-
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 5. 1984. ous s o i l . V e r y rare i n the Netherlands (Oegstgeest; W i j l r e ) , Sept.
S E L . D E S C R . & F I G S . - P . D . O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : 181- Scattered and very rare i n Europe, more c o m m o n i n the mediterranean
182. 1960; R o m a g n . i n B u l l . F e d . m y c o l . D a u p h . Savoie 100: 9-10. region. A l s o recorded from northern A f r i c a and Israel.
1986.
V E R N . N A M E - Beurschampignon. Agaricus gennadii can superficially resemble variants o f A. bemardii
without an annulus. It differs f r o m A. bemardii i n the larger spores,
Pileus 65-100 m m , at first hemispherical, expanding to convex, some- fusiform stipe w i t h pseudorhiza and the hardly d i s c o l o u r i n g context.
times w i t h depressed centre, w i t h first involute, later straight margin, Agaricus vQlvatus ( A . P e a r s ^ H e T n e m . is closely related or perhaps
w h i t i s h to y e l l o w i s h ( M u . 2.5 Y 8/2), later also w i t h b r o w n i s h spots conspecific. Agaricus slovenicus H l a v a c e k (in M y k o l . Sb., Praha 6 9 :
(10 Y R 8/6), not d i s c o l o u r i n g o n handling; surface smooth to slightly 129. 1992) is an i n v a l i d name since no type was designated. It m a y be
fibrillose, w i t h m u c h adhering soil when young. L a m e l l a e , L = 145- very close to or perhaps conspecific w i t h this taxon. Agaricus pequinii
175,1 = 1-5, crowded, free, subventricose, up to 5 m m broad, at first (Boud.) Konr. & M . ( = A. gennadii ssp. microspores B o h u s ) i s a close-
p i n k i s h , later dark b r o w n , w i t h paler, entire to pruinose edge. Stipe 40- l y related species, differing i n the smaller spores and larger basidio-
65 X 15-27 m m , fusiform, at base up to 25 m m broad, straight, stuffed, carps, w h i c h occurs more to the south i n E u r o p e . Agaricus rollanii
w h i t i s h , w i t h w h i t i s h velar sock up to 0.50 o f height o f stipe, w i t h Parra (in B o l . S o c . m i c o l . M a d r i d 20: 132. 1995) i s p r o b a b l y also a
ascending, patent, 3 m m broad margin, above velar sock w i t h greyish closely related taxon.
b r o w n (10 Y R 7/3) fibrillose-flocculose girdles, at base faintly y e l -
l o w i s h b r o w n (9 Y R 7.5/6) o n h a n d l i n g , w i t h pseudorhiza. A n n u l u s 10. Agaricus bisporus (J. Lange) Imbach i n M i t t , naturf. Ges. L u z e r n
15: 15. 1946. - F i g . 11. D a n s k bot. A r k . 4(12): 8. 1926; Psalliota bis-
pora (J. L a n g e ) M 0 l l e r & Schaeff. i n A n n l s m y c o l . 3 6 : 69. 1938. -
Agaricus campestris var. hortensis C o o k e , H a n d b . B r i t . F u n g i : 138.
1870; Psalliota hortensis (Cooke) J . L a n g e , i n D a n s k bot. A r k . 4(12):
8. 1926; Agaricus hortensis (Cooke) Pilat i n A c t a M u s . nat. P r a g . 7 B
(1): 6. 1951, non A. hortensis Pers. 1801:Fr. - Psalliota hortensis f.
albida J . L a n g e i n D a n s k bot. A r k . 4(12): 9. 1926.
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 7. 1984; D a h n c k e , 1200 P i l z e :
485. 1993; R . P h i l l i p s , Paddest. S c h i m m . : 163. 1981.
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 10-11. 1950.
V E R N . N A M E - Gekweekte champignon.
Pileus (35-)60-120(-130) m m , at first hemispherical or conico-convex,

later convex to plano-convex w i t h flattened or slightly depressed cen-
tre, w i t h inflexed, later deflexed margin, light b r o w n to b r o w n at cen-
tre ( M u . 5 Y R 5/6-4/3, 7.5 Y R 3/2, 5/4, 7/6, 10 Y R 6-7/4), towards
margin paler, b r o w n fibrillose or covered w i t h b r o w n appressed, fib-
rillose pointed r a d i a l l y arranged squames (5 Y R 4/3, 7.5 Y R 4/4, 10
Y R 6/3-4, 5/4, 8/3) o n w h i t i s h to pale b r o w n background, sometimes
entirely white (cultivated variant), not d i s c o l o u r i n g o n handling; mar-
gin exceeding lamellae for u p to 4 m m w i t h w h i t i s h rim. L a m e l l a e , L
= 90-175,1 = 1-3, crowded, free, straight, up to 8 m m broad, at first
p i n k i s h grey-brown to p i n k i s h b r o w n ( K . & W . 8 C - D 5 ; M u . 2.5 Y R
6/6-10 R 6/6), finally dark b r o w n (7.5 Y R 5/4, 5 Y R 4/4), w i t h w h i t i s h
F i g . 10. Agaricus gennadii. (habit X A). to paler b r o w n i s h flocculose to denticulate edge. Stipe (30-)40-60(-80)
AGARICUS (Nauta) 35
c o m m o n , widespread. M a y - N o v . C o m m o n and widespread i n Europe.

Cosmopolitan.
The cultivated variant w i t h the almost white pileus is sometimes dis-

tinguished at species level as A. hortensis, but has n o t a x o n o m i c a l
importance. T h i s variant is rarely found i n nature.
Agaricus brunnescens P e c k is considered a different taxon, differ-
ing i n the not discolouring context and 4-spored basidia. T h i s taxon is
by some authors regarded as conspecific w i t h A. bisporus and used as
the correct name for the cultivated m u s h r o o m . F o r a discussion see
K e r r i g a n (in Wuest et a l . , C u l t i v a t i n g edible fungi: 141-154. 1987),
M a l l o c h (in M y c o l o g i a 79: 839-846. 1987) and Singer ( i n M y c o t a x o n
20: 479-482. 1984).
11. Agaricus devoniensis P.D. Orton i n Trans. B r . m y c o l . S o c . 4 3 : 173.

1960. - F i g . 12
Psalliota arenicola W a k . & Pears, i n Trans. B r . m y c o l . S o c . 2 9 :
205. 1946; Agaricus arenicola (Wak. & Pears.) Pilat, K i t e urc. H u b
hfib. bedl.: 4 0 1 . 1951, n o n A. arenicola Berk. 1843; Agaricus
arenophilus H u i j s m a n i n Persoonia 1: 324. 1960 (superfluous).
S E L . I C O N . - Cappelli, Agaricus: p i . 4 . 1 9 8 4 ; D a l Savio i n B o l l . Gruppo
m i c o l . G . Bres. 27: 155. 1984; Gennari i n Riv. M i c o l . 37: 11. 1994.
S E L . D E S C R . & F I G S . - M . B o n i n D o c . m y c o l . 19(76): 7 5 . 1989; F .
F i g . 11. Agaricus bisporus. (habit X A). M 0 l l e r i n F r i e s i a 4: 199-201, f i g . 3 3 . 1952 (as Psalliota arenicola).
V E R N . N A M E - Zeeduinchampignon.
X (9-) 12-20 m m , annulate, c y l i n d r i c a l to subclavate, slightly attenuat-

ed or enlarged at base up to 23 m m broad, straight to curved, narrow- Pileus (25-)30-50(-55) m m , at first convex, later plano-convex to
l y fistulose, w h i t i s h , occasionally b r o w n or reddish b r o w n on handling, applanate o r convex w i t h flattened o r depressed centre, w i t h inflexed,
especially towards base, above annulus smooth, satiny, sometimes pale later straight margin exceeding lamellae, white to w h i t i s h ( M u . 10 Y R
p i n k i s h b r o w n , b e l o w annulus usually flocculose o r w i t h some floc- 8/1), sometimes w i t h pale b r o w n centre, sometimes later w i t h faintly
cose v e i l remnants, without rhizomorphs o r o c c a s i o n a l l y w i t h a few yellowish spots o r y e l l o w i s h at centre (2.5 Y 8/4); surface innately to
short, thick, white r h i z o m o r p h s . A n n u l u s at 0.50 - 0.80 o f height o f
stipe, 5-8 m m wide, ascending or descending, patent to pending, thick,
c. triangular i n transverse section, persistent, w h i t i s h , not discolouring,
w i t h striate or sometimes sulcate upperside; underside floccose, or
w i t h b r o w n i s h squames, sometimes w i t h i n c i s e d margin. Context 10-
15 m m thick i n pileus, firm, w h i t i s h , (faintly) d i s c o l o u r i n g p i n k i s h or
b r o w n i s h w h e n cut, especially i n pileus and upper part o f stipe, i n base
of stipe sometimes y e l l o w i s h pink. S m e l l herb-like, spicy, fungoid, like
nuts. Taste fungoid, not very distinct.
M a c r o c h e m i c a l reactions: K O H 10% negative (context o f stipe;
surface o f pileus); Schaeffer-reaction negative (surface o f pileus, sur-
face o f base o f stipe).
Spores 6.5-8.0 X (5.0-)5.5-6.5 m m , o n average 7.1-7.3 X 5.4-5.9
mm, Q = 1.10-1.45, Q a v = 1.20-1.30, broadly e l l i p s o i d , sometimes
ellipsoid, without germ pore. B a s i d i a 20-23 X 7.0-8.0 m m , 2-spored,
usually w i t h some 1-spored present. L a m e l l a edge w i t h a 40-75 m m
broad sterile layer; c h e i l o c y s t i d i a 16.5-33(-36) X (6.5-)8.5-12 m m ,
narrowly clavate to sometimes utriform, occasionally w i t h some basal
septa, f o r m i n g short chains o f 2-3 rectangular elements, 8.0-13 X 5.0-
5.5 m m , w i t h colourless or often b r o w n i s h contents. Pileipellis a slight-
l y gelatinized cutis o f unbranched, 3.0-6.0 m m w i d e hyphae w i t h
c y l i n d r i c a l elements, w i t h clavate to c y l i n d r i c a l terminal elements,
gradually passing into pileitrama, w i t h b r o w n i s h intracellular pigment.
F i b r i l s consisting o f curved, up to 5.0 m m w i d e hyphae, w i t h b r o w n -
ish intracellular and parietal pigment. Stipitipellis a regular cutis o f
unbranched, 3.0-5.5 m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h
pale y e l l o w intracellular pigment.
H A B I T A T & DISTR. - S o l i t a r y o r i n s m a l l groups, saprotrophic, ter-
restrial o n s o i l along roads, i n roadside verges, i n parks, more rarely i n
(fertilized) meadows o r i n d r y dunes, o n various s o i l types. Rather F i g . 12. Agaricus devoniensis.
appressed radially fibrillose, w i t h m u c h adhering sand. L a m e l l a e , L =

80-95,1 = l-3(-9), crowded, free, subventricose, up to 6 m m broad, at
first greyish brown to p i n k i s h grey (7.5 Y R 6/2,5/4), finally dark brown,
w i t h w h i t i s h to concolorous, entire edge. Stipe 25-50 X 5-10(-15) m m ,
usually annulate, occasionally w i t h additional velar zones below, some-
times o n l y w i t h floccose velar zones, c y l i n d r i c a l , sometimes slightly
enlarged at base, straight, narrowly fistulose, white, fibrillose, above
annulus smooth to subfloccose or w i t h few flocks, sometimes w i t h red-
d i s h hue, b e l o w annulus floccose, especially at base, often w i t h
m y c e l i a l c l o d , sometimes w i t h few thin white rhizomorphs. A n n u l u s at
c. 0.50 o f height o f stipe, narrow, c. 3 m m wide, ascending, slightly
patent, thin, fibrillose, white, w i t h smooth upperside; underside fibril-
lose. Context 7 m m thick i n pileus, white, not discolouring or dis-
colouring faintly p i n k i s h or brownish when cut, especially i n upper part
of stipe. S m e l l indistinct or faintly like radish. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus);
Schaeffer-reaction negative (surface o f pileus and base o f stipe).
Spores (6.0-)6.5-7.5 X (4.0-)4.5-5.5 m m , on average (6.1-)6.6-6.9 X
4.6-5.3 m m , Q = 1.20-1.40(-1.55), Qav = 1.25-1.30(-1.45), ellipsoid to
broadly ellipsoid, without germ pore. B a s i d i a 20-25 X 7.0-9.0 m m , usu-
ally 4-spored, occasionally also 2-spored basidia present. L a m e l l a edge
with an up to 40 m m broad sterile layer; cheilocystidia not i n chains, (15-
)20-33 X 9.0-13(-15.5) m m , clavate, hyaline or w i t h brown contents. F i g . 13. Agaricus subfloccosus. (habit X 'A).
Pileipellis a slightly gelatinized cutis o f slightly irregularly and loosely
arranged, 2.5-7.5 m m wide hyphae w i t h cylindrical elements, sometimes
inflated up to 9.5 |xm, gradually passing into pileitrama, w i t h very»pale white, pruinose edge. Stipe (30-)50-60(-70) X (12-)15-25(-30) m m ,
y e l l o w intracellular pigment. Stipitipellis a cutis of unbranched, 4.0-8.0 usually annulate, c y l i n d r i c a l to clavate, sometimes fusiform, at base up
m m w i d e hyphae w i t h cylindrical elements, w i t h intracellular very pale to 28 m m hroad, straight, stuffed to narrowly fistulose, white to pale
y e l l o w pigment, partly covered w i t h v e i l composed o f curved hyphae brown, sometimes b r o w n i s h or reddening at base o n handling, above
with short, inflated elements up to 9.5 u,m. annulus smooth to scarcely flocculose, b e l o w annulus fibrillose, floc-
H A B I T A T & D I S T R . - U s u a l l y solitary, saprotrophic, terrestrial o n cose to flocculose or fibrillose-squamulose, sometimes w i t h short
bare, sandy s o i l i n coastal sand dunes, also rarely i n l a n d o n sandy soil. white rhizomorphs. A n n u l u s at 0.60-0.80 o f height o f stipe, 8-13 m m
C o m m o n along the coast, very rare elsewhere i n the Netherlands, w i d e , sometimes r e m a i n i n g attached at m a r g i n o f pileus, ascending,
Sept.-Nov. R a r e but widespread i n Europe. patent to slightly pending, very thick, persistent, white to pale b r o w n ,
not d i s c o l o u r i n g , w i t h striate to sulcate upperside; underside fibrillose
12. Agaricus subfloccosus (J. L a n g e ) Pilat i n A c t a M u s . nat. Prag. 7 B to squamulose, w i t h thickened m a r g i n . Context 12-18 m m thick i n
(1): 6. 1 9 5 1 . - F i g . 13. pileus, firm, white, discolouring q u i c k l y , rarely slowly, greyish pink, or
Psalliota hortensis f. subfloccosa J. L a n g e i n D a n s k bot. A r k . reddish b r o w n to reddish i n pileus and upper part o f stipe when cut (10
4(12): 8. 1926; Psalliota subfloccosa (J. Lange) J. Lange, F l . agar. dan. R 8/3), i n stipe soon fading to b r o w n i s h or b r o w n i s h orange. S m e l l
4: 58. 1939. indistinct to agreeable, aromatic w i t h c h e m i c a l c o m p o u n d , fungoid,
S E L . I C O N . - Breitenb. & K r a n z L , P i l z e S c h w e i z 4: p i . 195. 1995; rarely unpleasant. Taste fungoid w i t h unpleasant c o m p o u n d .
C a p p e l l i , A g a r i c u s : p i . 26, 26bis. 1984; J. L a n g e , F l . agar. dan. 4: M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus
139D. 1939. and stipe); Schaeffer-reaction negative (surface o f pileus).
S E L . D E S C R . & FIGS. - C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 26: Spores 5.5-7.5 X 4.5-6.0 m m , o n average 6.0-6.8 X 5.0-5.2 m m ,
28. 1983; C a p p e l l i , A g a r i c u s : 202. 1984; J. L a n g e , F l . agar. dan. 4 : 58- Q = 1.15-1.45, Q a v = 1.25-1.30, e l l i p s o i d to broadly e l l i p s o i d , w i t h -
59. 1939; F. M 0 l l e r i n F r i e s i a 4 : 11-12. 1950. out g e r m pore. B a s i d i a 23-39 X (6.0-)7.5-10.5 m m , 4-spored.
V E R N . N A M E - Rafelige champignon. L a m e l l a edge w i t h a 55-80 m m broad sterile layer; c h e i l o c y s t i d i a not
i n chains, (26.5-)30-49 X 8.5-15.0(-16.5) m m , c y l i n d r i c a l to nar-
Pileus 65-110(-120) m m , at first convex, later convex w i t h flattened r o w l y clavate, rarely some large elements present u p to 7 0 X 3 0 |xm,
centre to plano-convex, sometimes w i t h depressed centre, w i t h h y a l i n e or w i t h b r o w n i s h content. P i l e i p e l l i s a cutis o f unbranched,
inflexed, later deflexed margin, at first greyish to b r o w n i s h white or i r r e g u l a r l y r a d i a l l y arranged, (2.0-)3.0-5.5(-7.5) m m wide hyphae
pale greyish brown, later usually at centre b r o w n i s h ( M u . 10 Y R 5/3, w i t h c y l i n d r i c a l , rarely s l i g h t l y inflated elements, gradually passing
6/4), towards m a r g i n paler b r o w n (2.5 Y 7/4, 10 Y R 6/3, 7/4, 8/2-3) into p i l e i t r a m a , w i t h pale y e l l o w intracellular pigment. F i b r i l s c o n -
w i t h darker b r o w n fibrils concolorous to centre, rarely remaining pale sisting o f branched, curved, 4.0-5.5 m m w i d e hyphae w i t h c y l i n d r i -
b r o w n i s h white, not d i s c o l o u r i n g or rarely reddish o n handling; sur- cal elements w i t h pale y e l l o w i n t r a c e l l u l a r pigment. S t i p i t i p e l l i s a
face radially fibrillose to fibrillose-flocculose, occasionally w i t h large cutis o f sometimes branched, (2.5-)3.5-5.5 m m w i d e h y p h a e w i t h
fibrillose appressed innate squames, i n centre occasionally areolate- c y l i n d r i c a l elements, w i t h some s l i g h t l y ascending, c y l i n d r i c a l to
rimose, sometimes towards m a r g i n fissurate, near m a r g i n also w i t h clavate, up to 6.5 (Jim w i d e terminal elements, w i t h very pale y e l l o w ,
some w h i t i s h velar f l o c k s ; m a r g i n exceeding lamellae, w i t h broad parietal a n d intracellular pigment.
adhering v e i l remnants. L a m e l l a e , L = (80-)100-165, 1 = 2-3(-7), H A B I T A T & D I S T R . - Solitary or i n s m a l l groups, saprotrophic, ter-
moderately crowded, free, arcuate, u p to 7 m m broad, at first reddish restrial i n Pi'ceo-woods or on roadside verges i n deciduous woods, also
b r o w n , later dark b r o w n (5 Y R 3/2; 7.5 Y R 3/2), w i t h pale b r o w n to in garden, o n h u m u s - r i c h and calcareous soil. Widespread but rare i n
AGARICUS (Nauta) 37
the Netherlands, M a y - J u n e and Sept.-Oct. Widespread but very rare i n w i t h at first inflexed margin, b r o w n i s h , b r o w n or dark greyish brown
Europe. A l s o k n o w n f r o m N o r t h A m e r i c a . to dark b r o w n ( M u . 10 Y R 7/4-6, 7.5-10 Y R 4/4, 7.5 Y R 4/2-5/5, 4/4
5-6/6), not discolouring o n handling, r a d i a l l y squamose w i t h unclear-
Recent research o f N o b l e et a l . (in M y c o l . Res. 99: 1453-1461. 1995) ly delimited, appressed, fibrillose (dark) b r o w n squames on pale brown
and K e r r i g a n et a l . (in M y c o l . R e s . 103: 1515-1523. 1999) revealed to w h i t i s h background (10 Y R 8/4); surface slightly v i s c i d when moist-
two genetically different groups w i t h i n this taxon, one from higher ele- margin slightly exceeding lamellae, white appendiculate-dentate.
vation and one f r o m lower elevation. T h e m o r p h o l o g i c a l differences L a m e l l a e , L = 100-210, 1 = l - 3 ( - 9 ) , crowded, free, straight to sub-
between the t w o groups proved rather s m a l l , and the m o r p h o l o g i c a l ventricose or undate, up to 9.5 m m broad, p i n k i s h b r o w n (5 Y R 3/4,7.5
Y R 5/4-10 Y R 4/4, 7.5 Y R 6/4, 10 Y R 5/3), later b r o w n to dark brown
variability o f the specimens studied f r o m the Netherlands seems to
(7.5 Y R 3/2), w i t h w h i t i s h denticulate edge. Stipe (35-)65-120 X (14-
cover the w h o l e range o f the t w o groups, even w i t h i n one collection.
)19-25(-29) m m , annulate, c y l i n d r i c a l , slightly enlarged at base up to
T h i s taxon requires further study.
30 m m , or slightly ventricose, straight, stuffed, w h i t i s h to p i n k i s h grey,
above annulus smooth to fibrillose, b e l o w annulus w i t h several, dis-
13. Agaricus subperonatus (J. L a n g e ) S i n g , i n L i l l o a 2 2 : 4 3 1 .
tant, often incomplete, (pale) b r o w n i s h girdles o f v e i l or b r o w n -
( ' 1 9 4 9 ' ) 1 9 5 1 . - F i g . 14.
squamose, sometimes w i t h pale b r o w n velar sock, rarely w i t h few thin
Psalliota hortensis f. subperonata J . L a n g e i n D a n s k bot. A r k .
white rhizomorphs. A n n u l u s at 0.70-0.85 o f height o f stipe, 9-15 m m
4(12): 8. 1926; Psalliota subperonata (J. L a n g e ) J . L a n g e , F l . agar,
wide, descending, pending to appressed, triangular i n transverse sec-
dan. 4: 5 9 . 1 9 3 9 . -Agaricus campestris (3 vaporarius Pers., S y n . meth.
tion, thick, persistent, white to brownish, not d i s c o l o u r i n g , w i t h striate
F u n g . : 4 1 8 . 1801; Psalliota vaporaria (Pers.) M 0 l l e r & Schaeff. i n
to smooth upperside; underside fibrillose to squamulose. Context 11-
A n n l s m y c o l . 36: 7 1 . 1938; Agaricus vaporarius (Pers.) C a p p e l l i ,
24 m m thick i n pileus, firm, white to pale b r o w n , d i s c o l o u r i n g (pale)
A g a r i c u s : 149. 1984, n o n A. vaporarius S c h r a n k 1789; Agaricus
p i n k i s h i n pileus and upper part o f stipe w h e n cut, fading to brownish,
pseudovillaticus Rauschert i n N o v a H e d w i g i a 54: 214. 1992.
rarely not discolouring. S m e l l indistinct to faint, disagreeable to fun-
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 15. 1984; D a h n c k e , 1200 P i l z e :
g o i d or pleasant. Taste not k n o w n .
487. 1993; J . L a n g e , F l . agar. dan. 4: M O D . 1939.
S E L . D E S C R . & FIGS. - J . L a n g e , F l . agar. dan. 4: 59. 1939; F. M 0 l l e r M a c r o c h e m i c a l reactions: a m m o n i a pale y e l l o w (context o f stipe);
i n F r i e s i a 4: 42. 1950. K O H 10% negative (context o f stipe and pileus); Schaeffer-reaction
VERN. NAME - Gordelchampignon.
negative (surface o f pileus).
Spores (5.5-)6.0-8.0 X 4.5-6.5 m m , o n average 6.6-7.0 X 5.2-5.6
Pileus (40-)60-125(-135) m m , at first h e m i s p h e r i c a l to convex, m m , Q = 1.10-1.40(4.50), Q a v = 1.20-1.30, broadly ellipsoid, some-
expanding to convex w i t h slightly flattened centre or plano-convex, times ellipsoid, without g e r m pore. B a s i d i a 18-27 X 6.5-9.0 m m , i n
majority 4-spored, usually also a variable amount o f 2-spored present.
L a m e l l a edge w i t h a 40-70 m m broad, sometimes agglutinated, often
sterile layer; cheilocystidia usually i n protruding clusters, 18-36(-45)
X (5.5-)7.5-13 m m , clavate, sometimes w i t h few basal septa, f o r m i n g
short chains o f rectangular elements o f 7.0-12 X 3.0-5.0 m m , hyaline
or rarely w i t h pale b r o w n contents. P i l e i p e l l i s a slightly gelatinized
cutis o f radially, sometimes irregularly arranged, (3.5-)4.5-8.0 m m
w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h b r o w n i s h intracellular
pigment, gradually passing into pileitrama. Squames and fibrils indis-
tinguishable. Stipitipellis a cutis o f sometimes branched, 4.5-6.5 m m
w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h c y l i n d r i c a l to narrowly
clavate terminal elements up to 7.0 (xm wide, w i t h pale y e l l o w intra-
cellular pigment.
H A B I T A T & DISTR. - U s u a l l y gregarious, saprotrophic, terrestrial on,

usually clayey, s o i l or debris i n roadside verges, city parks, woods,
riverdikes, often o n nutrient-rich s o i l . M o d e r a t e l y c o m m o n , but
increasing i n the Netherlands, J u l y - N o v . Widespread but u n c o m m o n i n
Europe. C o s m o p o l i t a n .
T h e differences between Agaricus pseudovillaticus (= Psalliota vapo-

raria) and A. subperonatus according to M 0 l l e r (in F r i e s i a 4: 40-43.
1950) are very subtle, and to be found i n the size, c o l o u r and stature o f
the basidiocarps, and i n the size o f the cystidia. N o such differences
were found i n the collections studied, w h i c h covered the w h o l e range
of character-states mentioned b y M 0 l l e r .
14. Agaricus bohusii M . B o n i n D o c . m y c o l . 13(49): 56. 1981. - F i g .

15.
Agaricus caespitipes H l a v a c e k i n M y k o l . Sb., Praha 59: 66. 1982
(invalid); Agaricus caespitipes H l a v a c e k i n M y k o l . Sb., Praha 74: 5 1 .
1999.
M I S A P P L . - Agaricus elvensis sensu C o o k e , 111. B r i t . F u n g i 4: p i .
F i g . 14. Agaricus subperonatus. (habit X A). 522. 1885.
slightly curved, deeply rooting w i t h pseudorhiza, stuffed to narrowly

fistulose, white to pale b r o w n , reddish y e l l o w to reddish b r o w n o n
handling, fibrillose, above annulus shiny, smooth to striate, b e l o w
annulus fibrillose to flocculose-squamulose, w i t h b r o w n i s h girdles o f
v e i l remnants. A n n u l u s at 0.60-0.85 o f height o f stipe, up to 12 m m
wide, descending, pending, thick, often triangular i n transverse section,
persistent, double, pale b r o w n (7.5 Y R 5/4), w i t h smooth to fibrillose
upperside; underside lanate-fibrillose to fibrillose-squamulose. Context
(11-) 15-22 m m thick i n pileus, f i r m , white to very pale b r o w n , dis-
c o l o u r i n g pale reddish b r o w n to reddish i n upper part o f stipe and
pileus when cut, sometimes not d i s c o l o u r i n g . S m e l l indistinct. Taste
not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% pale y e l l o w to reddish (con-
text o f pileus); Schaeffer-reaction y e l l o w to orange y e l l o w (surface o f
pileus).
Spores 5.5-7.0 X 4.5-6.0 m m , o n average 5.8-6.5 X (4.8-)5.3-5.5
m m , Q = 1.05-1.30, Q a v = 1.10-1.20, broadly ellipsoid to subglobose,
without germ pore. B a s i d i a 22-26 X 7.0-9.0 m m , 4-spored. L a m e l l a
edge w i t h a c. 5 0 m m broad sterile layer; cheilocystidia 15-25(-33) X
6.0-9.0 m m , clavate, colourless or w i t h b r o w n i s h contents, often w i t h
few basal septa, forming short chains o f rectangular, sometimes inflat-
ed elements o f 10-13 X 4.0-5.5 m m . P i l e i p e l l i s a slightly gelatinized
cutis o f unbranched, 4.5-8.0 m m w i d e hyphae w i t h c y l i n d r i c a l ele-
ments, w i t h intracellular y e l l o w i s h b r o w n pigment, gradually passing
into pileitrama. Squames consisting o f curved, up to 6.0 m m w i d e
hyphae w i t h c y l i n d r i c a l elements and recurved, narrowly clavate, up to
8.0 (Jim w i d e terminal elements, w i t h y e l l o w i s h b r o w n intracellular
pigment. S t i p i t i p e l l i s a cutis o f slightly irregularly arranged,
unbranched, 2.5-6.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h
pale y e l l o w intracellular pigment. V e i l consisting o f interwoven, 4.0-
5.0 m m w i d e hyphae w i t h relatively short, 14-30(-45) m m l o n g , c y l i n -
drical elements, w i t h y e l l o w i s h b r o w n , parietal pigment.
H A B I T A T & DISTR. - Fasciculate, i n clusters o f up to 25 basidiocarps,
saprotrophic, terrestrial i n estates or dikes o n clayey soil. Rare, i n river
region. Sept. Rare but widespread i n Europe. A l s o k n o w n from N o r t h
America.
F i g . 15. Agaricus bohusii. (habit X A).

Sect. Sanguinolenti J. Schaeff. & M 0 l l e r
\ K u -. \
S E L . I C O N . - B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: p i . 59. 1977 (as
Pileus not d i s c o l o u r i n g ck reddening or b r o w n i n g ; annulus relatively
A. elvensis sensu C o o k e ) ; C a p p e l l i , A g a r i c u s : p i . 22. 1984.
persistent^ w i t h thickened margin; context not d i s c o l o u r i n g or redden-
S E L . D E S C R . & FIGS. - B o h u s i n A n n l s hist.-nat. M u s . natn. hung.
i n g or b r o w n i n g o n exposure or w h e n damaged.
63: 7 7 . 1971; B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: 5-6, fig. 3. 1977
K O H - r e a c t i o n negative; Schaeffer-reaction negative.
(as A. elvensis sensu C o o k e ) .
Spores e l l i p s o i d to oblong, Q a v > (1.30-)1.40; without germ pore;
VERN. NAME - Spoelvoetchampignon.
l a m e l l a edge sterile; c h e i l o c y s t i d i a clavate to (sub)globose, up to 2 0
u-m wide.
Basidiocarps fasciculate. Pileus (55-)75-160(-220) m m , at first c o n i -
cal, later convex, w i t h at first involute margin, later reflexed and 15. Agaricus sylvaticus Schaeff., F u n g . Bavariae 4: 62.1774:Fr. - F i g .
exceeding lamellae and slightly appendiculate, at centre b r o w n to dark 16.
b r o w n ( M u . 6 Y R 4/3, 7.5 Y R 5/4, 8 Y R 4-5/4, 10 Y R 4-5/4, 5/3), Agaricus sylvaticus Schaeff.:Fr., Syst. m y c o l . 1: 282. 1821 (as A .
lanate-fibrillose w i t h s m a l l squames, towards m a r g i n covered w i t h silvaticus); Psalliota sylvatica (Schaeff.) K u m m . , Fiihr. P i l z k . : 7 3 .
large, fibrillose, pointed, usually recurved b r o w n to reddish b r o w n con- 1871 (as P. silvaticus) - Agaricus haemorrhoidarius S. S c h u l z . i n
centrically arranged squames, concolorous w i t h centre, on pale b r o w n K a l c h b r . , Ic. sel. H y m e n o m y c . H u n g . : 2 9 . 1874. - Agaricus san-
background (6 Y R 5/4). L a m e l l a e , L = 80-175, 1 = 1-3, moderately guinarius P. Karst., R y s s l . , F i n l . S k a n d . H a l f 0 n s Hattsvamp.: 2 3 2 .
crowded, free, segmentiform, slightly undate, up to 9 ( - l 1) m m broad, 1882. - Agaricus vinosobrunneus P.D. O r t o n i n Trans. B r . m y c o l . S o c .
at first pale b r o w n to greyish p i n k (10 Y R 6/4; 7.5 Y R 4/2), later red- 43: 183. 1960; Agaricus sylvaticus var. vinosobrunneus (P.D. Orton)
d i s h b r o w n (5 Y R 5/4), finally dark b r o w n , w i t h concolorous to paler H e i n e m . i n S y d o w i a 30: 7. ( ' 1 9 7 7 ' ) 1978. - Psalliota sylvatica var.
denticulate to serrulate edge. Stipe (75-)135-185 X 13-28(-36) m m , saturata F. M 0 l l e r i n F r i e s i a 4: 38. 1950; Agaricus sylvaticus var. sat-
annulate, fusiform or c y l i n d r i c a l and tapering towards base, straight to urate (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 203. 1952.
AGARICUS (Nauta) 39
appearing q u i c k l y , i n base o f stipe rarely y e l l o w i s h b r o w n . S m e l l like

nuts or indistinct. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus,
context pileus); Schaeffer-reaction negative (all parts).
Spores 4.5-6.0 X (3.0-)3.5-4.0 m m , o n average 5.0-5.5(-5.7) X 3.4-
3.7 m m , Q = 1.30-1.65(-1.70), Q a v = 1.40-1.55, e l l i p s o i d , without
germ pore. B a s i d i a 18-23 X 6.0-7.5 m m , 4-spored. L a m e l l a edge with
a 40-65 m m broad sterile layer; c h e i l o c y s t i d i a i n short chains o f glo-
bose to rounded rectangular elements o f 6.0-9.0(-17.5) X (4.0-)6.5-8.5
m m , w i t h clavate terminal elements o f (12-)14-25.5 X (5.5-)6.0-13
m m , sometimes w i t h b r o w n i s h contents. P i l e i p e l l i s a cutis o f 4.0-8.0
m m wide hyphae w i t h c y l i n d r i c a l elements, w i t h slightly ascending,
cylindrical terminal elements, gradually passing into pileitrama, w i t h
y e l l o w i s h , parietal pigment. Stipitipellis a, sometimes slightly gelati-
nized, cutis o f 4.5-6.0(-8.5) m m w i d e hyphae w i t h c y l i n d r i c a l ele-
ments. Squames consisting o f curved, up to 8.5 m m w i d e hyphae with
some inflated, up to 10.5 m m w i d e elements, and c y l i n d r i c a l terminal
elements, w i t h y e l l o w i s h , parietal pigment.
H A B I T A T & DISTR. - U s u a l l y i n groups, saprotrophic, terrestrial i n
deciduous and coniferous woods, parks, roadside verges o n humus-rich
soil, sometimes i n open dunes. Rather c o m m o n i n the Netherlands,
Sept.-Nov. Widespread and rather c o m m o n i n Europe. C o s m o p o l i t a n .
The identity o f the taxa Agaricus sylvaticus var. pallidus (F. M 0 l l e r ) F.

M 0 l l e r and A. sylvaticus var. fuscosquamatus (F. M 0 l l e r ) F. M 0 l l e r (in
F r i e s i a 4: 2 0 3 . 1952) is unclear. Unfortunately no type or o r i g i n a l
material has been preserved.
F i g . 16. Agaricus sylvaticus. (habit X A).
16. Agaricus impudicus (Rea) Pilat, K l i c urc. H u b hfib. bedl.: 403.

S E L . I C O N . - C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 26: 12. 1983; 1 9 5 1 . - F i g . 17.
C a p p e l l i , A g a r i c u s : p i . 1 7 , 1 7 b i s . 1984; R . P h i l l i p s , Paddest. S c h i m m . : Psalliota impudica R e a i n Trans. B r . m y c o l . S o c . 17: 37. 1932;
160. 1981. Agaricus reai M . B o n i n D o c . m y c o l . 11(44): 2 8 . 1981 (superfluous).
S E L . D E S C R . & F I G S . - C a p p e l l i , A g a r i c u s : 163-164. 1984. F. M 0 l l e r - Psalliota variegata F. M 0 l l e r i n F r i e s i a 4: 30. 1950; Agaricus varie-
i n F r i e s i a 4: 33-38, p i . 14-15. 1950 (resp. as P. haemorrhoidaria and gatus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B (1): 8. 1951, n o n A .
P. silvatica). variegatus Pers. 1801:Fr.; Agaricus variegans F. M 0 l l e r i n Friesia 4:
V E R N . N A M E - Schubbige boschampignon. 203. 1952. - Psalliota variegata var. koelerionensis M . Bon in Doc.
m y c o l . 3: 12. 1972; Agaricus koelerionensis ( M . Bon) M . B o n in Doc.
Pileus 30-100(-145) m m , at first hemispherical, expanding to convex m y c o l . 10(37-38): 9 1 . 1979.
to plano-convex, sometimes irregularly, sometimes w i t h slightly S E L . I C O N . - C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 26: 13. 1983;
depressed centre, w i t h involute m a r g i n , later w i t h m a r g i n s l i g h t l y C a p p e l l i , A g a r i c u s : p i . 18, 18bis. 1984; D a h n c k e , 1200 P i l z e : 4 8 8 .
exceeding lamellae, i n centre reddish b r o w n to dark y e l l o w i s h b r o w n , 1993.
rarely b r o w n ( M u . 5-7.5 Y R 4.5-6/4-5, 10 Y R 4/4, 5-6/3), for the rest
breaking u p or covered w i t h l i k e w i s e coloured, appressed, fibrillose
squamules o n paler b a c k g r o u n d , rarely squarrulose or covered w i t h
thick, wart-like squamules; q u i c k l y d i s c o l o u r i n g red w h e n damaged;
v e i l present as appendiculate white fringe. L a m e l l a e , L = (95-)105-
160, 1 = 1-3, moderately crowded, free, ventricose to undate, up to
8 ( - l l ) m m broad, dark g r e y i s h to p i n k i s h b r o w n (7.5 Y R 3/2, 4/2),
w i t h w h i t i s h , dentate to serrulate edge. Stipe (30-)40-140 X 6-12(-21)
m m , annulate, clavate to c y l i n d r i c a l w i t h bulbous, up to 28 m m broad
at base, straight, fistulose, w h i t i s h , above annulus greyish b r o w n , usu-
a l l y reddening w h e n damaged, sometimes i m m e d i a t e l y r e d , some-
times slightly b r o w n i n g o n h a n d l i n g , above annulus lengthwise s i l k y
striate, b e l o w annulus w h i t i s h to b r o w n i s h flocc(ul)ose, without r h i -
z o m o r p h s . A n n u l u s at 0.60-0.90 o f height o f stipe, 6-10(-13) m m
w i d e , descending, p e n d i n g to s l i g h t l y spreading, floppy, relatively
thin, w h i t e to b r o w n , w i t h thickened, double m a r g i n ; upperside
smooth to finely striate; underside fibrillose, or w i t h s m a l l b r o w n
squamules o r large teeth at edge. C o n t e x t 7 m m t h i c k i n pileus,
w h i t i s h to pale b r o w n , d i s c o l o u r i n g red(dish) to orange-red i n upper
part o f stipe w h e n cut, o c c a s i o n a l l y also i n pileus, especially above
lamellae, sometimes p i n k i s h i n apex o f stipe; c o l o u r o c c a s i o n a l l y dis- F i g . 17. Agaricus impudicus. (habit X 'A).
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 31-33, p i . l a & 13. clavate terminal elements, w i t h parietal, sometimes encrusting, y e l -
1950. l o w i s h pigment. S t i p i t i p e l l i s a s l i g h t l y irregular cutis o f sometimes
VERN. NAME - Panterchampignon. branched, 4.0-6.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, and
slightly ascending, c y l i n d r i c a l , u p to 8.0 m m w i d e terminal elements,
Pileus ( 3 0 - ) 4 0 - l 10(-125) m m , at first hemispherical to conico-convex, w i t h parietal, y e l l o w i s h pigment.
expanding to convex to plano-convex w i t h slightly depressed centre, H A B I T A T & DISTR. - In groups, saprotrophic, terrestrial o n soil or lit-
sometimes w i t h faint u m b o , w i t h margin exceeding lamellae, at centre ter i n woods o r along paths, also i n open dunes, usually o n moist,
usually dark greyish b r o w n to reddish b r o w n ( M u . 5 Y R 3/2-3, 6 Y R humus-rich s o i l . Rather c o m m o n i n southern L i m b u r g and i n the dune
4/4-6. 7.5 Y R 3/2, 4/2-4; 10 Y R 5/4), sometimes y e l l o w i s h b r o w n (10 region, rather rare i n the other parts o f the country, but increasing i n
Y R 6/4), fibrillose to fibrillose-squamulose; otherwise covered w i t h recent times according to C h r i s p i j n (in A r n o l d s et a l . , O v e r z . Paddest.
concentrically arranged dark red-brown (5 Y R 4/3), appressed fibril- Nederland: 55. 1995). Sept.-Dec. Widespread i n Europe, not very rare.
lose squames o n paler, w h i t i s h to p i n k i s h b a c k g r o u n d (7.5 Y R 8/4); A l s o k n o w n f r o m northern A f r i c a , South A m e r i c a and A s i a .
squames getting smaller towards margin; at margin, m a i n l y i n o l d
specimens, sometimes o n l y fibrillose; surface not discolouring o n han- Agaricus koelerionensis, distinguished b y B o n o n account o f the larg-
d l i n g ; v e i l often present as thin dark b r o w n i s h fibrillose squames some er spores, is considered synonymous here, since the spore range o f the
m i l l i m e t r e s f r o m m a r g i n , sometimes also present as white fringe or studied collections also encloses the spore size given for that species.
appendiculate flocks, later disappearing. L a m e l l a e , L = 90-130, 1 = A c c o r d i n g to Rauschert (in N o v a H e d w i g i a 54: 214. 1992)
1-5, c r o w d e d , free, segmentiform to subventricose, up to 9 m m broad, Agaricus brunneola (I. Lange) Pilat (in A c t a M u s . nat. Prag. 7 B (1):
at first p i n k i s h (5 Y R 5-6/4, 8/3; 7.5 Y R 5/4, 7/2-6/4), sometimes 23. 1951) is also synonymous.
slightly reddening w h e n damaged, later light reddish b r o w n to b r o w n -
ish p i n k (2.5 Y R 6/4; 5 Y R 4/3, 6/4) to b r o w n (10 Y R 5/3), with paler, 17. Agaricus langei (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 203. 1952. -
entire to denticulate edge. Stipe (35-)50-110(-125) X (5-)6-12 m m , F i g . 18.
annulate, subclavate to clavate, at base up to 23 m m broad, sometimes Psalliota langei F. M 0 l l e r i n Friesia 4: 28. 1950. - Psalliota medio-
w i t h bulbous base u p to 29 m m broad, straight, narrowly fistulose, fusca F . M 0 l l e r i n F r i e s i a 4: 32. 1950; Agaricus mediofuscus (F.
white, later b r o w n i s h o n handling, above annulus glabrous to striate, M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B (1): 8. 1951; Agaricus langei
b e l o w annulus usually fibrillose-squamulose to flocculose, sometimes var. mediofuscus (F. M 0 l l e r ) Wasser i n U k r . bot. Z h . 35: 516. 1978.
lanate, sometimes with b r o w n i s h squames (4 Y R 4/4) f o r m i n g girdles S E L . ICON.* C a p p e l l i i n B o l l . G r u p p o m i c o l . G . Bres. 26: 24. 1983;
at base, later o c c a s i o n a l l y smooth, occasionally w i t h some short or C a p p e l l i , Agaricus: p i . 2 7 , 2 7 b i s . 1984; Dahncke, 1200 Pilze: 487. 1993.
l o n g , narrow, white, not d i s c o l o u r i n g rhizomorphs. A n n u l u s at 0.65- S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 28-30. 1950.
0.85 o f height o f stipe, (7-)9-15 m m wide, descending, patent, thin to V E R N . N A M E - K n o l l o z e boschampignon.
moderately thick, persistent, w h i t i s h , not d i s c o l o u r i n g , w i t h usually
smooth or velvety, sometimes striate upperside; underside fibrillose- Pileus 35-90 m m , at first c o n i c a l to convex, expanding to convex w i t h
squamulose, velvety to floccose o r cottony, usually at m a r g i n w i t h flattened centre o r plano-convex, sometimes w i t h slight u m b o , w i t h
b r o w n i s h rim o r w i t h radially arranged, slightly thicker, w h i t i s h , y e l - appendiculate margin, at centre b r o w n to reddish o r vinac6ous b r o w n
l o w i s h b r o w n to b r o w n i s h squames. Context 7-10 m m thick i n pileus, ( M u . 5 Y R 3/2, 3-4/3, 4-5/4, 6-7.5 Y R 4/4), fibrillose, for the rest
white to w h i t i s h , not or hardly discolouring when cut, sometimes i n densely covered w i t h relatively broad, appressed, concolorous to cen-
pileus faintly and s l o w l y reddish to reddish brown, sometimes i n older tre o r slightly paler, fibrillose squamules o n paler background, not dis-
specimens faintly b r o w n i n upper half o f stipe (10 Y R 6/3), sometimes c o l o u r i n g o n h a n d l i n g ; v e i l present as white appendiculate fringe.
pale b r o w n to pale y e l l o w i s h b r o w n i n base o f stipe; i n base o f stipe L a m e l l a e , L = 70-115,1 = 1-3, relatively crowded, free, subventricose
faintly y e l l o w i s h b r o w n o n scratching (10 Y R 8/3-4). S m e l l indistinct to segmentiform, 5-8 m m broad, at first pale p i n k i s h brown, later dark-
or unpleasant w h e n cut: faintly like Lepiota cristata (metallic with
sweetish (fruity) component), sometimes slightly fruity with unpleas-
ant c o m p o u n d . Taste like nuts but with unpleasant aftertaste.
M a c r o c h e m i c a l reactions: a m m o n i a negative (surface o f pileus,
context i n base o f stipe); K O H 10% and 4 0 % pale y e l l o w i s h b r o w n
(context i n base o f stipe and pileus; rarely negative) or faintly y e l l o w
(surface o f pileus); Schaeffer-reaction negative (surface o f pileus and
stipe).
Spores (4.5-)5.0-7.0 X 3.5-4.0(-4.5) m m , o n average (5.1-)5.7-6.2
X 3.5-3.9 m m , Q = (1.30-)1.40-1.80(-2.00), Q a v = (1.40-) 1.50-1.70,
e l l i p s o i d to o b l o n g , sometimes amygdaliform, without germ pore.
B a s i d i a 17-23 X 6.5-9.5 m m , 4-spored. L a m e l l a edge w i t h a 65-70 m m
broad sterile layer, rarely not entirely sterile; cheilocystidia usually not
in chains, (12-)15-26 X 9-20(-24) m m , globose, soon collapsed, rarely
w i t h short chains o f rectangular elements o f 5-11 X 3-6 u.m, some-
times agglutinated. P i l e i p e l l i s a slightly irregular cutis o f sometimes
branched, s l i g h t l y interwoven, 4.0-8.5(-13.5) m m w i d e hyphae with
c y l i n d r i c a l , rarely slightly inflated up to 8.5 m m w i d e elements, with
c y l i n d r i c a l terminal elements, gradually passing into pileitrama, w i t h
y e l l o w i s h parietal, sometimes intracellular pigment. Squames consist-
i n g o f sometimes branched, curved, interwoven, (4.5-)5.5-12.5 m m
w i d e hyphae w i t h 21-30(-40) m m long, cylindrical elements and sub- F i g . 18. Agaricus langei. (habit X A).
AGARICUS (Nauta) 41
er p i n k i s h b r o w n (5 Y R 6-7/4,7.5 Y R 6/4), finally b l a c k i s h brown, with

entire, sometimes slightly paler edge. Stipe (30-)40-85(-115) X 7-12(-
20) m m , annulate, c y l i n d r i c a l to clavate, at base up to 16 m m broad,
usually straight, fistulose, w h i t i s h to greyish, reddish b r o w n o n han-
d l i n g , above annulus smooth or finely striate, b e l o w annulus usually
fibrillose to sometimes m i n u t e l y floccose or felted to lanate, at base
occasionally w i t h some larger b r o w n squames, without rhizomorphs.
A n n u l u s at 0.65-0.75 o f height o f stipe, 11-17 m m w i d e , descending,
pending to patent, thin to relatively thick, often at margin thicker, per-
sistent, w h i t i s h or b r o w n i s h , at underside b r o w n i s h , w i t h smooth to
striated upperside; underside felted or squamulose. Context 6-11 m m
thick i n pileus, soft, w h i t i s h , d i s c o l o u r i n g reddish to dark red w h e n cut
i n upper part o f stipe and pileus above lamellae, often immediately, i n
F i g . 19. Agaricus fuscoflbrillosus. (habit X A).
base o f stipe not d i s c o l o u r i n g or d i s c o l o u r i n g orange-red. S m e l l indis-
tinct to acidulous. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% negative (context); Schaeffer- Pileus 40-65(-100) m m , convex to p l a n o - c o n v e x a n d subumbonate,
reaction negative (all surfaces). dark reddish b r o w n at centre ( M u . 2.5 Y R 5/6; 5 Y R 5/4; 7.5 Y R 3-
Spores 7.0-9.0 X 4.0-6.0 m m , o n average (7.6-)7.8-8.3 X 4.4-5.4 4/2), fibrillose at centre, for the rest concolorous fibrillose, w h e n dry-
mm, Q = (1.35-)1.40-1.95(-2.00), Q a v = 1.50-1.85, e l l i p s o i d to ing sometimes fibrillose-squamulose, not d i s c o l o u r i n g o n handling.
oblong, without germ pore. B a s i d i a 23 X 8.5-9.0 m m , usually 4-spored, L a m e l l a e c r o w d e d , free, subventricose to segmentiform, u p to 9.5
occasionally also 2-spored present. L a m e l l a edge w i t h a 35-50 m m m m broad, at first p i n k , later dark b l a c k i s h b r o w n , w i t h paler, entire
broad sterile layer; cheilocystidia not i n chains, 19-46 X 14-20(-26) edge. Stipe 45-105(-150) X 6-14 m m , annulate, c y l i n d r i c a l to sub-
m m , clavate to globose, hyaline or with b r o w n i s h contents. Pileipellis a clavate, at base up to 18 m m broad, straight, fistulose, w h i t i s h , dis-
slightly irregular cutis o f (3.0-)6.5-11.0 m m w i d e hyphae with c y l i n - c o l o u r i n g b r o w n i s h to r e d d i s h b r o w n o n h a n d l i n g , above annulus
drical, rarely slightly inflated elements, w i t h scattered ascending, c y l i n - smooth, b e l o w annulus fibrillose, sometimes fibrillose-squamulose.
drical terminal elements, gradually passing into pileitrama, w i t h A n n u l u s at 0.70-0.75 o f height o f stipe, up to 18 m m w i d e , descend-
intracellular and parietal, pale b r o w n pigment. Stipitipellis a slightly ing, pending, thin, s i m p l e , often evanescent, w h i t e , not d i s c o l o u r i n g ,
gelatinized cutis o f 3.0-8.0 m m w i d e hyphae with sometimes inflated, w i t h smooth upperside; underside f i b r i l l o s e o r smooth. C o n t e x t up to
up to 12 m m w i d e elements, w i t h parietal, pale y e l l o w pigment. 8 m m t h i c k i n p i l e u s , soft, w h i t i s h , d i s c o l o u r i n g s l o w l y and not
H A B I T A T & DISTR. - S o l i t a r y or i n s m a l l groups, saprotrophic, ter- intensely reddish i n upper part o f stipe w h e n cut. S m e l l indistinct.
restrial i n parks, o n dikes planted with trees, i n deciduous woods, o n Taste not k n o w n .
nutrient-rich clayey or sandy soil. Rare i n the Netherlands, A u g . - O c t . M a c r o c h e m i c a l reactions: Schaeffer-reaction negative.
Relatively rare but widespread i n Europe. A l s o recorded from northern Spores 6.5-8.5 X 4.0-5.5 m m , o n average 6.9-7.7 X 4.3-4.5 m m , Q
A f r i c a and the A m e r i c a s . = 1.45-1.90, Q a v = 1.65-1.70, o b l o n g , sometimes e l l i p s o i d , without
germ pore. B a s i d i a 4-spored. L a m e l l a edge w i t h a c. 4 0 |xm broad ster-
T h e c o m p l e x o f species around Agaricus langei, w h i c h is character- ile layer; cheilocystidia not i n chains, 14-26.5 X (9-)12-20.5 m m , glo-
i z e d b y a reddish b r o w n squamulose pileus and spores larger than 7 bose, hyaline. P i l e i p e l l i s a cutis o f (5.5-)7.0-12.5 m m w i d e hyphae
m m , is as yet not fully understood. Several species are distinguished b y with c y l i n d r i c a l elements, gradually passing into pileitrama, w i t h pale
M 0 l l e r and others i n this group, m a i n l y o n account o f differences i n y e l l o w to brownish parietal pigment. Stipitipellis a cutis o f 4.0-7.5 m m
colour, amount o f squames and size o f the pileus, intensity o f the red- wide hyphae with some inflated, up to 10 m m w i d e elements, w i t h
d i s h discoloration o f the context, and the amount o f squames o n the parietal, pale y e l l o w pigment.
stipe. Further study is required, and m o l e c u l a r and experimental H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter-
research m a y make the relationships between the taxa more clear. restrial i n coniferous (Picea) or deciduous woods o n calcareous,
Agaricus babosi B o h u s (in A n n l s hist.-nat. M u s . natn. hung. 8 1 : 3 7 . humus-rich s o i l . Rather rare i n the Netherlands; Sept.-Oct. Rare but
1990) differs f r o m A . langei according to B o h u s i n a longer stipe w h i c h widespread i n E u r o p e . A l s o k n o w n f r o m A s i a , northern A f r i c a and
is dark fibrillose to squamose b e l o w the annulus. N o n e o f the studied South A m e r i c a .
collections fits the description o f B o h u s . Agaricus mediofuscus (F.
M 0 l l e r ) Pilat differs, according to M 0 l l e r (in Friesia 4: 32. 1950), i n the The above description is also based o n a c o l l e c t i o n f r o m B e l g i u m , sup-
pileus w i t h dark b r o w n squames o n the pileus, and the stipe w h i c h i s plemented w i t h the original description b y M 0 l l e r .
dark b r o w n scaly b e l o w the annulus. T h i s taxon seems to represent an Agaricus fuscoflbrillosus i s distinguished f r o m A. langei, apart
extreme variant o f A. langei, a n d a l l transitions c o u l d be found. from the fibrillose instead o f squamulose pileus and the less intense
Therefore it is regarded as conspecific. discoloration o f the context, b y the slightly smaller spores.
18. Agaricus fuscoflbrillosus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 19. Agaricus benesii (Pilat) Pilat i n A c t a M u s . nat. P r a g . 7 B (1): 56.
7 B (1): 19. 1 9 5 1 . - F i g . 19. 1 9 5 1 . - F i g . 20.
Psalliota fuscoftbrillosa F. M 0 l l e r i n F r i e s i a 4: 2 7 . 1950. Psalliota benesii Pilat i n M y k o l o g i a 2: 4 9 . 1925 (as P. benesi). -
S E L . I C O N . - C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 26: 20. 1983; Agaricus albosanguineus H o t s o n & Stuntz i n M y c o l o g i a 30: 2 1 7 .
C a p p e l l i , A g a r i c u s : p i . 2 1 . 1984; F. M 0 l l e r i n F r i e s i a 4: p i 2b. 1950. 1938. - Psalliota squamulifera F. M 0 l l e r i n F r i e s i a 4: 2 1 . 1950;
S E L . D E S C R . & F I G S . - B o h u s i n A n n l s hist.-nat. M u s . natn. hung. Agaricus squamuliferus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag. 7 B
61: 152-153. 1969; F. M 0 l l e r i n F r i e s i a 4: 27-28. 1950. (1): 7. 1951. - Agaricus caroli Pilat i n A c t a M u s . nat. P r a g . 7 B (1):
V E R N . N A M E - Fijnvezelige boschampignon. 130. 1951; Agaricus squamuliferus var. caroli (Pilat) Pilat i n Pilat &
U s a k , Nase H o u b y 2: 141. 1959 (invalid).
hyphae w i t h c y l i n d r i c a l , sometimes up to 14 m m inflated elements,

w i t h sometimes s l i g h t l y ascending, c y l i n d r i c a l terminal elements,
gradually passing into pileitrama, w i t h y e l l o w i s h parietal pigment.
Stipitipellis a slightly gelatinized cutis o f 4.0-5.5 m m w i d e hyphae
w i t h c y l i n d r i c a l elements w i t h y e l l o w i s h , parietal pigment.
H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic, terres-
trial i n roadside verges under deciduous trees, i n open vegetation under
trees o n nutrient-rich or calcareous soil. Very rare i n the Netherlands
(Maarsseveense plassen; Werkhoven, Laan van Beverweerd;
Amsterdam, Z w a a n s w i j k ) , Sept.-Oct. Rare but widespread i n Europe.
A l s o recorded from A s i a , northern A f r i c a and N o r t h A m e r i c a .
T h e above description is also based o n a c o l l e c t i o n from Denmark.

T h e colour o f the pileus varies from white" to pale brown, further-
more the amount o f squamules on the pileus is variable w i t h i n one c o l -
lection. Since this forms a continuous range Agaricus squamuliferus is
regarded as a s y n o n y m of A. benesii. Agaricus caroli Pilat, with a b r o w n
pileus already w h e n young, is not distinguished as a separate taxon.
F i g . 20. Agaricus benesii. (habit X A). 20. Agaricus depauperatus (F. M 0 l l e r ) Pilat i n A c t a M u s . nat. Prag.
7 B ( 1 ) : 18. 1 9 5 1 . - F i g . 2 1 .
Psalliota depauperata F. M 0 l l e r i n F r i e s i a 4: 24. 1950. - Agaricus
S E L . I C O N . - C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 26: 17 & 25 deylii Pilat i n A c t a M u s . nat. Prag. 7 B (1): 131. 1951.
(as A. squamuliferus). 1983; C a p p e l l i , A g a r i c u s : p i . 20, 24 & 25 (asL4. S E L . I C O N . - B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: p i . 58. 1977 (as
squamulifer). 1984; Wasser i n F u n g . rar. Ic. c o l . 10: p i . 75. 1979 (asM- A. deylii); C a p p e l l i , A g a r i c u s : p i . 32. 1984; F. M 0 l l e r i n Friesia 4: p i .
V
squamuliferus). 3a. 1950.
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 21-24. 1950 (as P. S E L . D E S C R . & FIGS. - B o h u s & B a b o s i n F u n g . rar. Ic. c o l . 8: 3-5.
squamulifera); P i l a t i n A c t a M u s . nat. Prag. 7 B (1): 56-57. 1 9 5 1 ; 1977 (as A. deylii); F . M 0 l l e r i n F r i e s i a 4 : 24-25. 1950; Pilat i n A c t a
Wasser i n F u n g . rar. Ic. c o l . 10: 3-7. 1979 (as A. squamuliferus). M u s . nat. Prag. 7 B (1): 53-56, figs. 21-23. 1951.
VERN. NAME - Tandringchampignon. V E R N . N A M E - Vale champignon.
P i l e u s 4 5 - 1 0 5 m m , at first h e m i s p h e r i c a l to convex, e x p a n d i n g to Pileus 50-60 m m , convex, w i t h inflexed margin, slightly exceeding

convex o r p l a n o - c o n v e x , often w i t h depressed centre, w i t h m a r g i n lamellae, white to pale brownish to greyish b r o w n w i t h pinkish hue or
s l i g h t l y e x c e e d i n g l a m e l l a e , w h i t i s h to pale g r e y i s h b r o w n o r y e l - greyish white, not d i s c o l o u r i n g o n handling, sometimes finally pale
l o w i s h b r o w n ( M u . 7.5 Y R 6/4, 10 Y R 8/1.5), reddening o n h a n d l i n g , brown, entirely fibrillose-squamulose w i t h small, appressed, concolor-
intensely r e d w h e n damaged, at m a r g i n often covered w i t h s m a l l , ous squamules o n paler background or w i t h faint appressed fibrillose
w h i t i s h to pale g r e y i s h b r o w n squamules, sometimes entirely squa- squames ( M u . 10 Y R 6-7/2); v e i l often present as appendiculate fringe,
m u l o s e , later often s m o o t h . L a m e l l a e , L = 130-170, 1 = l - 3 ( - 9 ) , later disappearing. L a m e l l a e , L = 90-105,1 = 1-5, crowded, free, sub-
c r o w d e d , free, segmentiform to ventricose, up to 8 m m broad, at first ventricose, up to 6 m m broad, at first greyish, finally dark brown, w i t h
g r e y i s h b r o w n to p i n k i s h b r o w n (2.5 Y R 5/4), later dark b l a c k i s h paler, denticulate edge. Stipe 40-70 X 10-14 m m , annulate, cylindrical,
b r o w n , w i t h w h i t i s h denticulate edge, r e d d e n i n g w h e n damaged. straight, stuffed or narrowly fistulose, white, later w i t h faint b r o w n to
S t i p e 6 0 - 1 3 0 X 10-27 m m , annulate, c y l i n d r i c a l to subbulbous, at reddish b r o w n tinge, sometimes slightly reddening o n handling, above
base u p to 30 m m broad, straight, fistulose, white, reddening w h e n
damaged, above annulus smooth, b e l o w annulus f i b r i l l o s e - s q u a m u -
lose, sometimes w i t h few pale g r e y - b r o w n squames (7.5 Y R 7/4).
A n n u l u s at c. 0.80 o f height o f stipe, 9-10 m m w i d e , descending,
p e n d i n g to s l i g h t l y spreading, thick, persistent, white, w i t h smooth
upperside; underside w i t h large, p i n k i s h b r o w n , recurved, teeth-like
squames, r a d i a l l y arranged near m a r g i n as a c o g w h e e l . Context 7-17
m m t h i c k i n p i l e u s , hard, w h i t i s h , d i s c o l o u r i n g q u i c k l y b r i g h t r e d
w h e n cut i n p i l e u s and upper part o f stipe, i n base o f stipe not chang-
i n g or d i s c o l o u r i n g y e l l o w i s h w h i t e . S m e l l i n d i s t i n c t to sweetish
(perfume). Taste not k n o w n .
M a c r o c h e m i c a l reactions: Schaeffer-reaction negative (all parts o f
basidiocarp).
= 1.30-1.65(-1.85), Q a v = 1.45-1.55, e l l i p s o i d , sometimes o b l o n g ,
without germ pore. B a s i d i a 21-26 X 6.5-7.5 m m , 4-spored. L a m e l l a
edge w i t h a 60-75 m m broad sterile layer; cheilocystidia i n short chains
of 2-3 rectangular elements o f 9.0-12.0 X 5-6.5 m m , w i t h clavate ter-
m i n a l elements o f 15.5-25.5(-28.5) X 7.5-12.5 m m , often w i t h b r o w n -
ish contents. P i l e i p e l l i s a slightly irregular cutis o f 3.5-6.5 m m w i d e F i g . 2 1 . Agaricus depauperatus. (habit X A).
AGARICUS (Nauta) 43
annulus smooth and later brownish, below annulus somewhat appressed Sect. Arvenses Konr. & M .
fibrillose, at base w i t h some faint girdles o f white squamules, without
rhizomorphs. A n n u l u s at c. 0.75 o f height o f stipe, 6-8 m m wide, Pileus y e l l o w i s h o n handling, especially o n d r y i n g ; annulus persistent,
descending, pending, relatively thick, persistent, white, sometimes with broad, up to 35 m m w i d e , w i t h remnants o f universal v e i l as t h i c k
pale b r o w n tinge, not discolouring, w i t h smooth upperside; underside flocks or warts at underside; context not d i s c o l o u r i n g o r d i s c o l o u r i n g
fibrillose, near edge double and w i t h s m a l l b r o w n i s h warts or larger slightly y e l l o w or orange to p i n k i s h o n exposure or w h e n damaged.
b r o w n i s h squames. Context 7 m m thick i n pileus, firm, white, dis- K O H - r e a c t i o n o n context usually positive and y e l l o w ; Schaeffer-
colouring s l o w l y b l i g h t but not intense red to vinaceous red when cut reaction o n pileus orange, rarely doubtful.
i n upper half o f stipe; colour q u i c k l y disappearing; i n base o f stipe y e l - Spores e l l i p s o i d to o b l o n g ; average spore length > 6.0 m m ; Q a v =
l o w i s h white, not discolouring. S m e l l indistinct. Taste not k n o w n . 1.35-1.60; without germ pore; l a m e l l a edge sterile; cheilocystidia cate-
M a c r o c h e m i c a l reactions: K O H 10% pale b r o w n (surface o f base nate, forming sterile layer around l a m e l l a edge (absent i n some taxa
o f stipe); Schaeffer-reaction negative (surface). not occurring i n the Netherlands).
= 1.20-2.00, Q a v = 1.30-1.75, e l l i p s o i d to o b l o n g , without germ pore. 21. Agaricus arvensis Schaeff., F u n g . Bavariae 4: 73. 1774. - F i g . 22.
B a s i d i a 18-25 X 7.0-8.0 m m , 4-spored, sometimes 2-spored also pres- Psalliota arvensis (Schaeff.) K u m m . , Fiihr. P i l z k . : 7 4 . 1871. -
ent. L a m e l l a edge w i t h a c. 35 p,m broad sterile layer; cheilocystidia Psalliota fissurata F. M 0 l l e r i n F r i e s i a 4: 165. 1952; Agaricus fissura-
usually not catenate, sometimes i n short chains o f 1-2 small, rectan- tus (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204. 1952. - Psalliota leucotricha
gular elements o f 5.5-7.0 X 4.0-6.0 u.m, w i t h clavate to globose ter- F. M 0 l l e r i n F r i e s i a 4: 159. 1952; Agaricus leucotrichus (F. M 0 l l e r ) F.
m i n a l elements o f 13.5-28 X 9-17 m m . P i l e i p e l l i s a cutis o f rarely M 0 l l e r i n F r i e s i a 4: 204. 1952.
branched, 4.5-8.0 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes up to E X C L . - Agaricus arvensis sensu C o o k e , 111. B r i t . F u n g i 4: p i . 523.
12 m m inflated elements, gradually passing into pileitrama, w i t h pale 1885 ( = Agaricus osecanus).
yellow, parietal pigment. Stipitipellis a cutis o f rarely branched, some- S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 33. 1984; Diihncke, 1200 P i l z e :
times curved, (2.5-)4.0-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, 498. 1993; J . L a n g e , F l . agar. dan. 4 : p i . 1 3 8 A . 1939; R . P h i l l i p s ,
w i t h scattered, up to 12.5 m m inflated elements, and sparse, ascending Paddest. S c h i m m . : 166. 1981.
c y l i n d r i c a l to clavate, u p to 9.0 m m w i d e terminal elements, w i t h y e l - S E L . D E S C R . & F I G S . - F . M 0 l l e r i n F r i e s i a 4 : 1 6 1 - 1 6 5 , f i g . 26, p i . 30;
l o w i s h intracellular and parietal pigment. 165-167, p i . 19b (as P. fissurata); 159-161, p i . 18a (as P. leucotricha).
H A B I T A T & D I S T R . - U s u a l l y solitary, sometimes i n s m a l l groups, 1952; Wasser, Tr. A g a r i c e a e Soviet U n i o n : 85-86. 1989.
saprotrophic, terrestrial i n parks, deciduous woods, i n woods o f Picea, V E R N . N A M E - Gewone anijschampignon.
or i n open dune grassland, usually o n sandy h u m u s - r i c h s o i l . R a r e but
probably often w r o n g l y identified i n the Netherlands; M a y , A u g . - N o v . Pileus (30-)60-105(-150) m m , at first h e m i s p h e r i c a l to convex,
R a r e but widespread i n E u r o p e . A l s o k n o w n from northern A f r i c a . expanding to plano-convex o r applanate, w i t h at first involute, later
deflexed, sometimes recurved m a r g i n , exceeding lamellae, at first
T h e above description is also based o n collections from southern white to w h i t i s h , soon y e l l o w i s h to b r o w n i s h i n centre ( M u . 10 Y R
England. 8/3-4, 8/6), d i s c o l o u r i n g y e l l o w o n h a n d l i n g , sometimes entirely y e l -
A c c o r d i n g to the type descriptions, Agaricus depauperatus and A. low o r y e l l o w i s h b r o w n (2.5 Y 8/4, 8/6, 10 Y R 8/4), m i n u t e l y f i b r i l -
deylii differ i n the c o l o u r o f the pileus, and the size o f the spores. lose at first, later smooth, often w i t h appressed concolorous squames
A c c o r d i n g to A . M . B r a n d ( L e i d e n ; pers. comment), however, w h o near margin, o c c a s i o n a l l y entirely squamulose o r r a d i a l l y fibrillose,
studied authentic material, the spore size is almost the same for both later sometimes radially fissurate; v e i l present w h e n y o u n g as appen-
species. W i t h i n the collections studied the pileus c o l o u r varied from diculate white floccose remnants. L a m e l l a e , L = 95-185,1 = l-3(-7),
completely white to white w i t h pale b r o w n fibrils, reason enough to very crowded, free, subventricose, up to 14 m m broad, at first w h i t i s h
consider the species conspecific. A l s o , the spore size w i t h i n a collec- to pale p i n k (7.5 Y R 8/4), later greyish p i n k to reddish b r o w n or b r o w n
tion shows a considerable amount o f variation. (5 Y R 4/3, 6/2-3, 10 Y R 4-5/3) to dark b r o w n , w i t h u s u a l l y paler,
F i g . 22. Agaricus arvensis. (habit X A).

entire, sometimes serrulate, edge. Stipe 55-120 X (8-)10-22 m m , H A B I T A T & D I S T R . - Solitary or i n groups, saprotrophic, terrestrial
annulate, c y l i n d r i c a l to attenuated towards apex, o c c a s i o n a l l y sub- i n meadows, roadside verges, c i t y parks, woods o n various soils.
clavate to subbulbous and then at base up to 34 m m broad, often bent C o m m o n and widespread i n the Netherlands. June-Nov. C o m m o n and
or s l i g h t l y flexuous, n a r r o w l y fistulose, white, y e l l o w i n g b e l o w annu- widespread i n E u r o p e . C o s m o p o l i t a n .
lus o n h a n d l i n g , above annulus glabrous, smooth, often shiny, b e l o w
annulus smooth to fibrillose or, especially towards base, velutinous to Agaricus fissuratus is considered an e c o l o g i c a l m o d i f i c a t i o n of, and
squamulose. A n n u l u s at 0.70-0.80 o f height o f stipe, (7-) 13-25 m m therefore conspecific w i t h , A. arvensis. Several intermediates were
w i d e , descending, p e n d i n g to sometimes spreading, thick, cream- found between A. leucotrichus and A. arvensis, w h i c h is therefore also
coloured, w i t h smooth to fibrillose upperside; underside when y o u n g considered conspecific. F o r an extensive discussion see N a u t a (in
w i t h t h i c k stellate-wise arranged squames (as cogwheel), later w i t h Persoonia 17: 458-459. ( ' 2 0 0 0 ' ) 2001).
pale reddish y e l l o w to b r o w n i s h floccose patches or warts. Context 6- Agaricus arvensis var. macrolepis Pilat & P o u z . (in A c t a M u s . nat.
15 m m t h i c k i n pileus, soft, white, sometimes y e l l o w i n g i n stipe w h e n Prag. 7 B (1): 134. 1951; as Agaricus macrolepsis) differs f r o m var.
cut. S m e l l l i k e anise, but sometimes faintly, or l i k e (sweet) almonds. arvensis i n the coarsely squamose surface o f the pileus, abruptly b u l -
Taste not k n o w n . bous stipe, w h i c h is floccose-squamose b e l o w the annulus, and the
M a c r o c h e m i c a l reactions: K O H 10% strongly c h r o m e - y e l l o w (sur- larger spores. T h i s taxon has not yet been found i n the Netherlands.
face o f pileus and stipe, upperside annulus) or negative to faintly y e l - T h e t a x o n o m i c a l status o f this variety needs reconsideration.
l o w (context); Schaeffer-reaction orange-red (surface o f pileus) to
faintly orange (surface o f stipe). 22. Agaricus augustus Fr., E p i c r i s i s : 212. 1838. - F i g . 23.
Spores 7.0-9.0(-9.5) X 5.0-6.0(-6.5) m m , o n average 7.9-8.6 X Psalliota augusta (Fr.) Q u e l , i n M e m . S o c . E m u l . M o n t b e l i a r d ser.
5.3-5.9 m m , Q = 1.30-1.65, Q a v = 1.40-1.55, sometimes aberrant II, 5: 255. 1872 ( C h a m p . Jura Vosges 1); Pratella augusta (Fr.) G i l l e t ,
spores present, e l l i p s o i d , w i t h o u t germ pore. B a s i d i a 15-20 X 9.0-11 H y m e n o m y c e t e s : 561. 1878. - Agaricus perrarus S. Schulz. i n Verh.
m m , 4-spored, rarely also 2-spored b a s i d i a present. L a m e l l a edge z o o l . bot. G e s . W i e n 29: 4 9 3 . 1880; Psalliota perrara (S. S c h u l z . )
w i t h a 6 0 - 1 1 0 m m broad, sterile layer; c h e i l o c y s t i d i a i n chains o f g l o - Bres., F u n g i trident. 1: 82. 1887; Agaricus augustus var. perrarus (S.
bose elements o f ( 8 . 5 - ) l l - 2 0 X 6.5-18 m m , w i t h clavate to some- Schulz.) B o n & C a p p e l l i i n D o c . m y c o l . 13(52): 16. 1983. -Agaricus
times l a g e n i f o r m or u t r i f o r m terminal elements o f (14-)20-36(-42), X peronatus M a s s . , B r i t . Fungus F l , 1:415. 1892, non Psalliota perona-
8.5-12(-18) m m , w i t h 4.0-9.0 m m w i d e obtuse apex, h y a l i n e or w i t h ta R i c h o n & R o z e 1885.
pale b r o w n contents. P i l e i p e l l i s a cutis o f sometimes s l i g h t l y gelati- M I S A P P L . w Psalliota subrufescens sensu J . Lange, F l . agar. dan. 4:
n i z e d , 4.0-8.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, gradually 55. 1939.
p a s s i n g into p i l e i t r a m a , w i t h pale y e l l o w , i n t r a c e l l u l a r pigment. S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 49, 50 (as A. augustus var. per-
S t i p i t i p e l l i s a cutis o f sometimes g e l a t i n i z e d , 4.0-8.0 m m w i d e rarus). 1984; K o n r . & M . , Ic. sel. F u n g . 1: p i . 27. 1925; J . Lange, F l .
hyphae w i t h often s l i g h t l y inflated elements, w i t h very pale y e l l o w agar. dan. 4: p i . 1 3 5 B , p i . 136B (as P. subrufescens). 1939; R . P h i l l i p s ,
intracellular pigment. Paddest. S c h i m m . : 164. 1981.
F i g . 23. Agaricus augustus. (habit X A).

AGARICUS (Nauta) 45
S E L . D E S C R . & F I G S . - H e i n e m . i n B u l l , trimest. S o c . m y c o l . Fr. 81: very pale-coloured variant w h i c h was p r o v i s i o n a l l y named by M o s e r

396-397. 1965; F. M 0 l l e r i n Friesia 4: 138-145, fig. 20, p i . 2 2 - 2 5 . 1 9 5 2 ; var. albus n o m . prov. ( R o h r l i n g e Blatterpilze, 3. A u f l . : 191. 1967) is
Wasser, Tr. A g a r i c e a e Soviet U n i o n : 61-63. 1989. probably not o f t a x o n o m i c a l importance.
VERN. NAME - Reuzenchampignon.
23. Agaricus osecanus Pilat i n A c t a M u s . nat. Prag. 7 B (1): 133. 1951.
Pileus (70-)90-220 m m , at first h e m i s p h e r i c a l , soon expanding to Psalliota nivescens F. M 0 l l e r i n F r i e s i a 4: 155. 1952; Agaricus
plano-convex, often w i t h depressed centre, rarely subumbonate, w i t h nivescens (F. M 0 l l e r ) F . M 0 l l e r i n F r i e s i a 4: 204. 1952. - Psalliota
straight margin, densely covered w i t h pointed, mostly appressed, ochre nivescens var. parkensis F. M 0 l l e r i n F r i e s i a 4 : 158. 1952; Agaricus
to reddish b r o w n fibrillose squames ( M u . 5 Y R 4/4, 7.5 Y R 4-5/6, 4- nivescens var. parkensis (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204. 1952.
5 / 4 , 1 0 Y R 8/6) o n a y e l l o w i s h white to pale ochraceous y e l l o w or light M I S A P P L . - Agaricus arvensis sensu C o o k e , 111. B r i t . F u n g i 4: p i .
b r o w n (7.5 Y R 7/6, 10 Y R 8/6) background, at centre densely 523. 1885.
squamose and ochraceous y e l l o w to dark reddish b r o w n (5 Y R 4/4, 7.5 S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 3 5 . 1984 (as A. nivescens);
Y R 3/2-3, 10 Y R 7/6), more intensely y e l l o w on handling; v e i l present C o o k e , 111. B r i t . F u n g i 4: p i . 523. 1885 (as A. arvensis); Essette, P s a l -
as w h i t i s h appendiculate remnants at margin. L a m e l l a e , L = 140-150, liotes: p i . 35. 1964 (as P. nivescens).
1 = 0-1, crowded, free, subventricose, up to 9 m m broad, at first pale S E L . D E S C R . & FIGS. - F. M 0 l l e r i n Friesia 4: 155-158, F i g . 24, p i .
greyish cream to flesh-coloured (2.5 Y 8/2, 7.5 Y R 6/4), finally dark 18b, 19. 1952 (as P. nivescens); Pilat i n A c t a M u s . nat. Prag. 7 B (1):
b r o w n , w i t h entire, w h i t i s h to rarely pale sulphur-yellow edge. Stipe 81-83, Figs. 41-42. 1951; Wasser, Tr. A g a r i c e a e Soviet U n i o n : 79-80
90-140(-175) X (12-)18-26 m m , annulate, subclavate to clavate, at (as A. nivescens), 86-87. 1989.
base up to 35 m m broad, straight to curved, fistulose, w h i t i s h to pale V E R N . N A M E - Sneeuwwitte anijschampignon.
ochraceous y e l l o w - b r o w n , above annulus glabrous and smooth, below
annulus usually densely squamose to floccose w i t h white to y e l l o w K E Y TO THE VARIETIES
r e c u r v i n g squames; towards base squames sometimes w i t h reddish 1. Stipe below annulus smooth to l o n g i t u d i n a l l y striate, o n l y near base
b r o w n tips; often deeply rooting. A n n u l u s at 0.80-0.85 o f height o f with scattered squames 23a. var. osecanus
stipe, 17-30(-41) m m w i d e , descending, pending, thin, easily torn, 1. Stipe below annulus floccose-squarrose . . . . 23b. var. squarrosipes
white, often d i s c o l o u r i n g y e l l o w o n h a n d l i n g , w i t h upperside finely
subfloccose to striate; underside w i t h t h i c k ochraceous to b r o w n i s h 23a. var. osecanus. - F i g . 2 4 a
floccose warts. Context 10-17 m m thick i n pileus, w h i t i s h , discolour- Pileus (40-)80-175(-190) m m , at first obtusely b r o a d l y c o n i c a l to
ing y e l l o w or light orange-brown w h e n cut o r bruised, i n base o f stipe hemispherical, expanding to convex o r plano-convex, sometimes
sometimes p i n k i s h b r o w n . S m e l l v a r y i n g from strongly nutty to faint- irregularly shaped, w i t h inflexed, rarely reflexed margin, white, slight-
ly o r strongly like anise, sometimes l i k e almonds, later unpleasantly ly and s l o w l y y e l l o w i n g o n h a n d l i n g , finally y e l l o w i s h ( K . & W . 2 A 4 )
sweetish. Taste not k n o w n . to light y e l l o w - b r o w n ( M u . 10 Y R 6/4), at centre y e l l o w (10 Y R 8/6),
M a c r o c h e m i c a l reactions: K O H 10% y e l l o w (surface o f pileus; also usually entirely m i n u t e l y fibrillose-squamulose at first, later usually
on context a c c o r d i n g to M 0 l l e r & Schaeff. i n A n n l s m y c o l . 36: 79. smooth and s i l k y at centre, floccose-squamose at m a r g i n , sometimes
1938), b r o w n i s h orange (context i n base o f stipe); Schaeffer-reaction remaining entirely but m i n u t e l y fibrillose-squamulose, rarely fissur-
usually orange-red, rarely negative (surface o f pileus). ate; v e i l usually present w h e n y o u n g as appendiculate, t h i c k denticles,
Spores 7.5-9.5(-10.0) X 5.5-6.51-7.0) m m , o n average 8.2-8.8(-9.0) later disappearing or rarely present as short fringe. L a m e l l a e , L = 9 5 -
X 5.5-5.8 urn, Q = (1.25-)1.30-1.65(-1.70), Q a v = 1.40-1.55, ellipsoid 120(-190), 1 = l - 4 ( - 7 ) , c r o w d e d , rarely partly anastomosing, free,
to sometimes amygdaliform, without germ pore. B a s i d i a 17-25 X 6.5- subventricose, up to 13 m m broad, at first very pale b r o w n (10 Y R
8.5 m m , 4-spored. L a m e l l a edge w i t h a 50-70 m m broad, sterile layer; 8/2-7/3), later p i n k i s h grey to reddish b r o w n (7.5 Y R 7/4, 5/2; 5 Y R
cheilocystidia clustered and protuberant, i n chains o f mostly subglobose 5/3, 8/4), finally dark b r o w n , w i t h entire, sometimes s l i g h t l y paler
elements o f ( l l - ) 1 5 - 1 7 ( - 2 2 ) X 5 . 5 - 1 1 ( 1 6 ) m m , w i t h subglobose to edge. Stipe (65-)80-120(-155) X (9-)15-28 m m , annulate, c y l i n d r i c a l
often irregularly broadly cylindrical, utriform to almost lecythiform ter- to gradually enlarged d o w n w a r d s , sometimes tapering downwards,
m i n a l elements o f (10-)15-22(-35) X (5.5-)6.5-10(-16)mm, with at first rarely subbulbous and then at base 21-24 m m w i d e , straight to curved,
hyaline, later y e l l o w to brownish contents. Pileipellis a slightly gelati- solid to narrowly fistulose, white, y e l l o w to pale b r o w n near baseband
nized cutis o f cylindrical, 4.5-6.5 m m w i d e hyphae, w i t h pale y e l l o w on h a n d l i n g , above annulus glabrous, smooth, s i l k y striate to shiny,
intracellular pigment, gradually passing into pileitrama. Squames con- b e l o w annulus at first often m i n u t e l y lanate to squamulose, later
sisting o f interwoven, sometimes agglutinated, ( 3 . 5 - ) 4 . 5 - l l m m wide smooth to finely l o n g i t u d i n a l l y striate o r fibrillose, near base o c c a -
hyphae w i t h 25-47 m m long, slightly inflated elements and ascending, sionally fibrillose-squamulose; squames sometimes b r o w n and
mostly clavate, up to 15 m m wide, terminal elements, w i t h intracellu- arranged i n girdles; sometimes w i t h rhizomorphs. A n n u l u s at (0.50-
lar y e l l o w i s h b r o w n pigment. Stipitipellis a cutis o f sometimes )0.70-0.80 o f height o f stipe, 18-33 m m w i d e , descending, pending to
branched, 5.5-12 m m wide hyphae o f cylindrical, slightly inflated ele- slightly spreading, thick, persistent, white, sometimes d i s c o l o u r i n g
ments. Squames o n stipe consisting o f interwoven, often curved, 5.5-11 y e l l o w o n h a n d l i n g , w i t h smooth upperside; underside w i t h thick
m m w i d e hyphae w i t h u p to 4 0 |xm l o n g inflated elements, w i t h often creamy buff to reddish y e l l o w squames, u s u a l l y arranged as a c o g -
clavate to subglobose, up to 17 u,m w i d e terminal elements. w h e e l along m a r g i n . C o n t e x t (6.5-) 10-16 m m t h i c k i n pileus, soft,
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic, terrestrial, white, hardly d i s c o l o u r i n g w h e n cut, rarely w i t h s l i g h t l y y e l l o w i s h
mostly i n deciduous woods or i n roadside verges under deciduous patches, i n base o f stipe o c c a s i o n a l l y s l o w l y ochraceous-brownish to
trees, also i n coniferous woods, o n sandy to clayey humus r i c h soil; sordid p i n k i s h b r o w n . S m e l l nutty, sometimes l i k e almonds, at first
rather c o m m o n i n the Netherlands, rare i n the Pleistocene regions. pleasant, o f cut basidiocarps more intense, f i n a l l y unpleasant. Taste
A u g . - S e p t . W i d e s p r e a d i n Europe, cosmopolitan. faintly to strongly nutty, pleasant.
M a c r o c h e m i c a l reactions: K O H 10% y e l l o w (surface o f pileus and
Agaricus perrarus S. S c h u l z . i s considered conspecific. F o r an exten- stipe), y e l l o w to faintly y e l l o w (context); Schaeffer-reaction orange-
sive discussion see N a u t a (in Persoonia 17: 459. ( ' 2 0 0 0 ' ) 2001). T h e red (surface o f pileus).
elements, w i t h very pale intracellular pigment, with scattered slightly

ascending, cylindrical to subclavate terminal elements; gradually pass-
ing into pileitrama. Stipitipellis a cutis o f sometimes irregular, 5.5-14.5
mm w i d e hyphae w i t h often inflated elements, w i t h pale intracellular
y e l l o w pigment.
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic, terrestrial
in grasslands, lawns, city parks, along avenues and, more rarely, i n
woods, o n relatively nutrient-rich soil, often o n clay. Rather rare, but
may have been confused i n the past with Agaricus arvensis. Probably
widespread i n the Netherlands. Jun.-Nov. Rather c o m m o n and w i d e -
spread i n Europe, also i n the Mediterranean area. A l s o k n o w n from A s i a .
Agaricus nivescens and A. osecanus are considered conspecific, for an

extensive discussion see N a u t a (in Persoonia 17: 460. ('2000') 2001).
23b. var. squarrosipes ( B o n & C o l l i n ) N a u t a i n Persoonia 17: 460.

('2000') 2 0 0 1 . - F i g . 24b.
Agaricus nivescens var. squarrosipes B o n & Collin in Doc. mycol.
17(67): 11. 1987
S E L . D E S C R . & FIGS. - C o l l i n i n B u l l , trimest. Soc. m y c o l . Fr. 109,
A t l a s : p i . 272. 1993 (as A. nivescens var. squarrosipes); Grilli in M i c o l .
Veget. medit. 9: 3-8. 1994 (as A . nivescens var. squarrosipes).
C H A R A C T E R I S T I C S - Differing from the typical variety i n the covering

of the stipe w h i c h is clearly, wtrite/-squarrose b e l o w the annulus.
M i c r o s c o p i c a l characters as i n typical variety.
H A B I T A T & DISTR. - Solitary, saprotrophic, terrestrial, occurring i n
the same k i n d o f habitats as var. osecanus, o n relatively nutrient-rich
soil. Very rare i n the Netherlands (Bergen; Utrecht; Willemstad); M a y ,
F i g . 24a. Agaricus osecanus var. osecanus. (habit X A).
A u g . - S e p t . Reported from garden i n France, also recorded from Italy.
24. Agaricus pseudoumbrella B o h u s i n M i k o l . K o z l e m . 34: 26. 1995.

-Fig. 25.
Agaricus arvensis var. umbrelloideus B o h u s i n A n n l s hist.-nat.
Mus. natn. hung. 66: 82. 1974.
SEL. D E S C R . & F I G S - B o h u s i n M i k o l . K o z l e m . 34: 26. 1995.
Pileus 65-95 m m , regularly convex, white, s l o w l y y e l l o w i n g o n han-

d l i n g , at first entirely finely fibrillose-floccose to squamulose, later
only at margin; v e i l present as appendiculate white remnants. L a m e l l a e
p i n k i s h b r o w n to greyish w i t h p i n k tinge. Stipe 50-80 X 8-18 m m ,
annulate, clavate o r w i t h a round bulb, at base (13-)20-26 m m , white,
sometimes y e l l o w i n g , above annulus smooth, below annulus finely fib-
rillose to striate, sometimes at base squamulose. A n n u l u s at 0.60-0.75
of height o f stipe, 10-22 m m w i d e , descending, pending, thick and per-
sistent, w i t h smooth upperside; underside w i t h large brownish squames
arranged as a c o g w h e e l . Context white, hardly discolouring, b e c o m i n g
faintly y e l l o w i s h to orange i n base o f stipe. S m e l l faintly l i k e anise,
sweetish. Taste not k n o w n .
M a c r o c h e m i c a l reactions: Schaeffer-reaction orange (surface o f
F i g . 24b. Agaricus osecanus var. squarrosipes. (habit X Vi).
pileus).
Spores (6.0-)6.5-8.0(-9.0) X 4.5-5.5(-6.0) m m , on average (6.5-)6.8- = 1.30-1.70, Q a v = 1.40-1.55, ellipsoid, sometimes oblong, without
7.5 X 4.8-5.3(-5.5) m m , Q = (1.20-)1.25-1.50(-1.65), Q a v = 1.35- germ pore. L a m e l l a edge w i t h c. 60 |xm broad sterile layer; cheilocys-
1.45, ellipsoid, without germ pore, often also aberrant spores present, tidia catenate, i n chains o f c y l i n d r i c a l to globose elements o f 10-19 X
differing i n size and/or shape. B a s i d i a 16-21 X 7.0-8.0 m m , 4-spored, 7.5-8.5 (Jim, w i t h globose to clavate or c y l i n d r i c a l , sometimes utriform
rarely partly 2-spored. L a m e l l a edge w i t h a 35-60(-100) m m broad, terminal elements o f (12-)15-28 X 8.5-11 m m . P i l e i p e l l i s a slightly
sterile layer; cheilocystidia i n sometimes agglutinated chains o f globose gelatinized cutis o f 3.5-7.5 m m w i d e hyphae w i t h c y l i n d r i c a l elements,
to clavate elements o f 6.0-15 X (4.0-)6.0-10 m m , w i t h clavate to c y l i n - w i t h pale y e l l o w intracellular pigment, gradually passing into pileitra-
drical or narrowly u n i f o r m terminal elements o f 15-31 X (5.5-)6.5-12 ma. Stipitipellis a cutis o f rarely branched, 3.0-8.0 | x m w i d e hyphae
mm, mostly with y e l l o w i s h b r o w n contents. Pileipellis a cutis o f 4.5- w i t h c y l i n d r i c a l elements, w i t h pale y e l l o w intracellular and parietal
10(-12) m m w i d e hyphae w i t h cylindrical, sometimes slightly inflated pigment.
AGARICUS (Nauta) 47
F i g . 25. Agaricus pseudoumbrella. (habit X A).
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic, ter-

restrial i n deciduous or coniferous woods or i n verges o f woods o n cal-
careous s o i l . V e r y rare i n the Netherlands, o n l y found i n southern
L i m b u r g (Bunderbos; C a d i e r & Keer, Riesenberg; Wijlre). Sept.-Oct. F i g . 26. Agaricus sylvicola var. sylvicola and var. squarrosus
O r i g i n a l l y described f r o m Robinia-v/oods. Rare i n Europe, but proba- (lower figs.) (habit X A).
bly widespread i n southern and eastern Europe.

25a. var. sylvicola
T h e macroscopical notes o f the D u t c h collections studied are i n c o m - S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 4 2 . 1984; R . P h i l l i p s . Paddest.
plete, therefore n o extensive m a c r o s c o p i c a l description can be given. S c h i m m . : 168. 1981; D i i h n c k e , 1200 P i l z e : 496, 497 (as A. essettei).
T h i s species can be mistaken for A . sylvicola. Agaricus pseudoum- 1993; J . Lange, F l . agar. dan. 4: p i . 1 3 8 B . 1939.
brella is more s l o w l y y e l l o w i n g o n handling, has an entirely fibrillose- S E L . D E S C R . & FIGS. - F. M 0 l l e r i n F r i e s i a : 146-148, f i g . 2 1 , p i . 2 6 ;
floccose to squamulose pileus w h e n y o u n g , slightly larger spores, and 151-153, f i g . 2 3 , p i . 2 7 (as P. abruptibulba). 1952; Wasser, Tr.
cheilocystidia o f a more irregular form. Agariceae Soviet U n i o n : 73-74. 1989.
25. Agaricus sylvicola (Vitt.) P e c k i n A n n . R e p . N . Y . State B o t . 2 3 : Pileus (55-)70-110 m m , at first h e m i s p h e r i c a l , later expanding to
97. 1872. (as A . silvicola). - F i g . 26. plano-convex o r applanate w i t h often l o w u m b o , sometimes w i t h
Agaricus campestris var. sylvicola Vitt., Descr. F u n g h i mang. Italia: depressed centre w i t h l o w u m b o , w i t h straight m a r g i n , exceeding
213. 1832; Psalliota sylvicola (Vitt.) R i c h o n & R o z e , A t l . C h a m p , lamellae, at first white, u s u a l l y very soon y e l l o w ( K . & W . 2 A 3 , 3 A 3 ;
comest. veneneux: p i . 7, figs. 13-16. 1885. - Pratella flavescens Gillet, M u . 1.5 Y 8/6), soon d i s c o l o u r i n g y e l l o w i s h to, especially i n centre,
Hymenomycetes: 564. 1878; Agaricus flavescens (Gillet) B r i t z . i n Ber. b r o w n i s h o n h a n d l i n g , later w i t h y e l l o w i s h b r o w n spots o n damaging,
naturh. Ver. A u g s b u r g 27: 168. 1883, non A . flavescens W a l l r . 1833. - usually smooth o r m i n u t e l y f i b r i l l o s e towards m a r g i n , sometimes
Agaricus essettei M . B o n i n D o c . m y c o l . 13(49): 56. 1983. radially fibrillose or, e s p e c i a l l y w h e n y o u n g , m i n u t e l y appressed to
M I S A P P L . - Agaricus abruptibulbus sensu F . M 0 l l e r i n F r i e s i a 4: sometimes patently squamulose to fibrillose-squamulose, later some-
151 1952; sensu H e i n e m . i n S y d o w i a 30: 21. ( ' 1 9 7 7 ' ) 1978. times areolate-rimose; v e i l present w h e n y o u n g as appendiculate
V E R N . N A M E - Slanke anijschampignon. white floccules at margin, later disappearing. L a m e l l a e , L = 80-120,
1 = 1-9, crowded, sometimes furcate, free, y o u n g segmentiform, later
subventricose, u p to 11 m m broad, at first white to pale grey, later
greyish p i n k ( K . & W . 6 C 2 , 6 D 4 ) to pale b r o w n to finally greyish dark
K E Y TO T H E VARIETIES brown, w i t h w h i t i s h , eroded edge. Stipe (45-)70-100(-135) X 7-13(-
15) m m , annulate, usually abruptly bulbous o r marginately bulbous
1. Stipe b e l o w annulus fibrillose, towards base sometimes minutely w i t h flattened underside, sometimes o n l y slightly enlarged at base or
squamulose 25a. var. sylvicola subbulbous, at base (12-)15-25 m m broad, often slightly bent, fistu-
1. Stipe b e l o w annulus strikingly squarrose . . . 25b. var. squarrosus lose, at first white to greenish y e l l o w above annulus, f i n a l l y slightly
b r o w n i s h , b e l o w annulus white, soon d i s c o l o u r i n g y e l l o w to orange-

y e l l o w o n h a n d l i n g , later b r o w n i s h y e l l o w , above annulus glabrous,
smooth, sometimes s i l k y striate, b e l o w annulus often fibrillose to
velutinous or sometimes lanate, towards base often minutely white-
squamulose to m i n u t e l y squarrose, sometimes arranged i n girdles,
rarely w i t h a t h i c k white fibrillose to velutinous sock, rarely w i t h
white or y e l l o w i s h basal tomentum, often w i t h short, w h i t i s h r h i z o -
morphs. A n n u l u s at (0.40-)0.65-0.75 o f height o f stipe, 9-15(-20) m m
w i d e , descending, patent or pending, thin, easily torn, white, w i t h
smooth to fibrillose upperside w h i c h is sometimes d i s c o l o u r i n g y e l -
l o w ; underside white-squamulose, w i t h c. 13 squames, to lanate-floc-
cose; squames sometimes patent and arranged as c o g w h e e l some
distance f r o m m a r g i n . Context 7-9 m m thick i n pileus, white, hardly
d i s c o l o u r i n g , sometimes faintly y e l l o w i n g i n base o f stipe when cut.
S m e l l w h e n fresh strongly l i k e anise, later faint and o n l y slightly l i k e
anise. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 10% y e l l o w (surface o f pileus, sur-
face o f stipe), negative to y e l l o w (context); Schaeffer-reaction strong-
l y orange (surface o f pileus, surface o f stipe).
Spores 5.5-7.5(-8.0) X 3.5-5.0 m m , o n average 6.0-6.7 X 4.1-4.7
mm, Q = 1.25-2.00, Q a v = 1.35-1.60, e l l i p s o i d to o b l o n g , rarely
b r o a d l y e l l i p s o i d , dark b r o w n , without germ pore. B a s i d i a 17-21 X
6.5-7.0 m m , 4-spored. L a m e l l a edge w i t h a 50-60 m m broad sterile
layer; cheilocystidia i n short chains o f globose to c y l i n d r i c a l elements
o f ( 8 . 5 - ) l l - 1 4 ( - 2 0 ) X 8.5-12 m m , w i t h predominantly globose-or
broadly clavate terminal elements o f (8.5-)14-26 X (5.5-)10-20 m m ,
usually w i t h y e l l o w contents. P i l e i p e l l i s a cutis o f 3.5-7.0 m m wide,
sometimes furcate hyphae w i t h c y l i n d r i c a l elements, w i t h pale y e l l o w
intracellular pigment gradually passing into pileitrama. Stipitipellis a
F i g . 27. Agaricus macrocarpus. (habit X A).
cutis o f 4.5-10 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes slightly
inflated elements, and ascending cylindrical terminal elements. Pileus 120-170 m m , at first paraboloid to hemispherical, expanding to
Squames consisting o f interwoven, 4.5-11 m m w i d e hyphae with convex or plano-convex, w i t h inflexed m a r g i n exceeding lamellae,
slightly inflated elements o f 17-37 m m long, w i t h clavate to sometimes white, usually gradually but intensely y e l l o w i n g o n handling, finally
irregularly shaped terminal elements o f 21-44 X 8.0-14 m m . y e l l o w ( M u . 10 Y R 8/5) to y e l l o w i s h white, sometimes w i t h b r o w n i s h
H A B I T A T & DISTR. - Solitary or gregarious, saprotrophic, terrestrial spots where touched, y o u n g radially fibrillose to silky, later smooth,
i n coniferous and deciduous woods o n nutrient-rich or calcareous soil. fibrillose or fibrillose-squamulose to flocculose near margin; partial
Rather rare but widespread i n the Netherlands, Sept.-Oct. C o m m o n v e i l present when y o u n g as white appendiculate teeth, later as irregu-
and widespread i n Europe. C o s m o p o l i t a n . lar fringe; universal v e i l w h e n y o u n g present as white patches o n sur-
face near margin, later disappearing. L a m e l l a e , L = 110-122, 1 =
Agaricus essettei is often regarded as a separate species, differing from l-2(-5), crowded, free, segmentiform to subventricose, up to 11 m m
A. sylvicola i n the stipe w i t h a more abrupt, marginate bulb and slight- broad, at first pale grey, later pale greyish p i n k ( K . & W . 7 C 2 , 7 D 2 - 3 )
l y larger spores. Since a l l intermediate variants can be found A. esset- to p i n k i s h b r o w n , w i t h entire, white edge. Stipe 100-200 X (15-)20-30
tei i s considered conspecific w i t h A. sylvicola. m m , annulate, c y l i n d r i c a l to subclavate, usually abruptly bulbous w i t h
flattened underside, at base up to 45 m m broad, straight, fistulose, at
25b. var. squarrosus N a u t a i n Persoonia 17: 4 6 1 . ('2000') 2001. first white, later downwards y e l l o w i n g to y e l l o w i s h b r o w n or orange
on handling, above annulus smooth, b e l o w annulus when y o u n g usu-
C H A R A C T E R I S T I C S - Differing from the typical variety i n the distinctly ally entirely floccose-squamulose, later w i t h smooth zone, below that
squarrose stipe b e l o w the annulus. minutely squamulose and near base white-lanate to white-fibrillose,
M i c r o s c o p i c a l characters as i n typical variety. sometimes w i t h some short rhizomorphs. A n n u l u s at 0.70-0.80 o f
H A B I T A T & DISTR. - In deciduous and coniferous woods o n nutrient- height o f stipe, 20-25 m m w i d e , descending, pending to slightly patent,
r i c h soil. V e r y rare i n the Netherlands (Bloemendaal; Noordoostpolder, thick, persistent, white, w i t h smooth upperside; underside usually w i t h
K u i n d e r b o s ; N i j m e g e n , estate Oosterhout), but probably more wide- many thick, cream to coloured ochraceous, fibrillose squames, some-
spread. A u g . - O c t . N o t yet recorded outside the Netherlands. times arranged as a cogwheel. Context 9-13 m m thick i n pileus, white,
hardly d i s c o l o u r i n g w h e n cut, sometimes reddish b r o w n or faintly y e l -
26. Agaricus macrocarpus (F. M 0 l l e r ) F . M 0 l l e r i n F r i e s i a 4 : 204. l o w i n g i n base o f stipe. S m e l l strongly to faintly like anise, sometimes
1952. - F i g . 27. unpleasant. Taste not k n o w n .
Psalliota macrocarpa F. M 0 l l e r i n F r i e s i a 4: 153. 1952. M a c r o c h e m i c a l reactions: Schaeffer-reaction orange (surface o f
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 182. 1995; pileus).
C a p p e l l i , A g a r i c u s : p i . 4 5 . 1984. Spores (6.5-)7.0-9.0(-9.5) X 5.0-6.5 m m , o n average 7.7-8.4 X 5.4-
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 153-155, p i . 28. 1952; 5.8 m m , Q = 1.30-1.65, Q a v = 1.40-1.45, e l l i p s o i d , without germ
Wasser, Tr. A g a r i c e a e Soviet U n i o n : 74-75. 1989. pore. B a s i d i a 16-23 X 6.5-8.5 m m , usually 4-spored, often partly 2 -
V E R N . N A M E - Forse anijschampignon. spored, rarely for the greater part 2-spored. L a m e l l a edge w i t h a 50-60
AGARICUS (Nauta) 49
mm broad, m a i n l y sterile layer o f s l i g h t l y agglutinated elements; 1978, n o n A. stramineus S c o p . 1772; Agaricus macrosporus var.
cheilocystidia usually i n short chains o f subglobose to c y l i n d r i c a l ele- excellens (F. M 0 l l e r ) B o h u s i n A n n l s hist.-nat. M u s . natn. hung. 8 1 :
ments o f (5.0-)8.5-16(-20) X (4.0-)6.0-12(-15) m m , w i t h mostly glo- 41. 1990, non A. stramineus S c o p . 1772; Agaricus albertii var. excel-
bose, to c y l i n d r i c a l terminal elements o f 14-25 X (8.0-)9.0-16.5 m m , lens (F. M 0 l l e r ) B o h u s i n A n n l s hist.-nat. M u s . natn. h u n g . 82: 5 1 .
mostly w i t h b r o w n contents. P i l e i p e l l i s a slightly gelatinized cutis o f 1990 (superfluous). - Agaricus kuehnerianus H e i n e m . i n B u l l . mens.
3.5-8.5 m m w i d e hyphae w i t h c y l i n d r i c a l to inflated elements up to 13 Soc. l i n n . L y o n , N o . spec. 4 3 : 183. 1974.
m m w i d e , w i t h scattered ascending c y l i n d r i c a l to clavate terminal ele-
ments u p to 10 m m , gradually passing into pileitrama, w i t h pale y e l - K E Y TO THE VARIETIES
low parietal pigment. Stipitipellis a s l i g h t l y gelatinized cutis o f 1. Pileus soon w i t h y e l l o w i s h tinges; ratio o f pileus w i d t h and stipe
sometimes branched, 3.5-8.0(-13) m m w i d e hyphae w i t h c y l i n d r i c a l length usually > 1; surface o f stipe b e l o w annulus floccose-
elements and scattered ascending clavate terminal elements up to 25 squamose; i n grasslands 27a. var. urinascens
mm w i d e . Squames consisting o f interwoven, c u r v e d 8.0-10.0 m m 1. Pileus whitish; ratio o f pileus w i d t h and stipe length usually < 1;
w i d e hyphae w i t h u p to 18 m m w i d e terminal elements, w i t h y e l l o w surface o f stipe b e l o w annulus smooth and striate, towards base
parietal pigment. squamose; i n woods 27b. var. excellens
H A B I T A T & DISTR. - Solitary o r gregarious, saprotrophic, terrestrial
i n coniferous woods o n nutrient-rich, calcareous s o i l . Rather rare, 27a. var. urinascens
m a i n l y i n the new polders. A u g . - O c t . R a r e but widespread i n Europe, S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 4 1 . 1984 (as A . macrosporus);
also occurring i n A s i a (former Soviet U n i o n ) . R. P h i l l i p s , Paddest. S c h i m m . : 165. 1981 (as A. macrosporus);
Dahncke, 1200 P i l z e : 500. 1993 (as A . macrosporus).
27. Agaricus urinascens ( M 0 l l e r & Schaeff.) S i n g , i n L i l l o a 22: 431. S E L . D E S C R . & FIGS. - F. M 0 l l e r i n F r i e s i a 4: 178-181, f i g . 29, p i . 33
( ' 1 9 4 9 ' ) 1 9 5 1 . - F i g . 28. (as P. macrospora); 177-178, p i . 1 9 A (as P. straminea). 1952.
Psalliota urinascens M 0 l l e r & Schaeff. i n A n n l s m y c o l . 36: 7 9 . V E R N . N A M E - Grootsporige champignon.
1938. - Psalliota arvensis subsp. macrospora M 0 l l e r & Schaeff. i n
A n n l s m y c o l . 3 6 : 7 8 . 1938; Agaricus macrosporus (M0ller & Pileus 105-265 m m , at first obtusely c o n i c a l to conico-convex, later
Schaeff.) P i l a t i n A c t a M u s . nat. P r a g . 7 B (1): 7 8 . 1951, n o n A. expanding to plano-convex or applanate, subumbonate o r w i t h slight-
macrosporus M o n t . 1837; Psalliota macrospora ( M 0 l l e r & Schaeff.) ly depressed centre, w i t h inflexed margin, at first white to cream-white,
F. M 0 l l e r i n F r i e s i a 4 : 181. 1952; Agaricus albertii M . B o n in Doc. y e l l o w to y e l l o w i s h b r o w n o n handling, p i n k i s h y e l l o w o n scratching,
m y c o l . 18(72): 6 3 . 1988. - Psalliota straminea M 0 l l e r & Schaeff. i n finally pale y e l l o w to y e l l o w - b r o w n ( M u . 2.5 Y 8/4, 8/3-4, 10 Y R 8/6),
A n n l s m y c o l . 36: 78. 1938; Agaricus stramineus ( M 0 l l e r & Schaeff.) i n centre pale b r o w n , at first entirely squamulose, later centre smooth
S i n g , i n L i l l o a 22: 4 3 2 . ( ' 1 9 4 9 ' ) 1951, n o n A . stramineus S c o p . 1772; and towards m a r g i n squamulose o r entirely s i l k y w i t h scattered
Agaricus macrosporus var. stramineus ( M 0 l l e r & Schaeff.) M . B o n i n squames, often o n l y near m a r g i n , o r entirely fibrillose, sometimes
D o c . m y c o l . 15(60): 25. 1985; Agaricus substramineus Courtecuisse areolate-rimose; squames at first often white, later often partly pale
i n D o c . m y c o l . 16(61): 4 9 . 1985; Agaricus stramineosquamulosus y e l l o w i s h b r o w n to pale b r o w n (7.5 Y R 7/6, 10 Y R 8/4); v e i l w h e n
Rauschert i n N o v a H e d w i g i a 54: 215. 1992 (superfluous).- Psalliota y o u n g present as appendiculate teeth, later m o s t l y disappearing but
collina V e l e n . , N o v i t . m y c o l . nov.: 8 2 . 1947. -Psalliota excellens F. sometimes r e m a i n i n g as white irregular v e i l patches near margin.
M 0 l l e r i n F r i e s i a 4 : 178. 1952; Agaricus excellens (F. M 0 l l e r ) F . L a m e l l a e , L = 165-220, 1 = 2-4, very c r o w d e d , free, linear to seg-
M 0 l l e r i n F r i e s i a 4 : 204. 1952; Agaricus macrosporus subsp. excel- mentiform, sometimes undate, sometimes furcate, up to 21 m m broad,
lens (F. M 0 l l e r ) B o h u s i n A n n l s hist.-nat. M u s . natn. hung. 7 0 : 105. at first p i n k i s h white to pale p i n k or grey-lilacinous (7.5 Y R 8/2, 5 Y R
0 00 0
F i g . 28. Agaricus urinascens. (habit X A).

ll
7/4, 8/3), later dark grey l i l a c i n o u s (5 Y R 6/4), finally dark reddish nat. M u s . natn. hung. 8 1 : 4 1 . 1990, n o n A . stramineus S c o p . 1772;
b r o w n to dark b r o w n (5 Y R 3-4/3, 7.5 Y R 3/2), w i t h paler to white, Agaricus albertii var. excellens (F. M 0 l l e r ) B o h u s i n A n n l s hist.-nat.
mostly irregular to minutely serrate edge. Stipe 80-180 X 1 8 - 5 5 m m , M u s . natn. hung. 82: 5 1 . 1990 (superfluous).
annulate, m o s t l y clavate to bulbous and at base up to 53 m m broad, S E L . I C O N - R . B r o w n i n M y c o l o g i s t 13(1): front cover. 1999;
sometimes tapering towards base, mostly straight, fistulose to stuffed, C a p p e l l i , A g a r i c u s : p i . 4 1 . 1984; R . P h i l l i p s , Paddest. S c h i m m . : 167.
at first creamy white, later w i t h b r o w n i s h tinges, above annulus most- 1981.
l y glabrous to silky, sometimes minutely squamulose, b e l o w annulus S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 178-181, fig. 29, p i . 33.
floccose to strongly squamose, w i t h m o s t l y concolorous, sometimes 1952; Wasser, Tr. A g a r i c e a e Soviet U n i o n : 67-68. 1989.
very pale b r o w n , sometimes pointed squames; squames sometimes V E R N . N A M E - Statige c h a m p i g n o n .
arranged i n girdles, later sometimes disappearing. A n n u l u s at 0.70-
0.80 o f height o f stipe, 30 m m wide, descending, pending, thick, white, C H A R A C T E R I S T I C S - Differing f r o m var. urinascens i n the whiter pileus,
w i t h smooth, sometimes m i n u t e l y sulcate upperside; underside strong- the slender habit, stipe mostly longer than pileus width, and the surface
l y floccose w i t h concolorous squames; sometimes along m a r g i n thick, of the stipe w h i c h is striate or smooth directly b e l o w the annulus
y e l l o w to b r o w n i s h y e l l o w squames present, arranged as a cogwheel. instead o f squamose as i n var. urinascens.
Context 20-31 m m thick i n pileus, soft, white, not discolouring to dis- M i c r o s c o p i c a l characters as i n t y p i c a l variety.
c o l o u r i n g p i n k i s h to pale reddish b r o w n i n centre o f pileus and base o f H A B I T A T & DISTR. - Solitary, saprotrophic, terrestrial i n coniferous
stipe w h e n cut, sometimes y e l l o w i n base o f stipe. S m e l l fresh faintly
and deciduous woods. V e r y rare i n the Netherlands (Hoogmade), but
like almonds, soon unpleasant. Taste nutty, pleasant.
probably more widespread; Sept.-Oct. K n o w n from nearby countries
M a c r o c h e m i c a l reactions: K O H 10% pale y e l l o w (surface o f as D e n m a r k , Germany, but rare.
pileus), y e l l o w (context o f base and surface o f stipe), negative (context
o f pileus; upperside annulus), faintly y e l l o w (underside annulus);
Schaeffer-reaction negative to orange (surface o f pileus), red to pur-
ple-red (surface o f stipe). Sect. Minores Fr.
Spores (9.0-)9.5-11.5(-12.0) X 6.0-7.5(-8.0) m m , o n average 10.1-
10.7 X 6.5-7.2 m m , Q = 1.35-1.70(-1.80), Q a v = 1.50-1.60, e l l i p - Pileus strongTy~yellowing w h e n touched or o n d r y i n g ; annulus narrow,
soid, sometimes o b l o n g , without g e r m pore. B a s i d i a 20-32 X 8.5-10.5 thin, up to 10 m m w i d e ; sometimes absent) fibrillose, without flocks or
m m , clavate, 4-spored, sometimes 2-spored, sometimes w i t h b r o w n - warts at underside; context discolouring slightly yellow, orange-yellow
ish contents. L a m e l l a edge w i t h a 55-70 m m broad, sterile layer; or y e l l o w i s h b r o w n o n exposure or when damaged.
c h e i l o c y s t i d i a i n often s l i g h t l y agglutinated chains o f c y l i n d r i c a l to K O H - r e a c t i o n y e l l o w to orange; Schaeffer-reaction o n surface o f
subglobose elements o f (7.0-)8.0-13.5(-16) X (5.0-)6.0-9.0 m m , w i t h pileus strongly orange.
clavate, rarely irregularly shaped, terminal elements o f (10-)15-28(- Spores ellipsoid; average spore length < 6.5 m m ; Q a v = 1.35-1.60;
32) X 6.5-13 m m , often w i t h y e l l o w i s h contents. P i l e i p e l l i s a some- without g e r m pore; l a m e l l a edge sterile; cheilocystidia shortly catenate
times irregular cutis o f 3.5-6.5 m m w i d e , sometimes branched, or not.
h y a l i n e hyphae w i t h c y l i n d r i c a l elements, w i t h pale y e l l o w intracel-
lular pigment, w i t h often ascending, c y l i n d r i c a l to narrowly clavate, A l t h o u g h sect. Minores seems to f o r m a homogeneous group, the
up to 7 m m w i d e , terminal elements, gradually passing into pileitra- d e l i m i t a t i o n o f the species is far f r o m clear. T h e species have been
ma. Squames c o n s i s t i n g o f i n t e r w o v e n hyphae with ascending, distinguished o n account o f size o f spores, and amount and size o f
clavate, 5.0-8.0 m m w i d e , terminal elements, w i t h intracellular, most- c h e i l o c y s t i d i a , but these characters have s h o w n to be variable. A l s o ,
l y pale y e l l o w i s h b r o w n pigment. S t i p i t i p e l l i s a cutis o f sometimes colours o f the p i l e u s are l i k e w i s e variable, w i t h often vinaceous fib-
branched, 3.5-5.0 m m w i d e hyphae o f c y l i n d r i c a l , s l i g h t l y inflated rils soon fading. M o d e r n methods are required to c o m e to a more b a l -
elements. Squames c o n s i s t i n g o f chains o f inflated to globose ele- anced t a x o n o m y w i t h i n this group and judge the t a x o n o m i c a l value
ments, 20-38 X 6.5-16 m m . o f the m o r p h o l o g i c a l characters l i k e p i l e u s surface and amount o f
H A B I T A T & DISTR. - Solitary or gregarious, saprotrophic, terrestrial, purple. H e r e a p r a c t i c a l approach is chosen, and the nomenclature
m o s t l y i n grasslands, sometimes i n roadside verges (rarely planted and interpretation o f the taxa f o l l o w the o r i g i n a l descriptions as
w i t h trees), o n nutrient-rich or calcareous s o i l . Rather rare i n the c l o s e l y as possible.
Netherlands, J u l y - O c t . W i d e s p r e a d but rare i n Europe. A l s o k n o w n
from A s i a . 28. Agaricus dulcidulus S. Schulz. i n K a l c h b r . , Ic. sel. H y m e n o m y c .
H u n g . : 29. 1 8 7 4 . - F i g . 29.
F o r an extensive discussion o f the s y n o n y m y see N a u t a (in Persoonia Psalliota purpurella F. M 0 l l e r i n F r i e s i a 4: 193. 1952; Agaricus
17: 461-463. ( ' 2 0 0 0 ' ) 2001). purpurellus (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204. 1952.
Agaricus macrosporoides B o h u s (in A n n l s hist.-nat. M u s . natn. M I S A P P L . - Agaricus semotus sensu B o h u s i n M i k o l . K o z l e m . 34:
hung. 66: 84. 1974) is a species w h i c h m a c r o s c o p i c a l l y resembles A. 21. 1995; sensu M . B o n i n D o c . m y c o l . 15(60): 22. 1985; Psalliota
urinascens i n every aspect, but differs i n the smaller spores o f 8.0-9.5 semota sensu F. M 0 l l e r i n F r i e s i a 4: 194. 1952. - Psalliota amethysti-
X 5.3-6.0 m m . A c c o r d i n g to B o h u s it occurs i n saline pastures. na sensu J . L a n g e , F l . agar. dan. 4: 1 3 5 A . 1939.
E X C L . - Agaricus dulcidulus sensu J . L a n g e i n D a n s k bot. A r k .
27b. var. excellens (F. M 0 l l e r ) N a u t a i n Persoonia 17: 4 6 2 . ( ' 2 0 0 0 ' ) 4(12): 11. 1926, F l . agar. dan. 4: 6 1 , p i . 135C. 1939.
2001. S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 191. 1995 (as
Psalliota excellens F. M 0 l l e r i n F r i e s i a 4: 178. 1952; Agaricus A. semotus); J . L a n g e , F l . agar. dan. 4: 1 3 5 A , 1 3 7 A . 1939 (as P.
excellens (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204. 1952; Agaricus amethystina and P. semota resp.).
macrosporus subsp. excellens (F. M 0 l l e r ) B o h u s i n A n n l s hist.-nat. S E L . D E S C R . & FIGS. - F. M 0 l l e r i n F r i e s i a 4: 195-196, p i . 17. 1952
Mus. natn. hung. 70: 105. 1978, n o n A. stramineus S c o p . 1772; (as P. semota).
Agaricus macrosporus var. excellens (F. M 0 l l e r ) B o h u s i n A n n l s hist.- V E R N . N A M E - Wijnkleurige champignon.
AGARICUS (Nauta) 51
hyphae with c y l i n d r i c a l elements, w i t h c y l i n d r i c a l terminal elements,

w i t h y e l l o w i s h , intracellular pigment, gradually passing into pileitra-
ma. Squames consisting o f curved, u p to 7.0 |j.m w i d e hyphae w i t h
brownish intracellular pigment. Stipitipellis a cutis o f sometimes
branched, 2.5-6.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h
pale, intracellular pigment.
H A B I T A T & DISTR. - Solitary or gregarious, saprotrophic, terrestrial
i n deciduous and coniferous woods, city parks, gardens, o n humus-rich
soil, also i n open dunes. Rather rare, c o m m o n i n coastal areas and
southern L i m b u r g ; J u l y - N o v . W i d e s p r e a d but not c o m m o n i n Europe.
A l s o k n o w n from A s i a , northern A f r i c a and the A m e r i c a s .
This taxon is i n modern literature often k n o w n as Agaricus semotus.

However, the o r i g i n a l description o f F r i e s ( M o n o g r . H y m e n o m y c .
Sueciae 2: 347. 1863) and his later plate (Ic. sel. H y m e n o m y c : t. 131.
1879) o f A. semotus are very doubtful, s h o w i n g a stipe covered with
reddish v e i l . These do not refer to the present taxon, and probably not
even to an Agaricus.
F i g . 29. Agaricus dulcidulus. Agaricus purpurellus i s a s m a l l variant o f A. dulcidulus, not worth
distinguishing as a separate taxon. It i s based o n a plate b y L a n g e ( F l .
P i l e u s (20-)30-60(-70) m m , at first c o n i c o - c o n v e x or truncately c o n - agar. dan. 4 : p i . 1 3 5 A . 1939), w h i c h shows a basidiocarp w i t h purple-
i c a l , later convex w i t h flattened centre to plano-convex, sometimes fibrillose pileus. T h e differences according to M 0 l l e r w i t h A. semotus
w i t h depressed centre, at first w i t h s l i g h t l y i n f l e x e d m a r g i n , later rev- are a more purple pileus and smaller basidiocarps. N o original materi-
olute, u s u a l l y w i t h vinaceous to p u r p l e - b r o w n or p i n k i s h grey dense- al o f M 0 l l e r has been preserved.
l y fibrillose to fibrillose-squamulose centre ( M u . 7.5 R 7/4, 5 Y R 3/3, Agaricus lutosus (F. M 0 l l e r ) F. M 0 l l e r (in F r i e s i a 4: 204. 1952) dif-
4/4, 6/3-4), towards m a r g i n paler a n d there w h i t i s h to y e l l o w i s h fers, according to M 0 l l e r , i n the c y l i n d r i c a l stipe w h i c h is tapering
b r o w n (10 Y R 7/6), sometimes g r e y i s h b r o w n , o c c a s i o n a l l y o n l y w i t h towards the base, and the pileus w h i c h i s y e l l o w i s h and w i t h some
some r e d d i s h b r o w n fibrils at centre; p u r p l e c o l o u r s often fading, vinaceous fibrils at the centre o n l y w h e n y o u n g .
e s p e c i a l l y towards m a r g i n ; sometimes later entirely y e l l o w i s h b r o w n
or ochraceous b r o w n , rather strongly y e l l o w i n g o n h a n d l i n g or 29. Agaricus luteomaculatus (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204.
scratching; m a r g i n y o u n g w i t h c. 2 m m r i m e x c e e d i n g lamellae, later 1952. - F i g . 30.
w i t h or w i t h o u t appendiculate v e i l remnants. L a m e l l a e , L = 55-75,1 Psalliota luteomaculata F . M 0 l l e r i n F r i e s i a 4: 192. 1952. -
= l ( - 3 ) , c r o w d e d , free, subventricose, u p to 9 m m broad, at first pale Agaricus lutosus var. macrosporus Parra i n B o l . S o c . m i c o l . M a d r i d
greyish to g r e y i s h b r o w n (10 Y R 7/3, 5 Y R 6-7/4, 6/1-2), later p i n k - 23: 98. 1998 (invalid).
ish b r o w n (10 Y R 5/3), f i n a l l y dark b r o w n (5 Y R 4/3), w i t h w h i t i s h S E L . I C O N . - F. M 0 l l e r i n F r i e s i a 4: p i . 21b. 1952; Parra i n B o l . Soc.
to paler, crenulate edge. Stipe (15-)20-45(-80) X (2-)4-7(-10) m m , m i c o l . M a d r i d 23: 99. 1998.
annulate, c y l i n d r i c a l w i t h bulbous base to clavate, at base u p to 11(- S E L . D E S C R . & FIGS. - F. M 0 l l e r i n Friesia 4: 192. 1952; Parra i n B o l .
13) m m b r o a d , straight, n a r r o w l y fistulose, w h i t e to w h i t i s h , often Soc. m i c o l . M a d r i d 23: 98-99. 1998 (as A. lutosus var. macrosporus).
towards base y e l l o w i s h to y e l l o w i s h b r o w n (2.5 Y 8/8, 10 Y R 6-7/8), V E R N . N A M E - Okerbruine dwergchampignon.
q u i c k l y y e l l o w i n g o n h a n d l i n g , above annulus glabrous, silky, b e l o w
annulus at first flocculose, later fibrillose to smooth, near base veluti- Pileus 55-65 m m , convex, y e l l o w i s h b r o w n , at centre purple b r o w n fib-
nous, o c c a s i o n a l l y w i t h several s m a l l short branched w h i t e r h i - rillose ( M u . 9 Y R 6/6), q u i c k l y y e l l o w i n g o n handling, later entirely
z o m o r p h s . A n n u l u s at 0.65-0.75 o f height o f stipe, narrow, ( l - ) 2 - 3 ochraceous brown; w i t h m a r g i n exceeding lamellae w i t h 1 m m broad
mm w i d e , d e s c e n d i n g , p e n d i n g to s l i g h t l y spreading, t h i n , fragile, white r i m . L a m e l l a e crowded, free, ventricose, 5.5-7 m m broad, at first
fibrillose, w h i t i s h , often d i s c o l o u r i n g y e l l o w i s h , w i t h smooth or stri- p i n k i s h grey, later b r o w n (5 Y R 4/3), w i t h w h i t i s h , fimbriate edge.
ate to sometimes sulcate upperside; underside f l o c c u l o s e to fibril- Stipe 60-85 X 11-12 m m , annulate, c y l i n d r i c a l to subclavate, at base
lose-floccose. C o n t e x t 3-4.5(-7) m m thick i n pileus, whitish, sometimes up to 16 m m broad, straight, narrowly fistulose, w h i t i s h to
d i s c o l o u r i n g s l i g h t l y y e l l o w i s h to ochraceous w i t h orange tinge i n y e l l o w i s h (10 Y R 7/8), near base y e l l o w i s h b r o w n , y e l l o w i n g o n han-
stipe w h e n cut. S m e l l l i k e almonds, sometimes also anise, often not d l i n g , above annulus smooth to fibrillose, b e l o w annulus fibrillose,
strong. Taste pleasant, l i k e nuts, or l i k e almonds. sometimes w i t h rhizomorphs. A n n u l u s at c. 0.60 o f height o f stipe, up
M a c r o c h e m i c a l reactions: K O H 10% & 4 0 % orange (base o f stipe), to 8 m m wide, descending, pending, thin, white, y e l l o w i n g , w i t h
y e l l o w (surface o f pileus); Schaeffer-reaction orange (surface o f pileus smooth upperside; underside fibrillose-flocculose. Context 7 m m thick
and stipe). in pileus, w h i t i s h , d i s c o l o u r i n g y e l l o w i s h b r o w n , sometimes w i t h
Spores 4.5-6.0 X 3.5-4.0 m m , o n average 5.1-5.6 X 3.4-3.7 m m , Q orange tinge i n stipe. S m e l l hardly distinct, s l i g h t l y l i k e anise or
= 1.35-1.65, Q a v = 1.35-1.50, ellipsoid, without germ pore. B a s i d i a almonds. Taste not k n o w n .
17-23 X 6.0-8.0 m m , 4-spored. L a m e l l a edge w i t h a 40-50 m m broad, M a c r o c h e m i c a l reactions: a m m o n i a y e l l o w (surface o f pileus);
usually sterile layer; c h e i l o c y s t i d i a usually i n short chains o f globose K O H 10% y e l l o w (surface o f pileus); Schaeffer-reaction orange (sur-
to quadrangular elements o f 4.0-5.5 X 5.0 m m , w i t h clavate, some- face o f pileus).
times globose terminal elements o f (10-)14-21(-30) X (7.0-)9.5-14(- Spores 5.5-6.5 X 4.0-5.0 m m , o n average 6.2-6.3 X 4.5-4.7 m m ,
17.5) m m , often w i t h y e l l o w i s h contents; sometimes intermixed w i t h Q = 1.25-1.50, Q a v = 1.35-1.40, e l l i p s o i d , without g e r m pore.
some basidia. P i l e i p e l l i s a sometimes s l i g h t l y gelatinized cutis o f B a s i d i a 20-23 X 6.0-8.0 m m , 4-spored. L a m e l l a edge almost sterile;
slightly irregularly arranged, unbranched, 3.5-6.5(-ll) m m wide c h e i l o c y s t i d i a not i n chains, 14-21 X 10-13 m m , clavate, w i t h
Pileus 17-35(-40) m m , at first convex w i t h s l i g h t l y flattened centre,

expanding to plano-convex w i t h depressed centre, rarely w i t h l o w
umbo, white to w h i t i s h ( M u . 10 Y R 7/2), rarely w i t h some faint pur-
p l i s h hue at centre, soon y e l l o w i s h or b r o w n i s h y e l l o w (10 Y R 7/8,
7/10), d i s c o l o u r i n g q u i c k l y y e l l o w o n h a n d l i n g ; surface m i n u t e l y
velutinous-fibrillose; remarkably orange i n dried material. L a m e l l a e ,
L = 55-70,1 = 1-3, crowded, free, subventricose to linear, up to 3.5
m m broad, at first pale p i n k i s h grey to b r o w n i s h grey (10 Y R 6/2), later
darker greyish b r o w n , finally b r o w n , w i t h entire to minutely fimbriate
white edge. Stipe 20-55 X 2-6 m m , annulate, c y l i n d r i c a l to subclavate,
at base 5-8(410) m m broad, straight to curved, stuffed to fistulose,
white, strongly y e l l o w o n h a n d l i n g , b e l o w annulus ochraceous w i t h
orange tinge, above annulus glabrous, smooth; b e l o w annulus f i b r i l -
lose to flocculose-fibrillose, without or w i t h short white rhizomorphs.
A n n u l u s at 0.62-0.70 o f height o f stipe, narrow, 1.5-3 m m w i d e ,
descending, pending, thin, fragile, white, d i s c o l o u r i n g yellow, w i t h
smooth upperside; underside fibrillose to velutinous. Context 2-3(-4.5)
m m thick i n pileus, w h i t i s h , not d i s c o l o u r i n g w h e n cut except for base
o f stipe w h i c h s l o w l y discolours faintly y e l l o w i s h orange. S m e l l l i k e
F i g . 30. Agaricus luteomaculatus. almonds. Taste pleasant, l i k e nuts.
M a c r o c h e m i c a l reactions: K O H 4 0 % y e l l o w (surface o f pileus,
context i n base o f stipe); Schaeffer-reaction strongly orange (surface o f
y e l l o w i s h contents. P i l e i p e l l i s a sometimes slightly gelatinized cutis pileus, surface o f base o f stipe).
o f 3.0-5.0 m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h some Spores 4.5-5.5(-6.0) X 3.5-4.0 m m , o n average 5.2-5.3 X 3.6-3.7
ascending, clavate terminal elements up to 8.5 u.m w i d e , w i t h y e l - m m , Q = 1.30-1.60, Q a v = 1.40-1.45, e l l i p s o i d , without g e r m pore.
l o w i s h i n t r a c e l l u l a r pigment, gradually passing into p i l e i t r a m a . B a s i d i a 17-18 X (4.5-)6.5-7.0 m m , 4-spored. L a m e l l a edge w i t h a
S t i p i t i p e l l i s a cutis o f unbranched, 5.0-6.5 m m w i d e hyphae w i t h mixture o f basidia and usually a minority o f cheilocystidia, cheilo-
c y l i n d r i c a l , sometimes up to 8.5 \xm inflated elements, w i t h y e l l o w - c y s t i d i a sometimes scarce; c h e i l o c y s t i d i a i n short chains o f inflated
ish intracellular pigment. rectangular elements o f 7-11 X 4.5-6.0(-7.0) m m , w i t h clavate ter-
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter- m i n a l elements o f ( l l - ) 1 6 . 5 - 2 4 X (6.0-)7.5-10.5(-16) m m , some-
restrial i n deciduous w o o d s o n calcareous sandy s o i l (according to times w i t h y e l l o w contents. P i l e i p e l l i s a cutis o f (2.5-)3.5-6.0 m m
M 0 l l e r (in F r i e s i a 4: 192. 1952) i n Ffcea-woods). Very rare, probably w i d e hyphae w i t h c y l i n d r i c a l , sometimes u p to 8 p,m elements, w i t h
confined to dune region (Wassenaar, B i e r l a p ; Overveen, E l s w o u t ) . s l i g h t l y ascending, clavate t e r m i n a l elements, w i t h y e l l o w i s h , intra-
O c t . - N o v . V e r y rare but widespread i n Europe. A l s o k n o w n from A f r i c a c e l l u l a r pigment, g r a d u a l l y p a s s i n g into p i l e i t r a m a . S t i p i t i p e l l i s a
and southern S o u t h A m e r i c a . cutis o f unbranched, 4.0-8.0 m m w i d e hyphae w i t h c y l i n d r i c a l ele-
ments, w i t h some ascending terminal elements, w i t h y e l l o w i s h intra-
T h e macroscopic description is incomplete because o f the rare occur- cellular pigment.
rence o f the taxon. T h e two collections examined differ from M 0 l l e r ' s H A B I T A T & D I S T R . - U s u a l l y solitary, saprotrophic, terrestrial i n
description i n the slightly larger pileus and the longer stipe w h i c h is at lawns, or roadside verges, o n sandy or c l a y e y s o i l . Rather rare but
most subclavate. widespread i n the Netherlands, m a i n l y i n dune region and southern
Agaricus luteomaculatus differs, according to the original descrip- L i m b u r g ; probably under-recorded. Sept.-Oct. Rather rare but w i d e -
tion b y M 0 l l e r , from the other species i n sect. Minores i n the shorter, spread i n E u r o p e . A l s o k n o w n from the A m e r i c a s , the P h i l i p p i n e s and
more floccose stipe b e l o w the ring, and i n the slightly larger spores o f southern South A m e r i c a .
5.5-6 p-m. A l t h o u g h M 0 l l e r ' s plate and description show a species with
a clavate stipe this character is not considered o f t a x o n o m i c a l i m p o r -
tance. T h e examined collections h a d a c y l i n d r i c a l to subclavate stipe.
Agaricus lutosus var. macrosporus refers to the same taxon, although
again w i t h c y l i n d r i c a l instead o f clavate stipe.
T h e recently described A. pseudolutosus ( M o r e n o et al.) M o r e n o et
al. (in M i c o l . Veget. medit. 14: 60. 1999) is, j u d g i n g from the plate and
description, without doubt closely related to the above described taxon.
T h e o n l y difference is the absence o f cheilocystidia i n A. pseudoluto-
sus and the c y l i n d r i c a l stipe instead o f the subclavate stipe of A. luteo-
maculatus o n the original plate o f M 0 l l e r .
30. Agaricus comtulus Fr., E p i c r i s i s : 215. 1838. - F i g . 3 1 .

Agaricus niveolutescens H u i j s m a n i n Persoonia 1: 321. 1960.
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 52, p i . 57 (as A . niveolutescens).
1984; J . L a n g e , F l . agar. dan. 4 : p i . 1 3 6 A . 1939.
S E L . D E S C R . & FIGS. - H u i j s m a n i n Persoonia 1: 321-322, figs. 1 &
2. 1960 (as A . niveolutescens); F. M 0 l l e r i n F r i e s i a 4: 185-186. 1952.
V E R N . N A M E - K l e i n e champignon. F i g . 3 1 . Agaricus comtulus.
AGARICUS (Nauta) 53
Agaricus comtulus is rarely collected a n d described f r o m the of stipe, up to 8 m m w i d e , descending, p e n d i n g to s l i g h t l y spreading,

Netherlands, therefore the above description is also based o n descrip- thin, fragile, e a s i l y torn, w h i t e , edge d i s c o l o u r i n g strongly y e l l o w ,
tions o f B r i t i s h material, and o n F r e n c h material b y H u i j s m a n (in w i t h smooth o r striate upperside; underside f i b r i l l o s e . C o n t e x t c. 5
Persoonia 1: 321-322. 1960). T h e flesh-coloured lamellae w h i c h are m m t h i c k i n pileus, white, d i s c o l o u r i n g s l o w l y faintly orange w h e n
mentioned i n the t y p i c a l description b y F r i e s ( E p i c r i s i s : 215. 1838), cut. S m e l l strongly l i k e bitter almonds, sometimes w i t h anise c o m -
c o u l d not be found i n the D u t c h material. L a n g e ( F l . agar. dan. 4 : 62. ponent. Taste l i k e nuts.
1939) c l a i m s that this taxon lacks c h e i l o c y s t i d i a . T h i s c o u l d not be M a c r o c h e m i c a l reactions: K O H 10% y e l l o w (surface) o r negative
confirmed, although sometimes the cheilocystidia are very scarce. to pale orange (context); Schaeffer-reaction strongly orange (surface
T h e identity o f Agaricus rusiophyllus L a s c h ( i n L i n n a e a 3: 37. of pileus).
1828), w h i c h is sometimes mentioned i n connection w i t h the above Spores 4.5-6.0 X 3.5-4.5 m m , on average 5.2-5.6 X 3.7-4.2 m m , Q
described taxon, is doubtful. = 1.20-1.50(-1.55), Q a v = 1.35-1.40, e l l i p s o i d , sometimes broadly
e l l i p s o i d , without g e r m pore. B a s i d i a (13-)16-21 X 5.5-7.5 m m , 4 -
31. Agaricus xantholepis (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 2 0 4 . 1 9 5 2 . spored. L a m e l l a edge sterile o r almost sterile, consisting o f cheilocys-
-Fig. 32. tidia and some basidia; c h e i l o c y s t i d i a sometimes i n short chains o f
Psalliota xantholepis F. M 0 l l e r i n F r i e s i a 4: 191. 1952. rounded, rectangular elements o f 5-7 X 4.5-6.5 m m , w i t h clavate ter-
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 58. 1984; F. M 0 l l e r i n F r i e s i a 4 : m i n a l elements o f 15-21.5(-29) X 8-15 m m , often w i t h y e l l o w c o n -
p i . 21c. 1952. tents. P i l e i p e l l i s a cutis o f sometimes branched, 3.0-5.5 m m w i d e
S E L . D E S C R . & F I G S . - F. M 0 l l e r i n F r i e s i a 4: 191. 1952; H u i j s m a n hyphae w i t h c y l i n d r i c a l elements, sometimes constricted at septa, w i t h
i n P e r s o o n i a 1: 3 2 3 , figs. 3-5. 1960; Wasser, Tr. A g a r i c e a e Soviet c y l i n d r i c a l terminal elements, w i t h y e l l o w i s h , parietal and intracellu-
U n i o n : 103-104. 1989. lar pigment, gradually passing into pileitrama. F i b r i l s consisting o f
V E R N . N A M E - Geelschubbige satijnchampignon. similar, curved hyphae with intracellular, brownish pigment.
Stipitipellis a cutis o f 4.0-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l ele-
P i l e u s 40-70(-85) m m , at first h e m i s p h e r i c a l , e x p a n d i n g to convex or ments, w i t h y e l l o w i s h intracellular pigment.
plano-convex, occasionally with umbo, w i t h deflexed margin, H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter-
w h i t i s h to y e l l o w to y e l l o w i s h b r o w n ( M u . 2.5 Y 8/4-6, 10 Y R 8/2, restrial i n m i x e d o r deciduous woods o n calcareous soil, also i n open
7/6), sometimes w i t h darker, b r o w n i s h centre (7.5 Y R 6/5, 8.5 Y R calcareous dunes. V e r y rare i n the Netherlands, occurring i n the c a l -
6 . 5 / 6 , 1 0 Y R 6/6), fibrillose, usually breaking u p into appressed squa- careous dune region and i n southern L i m b u r g (Vogelenzang, H u i s te
mules, especially at centre, d i s c o l o u r i n g b r i g h t l y y e l l o w o n h a n d l i n g ; Vogelenzang; Vogelenzang, A W D ; G r o n s v e l d , Savelsbos). A u g . - O c t .
v e i l present w h e n y o u n g as appendiculate w h i t e remnants. L a m e l l a e , Rare but widespread i n E u r o p e . A l s o k n o w n f r o m A s i a and South
L = 9 0 - 1 0 0 , 1 = 1-7, c r o w d e d , free, subventricose, u p to 8 m m America.
broad, at first grey to p i n k i s h grey (10 Y R 7/2), later g r e y i s h b r o w n
(7.5 Y R 7/2-6/4, 10 Y R 5/2), f i n a l l y b r o w n (5 Y R 4/2-3/3), w i t h paler T h e above description i s also based o n collections f r o m southern
to w h i t e , serrulate edge. Stipe 4 5 - 6 0 X 6-12 m m , annulate, some- E n g l a n d and western Germany.
times w i t h o u t annulus, c y l i n d r i c a l to subclavate o r subbulbous, at Agaricus comtulus Fr. is very close, differing i n smaller size, fib-
base u p to 14(-23) m m broad, rarely tapering towards base, straight rillose covering and white to y e l l o w colours o f the pileus, and the less
to c u r v e d , fistulose, w h i t e , y e l l o w i n g o n h a n d l i n g , at base often bulbous stipe.
strongly d i s c o l o u r i n g y e l l o w to orange-brown, above annulus smooth Agaricus lutosus (F. M 0 l l e r ) F . M 0 l l e r ( i n F r i e s i a 4: 2 0 4 . 1952)
to silky, b e l o w annulus f i b r i l l o s e , sometimes l o c a l l y m i n u t e l y white- seems to differ o n l y i n the presence o f a faint purplish b r o w n tinge i n
squamulose o r floccose-squamulose. A n n u l u s at 0.65-0.75 o f height the squames on the pileus w h e n young, and the stipe w h i c h is tapering
towards the base. It seems l i k e l y that this taxon is very close o r c o n -
specific, considering the variability o f the distinguishing characters.
33. Agaricus porphyrizon P D . O r t o n i n Trans. B r . m y c o l . Soc. 4 3 :

174. 1960. - F i g . 33.
Agaricus arvensis var. purpurascens C o o k e , 111. B r i t . F u n g i 4: p i .
584. 1885; Psalliota purpurascens (Cooke) F. M 0 l l e r i n F r i e s i a 4: 187.
1952; Agaricus purpurascens (Cooke) P i l a t i n A c t a M u s . nat. Prag. 7 B
(1): 10. 1951, non A. purpurascens A. & S. 1805:Fr.
S E L . I C O N . - M . B o n i n D o c . m y c o l . 19(76): p i . 2. 1989; C a p p e l l i ,
A g a r i c u s : p i . 5 1 , 51bis. 1984; D i i h n c k e , 1200 P i l z e : 502. 1993.
S E L . D E S C R . & F I G S . - M . B o n i n D o c . m y c o l . 19(76): 7 6 . 1989; F .
M 0 l l e r i n F r i e s i a 4: 187-188, p i . 21a, 3 5 . 1952 (as Psalliota
purpurascens).
V E R N . N A M E - Purperen champignon.
Pileus (20-)50-90 m m , at first convex w i t h obtuse centre o r truncately

c o n i c a l , expanding to convex, sometimes w i t h depressed centre, o r
plano-convex, at centre greyish b r o w n o r vinaceous b r o w n to purplish
brown ( M u . 10 R 4-5/4, 5/7, 5 Y R 3/3, 7.5 Y R 5/4), covered w i t h fib-
rillose, pointed, appressed, purple squames (10 R 3/3-4, 5/3, 2.5 Y R
5/4), occasionally o n pale, b r o w n i s h background, usually towards mar-
F i g . 32. Agaricus xantholepis. gin more fibrillose; later sometimes purple colours fading and pileus
8 (Jim inflated elements, w i t h pale y e l l o w intracellular pigment, w i t h

slightly ascending, clavate, u p to 9 u-m w i d e terminal elements.
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic, ter-
restrial, o n h u m u s - r i c h sandy s o i l i n deciduous woods, dune scrub,
dune grasslands. Rather c o m m o n i n the coastal dune region, rather rare
in the rest o f the Netherlands. Sept.-Nov. Rather rare but widespread
i n Europe. A l s o k n o w n f r o m A s i a , the A m e r i c a s and northern A f r i c a .
O n account o f the size this taxon is b y some authors placed i n sect.

Arvenses. However, the structure o f the annulus, i n combination w i t h
the strong y e l l o w discoloration, calls for placement i n sect. Minores.
Sect. Xanthodermi Sing.
Pileus not discolouring or sometimes y e l l o w on scratching, disappear-

ing on drying, rarely slightly p i n k ; stipe usually annulate; annulus frag-
ile and thin, at underside near margin thickened or w i t h squames;
surface o f and context i n base o f stipe usually strongly bright y e l l o w
on exposure or w h e n handled, damaged or cut.
K O H - r e a c t i o n i n context o f base o f stipe positive and y e l l o w to
F i g . 33. Agaricus porphyrizon. orange; Schaeffer-reaction o n surface o f pileus and base o f stipe
negative.
entirely reddish b r o w n (2.5 Y R 3/4, 5 Y R 5/4, 7.5 Y R 5/4-6, 10 Y*R Spores (broadly) e l l i p s o i d to o b l o n g , spore length < 6.5 m m o n
4/4); ( s l o w l y ) y e l l o w i n g o n handling, especially at centre; pellis easi- average; Q a v = (1.20-)1.30-1.60(-1.70); without germ pore; l a m e l l a
l y peeled off; margin exceeding lamellae, with appendiculate w h i t i s h , edge sterile; c h e i l o c y s t i d i a catenate or not.
sometimes yellowing, veil remnants, later sometimes incised.
L a m e l l a e , L = 90-120, 1 = ( l - ) 3 - 7 , crowded, free, segmentiform to 33. Agaricus x a n t h o d e r m a Genev. i n B u l l . S o c . bot. Fr. 23: 32. 1876.
subventricose, up to 10 m m broad, p i n k i s h grey at first (5 Y R 7/2; 10 - F i g . 34.
Y R 5/2), later p i n k i s h b r o w n to p i n k i s h grey or greyish p i n k (2.5 Y R Psalliota xanthoderma (Genev.) R i c h o n & R o z e , A t l . C h a m p ,
4-5/2, 5 Y R 4/2, 7.5 Y R 7 / 2 , 1 0 Y R 5/2), finally b r o w n (5 Y R 2.5/2, 7.5 comest. veneneux: 5 3 . 1885. - Psalliota flavescens Richon & Roze,
Y R 3/2), w i t h paler or white, entire or denticulate to crenulate edge. A t l . C h a m p , comest. veneneux: 4 2 , p i . 17, f i g . 17-21. 1885; Agaricus
Stipe (30-)40-80(-110) X 7-11 m m , usually annulate, clavate or c y l i n - pseudocretaceus M . B o n i n D o c . m y c o l . 15(60): 34. 1985. -Agaricus
d r i c a l w i t h subbulbous base, at base up to 19 m m , b r o a d , straight, jodoformicus Speg. i n A n . l v l u s . nac. Hist. nat. B . A i r e s 6: 141. 1899.
stuffed or narrowly fistulose, white, sometimes w i t h reddish tinge, y e l - -Agaricus xanthodermus var. lepiotoides M a i r e i n B u l l , trimest. S o c .
l o w i n g o n handling, especially at base, above annulus squamulose-fib- m y c o l . F r . 24: L V I I I . 1909. - Psalliota xanthoderma var. grisea A .
rillose, below annulus fibrillose or fibrillose-squamulose, later entirely Pears, i n Trans. B r . m y c o l . S o c . 2 9 : 204. 1946; Psalliota grisea ( A .
glabrous a n d smooth, o c c a s i o n a l l y w i t h short white rhizomorphs or Pears.) Essette, Psalliotes: p i . 4 2 . 1964 (invalid); Agaricus xanthoder-
white pseudorhiza. A n n u l u s at (0.40-)0.50-0.65 o f height o f stipe, (3- mus var. griseus ( A . Pears.) B o n & C a p p e l l i i n D o c . m y c o l . 13(52): 16.
)5-10 m m w i d e , descending, pending to patent, thin, easily torn, some- 1983.
times absent, white, w i t h smooth upperside; underside smooth to S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i , 59 & 6 0 . 1984; M a r c h a n d ,
slightly fibrillose, or slightly flocculose at margin, q u i c k l y y e l l o w i n g C h a m p . N o r d M i d i 2: p i . 113. 1973; J . Petersen i n Svampe 28: fig. 2.
on h a n d l i n g . Context 6-8 m m thick i n pileus, soft, white, not dis- 1993.
c o l o u r i n g to d i s c o l o u r i n g pale orange, orange-yellow or y e l l o w i s h S E L . D E S C R . & F I G S . - C a p p e l l i , A g a r i c u s : 313-324. 1984; F. M 0 l l e r
brown, rarely w i t h p i n k i s h blush, i n stipe when cut. S m e l l like almonds in F r i e s i a 4: 168-169. 1952.
w h e n cut, sometimes strongly, o c c a s i o n a l l y w i t h anise-compound. VERN. NAME - Karbolchampignon.
Taste not k n o w n .
M a c r o c h e m i c a l reactions: Schaeffer-reaction strongly orange (sur- Pileus (60-)70-150 m m , at first hemispherical to truncately conical,
face o f pileus and base o f stipe). expanding to convex, plano-convex or plano-concave, with inflexed,
Spores 5.0-6.0 X 3.5-4.0(-4.5) m m , o n average 5.4-5.8 X 3.7-4.2 later reflexed margin, white to cream-coloured ( M u . 2.5 Y 9/2), later
m m , Q = 1.30-1.65, Q a v = 1.35-1.55, ellipsoid, without germ pore. occasionally pale greyish b r o w n to grey-brown (7.5 Y R 7/2, 10 Y R 5/2,
B a s i d i a 2 1 - 2 2 X 6.5-7.0 m m , 4-spored, sometimes also 2-spored 6/2-4/3,6-8/3), at centre often darker, when fresh often brightly chrome-
basidia present. L a m e l l a edge w i t h a 40-75 m m broad sterile layer; y e l l o w o n scratching, at margin sometimes discolouring y e l l o w i s h o n
cheilocystidia not i n chains, sometimes i n clusters, (17.5-)20-33 X 9.5- handling, but colour soon fading away; when young minutely squamu-
13(-20.5) m m , clavate, with colourless or brownish contents. Pileipellis lose at centre, later smooth, soinetimes breaking up into squames, near
a cutis o f 4.0-7.0(-10) m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes margin often radially fibrillose and becoming slightly fissurate; margin
slightly inflated elements, gradually passing into pileitrama, w i t h pale exceeding lamellae, v e i l present when young as brownish patches at cen-
yellow, intracellular pigment. Squames and fibrils consisting o f curved, tre. Lamellae, L = 120-145,1 = 0-3, moderately crowded, free, ventri-
up to 8 m m w i d e hyphae w i t h short, slightly inflated elements, w i t h cose, up to 11 m m broad, at first pinkish to pinkish grey (5 Y R 8/2.5, 7.5
b r o w n i s h intracellular pigment. Stipitipellis a cutis o f sometimes Y R 8/2), later reddish brown (5 Y R 4/3, 7.5 Y R 3/2, 7/2), finally dark
branched, 3.0-5.0 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes up to brown (7.5 Y R 3/4), w i t h concolorous or whitish, entire to minutely den-
AGARICUS (Nauta) 55
a slightly gelatinized cutis o f 3.5-4.5 m m w i d e hyphae w i t h c y l i n d r i -

cal elements, at surface gradually passing into somewhat interwoven,
curved, sometimes branched, 5.0-5.5 m m w i d e hyphae w i t h 16-28 m m
long, irregularly inflated elements and free, u p to 6.5 m m w i d e termi-
nal elements, w i t h pale y e l l o w intracellular pigment.
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic, terrestrial
i n deciduous woods, parks, roadside verges, r i c h grasslands, but also
i n open dunes and dune grasslands o n nutrient-rich or calcareous sandy
or clayey soils. Widespread, rather c o m m o n i n the west and south-east
of the Netherlands, rare i n the rest o f the country. June-Nov.
Widespread i n Europe. C o s m o p o l i t a n .
A l t h o u g h the majority o f the collections has a bulbous stipe, several

specimens with a cylindrical stipe were found i n the dunes. These spec-
imens do not differ i n any other character from the typical ones, there-
fore no taxonomical importance was attached to this feature. T h e variety
lepiotoides M a k e w i t h an areolate pileus is considered without taxo-
nomical meaning and caused by ecological circumstances l i k e drought.
T h e variety griseus, supposedly differing f r o m var. xanthodermus
i n the greyish instead o f w h i t i s h pileus c o u l d not be distinguished.
A l t h o u g h some collections have o n l y white colours, many collections
consist partly o f specimens w i t h a grey pileus.
34. Agaricus pilatianus (Bohus) B o h u s i n A n n l s hist.-nat. M u s . natn.

hung. 66: 78. 1974. - F i g . 3 5 .
Agaricus xanthodermus var. pilatianus B o h u s i n A n n l s hist.-nat.
M u s . natn. hung. 63: 80. 1971.
S E L . I C O N . - C a p p e l l i i n S v a m p e 28: 17. 1993; C a p p e l l i i n B o l l .
G r u p p o m i c o l . G . B r e s . 28: 172. 1985.
S E L . D E S C R . & F I G S . - B o h u s i n A n n l s hist.-nat. M u s . natn. hung.
66: 78-79. 1974.
F i g . 34. Agaricus xanthodermis. (habit X A). VERN. NAME - Kraagchampignon.
ticulate edge. Stipe 60-125 X 10-18 m m , annulate, cylindrical to sub- Pileus 35-40 m m , broadly truncately c o n i c a l , at first white, later pale
clavate, often with a bulbous base up to 30 m m broad, straight to slight- greyish b r o w n w i t h slightly p i n k i s h tinge ( M u . 10 Y R 7/3-4), smooth
ly flexuose or curved, broadly fistulose, shiny and white, with reddish to very m i n u t e l y felted, w h e n y o u n g bright y e l l o w o n scratching.
tinges at lower part o f stipe o n handling, sometimes faintly yellowing, L a m e l l a e very c r o w d e d , free, at first pale b r o w n i s h y e l l o w (10 Y R
above annulus glabrous and smooth, lengthwise striate, sometimes grey- 8/3-4), later reddish b r o w n (5 Y R 4/3), w i t h a concolorous entire edge.
ish, below annulus smooth, sometimes fibrillose; surface at base o f stipe Stipe 35-40 X 7-9 m m , annulate, c y l i n d r i c a l to tapering towards base,
discolouring strongly y e l l o w on scratching; sometimes deeply rooted i n straight, fistulose, white at first, later turning pale y e l l o w i s h b r o w n i n
soil. A n n u l u s at 0.70-0.90 o f height o f stipe, 11-20 m m wide, descend- lower half, slightly y e l l o w i n g o n scratching, above and b e l o w annu-
ing, pending to slightly spreading, rather thin and floppy, near margin lus glabrous and smooth. A n n u l u s at c. 0.7 o f height o f stipe, 5-7 m m
thicker, white, with smooth, sometimes striate upperside; underside fib- wide, descending, patent, thin, white, at m a r g i n double and more or
rillose and near margin w i t h several, large squames. Context 8-15 m m less split i n two, f o r m i n g a k i n d o f triangle i n transverse section; w i t h
thick i n pileus, white, sometimes discolouring faintly y e l l o w when cut,
i n base o f stipe strongly yellow, soon fading. S m e l l faintly unpleasant,
rather strongly carbolic when cut. Taste not k n o w n .
M a c r o c h e m i c a l reactions: K O H 1 0 % y e l l o w (context i n and sur-
face o f base o f stipe), Schaeffer-reaction negative (surface and context
of pileus and stipe).
Spores 4.5-6.5(-7.0) X 3.5-5.5(-6.0) m m , o n average 5.3-6.3 X 3.9-
4.6(-5.1) m m , Q = 1.15-1.45, Q a v = (1.20-)1.30-1.40, broadly ellip-
soid to e l l i p s o i d , without g e r m pore. B a s i d i a 17-21 X 6.0-7.0 m m ,
mostly 4-spored, often also i n part 2-spored. L a m e l l a edge w i t h a c. 5 0
m m broad sterile, often agglutinated layer; cheilocystidia not i n chains,
15-26.5 X 10-19.5(-24) m m , sphaeropedunculate to broadly e l l i p s o i d
or broadly clavate, easily collapsing, hyaline, developed f r o m hyphae
consisting o f l o n g , s l i g h t l y inflated, u p to 5.0 m m w i d e elements.
P i l e i p e l l i s a cutis o f slightly interwoven, sometimes branched, 4.0-9.0
m m w i d e hyphae w i t h c y l i n d r i c a l to sometimes inflated elements up to
14.5 m m w i d e , w i t h ascending, clavate terminal elements, gradually
passing into pileitrama, w i t h pale y e l l o w parietal pigment. Stipitipellis F i g . 35. Agaricus pilatianus. (habit X A).
s m o o t h upperside; underside w i t h pale b r o w n , sometimes split r i m annulus greyish b r o w n , smooth; annulus at 0.50-0.65 o f height o f stipe,
near edge. C o n t e x t white, i n base o f stipe q u i c k l y c h r o m e - y e l l o w 2-3 m m w i d e , descending, pending, moderately thick, w i t h thickened
w h e n cut; c o l o u r fading away soon; elsewhere not discolouring. S m e l l edge, rather persistent, white, w i t h smooth upperside; underside
faintly unpleasant, strongly w h e n cut. Taste very unpleasant. smooth to fibrillose; context white, d i s c o l o u r i n g y e l l o w i n base o f
M a c r o c h e m i c a l reactions: N a O H 3 0 % bright y e l l o w o n base o f stipe, after several minutes reddish; s m e l l faintly carbolic, p h e n o l i c ;
stipe; Schaeffer-reaction negative (surface o f pileus; context). taste not k n o w n .
Spores 5.5-7.0 X 4.0-5.0 m m , o n average 5.8-6.0 X 4.5 m m , Q = M a c r o c h e m i c a l reactions: not k n o w n .
1.25-1.45, Q a v = 1.30-1.35, e l l i p s o i d , sometimes broadly e l l i p s o i d , Spores 5.0-7.0 X ( 3 . 5 - 1 4 ^ = 4 ^ ^ 1 1 1 ^ average 5.7-5.8 X 3.9-4.0
without g e r m pore. B a s i d i a 17-21 X 5.5-6.5 m m , 4-spored. L a m e l l a m m , Q = (1.35-)1.40-1.65(-1.75), Q a v - ^ 1 . 4 5 , e l l i p s o i d , sometimes
edge not entirely sterile; cheilocystidia i n chains o f broadly c y l i n d r i c a l oblong, without germ pore; basidia 4-spored; l a m e l l a edge almost ster-
to clavate elements o f 5.0-17 X 3.5-7.0 m m , w i t h clavate terminal ele- ile; cheilocystidia i n short chains, w i t h clavate terminal elements o f 17-
ments o f 13-22 X 6.5-12 m m . P i l e i p e l l i s a slightly irregular cutis o f 24 X (6-)8-12(-14) m m . P i l e i p e l l i s a slightly irregular cutis o f
2.5-5.5 m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h scattered sometimes branched, 3.5-7.5 m m w i d e hyphae w i t h c y l i n d r i c a l to
slightly ascending, clavate terminal elements o f 13-17 X 5.0-5.5 |xm, somewhat inflated elements up to 9 u,m w i d e , w i t h parietal, pale
gradually passing into pileitrama, w i t h intracellular, very pale y e l l o w - brownish or y e l l o w i s h pigment, gradually passing into pileitrama.
ish pigment. Stipitipellis a cutis o f slightly gelatinized, 3.0-6.0 m m H A B I T A T & D I S T R . - Solitary or i n groups, saprotrophic, terrestrial
w i d e hyphae w i t h sometimes s l i g h t l y inflated elements up to 8 u,m i n gardens, w o o d s or along roadsides o n sandy s o i l . N o t i n the
w i d e , w i t h intracellular, pale y e l l o w i s h pigment. Netherlands. V e r y rare i n E u r o p e , o c c u r r i n g i n E n g l a n d , the B a l k a n
H A B I T A T & D I S T R . - Gregarious, saprotrophic, terrestrial i n m i x e d area and mediterranean countries.
deciduous w o o d o n clayey s o i l or under scrub o n calcareous sandy soil
i n the coastal dune area. V e r y rare i n the Netherlands (Wassenaar, T h e above characteristics are extracted f r o m the description b y B o h u s
M e i j e n d e l ; Grootebroek); Oct. R a r e i n E u r o p e , m a i n l y o c c u r r i n g i n and notes a c c o m p a n y i n g collections f r o m southern E n g l a n d . Var.
eastern and southern regions. niveus B o h u s (in A n n l s hist.-nat. M u s . natn. hung. 72: 95. 1980) is dis-
tinguished f r o m the t y p i c a l variety b y the smooth pileus, w h i c h
T h e above description i s based o n two collections, o f w h i c h o n l y was remains white and does not break up into squames.
p r o v i d e d w i t h m a c r o s c o p i c a l notes. T h e basidiocarps are unusually The species has not yet been recorded from the Netherlands, but the
s m a l l compared to the original description o f B o h u s , and probably rep- t y p i c a l variety i s k n o w n f r o m southern E n g l a n d . T h e record f r o m
resent a s m a l l variant o f this species. In the o r i g i n a l description the S w e d e n ( K n u d s e n i n Jordstjarnan 18: 36-39. 1997) is based o n a
basidiocarps have a pileus diameter o f 60-120 m m , and a stipe o f 4 5 - misidentification (pers. c o m m . H . K n u d s e n , Copenhagen).
80 X 20-30 m m (Bohus i n A n n l s hist.-nat. M u s . natn. hung. 6 3 : 80. Because o f the habit it can be confused w i t h A . pilatianus or species
1971). D e c i s i v e characters are the b r o w n i s h - p i n k i s h discoloration o f i n the sect. Spissicaules. It differs from sect. Spissicaules i n the more
the pileus o n handling, the partly fertile l a m e l l a edge w i t h tufts o f nar- sturdy annulus w i t h thicker margin.
row, clavate cheilocystidia w h i c h are catenate, and the c y l i n d r i c a l stipe
w i t h a triangular annulus. 36. Agaricus moelleri Wasser i n N o v . Sist. v y s s h . n i z s h . Rast. 13: 7 7 .
1 9 7 6 . - F i g . 37.
35. Agaricus pseudopratensis (Bohus) Wasser i n U k r . bot. Z h . 33: Agaricus xanthodermus var. obscuratus M a i r e i n B u l l , trimest. Soc.
250. 1 9 7 6 . - F i g . 36. m y c o l . Fr. 26: 192. 1910; Psalliota meleagris var. obscurata (Maire)
Psalliota pseudopratensis B o h u s i n B o r b a s i a 1: 114. 1939; F. M 0 l l e r i n F r i e s i a 4: 173. 1952; Agaricus meleagris var. terricolor F.
Agaricus pseudopratensis (Bohus) B o h u s i n A n n l s hist.-nat. M u s . natn. M0ller i n Friesia 4: 2 0 8 . 1952 ( n o m . illeg.); Agaricus prae-
hung. 6 3 : 8 L 1971 (invalid). - Agaricus pseudopratensis var. niveus claresquamosus var. terricolor (F. M 0 l l e r ) B o n & C a p p e l l i i n D o c .
B o h u s i n A n n l s hist.-nat. M u s . natn. hung. 72: 94. 1980. m y c o l . 13(52): 16. 1983. - Psalliota meleagris J . Schaeff. i n Z . P i l z k .
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 6 8 . 1984; C a p p e l l i i n B o l l . 4: 28. 1925; Agaricus meleagris (J. Schaeff.) Imbach i n M i t t , naturf.
G r u p p o m i c o l . G . B r e s . 2 8 : 180. 1985. Ges. L u z e r n 15: 1 5 . 1 9 4 6 , n o n A . meleagris Sow. 1798; Agaricusprae-
S E L . D E S C R . & FIGS. - B o h u s i n A n n l s hist.-nat. M u s . natn. hung. claresquamosus A . E . F r e e m a n i n M y c o t a x o n 8: 9 0 . 1979 (superflu-
63: 81-82. 1971. ous); Agaricus moelleri var. meleagris (J. Schaeff.) Rauschert i n N o v a
H e d w i g i a 54: 214. 1992.
C H A R A C T E R I S T I C S - P i l e u s 25-80 m m , convex w i t h flattened centre, MISAPPL.- Agaricus placomyces sensu M o s . , Rohrlinge
w h i t i s h or greyish white w i t h a b r o w n centre, soon breaking up into Blatterpilze, 4. A u f L : 234. 1978.
large, appressed brownish to reddish b r o w n squames at centre, towards S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 62. 1984; J . Petersen i n Svampe
margin more fibrillose-squamulose; lamellae free, up to 4 m m broad, 28: f i g . 7. 1993 (as A. praeclaresquamosus); R . P h i l l i p s , Paddest.
at first p i n k i s h , later dark b r o w n ; stipe 20-65 X 7-15 m m , annulate, S c h i m m . : 169. 1981 (as A . meleagris).
c y l i n d r i c a l or clavate, sometimes tapering towards base, whitish, above S E L . D E S C R . & F I G S . - C a p p e l l i , A g a r i c u s : 325-332. 1984 (as A.
praeclaresquamosus); F. M 0 l l e r i n F r i e s i a 4: 172-173. 1952 (as P.
meleagris).
VERN. NAME - Parelhoenchampignon.
Pileus (50-)60-120(-140) m m , at first convex, expanding to plano-con-

vex to applanate, sometimes w i t h depressed centre, mostly w i t h umbo,
w i t h at first involute, later straight margin, densely covered w i t h minute,
appressed, brown to dark greyish b r o w n fibrillose squamules o n a white
to later pale b r o w n or light greyish b r o w n background ( M u . 7.5 Y R 8/2),
F i g . 36. Agaricus pseudopratensis. at centre u n i f o r m l y b r o w n or y e l l o w i s h b r o w n to dark grey-brown or
AGARICUS (Nauta) 57
of sometimes branched, 3.5-9.5(-17) m m w i d e hyphae w i t h cylindrical

to somewhat inflated elements, w i t h slightly ascending, c y l i n d r i c a l ter-
m i n a l elements, gradually passing into pileitrama, w i t h intracellular,
pale brown pigment. Squames consisting o f curved, 7.5-14.5 m m wide
hyphae with 17-43 m m long elements and clavate, up to 14 m m wide
terminal elements, with intracellular dark b r o w n pigment. Stipitipellis
a sometimes irregular cutis o f occasionally gelatinized, 4.0-8.0 m m
wide hyphae with frequently slightly inflated elements, and somewhat
protruding, curved cylindrical terminal elements.
H A B I T A T & DISTR. - U s u a l l y gregarious, saprotrophic, terrestrial, i n
m i x e d deciduous woods o n humus-rich, calcareous soil. F a i r l y c o m -
mon i n the calcareous dunes and i n southern L i m b u r g , very rare i n the
rest o f the Netherlands. M a y - N o v . Widespread but rare i n Europe. A l s o
recorded from the A m e r i c a s , A s i a and northern A f r i c a .
This species has been confused i n the past w i t h the A m e r i c a n species

A. placomyces P e c k . A c c o r d i n g to F r e e m a n ( i n M y c o t a x o n 8: 9 0 .
1979) the latter concerns a different species without cystidia.
The species was o r i g i n a l l y described by Schaeffer as h a v i n g a usu-
ally b l a c k squamose, sometimes b r o w n squamose, pileus covering.
M 0 l l e r described the dark variant as a n e w variety, var. obscuratus,
based on Agaricus xanthodermus var. obscurata M a i r e , and changed it
later into var. terricolor. T h e dark variant was for nomenclatural rea-
sons renamed as A. moelleri, necessitating a new name for the b r o w n
variety, v i z . var. meleagris.
In the D u t c h specimens no two such forms c o u l d be recognized.
F i g . 37. Agaricus moelleri. (habit X W).
37. Agaricus phaeolepidotus (F. M 0 l l e r ) F. M 0 l l e r i n F r i e s i a 4: 204.
1 9 5 2 . - F i g . 38.
blackish b r o w n (5 Y R 2.5/2, 7.5 Y R 4/2, 10 Y R 3/1-2, 4/3-3/2, 5/4), Psalliota phaeolepidotus F. M 0 l l e r i n F r i e s i a 4: 170-172. 1952. -
rarely slightly y e l l o w i n g when rubbed; v e i l present when young as nar- Agaricus meleagris var. perdicinus Pilat i n A c t a M u s . nat. Prag. 7 B (1):
row white fringe at margin. Lamellae, L = 135-150,1 = 1-3, crowded, 108. 1951 (invalid); Agaricus perdicinus Pilat i n A c t a M u s . nat. Prag.
free, segmentiform to subventricose, up to 10 m m broad, at first whitish 9 B (2): 24. 1953.
to pale greyish pink, later pinkish b r o w n to brown (7.5 Y R 4 / 2 - 4 , 1 0 Y R S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 64. 1984; F. M 0 l l e r i n Friesia
5/3), finally greyish chocolate-brown, with serrulate, whitish edge. Stipe 4: p i . 20b. 1952; J . Petersen i n Svampe 28: fig. 8. 1993.
(50-)60-115(-140) X 7-15(-18) m m , annulate, clavate to cylindrical with S E L . D E S C R . & FIGS. - F. M 0 l l e r i n F r i e s i a 4: 170-172, p i . 32. 1952;
an abruptly bulbous base, at base up to 22 m m broad, straight to slight- Pilat i n A c t a M u s . nat. P r a g . 9 B (2): 24-28, f i g . 30-34. 1953 (as A.
l y curved near base, fistulose, dirty white, at base strongly y e l l o w i n g on perdicinus).
handling, sometimes o n l y y e l l o w i s h b r o w n , later w i t h pinkish brown VERN. NAME - Hazelhoenchampignon.
spots, above annulus glabrous, shiny, smooth, sometimes finely striate,
below annulus glabrous and smooth, at base slightly fibrillose, some- Pileus 55-120 m m , at first truncately c o n i c a l to convex, expanding to
times tomentose, sometimes with pseudorhiza. A n n u l u s at 0.75 - 0.90 o f plano-convex, sometimes w i t h slightly depressed centre o r l o w umbo,
height o f stipe, 7-12 m m wide, descending, pending to slightly patent, w i t h deflexed margin, at centre u n i f o r m l y light b r o w n to b r o w n ( M u .
thin to thick w i t h mostly thickened margin, white, with smooth or often 7.5 Y R 5-6.5/4, 5.5/3, 6/4), sometimes reddish b r o w n (2.5 Y R 2.5/4),
striate upperside; underside smooth to fibrillose, thickened near margin with appressed, pale b r o w n fibrillose squames o n a white to isabella
or, occasionally, w i t h some radially arranged, sometimes brownish
squames near the margin. Context 6-11 m m thick i n pileus, white, slight-
ly discolouring y e l l o w when cut, later with pale reddish b r o w n patches,
especially i n pileus, i n base o f stipe slightly to strongly chrome-yellow.
S m e l l varying from weak to strong, unpleasant: like metal, ink, i o d -
oform, carbolic. Taste slightly bitter or sour, unpleasant.
M a c r o c h e m i c a l reactions: K O H 10% & 4 0 % (bright) y e l l o w (con-
text o f stipe; surface o f pileus); Schaeffer-reaction negative (surface o f
pileus; surface o f base o f stipe).
Spores 4.5-6.5 X 3.0-4.0 m m , o n average 5.4-5.6(-6.1) X 3.5-3.7
m m , Q = 1.40-1.90, Q a v = 1.50-1.60(-1.70), ellipsoid to oblong, with-
out germ pore. B a s i d i a 14-20 X 4.5-6.5 m m , mostly 4-spored, occa-
sionally a m i n o r i t y o f 2-spored basidia present. L a m e l l a edge w i t h a
75-105 m m broad, almost sterile layer; cheilocystidia often i n slightly
agglutinated chains o f subglobose elements o f 6-14 X 5-8 m m , w i t h
globose to clavate, hyaline terminal elements o f 11-23 X 7-15 m m ,
occasionally w i t h brownish contents. Pileipellis a slightly irregular cutis F i g . 38. Agaricus phaeolepidotus. (habit X A).
b a c k g r o u n d (7.5 Y R 3/4, 8/2, 10 Y R 6/4), towards m a r g i n more Spores (broadly) e l l i p s o i d to oblong, Q a v (1.30-)1.35-1.60, w i t h -
m i n u t e l y squamulose-fibrillose, not d i s c o l o u r i n g on handling, finally out germ pore; l a m e l l a edge sterile or fertile; cheilocystidia either glo-
at m a r g i n sometimes w i t h p i n k i s h tinge; v e i l sometimes present w h e n bose to clavate or inconspicuous (basidioliform), sometimes scarce.
y o u n g as pale, fibrillose squamules. L a m e l l a e , L = (105-)135-160,1
= 1-3, c r o w d e d , free, segmentiform to ventricose, 7-10 m m broad, at 38. Agaricus litoralis (Wak. & Pears.) Pilat, K l i c urc. H u b hfib. bedl.:
first p i n k i s h w h i t e or pale p i n k i s h grey (5 Y R 8/2, 7/3), later dark 403. 1 9 5 1 . - F i g . 3 9 .
b r o w n to chocolate-brown (7.5 Y R 4/2, 3/4), w i t h entire to denticulate, Psalliota litoralis W a k . & Pears, i n Trans. B r . m y c o l . Soc. 29: 206.
often w h i t i s h edge. Stipe (45-)60-130 X 7-14 m m , annulate, c y l i n d r i - 1946. - Psalliota spissa F. M 0 l l e r 4 r i f t i e s i a 4: 53. 1950; Agaricus spis-
cal to clavate, often w i t h bulbous base w i t h flattened underside, at sicaulis F. M 0 l l e r i n Friesia 4: 203. l$52r~~Âgaricus maskae Pilat i n
base u p to 27 m m broad, straight to curved, fistulose, white, later w i t h C e s k a M y k o l . 8: 165. 1954. - Agaricus maskae~*vai. imrehii B o h u s i n
pale b r o w n or p i n k i s h grey tinges, especially at apex, b r o w n i s h o n A n n l s hist.-nat. M u s . natn. hung. 66: 83. 1974.
scratching, sometimes pale y e l l o w at base, above annulus silky, S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 37 (as A. spissicaulis), p i . 38 (as
smooth, b e l o w annulus lengthwise fibrillose to satiny striate, shiny, A. maskae). 1984; D a n i e l - A r r a n z i n B o l . S o c . m i c o l . M a d r i d 22: 148,
smooth, at base sometimes w i t h dispersed w h i t i s h fibrillose squames; fig. 5. 1997 (as A. maskae); F. M 0 l l e r i n F r i e s i a 4: p i . 3b. 1950 (as P.
sometimes w i t h w h i t i s h r h i z o m o r p h s . A n n u l u s at (0.65-)0.80-0.90 o f spissa); Wasser i n F u n g . rar. Ic. c o l . 10: p i . 7 5 . 1979 (as A. spissi-
height o f stipe, 10-24 m m w i d e , descending, p e n d i n g to slightly caulis).
spreading, moderately thin, sometimes torn, white, sometimes dis- S E L . D E S C R . & FIGS. - C a p p e l l i , A g a r i c u s : 4 5 4 - 4 5 6 . 1 9 8 4 (as A . spis-
c o l o u r i n g y e l l o w , w i t h smooth to m i n u t e l y striate upperside; under- sicaulis and A. maskae resp.); F. M 0 l l e r i n F r i e s i a 4: 5 3 . 1950 (as P.
side w i t h some, often b r o w n i s h , rectangular, r a d i a l l y arranged spissa); Wasser i n F u n g . rar. Ic. c o l . 10: 7-8. 1979 (as A. spissicaulis).
squames at m a r g i n , o c c a s i o n a l l y arranged as c o g w h e e l , often w i t h V E R N . N A M E - Plompe champignon.
b r o w n i s h exudate. Context 7-8 m m thick i n pileus, soft, white, not dis-
c o l o u r i n g or turning s l o w l y b r o w n i s h i n some patches w h e n cut, i n Pileus (50-)60-125(-135) m m , at first hemispherical to convex, expand-
base o f stipe at first pale y e l l o w , sometimes hardly d i s c o l o u r i n g , later ing to irregularly convex or plano-convex, with flattened or sometimes
s l i g h t l y b r o w n i s h to w h i t i s h b r o w n or b r o w n i s h w i t h p i n k i s h tinge! depressed centre, w i t h margin young incurved, later largely appendic-
S m e l l w h e n c u t c l e a r l y i n k - l i k e , to w e a k l y carbolic, or l i k e canned ulate and exceeding lamellae, whitish, greyish white or pale brown ( M u .
apple-sauce, disagreeable. Taste fungoid. 10 Y R 6/3, 8/4, 9/2), occasionally w i t h y e l l o w i s h or b r o w n i s h spots,
M a c r o c h e m i c a l reactions: K O H 10% y e l l o w to orange (especially often slightly y e l l o w i s h {2,5 Y 8/4, 8/5; K . & W . 2 A 6 ) on handling, fib-
at base o f stipe; also often at surface and context); Schaeffer-reaction rillose, w i t h often faintly delimited, large, light grey to greyish b r o w n
negative (at a l l parts o f basidiocarp). or y e l l o w i s h b r o w n appressed fibrillose squames at centre; veil present
Spores 5.0-6.0 X 3.5-4.5 m m , o n average 5.3-5.7 X 3.7-4.0 m m , Q w h e n y o u n g as w h i t i s h fringe or arachnoid flocks at margin, later as
= 1.30-1.70, Q a v = 1.40-1.50, ellipsoid, sometimes oblong, without appendiculate remnants. L a m e l l a e , L = 9 0 - 1 2 0 , 1 = 1-3, moderately
germ pore. B a s i d i a 14-16 X 6.5-8.0 m m , 4-spored, sometimes w i t h a crowded, free, arcuate to undate, (5-)6-10 m m broad, at first p i n k or
few 2-spored basidia. L a m e l l a edge w i t h a 75-85 m m broad, almost pinkish b r o w n to pale b r o w n with p i n k i s h tinge or brownish grey (10
sterile layer; c h e i l o c y s t i d i a h y a l i n e , i n short chains o f 2-3, broadly Y R 5/3, 7/3), only later dark b l a c k i s h b r o w n (7.5 Y R 4/2-3/2, 10 Y R
c y l i n d r i c a l to subglobose elements o f 8.5-13.0 X 7.0-8.0 m m , w i t h 2/1), with paler, entire edge. Stipe (25-)35-60(-70) X ( l l - ) 1 4 - 1 9 ( - 2 3 )
clavate, o b o v o i d to globose or c y l i n d r i c a l terminal elements o f 15-43 mm, annulate, fusoid to irregularly c y l i n d r i c a l , tapering at base, at
X (6.5-)8.0-20(-23) m m . P i l e i p e l l i s a cutis o f 4.5-6.5 m m w i d e hyphae widest halfway u p to 28 m m broad, slightiy curved, stuffed to narrow-
w i t h c y l i n d r i c a l , sometimes slightly inflated, hyaline elements, gradu- ly fistulose, white, at base occasionally faintly ochraceous, often w i t h
ally passing into pileitrama. Squames consisting o f interwoven, 5.5-9.5 b r o w n i s h tinge, sometimes finally entirely slightly b r o w n i n g o n han-
m m w i d e hyphae w i t h 25-52 m m l o n g , inflated elements and ascend- dling, above annulus minutely fibrillose-striate and sometimes greyish
ing, c y l i n d r i c a l terminal elements, w i t h parietal pale y e l l o w i s h b r o w n or brownish, below annulus merely fibrillose to lanate-fibrillose, with
pigment, gradually passing into pileipellis. Stipitipellis a slightly gelat-
i n i z e d cutis o f 4.5-9.0(-10.0) m m w i d e hyphae w i t h c y l i n d r i c a l , slight-
ly inflated elements.
H A B I T A T & DISTR. - Gregarious, saprotrophic, terrestrial i n decidu-
ous woods o n nutrient-rich, calcareous clayey or sandy soil. Rare, pre-
dominantly i n the river region, southern L i m b u r g and the inner dune
area. A u g . - O c t . Widespread, but rare i n Europe. A l s o k n o w n from A s i a
and southern South A m e r i c a .
T h e description is partly based o n collections from D e n m a r k .
Sect. Spissicaules (Heinem.) Kerrigan
Pileus often slightly y e l l o w i n g ; annulus relatively thin, evanescent or

relatively persistent, underside smooth or flocculose; base o f stipe with
y e l l o w i n g rhizomorphs; context i n base o f stipe usually yellow, else-
where usually faintly reddish.
K O H - r e a c t i o n negative; Schaeffer-reaction o n surface o f pileus
negative, usually orange o n base o f stipe. F i g . 39. Agaricus litoralis. (habit X A).
AGARICUS {Nauta) 59
1-3, thick, white, y e l l o w d i s c o l o u r i n g (10 Y R 8/8) rhizomorphs.

A n n u l u s at 0.50-0.60(-0.80) o f height o f stipe, 3-5 m m wide, descend-
ing, pending, thin, easily torn, simple, white, not discolouring, w i t h
smooth to slightly striate upperside; underside smooth to fibrillose.
Context (8.5-)13-20(-24) m m thick i n pileus, relatively firm, white,
often w i t h brownish tinge, sometimes w i t h a reddish tinge, occasional-
ly s l o w l y and faintly discolouring to faintly p i n k i s h b r o w n to brownish-
reddish o r slightly orange-red, i n base o f stipe p i n k i s h orange to
b r o w n i s h when cut, i n pileus faintly brownish o r greyish. S m e l l faint,
often indistinct, sometimes like anise, o f context w h e n freshly cut
slightly l i k e almonds, later faintly unpleasant. Taste faintly like nuts.
Macrochemical reactions: K O H 1 0 % negative ( a l l parts);
Schaeffer-reaction usually negative (surface o f pileus and stipe), some-
times slightly orange (surface o f pileus), o r orange (surface o f base o f
stipe; rhizomorphs).
Spores (6.5-)7.0-8.5 X (4.5-)5.0-6.5 m m , o n average (7.1-)7.4-8.0
X (5.2-)5.5-5.8 m m , Q = (1.15-)1.20-1.55(-1.65), Q a v = 1.35-1.45,
e l l i p s o i d to broadly ellipsoid, without g e r m pore. B a s i d i a (21-)23-28(-
31) X 7 . 0 - 1 0 . 0 ( - l l ) m m , usually 4-spored, sometimes also 2-spored
present. L a m e l l a edge not entirely sterile, usually composed o f basidia
and a m i n o r i t y o f usually inconspicuous b a s i d i o l i f o r m clavate c h e i l o -
cystidia o f (14-)16.5-22(-28) X (4.5-)6-10(-13) u,m; sometimes l a m e l -
la edge almost sterile; rarely w i t h conspicuous, clavate c h e i l o c y s t i d i a F i g . 40. Agaricus lanipes. (habit X Vi).
up to 13 |x,m w i d e . P i l e i p e l l i s a cutis o f 4.0-5.5 m m w i d e hyphae w i t h
c y l i n d r i c a l elements, w i t h slightly erect, clavate terminal elements o f
18-24 X 5.5-6.5 |xm, gradually passing into pileitrama, w i t h pale y e l - discolouring o n handling; v e i l present w h e n y o u n g as w h i t i s h appen-
low, parietal pigment. Stipitipellis a slightly irregular cutis o f 3.0-6.5 diculate fringe. L a m e l l a e , L = 80-120, 1 = 1-5, crowded, free, sub-
m m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h curved, u p to 8 m m ventricose to undate, up to 9.5 m m broad, at first beige to p i n k i s h grey
wide, c y l i n d r i c a l terminal elements; w i t h y e l l o w i s h parietal pigment. (2.5 Y R 6/3, 7.5 Y R 7/2-4, 10 Y R 7/3), later b l a c k i s h b r o w n (7.5 Y R
H A B I T A T & DISTR. - Solitary o r i n groups, saprotrophic, terrestrial, 3/2, 4/2), w i t h w h i t i s h serrulate to denticulate edge. Stipe 40-70(-85)
often i n d r y pastures, or lawns o n sandy s o i l , i n mossy areas i n d r y X (8-) 10-18 m m , annulate, broadly clavate, at base up to 29 m m broad,
coastal dunes; sometimes along roads i n grass. W i d e s p r e a d and rather straight to slightly curved, stuffed, often y e l l o w o r b r o w n i s h y e l l o w at
rare i n the Netherlands, J u n e - N o v . W i d e s p r e a d and rare i n E u r o p e , base o n handling, sometimes s l o w l y pale b r o w n i s h red o n scratching,
probably more c o m m o n i n the B a l k a n area. A l s o k n o w n f r o m N o r t h usually elsewhere not d i s c o l o u r i n g o n handling, above annulus white
A m e r i c a and northern A f r i c a . to later greyish, smooth to very m i n u t e l y squamulose, b e l o w annulus
velutinous or fibrillose to floccose, w h i t i s h , towards base usually with
Agaricus litoralis is considered s y n o n y m o u s w i t h A. maskae and A. some girdles o f brownish, fibrillose, appressed squames, often at base
spissicaulis, o n account o f studies o f the type and o r i g i n a l material smooth again; sometimes entire surface o f stipe brownish (5 Y R 6/3,
(Nauta i n Persoonia 17: 227. 1999). 7/3; 7.5 Y R 6/4, 10 Y R 6/3); usually w i t h one or a few short to long
For an extensive d i s c u s s i o n o n the s y n o n y m y see N a u t a ( i n whitish, y e l l o w i n g rhizomorphs. A n n u l u s at (0.60-)0.70-0.75 o f height
Persoonia 17: 227. 1999). of stipe, (5-)10-18 m m wide, descending, slightly spreading to patent,
thin, relatively persistent, w h i t i s h to y e l l o w i s h b r o w n (10 Y R 6/4), with
39. Agaricus lanipes ( M 0 l l e r & Schaeff.) S i n g , i n L i l l o a 2 2 : 4 3 2 . smooth to striate upperside; underside floccose to squamose o r lanate.
('1949') 1 9 5 1 . - F i g . 40. Context 6-9 m m thick i n pileus, w h i t i s h to pale b r o w n , hardly dis-
Psalliota lanipes M 0 l l e r & Schaeff. i n A n n l s m y c o l . 36: 65. 1938. colouring to d i s c o l o u r i n g s l o w l y and slightly reddish o r b r o w n when
-Agaricus luteolorufescens P . D . O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : cut i n upper part o f stipe and pileus, occasionally y e l l o w i s h i n base o f
182. 1960. - Psalliota lanipes var. verecunda F. M 0 l l e r i n F r i e s i a 4: stipe. S m e l l like nuts to somewhat like almonds w h e n cut, especially
26. 1950; Agaricus lanipes var. verecundus (F. M 0 l l e r ) F. M 0 l l e r i n i n base o f stipe. Taste not k n o w n .
F r i e s i a 4: 203. 1952. M a c r o c h e m i c a l reactions: K O H 10% negative (surface o f pileus) or
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 2 3 . 1984; C a p p e l l i i n B o l l . y e l l o w i s h (base o f stipe), N a O H negative (surface o f pileus) or b r o w n -
G r u p p o m i c o l . G . B r e s . 26: 2 4 . 1983. ish to y e l l o w i s h (context i n base o f stipe); Schaeffer-reaction negative
S E L . D E S C R . & FIGS. - C a p p e l l i , A g a r i c u s : 188-189. 1984; F. M 0 l l e r (surface o f pileus and stipe) o r sometimes orange (surface o f base o f
i n F r i e s i a 4: 25-26, p i . 10. 1950; N a u t a i n Persoonia 17: 222-224, fig. stipe), usually i n connection w i t h a y e l l o w discoloration there.
1. 1999. Spores 5.5-8.0 X 3.5-5.5 m m , o n average (5.8-)6.6-7.1 X (3.6-)4.4-
V E R N . N A M E - Breedschubbige champignon. 4.7(-5.0) m m , Q = 1.30-1.75(-2.00), Q a v = (1.40-) 1.45-1.60, ellipsoid
to oblong, without germ pore. B a s i d i a 16-17.5 X 5.5-6.5 m m , usually
Pileus 50-100(-140) m m , at first convex, expanding to plano-convex, 4-spored, often some 2-spored also present. L a m e l l a edge w i t h a 75-80
often w i t h depressed centre, w i t h m a r g i n exceeding lamellae, w h e n m m broad sterile layer; cheilocystidia usually i n short chains o f more
y o u n g evenly b r o w n and fibrillose, soon i n centre densely dark b r o w n or less rounded rectangular elements o f 9-13 X 6 m m , w i t h globose to
fibrillose-squamose ( M u . 6 Y R 5/3, 7.5 Y R 5/4, 10 Y R 3-4/2), towards clavate hyaline terminal elements o f (12-)14-30 X 9-16(-22) m m .
m a r g i n w i t h broad, appressed, fibrillose, b r o w n to y e l l o w i s h b r o w n Pileipellis a cutis o f 4.0-9.5 m m w i d e hyphae w i t h c y l i n d r i c a l , slightly
squames (6 Y R 4-5/3, 10 Y R 4-7/4, 6/3) o n a paler background, not inflated elements, gradually passing into pileitrama and into squames,
w i t h parietal pigment; squames consisting o f curved, up to 11 m m w i d e later w i t h y e l l o w i s h spots; v e i l present w h e n y o u n g as appendiculate

hyphae w i t h 25-35 m m l o n g elements, sometimes w i t h slightly thick- fringe, later disappearing. L a m e l l a e , L = 80-95, 1 = 1-3, crowded,
ened walls, and c y l i n d r i c a l , up to 11 m m wide terminal elements, w i t h free, up to 10 m m broad, at first greyish p i n k , finally b l a c k i s h b r o w n ,
parietal pigment. Stipitipellis a cutis o f slightly gelatinized, 3.5-4.5 m m w i t h concolorous, entire edge. Stipe 30-70(-80) X 7.5-16(-20) m m ,
w i d e hyphae w i t h c y l i n d r i c a l elements w i t h parietal pigment; squames annulate, clavate to c y l i n d r i c a l w i t h bulbous base, at base up to 30 m m
consisting o f curved, 4.5-6.5 m m wide hyphae with 12-28.5 m m long, broad, stuffed, straight, white, d i s c o l o u r i n g y e l l o w to y e l l o w i s h b r o w n
slightly inflated elements and slightly clavate, up to 11 m m w i d e ter- at base, above annulus striate, b e l o w annulus fibrillose to fibrillose-
m i n a l elements, with parietal and sometimes intracellular pigment. squamulose, w i t h thick y e l l o w i n g pseudorhiza. A n n u l u s at 0.65-0.75
H A B I T A T & D I S T R . - Gregarious to sometimes caespitose, sapro- o f height o f stipe, narrow, c. ^^nrnjwide^descending, pending, simple,
trophic, terrestrial i n m i x e d woods or gardens o n nutrient-rich sandy or thin, often evanescent, white, w i t h striatelrpperside; underside smooth.
c l a y e y s o i l . Rather rare i n the Netherlands, restricted to regions o n Context 6-11 m m thick i n pileus, white, d i s c o l o u r i n g faintly reddish
nutrient-rich soil as the new polders, southern L i m b u r g , river region w h e n cut, i n base o f stipe faintly y e l l o w . S m e l l indistinct o r sometimes
and the western part o f the country. A u g . - Oct. Rare but widespread i n faintly like iodine. Taste not k n o w n .
the rest o f E u r o p e . M a c r o c h e m i c a l reactions: Schaeffer-reaction negative (surface o f
pileus), orange (surface o f base o f stipe).
M 0 l l e r ( i n F r i e s i a 4: 2 6 . 1950) described var. verecunda as differing Spores 5.5-7.5 X 4.0-5.0(-5.5) m m , o n average 6.0-6.7 X 4.2-4.7
f r o m the t y p i c a l variety i n the more gracile habit w i t h a pileus up to 8 m m , Q = 1.20-1.65(-1.75), Q a v =~(1.30-)1.40-1.50, ellipsoid, some-
c m i n diameter, the more diluted b r o w n squames, the stipe w i t h pale times broadly e l l i p s o i d o r oblong, without germ pore. B a s i d i a 17-25
squames instead o f b r o w n i s h b e l o w the ring, and the annulus w h i c h i s X 7.0-9.0 m m , 4-spored, sometimes some 2-spored. L a m e l l a edge usu-
completely white at the underside. These characters are not considered ally fertile, w i t h a variable amount o f sterile elements; cheilocystidia
sufficient for distinguishing a taxon, therefore the name is i n c l u d e d i n usually inconspicuous and resembling y o u n g basidia, 13-25 X 7-11(-
the synonyms. 13) m m , usually clavate, rarely more globose, sometimes i n very short
Agaricus luteolorufescens appeared to be conspecific w i t h A. chains o f rectangular elements o f 3.5-7.0 X 3.0-5.0 |xm, hyaline, rarely
lanipes (Nauta i n Persoonia 17: 224. 1999). Agaricus vinosobrunneus w i t h faintly b r o w n i s h contents. P i l e i p e l l i s a regular cutis o f r a d i a l l y
P.D. O r t o n (in Trans. B r . m y c o l . S o c . 4 3 : 183. 1960) is, according to arranged, 4.5-7.0 m m w i d e hyphae w i t h c y l i n d r i c a l , sometimes inflat-
O r t o n , closely related to A. lanipes, but proved to be conspecific w i t h ed u p to 8.5 p-m, elements, w i t h clavate, slightly ascending terminal
A. sylvaticus. elements, gradually passing into pileitrama, w i t h parietal pale y e l l o w
pigment. Stipitipellis a cutis o f sometimes curved, 4-8.5 m m w i d e
40. Agaricus bresadolanus B o h u s i n A n n l s hist.-nat. M u s . natn. hung. hyphae w i t h c y l i n d r i c a l elements, w i t h pale yellow, parietal pigment.
61: 154. 1969 (as A . bresadolanus). - F i g . 41. H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic, ter-
Agaricus campestris b radicatus Vitt., Descr. F u n g h i mang. Italia: restrial along paths or o n grassy spots i n deciduous woods o r i n gar-
42. 1835; Agaricus radicatus (Vitt.) R o m a g n . i n B u l l , trimest. S o c . dens, sometimes i n meadows o n nutrient-rich and calcareous soil. Rare
m y c o l . Fr. 5 3 : 129. 1938, non A. radicatus R e l h . 1785:Fr.; Agaricus i n the Netherlands, Apr., A u g . - N o v . Rare i n the rest o f Europe, proba-
romagnesii Wasser i n U k r . bot. Z h . 34: 305. 1977. - Psalliota infida bly not o c c u r r i n g i n northern E u r o p e . A l s o recorded from A s i a .
A l e s s i o i n M i c o l . ital. 4 ( 2 ) : 2 1 . 1975 (invalid); Agaricus infidus
(Alessio) M . B o n i n D o c . m y c o l . 11(44): 28. 1981 (invalid); A . romag- T h e description i s based o n collections from Great B r i t a i n , Hungary,
nesii var. infidus ( A l e s s i o ) B o n & C a p p e l l i i n D o c . m y c o l . 13(52): 16. the Netherlands and the type description b y B o h u s .
1983 (invalid). For a discussion o n the s y n o n y m y see N a u t a (in Persoonia 17: 229.
S E L . I C O N . - C a p p e l l i , A g a r i c u s : p i . 14, 70 (as A . romagnesii). 1984; 1999).
C a p p e l l i i n B o l l . G r u p p o m i c o l . G . B r e s . 28: 184. 1985 (as A . romag-
nesii); D . R e i d i n F u n g . rar. Ic. c o l . 6: p i . 42. 1972.
S E L . D E S C R . & F I G S . - D . R e i d i n F u n g . rar. Ic. c o l . 6: 6-8, f i g . 9.
1972; N a u t a i n Persoonia 17: 228, f i g . 4. 1999.
Subgen. Lanagaricus Heinem. emend. A . E .
V E R N . N A M E - Wortelende champignon. Freeman
Pileus 35-85(-100) m m , convex to plano-convex, sometimes w i t h V e i l c o m p o s e d o f hyphae w i t h m a i n l y inflated elements up to 11 | x m

depressed centre, at first white, later greyish white w i t h light b r o w n w i d e , intermixed w i t h loosely connected subglobose elements o f 10-
centre ( M u . 10 Y R 5/3, 6/5), fibrillose, later at centre w i t h indistinct 20 X 10-17 w,m.
darker greyish b r o w n , lanate squames; slightly y e l l o w i n g o n handling,
T h e d i v i s i o n o f Agaricus into subgenera requires further study, prefer-
ably o f m o r p h o l o g i c a l and m o l e c u l a r characters. T h e interpretation o f
F r e e m a n ( i n M y c o t a x o n 8: 7 2 . 1979) m a y not be the same as
H e i n e m a n n ' s . T h e subgenus i s m a i n l y confined to the tropics. T h e
species described b e l o w differs so strikingly i n the characters o f the
v e i l f r o m the other species found i n the Netherlands, that it is tenta-
tively placed i n subgen. Lanagaricus, awaiting further research. It i s
most closely related to representatives o f sect. Laeticolores Heinem.
41. Agaricus rufotegulis N a u t a i n Persoonia 17: 230. 1999. - F i g . 42.

S E L . I C O N . - N a u t a i n Persoonia 17: 335. 1999.
S E L . D E S C R . & FIGS. - N a u t a i n Persoonia 17: 231-232. 1999.
F i g . 4 1 . Agaricus bresadolanus. (habit X A). V E R N . N A M E - Bladhoopchampignon.
AGARICUS (Nauta) 61
23 m m , with broad annulus or without annulus, c y l i n d r i c a l to sub-

clavate, young subbulbous, at base up to 25 m m broad, straight to
curved, stuffed, white, at base y e l l o w i s h when scratched, pruinose to
minutely squamulose, usually w i t h more or less y e l l o w i n g , relatively
thick rhizomorphs. A n n u l u s at 0.75-0.85 o f height o f stipe, up to 25 m m
wide, descending, pending to slightly spreading, for a l o n g time partly
attached to pileus, often evanescent, white, w i t h smooth upperside;
underside with numerous often brownish floccose squames. Context 7-
10 m m thick i n pileus, white, not discolouring when cut, except for a
yellowish discoloration i n base o f stipe. S m e l l slightly like almonds or
anise when cut, sometimes unpleasant. Taste not k n o w n .
M a c r o c h e m i c a l reactions: Schaeffer-reaction y e l l o w i s h (surface o f
pileus) to orange (surface o f base o f stipe).
= 1.15-1.65, Q a v = 1.30-1.45, broadly e l l i p s o i d , without germ pore.
B a s i d i a l 8 - 2 4 X 7.0-8.5 m m , 4 - s p o r e d . L a m e l l a edge w i t h a 45-50 m m
broad sterile layer; cheilocystidia i n chains o f rounded rectangular to
globose elements o f 7-12 X 4.5-11 m m , w i t h globose to clavate ter-
m i n a l elements o f 9-21 X (6-)8-13 m m . P i l e i p e l l i s a cutis, pro parte a
transition to a trichoderm, o f r a d i a l l y arranged, 4.0-8.0 m m w i d e
hyphae w i t h c y l i n d r i c a l and often inflated elements o f 16-47 X 11-19
m m , w i t h slightly ascending clavate terminal elements o f 20-40 X 20-
F i g . 42. Agaricus rufotegulis. (habit X 'A). 23 urn, gradually passing into pileitrama, w i t h pale yellow, parietal pig-
ment. Stipitipellis a regular, slightly gelatinized cutis o f 4.5-7.5 m m
Pileus 80-115 m m , at first hemispherical, expanding to slightly irregu- w i d e hyphae w i t h c y l i n d r i c a l , sometimes slightly inflated elements,
larly convex w i t h sometimes a depressed centre, w i t h margin exceed- with pale yellow, parietal pigment, below annulus intermixed or part-
ing lamellae, y o u n g reddish b r o w n ( M u . 5 Y R 4/4 diluted), covered with l y covered w i t h clusters o f loosely connected, subglobose elements o f
fibrillose squamules, later at centre reddish brown, otherwise densely 10-20 X 10-17 m m .
covered w i t h reddish b r o w n to b r o w n fibrillose squamules o n paler H A B I T A T & DISTR. - Gregarious o n heaps o f rotting dead leaves i n
background, not discolouring to b e c o m i n g slightly copper-coloured on deciduous woods. Very rare i n the Netherlands, only k n o w n from the
handling; v e i l for a long time connecting pileus and stipe, later present type locality (Amersfoort); J u n e - A u g . A l s o k n o w n from one locality i n
as white fringe at margin o f pileus. L a m e l l a e , L = 140-180, 1 = 1-5, southern E n g l a n d , A u g . - N o v .
crowded, free, subventricose, up to 10 m m broad, at first pale p i n k i s h
brown, later darker, w i t h slightly paler, entire edge. Stipe 60-95 X 12- The above description is also based on material from E n g l a n d .
2. Allopsalliota Nauta & Bas
MARIJKE M . NAUTA
Agaricus sect. Magici Bas & Heinem. in Persoonia 13: 113. 1986; Allopsalliota Nauta & Bas, in Belg. J. Bot. 131:
189. ('1998') 1999.
S E L E C T E D LITERATURE - Bas & Heinem. in Persoonia 13: 113-121. 1986; Nauta in Belg. J. Bot. 131: 181-190.
('1998') 1999.
Basidiocarps collybioid, large, firm; pileus fleshy, usually fibrillose-squamose; lamellae free, crowded, relatively nar-
row, pale brown, finally reddish brown, blackening on drying; stipe thick, with remnants of general veil in form of gir-
dles or squames on lower half; context whitish, soon brightly yellow, then slowly vinaceous; spore print reddish brown.
Macrochemical reactions: ammonia green (context); Schaeffer-reaction purple (surface of pileus, stipe).
Spores ellipsoid or oblong to amygdaliform, microscopically yellowish brown in K O H , pale brown to brown in
water and ammonia, weakly dextrinoid, congophilous and cyanophilous when young, thick-walled, with apically thick-
ened endospore, smooth, minutely roughened as observed with S.E.M.; pleurocystidia absent; lamella edge sterile;
cheilocystidia not in chains, clustered, long, lageniform with capitate apex; hymenophoral trama regular; clamp-con-
nections lacking in all tissues. Development probably monovelangiocarpic, isocarpic or hymenocarpic. - Holotype
species: Agaricus geesterani Bas & Heinem.
HABITAT & DISTRIBUTION - Saprotrophic, terrestrial in woods and roadside verges, on nutrient-rich soil. Known from
The Netherlands, Germany and Israel.
The genus has been created to accommodate Agaricus geesterani Bas & Heinem., which is at present the only known
species in the genus. This species macroscopically resembles species of Agaricus, but is aberrant among other respects
in the macrochemical reactions. Microscopically it is more similar to representatives of the tropical genus
Micropsalliota Hohn. For a more detailed discussion of the differences and similarities of this genus and the genera
Agaricus and Micropsalliota see Nauta (in Belg. J. Bot. 131: 181-190. ('1998') 1999).
1. Allopsalliota geesterani (Bas & Heinem.) N a u t a & Bas i n B e l g . J . b r o w n tinges, without annulus, fibrillose to squamose w i t h reddish
Bot. 131: 189. ('1998') 1999. - F i g . 4 3 . brown squames, lower part often covered w i t h v e i l girdles; base fre-
Agaricus geesterani B a s & H e i n e m . i n Persoonia 13: 114. 1986. quently covered w i t h velar sock; often rooting i n s o i l . Context 20-30
S E L . I C O N . - B a s & H e i n e m . i n Persoonia 13: 121. 1986; Chrispijn, m m thick i n pileus, firm, white, immediately bright y e l l o w w h e n cut,
C h a m p . Jordaan: 68. 1999. after some minutes s l o w l y turning vinaceous red. S m e l l indistinct to
S E L . D E S C R . & F I G S . - B a s & H e i n e m . i n Persoonia 13: 114-117. faintly pleasant. Taste indistinct. Spore print reddish grey to reddish
1986; B e n d e r i n Z . P i l z k . 60: 19-20. 1994: N a u t a i n B e l g . J . B o t . 131: brown (5 Y R 4/ 2-3).
182-183. ('1998') 1999. M a c r o c h e m i c a l reactions: a m m o n i a green (context); K O H 5-30%
V E R N . N A M E - Toverchampignon. greyish to b r o w n i s h (context; B a s & H e i n e m . , i n Persoonia 13: 116.
1986); Schaeffer-reaction purple to dark p i n k (surface o f pileus, stipe).
Basidiocarps developed almost subterranean, i n late stage appearing. Spores 7.0-9.5(-10.0) X 4.5-6.0 u.m, o n average (7.5-)8.0-8.3(-8.7)
Pileus (70-)90-150 m m , at first (irregularly) hemispherical to convex, X 5.0-5.5 u,m, Q = 1.35-1.70(-1.80), Q a v = 1.50, e l l i p s o i d o r o b l o n g
expanding to convex or broadly truncately c o n i c a l w i t h depressed cen- to amygdaliform, without germ pore, with apically thickened
tre, w i t h irregularly involute margin, later straight, at first whitish, soon endospore. B a s i d i a 27-35 X 6.0-8.0 u,m, i n majority 4-spored, often i n
reddening, finally b e c o m i n g reddish b r o w n ( M u . 5 Y R 6/4, 5/6, 2.5 Y R part 2 - spored. L a m e l l a edge sterile; c h e i l o c y s t i d i a tufted, not i n
4-5/4, 10 R 4/3), often covered w i t h adhering soil, y o u n g fibrillose, chains, (40-)50-90 X (4.0-)5.5-7.5(-10.5) u,m, w i t h (2.0-)2.5-4.0(-4.5)
later often w i t h fibrillose appressed squames. L a m e l l a e , L = 160-200, u,m w i d e neck and (4.0-)5.0-6.5 u,m w i d e apex, filiform to lageniform
1 = 0-3, crowded, segmentiform, free, up to 6 m m broad, at first pale with capitate apex, near m a r g i n o f pileus irregularly shaped and u p to
y e l l o w i s h b r o w n (10 Y R 7/4), later p i n k i s h b r o w n , finally reddish 4 0 u,m long, hyaline or w i t h y e l l o w i s h parietal or intracellular pigment.
b r o w n (5 Y R 4/4-6), w i t h whitish, slightly denticulate edge. Stipe (55- P i l e i p e l l i s a cutis o f r a d i a l l y arranged, sometimes interwoven, (4.5-
)90-180 X 22-40 m m , irregularly c y l i n d r i c a l to slightly enlarged at )6.5-16.0 u,m w i d e hyphae w i t h c y l i n d r i c a l elements, w i t h slightly
base, usually slightly curved, stuffed, white at first, later w i t h p i n k i s h ascending, occasionally differentiated terminal elements, w i t h usually
62
ALLOPSALLIOTA {Nauta)
intracellular y e l l o w i s h b r o w n pigment, i n upper layer frequently w i t h

encrusting y e l l o w - b r o w n pigment. P i l e o c y s t i d i a clavate to lageniform
or narrowly u n i f o r m , 15-30 u,m long. T r a m a o f pileus often w i t h some,
relatively t h i c k - w a l l e d , 5.0-13 | x m w i d e , scarcely septate hyphae w i t h
yellow, oleiferous contents. Stipitipellis a regular cutis o f 3.0-15 (Jim
w i d e hyphae w i t h c y l i n d r i c a l elements w i t h intracellular y e l l o w i s h p i g -
ment. V e i l on stipe consisting o f curved, interwoven, 4.0-11 \xm w i d e
hyphae w i t h slightly inflated elements, and irregularly shaped, usual-
ly slightly ascending, up to 14 u,m w i d e terminal elements.
H A B I T A T & D I S T R . - In groups, saprotrophic, terrestrial i n m i x e d
deciduous woods and verges o f roads o n nutrient-rich clayey or sandy
soil. U s u a l l y o n slightly disturbed places. M a i n l y i n F l e v o l a n d and the
peat region i n the western part o f the country, occasionally elsewhere,
apparently spreading. July-Nov. Also recorded from Germany
(Nordrhein-Westfalen) and Israel.
F i g . 43. Allopsalliota geesterani. (habit X Vi).

3. Macrolepiota Sing.
E L S E C . VELLINGA
WITH COLLABORATION OF R.PJ. DE K O K
Macrolepiota Sing, in Papers Mich. Acad. Sc. Arts Letters 32: 141. ('1946') 1948.
SELECTED LITERATURE - M . Bon, Fl. mycol. Eur. 3, Lepiotes: 116-125. 1993; Candusso & Lanzoni, Lepiota: 505-
593. 1990; Migl. in Boll. Gruppo micol. G. Bres., n.S. 38: 131-148. 1995.
Basidiocarp pluteoid, with stipe protruding into pileus context, relatively big; pileus with coarse squames to fine gran-
ulose squamules; lamellae free and remote from stipe; stipe often with bulbous base; annulus present, in most species
made up of partial veil and universal veil; spore print white, cream, or pink.
Spores thick-walled, binucleate, dextrinoid, congophilous, cyanophilous, metachromatic in Cresyl Blue, with walls
swelling in ammonia (NH ( )) and acetic acid (CH COOH - q- ), with germ pore, with or without hyaline cap over
3 aq 3 ( a )
germ pore, glabrous; cheilocystidia always present; pleurocystidia absent; hymenophoral trama trabecular; velum uni- {
versale on pileus trichodermal or hymenidermal; clamp-connections in most species present, but very often rare and
difficult to find. Development bivelangiocarpic, and pileocarpic. - Type species Macrolepiota procera (Scop.: Fr.) Sing.
HABITAT & DISTRIBUTION - Gregarious, in fairy rings, saprotrophic, terrestrial, in grasslands, deciduous and conif-
erous woods and plantations, on roadsides and in nutrient-rich places like compost heaps and landfills. Widespread and
world-wide, though absent from arctic and high-alpine habitats.
The genus Macrolepiota consists of two groups of species, one characterized by a glabrous stipe, relatively small spores
without a hyaline cap over the germ pore and narrowly clavate, tightly packed elements in the pileus covering, and the
other by a banded-squamulose or granulose stipe, a hyaline cap over the germ pore and a more loosely arranged tri-
chodermal pileus covering, made up of cylindrical elements. Within those two sections, several well-defined species
complexes can easily be recognized: M. rachodes-growp, M. procera-group and M. mastoidea and allies; to determine
individual species within the complexes, is much more difficult.
The presence or absence of clamp-connections is often difficult to determine. For that reason it is only used as an addi-
tional character in the key.
The neutral term 'pileus covering' is used for the covering layers of the pileus, not taking into account their origin.
Some species may cause gastrointestinal problems; many are considered excellent edibles, though individual reac-
tions to them vary greatly.
K E Y TO T H E SPECIES ° ^ s 1<W
1. Stipe with granules, squamulose zig-zag bands, either concolorous with background, or contrasting in colour
2. Stipe covering not or hardly contrasting with background; pileus with granular squamules or patches or a star-
shaped patch covering most of the surface
3. Annulus thin and simple, white; pileus (pale) brown at centre, pale buff to cream around centre, with a pale
star-shaped pattern on whitish background; stipe covering indistinct; clamp-connections often present at base
of basidia 5. M . excoriata
3. Annulus thick, often with double crown, but upper part not always well developed, with coloured patches on
underside; pileus colour variable, with granular aspect, which may form a star-shaped pattern on whitish back-
ground; stipe with a banded pattern; clamp-connections rarely present at base of basidia . . 4. M . mastoidea
2. Stipe covering dark, brown or greyish; pileus either with a dark brown granulose, patchy covering, or with dis-
tinct squames and squamules on top of a radially fibrillose background
4. Pileus brown granulose, and overall pattern either star-shaped or more irregularly patchy . . . 4. M . mastoidea
4. Pileus with distinct squames and squamules lying on top of or at the end of radially oriented fibrils
64
MACROLEPIOTA (Vellinga) 65
5. Pileus greyish brown to pale beige, strongly radially fibrillose, with loose-lying grey-brown patches of uni-
versal veil around centre 3. M . fuliginosa
5. Pileus with brown to vinaceous brown squamules on pale background, around central calotte with concen-
trically arranged small squames, at the end of short fibrils
6. Pileus and stipe with brown squamules on whitish background; stipe not reddening on scratching
1. M . procera
6. Pileus and stipe with dark brown squamules with reddish, vinaceous tinges; stipe becoming red when
scratched 2. M . permixta
1. Stipe smooth, without granules, bands etc.
7. Lamellae and spores of mature specimens green; spores with wide truncate apex Chlorophyllum molybdites
7. Lamellae and spores not changing colour with aging; spores with narrow truncate or rounded apex
8. Annulus thin and simple, not thickened with a double edge
9. Pileus pale brown to whitish, with star-shaped granular to slightly squamulose covering on whitish back-
ground; spores 11.0-16.0 X (6.5-)7.5-10.5 pm, with rounded apex with hyaline cap over germ pore
5. M . excoriata
9. Pileus whitish with coarse brown squames; spores 9.5-11.5 X 6.5-8.0 pm, with truncate apex, without hya-
line cap 8. M . venenata
8. Annulus thickened, with double edge (crown), consisting of tissue of the velum universale on the underside,
and of the velum partiale fringed on the upperside
10. Stipe context becoming pale brown on exposure; pileus covering with ascending hyphae with loosely
arranged terminal, brown, cylindrical elements; cheilocystidia lageniform to narrowly clavate with apical
excrescence, colourless; spores with translucent cap on germ pore; clamp-connections absent
Leucoagaricus nympharum
10. Stipe context discolouring orange-red when cut; pileus covering (velum patches) trichodermal with close-
ly packed, narrowly clavate, terminal elements; cheilocystidia clavate, narrowly clavate or spheropedun-
culate, often with brown intracellular pigment; spores without translucent cap on germ pore;
clamp-connections present at the base of basidia
11. Pileus with contrasting velar squames (brown on white to cream often fibrillose background); spores
on average 9.8-11.1 X 6.3-7.7 pm 6. M . rachodes
11. Pileus squames (velar patches) not contrasting in colour with background fibrils, equally brown or
grey-brown in colour; spores on average 8.7-10.0 X 5.8-6.6 pm 7. M . olivieri
Sect. Macrolepiota Pileus c. 100-200(-300) m m , w h e n y o u n g more or less paraboloid,

expanding v i a campanulate and convex w i t h l o w u m b o to applanate
Stipe w i t h squamulose o r granulose c o v e r i n g ; those squamules w i t h deflexed margin and l o w umbo, w h e n very y o u n g w i t h complete-
arranged i n bands o r girdles, or irregular patches; spores w i t h hyaline ly closed, felted, b r o w n ( M u . 7.5 Y R 6-5/4) to pale b r o w n (10 Y R 8-7/4)
cap over g e r m pore. at margin covering; this covering breaks open, leaving a central calotte,
and squamules around it; i n mature specimen central part c. 30-50 m m
1. Macrolepiota procera (Scop.: Fr.) S i n g , i n Papers M i c h . A c a d . S c i . , i n diameter, felted, dark b r o w n (10 Y R 4-3/4-6, 7.5 Y R 4/6), around
A r t s Letters 32: 141. 1946. - F i g . 4 4 . central patch first concentrical, small squamular patches, concolorpus
Agaricus procerus Scop., F l . earn. 2: 418. 1772; Agaricus procerus with calotte, not o r slightly uplifted, towards margin bigger and paler
Scop.: Fr., Syst. m y c o l . 1: 20. 1821; Lepiota procera (Scop.: Fr.) S . F . brown patches, at the uplifted tips o f the cream to pale b r o w n , radially
Gray, Nat. Arr. B r i t . PI. 1: 601. 1821; Mastocephalusprocerus (Scop.: fibrillose-squamose background; i n marginal zone o n l y radial fibrils
Fr.) Pat., E s s a i tax. H y m . : 1 7 1 . 1 9 0 0 ; Lepiotophyllumprocerum (Scop.: present; margin fringed, w h i t i s h cream, exceeding lamellae. L a m e l l a e ,
Fr.) L o c q . i n B u l l . mens. S o c . l i n n . L y o n 11: 40. 1942. L = 120-160, 1 = 0-3, moderately crowded, free, remote from stipe,
EXCL. - Lepiota procera sensu C l e L , Toadst. M u s h r . South subventricose, 8-12 m m broad, white, later creamy b r o w n , w i t h white
A u s t r a l i a : 54. 1935 ( = Macrolepiota clelandii); Lepiota procera sensu floccose edge (visible without lens). Stipe c. 100-400 X 10-20 m m ,
L i n c o f f , F i e l d G u i d e N . A m e r . M u s h r . : p i . 172. 1981 ( = Macrolepiota widened at base to a rounded bulb o f 25-50 m m wide, c y l i n d r i c a l and
spec). slightly w i d e n i n g downwards, hollow, i n y o u n g specimens dark grey-
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 254. 1995; brown (10 Y R 5/3) w i t h very s m a l l 'patches' o n white background; i n
Candusso & L a n z o n i , L e p i o t a : p i . 6 3 . 1990; Cetto, G r . P i l z f . , 6. A u f l . , older specimens brownish (c. 10 Y R 7/4) and w i t h adnate zig-zag bands
1: p i . 2 0 . 1980; D i i h n c k e , 1200 P i l z e : 5 2 1 . 1993; J . L a n g e , F l . agar, or adnate squamulose girdles over w h o l e length, but vaguer and less
dan. 1: p i . 8b. 1935; M i g l . i n R i v . M i c o l . 41 (4): b a c k cover. ( ' 1 9 9 8 ' ) contrasting at apex than i n m i d d l e part, o n w h i t i s h background; bulb
1999; R . P h i l l i p s , Paddest. S c h i m m . : 24. 1981. white tomentose; surface when scratched not or slightly and slowly dis-
S E L . D E S C R . & F I G S . - Breitenb. & K r a n z l , P i l z e S c h w e i z 4 : 218. colouring yellow. A n n u l u s movable w i t h age, complex, thickened w i t h
1995; Candusso & L a n z o n i , L e p i o t a : 510-512, f i g . 113. 1990 (as M. a broad edge (double crown), upperside creamy and fringed; underside
procera var. procera); H o r a k i n S y n . G e n . Agar.: 363-365. 1968. brown as calotte. Context thick i n pileus, c. 15 m m high, white and dull,
V E R N . N A M E - Grote parasolzwam. buff-coloured i n stipe; stipe protruding into pileus; sometimes slightly
F i g . 44. Macrolepiota procera.
d i s c o l o u r i n g to y e l l o w i s h red (5 Y R 5/6), especially i n stipe. S m e l l H A B I T A T & D I S T R . - Gregarious, saprotrophic and terrestrial i n open
indistinct, a bit nutty fungoid, sometimes spermatic. Taste none to fun- places, i n dune grasslands, o n roadsides, i n light woods, etc. C o m m o n
goid. Spore print p i n k (Breitenbach & K r a n z l i n , P i l z e S c h w e i z 4: p i . and widespread i n the Netherlands, but a v o i d i n g the areas w i t h peat
254. 1995), but white according to other authors. and heavy clay; June-Nov. F o r more information o n ecology and dis-
Spores 12.5-16.5 X (7.5-)8.0-11.0um, o n average 13.8-15.7 X 9,2- tribution, see N a u t a & V e l l i n g a ( A t l . N e d e r l . Paddest.: 202. 1995;
9.9 u m , Q = 1.4-1.95, Q a v = 1.45-1.6, i n side-view e l l i p s o i d - a m y g - i n c l u d i n g M. fuliginosa). W i d e s p r e a d and c o m m o n i n Europe; it has
daliform, i n frontal v i e w e l l i p s o i d to o b l o n g , t h i c k - w a l l e d , w i t h apical been reported from N o r t h A m e r i c a , A f r i c a , Japan, A u s t r a l i a and N e w
germ pore and h y a l i n e cap over it, dextrinoid, congophilous, Zealand, but several closely related species may be involved.
cyanophilous, w i t h p i n k inner w a l l i n C r e s y l B l u e . B a s i d i a 38-54 X
10.5-16 u m , 4-spored, w i t h c l a m p - c o n n e c t i o n . L a m e l l a edge sterile Macrolepiota procera and the closely related species M. fuliginosa and
and f o r m i n g a b r o a d band. C h e i l o c y s t i d i a 17-62 X 9.0-18 u m , very M. permixta, are characterized by the peculiar structure o f the veil: the
variable i n size and shape, c y l i n d r i c a l , narrowly utriform, irregularly v e l u m universale is a trichoderm, made up o f erect cylindrical elements,
shaped, often catenate, colourless. P l e u r o c y s t i d i a not observed. Pileus w i t h an increasing amount o f pigment towards the top; last elements
c o v e r i n g (in the calotte) a trichoderm, w i t h more and more b r o w n , with widened apex, o n w h i c h thick-walled b r o w n , cylindrical elements
intracellular pigment towards apex, made up o f c y l i n d r i c a l elements, are situated, often w i t h several relatively thin septa. T h e strength o f the
5.0-12 u m wide, ending i n a capitate element; terminal elements o n top connection between the t h i c k - w a l l e d and the normal elements, deter-
of these elements, very t h i c k - w a l l e d , w i t h b r o w n pigment, w i t h 0-2 mines whetherthe v e i l is loosely l y i n g o n the surface or not.
thin septa, up to 2 0 0 X 8.0-10 u m , w i t h rounded apex; clamp-connec- In the Netherlands, two different variants o f M. procera occur, the
tions absent. Stipitipellis a cutis w i t h squames and bands o f pale b r o w n ' n o r m a l ' type, described above, and a greyer variant, w i t h a radially
hyphae, made up o f rather short thick elements; terminal elements 5.0- fibrillose pileus, o n w h i c h loose rather b i g squames are l y i n g . B o t h
10(-20) u m w i d e ; pigment parietal and sometimes intracellular. C l a m p - may d i s c o l o u r p i n k or reddish when scratched. T h e second variant was
connections present at base o f basidia. k n o w n as M. permixta, and D e K o k & V e l l i n g a (in Persoonia 17: 76-
77. 1998) considered the two identical. However, m o l e c u l a r evidence fuliginosa sensu C a n d u s s o & L a n z o n i , L e p i o t a : 5 1 4 - 5 1 7 , p i . 64.
points clearly to the existence o f two taxa, M. procera, and a second 1990.
taxon. T h e two can be distinguished o n m o r p h o l o g i c a l grounds, though S E L . I C O N . - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 25: 121. 1982
there are no m i c r o s c o p i c a l differences. It appeared, however, that M. (as M. fuliginosa); Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 253. 1995;
permixta is not the correct name for the second taxon, but that the name Candusso & L a n z o n i , L e p i o t a : p i . 64 (based o n p. 121 i n B o l l . G r u p p o
M. fuliginosa is more appropriate. C o n f u s i o n concerning the name o f m i c o l . G . B r e s . 25. 1982). 1990 (as M. procera var. fuliginosa).
the species is caused b y different opinions i n the literature: the Italian S E L . D E S C R . & FIGS. - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 25:
authors B e l l i i (in B o l l . G r u p p o m i c o l . G . B r e s . 25: 115,117-118. 1982) 117-118. 1982 (as M. fuliginosa); Candusso & L a n z o n i , L e p i o t a : 514-
and Candusso & L a n z o n i (Lepiota: 514-521, p i . 64 & 65. 1990) used 516, fig. 114. 1990 (as M. procera var. fuliginosa); Pacioni i n M i c o l .
'fuliginosa' for a red-brown taxon, and ' p e r m i x t a ' for the more grey- ital. 8 (3): 13. 1979.
b r o w n specimens. B o n (e.g. F l . m y c o l . E u r 3., Lepiotes: 125. 1993) o n
the other hand, stuck to the original descriptions by B a r l a (Lepiota per- C H A R A C T E R I S T I C S - P i l e u s up to 150-200 m m , applanate w i t h l o w
mixta i n B u l l . S o c . m y c o l . Fr. 2: 114. 1886; and Lepiota procera var. umbo, w i t h b r o w n reddish to b r o w n b l a c k i s h centre and s m a l l b r o w n
fuliginosa i n C h a m p . A l p e s - M a r i t i m e s : 2 1 . 1888), and regarded grey- squamules concentrically arranged o n radially fibrillose background;
ish b r o w n specimens as M. fuliginosa (hence the name), and reserved lamellae often b r o w n - p i n k discoloured; stipe up to c. 2 0 0 X 10 m m ,
the name M. permixta for the red-brown species. It is not clear whether w i t h up to 30 m m w i d e bulbous base, w i t h b r o w n reddish to dark
M. permixta i n this sense does o c c u r i n the Netherlands, but a short brown irregular bands, sometimes reddish at bulb; annulus w i t h dou-
characterization is given to facilitate recognition. ble c r o w n , w h i t i s h o n upperside, and dark b r o w n o n underside; context
P a z m a n y ( i n N o t . bot. H o r t i agrobot. C l u j - N a p o c a 18-19: 5-22. reddening i n pileus o n exposure, b e c o m i n g red-brown i n stipe; spore
1988/1989) described several varieties and formae o f M. procera from print white to p i n k i s h cream.
R o m a n i a , o n account o f differences i n colour, stature, and squamosity Spores 12-18 X 8.0-11 u m , w i t h g e r m pore; b a s i d i a 4-spored;
of the pileus; these taxa have not yet been recorded f r o m elsewhere i n cheilocystidia clavate to c y l i n d r i c a l , often ending i n l o n g excrescences;
Europe. pileus covering (in calotte) a trichoderm; clamp-connections present at
Macrolepiota procera var. pseudoolivascens B e l l u & L a n z o n i (in base o f basidia.
Beitr. K e n n t n . P i l z e Mitteleur. 3: 190-191. 1987) differs from the typ- H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial i n open
i c a l variety o f M. procera i n the olivaceous-greenish discoloration o f places i n woods, city-parks etc. Reported from central and southern
pileus surface and stipe b y scratching; the context c o l o u r changes into Europe, not k n o w n w i t h certainty from the Netherlands.
p i n k i s h . T h i s variety has a mediterranean distribution.
Macrolepiota procera var. mediterranea M . B o n (in D o c . m y c o l . 23 T h e characteristics given above, are based o n the selected descriptions
(89): 62. 1993) has been described from the Mediterranean, m a i n l y dif- and figures.
fering from typical M. procera i n the huge size (pileus up to 4 0 c m and The presence o f Macrolepiota permixta i n the Netherlands is not
a stipe reaching 60 cm) and the labile, b r o w n squames o n the pileus. certain. If it occurs, it is a very rare species indeed. T h e data available
Macrolepiota olivascens M o s . ex M o s . & S i n g , (in S c h w e i z . Z . indicate that M. permixta is a species w i t h a predominantly southern
P i l z k . 39: 154. 1961) resembles M. procera, but stipe and pileus turn distribution i n Europe.
green w h e n maltreated, and the context discolours reddish; it has a T h e taxon w h i c h was k n o w n as M. permixta i n the Netherlands has
p i n k spore print. It is a montane species o f central E u r o p e and to be c a l l e d M. fuliginosa (see for an extensive discussion under M.
S c a n d i n a v i a , g r o w i n g under conifers. M o r e research is needed to procera).
establish the value o f these intraspecific and specific taxa.
A s M. procera is often collected for food, it has often been tested 3. Macrolepiota fuliginosa (Barla) M . B o n i n D o c . m y c o l . 11 (43):
for the amount o f heavy metals and radioactive material. It appears that 75. 1 9 8 1 . - F i g . 45.
M. procera is not an accumulator o f those elements, but consumption Lepiota procera var. fuliginosa Barla, Champ. Alpes-Maritimes:
of basidiocarps f r o m contaminated localities (roadsides, landfills, i n 21. 1888; Leucocoprinus fuliginosus (Barla) L o c q . i n B u l l . mens. Soc.
the neighbourhood o f smelters o f heavy metals) is to be discouraged l i n n . L y o n 14: 92. 1945; Macrolepiota fuliginosa (Barla) M . B o n i n
(Seeger i n Beitr. K e n n t n . P i l z e Mitteleur. 3: 2 8 9 - 2 9 8 . 1 9 8 7 ; T h o m a s i n D o c . m y c o l . 7 (27-28): 2 0 . 1 9 7 7 (not valid); Macrolepiota procera yar.
M y c o l o g i s t 6: 195-196. 1992; K a l a c et a l . i n S c . tot. E n v . 177: 2 5 1 - fuliginosa (Barla) B e l l u & L a n z o n i i n Beitr. K e n n t n . P i l z e Mitteleur. 3:
258. 1996). 190. 1987. - Lepiota rhodosperma P D . O r t o n i n Notes R . bot. G d n
Manz (Unters. Kultur Lebensz. Leucoagaricus naucinus, E d i n b . 4 1 : 591. 1984 (as Lepiota 'Macrolepiota' rhodosperma);
M a c r o l e p i o t a procera. 1971) was successful i n culturing Macrolepiota Macrolepiota rhodosperma (P.D. Orton) M i g l . i n B o l l . G r u p p o m i c o l
procera f r o m spore to basidiocarp, though it is not an easy procedure. G . Bres., n . S . 38: 140-141. 1995.
E X C L . - Macrolepiota fuliginosa sensu B e l l u i n B o l l . G r u p p o
2. Macrolepiota permixta (Barla) Pacioni i n M i c o l . ital. 8 (3): 13.1979. m i c o l . G . B r e s . 2 5 : 117-118, 121. 1982; Macrolepiota procera var.
Lepiota permixta B a r l a i n B u l l . S o c . m y c o l . Fr. 2: 114. 1886; fuliginosa sensu Candusso & L a n z o n i , L e p i o t a : 514-517, p i . 64. 1990
Leucocoprinus permixtus (Barla) L o c q . i n B u l l . mens. Soc. l i n n . L y o n ( = i n both cases M. permixta).
14: 91. 1945; Macrolepiota permixta (Barla) M o s . , Blatter-, M I S A P P L . - Macrolepiota konradii sensu Breitenb. & K r a n z l , P i l z e
B a u c h p i l z e , 1. A u f l . : 114. 1953 (not valid); Macrolepiota procera var. S c h w e i z 4: p i . 250. 1995; Macrolepiota permixta sensu B e l l u i n B o l l .
permixta (Barla) Candusso i n Candusso & L a n z o n i , L e p i o t a : 518. 1990. G r u p p o m i c o l . G . B r e s . 25: 115, 117 & 120. 1982; Macrolepiota pro-
E X C L . - Macrolepiota permixta sensu B e l l u i n B o l l . G r u p p o m i c o l . cera var. permixta sensu Candusso & L a n z o n i , L e p i o t a : 518-521, p i .
G . B r e s . 25: 115, 117 & 120. 1982; Macrolepiota procera var. per- 65. 1990.
mixta sensu Candusso & L a n z o n i , L e p i o t a : 518-521, p i . 65. 1990 ( = S E L . I C O N . - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 25: 120. 1982
M. fuliginosa). (as M. permixta); Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 250. 1995
M I S A P P L . - Macrolepiota fuliginosa sensu B e l l u i n B o l l . G r u p p o (as M. konradii); Candusso & L a n z o n i , L e p i o t a : p i . 65. 1990 (as M.
m i c o l . G . B r e s . 2 5 : 117-118, 121. 1982; Macrolepiota procera var. procera var. permixta).
P i l z e S c h w e i z 4: p i . 250. 1995), or pale clay-pink, pale salmon to pale

p i n k (Orton i n Notes R . bot. G d n E d i n b . 4 1 : 592. 1984).
Spores (11.5-) 12-20 X 7.5-11.5 u m , o n average 13.6-15.2 X 8.6-
10.5 u m , Q = 1.2-1.85, Q a v = 1.4-1.6, e l l i p s o i d to o b l o n g , slightly
a m y g d a l i f o r m i n side-view, t h i c k - w a l l e d , w i t h g e r m pore and hyaline
cap over it, dextrinoid, c o n g o p h i l o u s , cyanophilous, w i t h p i n k inner
w a l l i n C r e s y l B l u e . B a s i d i a 30-52 X 10-16 u m , 4-spored, sometimes
i n t e r m i x e d w i t h 2-spored b a s i d i a , w i t h basal clamp-connection.
L a m e l l a edge sterile, w i t h a b a n d o f cystidia. C h e i l o c y s t i d i a 16-60 X
7.0-17 u m , very variable i n shape and size, n a r r o w l y clavate, c y l i n -
d r i c a l , l a g e n i f o r m , often irregularly shaped, often catenate, rather
t h i n - w a l l e d and colourless. P l e u r o c y s t i d i a not observed. Pileus c o v -
ering (in calotte) a trichoderm o f erect c y l i n d r i c a l elements, c. 8.0-12
um w i d e , w i t h towards the uppersiaexmore b r o w n intracellular p i g -
ment, also w i t h bands o f b r o w n , e n c r u s f f a g p i g m e n t ; terminal ele-
ments o n top o f this trichoderm, b r o w n , very t h i c k - w a l l e d , w i t h 0-4
t h i n septa, w a v y often l y i n g h o r i z o n t a l l y o n top o f the trichoderm, up
to 2 5 0 X 12 u m , w i t h r o u n d e d apex; clamp-connections absent.
S t i p i t i p e l l i s a cutis, w i t h bands o f squamules o f pigmented p a r a l l e l
hyphae; terminal elements 16-50 X 5.0-12 u m , w i t h parietal or
encrusting pigment, w i t h rounded apex; thinner c y l i n d r i c a l elements
w i t h intracellular pigment present as w e l l . C l a m p - c o n n e c t i o n s pres-
ent, at least at the base o f basidia.
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic and
terrestrial, i n deciduous woods o n loamy soil, i n the dunes, and i n road-
sides, scattered and rare i n the Netherlands, k n o w n from southern
L i m b u r g , and some localities i n the Pleistocene areas, also recorded
F i g . 45. Macrolepiota fuliginosa.
from the F r i e s i a n islands, (July) Sept.-Oct. Widespread i n Europe, but
everywhere rare.
S E L . D E S C R . & F I G S . - B e l l u i n B o l l . G r u p p o m i c o l . G . Bres. 25:
115, 117. 1982 (as M. permixta); Candusso & L a n z o n i , L e p i o t a : 518- Macrolepiota fuliginosa is f a t h e r variable i n size, i n its reddening
520, f i g . 115. 1990 (as M. procera var. permixta); P . D . O r t o n i n Notes reactions and i n the c o l o u r o f the spore print. Its m a i n characteristics
R . bot. G d n E d i n b . 4 1 : 592. 1984 (as Lepiota rhodosperma). are the rather g r e y - b r o w n colours o f the velar patches o n the pileus,
w h i c h l i e l o o s e l y o n top o f a strongly r a d i a l l y - f i b r i l l o s e , pale grey-
Pileus 68-220 m m , plano-convex, applanate w i t h shallow depression to brown covering.
subconcave, w i t h l o w central umbo, w i t h deflexed marginal zone, w i t h D u t c h m y c o l o g i c a l literature and tradition have used the name M.
a calotte w h i c h i s grey-brown to rather dark b r o w n ( M u . 7.5 Y R 5/4 to permixta for a l l M. procera-liks specimens w h i c h , by turning red when
3/4, 2.5 Y R 2.5/2), central, r o u n d to star-shaped up to 1/3 o f radius bruised, d i d not fit the concept o f M. procera. Macrolepiota permixta is
w i d e , and felted-subsquamose; around central calotte w i t h rather covered i n red-brown squamules on a m u c h less fibrillose background,
s m a l l , 3-8 m m w i d e and very easily removable velar patches, w i t h and discolours vinaceous red-brown i n a l l its parts (see selected icones
recurving edges o n a l l sides, o n a radially coarsely fibrillose or floc- for representative pictures o f both species). In Italian literature the name
cose-fibrillose covering o f greyish hazel-brown or pale b r o w n or pale M. permixta has been used for M. fuliginosa and vice versa (see m i s -
beige (7.5 Y R 8-7/4, 10 Y R 7/4) fibrillose strands o n a p a l l i d to pale applied and excluded names). T h e notes under M. procera give more
beige background; m a r g i n slightly to distinctly fringed and exceeding information o n the two different applications o f these names.
lamellae. L a m e l l a e , L = c. 80-120,1 = ( 0 - ) l - 3 , moderately crowded Macrolepiota rhodosperma (P.D. Orton) M i g l . appeared to be iden-
to crowded, free and remote f r o m stipe, near stipe often anastomosing, tical w i t h M. fuliginosa, though it was o r i g i n a l l y described because o f
slightly ventricose, up to 12 m m w i d e , whitish w i t h p i n k i s h sheen, dis- its striking spore print colour, v i z . pale salmon-pink; this, however, is
tinctly cream-beige, w i t h age sordid pale cream w i t h sordid vinaceous not unusual i n the genus (see the spore print colours i n Breitenbach &
b r o w n stains a l o n g the edges, w i t h white, eroded to flocculose edge. K r a n z l i n , P i l z e S c h w e i z 4: p i . 252 and 254. 1995).
Stipe 95-270 X 4-16 m m w i t h clavate to rather abrupt basal bulb (14-
4 6 m m w i d e ) , h o l l o w , w i t h crowded, appressed, dark, slightly pale 4. Macrolepiota mastoidea (Fr.: Fr.) S i n g , i n L i l l o a 22: 417. ('1949')
beige greyish to greyish b r o w n squamules i n transverse bands o n l o w e r 1 9 5 1 . - F i g . 46.
half, but lighter, smaller and very crowded above (7.5 Y R 5/4 to 5 Y R Agaricus mastoideus Fr.: Fr., Syst. m y c o l . 1: 20. 1821; Lepiota
5/3-4/4) on pale buff background, often d i s c o l o u r i n g o n scratching mastoidea (Fr.: Fr.) K u m m . , Fiihr. P i l z k . : 135. 1871; Lepiota excoria-
(especially at base) to slightly vinaceous p i n k or red. A n n u l u s not real- ta subsp. mastoidea (Fr.: Fr.) Q u e l . , F l . m y c o l . France: 301. 1881;
ly movable, descending, w i t h cuff around stipe, creamy above; under- Leucocoprinus mastoideus (Fr.: Fr.) S i n g , i n Rev. M y c o l . 4: 67. 1939;
side concolorous w i t h calotte o n pileus; w i t h double c r o w n , but not Lepiotophyllum mastdideum (Fr.: Fr.) L o c q . i n B u l l . mens. Soc. l i n n .
always o b v i o u s l y so, as upperside may be under-developed. Context L y o n 11: 4 0 . 1942. - Agaricus gracilentus K r o m b h . , Naturg. A b b .
thick i n pileus, w h i t i s h or cream, often w i t h p i n k i s h - b r o w n i s h tinge, Beschr. essb. schadl. verd. S c h w a m m e : 8, p i . 24. figs. 13 & 14. 1836;
not changing colour or b e c o m i n g y e l l o w - b r o w n i n stipe; stipe context Lepiota gracilenta (Krombh.) Quel, i n M e m . Soc. E m u l . Montbeiiard,
protruding into pileus. S m e l l indistinct, nutty-fungoid, or slightly rub- ser. II, 5: 71. 1872 ( C h a m p . Jura Vosges 1); Macrolepiota gracilenta
ber-like. Taste like s m e l l . Spore print cream (Breitenbach & K r a n z l i n , ( K r o m b h . ) M o s . , Blatter-, B a u c h p i l z e , 1. A u f l . : 114. 1953 (not valid);
toidea^. rickenii, and M. konradii); Potz i n C a r i n t h i a II 181/101:412.

1991 (as M. konradii).
V E R N . N A M E - Tepelparasolzwam, i n c l . D o n k e r e parasolzwam.
Pileus 40-120 m m , w h e n y o u n g hemispherical to campanulate w i t h

inflexed margin, later w i d e - c o n i c a l to applanate, w i t h distinct umbo,
with margin exceeding lamellae, w h e n very y o u n g totally covered i n
pinkish-greyish-brownish ( M u . 1 0 Y R 5-8/3-4(-6); 7.5 Y R 5-8/4-6), to
dark b r o w n (e.g. K . & W . 7F7) velvety covering, later still w i t h velvety,
or plush-like covering at centre, towards margin more and more granu-
lose-patchy, w i t h bigger patches towards margin, sometimes i n a dis-
tinct star-shaped pattern, brownish yellow, b r o w n or p i n k i s h b r o w n (7.5
Y R 7-6/4), to dark b r o w n (7.5 Y R 4/4, paler than 6 D 4 near margin), on
cream-coloured to pale to very pale b r o w n ( M u . 10 Y R 7-8/2-4), fibril-
lose-squamulose background, showing white context i n between fibrils.
Lamellae, L = c. 60-85,1 = 0-3, crowded, moderately crowded, free,
remote from stipe, segmentiform to ventricose, 6-15 m m wide, some
furcate, white, creamy to pale brownish y e l l o w (10 Y R 6/4, 2.5 Y 8/1),
not discolouring on touch, w i t h even to eroded, whitish edge. Stipe (50-
)85-140(-210) X ( 5 - ) 7 - l l ( - 1 4 ) m m , bulbous, w i t h 1 5 - 2 3 ( - 3 5 ) m m w i d e
bulb, cylindrical to slightly tapering towards apex, hollow, whitish, dark
brown after touch (7.5 Y R 3/4), w i t h pale brownish, greyish brown, or
brownish p i n k ( 5 A B 3 ) , or creamy small bands o f patches or floccules
over total length o f stipe, the palest at apex, sometimes w i t h white
tomentose layer on bulb. A n n u l u s m o b i l e w i t h age, w i t h cuff-like part
around stipe, either ascending or descending, w i t h double c r o w n , but
often without, 12-40 m m i n diameter; underside brownish, as pileus
covering, or w i t h dark patchy r i m ; i f developed as double c r o w n , cream-
coloured at upperside, w i t h fringed margin. Context compact, up to 13
mm thick i n pileus, white to cream-coloured, after cutting not dis-
colouring or becoming reddish y e l l o w (5 Y R 7/6, 7.5 Y R 6/4) i n pileus
or reddish i n stipe; stipe context protruding into u m b o o f pileus. S m e l l
none, indistinct (also i n cut specimens) to slightly unpleasant, or fun-
goid. Taste faint. Spore print ' w h i t e ' .
Spores (11.0-)11.5-18.5(-20) X 7.5-12 urn, o n average 13.1-14.7 X
8.0-9.5 urn, Q = 1.35-1.8, Q a v = 1.45-1.65, e l l i p s o i d to oblong, ovoid,
F i g . 46. Macrolepiota mastoidea.
with central germ pore w i t h hyaline cap, w i t h 0.7-0.8 p m thick w a l l ,
dextrinoid, congophilous, cyanophilous, metachromatic in Cresyl
Macrolepiota gracilenta ( K r o m b h . ) Wasser i n U k r . bot. Z h . 35: 516. B l u e . B a s i d i a 31-68 X 10-16 p m , predominantly 4-spored, without,
1978. -Lepiota rickeniiWelen., N o v i t . m y c o l . : 47. 1939;Macrolepiota rarely with, basal clamp-connection. Lamella edge sterile.
rickenii (Velen.) B e l l u & L a n z o n i i n Beitr. K e n n t n . P i l z e Mitteleur. 3: C h e i l o c y s t i d i a 11-43 X 6.0-16 um, clavate, narrowly clavate, (broad-
196. 1987. - Lepiota excoriata var. konradi H u i j s m a n i n M e d e d . N e d . ly) fusiform to (narrowly) utriform, sometimes c y l i n d r i c a l and/or ros-
m y c o l . Vereen. 28: 18. 1943 (not valid); Lepiota konradii H u i j s m a n ex trate, or lageniform, thin-walled, colourless. P i l e u s covering at centre
P.D. O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : 283. 1960; Macrolepiota kon- an intricate trichoderm, made up o f c y l i n d r i c a l elements; terminal ele-
radii (Huijsman ex P.D. Orton) M a r c h a n d , C h a m p . N o r d M i d i 1: 224. ments 14-122 X 3.5-10 p m , c y l i n d r i c a l , often l y i n g o n surface, w i t h
1971 (not valid); Macrolepiota konradii ( H u i j s m a n ex P . D . Orton) parietal, sometimes intracellular pale to b r o w n pigment, often encrust-
Mos., R o h r l i n g e Blatterpilze, 3. A u f l . : 185. 1967. ing at base o f elements, and i n elements l o w e r d o w n . S t i p i t i p e l l i s a
E X C L . - Macrolepiota konradii sensu Breitenb. & K r a n z l . , P i l z e cutis o f narrow, c y l i n d r i c a l , colourless elements, 2.0-10 u m i n diame-
S c h w e i z 4: p i . 250. 1995 ( = M. fuliginosa). ter, w i t h clusters o f irregularly twisted and shaped hyphae, w i t h termi-
M I S A P P L . - Macrolepiota excoriata sensu Breitenb. & K r a n z l . , P i l z e nal elements 20-70 X 8.0-17 u m ; pigment pale and diffuse; b r o w n and
S c h w e i z 4: p i . 248. 1995; sensu Johnson & V i l g a l y s i n M y c o l o g i a 90: refractive i n an occasional narrow hypha. C l a m p - c o n n e c t i o n s absent
974-976. 1998. to rare at the base o f basidia.
S E L . I C O N . - B a r l a , F l . m y c o l . i l l . : p i . 11. figs. 1-10. 1889; M . B o n , H A B I T A T & DISTR. - Solitary, gregarious, never i n b i g groups, sapro-
C h a m p . Eur. o c c i d . : 2 9 1 . 1989 (as M. gracilenta & M. konradii); trophic and terrestrial i n (dune) grasslands, open places i n woods,
Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 251. 1995; Bres., Iconogr. rather rare i n the Netherlands, especially o n sandy soils (dune area,
m y c o l . 1: p i . 23 & 2 1 . 1927; Candusso & L a n z o n i , L e p i o t a : p i . 7 3 , 74 Pleistocene regions), and i n southern L i m b u r g . F o r more information
& 79. 1990 (as M. mastoidea, M. rickenii, and M. konradii resp.); on ecology and distribution see N a u t a & V e l l i n g a ( A t l . N e d e r l .
Cetto, F u n g h i V e r o 7: p i . 2597. 1993 (as M. konradii); C o o k e , 111. B r i t . Paddest.: 2 0 1 . 1995 (as M. konradii and M. mastoidea)). Sept.-Nov.
F u n g . 1: fig. 24 (23). 1881 (as Agaricus mastoideus); J . L a n g e , F l . agar, Widespread i n Europe.
dan. 1: p i . 8 C . 1935 (as Lepiota umbonata).
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 552-556, 5 6 3 - Macrolepiota mastoidea i n the current sense covers pale variants, for-
566, 584-586, figs. 123, 126 & 132 (resp. as M. mastoidea var. mas- merly recognized as M. mastoidea, m i d d l e b r o w n variants, formerly
k n o w n as M. rickenii, and dark b r o w n variants like M. konradii. T h e

annulus can be c o m p l i c a t e d w i t h a double c r o w n to almost simple w i t h
the upper part o f the c r o w n hardly developed. Contrary to what was
found b y D e K o k ( i n C o o l i a 34: 9 8 - 1 0 1 . 1991), clamp-connections
were very rarely observed o n the basis o f basidia i n the collections
studied. Candusso & L a n z o n i ( L e p i o t a : 556, 566. 1990) mentioned
that i n M. mastoidea var. mastoidea, and M. rickenii resp., c l a m p - c o n -
nections are present but difficult to find. A n o t h e r character thought to
distinguish M. konradii from M. mastoidea was the k i n d o f pigmenta-
tion o f the velar elements: encrusted i n M. konradii, and non-encrust-
ed i n the paler species. However, incrustations are also found i n typical
pale M. mastoidea-coWsctions,. M o l e c u l a r evidence supports the treat-
ment o f this group o f species as one taxon.
A discussion o n the s y n o n y m y o f M. mastoidea and M. gracilenta
can be found i n D e K o k & V e l l i n g a (in Persoonia 17: 74-76. 1998).
Several other taxa are closely related to M. mastoidea o r might be
identical to M. mastoidea: M. affinis (Velen.) M . B o n (see for a
description Candusso & L a n z o n i , L e p i o t a : 545-549. 1990), M. promi-
nens (Viv. ex Fr.) M o s . (a description can be found b y B e l l u & L a n z o n i
i n Beitr. K e n n t n . P i l z e Mitteleur. 2: 19-26. 1986), and M. subsquarrosa
( L o c q . ) M . B o n ( i n D o c . m y c o l . 11 (43): 7 2 . 1981). Whether these
species are identical to M. mastoidea has still to be proven.
Macrolepiota psammophila G u i n b . ( i n D o c . m y c o l . 24 (102): 2.
1996), described from the southern A t l a n t i c coast i n France, belongs
here as w e l l , o n account o f the granular pileus covering, and thg
absence o f clamp-connections at the base o f basidia. It is a very robust F i g . 47. Macrolepiota excoriata.
fungus, g r o w i n g i n calcareous dune sand.
A variety w h i c h reddens at the base o f the stipe has been described P i l e u s 60-100 m m , plano-convex to applanate w i t h u m b o and
f r o m France: M. mastoidea var. coccineobasalis (Locq.) M . B o n (in deflexed m a r g i n , w i t h utmost m a r g i n (velar remnants) exceeding
D o c . m y c o l . 11 (43): 7 2 . 1981). Macrolepiota mastoidea var. atro- lamellae, b r o w n at centre ( M u . 7.5 Y R 5/6, rarely 5/4 or 5/8, 5 Y R
brunnea D e r m e k (in F u n g . rar. Ic. c o l . 14: 11. 1985) is characterized 4/6), around centre pale cream-buff (c. 10-7.5 Y R 8/4), finely tomen-
by dark colours o n pileus, and fits w e l l into the present concept o f M. tose-velutinous, o r fine-granulose; c o v e r i n g towards m a r g i n breaking
mastoidea. up i n irregular patches or l e a v i n g a star-shaped velar patch; m a r g i n a l
To complete the o v e r v i e w o f the genus as it is represented i n zone, where the v e i l is not present, cream-coloured to white and r a d i -
Europe, two species w i t h dark squamulose-fibrillose pileus have to be ally fibrillose to squamulose. L a m e l l a e , L = c. 120,1 = ( 0 - ) l - 3 , m o d -
mentioned, both described from the mediterranean region: M. fuligi- erately c r o w d e d , c r o w d e d , d i s t i n c t l y free, segmentiform o r s l i g h t l y
neosquarrosa M a l e n 9 . (in B e i h . S y d o w i a 8: 261. 1979) from M o r o c c o , ventricose, u p to 7.5 m m w i d e , pale greyish-brownish to cream-beige
and M. phaeodisca B e l l u (in B o l l . G r u p p o m i c o l . G . Bresadola 27: 6. (7.5 Y R - 2.5 Y R 8/2), w i t h white flocculose edge. Stipe 90-120 X 8-
1984) f r o m Sardinia. See the latter publication for a comparison o f the 10 m m , c y l i n d r i c a l w i t h slightly bulbous to bulbous base, up to 18 m m
two species, illustrated b y photographs. T h e former has been recorded w i d e , h o l l o w , c o m p l e t e l y pale cream, b e l o w annulus w h i t i s h tomen-
f r o m Great B r i t a i n ( R e i d i n B o l l . G r u p p o m i c o l . G . Bres., n.S. 4 0 (2- tose-banded, b e c o m i n g s l i g h t l y b r o w n o r reddish b r o w n when
3): 399-404. ( ' 1 9 9 7 ' ) 1998). touched. A n n u l u s thin a n d s i m p l e , v a n i s h i n g w i t h age, descending;
underside concolorous w i t h stipe; upperside paler, or both sides cream
5. Macrolepiota excoriata (Schaeff.: F r ) Wasser i n U k r . bot. Z h . 3 5 : w i t h shaggy b r o w n i s h edge. Context white to white-cream i n pileus,
516. 1 9 7 8 . - F i g . 4 7 . t h i c k (up to 12 m m ) a n d d u l l , w h i t i s h to creamy, sometimes r e d i n
Agaricus excoriatus Schaeff., F u n g . Bavariae 4: p i . 18, 19. 1774; places; stipe protruding into pileus, context i n stipe shiny, d u l l i n base
Agaricus excoriatus Schaeff.: Fr., Syst. m y c o l . 1: 2 1 . 1821; Lepiota o f stipe. S m e l l none to strong and lepiotoid-fungoid. Taste not k n o w n .
excoriata (Schaeff.: Fr.) K u m m . , Fiihr. Pilzk.: 135. 1871; Spore print ' w h i t e ' .
Lepiotophyllum excoriatum (Schaeff.: Fr.) L o c q . i n B u l l . mens. Soc. Spores 11.0-16.0 X (6.5-)7.5-10.5 u m , o n average 11.9-13.6 X 8.1-
l i n n . L y o n 11: 40. 1942; Leucoagaricus excoriatus (Schaeff.: Fr.) S i n g , 9.2 u m , Q = 1.3-1.85, Q a v = 1.4-1.55, e l l i p s o i d to o b l o n g , slightly
in S y d o w i a 2: 3 5 . 1948; Macrolepiota excoriata (Schaeff.: Fr.) M o s . a m y g d a l i f o r m i n side-view, w i t h adaxial side less convex than abaxial
R o h r l i n g e , Blatter-, B a u c h p i l z e , 2. A u f L : 130. 1955 (not valid). side, thick-walled, w i t h apical germ pore and hyaline cap o n pore, dex-
E X C L . - Macrolepiota excoriata sensu Breitenb. & K r a n z l . , P i l z e trinoid, c o n g o p h i l o u s , cyanophilous, metachromatic i n C r e s y l B l u e .
S c h w e i z 4: p i . 248. 1995; sensu Johnson & V i l g a l y s i n M y c o l o g i a 90: B a s i d i a 3 5 - 5 7 X 11.5-14.5 u m , 4-spored, often with clamp-connection
974-976. 1998 ( = i n both cases Macrolepiota mastoidea). at base. L a m e l l a edge sterile. C h e i l o c y s t i d i a 18-41 X 6.0-13 p m , nar-
S E L . I C O N . - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 25: 103. 1982; r o w l y clavate, c y l i n d r i c a l , narrowly utriform, not coloured, w i t h hard-
Candusso & L a n z o n i , L e p i o t a : p i . 76. 1990; V e l l i n g a & H u i j s e r i n l y thickened w a l l . P l e u r o c y s t i d i a not observed. P i l e u s c o v e r i n g at
C o o l i a 40: p i . 8. 1997. centre a trichoderm made up o f c y l i n d r i c a l hyphae; terminal elements
S E L . D E S C R . & FIGS. - B e l l u i n B o l l . Gruppo micol. G . Bres. 25: 60-145 X 5.0-17 u m , slightly tapered at apex, and l y i n g o n surface,
102. 1982; Candusso & L a n z o n i , L e p i o t a : 572-574, fig. 129. 1990 (as colourless; pigment light-brown situated i n lower trichodermal hyphae.
M. excoriata var. excoriata). Stipitipellis a cutis o f colourless, c y l i n d r i c a l , hyphae, 3.0-6.0 u m w i d e ,
V E R N . N A M E - Rafelige parasolzwam. w i t h clusters o f irregularly twisted, rather narrow hyphae, 3.0-10 u m
wide, w i t h pale, slightly refractive intracellular pigment. C l a m p - c o n - Sect. Laevistipedes (Pazmany) M . B o n

nections present at the base o f basidia.
H A B I T A T & D I S T R . - Gregarious, solitary, terrestrial and saprotroph- Stipe smooth, not covered i n granules, squamules, bands etc.; spores
ic i n dune grasslands, o n dikes, i n roadsides and i n other types o f nutri- without hyaline cap over g e r m pore; pileus c o v e r i n g w i t h tightly
ent-poor grasslands; i n the Netherlands scattered and u n c o m m o n , and packed narrowly clavate terminal elements.
strongly declined; June-Nov. F o r more information o n ecology and dis-
tribution, and changes throughout time, see N a u t a & V e l l i n g a ( A t l . 6. Macrolepiota rachodes (Vitt.) S i n g , i n L i l l o a 22: 4 1 7 . ('1949')
N e d e r l . Paddest.: 200. 1995). Widespread i n E u r o p e . 1 9 5 1 . - F i g . 48.
Agaricus rachodes Vitt., Descr. F u n g h i mang. Italia: 158. 1835;
T h e basidia o f M. excoriata are often p r o v i d e d w i t h a distinct c l a m p - Agaricus procerus var. rachodes Rab., Deutschl. K r y p t . F l . : 574. 1844;
connection, though several authors e x p l i c i t l y mention the absence o f Lepiota rachodes (Vitt.) Quel, i n M e m . Soc. E m u l . Montbeliard. ser. II,
clamp-connections (Candusso & L a n z o n i , L e p i o t a : 5 8 0 . 1 9 9 0 ; B o n , F l . 5: 5.1872 (Champ. Jura Vosges 1); Leucocoprinus rachodes (Vitt.) Pat.,
m y c o l . Eur. 3, L e p i o t e s : 117. 1993). H o w e v e r , K a s p a r e k (in M i t t b l . Ess. tax. H y m . : 171. 1900 (as L. rhacodes); Lepiotophyllum rachodes
A r b e i t s g e m . P i l z k . N i e d e r r h e i n 10: 85. 1992) also mentioned the pres- (Vitt.) L o c q . in B u l l . mens. Soc. linn. L y o n 1 1 : 4 0 . 1 9 4 2 (as L. rhacodes).
ence o f clamp-connections. -Agaricus subtomentosus K r o m b h . , Naturg. A b b . Beschr. essb. schadl.
Macrolepiota heimii L o c q . ex M . B o n (in B e l l u i n B o l l . G r u p p o verd. Schwamme: 9, p i . 4. 1836; Agaricus procerus var. subtomentosus
m i c o l . G . Bres. 27: 18. 1984) is also a pale species. That it has c l a m p - (Krombh.) Rab., Deutschl. K r y p t . F L : 574. 1844. - Lepiota subprocera
connections was thought to separate it from M. excoriata, but this dis- Saut. i n H e d w i g i a 15: 152. 1876. - Lepiota bohemica Wichansky in
tinction is no longer v a l i d . B o t h species o c c u r i n grasslands, and M y k o l . Sb., Praha 38: 103. 1961; Macrolepiota bohemica (Wichansky)
despite the fact that the pileus c o v e r i n g is hardly star-shaped i n M. Krieglst. i n Z . M y k o l . 47: 83. 1981 (not valid); Macrolepiota bohemica
heimii, the two species appear to be identical. (Wichansky) Krieglst. & Pazmany i n P a z m a n y i n Z . M y k o l . 51: 52.
Several varieties o f M. excoriata have been described. Candusso & 1985; Macrolepiota rachodes var. bohemica (Wichansky) B e l l u &
L a n z o n i (Lepiota: 576-578. 1990) gave a g o o d overview. O n e i n w h i c h L a n z o n i i n Beitr. Kenntn. P i l z e Mitteleur. 3: 191. 1987.
the context reddens at exposure to air, is c a l l e d M. excoriata var. M I S A P P L . - Lepiota rachodes var. hortensis sensu P i l . & U s a k , Nase
rubescens (Dufour) M . B o n . H o u b y 1: 111. 1952. - Lepiota badhamii sensu M i c h a e l & H e n n i g ,
Because o f changes i n farming practice and land use especially i n Handb. Pilzfr. 3. A u f l . , 3: 139, p i . 16. 1964. - Macrolepiota olivieri
the second half o f the 20th century, this species has drastically declined sensu Wasser, F l . F u n g . R . S . S . U c r a i n i c a e , A g a r i c a c e a e : 298. 1980;
in the Netherlands (Nauta & Vellinga, A t l . N e d e r l . Paddest.: 2 0 0 . 1 9 9 5 ) . sensu Wasser i n L i b r i bot. 9: 87. 1993.
F i g . 48. Macrolepiota rachodes.

S E L . I C O N . - B a r l a , F l . m y c o l . i l l . : p i . 9bis, figs. 1-5. 1889; B e l l u i n pigment encrusting i n l o w e r elements. S t i p i t i p e l l i s a cutis o f 3.5-8.0

B o l l . G r u p p o m i c o l . G . B r e s . 25: 116. 1982 (as M. rhacodes var. hort- um w i d e , c y l i n d r i c a l hyphae, colourless; p i g m e n t situated i n stipi-
ensis); Candusso & L a n z o n i , L e p i o t a : p i . 69 1990 (as M. rachodes var. titrama. C l a m p - c o n n e c t i o n s present, at basidia, cystidia, and trama
bohemica); Cetto, F u n g h i Vero, E d . 5, 1: p i . 2 1 . 1975; R . P h i l l i p s , hyphae.
Paddest. S c h i m m . : 2 5 . 1981 (as M. rhacodes var. hortensis). H A B I T A T & D I S T R . - Solitary or gregarious, i n fairy rings and
S E L . D E S C R . & FIGS. - B e l l u i n B o l l . Gruppo micol. G . Bres. 25: swarms, saprotrophic a n d terrestrial, o n rather nutrient-rich s o i l , o n
115. 1982 (as M. rhacodes var. hortensis); B e l l u & L a n z o n i i n Beitr. compost heaps, i n gardens, orchards, deciduous and coniferous woods,
K e n n t n . P i l z e Mitteleur. 3: 191, 193, fig. 2. 1987 (as M. rachodes var. and o n roadsides, also i n greenhouses; c o m m o n , (Apr.-)Sept.-Dec.
bohemica); Candusso & L a n z o n i , L e p i o t a : 536-540, fig. 120. 1990 (as W i d e s p r e a d i n the subtropical to temperate regions o f the Northern
M. rachodes var. bohemica); L a v o r a t o i n Riv. M i c o l . 32: 273-274, fig. Hemisphere. R e c o r d e d f r o m the Southern Hemisphere, but c l o s e l y
1; 276-277, fig. 2. 1989 (as M. rachodes var. rachodes and M. rachodes related species might be i n v o l v e d . /
var. bohemica resp.).
VERN. NAME - Knolparasolzwam. Two ways to spell the epithet o f this species can be found i n the m y c o -
l o g i c a l literature: rachodes and rhacodes. T h e o r i g i n a l author, Vittadini
Pileus 42-220 m m , w h e n y o u n g globose to hemispherical, expanding (Descr. F u n g h i mang. Italia: 158. 1835) used the first spelling consis-
to plano-convex, w i t h shallow depression or l o w broad umbo, w h e n tently, though it is without obvious meaning, whereas 'rhacodes' is
very y o u n g completely ochraceous y e l l o w to dark b r o w n , soon w i t h derived from a G r e e k w o r d meaning ' r a g ' . T h e original author i s f o l -
central rounded to star-shaped or r a d i a l l y cracked patch, up to c. 60 l o w e d i n his s p e l l i n g o f the name.
mm i n d i a m . , surrounded b y irregular concentric rings o f squamular Macrolepiota rachodes is, w i t h M. procera, very often depicted, but
patches, flat or c u r v e d upwards, dark r e d to y e l l o w i s h r e d or (dark) g o o d descriptions o f this species, i n the sense used here, e x c l u d i n g
b r o w n (2.5 Y R 3/6, 5 Y R 3 - 6 / 2 - 8 , 1 0 Y R 4 - 5 / 3 - 4 , 1 0 Y R 6/3-6, 5/6, 7.5 dark variants recognized as M. olivieri, are hard to find. F o r instance,
Y R 5/8, 3/4, 6/4, 5/6, 4/4), those patches vanishing towards margin or the description o f M. rachodes b y Candusso & L a n z o n i , L e p i o t a : 530-
o n l y present at the tips o f r a d i a l l y fibrillose squamules, o n cream to 534. 1990 (as var. rachodes) includes M. olivieri. \
(very) light b r o w n or reddish yellow, sometimes light y e l l o w or y e l - Macrolepiota olivieri differs from M. rachodes i n that velar patch-
l o w i s h red (e.g. 10 Y R 8/2, 10 Y R 5-8/3-4, 7.5 Y R 6/4, 7.5 Y R 7/6, 2s5 es and background hyphae are both dark coloured, and that the spores
Y 7/4, 6/6, 5 Y R 5/8, 6/4) background, when y o u n g more or less are slightly smaller. C l a i m e d differences i n habitat, e x c l u s i v e l y i n
smooth, later distinctly r a d i a l l y fibrillose-squamose; margin fringed Pi'cea-plantations for M. olivieri versus a w i d e range o f habitats,
and exceeding lamellae; background bruising red, turning into black. i n c l u d i n g Pfcea-plantations f o r M rachodes (De K o k i n C o o l i a 34: 9 9 .
L a m e l l a e , L = c. 9 0 - 1 0 0 , 1 = 0-3, moderately c r o w d e d to crowded, 1991), c a n no longer be used to separate them, as grey variants o f M.
free, 1-4 m m remote f r o m stipe, sometimes anastomosing or furcate olivieri have been found i n deciduous woods i n southern L i m b u r g .
near stipe, (sub)ventricose, 4-18.5 m m w i d e , white (10 Y R 8/3) to Macrolepiota bohemica is often recognized as a separate taxon
cream, w h e n touched reddish y e l l o w (5 Y R 7/6) to light red (10 R 6/8), (either as a species, more often as a variety o f M. rachodes). A s was
orange (7.5 Y R 7/8), or yellow, later b r o w n (7.5 Y R 5/4), w h e n young shown b y D e K o k & V e l l i n g a (in Persoonia 17: 70-71. 1998), a l l the
w i t h white, later w i t h b r o w n , even to eroded edge. Stipe 26-160(-225) characters used to distinguish the t w o species form a continuum, and
X 7-30 m m , c y l i n d r i c a l w i t h 17-50 m m wide, sometimes slightly mar- vary independently. Preliminary molecular studies have failed to sepa-
ginate, bulb, h o l l o w , white, pale greyish-brownish, when touched light rate the two taxa. F o r a discussion on the name 'hortensis', also used for
b r o w n or y e l l o w i s h red (10 Y R 7/4, 5/4, 6/6, 5 Y R 5/6, 7.5 Y R 6/4, 5/6, this taxon, see again D e K o k & V e l l i n g a (in Persoonia 17: 70-71.1998).
7/6, 4/4, 5/8), smooth or l o n g i t u d i n a l l y striate, sometimes w i t h r h i - Macrolepiota venenata is quite s i m i l a r to M. rachodes, but differs
zomorphs at base; bulb white tomentose. A n n u l u s w i t h double c r o w n , m a i n l y i n the absence o f clamp-connections, a s i m p l e r i n g , and the
18-57 m m w i d e , 1-12 m m thick, w h i t i s h to buff-brown (5 Y R 7/3) more r a d i a l l y oriented velar arrangement o n the pileus. F i l i p p i &
above, sometimes adhered, concolorous w i t h pileus centre at under- B a r b i n i ( i n R i v . M i c o l . 3 2 : 266-271. 1989) proposed that the t w o
side. Context i n pileus 5-18 m m thick, white to w h i t i s h , turning b r o w n species c o u l d be identical.
w h e n aging (7.5 Y R 5/4), w h e n cut discolouring, sometimes first turn- Macrolepiota abruptibulba (R. H e i m ) H e i n e m . , a species described
ing y e l l o w (10 Y R 7-8/6), then reddish y e l l o w (7.5 Y R 8-6/6-8, 5 Y R from tropical A f r i c a , was found i n a greenhouse i n E n g l a n d (Pegler et
6-7/6-8 or 5 Y R 5/8) to w e a k l y r e d (10 R 5-6/6-8 or 2.5 Y R 6/6, 4/8, al. i n B o t . J . S c o t l . 5 0 : 191-197. 1998). T h i s species also belongs to
6/8), and finally b r o w n (5 Y R 3/3, 10 R 5/8). S m e l l fungoid, earth-like sect. Laevistipedes, because o f the smooth stipe, and the shape o f the
or l i k e rubber, slightly unpleasant. Taste fungoid or earth-like. Spore spores. T h e most striking characters o f this species are the strongly bul-
print probably white to cream. bous base, the dark purplish b r o w n pileus covering, and the pale pink-
Spores 9.0-12 X 6.0-7.5 u m , o n average 9.8-11.1 X 6.3-7.7 um, Q ish b r o w n pileus surface i n between the squames, and the p u r p l i s h
= 1.35-1.75, Q a v = 1.45-1.6, i n side-view e l l i p s o i d - a m y g d a l i f o r m , brown discolouration o f the stipe w h e n bruised.
o b l o n g - a m y g d a l i f o r m , i n frontal v i e w o v o i d , w i t h rounded to truncate C l a m p - c o n n e c t i o n s are often present, at least at the base o f the
apex, w i t h g e r m pore, w i t h distinct h i l a r appendage, t h i c k - w a l l e d , basidia i n M. rachodes, but sometimes are reported to be absent
d e x t r i n o i d , c o n g o p h i l o u s , c y a n o p h i l o u s , metachromatic in Cresyl (Kasparek i n M i t t b l . A r b e i t s g e m . P i l z k . Niederrhein 10: 84-85. 1992).
B l u e . B a s i d i a 32-49 X 8.0-14 u m , 4-spored, rarely 2-spored, often Macrolepiota rachodes is considered a g o o d edible species, but
w i t h b r o w n p i g m e n t near l a m e l l a edge, w i t h clamp-connection. specimens g r o w i n g i n the v i c i n i t y o f roads, i n areas where chemicals
L a m e l l a edge sterile. C h e i l o c y s t i d i a i n a broad band, 16-43 X 8.5-25 are sprayed, near heavy metal smelters etc. should be avoided. T h e
um, clavate, broadly clavate, spheropedunculate, often catenate, after species accumulates the heavy metals lead and to a lesser extent mer-
a frost-period sometimes rostrate, mostly w i t h brown(ish) intracellu- cury ( K a l a c & S v o b o d a i n F o o d C h e m . 69: 273-281. 2000).
lar pigment, rarely colourless. P l e u r o c y s t i d i a not observed. P i l e u s Sagara (in Trans, m y c o l . Soc. Japan 33: 487-496. 1992) reported
covering i n centre o f pileus an intricate trichoderm, w i t h clavate ter- on the occurrence o f M. rachodes o n o l d wood-ant nests i n E n g l a n d .
m i n a l elements, 20-65 X 8-14 u m , w i t h pale b r o w n intracellular p i g - However, there i s no evidence that the ants cultivate this m u s h r o o m
ment; some narrow elements w i t h b r o w n refracting pigment present; species.
7. Macrolepiota olivieri (Barla) Wasser, F l . F u n g . R . S . S . U c r a i n i c a e , amygdaliform, slightly o v o i d i n frontal v i e w , w i t h rounded to slight-

Agaricaceae: 298. 1980. - F i g . 4 9 . ly truncate apex w i t h s m a l l g e r m pore, t h i c k - w a l l e d , dextrinoid, c o n -
Lepiota olivieri B a r l a i n B u l l . S o c . m y c o l . Fr. 2 : 113. 1886; Lepiota gophilous, cyanophilous, metachromatic i n Cresyl Blue. Basidia
rachodes var. olivieri (Barla) B a r l a , F l . m y c o l . i l l . : 2 7 . 1889; 29-40 X 8.0-11 p m , 4-spored, sometimes i n t e r m i x e d w i t h 2 - and 3-
Macrolepiota rachodes f. olivieri (Barla) R . de K o k i n K o k & V e l l i n g a spored basidia, w i t h c l a m p - c o n n e c t i o n . L a m e l l a edge sterile w i t h a
i n Persoonia 17: 74. 1998. broad band o f cystidia. C h e i l o c y s t i d i a 16-45 X 35-45 u m , clavate to
E X C L . - Macrolepiota olivieri sensu Wasser, F l . F u n g . R . S . S . ovoid, not rostrate, many w i t h c l o u d e d b r o w n intracellular pigment.
Ucrainicae, Agaricaceae: 298. 1980; sensu Wasser i n L i b r i bot. 9: 87. P l e u r o c y s t i d i a not observed. P i l e u s c o v e r i n g at centre o f pileus an
1993 ( = i n both cases M. rachodes). intricate trichoderm w i t h n a r r o w l y clavate terminal elements, 25-50
S E L . I C O N . - B a r l a , F l . m y c o l . i l l . : p i . 9bis. figs. 6-10. 1889; B e l l u X 6.0-11 p m , w i t h b r o w n intracellular pigment; p i l e i t r a m a hyphae
i n B o l l . G r u p p o m i c o l . G . Bres. 25: 113 upper photo. 1982 (as M. rha- also pigmented. S t i p i t i p e l l i s a cutis o f 3.0-5.0 p m w i d e c y l i n d r i c a l
codes var. rhacodes); Candusso & L a n z o n i , L e p i o t a : p i . 6 7 . 1990 (as hyphae, w i t h b r o w n , often refractive pigment. Clamp-connections
M. rachodes var. rachodes); J . L a n g e , F l . agar. dan. 1: p i . 9 C . 1935 (as present, at least at base o f basidia.
Lepiota rhacodes); R . P h i l l i p s , Paddest. S c h i m m . : 2 5 . 1981 (as M. H A B I T A T & DISTR. - Solitary or gregarious, saprotrophic and terres-
rachodes var. rachodes); P i l . & U s a k , N a s e H o u b y 1: p i . 110. 1980 (as trial on l o a m y soils, i n woods and plantations w i t h coniferous trees (in
Lepiota rhacodes). the Netherlands Picea), i n southern L i m b u r g also i n deciduous o r
SEL. D E S C R . & F I G S . - K o k & V e l l i n g a i n P e r s o o n i a 17: 7 4 . 1998 (as m i x e d woods. Rather rare and scattered, Sept.-Nov. W i d e s p r e a d i n
M. rachodes f. olivieri). Europe, and probably quite c o m m o n .
V E R N . N A M E - Sombere knolparasolzwam.
Macrolepiota olivieri differs from M. rachodes i n the slightly smaller
Pileus 46-130 m m , when mature plano-convex, applanate, sometimes spores, and the fact that the velar squames and the fibrillose back-
(sub)umbonate; central velar patch star-shaped to circular, w i t h indistinct ground are o f the same colour.
border, 21-38 m m wide, w i t h concentrical zones o f applanate or upwards
curving squamulose patches, dark brown to reddish brown at centre to 8. Macrolepiota venenata M . B o n i n M . B o n et a l . i n D o c . m y c o l . 9
paler brown at margin (7.5 Y R 5/6,4-3/4, 3/2, 5 Y R 4-3/4, 10 Y R 6-5/4- (35): 13. 1979.
6, 7/3); background y e l l o w i s h brown to light brown or greyish brown S E L . I C O N . - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 24: 103, 105,
(10 Y R 5-6/3-5, 7/3, 7.5 Y R 6/4, 3/2), not or hardly paler than patches, 106. 1982; B o n et a l . i n D o c . m y c o l . 9 (35): p i . 1-3. 1979.
smooth to radially fibriUose-ragged. L a m e l l a e , L = c. 80-100,1 = 1-3, S E L . D E S C R . & FIGS. - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 24:
moderately crowded, free, 1-7 m m remote from stipe, 8.5-15 m m wide, 104, 107. 1982; B o n et a l . i n D o c . m y c o l . 9 (35): 13-16, f i g . 1. 1979;
whitish, later cream-beige, or y e l l o w i s h discoloured, when touched light Lavorato i n R i v . M i c o l . 32: 277-280, fig. 3. 1989.
red (2.5 Y R 6/8) to y e l l o w i s h red, w i t h dark brown eroded edge. Stipe
(45-)80-190 X 7-13 m m , bulbous to subbulbous, w i t h 21-32 m m wide C H A R A C T E R I S T I C S - Pileus 120-170 c m , convex, without umbo, w i t h
bulb, hollow, white to cream coloured, when scratched yellowish brown, central light b r o w n calotte u p to 70 m m i n diam., and w i t h light brown
saffron-orange to darker reddish brown (10 Y R 4/4, 10 Y R 5/4-6, 5 Y R appressed squames, w i t h towards margin a radial-fibrillose aspect, o n
3/4, 7.5 R 3/4), lengthwise marmorate-striate, white-tomentose on bulb. pale cream background; lamellae crowded, free, white to cream, dis-
A n n u l u s mobile w i t h age, w i t h double crown, 25-30 m m i n diameter, 1- colouring reddish b r o w n w i t h age; stipe 100-160 X 10-15 m m , w i t h
5 m m thick, very light b r o w n (10 Y R 7/4) at upper side, sometimes bulbous base, glabrous, pale; annulus simple, descending; context
adhered, lower side coloured as pileus centre. Context white, 7-17 m m white, w h e n cut d i s c o l o u r i n g p i n k - b r o w n ; s m e l l indistinct, fungoid;
thick i n pileus, when cut turning reddish y e l l o w or orange (7.5 Y R 6-7/8, spore print white.
5 Y R 6/8), later red (10 R 4-6/6-8 o r 2.5 Y R 5/6). S m e l l none, earth-like Spores 9.5-11.5 X 6.5-8.0 p m , on average 10.4 X 7.4 p m , Q = 1.3-
or fungoid. Taste unpleasant. Spore print colour unknown. 1.5, Qav = 1.4, e l l i p s o i d - a m y g d a l i f o r m , w i t h truncate apex, i n w h i c h
Spores (7.5-)8.0-11.0 X 5.5-7.0 u m , o n average 8.7-10.0 X 5.8- germ pore i s situated, t h i c k - w a l l e d ; b a s i d i a 28-37 X 11-15 u m , 4-
6.6 p m , Q = 1.3-1.65, Q a v = 1.45-1.55, i n side-view e l l i p s o i d - spored; cheilocystidia 26-35 X 13-15 p m , clavate; v e l u m o n pileus a
trichoderm, w i t h clavate terminal elements; pigment b r o w n and intra-
cellular i n c y l i n d r i c a l hyphae; stipitipellis a cutis; clamp-connections
not observed.
H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial i n
nitrogen-rich grasslands. K n o w n f r o m northern France and Italy.
Macrolepiota venenata is very c l o s e l y related to M. rachodes, from

w h i c h it differs i n the absence o f clamp-connections, the simple annu-
lus, and the radially fibrillose pileus surface.
T h e macroscopic characteristics given above, are based o n the
description b y B o n et a l . ( i n D o c . m y c o l . 9 (35): 14-15. 1979); the
microscopy is based o n observations o n the type material.
Ingestion o f M. venenata has been the cause o f gastrointestinal
problems ( B o n et al. i n D o c . m y c o l . 9 (35): 13-21. 1979; M a z z o l a i i n
CH
R i v . M i c o l . 32: 264-265. 1989).
F i g . 4 9 . Macrolepiota olivieri.
4. Chlorophyllum Mass.
ELSE C . VELLINGA
Chlorophyllum Mass. in Kew Bull. 1898: 136. 1898, non Chlorophyllum Murrill, 1910; Macrolepiota sect.
Chlorophyllum (Mass.) Moreno et al. in Mycotaxon 55: 469. 1995 (not valid).
SELECTED LITERATURE - Heinem. in Bull. Jard. bot. natn. Belg. 38: 195-206. 1968; Reid & Eicker in Bot. Bull. Acad.
sinica32: 317-333. 1991.
Basidiocarp pluteoid; pileus with coarse squames; lamellae free and remote from stipe; stipe with bulbous base; annu-
lus present, made up of partial veil and universal veil; spore print pale green to grey green.
Spores thick-walled, when young dextrinoid, congophilous, cyanophilous, metachromatic in Cresyl Blue, but when
mature not visibly so, with germ pore, glabrous; cheilocystidia present; pleurocystidia absent; hymenophoral trama
trabecular; velum universale on pileus trichodermal; clamp-connections inmost species present, but very rare and dif-
ficult to find. Development probably bivelangiocarpic, possibly hemiangiocarpic, and pileocarpic. - Holotype species
Chlorophyllum esculentum Mass. *
HABITAT & DISTRIBUTION - Saprotrophic, terrestrial, gregarious, in fairy rings, probably in woods in the Tropics, but
outside the Tropics in irrigated lawns and fields, in temperate areas only indoors. Widespread in the Tropics.
The genus Chlorophyllum is closely related to Macrolepiota sect. Laevistipedes (Pazmany) M . Bon, and can be dis-
tinguished by the green spores. >
Chlorophyllum molybdites ( G . M e y e r : Fr.) M a s s . i n K e w B u l l . 1898: Macrolepiota molybdites); R e i d & E i c k e r i n B o t . B u l l . A c a d , sinica 32:

136. 1898. 318-320, figs. 1 & 2. 1991; Sundb. i n M a d r o n o 2 1 : 15-20, f i g . 1. 1971;
Agaricus molybdites G . M e y e r , Pr. F l . essequ.: 300. 1818; Agaricus V e l l i n g a i n C o o l i a 33: 78-79, figs. 1 & 2. 1990; W a t l i n g i n M y c o l o g i s t
molybdites G . M e y e r : Fr., Syst. m y c o l . 1: 308. 1821; Mastocephalus 5: 23. 1991.
molybdites ( G . M e y e r : Fr.) O . K u n t z e , Rev. G e n . P I . 2 : 8 6 0 . 1891; VERN. NAME - Groenspoorknolparasolzwam.
Leucocoprinus molybdites ( G . M e y e r : Fr.) Pat. i n B u l l , trimest. S o c .
m y c o l . F r . 2 9 : 215. 1913; Macrolepiota molybdites ( G . M e y e r : Fr.) C H A R A C T E R I S T I C S - Pileus c. 9 0 m m , plano-convex w i t h very l o w ,
M o r e n o et a l . i n M y c o t a x o n 55: 467. 1995. - Agaricus morganii Peck broad umbo; v e l u m closed o n umbo, dark chocolate b r o w n ( M u . 10 Y R
in B o t . G a z . 4: 137. 1879; Lepiota morganii (Peck) S a c c , S y l l . F u n g . 3/4), pallescent towards m a r g i n and slightly c r a c k i n g into s m a l l
5: 30. 1887; Mastocephalus morganii (Peck) O . K u n t z e , Rev. G e n . PI. velutinuous patches around centre, w i t h some scattered 4-5-sided dark
2: 8 6 0 . 1891; Chlorophyllum morganii (Peck) M a s s . i n K e w B u l l . b r o w n patches o n cream coloured, radially fibrillose background, w i t h
1898: 136.1898. - Lepiota ochrosporaCooke & Mass. inGrevillea.21: at m a r g i n also some dark b r o w n patches; lamellae crowded, free,
73. 1893. - Chlorophyllum esculentum M a s s . i n K e w B u l l . 1898: 136. brownish w i t h greenish sheen; l a m e l l a edge eroded and dark b r o w n i n
1898; Lepiota esculenta (Mass.) Sacc. & S y d . i n S a c c , S y l l . F u n g . 16: places; stipe 84 X 11 m m , w i d e n i n g downwards to 2 2 m m w i d e trun-
2 . 1 9 0 1 . -Agaricus guadelupensis Pat. i n B u l l . Soc. m y c o l . Fr. 15: 197. cate base, ascending into pileus, h o l l o w , pale creamy buff, w i t h white
1 8 9 9 . - A n n u l a r i a camporum Speg. i n A n . M u s . nac. Hist. nat. B . A i r e s b l o o m , darkening o n touch, w i t h ascending annulus; context pale buff
6: 117. 1899; Lepiota camporum (Speg.) Speg. i n B o l n A c a d . nac. in pileus, b r o w n i s h i n stipe and reddish i n base o f stipe; smell o f cut
C i e n c . C o r d o b a 29: 114. 1926. - Agaricus congolensis Beeli i n Bull. basidiocarps like carrots; taste not reported; spore print 'pale green to
Soc. roy. bot. B e l g . 6 1 : 92. 1928; Chlorophyllum molybdites var. con- grey green' w h e n fresh (see Sundberg i n M a d r o n o 2 1 : 18. 1971).
golense ( B e e l i ) H e i n e m . i n F l . i c o n . C h a m p . C o n g o 16: 323. 1967. - Spores 8.7-11.2 X 6.7-8.3 um, o n average 9.9 X 7.6 um, Q = 1.2-
Chlorophyllum molybdites var. luteolosperma S i n g , i n Papers M i c h . 1.45, Qav = 1.35-1.4, broadly ellipsoid to ellipsoid, slightly amygdali-
A c a d . S c i . , A r t s Letters 32: 139. 1946. - Lepiota molybdites var. mar- form i n side view, w i t h conspicuous hilar appendix, green- and
ginata A . H . S m i t h , M u s h r . nat. H a b . : 429. 1949. thick-walled, with broad truncate apex and germ pore; mature spores not
S E L . I C O N . - Imaz. et a l , F u n g i Japan: 181. 1988; L i n c o f f , F i e l d reacting i n C o n g o R e d , M e l z e r ' s Reagent, nor i n C r e s y l B l u e ; immature
G u i d e N . A m e r . M u s h r . : p i . 169, 170. 1981; M i g l . i n B o l l . G r u p p o spores, of w h i c h the w a l l is not yet green, do colour i n the reagents men-
m i c o l . G . Bres., n.S. 39: 109. 1996; Serafin i n R i v . M i c o l . 39 (3): front tioned; basidia 29-40 X 9.0-11.5 um, 4-spored, without clamp-connec-
& back cover. 1996; W a t l i n g i n M y c o l o g i s t 5: 23. 1991. tion; lamella edge sterile; cheilocystidia 32-65 X 13-22 um, clavate,
S E L . D E S C R . & FIGS. - C h e r u b i n i i n B o l l . A s s . m i c o l . e c o l . R o m a n a narrowly clavate, thick-walled, many w i t h b r o w n contents; pileus cover-
45: 20-22. 1998; M o r e n o et a l . i n M y c o t a x o n 55: 468, fig. 1. 1995 (as ing (velar patches) made up o f erect brown-coloured hyphae, densely
74
CHLOROPHYLLUM (Vellinga) 75
packed; terminal elements 4.0-10 u m w i d e ; clamp-connections not Chlorophyllum molybdites is variable i n its appearance; the D u t c h
observed, but rare i n pileitrama (Sundberg i n M a d r o n o 21:18. 1971). specimen, on w h i c h the macroscopic description given here is based,
H A B I T A T & D I S T R . - Gregarious, i n fairy rings, saprotrophic, terres- is characterized b y a dark extensive velar patch o n the pileus; speci-
trial. In the Netherlands o n l y recorded once, g r o w i n g indoors, i n the mens with smaller, and loose-lying patches, and a very fibrillose c o v -
s o l a r i u m o f a tropical s w i m m i n g p o o l , w i t h imported plants. A l s o ering are often depicted (e.g. I m a z e k i et al., F u n g i Japan: 181. 1988;
reported f r o m Scotland, g r o w i n g i n a s i m i l a r habitat. A pantropical Lincoff, F i e l d G u i d e N . A m e r . Mushr.: p i . 169, 170. 1981).
species, w i t h a w i d e distribution, also i n artificially irrigated lawns etc. There are many reports i n literature o n the toxicity o f this species
i n subtropical areas (e.g. Israel, Canary Islands, N o r t h A m e r i c a ) . (for an overview see R e i d & E i c k e r i n B o t . B u l l . A c a d , sinica 32: 317-
333. 1991). It causes gastrointestinal problems, especially w h e n eaten
A very extensive literature survey o f this species is given by R e i d & raw. T h e effect o n c h i l d r e n is i n general more severe than o n adults.
E i c k e r (in B o t . B u l l . A c a d , sinica 32: 317-333. 1991); both the mor- One report mentioned the presence o f amatoxins (Feinfeld et a l . i n J .
p h o l o g i c a l and the toxic characteristics o f the species are broadly dis- T o x i c o l . C l i n . T o x i c o l . 32: 715-721. 1994).
cussed. B e c a u s e o f its w i d e distribution, and its toxic effects,
Chlorophyllum molybdites is the species o f the lepiotaceous fungi most
often mentioned i n literature.
5. Leucocoprinus Pat.
Leucocoprinus Pat. in J. Bot., Paris 2: 16. 1888. - Mastocephalus Batt. ex O. Kuntze, Rev. Gen. PI. 2: 857. 1891. -
Lepiota subgen. Leucobolbitius J. Lange, Fl. agar. dan. 1: 23. 1935 (not valid). - Lepiota sect. Striatae Murrill in N .
Amer. Fl. 10 (1): 42. 1914.
SELECTED LITERATURE - Candusso & Lanzoni, Lepiota: 458-504. 1990; Migl. in Boll. Gruppo micol. G. Bres., n.S.,
39:14-16; 73-82. 1996.
Basidiocarp pluteoid. Pileus with striate-sulcate marginal zone, thin-fleshed; lamellae free, not changing colour with
ammonia vapour; stipe often bulbous-clavate; annulus present; spore print white to pale pink, pale lilacinous, or cream.
Spores smooth, thick-walled, binucleate, dextrinoid, congophilous, with metachromatic inner wall and tract (if pres-
ent) in Cresyl Blue, cyanophilous, with or without germ pore, with walls swelling in NH3( q) (ammonia) and
a
C H C O O H ( q ) (acetic acid); in most species basidia starting short and squat, growing out to be long and slender,
3 a
always surrounded by pseudoparaphyses; cheilocystidia present; pleurocystidia absent; lamellar trama trabecular;
clamp-connections absent. Development bivelangiocarpic, and pileostipitocarpic. - Type species Lepiota cepaestipes
(Sow.: Fr.) Kumm.
HABITAT & DISTRIBUTION - Solitary to gregarious, sometimes in clumps, saprotrophic and terrestrial, on compost
heaps, more rarely on rotten wood or woodchips, mainly tropical, a few species outside in temperate regions, but also
some species introduced in greenhouses, and appearing indoors with potted plants.
Only the most commonly encountered greenhouse and indoor species are treated here, viz. L. birnbaumii, L.
cepaestipes, L. cretaceus, L. ianthinus, and L. straminellus. Some of these occasionally fruit outside on compost heaps,
piles of leaves or sawdust and woodchips etc. in warm and humid summers. Additional species might be encountered
in hothouses. The selected literature is restricted to European references only.
As the systematics of this genus is under study, a formal infrageneric classification would be premature. There are,
however, two morphologically distinct groups of species, one with big, thick-walled spores with a germ pore, and a
second group with relatively small, thinner walled spores without a germ pore. The sulcate pileus margin, and the pres-
ence of pseudoparaphyses distinguish the second group from species in Cystolepiota and Leucoagaricus with which
they could be confused.
The neutral term 'pileus covering' is used for all the covering layers of the pileus, not taking into account their origin.
In the Neotropics several species belonging to the genera Leucocoprinus and Leucoagaricus are cultivated by attine
ants; for more information on this interesting topic, see e.g. Chapela et al. (in Science 266: 1691-1694. 1994); and
Mueller et al. (in Science 281: 2034-2039. 1998).
K E Y TO T H E SPECIES
1. Elements of pileus covering globose; clamp-connections present Cystolepiota

1. Elements of pileus covering globose, inflated, irregular, cylindrical, lageniform; clamp-connections absent
2. Pileus and stipe turning red on bruising; lamellae green or red with ammonia vapour
see Leucoagaricus Key one
2. Pileus and stipe not turning red on bruising; lamellae not changing colour with ammonia vapour
3. Basidiocarp yellow, sulphur-yellow, or pale yellow
4. Elements on pileus elongate, not globose; spores 8.0-12 X 5.0-9.0 pm, with germ pore 3. L . birnbaumii
4. Elements on pileus surface globose; spores 5.0-8.5 X 4.0-7.0 um, without germ pore 7. L . straminellus
3. Basidiocarp white, cream, with or without coloured (brown, lilac, black) squamules and/or fibrils on pileus or
with brown pileus
76
LEUCOCOPRINUS (Vellinga) 77
5. Pileus brown, especially at centre, or cream with lilac tinges

6. Pileus with lilac centre; stipe lilac in lower part 4. L . ianthinus
6. Pileus with brown or buff tinges
7. Pileus with brownish centre and uplifted squamules; terminal elements of pileus squamules narrowly
lageniform with long, often flexuous neck 1. L . cepaestipes sensu J. Lange
7. Pileus white, at centre cream, and with age with cream buff tips of cottony flocculose warts or
squames; elements of pileus covering cylindrical, often branched and H-shaped or T-shaped, and bone-
like in appearance 2. L . cretaceus
5. Pileus totally white, or white with grey to blackish squamules or fibrils
8. Pileus with black or dark grey, rarely grey, tinges
9. Pileus with dark calotte, contrasting with white surface (set with squamules), rarely with faded, pale
grey, calotte; spores 8.5-13 X 5.0-8.0 pm, with germ pore; growing outdoors; basidiocarps medium-
sized (pileus 15-40(-80) mm) 5. L . brebissonii
9. Pileus with gradual transition between centre and rest, with dark fibrils and small squamules; spores
5.5-9.0(-9.5) X 3.0-4.5(-5.0) pm, without germ pore; growing either in greenhouses or outdoors;
basidiocarps small (pileus 7-20 mm)
10. Spores 6.0-9.0(-9.5) X 3.0-4.5(-5.0) pm, Qav = 1.8-1.85; pileus sulcate in marginal zone; grow-
ing in greenhouses 8. L . heinemannii
10. Spores 5.5-7.0 X 3.5-4.5(-5.0) pm, Qav = 1.55-1.65; pileus not sulcate; growing outdoors
Leucoagaricus melanotrichus var. melanotrichus
8. Basidiocarps totally white8
11. Basidiocarps big (pileus 35-90 mm; stipe 28-110 X 2-13 mm), and warty-floccose on pileus, floccose
on stipe; spores 8.0-12 X 5.5-7.5 pm, with germ pore 2. L . cretaceus
11. Basidiocarps small (pileus 10-35 mm; stipe 9-50 X 1-3 mm), fine-granular-floccose all over or only
on pileus centre; spores 5.0-8.5 X 3.5-7.0 pm, without germ pore
12. Pileus covering rather smooth, made up of radially oriented hyphae, with occasional round
elements; spores ellipsoid-amygdaliform in side-view; growing in natural habitats, most often
on rotten wood 6. L . cygneus
12. Pileus powdery of globose elements; spores subglobose to broadly ellipsoid; growing in green-
houses, rarely outside and then on compost heaps etc.
L. straminellus var. albus (see notes under 7. L. straminellus)
1. Leucocoprinus cepaestipes (Sow.: Fr.) Pat. sensu J . L a n g e , F l . agar, are further apart near margin than near centre, o n creamy background;
dan. 1: p i . 14F. 1 9 3 5 . - F i g . 50. sulcate i n marginal, 5-7 m m wide, zone. L a m e l l a e , L = c. 80,1 = 0-1,
Leucocoprinus cepaestipes f. macrosporus M i g l . in Boll. Ass. rather crowded to crowded, free and remote f r o m stipe, segmentiform
m i c o l . ecol. R o m a n a 6-7: 18. 1986. to ventricose, up to 4 m m w i d e , cream, pale b r o w n i s h cream, dis-
E X C L . - Leucocoprinus cepaestipes sensu D e n n i s i n K e w B u l l . 7: colouring b r o w n w i t h age, w i t h w h i t i s h , fimbriate to flocculose (lens)
462-488. 1952; sensu H o r a k , S y n . G e n . Agar.: 346-348. 1968; sensu edge. Stipe 30-130 X 2.5-5 m m , slightly broadening downwards and
P e g l . , P r e l . A g a r i c F l . East A f r i c a : 318. 1977; sensu C h i u s a i n R i v . tapering upwards, h o l l o w , w h e n y o u n g white, later cream to pale
M i c o l . 4 1 : 514-157. 1998; sensu P e g l . , A g a r i c F l . Lesser A n t i l l e s : 417. b r o w n i s h cream, densely fine-pubescent a l l over, b e l o w annulus w i t h
1983; sensu P e g l . , A g a r i c F l . S r i L a n k a : 324-325. 1986; sensu Pegl. & some small squamules, w h e n y o u n g turning y e l l o w w h e n bruised, later
C a l o n g e i n B o l . S o c . m i c o l . M a d r i d 22: 49-50, f i g . 2. 1997 ( = i n a l l y e l l o w - b r o w n to b r o w n w h e n damaged, w h e n fresh and y o u n g exud-
cases L. cretaceus). ing drops. A n n u l u s ascending, white w h e n y o u n g , w i t h y e l l o w exuda-
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 246. 1995; J . tion drops, w i t h at the margin some thickened squamules, concolorous
L a n g e , F l . agar. dan. 1: p i . 14F. 1935. w i t h those o n pileus, evanescent. Context i n y o u n g specimen white i n
S E L . D E S C R . & F I G S . - Candusso & L a n z o n i , L e p i o t a : 472-474, fig. pileus and stipe, later cream. S m e l l o f cut basidiocarp slightly like the
104. 1990 (as L. cepistipes); H a l l e r i n S c h w e i z . Z . P i l z k . 29: 33-35. smell o f Lepiota cristata, or fruity w i t h some soapy component. Taste
1951 (as Lepiota cepaestipes); M i g l . in B o l l . Ass. micol. ecol. Romana like rubber. Spore print probably white.
6-7: 10-14. 1986 (as L. cepaestipes f. macrosporus). Spores 7.5-13 X 6.0-8.0 um, o n average 8.5-11.1 X 6.6-7.5 um, Q
V E R N . N A M E - Dikvoetplooiparasol. = 1.1-1.7, Q a v = 1.25-1.55, i n side-view e l l i p s o i d , o b l o n g , slightly
amygdaliform, i n frontal v i e w e l l i p s o i d to oblong, t h i c k - w a l l e d , w i t h
P i l e u s w h e n expanded 30-65 m m , w h e n very y o u n g paraboloid, germ pore, congophilous, dextrinoid, cyanophilous, w i t h p i n k inner
expanding to campanulate, and finally plano-convex w i t h flattened w a l l and tract i n C r e s y l B l u e . B a s i d i a 16-37 X 8.0-12 u m , 4-spored,
centre, to applanate, w h e n y o u n g totally granular plush-like, b r o w n - rarely 2-spored, surrounded by 4-6 pseudoparaphyses. L a m e l l a edge
coloured, later at centre plush-like, velvety (lens), dark b r o w n to pale sterile; c h e i l o c y s t i d i a 25-60 X 8-15 u m , clavate, without (in y o u n g
beige-brown ( M u . 10 Y R 4/4 - 7/4), around centre w i t h r a d i a l l y specimens) or w i t h (in mature specimens) apical excrescence, up to 20
arranged, uplifted, s m a l l beige-brown or ochraceous b r o w n (10 Y R 6- X 2.0-3.0 p m , lageniform, colourless. P l e u r o c y s t i d i a absent. Pileus
7/6) squamules, d i s c o l o u r i n g y e l l o w w h e n touched; those squamules covering w i t h squamules, made up o f short squat elements; terminal
w h i c h L. cretaceus is the correct name (see excluded names). However,

the present species, w h i c h must be widespread i n the tropics and is
clearly separated f r o m L. cretaceus, is not represented b y its o w n
description i n Pegler's w o r k . Sowerby ( C o l . F i g s . engl. F u n g i 1: p i . 2.
1796) introduced Agaricus cepaestipes to accommodate both Agaricus
cretaceus B u l l , and A . luteus B o l t , w h i c h he considered colour variants
o f the same species.
2. Leucocoprinus cretaceus ( B u l l A F r . ) L o c q . i n B u l l . mens. Soc. l i n n .

L y o n 14: 93. 1945. - F i g . 5 1 . \
Agaricus cretaceus B u l l . , Herb. France: p i . 374. 1788; Agaricus
cretaceus B u l l . : Fr., Syst. m y c o l . , Ind. g e k : 14. 1832, non Agaricus
cretaceus Pers., 1801, ntc Agaricus cretaceus Pers.: Fr., 1821; Lepiota
cretacea ( B u l l . : Fr.) Q u e l . , F l . m y c o l . France: 2 9 8 . 1 8 8 1 . -Lepiota cre-
tata L o c q . i n H a l l e r i n M i t t , aargau. naturf. G e s . 2 3 : 82. 1950 (not
valid); Leucocoprinus cretatus ( L o c q . ) M o s . , Blatter-, B a u c h p i l z e , 1.
A u f l . : 116. 1953 (not valid); Leucocoprinus cretatus L o c q . ex L a n z o n i
i n A t t i X I X C o m . scient. naz. Serina (Bergamo): 30-31. 1986.
EXCL. - Leucoagaricus cretaceus sensu M o s . , Rohrlinge
Blatterpilze, 3. A u f l . : 185. 1967; sensu K n u d s e n i n H a n s e n &
Knudsen, Nordic M a c r o m y c . 2: 2 2 3 . 1992 ( = i n both cases
Leucoagaricus leucothites). - Lepiota cretacea sensu M a i . & Bert., F l .
C h a m p , sup. M a r o c 1: 120-121, fig. 16. 1970 ( = u n k n o w n species). I
M I S A P P L . - Leucocoprinus cepaestipes sensu C h i u s a i n R i v . M i c o l .
41: 5 1 4 - 1 5 7 . 1 9 9 8 ; sensu Pegl., Prel. A g a r i c F l . East A f r i c a : 3 1 8 . 1 9 7 7 ;
sensu P e g l . , A g a r i c F l . Lesser A n t i l l e s : 417. 1983; sensu Pegl., A g a r i c
F l . S r i L a n k a : - 3 2 4 - 3 2 5 . 1986; sensu P e g l . & C a l o n g e i n B o l . S o c .
m i c o l . M a d r i d 2 2 : 49-50, fig. 2. 1997. - Lepiota cepaestipes sensu
F i g . 50. Leucocoprinus cepaestipes. Lincoff, F i e l d G u i d e N . A m e r . M u s h r . : 515, p i . 179. 1981
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 60. 1990 (as L. cre-
elements 31-110 X 5-9 p m , 3.0-5.0 u m at apex, cylindrical, lageniform tatus); Lincoff, F i e l d G u i d e N . A m e r . Mushr.: p i . 179. 1981 (as Lepiota
with l o n g flexuous neck, w i t h y e l l o w i s h intracellular pigment. cepaestipes); M i g l . i n B o l l . G r u p p o m i c o l . G . Bres. 39: 4, 6. 1996 (as
Stipitipellis a cutis o f narrowly c y l i n d r i c a l hyphae, 2.0-4.0 u m wide, L. cretatus); P e g l . & C a l o n g e i n B o l . S o c . m i c o l . M a d r i d 22: f i g . 2.
w i t h colourless c y s t i d i o i d elements i n tufts, as terminal elements o f 1997 (as L. cepaestipes).
pileus squamules: 45-90 X 6.5-9.0 u m , 3.0-4.0 u m at apex, narrowly S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 477-478, fig.
lageniform w i t h l o n g tapering neck. Clamp-connections absent. 105. 1990 (as L. cretatus); M i g l . i n B o l l . G r u p p o m i c o l . G . B r e s . 39:
H A B I T A T & DISTR. - In s m a l l c l u m p s and groups, saprotrophic and 5-8, figs. 1-3.1996 (as L. cretatus); R e i d & E i c k e r i n S. Afr. J . B o t . 59:
terrestrial, o n s o i l i n greenhouses, a n d o n w o o d chips and compost 92-95, figs. 16-19. 1993.
heaps outside. I n the Netherlands rather rare, and recently repeatedly V E R N . N A M E - Krijtwitte plooiparasol.
recorded f r o m the outside (Wassenaar, A p e l d o o r n ) . E n d o f July-begin-
n i n g o f Sept. (both inside and outside). Widespread i n Europe, m a i n l y Pileus 38-90 m m , at first hemispherical to obtusely c o n i c a l or c o n i c o -
i n greenhouses etc., o f tropical o r i g i n . convex or convex, w i t h m a r g i n i n f l e x e d at first, later expanding to
plano-concave w i t h or without a vague l o w umbo, w h e n fully mature
Leucocoprinus cepaestipes i n the present sense has the f o l l o w i n g w i t h sulcate m a r g i n a l 2-3 m m w i d e zone, thin-fleshed (1-2.5 m m
m i c r o s c o p i c a l characters: c h e i l o c y s t i d i a clavate w i t h i n full-grown above m i d d l e o f lamellae), white w i t h cream-buff tinges at centre, at
specimens abrupt apical, c y l i n d r i c a l excrescence; narrowly lageniform first completely covered w i t h c o n i c a l , about 2 m m i n d i a m . and u p to
to c y l i n d r i c a l elements on pileus and stipe surfaces. T h e spores are, as 2.5 m m h i g h soft c o n i c a l warts to merely irregularly floccose-squa-
i n m a n y species o f the genus, variable i n size and shape, depending o n mulose and these warts white to cream-buff tinged, usually at centre
age and g r o w i n g conditions o f the basidiocarps. F o r this reason w i t h pale b r o w n i s h tips; pileus surface dry between warts, r a d i a l l y
M i g l i o z z i ' s forma macrosporus (described i n i n B o l l . A s s . m i c o l . ecol. w o o l l y - f i b r i l l o s e (lens); m a r g i n h u n g w i t h fringes and so exceeding
R o m a n a 6 - 7 : 10-14. 1986) i s not recognized here. lamellae. L a m e l l a e , L = 85-100,1 = 0-3, rather crowded, free and dis-
Leucocoprinus cepaestipes var. rorulentus ( P a n i z z i ) B a b o s is a tant f r o m stipe, very thin, (sub)ventricose, u p to 7 m m broad, white to
white variant, i n w h i c h the lamellae turn p i n k w i t h age ( P a n i z z i i n very pale sordid-creamy whitish, w i t h entire, concolorous edge. Stipe
C o m m . S o c . critt. ital. 1: 172. 1861). 28-110 X 2-13 m m , attenuate upwards, w i t h strongly enlarged bulbous
Variants w i t h lilac-grey to greyish b r o w n tinges i n the pileus have or clavate to fusiform base (7-19 m m w i d e ) and somewhat rooting,
been found, both indoors i n heated and non-heated glasshouses, and solid, but later stuffed, white at first but taking very slight ochraceous
on a p i l e o f leaves outside (in a botanical garden). T h e cheilocystidia buff tinge, particularly i n upper part, minutely felted-flocculose above
are different f r o m the t y p i c a l L. cepaestipes, but c a u l o c y s t i d i a and annulus, w i t h rather t h i c k fine-warted c o v e r i n g to finely flocculose-
pileus c o v e r i n g elements are similar. These collections have been squamulose i n transverse rows below annulus, w i t h stipe surface under
excluded f r o m the above description. covering d i s c o l o u r i n g y e l l o w i s h w h e n bruised, w i t h white m y c e l i u m .
E n g l i s h authors, e.g. Pegler ( A g a r i c F l . S r i L a n k a : 324-325. 1986), A n n u l u s c. halfway up stipe, first flaring, but then pending, 5-10 m m
have used the name L. cepaestipes for a white tropical species, for wide, thin, felted-submembranaceous, smooth at upper surface,
F i g . 5 1 . Leucocoprinus cretaceus.
minutely flocculose at l o w e r surface, w i t h b r o k e n thicker r i m at inside non-heated glasshouses, but i n recent years also outside ( A p e l d o o r n ,
of m a r g i n . C o n t e x t w h i t i s h to pale buff, w i t h age somewhat sordid, Leusden), but rare. Outside ( A u g . ) Sept., inside the w h o l e year
rather soft and pliable except i n cortex o f stipe. S m e l l variable, record- through. Scattered i n E u r o p e , m o s t l y recorded f r o m greenhouses,
ed as weak, t y p i c a l l y l e p i o t o i d and later l i k e the s m e l l o f Inocybe bon- widespread i n the tropics.
gardii, or as rather strong, unpleasant, rather nauseating (also i n y o u n g
specimens); w h e n crushed different, more herbaceous, suggestive o f Leucocoprinus cretaceus is taken here i n the original sense o f B u l l i a r d
Pelargonium. Taste strong, unpleasant, astringent (lepiotoid), not bit- (Herb. France: p i . 374. 1788). B u l l i a r d ' s description under the plate
ter. Spore print off-white to pale c r e a m ( R o m a g n e s i l b - 2 a ) . clearly stated that the pileus c o v e r i n g was cottony or p l u s h - l i k e and
Spores 8.0-12 X 5.5-7.5 p m , o n average 8.7-10.5 X 5.9-6.8 um, Q chalk-white. Some authors (e.g. H a l l e r i n M i t t , aargau. naturf. Ges. 23:
= 1.3-1.95, Q a v = 1.4-1.65, i n side-view e l l i p s o i d to o b l o n g , slight- 82-85. 1950) have argued that B u l l i a r d ' s somewhat stylized plate
ly a m y g d a l i f o r m , sometimes w i t h s h a l l o w suprahilar depression, i n depicts a species w i t h a smooth pileus surface, n o w c o m m o n l y k n o w n
frontal v i e w e l l i p s o i d to o b l o n g , or slightly o v o i d , t h i c k - w a l l e d , w i t h as Leucoagaricus leucothites (Vitt.) Wasser, and consequently have
g e r m pore, often w i t h b u l g i n g h y a l i n e cap over germ pore, dextrinoid, used a different name for the present species. K n u d s e n (in Hansen &
congophilous, c y a n o p h i l o u s , w i t h p i n k inner w a l l and tract i n C r e s y l K n u d s e n , N o r d i c M a c r o m y c . 2: 223. 1992) still used Leucoagaricus
B l u e . B a s i d i a 18-28 X 8.5-12 p m , 4-spored, surrounded b y 4-6 cretaceus for the species c a l l e d L. leucothites i n the present work.
pseudoparaphyses. L a m e l l a edge sterile; c h e i l o c y s t i d i a (30-)40-100 The name Agaricus cretaceus B u l l , was sanctioned b y Fries i n the
X 7-14(-21) p m , n a r r o w l y clavate to fusiform, n a r r o w l y lageniform, Index to the Systema m y c o l o g i c u m (1832: 14), and so has priority over
c y l i n d r i c a l , sometimes w i t h short a p i c a l excrescence, slightly m o n i l - Agaricus cretaceus Pers. ( S y n . meth. F u n g . : 349. 1801), w h i c h was
i f o r m , rarely clavate, colourless, and rather t h i n - w a l l e d to s l i g h t l y also sanctioned b y Fries (Syst. m y c o l . 1: 95. 1821).
thick-walled, sometimes with amorphous matter at apex. L o c q u i n (in H a l l e r i n M i t t , aargau. naturf. G e s . 23: 82. 1950) pro-
P l e u r o c y s t i d i a absent. P i l e u s c o v e r i n g w i t h warts and floccules, made posed the name Lepiota cretata for the present species. T h i s has been
up o f elongate colourless elements, c y l i n d r i c a l , branched, w i t h excres- w i d e l y followed, o v e r l o o k i n g the many older synonyms, based on trop-
cences, H - s h a p e d or T-shaped, w i t h straight and often w i d e n e d ends, i c a l collections, w h i c h are available (see Pegler, A g a r i c F l . S r i L a n k a :
often a bit b o n e - l i k e i n appearance, 5.0-10 p m w i d e . S t i p i t i p e l l i s a 323-324. 1986 for an extensive list, under L. cepaestipes).
cutis o f narrow, colourless, c y l i n d r i c a l hyphae, 2.0-4.0 p m w i d e , w i t h Pegler (e.g. A g a r i c F l . S r i L a n k a : 323-324. 1986), and other E n g l i s h
floccules made up o f elements similar to those o f squamules o n pileus. authors, have used the name L. cepaestipes (Sow.: Fr.) Pat. for this
C l a m p - c o n n e c t i o n s absent. white species. T h e name Leucocoprinus cepaestipes is nowadays most
H A B I T A T & D I S T R . - In bundles, i n groups, saprotrophic and terres- c o m m o n l y used for a species w i t h b r o w n squamules (e.g. Candusso &
trial, often o n sawdust or w o o d c h i p s , compost heaps etc., i n green- L a n z o n i , L e p i o t a : 472-474. 1990), f o l l o w i n g L a n g e ' s interpretation
houses, or outdoors. I n the Netherlands k n o w n f r o m heated and ( F l . agar. dan. 1: p i . 14F. 1935).
Cystolepiota pulverulenta is also a clampless, white, floccose- to 4 m m wide, pale-yellow, w i t h strongly eroded, more intense y e l l o w
warty species, but the basidiocarps discolour o n bruising, the spores to b r o w n i s h edge. Stipe 30-80 X 2-5 m m , at base distinctly clavate-
are s m a l l ( v i z . (3.5-)4.0-5.5(-6.0) X (2.0-)2.5-3.5 um) and they lack a bulbous, up to 7 m m wide, hollow, first l e m o n - y e l l o w to sulphur-yel-
germ pore. T h i s species occasionally grows i n pots o f houseplants and low, later b r o w n i s h , over total length fine-squamulose, except for just
in greenhouses. above base and there distinctly yellow. A n n u l u s membranaceous, y e l -
l o w o n upper side, w h i t i s h o n underside, evanescent. Context thin and
3. Leucocoprinus birnbaumii (Corda) S i n g , i n S y d o w i a 15: 67 soft, d u l l and w h i t i s h i n pileus, shiny w h i t i s h i n stipe. S m e l l indistinct.
( ' 1 9 6 1 ' ) 1962. - F i g . 5 2 . Taste indistinct. Spore print white.
Agaricus birnbaumii C o r d a , Icon. F u n g . 3: 4 8 . 1839. - Agaricus Spores 8.0-12 X 5.0-9.0 u m \ o n average 8.9-9.9 X 6.3-7.2 um, Q =
flos-sulphuris S c h n i z l . i n Sturm, Deutschl. F l . 3, A b t . 31. 32. Heft: 2. 1.25-1.6, Q a v = 1.35-1.45, i n side-view amygdaliform-elhpsoid, ellip-
1851; Lepiota flos-sulphuris (Schnizl.) M a t t i r o l o i n A t t i A c c a d . naz. soid, i n frontal v i e w ellipsoid or ovoid, thick-walled, w i t h germ pore and
L i n c e i M e m o r i e , Ser. V , 12 (11): (566) 34. 1918; Leucocoprinus flos- hyalinous cap over germ pore, dexninoid, congophilous, cyanophilous,
sulphuris (Schnizl.) C e j p i n C e s k a M y k o l . 2: 78. 1948; Lepiota with p i n k inner w a l l and tract i n C r e s y l B l u e . B a s i d i a 16-40 X 7.5-10
cepaestipes vex. flos-sulphuris (Schnizl.) J . R i c k i n Iheringia, B o t . 8: um, 4-spored, surrounded by 4-6 pseudoparaphyses. L a m e l l a edge ster-
314. 1961 (not v a l i d ) . - A g a r i c u s luteus B o l t . , Hist. F u n g . Halifax 2: p i . ile; cheilocystidia 25-70 X 11-17 um, variable i n shape and size, lageni-
50. 1788, non Agaricus luteus Huds., 1778, nec Agaricus luteus Huds.: form w i t h 3.0-6.0 u m wide neck, utriform, narrowly clavate, thin-walled
Fr., 1821; Lepiota lutea (Bolt. 0 ) G o d f r i n i n B u l l . Soc. m y c o l . Fr. 13: and colourless. Pleurocystidia absent. Pileus covering w i t h squamules
33. 1897; Leucocoprinus luteus (Bolt. 0 Godfrin) L o c q . i n B u l l . mens. made up o f elongate, inflated, sometimes branched elements, 5.0-15 u m
Soc. l i n n . L y o n 14: 9 3 . 1 9 4 5 . - Lepiota aurea M a s s . i n K e w B u l l . 1912: wide, and some scattered ellipsoid and clavate elements, up to 35 X 23
189. 1912, n o n Lepiota aurea (Mattuschka: Fr.) S . F Gray, 1821. - um. Pigment yellowish-brownish, granular and diffuse, intracellular.
Lepiotapseudolicmophora R e a , B r i t . B a s i d i o m y c : 74. 1922. - Lepiota Stipitipellis a cutis o f narrow cylindrical, yellowish, hyphae, 2.0-5.0 um,
coprinoides B e e l i i n F l . icon. C h a m p . C o n g o 2: 42. 1936. with tufts o f irregular tangled hyphae, with rather short elements, c y l i n -
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 244. 1995; drical, almost cylindrical, often branched, around 5.0 p m i n diam., with
D a h n c k e & D a h n c k e , 700 P i l z e : 318. 1979; Imaz. et al., F u n g i Japan: yellowish intracellular pigment. Clamp-connections absent.
185. 1988; J . L a n g e , F l . agar. dan. 1: p i . 1 4 G . 1935 (as Lepiota luteal?, H A B I T A T & DISTR. - Solitary and i n clumps o f a few specimens, sapro-
M i g l . et a l . i n R i v . M i c o l . 32: 2 4 . 1 9 8 9 ; Rocabruna i n Bolets Catalunya trophic and terrestrial i n houseplant pots indoors, and i n greenhouses.
3: p i . 130. 1984. C o m m o n , throughout the year, widespread i n Europe, of tropical origin.
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 465-466, fig.
102. 1990; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 174-175. 1994); Annotations w i t h D u t c h material stated that the spore print is white;
M i g l . et a l . i n R i v . M i c o l . 32: 24-26. 1989. surprisingly Breitenbach & K r a n z l i n (Pilze S c h w e i z 4: p i . 244. 1995)
V E R N . N A M E - Goudgele plooiparasol. said that the spore print is pale pink.
Pileus w h e n expanded 20-60 m m , when very y o u n g paraboloid, then 4. Leucocoprinus ianthinus ( C o o k e 0 Sacc.) L o c q . i n B u l l . mens.
hemispherical, and expanding to applanate, or plano-convex, w i t h or Soc. l i n n . L y o n 14: 94. 1945. - F i g . 5 3 .
without umbo, bright sulphur y e l l o w w h e n young, w i t h easily remov- Agaricus ianthinus C o o k e i n G r e v i l l e a 16: 101. 1888, non Agaricus
able floccules, pallescent to pale y e l l o w - b r o w n when o l d , w i t h y e l l o w - janthinus Fr.: Fr., 1821; Lepiota ianthina (Cooke 0 ) S a c c , Syll. Fung.
b r o w n squamules, f o r m i n g a patch at centre, i n o l d specimens w i t h 9: 10. 1891; Leucocoprinus ianthinus (Cooke 0 Sacc.) M o h r i n
distinctly r a d i a l l y grooved marginal zone, w h e n y o u n g w i t h y e l l o w Boletus 18: 4 8 . 1992 (superfluous). - Lepiota lilacinogranulosa P.
fringes at m a r g i n . L a m e l l a e , L = 50-70, 1 = 1, rather c r o w d e d to H e n n . i n Verh. bot. ver. Prov. Brandenb. 40: 145. 1898; Leucocoprinus
crowded, free and remote f r o m stipe, segmentiform to ventricose, up lilacinogranulosus (P. Henn.) L o c q . i n B u l l . mens. S o c . l i n n . L y o n 12:
94. 1943; Leucocoprinus lilacinogranulosus (P. Henn.) M . B o n et a l .
in D o c . m y c o l . 19 (75): 5 3 . 1989 (superfluous); Hiatula cepaestipes
var. lilacinogranulosa (P. Henn.) H e i m & R o m a g n . i n B u l l , trimest.
Soc. m y c o l . Fr. 50: 184. 1934.
S E L . I C O N . - H e i m & R o m a g n . i n B u l l , trimest. S o c . m y c o l . Fr. 50:
p i . 11, f i g . 1. 1934 (as Hiatula cepaestipes var. lilacinogranulosa);
M i g l . et a l . i n R i v . M i c o l . 32: 21. 1989 (as L. lilacinogranulosus);
S E L . D E S C R . & F I G S . - B a b o s i n A g a r i c a 6 (12): 212-213, fig. 14.
1985 (as L. lilacinogranulosus); Candusso & L a n z o n i , L e p i o t a : 4 8 4 -
486, fig. 107. 1990 (as L. lilacinogranulosus); G . Fischer i n U l m e r
P i l z f l . 1: 136. 1986; H e i m & R o m a g n . i n B u l l , trimest. S o c . m y c o l . Fr.
50: 184-185, fig. 12. 1934 (as Hiatula cepaestipes var. lilacinogranu-
losa); M i g l . et al. i n R i v . M i c o l . 32: 20-23. 1989 (as L. lilacinogranu-
losus); D . R e i d i n M y c o l . Res. 9 3 : 418. 1989 (as L. ianthinus and L.
lilacinogranulosus resp.).
V E R N . N A M E - Lilakorrelige plooiparasol.
Pileus i n expanded specimens 17-62 m m , w h e n y o u n g closed and

p a r a b o l o i d w i t h flattened apex to o v o i d , expanding to broadly c a m -
panulate, conical-campanulate and finally plano-convex w i t h b i g , trun-
cate umbo, and uplifted margin, at centre finely fibrillose-squamulose
F i g . 52. Leucocoprinus birnbaumii. felted, purplish, reddish b r o w n ( K . & W . 8 E 6 , 9 D 6 , 7 E 5 ) , around cen-
27-80 X 9.0-35 p m , v a r y i n g f r o m c y l i n d r i c a l , u t r i f o r m to broadly

clavate, thin-walled, colourless. P l e u r o c y s t i d i a absent. Pileus covering
made up o f irregular chains o f c y l i n d r i c a l to e l l i p s o i d to globose ele-
ments, w i t h adnate to ascending elements, ranging from 15-35 X 5.0-
9.0 p m to 25 p m i n d i a m . ; pigment intracellular and l i l a c - b r o w n to
greyish lilac. Stipitipellis a cutis o f c y l i n d r i c a l colourless hyphae, 4.0-
13 p m i n diam., w i t h at base o f stipe s i m i l a r elements as o n pileus sur-
face. Clamp-connections absent.
H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial i n
pots o f plants, i n greenhouses. Rather rare i n the Netherlands, found
throughout the year, widespread i n Europe, probably o f tropical origin.
T h e name L. ianthinus is used for the species, more c o m m o n l y k n o w n

as L. lilacinogranulosus, f o l l o w i n g B o n ( F l . m y c o l . Eur. 3, Lepiotes:
112. 1993); R e i d (in M y c o l . Res. 93: 418. 1989) gave arguments for
and against this synonymy, but he d i d not reach a c o n c l u s i o n .
R e i d (in M y c o l . R e s . 93: 420. 1989) distinguished L. lilacinogran-
ulosus var. subglobisporus w i t h smaller, subglobose spores; this vari-
ety has not yet been recorded f r o m the Netherlands. Z u c c h e r e l l i &
M i g l i o z z i (in B o l l . G r u p p o m i c o l . G . Bres., n.S., 4 1 : 197-203. 1998)
reported o n its occurrence i n Italy.
Specimens w i t h grey-lilac to l i l a c - b r o w n tinges and narrowly
lageniform to c y l i n d r i c a l elements o n pileus and stipe are described
under L. cepaestipes.
5. Leucocoprinus brebissonii (Godey) L o c q . i n B u l l . mens. S o c . l i n n .

L y o n 12: 41. 1 9 4 3 . - F i g . 54.
Lepiota brebissonii G o d e y i n G i l l e t , H y m e n o m y c e t e s : 64. 1874. -
Leucocoprinus otsuensis H o n g o i n J . Jap. B o t . 28: 70. 1953.
M I S A P P L . -Agaricus clypeolarius sensu Sow., C o l . 111. E n g l . F u n g i
2: pi. 171. 1798.
S E L . I C O N . - A r n o l d s & V e e r k a m p , G i d s Paddest. Meetnet: 76.
1999; Krieglst. i n Z . M y k o l . 48: opp. p. 64. 1982; J . L a n g e , F l . agar,
dan. 1: p i . 1 4 H . 1935; R . P h i l l i p s , Paddest. S c h i m m . : 30. 1981.
F i g . 53. Leucocoprinus ianthinus.
tre w i t h s m a l l p u r p l i s h fibrils, especially o n the ridges, but mostly

white or cream especially i n outer half, and there without p u r p l i s h fib-
rils, costate-plicate up to 2/3 o f radius or up to umbo, at utmost mar-
gin y e l l o w i n g w i t h age, easily r a d i a l l y splitting, very thin-fleshed.
L a m e l l a e , L = 35-55,1 = (0-)l(-2), moderately distant to moderately
c r o w d e d , free and remote f r o m stipe, segmentiform-linear to ventri-
cose, narrow, up to 3 m m w i d e , sometimes anastomosing-furcate,
white w i t h vague l i l a c i n o u s sheen o n some places w h e n older, w i t h
finely flocculose white edge. Stipe 35-70(-150) X 2-5 m m , broaden-
ing downwards and there bulbous-clavate, h o l l o w , i n apical part y e l -
l o w i s h white, shiny, lengthwise fibrillose, b e l o w annulus white, or pale
y e l l o w i s h white w i t h age, i n lower half and at at bulb w i t h purplish-
violaceous fibrils to complete l i l a c c o v e r i n g (e.g. 1 1 C 4 ) , w i t h white
tomentum at bulb itself, d i s c o l o u r i n g y e l l o w w h e n bruised. A n n u l u s
ascending, and slightly flaring, white, w i t h p u r p l i s h m a r g i n , evanes-
cent. Context very thin and soft i n pileus, white, concolorous w i t h sur-
face i n stipe (but not lilac-tinged). S m e l l o f cut basidiocarp fungoid,
strong, slightly astringent. Taste fungoid. Spore print probably pale
lilacinous to white.
Spores 8.0-12 X 5.5-7.5 um, o n average 9.4-10.4 X 6.5-6.7 p m , Q
= 1.25-1.7, Q a v = 1.4-1.55, i n side-view e l l i p s o i d - a m y g d a l i f o r m ,
some w i t h a p i c a l p a p i l l a , i n frontal v i e w e l l i p s o i d to slightly o v o i d ,
t h i c k - w a l l e d , w i t h g e r m pore and h y a l i n e cap over pore, dextrinoid,
congophilous, cyanophilous, w i t h p i n k inner w a l l and tract i n C r e s y l
B l u e . B a s i d i a 17-42 X 8.0-11 p m , 4-spored, rarely 2-spored, sur-
rounded b y 4-6 pseudoparaphyses. L a m e l l a edge sterile; cheilocystidia F i g . 54. Leucocoprinus brebissonii.
•J
S E L . DESCR. & FIGS. - B a b o s i n A g a r i c a 6 (12): 208, 2 1 0 . 1985;

Candusso & L a n z o n i , L e p i o t a : 469-470, fig. 103. 1990; H o t z i n
S c h w e i z . Z . P i l z k . 6 1 : 89. 1983; K r i e g l s t . i n Z . M y k o l . 4 8 : 54-55.
1982; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 176-177. 1994; K u h n e r
in B u l l , trimest. S o c . m y c o l . F r . 52: 237-238. 1936.
V E R N . N A M E - Spikkelplooiparasol.
Pileus when expanded 20-60(-80) m m , when very young paraboloid,

then c o n i c a l , w i t h rounded apex and expanding v i a campanulate or con-
vex to plano-convex and applanate w i t h l o w umbo, or without any
F i g . 55. Leucocoprinus cygneus (habit del. P. Kelderman).
umbo, very dark grey-brown, grey-brown, dark b r o w n to almost black
( M u . 10 Y R - 2/5 Y 3/2), and velvety or finely squamulose-granular at
centre; centre w e l l - l i m i t e d ; around centre w i t h small concentrical dark cygnea (J. Lange) L o c q . i n B u l l , trimest. Soc. m y c o l . Fr. 68: 169. 1952;
grey-brown to blackish squamules and more towards margin w i t h fine Sericeomyces cygneus (J. L a n g e ) H e i n e m . i n B u l l . Jard. bot. natn.
dark brown-grey to b l a c k i s h fibrils on cream-coloured or whitish back- B e l g . 48: 405. 1978; Cystolepiota cygnea (J. L a n g e ) M o s . , R o h r l i n g e
ground, rarely w i t h faded colours on centre and squamules, slightly y e l - Bliitterpilze, 4. A u f l . : 236. 1978.
l o w i n g w h e n damaged, when y o u n g scarcely sulcate at m a r g i n later S E L . I C O N . - Boisselet & M i g l . i n B u l l . F e d . m y c o l . D a u p h i n e -
more distinctly so, up to 1/4 o f radius. Lamellae, L = 35-60,1 = 0-3, Savoie 138: 9. 1995; J . L a n g e , F l . agar. dan. 1: p i . 1 3 A . 1935 (as
moderately crowded, free, some furcate-anastomosing, segmentiform Lepiota cygnea); M i g l . et al. i n R i v . M i c o l . 32: 18. 1989.
to slightly ventricose, up to 3.5 m m broad, white, pale cream-coloured S E L . DESCR. & FIGS. - Boisselet & M i g l . i n B u l l . F e d . mycol.
or slightly y e l l o w i s h , w i t h white, eroded to flocculose edge. Stipe 35- Dauphine-Savoie 138: 4-5, figs. 1 & 2. 1995; Candusso & L a n z o n i ,
80 X 2-5 m m , gradually w i d e n i n g towards clavate, up to 8 m m wide L e p i o t a : 482, fig. 106. 1990; H u i j s m a n i n M e d e d . N e d . m y c o l . Vereen.
base, h o l l o w , white and pinkish-brownish i n lower part, finely floccu-
28: 51-53. 1943; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 178-179.
lose-pruinose to dark fibrillose-flocculose, especially i n lower part,
1994; M i g l . et a l . i n R i v . M i c o l . 32: 18-20. 1989; R o m a g n . i n K u h n e r
white tomentose on base. A n n u l u s ascending, white, tomentose, evanes^*
& R o m a g n . i n B u l l . S o c . nat. O y o n n a x 10-11: 92-94. 1957 ( C o m p l . F l .
cent. Context white i n pileus, i n stipe white i n upper part and brownish
anal. 7).
i n lower part. S m e l l indistinct, either fungoid, or reminiscent o f the
V E R N . N A M E - Gladde plooiparasol.
smell o f Lepiota cristata. Taste not unpleasant to indistinct. Spore print
pale cream-coloured.
Pileus 4-15 m m , when young spherical, expanding v i a plano-convex
Spores 8.5-13 X 5.0-8.0 p m , o n average 9.3-11.5 X 5.6-7.0 um, Q w i t h umbo and uplifted margin to applanate, white, slightly ochraceous
= 1.35-2.1, Q a v = 1.6-1.8, i n side-view e l l i p s o i d - a m y g d a l i f o r m to w i t h age, w h e n y o u n g w i t h scattered greyish granulose flocks, later
c y l i n d r i c a l - a m y g d a l i f o r m , w i t h or without elongated papilla-like apex, only slightly granulose-farinose at centre, and towards margin radial-
in frontal v i e w e l l i p s o i d - o v o i d to c y l i n d r i c a l - o v o i d , thick-walled, w i t h ly fibrillose, sulcate i n outer half o f radius. L a m e l l a e , L = c. 4 0 , 1 =
g e r m pore, dextrinoid, congophilous, cyanophilous, w i t h p i n k inner 0-1, rather crowded to very crowded, free, ventricose, white w i t h c o n -
w a l l and tract i n C r e s y l B l u e . B a s i d i a 20-43 X 8.0-11 p m , 4-spored, colorous flocculose edge. Stipe 9-28 X 0.5-2 m m , c y l i n d r i c a l w i t h
x
rarely 2-spored, surrounded by 4-6 pseudoparaphyses. L a m e l l a edge widened clavate base, h o l l o w , white, pruinose i n upper part, slightly
sterile; c h e i l o c y s t i d i a i n tufts, 34-80 X 8.0-18.5 p m , c y l i n d r i c a l to nar- hairy-pubescent below annulus. A n n u l u s ascending, membranaceous,
r o w l y clavate, or subutriform, or sublageniform, thin-walled to slight- white. Context white and very thin i n pileus, white i n stipe. S m e l l
ly t h i c k - w a l l e d , especially i n apical part, some w i t h some s m a l l indistinct. Taste not k n o w n . Spore print probably white.
crystals at apex, colourless. P l e u r o c y s t i d i a absent. P i l e u s covering Spores (5.5-)6.0-7.0(-7.5) X 3.5-4.5 u m , on average 6.5 X 4.0, Q
made up o f c y l i n d r i c a l and short to ellipsoid-clavate to globose ele- = 1.4-1.8(-1.9), Q a v = 1.6, ellipsoid-amygdaliform i n side-view, ellip-
ments, i n chains or disorderly arrangements; elements up to 15-50(- soid to o v o i d i n frontal view, dextrinoid, congophilous, w i t h p i n k inner
70) X 8.0-30 p m ; towards m a r g i n ascending, narrowly clavate w a l l i n C r e s y l B l u e ; i n a m m o n i a germ pore not visible, after treatment
elements are predominant; pigment dark grey-brown and intracellular.
with acetic a c i d i n some spores faintly visible. B a s i d i a 15-20 X 6.5-8.0
Stipitipellis a cutis o f colourless, cylindrical to slightly inflated hyphae,
pm, 4-spored. L a m e l l a edge sterile; c h e i l o c y s t i d i a 24-65(-100) X 6-
6.0-18 p m w i d e , w i t h scattered clavate and narrowly clavate c y s t i d i o i d
15(-18) p m , lageniform w i t h l o n g neck (up to 5 0 X 3.0-7.0 p m ) and
elements, c. 16-20 u m w i d e , w i t h brown-grey intracellular pigment.
w i t h subcapitate apex, rarely clavate, or w i t h short excrescence, not-
Clamp-connections absent.
coloured. Pleurocystidia absent. Pileus covering w i t h radial c y l i n d r i c a l
H A B I T A T & DISTR. - Gregarious i n small groups, saprotrophic and hyphae, 3.0-10 p m w i d e , w i t h colourless encrusting pigment, and w i t h
terrestrial, i n woods o n various soil types, from sand to clay, probably chains o f e l l i p s o i d to globose elements, 15-35 X 15-25 p m , w i t h intra-
always relatively nutrient-rich. N o t u n c o m m o n i n the Netherlands, cellular greyish pigment i n floccules, and at centre o f pileus. C l a m p -
scattered, A u g . - O c t . ; widespread i n Europe. connections absent.
Leucocoprinus brebissonii is very easy to recognize i n the field, because terrestrial or o n rotting w o o d o f stumps; i n the Netherlands very rare,
of its contrasting dark centre on the mostly white pileus. It is possibly recorded from the dunes on Walcheren (on a stump) and from south-
one o f those species whose numbers are increasing i n the Netherlands, ern L i m b u r g (in a deciduous wood), beginning o f Sept. R a r e l y record-
because o f ongoing nitrogen enrichment o f the environment. ed i n the rest o f E u r o p e ; mentioned from a greenhouse i n Italy.
6. Leucocoprinus cygneus (J. L a n g e ) M . B o n i n D o c . m y c o l . 8 (30- T h e description is based o n the selected descriptions, as cited.
31): 7 0 . 1 9 7 8 . - F i g . 5 5 . Leucocoprinus cygneus is aberrant f r o m the other European
Lepiota cygnea J . L a n g e , F l . agar. dan. 1: 3 5 . 1935 (not valid); Leucocoprinus taxa i n the shape o f the spores and the peculiar archi-
Lepiota cygnea J . L a n g e , F l . agar. dan. 5: V . 1940; Pseudobaeospora tecture o f the pileus covering.
LEUCOCOPRINUS {Vellinga) 83
7. Leucocoprinus straminellus ( B a g l . ) N a r d u c c i & Caroti in Spores 5.0-8.5 X 4.0-7.0 u m , o n average 5.4-7.1 X 4.2-5.3 um, Q
M e m o r i e S o c . tosc. S c i . nat., Ser. B , 102: 4 9 . ('1995') 1996. - F i g . 56. = 1.1-1.6, Q a v = 1.25-1.35, broadly e l l i p s o i d to e l l i p s o i d i n side-view
Agaricus straminellus B a g l . i n C o m m . S o c . critt. ital. 2 (2): 2 6 3 . and i n frontal view, without g e r m pore, dextrinoid, congophilous,
1865; Lepiota straminella (Bagl.) S a c c , S y l l . F u n g . 5: 4 4 . 1887; cyanophilous, w i t h p i n k inner w a l l i n C r e s y l B l u e . B a s i d i a 15-31 X
Lepiota cepaestipes var. straminella (Bagl.) Konr. & M . , Ic. sel. F u n g . 6.0-8.0 p m , 4-spored, surrounded b y 4-6 pseudoparaphyses. L a m e l l a
6: 4 1 . 1924. - Agaricus denudatus R a b . i n H e d w i g i a 6: 4 5 . 1867; edge sterile or almost entirely so; c h e i l o c y s t i d i a 13-48 X 6.0-15 p m ,
Lepiota denudata (Rab.) S a c c , S y l l . F u n g . 5: 5 2 . 1 8 8 7 ; Leucocoprinus extremely variable, rather irregular, lageniform w i t h 3.0-6.5 p m w i d e
denudatus (Rab.) S i n g , i n L i l l o a 2 2 : 4 2 4 . ( ' 1 9 4 9 ' ) 1951. - Lepiota neck, clavate-cylindrical, c y l i n d r i c a l , w i t h or without p e d i c e l , colour-
boudieri G u e g u e n i n B u l l , trimest. S o c . m y c o l . Fr. 24: 131. ( ' 1 9 0 8 ' ) less, but i n fresh material f u l l o f y e l l o w granules. P l e u r o c y s t i d i a
1909, n o n Lepiota boudieri B r e s . , 1884; Lepiota gueguenii Sacc. & absent. Pileus covering squamules made up o f globose elements, 15-
Trott. i n S a c c , S y l l . F u n g . 2 1 : 21. 1912. 45 p m i n diam., some ellipsoid, some w i t h broad connecting excres-
S E L . I C O N . - M i g l . et a l . i n R i v . M i c o l . 32: 9-11. 1989; V e l l i n g a & c e n c e ^ ) to c y l i n d r i c a l elements, 3.0-5.0 p m w i d e ; the bigger elements
Huijser i n C o o l i a 40: p i . 7. 1997 (both as L. denudatus). slightly t h i c k - w a l l e d ; pigment pale y e l l o w i n drops and granules situ-
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 4 9 3 , f i g . 109. ated i n the c y l i n d r i c a l elements. Stipitipellis a cutis o f narrow c y l i n -
1990 (as L. denudatus); G u e g u e n i n B u l l , trimest. S o c . m y c o l . Fr. 24: drical elements, 2.0-5.0 p m wide, w i t h scattered squamules made up o f
131. ( ' 1 9 0 8 ' ) 1909 (as Lepiota boudieri); H e r i n k i n C e s k a M y k o l . 13: rounded and e l l i p s o i d elements, as o n pileus surface. C l a m p - c o n n e c -
110-115. 1959 (as L. denudatus); K e l d e r m a n , Parasolzw. Z u i d - tions absent.
L i m b u r g : 180-181. 1994 (as L. denudatus); M i g l . et a l . i n R i v . M i c o l . H A B I T A T & DISTR. - In c l u m p s and groups, rarely solitary, sapro-
32: 10. 1989 (as L. denudatus). trophic and terrestrial, i n pots o f plants i n greenhouses or o n compost
V E R N . N A M E - Zwavelgele plooiparasol. heaps and woodchips outdoors. In the Netherlands rather rare, i n recent
years recorded f r o m several places outside ( E i n d h o v e n , Eerbeek).
P i l e u s , w h e n expanded 15-33 m m , w h e n y o u n g p a r a b o l o i d - c y l i n d r i - Aug.-Sept. i n the w i l d , and the whole year through inside. Widespread
cal, expanding to plano-convex w i t h deflexed m a r g i n to applanate with and recorded from a l l over E u r o p e , probably o f tropical o r i g i n .
s m a l l u m b o i n shallow central depression, w i t h or without uplifted
margin, pale l e m o n - y e l l o w ( K . & W . 1 A 4 at centre, 1 A 3 i n rest o f N a r d u c c i & C a r o t i (in M e m o r i e S o c . tosc. S c i . nat., Ser. B , 102: 4 9 .
pileus), sometimes cream-coloured at margin, w i t h floccose granules ('1995') 1996) showed that Agaricus straminellus B a g l . is identical
to fine-granulose (lens) a l l over, or even m a k i n g a smooth impression, with A. denudatus R a b . T h e former name was the first published, so the
outer half to 3/4 o f radius radially sulcate. L a m e l l a e , L = c. 55,1 = 0- w e l l - k n o w n name L. denudatus has to be replaced b y the virtually
1, moderately crowded, free and remote from stipe, subventricose, up u n k n o w n name L. straminellus.
to 2 m m w i d e , white w i t h slightly sulphur tinge to pale lemon-yellow, F r o m France and Italy a white variety o f this species i s k n o w n , v i z .
w i t h concolorous, eroded to finely fimbriate edge. Stipe 17-50 X 1-3 L. straminellus var. albus (Joss.) M i g l . & R a v a (in M i c o l . Veget. medit.
m m , c y l i n d r i c a l , slightly w i d e n e d i n basal part, h o l l o w , pale l e m o n - 14 (1): 25. 1999); this has not yet been found i n the Netherlands.
y e l l o w , c o n c o l o r o u s w i t h pileus surface, totally pruinose to covered
w i t h loose floccules o f granules. A n n u l u s m i d w a y o n stipe, small, 8. Leucocoprinus heinemannii M i g l . i n M i c o l . ital. 16 (2): 8. 1987.
ascending, pale lemon-yellow. Context thin and concolorous w i t h sur- S E L . I C O N . - M i g l . et a l . i n R i v . M i c o l . 32: 13. 1989; M i g l . &
faces. S m e l l fungoid, unpleasant. Taste not k n o w n . Spore print white. Z e c c h i n i n B e l g . J . B o t . 131: 172. ( ' 1 9 9 8 ' ) 1999.
SEL. D E S C R . & F I G S . - F o u r r e i n D o c . m y c o l . 24 (93): 6 3 . 1994;
M i g l . i n M i c o l . ital. 16 (2): 8-13. 1987; M i g l . et a l . i n R i v . M i c o l . 3 2 :
12. 1989; M o h r i n M y k o l . M i t t b l . 35: 87-89. 1992.
C H A R A C T E R I S T I C S - Pileus 15-20 m m , plano-convex to plano-concave

w i t h age, w i t h flattened centre, at centre dark grey to almost black,
w i t h small fibrils, around centre w i t h irregular dark grey to b l a c k ,
plush-like fibrils o n pale, w h i t i s h to y e l l o w i s h background, sulcate, i n
outer 3-10 m m o f radius, sometimes w i t h remnants o f annulus at mar-
gin; lamellae, L = 35-40,1 = 0-1, moderately to rather crowded, free,
subventricose, whitish to cream, w i t h even and concolorous edge; stipe
15-35 X 1.5-2 m m , gradually w i d e n i n g downwards to 3 m m at base,
hollow, shiny white-cream to very pale p i n k i s h - l i l a c i n o u s w i t h white
fibrils; annulus fugacious, white; context white i n pileus and stipe;
smell fungoid or none.
Spores 6.0-9.0(-9.5) X 3.0-4.5(-5.0) u m , o n average 6.5-7.7 X 3.6-
4.2 p m , Q = 1.55-2.0, Q a v = 1.8-1.85, i n side-view o b l o n g - a m y g d a l -
i f o r m , i n frontal v i e w o b l o n g , without g e r m pore; b a s i d i a 14-21 X
6.5-8.5 um, 4-spored, surrounded b y pseudoparaphyses; l a m e l l a edge
sterile; c h e i l o c y s t i d i a 13-33 X 7.0-11 p m , n a r r o w l y clavate to narrow-
ly u n i f o r m , variable, sometimes slightly capitate; pileus covering made
up o f loosely attached irregular hyphae w i t h short and l o n g elements,
up to 11 u m wide, w i t h grey-brown or greenish b r o w n intracellular p i g -
ment; clamp-connections absent.
H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial, i n
F i g . 56. Leucocoprinus straminellus. pots o f plants, i n greenhouses. In the Netherlands a few times record-
ed ( L e i d e n , botanical garden; 's-Graveland, Boekesteyn), Sept.-Jan. It differs f r o m L. melanotrichus i n m i c r o s c o p i c a l characters (see

Reported f r o m several European countries (Italy, France, G e r m a n y ) , M i g l i o z z i & Z e c c h i n i n B e l g . J . B o t . 131: 174. ( ' 1 9 9 8 ' ) 1999. for a
probably o f tropical o r i g i n . tabular overview). T h e easiest character to tell the two taxa apart is the
spore shape, as expressed by the average length w i d t h ratio: 1.8-1.85
Leucocoprinus heinemannii macroscopically resembles Leucoagaricus i n L . heinemannii versus 1.55-1.65 i n L. melanotrichus. Leucocoprinus
melanotrichus ( M a i . & Bert.) T r i m b a c h var. melanotrichus. Because heinemannii shows a sulcate pileus margin, whereas the pileus is not
o f this resemblance the characteristics o f this greenhouse species are grooved i n L. melanotrichus.
given here.
6. Leucoagaricus (Locq. ex) Sing.
Leucocoprinus subgen. Leucoagaricus Locq. in Bull. mens. Soc. linn. Lyon 12: 92. 1943 (not valid); Leucoagaricus
(Locq. ex) Sing, in Sydowia 2: 35. 1948. - Sericeomyces Heinem. in Bull. Jard. bot. natn. Belg. 48: 401. 1978.
SELECTED LITERATURE - M . Bon in Doc. mycol. 11 (43): 50-65. 1981; M . Bon, F l . mycol. Eur. 3, Lepiotes: 85-110.
1993 (as Sericeomyces and Leucoagaricus resp.); Candusso & Lanzoni, Lepiota: 329-457. 1990; Kelderman,
Parasolzw. Zuid-Limburg: 141-171. 1994.
Basidiocarps pluteoid, varying from fragile and slender to sturdy and fleshy; pileus not or rarely sulcate; lamellae free;
annulus present, though often fugacious, made up of partial, more rarely also universal veil; spore print white, whitish.
Some species discolouring with ammonia vapour, and some changing colour on drying.
Spores smooth or minutely roughened, binucleate, with or without germ pore, dextrinoid though slowly and vague-
ly in some species, and metachromatic in Cresyl Blue, in some species with metachromatic tract, congophilous,
cyanophilous, with walls swelling in ammonia (NH ( q)) and acetic acid (CH^COOH^q); basidia 4-spored, rarely 2-
3 a
spored, not surrounded by pseudoparaphyses; cheilocystidia present; pleurocystidia rarely present; hymenophoral trama
trabecular or regular; pileus covering very variable, from a cutis to a complicated trichoderm; clamp-connections
absent. Development bivelangiocarpic and stipitocarpic. - Type species Leucoagaricus macrorhizus Locq. ex Horak.
HABITAT & DISTRIBUTION - Saprotrophic and terrestrial, on wood chips and sawdust, in woods, grasslands and on
road verges, and in dune vegetation. Widespread, and world-wide, but represented by more species in the subtropical
than in cool-temperate regions, absent from arctic and alpine habitats.
The genus Leucoagaricus was split off from Leucocoprinus Pat. which thereby became more homogeneous.
Leucoagaricus, however, accommodates species which differ widely in macroscopical and microscopical characters.
The position of some species has been disputed. The group of species which turn green or red in ammonia vapours has
been placed both in Leucocoprinus and in Leucoagaricus. The latter approach is followed here. The species, charac-
terized by a white sericeous pileus covering, have been placed in a separate genus, Sericeomyces Heinem., but that is
not recognized here. There are strong indications that most species of Sericeomyces form a monophyletic group togeth-
er with species like L. rubrotinctus (Peck) Sing., L. sublittoralis, and L. purpureolilacinus. Macrolepiota nympharum
(Kalchbr.) Wasser is included in Leucoagaricus, on account of its molecular and morphological characteristics.
The existing infrageneric classifications of the genus are very inconsistent, as closely related species are placed in
different sections (e.g. L. americanus (as L. bresadolae) in sect. Annulati and L. meleagris in sect. Piloselli; L. bars'sii
(as L. pinguipes and L. macrorhizus) in sect. Leucoagaricus and L. leucothites in sect. Annulati) (see e.g. Bon, F l .
mycol. Eur. 3, Lepiotes: 90-110. 1993). Furthermore, widely different species, like L. barssii and L. melanotrichus,
were placed in the same section. As more research on species from all over the world, especially on the molecular
level, is needed to establish a well-founded and sound infrageneric classification, none is given here.
A thorough monographic approach to this group is badly needed. Many species are known from a single or a few
reports in literature, without a critical character evaluation.
The genus is represented by a relatively small number of species in the northern parts of Europe, whereas southern
Europe harbours a much greater diversity. This suggests even greater diversity in the tropics, but the actual species
numbers are unknown.
Note that species not (yet) recorded from the Netherlands are not included in the keys, although some are treated in
notes under the species which most resemble them.
Several species are cultivated by attine ants in the Neotropics (see e.g. Chapela et al. (in Science 266: 1691-1694.
1994), and Mueller et al. (in Science 281: 2034-2039. 1998)). These fungi are mostly known from a vegetative stage,
as basidiocarp formation is inhibited by chemicals exuded by the ants.
85
1. Pileus and stipe turning red on bruising; lamellae green or red with ammonia vapour Key one
1. Pileus and stipe not changing colour when bruised, or some yellow or brown with age; lamellae not discolouring
with ammonia vapour
2. Basidiocarps big, fleshy, predominantly white (some greyish, or brown-squamulose); spores with or without germ
pore, with rounded apex Key two
2. Basidiocarps thin-fleshed, fragile; pileus white, white with brown or black, grey or pinkish fibrils or squamules
or totally coloured; spores without germ pore; spores amygdaliform with rather acute apex, or ellipsoid to oblong,
with rounded apex
3. Basidiocarps when fresh predominantly white, with sometimes some ochre or brown at pileus centre
Key three
3. Basidiocarps with coloured pileus (lilac, pink, purple, brown, grey or black) Key four
K E Y ONE
Species which turn green or red in ammonia vapour, and/or bruise red
1. Basidiocarps turning brick-red in ammonia vapour and on bruising, staying this colour, not changing to black
8. L . croceovelutinus
1. Basidiocarps turning green in ammonia vapour or not showing any reaction at all in ammonia; turning red or brown-
ish when damaged I
2. Basidiocarps when cut first turning yellow, saffron-yellow, then red; spores 7.0-12 X 5.5-8.0 pm, Qav — 1.25-
1.45, (broadly) ellipsoid, relatively thick-walled, with distinct or indistinct germ pore
3. Pileus (70-)100-135(-230) mm; germ pore distinct; elements on pileus gradually tapering towards apex
4. L . americanus
3. Pileus 13-45(-80) mm; germ pore indistinct or absent; elements on pileus with more or less abrupt appendix
5. L . meleagris
2. Basidiocarps directly turning red on bruising and cutting, no yellow discoloration happening; spores (5.5-)6.0-
10.0 X 3.5-5.0(-6.0) pm, Qav = 1.5-1.9, often amygdaliform, without germ pore
4. Basidiocarps whitish with lilacinous tinges, hardly staining or only browning at stipe, when bruised; pileus cov-
ering made up of more or less adnate hyphae 10. L . marriagei
4. Basidiocarps whitish when young, changing rapidly to red or black when touched, or brownish when young
and remaining so; pileus covering with erect cystidioid elements
5. Cheilocystidia clavate, without apical excrescence ( 9. L . pilatianus
5. Cheilocystidia clavate, narrowly clavate to narrowly fusiform, with short to long apical excrescence, though
mixed with clavate elements
6. Basidiocarps medium-sized to robust (pileus 25-65 mm; stipe 35-105 mm long) 6. L . badhamii
6. Basidiocarps slender, small (pileus not exceeding 20 mm; stipe not longer than 35 mm)
7. L . georginae
K E Y TWO
Species with white or grey, big, and fleshy basidiocarps; spores with rounded apex, with or without germ pore
1. Pileus covering made up of adnate cylindrical hyphae; spores without a germ pore
2. Cheilocystidia smooth-walled, without crystals; basidiocarps grey-fibrillose to pale brown-squamulose
1. L . barssii
2. Cheilocystidia with crystals at apex; basidiocarps predominantly white see notes under 21. L. menieri
1. Pileus covering made up of erect elements; spores with a germ pore
3. Ring complex, double; pileus surface coarsely squamose 3. L . nympharum
3. Ring simple, though variable; pileus surface smooth or granular, without or with star-shaped pale pattern
4. Spores 7.5-10(-11.5) X 5.0-7.0(-7.5) um; clamp-connections absent; pileus without star-shaped pattern, vary-
ing in colour from white to grey or grey-brown, often discolouring yellow when touched
2. L . leucothites
4. Spores 11-16 X (6.5-)7.5-10.5 pm; basidia often with clamp-connection; pileus often with star-shaped pattern,
brown at centre, cream around centre, not discolouring when bruised or turning red in stipe with damage
Macrolepiota excoriata
LEUCOCAGARICUS (Vellinga) 87
K E Y THREE
Species with predominantly white, sericeous, radially innate fibrillose pileus

1. Cheilocystidia clavate, rather thin-walled, with long pedicel, without crystals 19. L . serenus
1. Cheilocystidia lageniform, narrowly lageniform, narrowly clavate to cylindrical, with optionally some clavate cys-
tidia present, in most cases with crystals at apex
2. Pileus covering made up of adnate to ascending cylindrical hyphae, with ellipsoid to globose elements at centre
of pileus and in squamules Leucocoprinus cygneus
2. Pileus covering made up of adnate to ascending cylindrical hyphae only
3. Cheilocystidia distinctly lageniform, without crystals or occasionally with some crystals; basidia 4- or 2-
spored; basidiocarps turning brown on drying 18. L . sericifer
3. Cheilocystidia narrowly clavate, cylindrical, narrowly lageniform, with crystals at apex; basidia 4-spored;
basidiocarps retaining cream colours on drying
4. Basidiocarp relatively thin-fleshed; cheilocystidia narrowly clavate, cylindrical to lageniform, often with
coarse crystals at apex 20. L . crystallifer
4. Basidiocarp relatively thick-fleshed, with abrupt bulbous base; cheilocystidia irregularly shaped, lageni-
form, with crystals at apex 21. L . menieri
K E Y FOUR
Species which are thin-fleshed, with coloured sericeous or fibrillose-squamulose covering

1. Pileus rather smooth, with radially innate fibrils
2. Pileus with a white velar patch at centre; stipe with abrupt bulb at base; cheilocystidia narrow, 4.0-11.5 pm wide,
with pale grey contents, sometimes with some small crystals 15. L . wichanskyi
2. Pileus without velar patch at centre; stipe not abruptly bulbous, sometimes rooting in sand; cheilocystidia narrow
to wide, 6.0-15 um pm wide, with or without crystals at apex
3. Cheilocystidia rarely with some small crystals at apex, mostly inside cystidia; pileus pinkish brown at centre
14. L . sublittoralis
3. Cheilocystidia with crystals at apex; pileus purplish brown, dark purplish red-brown, dark vinaceous brown at
centre 16. L . purpureolilacinus
1. Pileus distinctly fibrillose to squamose
4. Pileus with pink, lilac, or purple colours (often fading to brown with age)
5. Spores amygdaliform with acute apex and often with apical papilla
6. Pileus squamules made up of erect elements, 20-130 X 6.0-14 pm, situated on short elements, 5.0-10 pm
wide 17. L . gauguei
6. Pileus squamules made up of elongate, ascending elements, 60-130 X 14-24 pm
see notes under 14. L. sublittoralis
5. Spores ellipsoid to oblong, not distinctly amygdaliform
7. Pileus pink, discolouring brownish with age; pileus covering made up of adnate cylindrical elements, with
terminal elements 40-145 X 10-17 pm 10. L . marriagei
7. Pileus purple-tinged (dark purple, violaceous purple); pileus covering with long, tapering elements, 110-300
X 8.0-12 pm 11. L . ionidicolor
4. Pileus brown, black, or dark grey
8. Pileus with black or dark grey fibrils (with or without green or purplish hue)
9. Spores 5.5-7.0 X 3.5-4.5(-5.0) pm, Qav = 1.55-1.65; pileus not sulcate; growing outdoors
12a. L . melanotrichus var. melanotrichus
9. Spores 6.0-9.0(-9.5) X 3.0-4.5(-5.0) pm, Qav = 1.8-1.85; pileus sulcate in marginal zone; growing in green-
houses Leucocoprinus heinemannii
8. Pileus brown-fibrillose
10. Pileus dull brown; fresh basidiocarps not exuding drops 12b. L . melanotrichus var. fuligineobrunneus
10. Pileus orange-red-brown, red-brown, or dark cinnamon-brown; fresh basidiocarps exuding drops, on stipe
and pileus 13. L . tener
1. Leucoagaricus barssii (Zeller) V e l l i n g a i n M y c o t a x o n 7 6 : 4 3 1 . fibrillose-squamulose, slightly b r o w n i s h grey, sordid pale grey-brown

2000. - F i g . 57. ( M u . 10 Y R 7/2) at centre, grey to grey-brown (10 Y R 7/2, 6-5/3)
Lepiota barssii Z e l l e r i n M y c o l o g i a 26: 211. 1934. - Lepiota pin- around it, later more fibrillose-squamose to squamose, and then sordid
guipesA. Pears, i n Trans. B r . m y c o l . S o c . 35: 97. 1952; Leucoagaricus dark b r o w n at centre to sordid pale b r o w n at margin, coarse white-fib-
pinguipes ( A . Pears.) M . B o n i n D o c . m y c o l . 11 (43): 54. 1981; rillose at margin, when y o u n g w i t h white teeth at margin and under-
Leucoagaricus macrorhizus var. pinguipes ( A . Pears.) A l e s s i o i n neath the fibrillose c o v e r i n g a white fringed m a r g i n (connected to
M i c o l . ital. 17(2): 8. 1988 (not valid). - Leucocoprinus macrorhizus annulus), exceeding lamellae. L a m e l l a e , L = c. 95,1 = ( 0 - ) l , c r o w d -
L o c q . i n B u l l . mens. S o c . l i n n . L y o n 12: 80. 1943 (not valid); ed, free and remote from stipe, ventricose, up to 8 m m wide, white-
Leucoagaricus macrorhizus ( L o c q . ) S i n g , i n S y d o w i a 2: 35. 1948 (not cream, w i t h distinctly white rather even edge. Stipe 60-165 X 8-15 m m
v a l i d ) ; Lepiota macrorhiza ( L o c q . ) K i i h n e r & R o m a g n . , F l . anal. i n the m i d d l e , tapering upwards and w i t h fusiform base, buried w i t h
C h a m p , sup.: 4 0 6 . 1953 (not v a l i d ) ; Leucoagaricus macrorhizus lower part (up to 2/3 o f total length) i n the soil, compressed, hollow,
( L o c q . ex) H o r a k , S y n . G e n . A g a r . : 344. 1968; Leucocoprinus white at superterranean part, d i s c o l o u r i n g b r o w n i s h o n handling,
macrorhizus ( L o c q . ex H o r a k ) D . R e i d i n M y c o l . Res. 93: 4 2 1 . 1989. brownish lower d o w n , hirsute-hairy i n upper part or shiny-fibrillose,
- Leucoagaricus macrorhizus var. pseudocinerascens M . B o n in Doc. and fibrillose i n lower part. A n n u l u s , easily removable from stipe, and
m y c o l . 20 (78): 58. 1990; Leucoagaricus pseudocinerascens ( M . Bon) c l i n g i n g to pileus m a r g i n , w i t h white upper side, and fringed, thick-
M . B o n i n D o c . m y c o l . 23 (89): 6 1 . 1993. ened margin, brownish-fibrillose underneath, i n appearance reminis-
S E L . I C O N . - Courtec. & D u h e m , G u i d e C h a m p . France Europe: fig. cent o f the annulus o f a Macrolepiota species. Context very thick i n
697. 1994 (as L. macrorhizus); Tabares i n Bolets Catalunya 2: p i . 80. pileus, white, slightly cream-beige, i n stipe brownish, tough and slight-
1983 (as L. macrorhizus); W i l h e l m i n S c h w e i z . Z . P i l z k . 72: 259. 1994 l y shiny. S m e l l strongly fungoid-lepiotoid. Taste pleasant-fungoid, like
(as L. macrorhizus var. pinguipes and var. macrorhizus resp.); Z a l i n & cultivated Agaricus bisporus. Spore print 'pale c r e a m ' .
M i g l . i n M i c o l . veneta 7 (3): 2 0 . 1991 (as L. macrorhizus var. Spores 6.5-9.5 X 4.0-5.5 p m , o n average 7.4-8.3 X 4.7-5.1 u m , Q
pseudocinerascens). = 1.4-1.85, Q a v = 1.5-1.65, e l l i p s o i d to oblong i n side-view (remi-
S E L . D E S C R . & F I G S . - H o r a k , S y n . G e n . Agar.: 344-346. 1968 (as L. niscent o f P / w / e i « - s p o r e s ) , without g e r m pore, dextrinoid, c o n - I
macrorhizus); M i g l . & C l e r i c u z i o i n M i c o l . Veget. medit. 4 (1): 29-33 gophilous, cyanophilous, w i t h p i n k inner w a l l i n C r e s y l Blue,
(as L. macrorhizus var. pinguipes); A . Pears, i n Trans. B r . m y c o l . SQC. thick-walled. B a s i d i a 18-42 X 6.5-10 p m , 4-spored, intermixed w i t h 2-
35: 9 7 . 1952 (as L. pinguipes); D . R e i d i n M y c o t a x o n 69: 124-126, spored basidia, sometimes t h i c k - w a l l e d and cystidioid. L a m e l l a edge
figs. 1-6. 1998 (as L. pinguipes). heterogeneous. C h e i l o c y s t i d i a 20-85 X 6.0-25 u m , varied i n shape and
V E R N . N A M E - Wortelende champignonparasol. size, clavate, c y l i n d r i c a l , broadly utriform, often w i t h thickened
colourless w a l l . P l e u r o c y s t i d i a present near l a m e l l a edge, similar to
Pileus 35-80 m m , w h e n y o u n g hemispherical-convex, truncate-coni- cheilocystidia. Pileus covering made up o f repent hyphae, up to 15 u m
cal, expanding to campanulate to plano-convex, w i t h l o w and flattened w i d e , w i t h an occasional short element; terminal elements 10-16 p m
u m b o , a n d s l i g h t l y deflexed m a r g i n , w h e n y o u n g completely hairy- wide, tapering towards acute apex; i n older material difficult to study,
because o f bacteria and gelatinization. Pigment either b r o w n and dif-
fuse, or grey and granular, intracellular; some elements w i t h internal-
l y thickened walls. Stipitipellis a cutis o f colourless c y l i n d r i c a l hyphae,
5-15 p m w i d e , w i t h loosely arranged colourless, c y l i n d r i c a l flexuous
hyphae i n tuft?. Clamp-connections absent.
H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic and ter-
restrial i n open mossy dune vegetation, i n open sand close to shrubs, or
i n road verge, also i n deciduous w o o d o n loamy soil, mostly i n the dune
area (from Texel southwards), once found i n southern L i m b u r g , end o f
Sept.-Oct. Reported from various countries i n Europe (e.g. Great Britain,
Italy, Spain, Switzerland, Hungary), also found i n N o r t h A m e r i c a .
Lepiota barssii, described f r o m the western part o f the U . S . A . , has

been demonstrated to be synonymous w i t h L. pinguipes (Vellinga i n
M y c o t a x o n 76: 431-433. 2000).
R e i d (in M y c o t a x o n 69: 117-128. 1998) gave an elaborate overview
of the literature on L. pinguipes and L. macrorhizus, covering the nomen-
clatorial issues involved, and the differences between the two taxa. H i s
conclusion, that differentiation between the two taxa o n species level is
not possible, is supported by observations on the D u t c h material.
D u t c h specimens w i t h the appearance o f L. macrorhizus are o l d .
These specimens are b i g w i t h a squamose, brownish, pileus covering,
and relatively l o n g spores. Y o u n g and fresh specimens fit the descrip-
tions o f L. pinguipes very w e l l . Therefore, the distinction o f varieties
w i t h i n L. barssii i s superfluous.
A germ pore was not observed, neither under the light microscope,
nor w i t h the S c a n n i n g E l e c t r o n M i c r o s c o p e , though some spores show
a slight thinning at the apex.
Pleurocystidia are often present close to the l a m e l l a edge, and some
F i g . 57. Leucoagaricus barsii. basidia near the edge become very b i g and t h i c k - w a l l e d .
2. Leucoagaricus leucothites (Vitt.) Wasser i n U k r . bot. Z h . 34: 308. mens. S o c . l i n n . L y o n 14: 9 2 . 1945; Leucoagaricus olgae (Velen.)
1977 (as L. leucothitus). - Fig. 58. Mos., Blatter-, B a u c h p i l z e , 1. A u f l . : 115. 1953 (not valid). -
Agaricus leucothites Vitt., Descr. F u n g h i mang. Italia: 310. 1835; Leucoagaricus subcretaceus M . B o n i n D o c . m y c o l . 7 (27-28): 17.
Lepiota leucothites (Vitt.) P.D. O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : 177. 1977 (not valid); Leucoagaricus subcretaceus M . B o n i n Doc. mycol.
1960; Leucoagaricus carneifolius var. leucothites (Vitt.) M . B o n i n 11 (43): 64. 1981 (not valid); Leucoagaricus subcretaceus M . Bon in
D o c . m y c o l . 7 (27-28): 2 1 . 1977 (not valid). - Agaricus holosericeus B o n & H a l u w i j n i n D o c . m y c o l . 13 (49): 4 9 . 1983.
Fr., E p i c r i s i s : 16. 1838; Leucocoprinus holosericeus (Fr.) L o c q . i n MISAPPL. - Leucoagaricus cretaceus sensu M o s . , Blatter-,
B u l l . mens. S o c . l i n n . L y o n 12: 9 5 . 1943; Leucoagaricus holosericeus B a u c h p i l z e , 1. A u f l . : 115. 1953; sensu M o s . , R o h r l i n g e Blatterpilze, 3.
(Fr.) M o s . , Blatter-, B a u c h p i l z e , 1. A u f l . : 115. 1953 (not valid); Aufl.: 185. 1967; sensu K n u d s e n i n H a n s e n & K n u d s e n , N o r d i c
Leucoagaricus holosericeus (Fr.) M o s . , R o h r l i n g e , Blatterpilze, 3. M a c r o m y c . 2: 2 2 3 . 1992.
A u f l . : 1 8 5 . 1 9 6 7 . -Agaricus naucinusFi., E p i c r i s i s : 16. 1838; Lepiota EXCL. - Lepiota naucina sensu M o s . , Rohrlinge, Blatter-
naucina (Fr.) K u m m . , Fiihr. P i l z k . : 1 3 6 . 1 8 7 1 ; Leucoagaricus naucinus Bauchpilze, 2. A u f l . : 135. 1955 ( = Lepiota subalba).
(Fr.) S i n g , i n L i l l o a 22: 4 2 3 . ('1949') 1951. -Lepiota densifolia Gillet,
C h a m p . France: 6 8 . 1874; Leucoagaricus densifolius (Gillet) B a b o s i n K E Y TO THE VARIETIES
S z u j k o - L a c z a , F l . Hortobagy natn. Park: 8 2 . 1982. -Lepiota carneifo- 1. Pileus white to cream, not grey, y e l l o w discolouring or not; elements
lia G i l l e t , H y m e n o m y c e t e s : 6 5 . 1874; Leucocoprinus carneifolius of pileus covering elongate (20-145 X 6.0-15 um), without, some-
(Gillet) Locq. in Bull. mens. Soc. linn. Lyon 14: 93. 1945; times w i t h pale intracellular pigment 2a. var. leucothites
Leucoagaricus carneifolius ( G i l l e t ) M o s . , Blatter-, B a u c h p i l z e , 1. 1. Pileus grey, grey-brown, brownish, especially w h e n y o u n g , and at
A u f l . : 115. 1953 (not valid); Leucoagaricus carneifolius (Gillet) M . centre when full-grown; elements o f pileus covering often ellipsoid,
B o n i n D o c . m y c o l . 7 (27-28): 2 1 . 1977; Leucoagaricus carneifolius l l - 7 0 ( - 1 2 5 ) X 8.0-15 p m , w i t h intracellular pigment
(Gillet) Wasser i n U k r . bot. Z h . 34: 307. 1977. - Lepiota olgae Velen., 2b. var. carneifolia
C e s k e H o u b y : 210. 1920; Leucocoprinus olgae (Velen.) L o c q . i n B u l l .
Fig. 58. Leucoagaricus (b: pc o f var. carneifolius).

2a. var. leucothites In older E u r o p e a n literature, this species is often k n o w n under the
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 240, p i . 243. name L. pudicus (e.g. M o s e r , R o h r l i n g e B l a t t e r p i l z e , 3. A u f l . : 185.
1995 (as L. leucothites a n d L. subcretaceus resp.); Candusso & 1967). However, B u l l i a r d (Herb. France: p i . 579. 1791) described and
L a n z o n i , L e p i o t a : p i . 5 2 - 5 4 . 1 9 9 0 (as L. leucothites a n d L . subcretaceus depicted two totally different species under that name, one w i t h white
resp.); Courtec. & D u h e m , G u i d e C h a m p . France Europe: p i . 704. lamellae, a striped annulus, and a stuffed stipe, not fitting the present
1994; J . Lange, F l . agar. dan. 1: p i . 9 A . 1935 (as Lepiota naucina). concept o f L. leucothites at a l l ; the other w i t h b r o w n lamellae, white
S E L . D E S C R . & F I G S . - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 170- patches o n the pileus, a n d also w i t h a striped annulus, resembling a
171. 1994. species o f the genus Rhozites.
V E R N . N A M E - B l a n k e champignonparasol. Leucoagaricus leucothites i s taken here i n a w i d e sense, i n c l u d -
ing a l l the w h i t e variants, v a r y i n g f r o m s m a l l to b i g . S p e c i m e n s
Pileus 35-200 m m , at first hemispherical w i t h straight deflexed mar- w h i c h turn y e l l o w o n a g i n g o r b r u i s i n g are also i n c l u d e d , as no char-
gin o r wide-truncate-conical w i t h inflexed margin, later expanding to acters were found to discriminate between the variants. G r e y variants
p l a n o - c o n v e x w i t h flattened o r depressed centre, plano-convex, o r c o m e very close, and are r e c o g n i z e d here as a variety o f L. leucoth-
more truncate-conical, f i n a l l y w i t h uplifted m a r g i n , white at first, ites ( v i z . var. carneifolius). T h i s taxon differs f r o m L. leucothites in
cream, i s a b e l l a cream to cream at m a r g i n (overall impression e.g. M u . the p r e d o m i n a n t l y grey o r g r e y - b r o w n p i l e u s , the m o r e p i n k l a m e l -
10 Y R 7/2), w i t h some pale greyish tinges, e.g. finely grey-punctate, lae, a n d the i n general shorter elements i n the pileus c o v e r i n g . A l l
often, but not always, irregularly breaking up into s m a l l to b i g patch- those characters show a certain amount o f variation; there are slight-
es, w i t h r a d i a l fissures, tomentose (lens) o r granulose-fibrillose a n d l y grey variants o f L. leucothites; p i g m e n t i s not totally absent f r o m
more r a d i a l l y f i b r i l l o s e at m a r g i n ; often y e l l o w i n g (paler than 5-2.5 Y w h i t i s h b a s i d i o c a r p s , a n d the l e n g t h o f the elements i n the pileus
8/4) at m a r g i n a n d i n spots w h e n touched; w h e n y o u n g m a r g i n c o v e r i n g varies greatly (see e.g. C a n d u s s o & L a n z o n i , L e p i o t a : f i g .
fringed, e x c e e d i n g l a m e l l a e . L a m e l l a e , L = c. 50-100, 1 = 1-5, 95. 1990).
c r o w d e d , rather c r o w d e d , rarely moderately distant, free and remote T h e y e l l o w i n g reaction o f the basidiocarps o n bruising is not c o n -
f r o m stipe, often anastomosing, segmentiform, subventricose to v e n - sidered o f any t a x o n o m i c a l importance. It s h o u l d be borne i n m i n d ,
tricose, 5-14 m m broad, white, pale creamy, greyish white, often w i t h that i n a l l the o r i g i n a l diagnoses o f the species i n this group this char-
p i n k i s h sheen, sometimes reddening w h e n bruised, w i t h white, b r o w n acter is not mentioned. T h i s is a pervasive p r o b l e m i n this species c o m -
w i t h age i n some specimens, finely flocculose edge (lens). Stipe 4 5 - plex, as m o d e r n authors (e.g. B o n , F l . m y c o l . E u r . 3, L e p i o t e s :
125 X 6-15(-40) m m , c y l i n d r i c a l , i n some specimens broadened into 108-110. 1993) have interpreted the species i n a different manner than
bulb (up to 25 m m w i d e ) , h o l l o w , w h e n y o u n g white a l l over, later the o r i g i n a l authors, and have distinguished species using characters,
o n l y white at apex, i n older specimens b e l o w this white zone greyish- w h i c h are not mentioned b y the older authors. O f course, i n almost a l l
b r o w n i s h , and b e l o w annulus short-striate or lengthwise creamy w i t h cases, type material i s l a c k i n g . Leucoagaricus densifolius exemplifies
some g r e y i s h fibrils i n l o w e r part, at bulb p i n k i s h o r y e l l o w i s h w h e n this. G i l l e t ( C h a m p . France: 68. 1874) described i t as a white fungus:
touched. A n n u l u s white o r cream, sometimes w i t h greyish underside, white pileus, white lamellae, and a white stipe, but i n modern concepts
variable, ascending w i t h tomentose-membranous cuff and thickened it has p i n k lamellae, and discolours q u i c k l y v i a y e l l o w to b r o w n . A n d
margin, descending w i t h f l a r i n g l i m b , o n l y a cuff around stipe w i t h a Candusso & L a n z o n i ( L e p i o t a : p i . 5 1 . 1990) even depicted i t as a
s m a l l flaring part halfway, or without any cuff, but just an evanescent species w i t h a brown-squamulose pileus.
flaring part. Context very thick i n pileus, white and d u l l , w i t h glassy Leucoagaricus cinerascens (Quel.) B o n & Boiffard is a distinct and
line above lamellae, turning y e l l o w i n some specimens, i n stipe shiny different ^pecies, w i t h a p e c u l i a r pileus c o v e r i n g , made up o f w i d e ,
and white, sometimes y e l l o w i n g w h e n cut, and p i n k i s h or b r o w n i s h i n globose elements, o n w h i c h narrow c y l i n d r i c a l hyphae are set (see
bulb, a n d stipe sometimes protruding into pileus. S m e l l strong a n d Candusso & L a n z o n i , L e p i o t a : f i g . 9 3 . 1990 for a s t y l i z e d picture).
fungoid (a b i t aniseed-like, l i k e Agaricus bisporus), a bit astringent, T h i s is probably a rare species, w i t h a mediterranean distribution.
m u s t y to m u s t y fungoid, indistinct. Taste fungoid, indistinct. Spore Leucoagaricus moseri (Wasser) Wasser, described f r o m the
print white to very pale p i n k . U k r a i n e , is probably also separate f r o m L. leucothites o n account o f
Spores 7.5-11 X 5.0-7.0 u m , o n average 8.0-9.2 X 5.3-6.4 u m , Q the absence o f c h e i l o c y s t i d i a (see description b y Wasser i n L i b r i bot.
= 1.25-1.9, Q a v = 1.35-1.6, e l l i p s o i d - a m y g d a l i f o r m , o b l o n g - a m y g - 9: 72-73. 1993).
d a l i f o r m , w i t h rounded o r w i t h slightly acute apex, t h i c k - w a l l e d , w i t h B o t h varieties o f Leucoagaricus leucothites are very widespread
g e r m pore, d e x t r i n o i d , c o n g o p h i l o u s , cyanophilous, w i t h inner w a l l and cosmopolitan. It is not k n o w n whether the distribution is natural
and p l u g i n g e r m pore p i n k i n C r e s y l B l u e . B a s i d i a 17-39 X 8.0-13 or anthropogenic. T h e species grows especially i n man-made habitats
pm, 4-spored, rarely 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a like grasslands, lawns, meadows, and road verges.
22-70 X 7.0-15 u m , c y l i n d r i c a l , narrowly clavate, often irregular, w i t h
excrescence at apex o r acuminate apex, o r subcapitate apex, not-
coloured, and slightly t h i c k - w a l l e d . P l e u r o c y s t i d i a absent. P i l e u s c o v - 2b. var. carneifolius (Gillet) V e l l i n g a i n prep.
ering made u p o f erect to ascending (close to margin) c y l i n d r i c a l M I S A P P L . - Leucoagaricus cinerascens sensu M i g l . & C o c c i a i n B o l l .
hyphae, often arranged into tufts, w i t h terminal elements 20-145 X A s s . m i c o l . ecol. r o m a n a 28: 7-14. 1993; sensu Wasser i n L i b r i bot. 9:
6.0-15 p m , often w i d e r than penultimate elements, without o r w i t h 7 6 . 1 9 9 3 ; sensu K n u d s e n i n H a n s e n & K n u d s e n , N o r d i c M a c r o m y c . 2:
pale greyish o r b r o w n i s h intracellular pigment. S t i p i t i p e l l i s a cutis o f 223. 1992.
c y l i n d r i c a l colourless hyphae, 4.0-7.0 p m i n d i a m . Clamp-connections S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 239. 1995 (as
not observed. L. cinerascens); M i g l . & C o c c i a i n B o l l . A s s . m i c o l . e c o l . R o m a n a 28:
H A B I T A T & DISTR. - Gregarious, solitary, saprotrophic and terrestri- 12. 1993 (as L. cinerascens).
al, m a i n l y i n lawns, grasslands, road verges, dune grasslands etc. A u g . - S E L . D E S C R . & FIGS. - M i g l . & C o c c i a i n B o l l . A s s . micol. ecol.
Nov. W i d e s p r e a d a n d c o m m o n i n the Netherlands. Widespread R o m a n a 28: 7-10. 1993 (as L. cinerascens).
throughout the Northern Hemisphere, also i n the Southern Hemisphere V E R N . N A M E - Grijze champignonparasol.
(e.g. southern Australia).
C H A R A C T E R I S T I C S - Pileus grey, grey-brown, especially when young;

lamellae p i n k w i t h age, w i t h white flocculose edge; stipe discolouring
y e l l o w w h e n touched.
Spores 8.0-11.5 X 5.0-7.0(-7.3) p m , o n average 9.0-9.5 X 5.7-6.5
um, Q = 1.3-1.8, Qav = 1.45-1.6, e l l i p s o i d - a m y g d a l i f o r m w i t h round-
ed apex, t h i c k - w a l l e d , w i t h g e r m pore, dextrinoid, congophilous, w i t h
p i n k inner w a l l and tract i n C r e s y l B l u e , cyanophilous; pileus covering
trichodermal, made up o f relatively short elements, 11-70(-125) X 8.0-
15 p m , w i t h greyish or b r o w n i s h intracellular pigment.
HABITAT & DISTR. - A s the t y p i c a l variety, but rare i n the
Netherlands. Sept.-Nov. Widespread i n the Northern Hemisphere, but
rarer than the t y p i c a l , white variety.
Leucoagaricus cinerascens is an often used name for this grey to brown

variety, but the concept o f B o n ( F l . m y c o l . Eur. 3, Lepiotes: 110. 1993)
and Candusso & L a n z o n i (Lepiota: 422-424. 1990) is f o l l o w e d here.
This taxon is characterized b y the interesting and unique structure o f its
pileus covering, i n w h i c h globose to w i d e - c y l i n d r i c a l elements give rise
to slender c y l i n d r i c a l elements. Furthermore, the cheilocystidia have a
distinct apical excrescence. T h i s is probably a species w i t h a mediter-
ranean distribution; it has not been found yet i n the Netherlands.
Leucoagaricus cinereolilacinus (Barbier) B o n & Boiffard may be
a s y n o n y m o f L. leucothites var. carneifolius, but i t is not quite clear
what its characters are, as different authors reported different spore
sizes; M i g l i o z z i & C o c c i a (in B o l l . A s s . m i c o l . e c o l . romana 28: 12.
1993) gave a g o o d overview.
3. Leucoagaricus nympharum (Kalchbr.) M . B o n i n D o c . m y c o l . 7

(27-28): 19. 1977. - F i g . 59.
Agaricus nympharum K a l c h b r . , Ic. sel. H y m e n o m y c . Hungariae:
10, p i . 2, fig. 2 . 1 8 7 3 ; Lepiota nympharum (Kalchbr.) K a l c h b r . i n M a g y
tudom. A k a d . Ertek. termeszettud. K o r e b . 1878: 7. 1878; Macrolepiota
nympharum (Kalchbr.) Wasser, A g a r . G r i b y S . S . S . R . : 114. 1985. -
Lepiota densesquamosaVclen., C e s k e H o u b y : 206. 1920. - Agaricus
rachodes var. puellaris Fr., M o n o g r . H y m e n o m y c . Suec. 2: 285. 1863;
Lepiota rachodes var. puellaris (Fr.) S a c c , S y l l . F u n g . 5: 29. 1887;
Lepiota procera var. puellaris (Fr.) M a s s . , B r i t . F u n g . F l . 3: 235. 1893;
Lepiota puellaris (Fr.) R e a , B r i t . Basidiomyc: 6 5 . 1922;
Lepiotophyllum rachodes var. puellaris (Fr.) L o c q . i n B u l l . mens. Soc.
l i n n . L y o n 11: 40. 1942; Leucocoprinus puellaris (Fr.) L o c q . i n B u l l ,
mens. S o c . l i n n . L y o n 14: 91. 1945; Macrolepiota puellaris (Fr.) M o s . ,
Blatter-, B a u c h p i l z e , 1. A u f l . : 114. 1953 (not v a l i d ) ; Macrolepiota
puellaris (Fr.) M o s . , R o h r l i n g e Blatterpilze, 3. A u f l . : 184. 1967.
M I S A P P L . - Lepiota cepaestipes sensu M i c h a e l , Fiihr. Pilzfr., A u s g .
B , 2: p i . 194. 1918.
E X C L . - Leucoagaricus nympharum sensu M . B o n i n D o c . m y c . 7 F i g . 59. Leucoagaricus nympharum.
(27-28): 19. 1977 ( = u n k n o w n species); Lepiota puellaris sensu R e a ,
B r i t . B a s i d i o m y c : 65. 1922 ( = u n k n o w n species); Macrolepiota puel-
laris sensu M . B o n i n D o c . m y c . 7 (27-28): 19. 1977 ( = Macrolepiota A m b . 66: 10-11. 1994 (as Macrolepiota puellaris); Pegl. & Legon in
rachodes). M y c o l o g i s t 6: 187. 1992 (as Macrolepiota puellaris).
S E L . I C O N . - B e l l u i n B o l l . G r u p p o m i c o l . G . B r e s . 25: 112. 1982 V E R N . N A M E - Witschubbige parasolzwam.
(as Macrolepiota puellaris); Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i .
255. 1995 (as Macrolepiota puellaris); Candusso & L a n z o n i , L e p i o t a : Pileus 38-90 m m , conico-convex, convex or slightly truncate broadly
p i . 66. 1990 (as Macrolepiota puellaris); Cetto, F u n g h i Vero, E d . 5, 1: c o n i c a l to plano-convex, without u m b o o r p a p i l l a , without depressed
p i . 23. 1975 (as L. puellaris); J . L a n g e , F l . agar. dan. 1: p i . 9 B . 1935 centre, w i t h margin inflexed w h e n y o u n g , later w i t h deflexed margin
(as L. rhacodes var. puellaris); K a l c h b r . , Ic. s e l . H y m e n o m y c . exceeding lamellae, white to very pale buff to very pale b r o w n ( M u .
Hungariae: p i . 2, fig. 2. 1873; K o r h o n e n i n S i e n i L e h t i 41 (3): cover. 10 Y R 8/3), dry and r a d i a l l y fibrillose-squamose; squames concentri-
1989 (as Macrolepiota puellaris); M i g l . & B i z i o i n F u n g h i A m b . 66: cally arranged w i t h uplifted tips; w i t h 13-31 m m wide, pale to m o d -
14. 1994 (as M. puellaris); P e g l . & L e g o n i n M y c o l o g i s t 6: 187. 1992 erately dark g r e y - b r o w n calotte (7.5 Y R 6/6 - 10 Y R 5/4.-7/3),
(as Macrolepiota puellaris). surrounded b y s m a l l patches o f v e l u m l y i n g o n top o f the often
S E L . D E S C R . & FIGS. - J . Favre, Cat. descr. C h a m p , sup. Z o n e sub- recurved squames. L a m e l l a e , L = c. 70-80,1 = 1, moderately c r o w d -
alp.: 556-557. 1960 (as Lepiota puellaris); M i g l . & Bizio i n Funghi ed, free and 1-1.5 m m remote f r o m stipe, ventricose, 5-10 m m wide,
white, creamy white w i t h pale p i n k sheen, w i t h entire to subfloccose F o r more information o n distribution and ecology see N a u t a & V e l l i n g a
w h i t i s h edge. Stipe 50-115 X 5-9 m m , tapering towards apex, w i t h ( A t l . N e d e r l . Paddest.: 204. 1995). Sept.-Oct. Rare and widespread i n
subbulbous to bulbous base, 16-42 m m i n diameter, fistulose w i t h Europe.
some fibrils inside, w h i t i s h , but soon sordid p i n k i s h b r o w n to very
pale b r o w n to b r o w n i s h y e l l o w (10 Y R 7/4 - 6/8), glabrous and dry, T h e above description is based o n material f r o m G e r m a n y and
w i t h white tomentum at bulb and base. A n n u l u s at 2/3 from base, first Switzerland, as no material o f the D u t c h c o l l e c t i o n was conserved.
i m m o b i l e , b e c o m i n g m o b i l e , c. 13 m m i n diameter, w h i t i s h to very Leucoagaricus nympharum strongly resembles members of
pale p i n k i s h - g r e y i s h - b r o w n i s h , slightly darker at edge, w i t h smooth Macrolepiota sect. Laevistipedes. M i g l i o z z i & B i z i o (in F u n g h i A m b .
upper and l o w e r side, and w i t h double, rarely single, c r o w n , w i t h side 66: 9-16. 1994) suggested that this species is an intermediate between
as pileus surface. Context compact, 7.5-9 m m thick i n pileus centre, the genus Leucoagaricus and Macrolepiota, because o f the relatively
w h i t e i n pileus and stipe, pale b r o w n after being cut (10 Y R 8/4). small spores, and the lageniform or appendiculate cheilocystidia. Other
S m e l l faint and indistinct. Taste indistinct to slightly fungoid. Spore m o r p h o l o g i c a l characters, like the absence o f clamp-connections, and
print pale cream ( R o m a g n e s i 2 A ) . the structure o f the pileus c o v e r i n g , as w e l l as m o l e c u l a r evidence,
Spores 8.0-12.5 X (5.5-)6.0-8.0 p m , i n average 8.5-9.7 X 6.3-6.4 point to a close relationship to Leucoagaricus leucothites, rather than
p m , Q = 1.3-1.65, Q a v = 1.35-1.55, broadly ellipsoid to oblong, thick- to the M. rachodes-group.
w a l l e d (up to 0.8 p m thick), w i t h central germ pore and rounded apex, In most E u r o p e a n literature this species is k n o w n as Macrolepiota
with hyaline cap over germ pore, dextrinoid, congophilous, puellaris, b u t D e K o k & V e l l i n g a (in Persoonia 17: 77-78. 1998) point-
cyanophilous, metachromatic i n C r e s y l B l u e . B a s i d i a 21-40 X 8.5-12 ed out that the epithet ' n y m p h a r u m ' has priority o n species level.
pm, 4-spored, without clamp-connection. Lamella edge sterile. Macrolepiota citrinascens Vasas (in A n n l s hist.-nat. M u s . natn.
C h e i l o c y s t i d i a 34-65 X 10-19 p m , subglobose to o b o v o i d or lageni- hung. 81: 46. 1990), described from P/cea-plantations i n Hungary, dif-
form, pedicellate, often rostrate, w i t h 3.0-22 X 3.0-6.0 c y l i n d r i c a l to fers from L. nympharum i n staining yellow, w h e n damaged, but is iden-
slightly m o n i l i f o r m excrescence, t h i n - w a l l e d , colourless, some w i t h tical i n a l l other respects.
pale b r o w n contents. P i l e u s c o v e r i n g a cutis; v e l u m w i t h ascending Leucoagaricus nympharum m a y cause gastrointestinal problems
hyphae, w i t h terminal elements 25-45 X 8.0-10 um, (irregularly) c y l i n - ( K i i h n l i n Siidwestd. Pilzrundschau 29: 21. 1993).
drical; pigment b r o w n , predominantly intracellular, situated i n ultimate
and optionally penultimate elements. Stipitipellis a cutis o f 5.0-10 u m 4. Leucoagaricus americanus (Peck) V e l l i n g a i n M y c o t a x o n 76: 433.
w i d e c y l i n d r i c a l colourless hyphae. C l a m p connections not observed; 2000. - F i g . 60.
according to M i g l i o z z i & B i z i o (in F u n g h i A m b . 66: 11. 1994) very Agaricus americanus P e c k i n R e p . N . Y . St. M u s . nat. H i s t . 23: 7 1 .
rare, m o s t l y i n the h y m e n i u m . ( ' 1 8 7 0 ' ) 1872; Lepiota americana (Peck) S a c c , S y l l . F u n g . 5: 43.
H A B I T A T & D I S T R . - Gregarious, saprotrophic and terrestrial i n 1887; Leucocoprinus americanus (Peck) R e d h e a d i n Groves, E d i b l e
P/cea-forests, also reported for other types of coniferous forests, rarely pois. M u s h r . C a n a d a , E d . 2: 323. 1979. -Agaricus cupreus S. S c h u l z .
i n deciduous woods. R e c o r d e d once f r o m a deciduous w o o d o n nutri- i n V e r h . zool.-bot. G e s . W i e n 27: 112. 1877, n o n Agaricus cupreus
ent-poor, dry soil i n southeastern L i m b u r g , but no material conserved. (Krombh.) Krombh., 1846; Lepiota bresadolae S. Schulz. in
F i g . 60. americanus
H e d w i g i a 2 4 : 132. 1885, n o n Lepiota bresadolae P. H e n n . , 1890; Spores (7.0-)8.0-11.5 X (5.5-)6.0-8.0 p m , o n average (7.7-)9.1-9.7
Leucoagaricus bresadolae ( S . S c h u l z . ) M . B o n i n D o c . m y c o l . 7 (27- X (6.0-)6.4-7.0 u m , Q = 1.2-1.6, Q a v = (1.25-)1.35-1.45, broadly
28): 15. 1977; Leucocoprinus bresadolae ( S . S c h u l z . ) Wasser i n N o v . ellipsoid, ellipsoid, t h i c k - w a l l e d , w i t h germ pore, dextrinoid, but not
Sist. vyssh. n i z s h . Rast. 1977: 221. ( ' 1 9 7 7 ' ) 1978; Leucocoprinus bre- strongly so, congophilous, cyanophilous, w i t h p i n k inner w a l l and pink
sadolae ( S . S c h u l z . ) M o s . , R o h r l i n g e B l a t t e r p i l z e , 4. A u f l . : 248. 1978 p l u g i n germ pore i n C r e s y l B l u e . B a s i d i a 28-39 X 9.5-11 p m , pre-
(superfluous). dominantly 4-spored, but often i n t e r m i x e d w i t h 2-spored basidia.
E X C L . - Lepiota americana sensu Johnson & V i l g a l y s i n M y c o l o g i a L a m e l l a edge sterile. C h e i l o c y s t i d i a abundant, 30-74 X 9.0-18 p m ,
90: 971-979. 1998 ( = Leucoagaricus s p e c ) . - Leucocoprinus bre- narrowly clavate, or gradually w i d e n i n g upwards, without o r w i t h
sadolae sensu Imaz. et a l . , F u n g i Japan: 184. 1988 ( = L. meleagris). abrupt, slightly m o n i l i f o r m appendix, 2.0-32 X 3.0-6.0 u m , sometimes
M I S A P P L . - Lepiota badhamii sensu H u i j s m a n i n M e d e d . N e d . with wider capitulum, w i t h green-brown contents and sometimes dark
m y c o l . Vereen. 28: 6-8. 1943; sensu M e l z e r i n C e s k a M y k o l . 1 3 : 1 1 7 - red granules i n ammonia. P l e u r o c y s t i d i a not observed. Pileus set w i t h
119; Leucoagaricus badhamii sensu R o c a b r u n a i n Bolets Catalunya 8: squamules made up o f erect elements, 100-160 X 8.0-12 p m , sinuous,
p i . 378. 1989; Leucocoprinus badhamii sensu Cetto, G r . Pilzf., 6. A u f l . , w i t h abrupt or gradual apical appendix, flexuous, sometimes capitu-
1: p i . 2 2 . 1980; sensu S c h w o b e l i n Z . P i l z k . 3 2 : 1-2. 1966; sensu late, or slightly m o n i l i f o r m , t h i c k - w a l l e d , green-brown i n ammonia,
W i c h a n s k y i n M y k o l . Sb., Praha 4 1 : 12, p i . 17. 1964. also w i t h b r o w n walls i n lower part o f elements; pigment i n connect-
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 238. 1995 (as ing hyphae sometimes encrusting. Stipitipellis a cutis o f c y l i n d r i c a l
L. bresadolae); B r u n e l l i i n S c h w e i z . Z . P i l z k . 63: opp. p. 174. 1985 (as hyphae, 2.0-4.0 p m i n diam., set (over total length, above and b e l o w
Leucocoprinus bresadolae); Gerhardt, G r . Pilzf.: 36. 1997 (as L. bre- annulus) with cystidioid elements: 65-180 (including appendix) X 9.0-
sadolae); Imler i n Icones mycol. 1985: p i . 102. 1985 (as 13 um, tapering into o r more abrupt ending i n flexuous appendix; p i g -
Leucocoprinus bresadolae); Massart, C h a m p i g n o n s : 3 1 . 1996 (as L. ment, also i n connecting hyphae, green-brown i n ammonia.
bresadolae); R o c a b r u n a i n B o l e t s C a t a l u n y a 8: p i . 378. 1989 (as L. Clamp-connections not observed.
badhamii). H A B I T A T & DISTR. - Clustered and i n groups, saprotrophic o n saw-
S E L . D E S C R . & FIGS. - D . R e i d i n M y c o l . Res. 94: 649-650, figs. 31- dust, piles o f w o o d chips etc. I n the Netherlands k n o w n f r o m a find
36. 1990 (as Leucocoprinus bresadolae); S c h w o b e l i n Z . P i l z k . 32: 1- near Putten i n A u g . 1895, and a second locality near A m s t e r d a m where
2, fig. 1. 1966 (as Lepiota badhami); Weholt i n A g a r i c a 13: 39, fig. 2. the species was found i n 1999, probably o n o l d w o o d , i n a meadow.
1986 (as Leucocoprinus bresadolae). Aug.-Sept. R e c o r d e d from a l l over Europe, except the boreal, montane
V E R N . N A M E - Gebundelde champignonparasol. and alpine zones, but not c o m m o n , possibly o f tropical o r i g i n .
Pileus (70-)100-135(-230) m m , at first cylindrical, subcylindrical with T h e macroscopic description is partly based o n notes accompanying
slightly convex sides w i t h truncate apex, then broadly conical w i t h extra-limital material and o n descriptions from literature (as cited).
rounded to almost flat centre, flattening to plano-convex to applanate, There seem to be two variants o f this species, one w i t h small patch-
w i t h very slight central depression or w i t h l o w umbo, first with uniform es on pileus surface, as depicted by B r u n e l l i (in S c h w e i z . Z . P i l z k . 63:
covering layer, later w i t h covering layer minutely granulose-squamu- opp. p. 174. 1985), the other variant w i t h a pileus surface more r e m i -
lose at centre, around centre breaking into small or bigger, concentric niscent o f a Macrolepiota species (as depicted b y Gerhardt, G r . P i l z f . :
to scattered patches, sometimes w i t h small fibrillose pointed scales at 36. 1997).
margin, and this covering at centre b r o w n i s h to dark purplish b r o w n T h e range o f spore sizes is quite large, due to the presence o f a vari-
( M u . 5 Y R 4/3, 10 Y R 7/5), or paler b r o w n , and around centre purple- able number o f 2-spored basidia.
brown, vinaceous b r o w n to m u c h paler ochraceous, (as pale as 10 Y R The elements o n the pileus are shorter than reported b y R e i d ( i n
7/6), turning pale vinaceous p i n k where bruised and with age, excori- M y c o l . Res. 94: 650. 1990), w h o specified elements up to 2 6 0 p m
ate at margin and thus leaving there whitish, radially fibrillose, sulcate long.
margin bare, i n some specimens w i t h pieces o f annulus adhering to Leucoagaricus americanus from N o r t h A m e r i c a and L. bresadolae
margin. L a m e l l a e , L = c. 100,1 = 0-7, crowded, fairly crowded, free from Europe proved to be identical ( V e l l i n g a i n M y c o t a x o n 76:433-
and remote from stipe, moderately broad, up to 18 m m , creamy whitish, 436. 2000), though S m i t h and Weber (in Contr. U n i v . M i c h . Herb. 16:
later pale sordid y e l l o w i s h (paler than 2.5 Y 8/4), w i t h y e l l o w reflection 219. 1987) stated that they were different. In fact, there are no rnor-
when looked into from below, with a minutely flocculose concolorous p h o l o g i c a l characters to distinguish the two, and m o l e c u l a r research
to w h i t i s h edge, but locally dark vinaceous brown-red (probably after has confirmed the occurrence o f identical species o n both sides o f the
damage). Stipe 95-180 X 12-20 m m , w i d e n i n g downwards, but with Atlantic Ocean.
tapering foot, hollow, white above to p i n k i s h brown, or purplish b r o w n In older literature the names o f L. badhamii and L. bresadolae are
(5 Y R 8/3 to 5 Y R 4/5) (from handling) below, somewhat flocculose at often confused (see the misapplied names). F o r a full discussion o f this
apex, almost glabrous downwards to coarsely lengthwise fibrillose, confusion see D e m o u l i n ( i n L e j e u n i a , n . S. 3 9 : 1-15. 1966) or
whitish to p i n k i s h b r o w n , white-subtomentose at base. A n n u l u s ascend- Krieglsteiner (in Beitr. K e n n t n . P i l z e Mitteleur. 7: 39-60. 1991).
ing, w i t h cuff-like part, and flaring 3-11 m m w i d e part, w h i t i s h then Leucoagaricus meleagris differs from L. americanus i n the small-
dark vinaceous or p u r p l i s h b r o w n o n both sides. Context white or er size o f the basidiocarps, the different aspect o f the pileus surface,
whitish, after cutting immediately o r rather rapidly turning i n spots, or and the shape o f the pilear and stipital elements. Furthermore, the germ
all over, bright yellow, o r sulphur y e l l o w (c. 5 Y 8/6), then after one pore is indistinct.
minute, or more slowly changing to bright brick-red to brownish orange D r i e d specimens o f L. americanus turn vinaceous pink, a feature
(c. 2.5 Y R 6/8), and then to vinaceous r e d to sordid purplish b r o w n . considered to be specific for L. biornatus sensu C o o k e b y Babos (in L .
S m e l l weak, pleasant, more or less chocolate-like. Taste indistinct, or americanus the exsiccates should turn ' w i n e reddish b r o w n ' ) (Babos
slightly astringent. Spore print colour not k n o w n . in B e i h . S y d o w i a 8: 34. 1979).
A m m o n i a solution o n context o f pileus and o n lamellae blue-green, R e i d (in M y c o l . R e s . 90: 1994) described two other c l o s e l y relat-
changing to very dark blue. E x s i c c a t e vinaceous p i n k , fading after ed species, v i z . Leucocoprinus caldariorum D . R e i d (not valid) and L .
some years o f storage. holospilotus ( B . & Br.) D . R e i d ( i n c l u d i n g Lepiota biornata ( B . & Br.)
S a c c ) . B o t h species are said to have smaller basidiocarps (pileus u p

to 9 c m ) ; L. caldariorum does not stain yellow, but immediately red,
and has relatively short, squat hairs o n the pileus. T h e context o f
Leucocoprinus holospilotus does not stain red at a l l , though the stem
discolours red-brown o n h a n d l i n g , and the spores are 7.0-8.5 X 5.0-
6.0 p m .
A l l these species seem very closely related, and more research is
definitely needed to clarify the relationships and similarities between
those taxa. T h e fact that they occur i n temperate regions only i n green-
houses and o n compost heaps and piles o f discarded w o o d and sawdust
near saw m i l l s etc., suggests a (sub)tropical origin.
Leucoagaricus americanus is the type species o f sect. Annulati and
its subsect. Rubescentes, but the closely related L. meleagris, has been
placed i n sect. Piloselli w i t h other red-staining species, like L. bad-
hamii ( B o n , F l . m y c o l . Eur. 3, Lepiotes: 104. 1993). Section Annulati
harbours the non-staining L. leucothites as w e l l .
S o m e E u r o p e a n authors (e.g. Breitenbach & K r a n z l i n , Pilze
S c h w e i z 4: 209. 1995; Massart, C h a m p i g n o n s : 3 1 . 1996; as L. bre-
sadolae) consider L. americanus a toxic species. A m e r i c a n authors, o n
the other hand, regard it as a g o o d edible (e.g. A r o r a , M u s h r . demyst.:
301 1986).
5. Leucoagaricus meleagris (Sow.) Sing, i n L i l l o a 22: 422. ('1949')

1 9 5 1 . - F i g . 61
Agaricus meleagris Sow., C o l . F i g . E n g l . F u n g i 2: p i . 171. 179,9;
Gymnopus meleagris (Sow.) S.F. Gray, Nat. Arr. B r . PI. 1: 609. 1821;
Lepiota meleagris (Sow.) Q u e l , i n M e m . S o c . E m u l . M o n t b e l i a r d , Ser.
II, 5: 3 3 8 . 1873 ( C h a m p . Jura Vosges 2); Hiatula meleagris (Sow.)
S i n g , i n A n n l s m y c o l . 34: 4 3 1 . 1936; Leucocoprinus meleagris (Sow.)
L o c q . i n B u l l . mens. S o c . l i n n . L y o n 14: 9 3 . 1943; Leucocoprinus
meleagris (Sow.) Zschieschang i n M y k o l . M i t t b l . 31: 5. 1988 (super-
fluous); Leucocoprinus meleagris (Sow.) Raithelhuber i n M e t r o d i a n a
16: 16. 1988 (superfluous); Leucoagaricus meleagris (Sow.) M . B o n i n
D o c . m y c o l . 7 (27-28): 19. 1977 (superfluous); Leucoagaricus melea-
gris (Sow.) M . B o n i n D o c . m y c o l . 9 (35): 44. 1979 (superfluous).
M I S A P P L . - Leucocoprinus bresadolae sensu Imaz. et a l . , F u n g i
F i g . 6 1 . Leucoagaricus meleagris.
Japan: 184. 1988.
S E L . I C O N . - V e l l i n g a & Huijser i n C o o l i a 40: p i . 6. 1997; Imaz. et dark purple to dark wine-red i n o l d specimens, sometimes pale pinkish,
al., F u n g i Japan: 184. 1988 (as Leucocoprinus bresadolae). discolouring brown when touched, with white m y c e l i a l cords at base.
S E L . D E S C R . & FIGS. - B a b o s i n A n n l s hist.-nat. M u s . natn. hung. Annulus present, mostly incomplete, sometimes only at pileus margin,
72: 84-87, figs. 3 & 4. 1980; B e n d e r i n M i t t b l . A r b e i t s g e m . P i l z k . ascending, white o n upper side, with dark b r o w n underside. Context i n
N i e d e r r h e i n 13: 137-139. 1995; A . v.d. B e r g & V e l l i n g a i n C o o l i a 4 1 : pileus and stipe whitish, discolouring at first bright y e l l o w (fading after
35-38, fig. 1. 1998; M i g l . & C o c c i a i n M i c o l . ital. 21 (2): 47-49, fig. 5 minutes), then reddish to vinaceous red, but disappearing, slightly
24. 1992; Z s c h i e s c h a n g i n M y k o l . Mittbl. 3 1 : 5-8. 1988 (as shiny i n stipe. S m e l l unpleasant, sweetish-fungoid. Taste unpleasant, bit-
Leucocoprinus meleagris). terish, or fungoid. Spore print probably cream.
VERN. NAME - Compostchampignonparasol. A m m o n i a vapour and solution o n pileus, lamellae and stipe, and
context green ( K . & W . 2 8 E F 4 - 5 ) . E x s i c c a t e vinaceous pink, fading
Pileus 13-45(-80) m m , when young ellipsoid with truncate apex, after a year o f storage.
expanding v i a campanulate to plano-convex, when very young with Spores 7.5-11.5 X 5.5-8.0 p m , o n average 8.6-9.7 X 6.3-7.5 p m , Q
closed covering, soon with plush-like to fibrillose squamules, i n undam- = 1.2-1.6, Q a v = 1.35-1.4, broadly e l l i p s o i d , e l l i p s o i d , w i t h i n c o n -
aged state pure white w i t h dark red-brown fibrillose squamules, espe- spicuous germ pore, dextrinoid, congophilous, cyanophilous, w i t h p i n k
c i a l l y at centre, more dispersed towards margin, o n fibrillose-silky inner w a l l and p l u g i n g e r m pore i n C r e s y l B l u e . B a s i d i a 20-39 X 9.0-
background, b e c o m i n g red, later wine-red ( M u . 7.5-10 Y R 3/4); o l d 12 p m , predominantly 4-spored, but 2-spored basidia may be present
specimens completely dark purple, w i t h darker squamules; marginal i n some places o n a lamella. L a m e l l a edge sterile, set w i t h c h e i l o c y s -
zone up to 8 m m wide, radially sulcate, i n some specimens y e l l o w at tidia, 20-65 X 8.0-17 p m , narrowly clavate, clavate, narrowly fusiform,
margin. Lamellae, L = 50-60,1 = 0-3, crowded, free, segmentiform or without, but m o s t l y w i t h a p i c a l excrescence(s) 2-25 p m long, some-
subventricose, up to 4 m m wide, white-cream at first, later lemon-yellow times branched and/or m o n i l i f o r m , w i t h ( i n ammonia) green-brown
(c. 5 Y 8/3), reddish o r reddish-vinaceous when damaged, with finely intracellular pigment and some red granulose structures. Pleurocystidia
flocculose edge, w h i c h turns red easily o n damaging. Stipe 20-65 X 3- absent. Pileus squamules made up o f cystidioid, slightly t h i c k - w a l l e d
7(-12) m m , narrower at apex and at base, slightly eccentrically fusiform elements, 35-170 X 10-20 p m , clavate short elements intermixed w i t h
with widest part 1/4 o f length above base, sometimes cylindrical, white- narrowly clavate to more c y l i n d r i c a l (depending o n age o f basidiocarp)
cream, completely set with fine, cobwebby fibrils, slightly squamulose, c y s t i d i o i d elements, slightly truncate at apex, w i t h or rarely without
apical excrescences, w i t h same kinds o f pigment as c h e i l o c y s t i d i a ;

connecting hyphae w i t h encrusting pigments. Stipitipellis a cutis o f
c y l i n d r i c a l hyphae, 3.0-5.0 u m i n diam., over total length o f stipe set
w i t h caulocystidia, 33-150 X 8.0-17 p m , narrowly clavate to narrow-
ly fusiform, w h e n y o u n g without, later w i t h narrow l o n g , sometimes
branched, appendices (included i n length), w i t h the same k i n d o f p i g -
ments as cheilocystidia. Clamp-connections not observed.
H A B I T A T & DISTR. - Fasciculate and often gregarious, saprotrophic
and terrestrial o n compost and leaf heaps, o n paths w i t h w o o d chips
etc., J u l y - O c t . , i n the Netherlands not c o m m o n , scattered. K n o w n i n
Europe f r o m hothouses, and similar substrates as i n the Netherlands,
very rarely recorded; also k n o w n from Japan, eastern U . S . A . and west-
ern Canada.
Leucoagaricus meleagris differs from L. americanus i n size o f the
basidiocarps and the structure o f the squamules o n the pileus; m i c r o -
scopically the c y s t i d i o i d elements are shorter and often more truncate
at the apex than i n L. americanus, and the germ pore i n the spores is
not as distinctly visible as i n L. americanus.
Krieglsteiner (in Beitr. K e n n t n . P i l z e Mitteleur. 7: 4 8 . 1991) sug-
gested that L. meleagris is merely a stunted and nutrient-deprived vari-
ant o f L. americanus ( w h i c h he c a l l e d L. bresadolae), but the
differences, both morphological and molecular, warrant the recognition
of t w o species. Differences i n spore size, as proposed b y several
authors, do not exist between the two species. B o t h show a wide varia-
tion i n spore size, partly due to a variable proportion o f 2-spored
basidia. Weather conditions prior to collecting, and refrigerated storage
before examining the specimens, also strongly influence spore size (and
shape).
Lepiota sanguiflua M u r r i l l (see modern description b y S m i t h &
Weber i n Contr. U n i v . M i c h . Herb. 16: 213-214. 1987) appears to be F i g . 62. Leucoagaricus badhamii.
very close o r identical w i t h L. meleagris, o n account o f its general
shape, discoloration (first yellow, then red), and shape and size o f cys- Clausen i n Svampe 26: 18. 1992 (as Leucocoprinus badhamii); Joss, i n
tidia and spores, w h i c h are p r o v i d e d w i t h an inconspicuous germ pore. B u l l . mens. Soc. l i n n . L y o n 4 3 : 210-213, fig. 4. 1974; K e l d e r m a n ,
Parasolzw. Z u i d - L i m b u r g : 162-163. 1994; D . R e i d i n M y c o l . Res. 94:
6. Leucoagaricus badhamii ( B . & B r . ) S i n g , i n L i l l o a 2 2 : 4 1 9 . 643-647. 1990.
('1949') 1 9 5 1 . - F i g . 62 V E R N . N A M E - Bloedende champignonparasol.
Agaricus badhamii B . & B r . i n A n n . M a g . nat. Hist., Ser. II, 13:
397. 1854 (Notic. B r . F u n g i 664); Lepiota badhamii ( B . & Br.) Q u e l , Pileus 25-65 m m , at first conico-convex o r hemispherical to convex
in M e m . S o c . E m u l . M o n t b e l i a r d , Ser. II, 5: 231. 1872 ( C h a m p . Jura with dentate-fringed margin, expanding to plano-convex, applanate,
Vosges 1); Lepiotophyllum badhamii ( B . & Br.) L o c q . i n B u l l . mens. w i t h broad, flattened or l o w u m b o , w i t h straight, non-sulcate margin,
Soc. l i n n . L y o n 11: 4 0 . 1942; Leucocoprinus badhamii (B. & Br.) either totally set with small plush-like squamules or with distinct calotte
L o c q . i n B u l l . mens. S o c . l i n n . L y o n 12: 15. 1943; Leucocoprinus bad- and around centre w i t h radial fibrillose squames, white when fresh and
hamii ( B . & Br.) Wasser i n N o v . Sist. v y s s h . n i z s h . Rast. 1977: 217. untouched, as soon as touched red, turning dark b r o w n , b l a c k i s h i n
('1977') 1978 (superfluous); Leucoagaricus badhamii ( B . & Br.) some minutes, often greyish spotted. L a m e l l a e , L = c. 70-100,1 = (0)1,
H e r i n k i n M y k o l . Sb., Praha 2 1 : 11. ( ' 1 9 8 5 ' ) 1986 (superfluous). - moderately crowded to crowded, free, remote from stipe, hardly ventri-
Lepiota rufovelutina Velen., C e s k e H o u b y : 2 1 7 . 1920. - Lepiota cose to ventricose, up to 5.5 m m wide, cream, w i t h reddish sheen, with
meleagroides H u i j s m a n i n M e d e d . N e d e r l . m y c o l . Vereen. 28: 11.1943 flocculose edge, w h i c h becomes dark. Stipe 35-105 X 3-7 m m , widen-
(not valid). - Lepiota rufovelutina var. sanguinescens Pilat i n A c t a ing gradually towards bulbous base, up to 17 m m w i d e , hollow, at
Mus. nat. Prag. 11B (2): 16. 1955. utmost apex smooth, rest o f stipe w i t h flocculose fibrils, at first white,
E X C L . - Lepiota badhamii sensu H u i j s m a n i n M e d e d . N e d . m y c o l . turning red and then dark brown. A n n u l u s ascending, rather thin, and
Vereen. 2 8 : 6-8. 1943; sensu M e l z e r i n C e s k a M y k o l . 13: 117-119; w i t h 2 m m wide r i m , white o n upper side, discolouring dark o n lower
Leucoagaricus badhamii sensu R o c a b r u n a i n Bolets C a t a l u n y a 8: p i . side, disappearing with age. Context i n pileus and stipe whitish, shiny
378. 1989; Leucocoprinus badhamii sensu Cetto, G r . P i l z f . , 6. A u f l . 1: and brittle i n stipe, immediately red when basidiocarp cut; this discol-
pi. 2 2 . 1980; sensu S c h w o b e l i n Z . P i l z k . 3 2 : 1-2. 1966; sensu oration vanishes after 0.5 minutes. S m e l l none, fungoid-lepiotoid or
W i c h a n s k y i n M y k o l . Sb., Praha 4 1 : 12, p i . 17. 1964 ( i n a l l cases L . unpleasant, rancid. Taste none or unpleasant. Spore print 'pale cream'.
americanus); Lepiota badhamii sensu M i c h a e l & H e n n i g , Handb. A m m o n i a vapour o n lamellae, pileus surface a n d context (olive)
Pilzfr., 3. A u f l . , 3: 139, p i . 16. 1964 ( = Macrolepiota olivieri). green, s l o w l y changing into grey-red. E x s i c c a t e b l a c k i s h .
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 237. 1995; Spores 6.0-9.5(-10.0) X 3.5-6.0 p m , o n average 6.4-8.8 X 4.0-5.1,
Enderle i n B o l l . G r . m i c o l . G . Bres. 42: 176. 1999; Enderle & L a u x i n Q = 1.25-2.0, Q a v = 1.5-1.85, distinctly a m y g d a l i f o r m i n side-view,
M i t t b l . Arbeitsgem. P i l z k . Niederrhein 11 (without page number). 1993. some w i t h an indistinct a p i c a l papilla, a n d a m y g d a l i f o r m i n frontal
S E L . D E S C R . & FIGS. - B a b o s i n S y d o w i a B e i h . 8:45-48, fig. 5 . 1 9 7 9 ; view, without g e r m pore, dextrinoid, c o n g o p h i l o u s , cyanophilous,
Enderle i n B o l l . G r . m i c o l . G . B r e s . 4 2 : 175-177. 1999; H e i l m a n n - metachromatic i n C r e s y l B l u e ; w a l l strongly s w e l l i n g i n a m m o n i a and
acetic acid. B a s i d i a 15-29 X 6.0-9.0 urn, predominantly 4-spored.

L a m e l l a edge sterile. C h e i l o c y s t i d i a abundant, 23-50 X 9.0-18 p m ,
clavate, without but mostly w i t h abrupt apical excrescence, 2.0-25 X
3.0-5.5 um, c y l i n d r i c a l and flexuous, sometimes capitate; green-brown
i n ammonia, often w i t h red-brown concrete granules (these disappear-
i n g after a w h i l e ) and w i t h s m a l l colourless crystals. Pleurocystidia not
observed. P i l e u s completely set w i t h pileocystidia, 90-350 X 8.0-22
pm, w i t h some shorter elements present as w e l l , c y l i n d r i c a l w i t h
rounded apex, w i t h green-brown pigment i n ammonia. Stipitipellis a
cutis o f c y l i n d r i c a l elements, 3.0-5.0 p m i n diam., some w i t h green-
b r o w n pigment i n ammonia, some w i t h refractive walls, set w i t h patent
to adnate c a u l o c y s t i d i a (over w h o l e length o f stipe), v a r y i n g f r o m
clavate to c y l i n d r i c a l , 74-260 X 9-30 p m , w i t h green-brown contents
i n ammonia. Clamp-connections not observed.
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic and
terrestrial i n nutrient-rich habitats, deciduous woods, copses, scattered
throughout the country, f r o m the outer dune ridge to the province o f
Drenthe, rather rare, S e p t . - m i d N o v . W i d e s p r e a d i n Europe, but not
c o m m o n , and preferring temperate to subtropical regions.
F i g . 63. Leucoagaricus georginae.
There is an extremely w i d e range o f spore sizes i n this species, though
two-spored b a s i d i a are not as c o m m o n as i n L. americanus o r L. c o l o u r i n g red i n both pileus and stipe. S m e l l a bit fruity-acrid, faintly
meleagris. reminiscent o f the s m e l l o f Lepiota cristata, but without the rubber
In E u r o p e a n literature o f the first two thirds o f the 20th century this ' component. Taste not k n o w n . Spore print colour not k n o w n .
;
species is often k n o w n as L. bresadolae (for a full discussion see* A m m o n i a vapour o n pileus, lamellae, stipe and context (glaucous)
D e m o u l i n i n L e j e u n i a , n . S. 39: 1-15. 1966; or Krieglsteiner i n Beitr. green. E x s i c c a t e b l a c k i s h .
Kenntn. P i l z e Mitteleur. 7: 39-60. 1991). Spores (5.5-)6.0-8.5 X 3.5-5.0 p m , o n average 6.7-7.7 X 3.9-4.3
R e i d (in M y c o l . R e s . 94: 647. 1990) considered L. badhamii var. um, Q = 1.45-2.25, Q a v = 1.7-1.85, ellipsoid to subcylindrical, slight-
erubescens (Babos) M . B o n a not-blackening variant o f L. badhamii ly fusiform to a m y g d a l i f o r m i n side-view, without germ pore, though
s.str. some w i t h an indistinct pore, dextrinoid, congophilous, cyanophilous,
many not o r hardly c o l o u r i n g i n C r e s y l B l u e , some spores w i t h p i n k
7. Leucoagaricus georginae ( W . G . Sm.) Candusso i n R i v . M i c o l . 3 3 : inner w a l l ; w a l l s w e l l i n g , but not strongly so, i n a m m o n i a and acetic
10. 1990. - F i g . 63 acid. B a s i d i a 15-25.5 X 6.0-8.0 u m , 4-spored. L a m e l l a edge sterile.
Agaricus georginae W . G . S m . i n Seem. J . B o t . 9: 1. 1871; Lepiota C h e i l o c y s t i d i a (13-)17-53 X (8.0-)9.0-18 um, ellipsoid, rarely slight-
georginae ( W . G . Sm.) S a c c , S y l l . F u n g . 5: 7 1 . 1887; Leucocoprinus l y utriform; appendix absent, most often present, rather abrupt, a n d
georginae ( W . G . Sm.) M o s . , R o h r l i n g e , Blatter-, B a u c h p i l z e , 2. A u f l . : then 3.0-25 X 2.5-6.0 p m , slightly m o n i l i f o r m or c y l i n d r i c a l , often
132. 1955 (not v a l i d ) ; Leucocoprinus georginae ( W . G . Sm.) B o n & w i t h c a p i t u l u m , up to 8.5 u m w i d e ; first w i t h reddish pigment i n
B o i f f a r d i n B u l l , trimest. S o c . m y c o l . Fr. 86: 28. 1972 (not valid); a m m o n i a , turning green-brown w i t h some r e d concrete granules.
Leucocoprinus georginae ( W . G . Sm.) Wasser, A g a r . G r i b y S . S . S . R . : Pleurocystidia not observed. Pileus covering w i t h clusters o f c y s t i d i o i d
108. 1985. elements, 4 5 - 2 4 0 X 10-20 p m , narrowly e l l i p s o i d , often w i t h l o n g
S E L . I C O N . - Charbonnel i n D o c . m y c o l . 28 (109-110): p i . I B . 1998. abrupt o r tapering into l o n g appendix, i n t e r m i x e d w i t h clavate ele-
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 401-402, fig. ments c. 30-40 X 10-14 p m , i n some basidiocarps w i t h predominant-
87. 1990; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 166-167. 1994; D . l y clavate elements, i n others w i t h a relatively h i g h number o f
R e i d i n Beitr. K e n n t n . P i l z e Mitteleur. 3: 205-206, figs. 1 & 2. 1987. c y s t i d i o i d elements; pigment parietal and dark b r o w n , especially i n
V E R N . N A M E - Glinsterende champignonparasol. lower part o f cystidioid elements, encrusting i n connecting hyphae, but
also green-brown contents and red granules (as observed i n ammonia)
Pileus 11-30 m m , truncately c o n i c a l to hemispherical w h e n y o u n g , present, also i n connecting repent hyphae. Stipitipellis a cutis o f c y l i n -
expanding v i a plano-convex without o r w i t h l o w u m b o to undulating- drical elements, 3.0-8.0 p m wide, over w h o l e length set w i t h clusters
applanate w i t h flattened centre, set w i t h glittering hairs (lens), m a k i n g o f patent c a u l o c y s t i d i a , 50-270 X 11-22 p m , narrowly e l l i p s o i d to
a granulose-fibrillose impression, white at first, easily turning red to c y l i n d r i c a l w i t h flexuous, slightly m o n i l i f o r m apex; some clavate ele-
red-brown and consequently dark b r o w n w i t h age, when young w i t h ments, as o n pileus, present as w e l l ; pigmentation as i n pileus ele-
fringed, overhanging m a r g i n , exceeding lamellae. L a m e l l a e , L = 2 5 - ments. Clamp-connections absent.
35,1 = 1-3, moderately distant to rather crowded, free, segmentiform H A B I T A T & DISTR. - Gregarious i n s m a l l groups, saprotrophic and
to hardly ventricose, w h i t i s h or cream, sometimes w i t h faint p i n k i s h terrestrial i n several vegetation types, such as deciduous woods o n clay
tinge, w i t h eroded or floccose concolorous edge, w h i c h easily dis- or l o a m y soils, Picea plantations, and dune copses; i n the Netherlands
colours red. Stipe 13-35 X 0.7-2 m m , slightly broadened at base, c y l i n - u n c o m m o n , scattered throughout the country, Oct. D u e to its s m a l l size
drical and flexuous, h o l l o w , white i n untouched places, when touched and inconspicuous colours not recorded often, but apparently w i d e -
orange-red or red, soon turning dark b r o w n to black, over total length spread, not k n o w n f r o m the nordic countries.
densely set w i t h minute, glittering hairs. A n n u l u s s m a l l and membra-
nous and evanescent o r persistent w i t h age, as a narrow band around G o o d photographs o r paintings o f Leucoagaricus georginae hardly
stipe, sometimes w i t h flaring r i m , concolorous w i t h stipe surface, not exist, because o f the s m a l l size and the immediate discoloration o f the
set w i t h hairs. Context white i n pileus, cream and shiny i n stipe, dis- basidiocarps o n the slightest touch.
8. Leucoagaricus croceovelutinus ( B o n & Boiffard) B o n & Boiffard Spores 6.0-9.0 X 3.5-5.0 pm, o n average 7.0-7.7 X 4.2-4.5 pm, Q
i n D o c . m y c o l . 6 (24): 4 5 . 1976. - F i g . 64 = 1.5-2.0, Q a v = 1.6-1.75, o b l o n g - a m y g d a l i f o r m , some w i t h apical
Leucocoprinus croceovelutinus B o n & B o i f f a r d i n B u l l , trimest. papilla, as seen i n side-view, dextrinoid, congophilous, cyanophilous,
Soc. m y c o l . F r . 88: 2 6 . 1972. - Leucocoprinus croceovelutinus var. metachromatic i n C r e s y l B l u e , without germ pore, or w i t h indistinct
diversisporus D . R e i d i n M y c o l . R e s . 94: 658. 1990 (not valid). germ pore (especially as seen i n a m m o n i a and acetic acid), some with
S E L . I C O N . - C h i u s a i n R i v . M i c o l . 42: 47. 1999; H e i l m a n n - C l a u s e n brown contents. B a s i d i a 15-22 X 6.0-8.0 p m , 4-spored. L a m e l l a edge
i n Svampe 26: 19. 1992 (as Leucocoprinus croceovelutinus); Mos. & sterile. Cheilocystidia 26-50 X 8.5-22 p m , ellipsoid, clavate, narrowing
Jul., Farbafl. B a s i d i o m y c . 15: III L e u c o c o p r i n u s 3. 1997. into appendix, or w i t h abrupt appendix 4.0-40 X 3.0-5.0 p m , rarely
S E L . D E S C R . & F I G S . - B a b o s i n S y d o w i a B e i h . 8: 42-45, fig. 4. without appendix; appendix often m o n i l i f o r m , w i t h up to 8.0 p m wide,
1979; B o n & Boiffard i n B u l l , trimest. S o c . m y c o l . F r . 88: 23-26, fig. sometimes branched, capitulum; cystidia w i t h red contents i n ammo-
3. 1972 (as Leucocoprinus croceovelutinus); Heilmann-Clausen i n nia. Pleurocystidia not observed. Pileus covering with globose, ellipsoid
Svampe 2 6 : 20-21. 1992 (as Leucocoprinus croceovelutinus); to clavate elements, 25-55 X 18-33 u m , i n clusters; o n top o f this layer
K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 164-165. 1994; D . R e i d i n squamules w i t h terminal, slightly inflated elements, 130-260 X 16-33
M y c o l . R e s . 9 4 : 655-658, figs. 4 6 - 5 1 . 1990 (as Leucocoprinus cro- pm, and c y l i n d r i c a l connecting hyphae; pigment i n the terminal ele-
ceovelutinus); Schreurs i n C o o l i a 25: 33-38. 1982. ments red-brown i n plates along the inner walls, and fine-encrusting i n
V E R N . N A M E - Verkleurende champignonparasol. connecting hyphae. Stipitipellis a cutis o f c y l i n d r i c a l elements, 3.0-5.0
p m wide, w i t h over w h o l e length o f stipe 10-15 p m w i d e , flexuous
Pileus 30-50 m m , plano-convex to applanate w i t h deflexed margin and hyphae, with red-brown pigment. Clamp-connections not observed.
l o w umbo, w h e n very y o u n g completely w h i t i s h - p i n k i s h velvety, later H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic and
w i t h purple-brown, tomentose-velvety calotte o f the pileus covering, terrestrial i n nutrient-rich places, along sunken roads a n d tracks, i n
around calotte w i t h concentrical zones o f radially arranged triangular deciduous woods a n d dune copses; rather rare i n the Netherlands,
adnate fibrillose squamules, closer to margin w i t h more fibrillose squa- k n o w n from the calcareous dune area, a n d southern L i m b u r g , Sept.-
mules; centre and squamules dark red-brown ( M u . 5 Y R 3/3), paler at Oct. Widespread and rare i n temperate Europe, absent f r o m boreal and
m a r g i n o n c r e a m background; b o t h squamules and b a c k g r o u n d dis- alpine habitats.
colouring orange-red (2.5 R 5/8); m a r g i n exceeding lamellae.
L a m e l l a e , L = c. 8 0 , 1 = 1-3, crowded, free, segmentiform, u p to 4 Leucoagaricus croceovelutinus is easily recognizable because o f the
m m w i d e , pale c r e a m to p i n k w i t h white edge, d i s c o l o u r i n g orange- following combination o f characters: a red discoloration o f a l l parts i n
red ( K . & W . 8 A B 7 to 9 E 8 ) w i t h age, and o n bruising. Stipe 50-85 X ammonia vapour, the spores w i t h an indistinct apical papilla, the pecu-
3-6 m m , tapering upwards, w i t h u p to 15 m m w i d e basal bulb, h o l l o w liar cheilocystidia and the pileus covering, made u p o f two layers o f
and fragile, white, p i n k i s h cream fibrillose, strongly discolouring vina- w i d e cells. C l o s e relatives, l i k e L. badhamii a n d L. georginae, dis-
ceous red (10 R 4/4) w h e n touched. A n n u l u s membranous and evanes- colour green i n a m m o n i a vapour and the pileus covering is made up o f
cent (not always present), and a second ascending annulus w i t h red r i m erect cystidioid elements.
b e l o w the membranous one. Context white, greyish white, shiny i n
stipe, immediately discolouring orange w h e n cut, then p i n k - r e d to dark 9. Leucoagaricus pilatianus ( D e m o u l i n ) B o n & B o i f f a r d i n D o c .
purple-brown, temporarily y e l l o w w i t h a m m o n i a vapour. S m e l l none. m y c o l . 6 (24): 45. 1976. - F i g . 6 5 .
Taste none to slightly bitter. Spore print colour not k n o w n . Lepiota pilatiana D e m o u l i n i n L e j e u n i a , n . S . 39: 11. 1966;
A m m o n i a vapour (not the solution) o n lamellae orange-red. Leucocoprinus pilatianus ( D e m o u l i n ) M o s . , R o h r l i n g e Blatterpilze, 3.
Exsiccate dark red-brown. A u f l . : 186. 1967 (not valid); Leucocoprinus pilatianus (Demoulin)
B o n & Boiffard i n B u l l , trimest. S o c . m y c o l . Fr. 88: 2 6 . 1972 (not
valid); Leucocoprinus pilatianus ( D e m o u l i n ) Wasser i n N o v . Sist.
vyssh. nizsh. Rast. 1977: 219. ( ' 1 9 7 7 ' ) 1978; Leucocoprinus badhamii
var. pilatianus ( D e m o u l i n ) Krieglst. i n Beitr. K e n n t n . P i l z e Mitteleur.
7: 57. 1991. - Leucoagaricus salmoneophyllus B o n & Guinb. i n Doc.
m y c o l . 2 2 (88): 3 1 . 1993; Leucoagaricus pilatianus var. salmoneo-
phyllus (Bon & Guinb.) M i g l . & Gennari i n Gennari & M i g l . i n R i v .
M i c o l . 4 1 : 293. ( ' 1 9 9 8 ' ) 1999. - Lepiota rubens K i i h n e r & M a i r e i n
B u l l . Soc. H i s t . nat. A f r . N . 28: 109. 1937 (not valid).
M I S A P P L . - Lepiota rufovelutina sensu Pilat i n A c t a M u s . natn.
Prag. 9 B (2): 12-18. 1953.
S E L . I C O N . - Cetto, F u n g h i Vero 7: p i . 2 6 0 3 . 1993.
Pileus 22-58 m m , convex to applanate, not umbonate, w h e n y o u n g

with slightly i n c u r v e d margin, finely b l a c k i s h o r dark b r o w n squarru-
lose around disc, b r o w n i s h to pale beige-brown, outwards squamose,
almost smooth or finely punctate, but more c o m m o n l y over w h o l e sur-
face very finely fibrillose o r slightly erect-hairy, hardly s h o w i n g under-
l y i n g context (lens) a n d (greyish) b r o w n , dark at centre, with
orange-brown sheen (fox-tinged), later, exposed and o n damaging, dis-
colouring to dark reddish b r o w n , vinaceous b r o w n to slightly purplish
b r o w n . L a m e l l a e , L = 80-120, 1 = 0 - 1 , very c r o w d e d , free, thin,
crowded, slightly ventricose, 4-7 m m broad, w h i t i s h w i t h faint creamy
F i g . 64. Leucoagaricus croceovelutinus. p i n k tinge, w i t h w h i t i s h fimbriate edge, reddish o r b r o w n i s h w i t h age
1
reddish crystals; some short clavate elements rarely present; connect-

ing hyphae sometimes w i t h encrusting pigment. Stipitipellis a cutis o f
c y l i n d r i c a l to slightly inflated hyphae, 3-12 p m w i d e , b r o w n i s h i n
a m m o n i a , irregularly set w i t h often flexuous, curved very narrowly
clavate to c y l i n d r i c a l c y s t i d i o i d hairs, 35-150 X 7-13 p m , w i t h or w i t h -
out b r o w n contents i n ammonia. Clamp-connections not observed.
H A B I T A T & DISTR. - Solitary to gregarious i n small groups, sapro-
trophic and terrestrial, i n deciduous copse o n l o a m y nutrient-rich soil.
V e r y rare i n the Netherlands, k n o w n f r o m one l o c a l i t y i n southern
L i m b u r g ( E l s l o o - G e u l l e , Bunderbos). Sept. Rare and scattered i n
Europe, not k n o w n f r o m the nordic countries.
T h e description given above i s based o n one c o l l e c t i o n from the

Netherlands, supplemented w i t h data from H u n g a r i a n collections.
Y o u n g specimens, and freshly p i c k e d basidiocarps hardly discolour
w h e n bruised o r treated w i t h a m m o n i a vapour, whereas older speci-
mens do show a distinct discolouration. It was probably this variation
i n discolouration that l e d to the distinction o f a separate variety
(Leucocoprinus pilatianus var. subrubens ( W i c h . ) Wasser). R e i d (in
M y c o l . R e s . 94: 665. 1990) distinguished this variety o n account o f
spore shape (with a blunt apex; more a m y g d a l i f o r m i n the typical vari-
ety). T h e Netherlands material showed more e l l i p s o i d than a m y g d a l i -
form spores; spores were o f both types w i t h i n the H u n g a r i a n \
collections studied.
T h e s m e l l i n the D u t c h c o l l e c t i o n was strongly like cedar w o o d , as
in L. pilatianus var. salmoneophyllus ( B o n & Guinb.) M i g l . & Gennari.
Leucoagaricus aurantiovergens G e n n a r i & M i g l . (in R i v . M i c o l .
41: 291-292. ( ' 1 9 9 8 ' ) 1999), described f r o m Italy comes very close,
but has longer spores a n d cystidia, a n d the stem discolours orange
w h e n scratched.
Leucoagaricus jubilaei (Joss.) M . B o n is a s m a l l species w i t h
clavate cystidia, i n w h i c h the basidiocarps first turn citric yellow, then
red (or not) w h e n cut. Josserand (in B u l l . mens. S o c . l i n n . L y o n , N o .
spec. 43: 205-216. 1974) gave an extensive description o f this taxon
and comparison w i t h other reddening species.
10. Leucoagaricus marriagei ( D . R e i d ) M . B o n i n D o c . m y c o l . 6

F i g . 65. Leucoagaricus pilatianus. (24): 44. 1976. - F i g . 6 6
Lepiota marriagei D . R e i d i n F u n g . rar. i c . C o l . 1: 2 4 . 1966. -
Leucoagaricus marriagei var. ammovirescens M . B o n in Doc. mycol.
or after bruising near pileus margin. Stipe 35-75 X 3-7 m m , up to 5 22 (88): 31. 1993.
m m w i d e at apex, and w i t h gradually bulbous 10 m m w i d e base, taper- S E L . I C O N . - Candusso i n R i v . M i c o l . 3 3 : 1 2 . 1990; M i g l . & Perrone
i n g upwards o r c y l i n d r i c a l , fistulose, w i t h a distinct ring, white a n d in B o l l . A s s . m i c o l . e c o l . R o m a n a 22: 27. 1991; D . R e i d i n F u n g . rar.
smooth above annulus, slightly fibrillose i n l o w e r part, d i s c o l o u r i n g Ic. c o l . 1: p i . 6c & d . 1966.
dark red-brown w h e n rubbed. A n n u l u s hanging, w i t h b r o w n i s h under- S E L . D E S C R . & F I G S . - Candusso i n R i v . M i c o l . 3 3 : 11-14. 1990;
side. Context white, reddening o n exposure (this reaction c a n vary M i g l . & Perrone i n B o l l . A s s . m i c o l . e c o l . R o m a n a 22: 23-30. 1991; D .
among basidiocarps). S m e l l absent, o r strong a n d l i k e cedar w o o d . R e i d i n F u n g . rar. Ic. c o l . 1: 20-22, figs. 11, 12, 14a & b. 1966.
Taste none. Spore print colour not k n o w n . V E R N . N A M E - Gespikkelde champignonparasol.
A m m o n i a o n pileus o f older specimens green, o n lamellae and c o n -
text glaucous green. Pileus 12-25 m m , mostly plano-convex w i t h distinct umbo, i n some
Spores 5.5-9.0 X 3.5-4.5 p m , o n average 6.2-7.2 X 3.9-4.0 u m , Q specimens concave w i t h rounded margin and prominent umbo, p i n k (c.
= 1.5-2.0 (-2.2), Q a v = 1.6-1.8, e l l i p s o i d to oblong, slightly o v o i d to K . & W . 9 D 5 to 8 D 5 ) at centre, paler at margin, d i s c o l o u r i n g to l i l a c
a m y g d a l i f o r m , without g e r m pore, pale orange-brown i n M e l z e r ' s or brownish i n older specimens, velvety at centre, w i t h adnate fibrils
Reagent, congophilous, cyanophilous, w i t h p i n k inner w a l l i n C r e s y l towards margin. L a m e l l a e , L = 45-50,1 = 1-3, moderately crowded,
B l u e ; w a l l strongly s w e l l i n g i n a m m o n i a and acetic acid. B a s i d i a 15- free, sometimes w i t h distinct c o l l a r i u m , whitish to creamy w i t h orange
27 X 6-8.5 p m , 4-spored, i n older specimens slightly t h i c k - w a l l e d . tinge, w i t h fine-flocculose white edge (lens). Stipe 25-92 X 2-4 m m ,
L a m e l l a edge sterile. C h e i l o c y s t i d i a 21-45(-52) X 8.5-13(-17) p m , nar- gradually w i d e n i n g downwards, h o l l o w , glassy cream w h e n moist,
r o w l y clavate, clavate, without apical excrescences, rarely mucronate, white w h e n dry, pubescent over w h o l e length, i n older specimens o n l y
reddish or brown-green i n ammonia. Pleurocystidia absent. Pileus c o v - v i s i b l e above annulus, w i t h s m a l l fibrils turning b r o w n w i t h age i n
ering, relatively densely set w i t h erect, c y s t i d i o i d hairs, 60-340(-400) lower part. A n n u l u s evanescent, w i t h l i l a c edge. Context white i n
X (9-) 12-20 p m , widest just above base, and gradually narrowing to pileus, concolorous w i t h surface i n stipe. S m e l l fungoid. Taste not
rounded apex, pale p i n k i s h to reddish b r o w n i n ammonia, w i t h some k n o w n . Spore print probably white.
Pileus 23-50 m m , campanulate-hemispherical when young, expanding

to applanate without umbo, w i t h more or less closed centre and radially
arranged, small fibrils around centre, dark purple at centre ( K . & W .
14EF4), violaceous purple (c. 1 3 A 3 ) around centre o n white back-
ground; margin fringed and exceeding lamellae. Lamellae, L = 52,1 =
1, moderately crowded, free (but barely), slightly ventricose, up to 6 m m
wide, white with fimbriate, concolorous edge. Stipe 35-44 X 2-5 m m ,
widened to 8 m m w i d e base, hollow, over total length, o r especially
below annulus, set w i t h short lengthwise oriented fibrils, pinkish-pur-
plish (11A5-4), w i t h y e l l o w i s h background shining through. A n n u l u s
ascending, disappearing i n age, with white, hairy-tomentose upper side,
fringed upper margin, and violaceous underside (less p i n k than stipe)
(14-15 A 4-5). Context white i n pileus and brittle, w h i t i s h i n stipe. S m e l l
fungoid-lepiotoid, not distinctive. Taste not k n o w n . Spore print white.
Spores 6.0-7.0x (3.5-)4.0-4.5 um, o n average 6.7 X 4.0 um, Q =
1.5-1.75, Q a v = 1.65, e l l i p s o i d to o b l o n g , w i t h rounded, sometimes
w i t h subacute apex, dextrinoid, congophilous, strongly metachromat-
ic, t h i c k - w a l l e d without g e r m pore. B a s i d i a 14-21 X 5.5-6.5 p m , 4 -
spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 23-40 X 8.0-10 p m ,
narrowly clavate, often a bit irregular, w i t h curved p e d i c e l ; most cys-
tidia w i t h granular hyaline contents. P l e u r o c y s t i d i a absent. Pileus c o v -
F i g . 66. Leucoagaricus marriagei.
ering made up o f adnate to ascending l o n g tapering elements, 110-300
X 8.0-12 um, widest just above base, w i t h intracellular diffuse p i g -
L a m e l l a e slightly turning green w i t h a m m o n i a vapour. ment, and refractive apical part; connecting elements short, 10-18 X
Spores 7.0-9.5 X 4.0-5.0 u m , o n average 8.0-8.4 X 4.4 p m , Q = 5.0-8.0 p m , sometimes w i t h encrusting pigments. Stipitipellis a cutis
I. 6-2.1, Q a v = 1.85-1.9, o b l o n g , often w i t h tapering apex, dextrinoid, of c y l i n d r i c a l , colourless hyphae, c. 5.0 u m w i d e , covered (below
congophilous, cyanophilous, w i t h p i n k inner w a l l i n C r e s y l B l u e , w i t h - annulus) with wider, c. 10-15 p m w i d e , coloured hyphae, w i t h intra-
out germ pore. B a s i d i a 16-25 X 6.5-9 urn, 4-spored. L a m e l l a edge ster- cellular b r o w n i s h - l i l a c i n o u s pigment, w i t h terminal elements w i t h
ile. C h e i l o c y s t i d i a 20-40 X 8-17 p m , broadly utriform, c y l i n d r i c a l o r abrupt l o n g and thin excrescence; terminal elements u p to 2 0 0 p m
slightly fusiform, not coloured, and rather thin-walled. P l e u r o c y s t i d i a long. Clamp-connections not observed.
not observed. P i l e u s c o v e r i n g w i t h fibrils o f c y l i n d r i c a l to slightly H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic and
inflated elements w i t h terminal elements w i t h rounded, slightly taper- terrestrial, o n clayey soil. In the Netherlands recorded f r o m one l o c a l -
i n g apex; terminal elements 40-145 X 10-17 p m , w i t h b r o w n i s h intra- ity i n southern L i m b u r g (Stokhem), where it was g r o w i n g under Picea
cellular pigment; pigment slightly encrusting i n penultimate elements. covered w i t h Clematis vitalba. O c t . R a r e throughout E u r o p e , and
Stipitipellis a cutis o f c y l i n d r i c a l colourless hyphae, 3.0-8.0 p m i n recorded from southern S w e d e n to S p a i n and southern Italy, and from
d i a m . , w i t h some ascending c y l i n d r i c a l , non-differentiated hyphae. Great B r i t a i n to the C z e c h R e p u b l i c .
Clamp-connections absent i n a l l tissues.
H A B I T A T & D I S T R . - Solitary o r i n s m a l l groups, saprotrophic and Leucoagaricus ianthinosquamulosus G u i n b . comes close, but differs
terrestrial i n deciduous woods o n l o a m y o r sandy soils, r i c h i n humus i n longer spores (6.5-10 (-12.5) p m long), 2-spored basidia, and
and nutrients; very rare i n the Netherlands, k n o w n f r o m southern encrusting pigment i n the pileus covering (Guinberteau i n D o c . m y c o l .
L i m b u r g ( E l s l o o - G e u l l e , Bunderbos) and one locality i n the province 22 (88): 10. 1993).
of N o o r d - H o l l a n d ( H e i l o o , H e i l o o e r B o s c h ) ; Sept.-Oct. N o t c o m m o n , A n o t h e r species w i t h violaceous tinges i s L. ianthinophaeus Locq.
but recorded f r o m several countries i n E u r o p e . (in B u l l , trimest. S o c . m y c o l . F r . 68: 177. 1952.), w i t h smaller spores
(up to 6.0 p m long), a n d a pileus c o v e r i n g w i t h tufts o f c y l i n d r i c a l
Leucoagaricus gauguei comes quite close i n general aspects, but dif- elements, intermixed w i t h some inflated elements.
fers i n the structure o f the pileus covering, and the shape o f the spores
and cheilocystidia.
II. Leucoagaricus ionidicolor B e l l u & L a n z o n i i n R i v . M i c o l . 3 1 :

107. 1988. - F i g . 67
Leucocoprinus caeruleoviolaceus D . R e i d i n M y c o l . R e s . 9 3 : 413.
1989; Leucoagaricus caeruleoviolaceus (D. Reid) M . B o n i n Doc.
m y c o l . 23 (91): 3 3 . 1993; Leucoagaricus ionidicolor var. caeruleovi-
olaceus ( D . R e i d ) D . R e i d i n M y c o t a x o n 53: 327. 1995.
S E L . I C O N . - B e l l i ! & L a n z o n i i n R i v . M i c o l . 31: 108. 1988; C o n t u
& Serra i n M i c o l . Veget. medit. 13: 55. 1998.
S E L . D E S C R . & FIGS. - B e l l u & L a n z o n i i n R i v . M i c o l . 31: 107-110.
1988; C o n t u & Serra i n M i c o l . Veget. medit. 13: 55-56. 1998; D . R e i d
i n M y c o l . R e s . 93: 4 1 3 , fig. 1. 1989 (as Leucocoprinus caeruleovio-
laceus); V i l a et a l . i n Rev. catal. M i c o l . 20: 172. 1997.
VERN. NAME - Elfjeschampignonparasol.
F i g . 67. Leucoagaricus ionidicolor.
12. Leucoagaricus melanotrichus ( M a i . & Bert.) T r i m b . i n D o c . Pileus covering w i t h irregularly arranged and sometimes anastomos-
m y c o l . 5 (20): 4 2 . 1975. - F i g . 7 0 . ing, repent to ascending hyphae made up o f u p to 5 coloured, c y l i n -
Lepiota melanotricha M a i . & Bert., F l . C h a m p , sup. M a r o c 1: 134. drical elements; terminal elements not widened, 20-55 X 7.0-10 p m ,
1970. w i t h pale to dark b r o w n intracellular pigment; w a l l s sometimes slight-
V E R N . N A M E - Elegante champignonparasol. l y thickened. Stipitipellis a cutis o f c y l i n d r i c a l , non-coloured hyphae,
3.0-7.0 p m i n d i a m . , i n lower part w i t h patent brown-coloured c y l i n -
K E Y TO THE VARIETIES drical elements 25-80 X 5.0-9.0 p m . Clamp-connections absent i n a l l
1. P i l e u s very dark grey, w i t h p u r p l i s h o r greenish hue tissues.
12a. var. melanotrichus H A B I T A T & DISTR. - Gregarious i n s m a l l groups, saprotrophic and
1. Pileus pale grey-brown at centre 12b. var. fuligineobrunneus terrestrial i n coniferous litter, a n d i n deciduous woods, o n clayey,
rarely sandy, soils. Rare i n the Netherlands, scattered; A u g . - O c t . N o t
12a. var. melanotrichus. - F i g . 68. c o m m o n , but widespread i n Europe, probably absent from boreal and
Leucoagaricus melanotrichus var. septentrionalis D . Reid in (sub-)alpine regions.
M y c o t a x o n 53: 331. 1995.
S E L . I C O N . - C o n t u & C u r r e l l i i n M i c o l . Veget. medit. 11 (2): cen- A variant w i t h relatively s m a l l basidiocarps and slightly a m y g d a l i f o r m
tral page. 1996; Tabares i n B o l e t s Catalunya 10: p i . 483. 1991. spores was described as var. septentrionalis b y R e i d (in M y c o t a x o n 53:
S E L . D E S C R . & F I G S . - B a b o s i n A g a r i c a 6 (12): 206. 1985; K u y p . & 331. 1995). B o t h size o f basidiocarps and spore shape are variable i n
Schreurs i n C o o l i a 26: 74-76. 1983; M a i . & Bert., F l . C h a m p , sup. L. melanotrichus, and R e i d ' s variety is not recognized here.
M a r o c 1: 132-134. 1970; T r i m b . i n D o c . m y c o l . 5 (20): 42-45. 1975. Leucocoprinus heinemannii M i g l . (in M i c o l . ital. 16 (2): 8. 1987)
resembles L. melanotrichus, but has different warmth requirements, as
Pileus 7-20 m m , b l u n t l y c o n i c a l w i t h i n f l e x e d m a r g i n w h e n y o u n g , i n E u r o p e it has o n l y been found i n greenhouses, and the spores are
expanding to applanate or plano-concave w i t h or without broad round- longer: 6.0-9.0(-9.5) X 3.0-4.5(-5.0) p m , Q a v = 1.8-1.85. Migliozzi
ed umbo, at centre dark grey (almost closed) w i t h purplish or greenish & Z e c c h i n (in B e l g . J . B o t . 131: 174. ( ' 1 9 9 8 ' ) 1999) gave a tabular
hue, o r very dark p u r p l i s h b r o w n , felted subtomentose, towards m a r i overview o f the differences between the two species.
gin w i t h squamulose fibrils o n y e l l o w i s h - w h i t i s h or white background^ Two other dark b r o w n to b l a c k i s h , bigger species have been
not or faintly sulcate, thin-fleshed. L a m e l l a e , L = 25-30,1 = 0-3, m o d - described f r o m E u r o p e : L. fuligineodiffractus B e l l u & L a n z o n i (see
erately distant to fairly crowded, free, ventricose a n d up to 3 m m Candusso & L a n z o n i , L e p i o t a : 363-366. 1990), and L. griseodiscus
broad, white-cream w i t h p i n k i s h sheen or distinctly y e l l o w i s h ( M u . 5 ( M . B o n ) B o n & M i g l . B o t h have spores w i t h an apical p a p i l l a , and
Y 8/4) w i t h age, w i t h white (minutely) flocculose edge, w h i c h might differentiated terminal elements i n the pileus c o v e r i n g (for further
turn darker w i t h age. Stipe 10-29 X 1-2 m m , c y l i n d r i c a l o r slightly information see: B o n , F l . m y c o l . Eur. 3, Lepiotes: 94. 1993).
broadened downwards, rarely attenuated upwards, often bent, h o l l o w ,
white or cream at apex, i n l o w e r half w i t h more and more grey to pur- 12b. var. fuligineobrunneus B o n & Boiffard i n M . B o n i n D o c . m y c o l .
p l i s h b l a c k fibrils, red-brown d i s c o l o u r i n g on touching. A n n u l u s m e m - 8 (29): 38. 1978.
branous and evanescent, often not present, thin, w h i t i s h w i t h b r o w n i s h Leucoagaricus melanotrichus f. fuligineobrunneus ( B o n & Boiffard)
to b l a c k i s h edge. Context thin and white i n pileus, concolorous to sur- M . B o n i n D o c . m y c o l . 23 (91): 3 3 . 1993. - Leucoagaricus melan-
face i n stipe. S m e l l indistinct. Taste not k n o w n . Spore print white. otrichus var. fuligineobrunneus f. citrifolius Courtecuisse i n D o c .
Spores 5.5-7.0 X 3.5-4.5(-5.0) p m , o n average 6.1-6.6 X 3.9-4.0 m y c o l . 19 (75): 4. 1989 (not valid).
pm, Q = 1.4-1.8, Q a v = 1.55-1.65, e l l i p s o i d to o b l o n g w i t h straight SEL. D E S C R . & F I G S . - B a s i n C o o l i a 34: 108-109. 1991.
adaxial side i n side-view, not or some spores slightly amygdaliform,
not dextrinoid, though some b e c o m i n g pale orange i n M e l z e r ' s C H A R A C T E R I S T I C S - Pileus w i t h grey-brown colours; lamellae very pale
Reagent, c o n g o p h i l o u s , some spores cyanophilous, w i t h pale p i n k greyish buff or pale beige; stipe w h i t i s h , ochraceous near base.
inner w a l l i n C r e s y l B l u e , without germ pore; w a l l s slightly s w e l l i n g Spores 6.0-7.0 X 3.5-4.5 p m , o n average 6.4-6.7 X 3.9-4.1 u m , Q
i n a m m o n i a a n d acetic a c i d . B a s i d i a 17-23 X 6.5-8.0 p m , 4-spored. = 1.4-1.8, Q a v = 1.65, e l l i p s o i d to o b l o n g w i t h straight abaxial side,
L a m e l l a edge sterile, loosely set w i t h cheilocystidia. Cheilocystidia 28- some slightly a m y g d a l i f o r m ; pigment i n pileus covering elements pale
50 X 7.0-17 p m , broadly c y l i n d r i c a l , sometimes a b i t strangulate o r greyish b r o w n .
flexuous, rarely clavate, without contents. Pleurocystidia not observed. H A B I T A T & D I S T R . - A s t y p i c a l variety, but m u c h rarer, k n o w n from
southern L i m b u r g and M u n t e n d a m (prov. Groningen); Sept.-Oct. Rare
and hardly recorded i n Europe.
T h e rather pale grey-brown colours distinguish this variety f r o m the

t y p i c a l variety. M i c r o s c o p i c a l characters, except for the colour o f the
pigment i n the pileus covering, are similar.
13. Leucoagaricus tener (P.D. Orton) M . B o n i n D o c . m y c o l . 7 (27-

28): 54. 1977. - F i g . 69
Lepiota tenera P . D . O r t o n i n Trans. B r . m y c o l . Soc. 4 3 : 288. 1960;
Leucoagaricus tener (P.D. Orton) M . B o n i n B u l l , trimest. S o c . m y c o l .
Fr. 92: 329. 1976 (not valid).
E X C L . - Leucoagaricus tener sensu M i g l . & C o c c i a i n M i c o l . ital.
19 (1): 22-24. 1990 ( = Leucoagaricus spec).
S E L . D E S C R . & F I G S . - M . B o n i n B u l l , trimest. S o c . m y c o l . Fr. 9 2 :
F i g . 68. Leucoagaricus melanotrichus. 329-331, f i g . 8. 1976; P . D . O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : 288-
provinces (Amsterdam, A m s t e l p a r k and Amsterdamse B o s ; R i j s w i j k ,

Ter Werve; Ter A a r , D e Put). Sept.-Oct. I n E u r o p e rare and scattered.
N o t k n o w n from the nordic countries.
The cheilocystidia i n the D u t c h material are cylindrical, though i n lit-

erature (see selected descriptions) they have been described as clavate.
The cystidia are arranged i n tufts, clearly v i s i b l e i n mature basidiocarps.
Leucoagaricus brunneocingulatus (P.D. Orton) M . B o n i s another
fragile red-brown species, characterized by the absence o f guttules, and
a brown-bordered annulus. S i n c e its o r i g i n a l description (Orton i n
Trans. B r . m y c o l . S o c . 43: 282. 1960) it has been reported only once
from continental E u r o p e ( M i g l i o z z i & Perrone i n B o l l . A s s . m i c o l .
ecol. R o m a n a 23: 3-6. 1991).
14. Leucoagaricus sublittoralis ( K i i h n e r ex H o r a ) S i n g , i n B e i h .

F i g . 69. Leucoagaricus tener. N o v a H e d w i g i a 29: 163. 1969. - F i g . 7 0 .
Lepiota sublittoralis K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 5 2 :
289. 1960; D . R e i d i n M y c o t a x o n 53: 334-335, figs. 2c-f, 3a-d. 1995; 216. 1936 (not valid); Leucocoprinus sublittoralis (Kuhn.) Locq. i n
U l j e i n C o o l i a 27: 10-12. 1984. B u l l . mens. Soc. l i n n . L y o n 14: 9 3 . 1945 (not valid); Leucoagaricus
V E R N . N A M E - Tere c h a m p i g n o n p a r a s o l . sublittoralis (Kiihner) Sing, i n L i l l o a 22: 422. ('1949') 1951 (not
valid); Lepiota sublittoralis K i i h n e r ex H o r a i n Trans. B r . m y c o l . Soc.
Pileus 11-36 m m , when y o u n g hemispherical-paraboloid, w i d e - c o n i - 43: 450. 1960.
cal, expanding v i a campanulate to plano-convex, o r even plano-con- E X C L . - Leucoagaricus sublittoralis sensu B a b o s i n C e s k a M y k o l .
cave, w i t h or without l o w u m b o , at centre densely set w i t h s m a l l 24: 217-219. 1970; sensu D . R e i d i n F u n g . rar. Ic. c o l . 2: 1-2. 1967 ( =
fibrillose squamules, orange-red-brown, red-brown, or dark cinnamon- in both cases L. wichanskyi).
brown, ( M u . 5 Y R 4/4, 7.5 Y R 5/6, between 7/5 Y R 4/6 and 5 Y R 3/4, S E L . I C O N . - M i g l . & M o h r i n M i c o l . ital. 21 (1): p i . 63, 64. 1992;
rarely 7.5 Y R 3/4; K . & W . 6 D 6 - 6 F 6 ) , around centre w i t h s m a l l b r o w n , Partacini i n R i v . M i c o l . 38: 2 8 1 . 1995.
radially arranged fibrils o n white-cream background, slightly sulcate i n S E L . D E S C R . & F I G S . - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 156-
m a r g i n a l zone, w i t h b r o w n guttules w h e n fresh, w h e n y o u n g w i t h 157. 1994; K i i h n e r i n B u l l , trimest. S o c . m y c o l . F r . 52: 216-218. 1936
white velar remnants adhered to margin. Surface b e c o m i n g orange (as Lepiota sublittoralis); Partacini i n R i v . M i c o l . 38: 280-282. 1995.
w h e n touched. L a m e l l a e , L = 45-50, 1 = 1, moderately c r o w d e d to V E R N . N A M E - Duinboschampignonparasol.
moderately distant, free, ventricose, up to 6 m m w i d e , white, slightly
greyish white, sometimes w i t h a sheen o f pink, w i t h white flocculose Pileus 30-55(-70) m m , c. hemispherical expanding to plano-convex,
edge. Stipe 20-40(-70) X 1.5-4 m m , c y l i n d r i c a l and often i n basal part applanate, w i t h l o w broad umbo, and slightly deflexed margin, at cen-
w i d e n e d up to 4.5 m m , often curved i n basal part, h o l l o w , cream and tre p i n k i s h b r o w n ( M u . 7.5 Y R 6/5, 6/4), around centre pale p i n k i s h -
glabrous, and even shiny i n upper part, felted-fibrillose a n d orange- brownish, at centre closed a n d slightly felted, sometimes w i t h fine
p i n k o r b r o w n - c o l o u r e d i n l o w e r part, w i t h white tomentum and cracks, towards m a r g i n r a d i a l l y f i b r i l l o s e to squamulose, adnate o n
m y c e l i u m cords at base, w h e n fresh w i t h y e l l o w to b r o w n guttules, w h i t i s h to cream background, w i t h fringed m a r g i n especially w h e n
especially o n apex. A n n u l u s s m a l l , halfway up stipe, or higher, fuga- young, exceeding lamellae. L a m e l l a e , L = c. 70-90,1 = 0 - l ( - 3 ) , m o d -
cious, patent, w h i t i s h w i t h b r o w n m a r g i n . Context w h i t i s h and very erately distant to c r o w d e d , free, a n d more than 1 m m remote from
thin i n pileus, concolorous w i t h surface i n stipe. S m e l l indistinct, or stipe, segmentiform to ventricose, 3-5 m m broad, white w h e n young,
distinct and then astringent l i k e rubber, suggestive o f L. cristata. Taste slightly cream w i t h age, w i t h white flocculose edge. Stipe 50-65(-100)
not k n o w n . Spore print c o l o u r not k n o w n . X 4-5.5 m m , w i t h (slightly) broadened base up to 12 m m wide, h o l -
Spores 6.5-8.0 X (3.5-)4.0-4.5(-5.0) u m , o n average 7.0-7.4 X 3.9- low, shiny white a l l over, somewhat b r o w n i s h i n lower part w i t h age,
4.2 um, Q = 1.4-2.0, Q a v = 1.6-1.8, ellipsoid to oblong, often amyg-
daliform i n side-view, without germ pore, dextrinoid, congophilous,
cyanophilous, a n d metachromatic in Cresyl Blue; wall strongly
swelling i n a m m o n i a and acetic acid, or hardly s w e l l i n g . B a s i d i a 18-26
X 6.5-8.5 p m , 4-spored. L a m e l l a edge sterile i n y o u n g specimens, later
w i t h tufts o f cheilocystidia. C h e i l o c y s t i d i a clustered, 19-75 X 6.5-13
um, c y l i n d r i c a l , very narrowly clavate, slightly fusiform, sometimes
w i t h strange appendix o n apex, colourless a n d slightly t h i c k - w a l l e d .
Pleurocystidia not present. Pileus covered w i t h repent and ascending
hyphae, made up o f c y l i n d r i c a l elements; terminal elements 30-130 X
6.0-9.0 p m , w i t h b r o w n intracellular pigment; pigment intracellular o r
encrusting i n penultimate elements. Stipitipellis a cutis o f cylindrical,
colourless hyphae, c. 5 p m i n diam., w i t h i n lower part o f stipe, irregu-
lar, flexuous, brown-coloured hyphae, w i t h terminal elements narrow-
ly clavate, 10-12 p m i n d i a m . Clamp-connections absent i n a l l tissues.
H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial o n clay
or humus-rich sandy soil, i n both coniferous and deciduous woods. I n
the Netherlands very rare, k n o w n from several localities i n the western F i g . 70. Leucoagaricus sublittoralis.
innately fibrillose. A n n u l u s ascending, white, w i t h p i n k i s h brownish of L. wichanskyi, and a bulbous stipe base i n the same species), and
edge, disappearing w i t h age. Context white i n stipe and pileus. S m e l l some m i c r o s c o p i c a l characters, especially shape o f the cheilocystidia
fungoid, l i k e the rubber component o f s m e l l o f L. cristata. Taste as and construction o f the pileus covering.
smell. Spore print probably white. O c c a s i o n a l l y squamose variants o f L. sublittoralis are found. These
Spores 6.5-10.0 X 3.5-5.0 u m , o n average 7.4-8.5 X 3.8-4.3 u m , Q have been excluded f r o m the above description. T h e squamosity may
= 1.5-2.2, Q a v = 1.7-2.0, o b l o n g - a m y g d a l i f o r m , often w i t h apical be caused b y dry g r o w i n g conditions.
papilla, rarely w i t h rounded apex, without germ pore, dextrinoid, c o n - Leucoagaricus gaillardii B o n & B o i f f a r d (in B u l l , trimest. S o c .
gophilous, cyanophilous, metachromatic i n C r e s y l B l u e . B a s i d i a 17- m y c o l . Fr. 90: 295. ('1974') 1975) is a b i g , fleshy representative o f this
26 X 6.0-8.0 p m , 4-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 21-70 group, w i t h a rather squamose pileus covering and non-amygdaliform
X 6.0-14 p m , c y l i n d r i c a l , narrowly clavate, narrowly utriform, some- spores. T h i s species has been found along the southern F r e n c h Atlantic
times subcapitate, colourless, most without, rarely w i t h some crystals and the mediterranean coasts. P r i o u et al. (in D o c . m y c o l . 25 (98-100):
on apex, often inside the cystidia. Pleurocystidia absent. Pileus cover- 349-358. 1995) gave an extensive overview o n the morphology, e c o l -
ing w i t h l o n g , radially arranged adnate c y l i n d r i c a l hyphae, 3.0-6.0 u m ogy and distribution o f this species.
wide, w i t h encrusting pigment, l o c a l l y w i t h clusters o f short hyphae Leucoagaricus glabridiscus (Sundb.) Wuilbaut, originally
and n a r r o w l y clavate-fusiform elements, 60-130 X 14-24 p m . described f r o m the western U . S . A . , and reported f r o m B e l g i u m b y
Stipitipellis a cutis o f c y l i n d r i c a l , colourless hyphae. C l a m p - c o n n e c - W u i l b a u t (in D o c . m y c o l . 17 (65): 46. 1986), is close to L. sublittoralis,
tions absent. but differs i n the copper to brown-red colours o f the pileus.
H A B I T A T & DISTR. - Solitary to gregarious i n s m a l l groups, sapro- Leucoagaricus subolivaceus M i g l . & Perrone (in B o l l . A s s . m i c o l .
trophic and terrestrial o n both sandy and clayey-loamy soils, i n copses, ecol. romana 9 (27): 44-45. 1992) is also close, but characterized b y
deciduous woods etc. In the Netherlands recorded from the dune area olivaceous colours i n the pileus; i t occurred i n sandy terrain under
(especially the inner dune region) and from several localities i n south- Quercus ilex i n the mediterranean region.
ern L i m b u r g . Sept.-Nov. R e c o r d e d f r o m several countries i n Europe,
where the species occurs i n l a n d as w e l l as i n the coastal regions, not 15. Leucoagaricus wichanskyi (Pilat) B o n & B o i f f a r d i n B u l l . ^,
recorded north o f Denmark. » trimest. S o c . m y c o l . Fr. 9 0 : 303. ('1974') 1975 (as L. wychanskyi). -
F i g . 71
Leucoagaricus sublittoralis is very close to or perhaps identical w i t h Lepiota wichanskyi P i l a t i n A c t a M u s . natn.. P r a g . 9 B (2): 4 - 1 1 .
L. littoralis ( M e n i e r ) M . B o n . K u h n e r (in B u l l , trimest. Soc. m y c o l . Fr. 1953.
52: 2 1 6 - 2 1 8 . 1 9 3 6 ) described L. sublittoralis from inland, and not from M I S A P P L . - Lepiota sublittoralis sensu D . R e i d i n F u n g . rar. Ic. c o l .
the coast as M e n i e r ' s L. littoralis. A c c o r d i n g to B o n ( F l . m y c o l . Eur. 2: 1-2. 1967.
3, Lepiotes: 9 8 . 1993) the m a i n difference between L. littoralis and L . S E L . I C O N . - Cetto, F u n g h i V e r o 7: p i . 2 6 0 1 . 1993; Z u c c h e r e l l i ,
sublittoralis i s the w i d t h o f the pilear elements: 6-10(-15) p m i n L. lit- F u n g h i Pinete Z o n e medit.: p i . 275. 1993.
toralis, and (10-)15-20(-25) u m i n L. sublittoralis. Furthermore, L. lit- S E L . D E S C R . & FIGS. - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 158-
toralis is the more sturdy o f the t w o taxa. A c c o r d i n g to K u h n e r the 159. 1994; P i l a t i n A c t a M u s . natn. Prag. 9 B (2): 4-11. 1953.
elements are 10-19 p m w i d e i n L. sublittoralis. In the D u t c h speci- VERN. NAME - Knolchampignonparasol.
mens, originating from both i n l a n d and the coastal area, the terminal
elements are n a r r o w l y clavate-fusiform, 14-25 p m w i d e . H o r a (in Pileus 35-80 m m , w h e n y o u n g hemispherical, w i t h age expanding to
Trans. B r . m y c o l . S o c . 4 3 : 450. 1960), w h o v a l i d l y described L. sub- plano-convex and (sub-)umbonate, sometimes w i t h shallow depression
littoralis, l i t e r a l l y translated K i i h n e r ' s description, but indicated a around umbo, flattened at umbo, w h i c h is covered w i t h some white
B r i t i s h c o l l e c t i o n as type, w h i c h subsequently got lost. T h e neotype, v e l u m patches, reddish b r o w n , p i n k - b r o w n , p u r p l i s h b r o w n ( M u . 2.5
designated b y R e i d (in F u n g . rar. Ic. c o l . 2: 1-2. 1967), represents L. Y R 6/4, 5-10-YR 7/6), paler at margin, to slightly ochraceous b r o w n or
wichanskyi. /
almost w h i t e ( 5 Y R 7/3-8/4 and paler); at first covering made up o f
Some authors (see excluded names) have considered L. sublittoralis 'closed' fibrils, later, except for centre, breaking up into looser fibrils or
and L. wichanskyi synonyms. B u t the two taxa are easily separated b y squamules, showing the white to p i n k - w h i t e context, under dry cir-
some m a c r o s c o p i c a l characters (presence o f a velar patch o n the pileus cumstances more squamose, under wet conditions more fibrillose; mar-
gin sometimes fringed w i t h white velar flocks. L a m e l l a e , L = 70-110, 16. Leucoagaricus purpureolilacinus H u i j s m a n i n Fungus 25: 33.
1 = 1-5, crowded, rather crowded, free and forming a collarium, slight- 1955. - F i g . 72.
ly ventricose, 2.5-6.0 m m wide, first white, later creamy, often w i t h sor- Leucoagaricus purpureorimosus B o n & Boiffard i n M . B o n i n D o c .
did grey tinge, w i t h even or slightly flocculose (lens), concolorous edge. m y c o l . 8 (29): 37. 1978. - Leucoagaricus purpureorimosus f. pallidus
Stipe 30-85 X 4-10 m m , w i t h distinctly widened abrupt, rarely clavate, B o n & Boiffard i n M . B o n i n D o c . m y c o l . 8 (29): 37. 1978.
often submarginate bulb, 10-20 m m w i d e , i n very y o u n g specimens S E L . I C O N . - L o n a t i i n M i c o l . Veget. medit. 6: p i . 8. 1991; M i g l . &
w i t h some v e l u m remnants o n bulb margin, tapering towards apex, h o l - M o h r i n M i c o l . ital. 21 (1): p i . 64. 1992 (both as L. purpureorimosus).
low, white, not o r hardly discolouring (brown) o n bruising, shiny and S E L . D E S C R . & F I G S . - H u i j s m a n i n F u n g u s 2 5 : 3 3 . 1955.
lengthwise innately fibrillose. A n n u l u s descending, rarely ascending, V E R N . N A M E - Purperen c h a m p i g n o n p a r a s o l .
evanescent w i t h age, white w i t h ultimate edge p i n k - b r o w n as o n pileus.
Context i n pileus white, rather thick, white i n stipe, w i t h age yellowish Pileus 35-60 m m , paraboloid-hemispherical at first, b l u n t l y c o n i c a l ,
or slightly (orange-)brown i n bulb. S m e l l o f y o u n g specimens none, expanding to applanate w i t h l o w broad u m b o , p u r p l i s h b r o w n , dark
later faintly like the smell o f L. cristata. Taste none. Spore print white. purplish red-brown, dark vinaceous b r o w n ( M u . 5 Y R 5/3, near 10 R
Spores (6.5-)7.0-9.5(-10.0) p m , o n average 7.4-8.5 X 4.1-4.6 p m , 3/1.5) at centre, i n rest o f pileus p i n k - b r o w n , pale b r o w n , pale ochra-
Q = 1.5-2.25, Q a v = 1.65-1.9, i n side-view a m y g d a l i f o r m , often w i t h ceous (5 Y R 6/4, 10 Y R 6/3), finely fibrillose-squamulose, i n bigger
indistinct a p i c a l p a p i l l a , very rarely o b l o n g and not a m y g d a l i f o r m , specimens w i t h r a d i a l l y radiating, reticulately connected, slightly
dextrinoid, c o n g o p h i l o u s , cyanophilous, w i t h metachromatic inner raised to sharp ridged darker venation, w i t h a covering w h i c h i s bro-
w a l l i n C r e s y l B l u e , without germ pore. B a s i d i a 18-27 X 6.5-10 p m , ken up at pileus margin, showing the white underground, w i t h white
predominantly 4-spored; number o f 2-spored basidia variable. L a m e l l a velar remnants at margin, at least w h e n y o u n g . L a m e l l a e , L = 65-80,
edge sterile. C h e i l o c y s t i d i a 18-55 X 4.0-11.5 u m , c y l i n d r i c a l , narrow- 1 = 1-3, crowded, free and remote f r o m stipe, segmentiform, subven-
ly lageniform, narrowly clavate, or narrowly utriform, sometimes some tricose, 4-6 m m w i d e , white to pale cream, w i t h concolorous or white
subcapitate, i n dried specimens w i t h pale grey contents, sometimes flocculose edge. Stipe 45-75 X 6-14 m m , gradually widened, but more
w i t h some s m a l l external apical crystals. Pleurocystidia not observed. often tapering towards base, straight o r flexuous, h o l l o w , w i t h rooting
Pileus covering made up o f l o n g , tapering colourless elements, w i t h or basal part, white, w i t h faint lilacinous tinge at base, finely fibrillose,
without septa, 75-250 X 5.0-16 u m , originating i n groups from short slightly shiny, glabrous. A n n u l u s evanescent, ascending, o r descend-
connecting elements, w i t h densely encrusting brown pigment. ing, membranous-felted, white, later b r o w n i s h . Context i n pileus thick,
Stipitipellis a cutis o f non-coloured c y l i n d r i c a l elements, 3.0-6.0 p m white and dull, white and shiny i n stipe, not d i s c o l o u r i n g . Spore print
wide. Clamp-connections not observed. white w i t h a pale p i n k i s h tinge.
H A B I T A T & D I S T R . - Gregarious i n s m a l l groups o r solitary, sapro- Spores 7.5-11.5 X 4.0-6.5 u m , o n average 8.7-10.3 X 4.2-5.3 p m ,
trophic and terrestrial, o n sandy and o n c l a y e y - l o a m y soils, i n decidu- Q = 1.6-2.3, Q a v = 1.8-1.95, a m y g d a l i f o r m i n side-view, often w i t h
ous woods, i n a l a w n , i n copses. I n the Netherlands very rare, k n o w n apical papilla, rounded-fusiform i n frontal view, without germ pore,
f r o m a few localities i n the dune region ( H e i l o o , H e i l o o e r bosch; dextrinoid, congophilous, cyanophilous, w i t h p i n k inner w a l l i n C r e s y l
Heemstede, L e y d u i n ) and w i t h certainty f r o m one locality i n southern B l u e . B a s i d i a 18-29 X 6.5-10 u m , 4-spored, rarely 2-spored. L a m e l l a
L i m b u r g (Neercanne, Cannerberg), perhaps more widespread i n that edge sterile. C h e i l o c y s t i d i a 19-41 X 6.0-15 u m , variable i n shape,
area; Sept.-Oct. R a r e l y recorded i n Europe, probably because i t is not
always recognized; w i t h distribution centres both i n central Europe and
i n coastal areas.
T h e above description i s based o n Netherlands a n d extralimital

material.
Leucoagaricus wichanskyi is m a c r o s c o p i c a l l y characterized b y the
remains o f the v e i l o n the pileus centre, and the bulbous base w i t h velar
remnants o n its rim. M i c r o s c o p i c a l l y the rather narrow cheilocystidia
w i t h grey contents and the structure o f the pileus covering are remark-
able. Spore sizes can not be used to distinguish species i n this group,
as the number o f 2-spored basidia varies, a n d spores are easily m i s -
shapen b y storage.
Some authors (e.g. B a b o s i n C e s k a M y k o l . 24: 217-219. 1970; R e i d
i n F u n g . rar. Ic. c o l . 6: 16-17. 1972) considered L. sublittoralis and L .
wichanskyi conspecific. H o w e v e r , the differences between the t w o
species are quite obvious, both i n m a c r o s c o p i c a n d i n m i c r o s c o p i c
characters.
Leucoagaricus idae-fragum G u i n b . et a l . , described from France,
(Guinb. et a l . i n B u l l , trimest. S o c . m y c o l . Fr. 114: 1-18. 1998) i s also
characterized by a velar patch o n the pileus, but differs from L. wichan-
skyi b y the raspberry-red pileus, the spores w i t h rounded apex, and the
clavate cheilocystidia.
Leucoagaricus rubrotinctus (Peck) S i n g , i s a N o r t h A m e r i c a n
species, o c c a s i o n a l l y reported f r o m E u r o p e , especially f r o m green-
houses and the mediterranean region ( B o n , F l . m y c o l . Eur. 3, Lepiotes:
100. 1993), a n d characterized b y a reddish pileus, without a velar
patch, n o n - a m y g d a l i f o r m spores, and l o n g c y l i n d r i c a l cheilocystidia.
clavate, rarely broadly clavate, c y l i n d r i c a l , often w i t h median c o n - Pileus 14-60 m m , w h e n y o u n g campanulate, expanding to applanate
striction, n a r r o w l y lageniform, slightly thick-walled, w i t h crystals i n w i t h l o w u m b o and deflexed m a r g i n , first totally white to cream, then
upper part o r o n apex only. Pleurocystidia absent. Pileus covering i n at centre w i t h granular-erect y e l l o w - b r o w n - p i n k , p i n k - b r o w n to l i l a c -
centre o f pileus w i t h gelatinized adnate hyphae, c. 3.0 u m i n d i a m . , b r o w n squamules, towards m a r g i n more and more spread out, show-
l y i n g o n top o f ellipsoid, coloured elements; near pileus margin w i t h ing the w h i t i s h f i b r i l l o s e and squamulose b a c k g r o u n d , f i n a l l y b r o w n
l o n g c y l i n d r i c a l adnate elements, up to 300 X 15 pm, w i t h encrusting to dark b r o w n , g r o o v e d and fringed i n m a r g i n a l 0.5 c m . L a m e l l a e , L
pigment. S t i p i t i p e l l i s a cutis o f narrow, 3.0-10 p m wide, colourless, = c. 6 0 , 1 = 1-2, c r o w d e d , pale cream-s'white, free, s l i g h t l y ventri-
c y l i n d r i c a l elements. Clamp-connections absent. cose, o r segmentiform, u p to 4.5 m m w i d e , w i t h w h i t e f l o c c u l o s e
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic and edge. Stipe 35-65 X 2-5 m m , tapering towards apex, o r c y l i n d r i c a l ,
terrestrial, i n dune grasslands, i n woods o n the inner side of the dunes, straight o r bent, h o l l o w , p i n k i s h cream, w i t h annulus, adnately
probably also i n woods o n l o a m y calcareous soil i n L i m b u r g , rare i n tomentose-fibrillose, b e l o w annulus s l i g h t l y squamulose, b e c o m i n g
the Netherlands. O c t . - N o v . R a r e and widespread along the coasts i n b r o w n i s h w h e n touched. A n n u l u s halfway u p stipe, w i t h b a n d as cuff
E u r o p e f r o m the Netherlands southwards. around stipe, w i t h r i m up to 2 m m w i d e , w i t h s l i g h t l y orange tinge at
fringed m a r g i n ; upper side w o o l l y - g r a n u l o s e . C o n t e x t w h i t i s h i n
Leucoagaricus purpureolilacinus i s characterized b y the dark pur- pileus, i n stipe s i l k y shiny and b e c o m i n g ochraceous b r o w n . S m e l l
p l i s h c o l o u r s at the centre o f the pileus, c y s t i d i a c r o w n e d w i t h crys- indistinct, s l i g h t l y f u n g o i d . Taste not distinct. Spore print w h i t e
tals, a n d rather b i g spores w h i c h resemble those o f Lepiota creamy.
subgracilis i n shape. Spores 6.5-8.5 X 4.0-5.0 p m , o n average 7.2 X 4.2 um, Q = 1.55-
Leucoagaricus purpureorimosus B o n & B o i f f a r d is considered a 1.9, Q a v = 1.8, a m y g d a l i f o r m , w i t h , rarely without, small apical papil-
s y n o n y m , as n o discriminating characters c o u l d be found. B o t h taxa la, dextrinoid, congophilous, cyanophilous, metachromatic i n C r e s y l
are reported f r o m dune areas. B l u e , without g e r m pore;. B a s i d i a 15-24 X 6.0-9.0 p m , predominant-
ly 4-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 24-57 X 6.5-10 u m ,
17. Leucoagaricus gauguei B o n & Boiffard i n i n B u l l , trimest. Soc. c y l i n d r i c a l , often slightly thickened at apex. P l e u r o c y s t i d i a absent. (
m y c o l . Fr. 90: 290. ( ' 1 9 7 4 ' ) 1975. - F i g . 73 Pileus squamules made up o f erect elements, 20-130 X 6-14 pm, c y l i n -
Sericeomyces gauguei ( B o n & Boiffard) H e i n e m . i n B u l l . Jard. bgst. drical w i t h rounded apex, or tapering towards apex, w i t h diffuse, pale
natn. B e l g . 48: 405. 1978. intracellular pigment; these terminal elements are situated o n short,
E X C L . - Leucoagaricus gauguei sensu M i g l . & Perrone i n M i c o l . isodiagonal elements, e.g. 5 . 0 1 0 p m w i d e , w i t h refringent w a l l s .
:
ital. 21 (1): 37-39. 1991 ( = Leucoagaricus spec.) Clamp-connections absent.

S E L . I C O N . - L o n a t i i n B o l l . A s s . m i c o l . ecol. R o m a n a 8-9: 8. 1986. H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic and ter-
S E L . D E S C R . & FIGS. - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 152- restrial i n deciduous woods, both on humus-rich sand, and o n calcare-
153. 1994. ous clay. Very rare i n the Netherlands, recorded from the dune region
V E R N . N A M E - Lilavezelige champignonparasol. (Bloemendaal, O u d Woestduin; Amsterdamse Waterleidingduinen), and
southern L i m b u r g (Bemelen). Sept.-Oct. Rare and scattered i n Europe,
but not k n o w n from the nordic countries.
T h e spore sizes reported b y K e l d e r m a n (Parasolzw. Z u i d - L i m b u r g :

153. 1994) for this species are smaller than given here, as his descrip-
tion is based o n y o u n g material, without fully mature spores.
18, Leucoagaricus sericifer ( L o c q . ) V e l l i n g a i n Persoonia 17: 4 7 7 .

2000. - F i g . 74.
Lepiota cristata var. sericea C o o l i n M e d e d . N e d . m y c o l . Vereen.
12: 23. 1922; Lepiota sericea ( C o o l ) H u i j s m a n i n M e d e d . N e d . m y c o l .
Vereen. 28: 4 6 . 1 9 4 3 , non Lepiota sericea M a s s . , 1912; Leucoagaricus
sericeus ( C o o l ) B o n & Boiffard i n M . B o n i n D o c . m y c o l . 9 (35): 4 0 .
1979 (not v a l i d ) ; Sericeomyces sericeus (Cool) Contu i n Cryptog.,
M y c o l . 12: 6. 1991 (not valid); Pseudobaeospora sericifera Locq. in
B u l l , trimest. Soc. m y c o l . Fr. 68: 169. 1952; Lepiota sericifera (Locq.)
L o c q . i n F r i e s i a 5: 294. 1956; Sericeomyces sericifer (Locq.) D0ssing
in K n u d s e n & H a n s e n i n N o r d i c J . B o t . 11: 4 8 1 . 1991; Lepiota seri-
cata K i i h n . & R o m a g n . , F l . anal. C h a m p , sup.: 405. 1953 (superflu-
ous); Sericeomyces sericatus (Kiihn. & Romagn.) Heinem. i n B u l l .
Jard. bot. natn. B e l g . 48: 404. 1978. - Lepiota sericatella Malenc. i n
M a i . & Bert., F l . C h a m p , sup. IVJaroc 1: 152. 1970; Leucoagaricus ser-
icatellus ( M a l e n c . ) M . B o n i n D o c . m y c o l . 9 (35): 4 0 . 1979;
Sericeomyces sericatellus ( M a l e n c . ) M . B o n i n B u l l , trimest. S o c .
m y c o l . F r . 9 6 : 172. 1980; Sericeomyces sericatus var. sericatellus
(Malenc.) H e i n e m . i n B u l l . Jard. bot. natn. B e l g . 4 8 : 4 0 4 . 1978;
Leucoagaricus sericifer f. sericatellus (Malenc.) V e l l i n g a i n Persoonia
17: 479. 2000.
M I S A P P L . - Lepiota serena sensu J . L a n g e , F l . agar. dan. 1: p i . 1 I B .
F i g . 73. Leucoagaricus gauguei. 1935.
age, w i t h white, entire to eroded edge. Stipe 20-90 X 2.5-4 m m ,

broadening d o w n w a r d to slightly inflated to subclavate base, 3-9 m m
w i d e , h o l l o w , white at apex, to pale p i n k i s h o r p i n k i s h - b r o w n i s h
below (by contact), subpruinose at apex to glabrous and shiny, w i t h o r
without annulus. A n n u l u s evanescent, ascending, w i t h c u f f around
stipe, and narrow flaring part, w h i t i s h w i t h slightly darker edge, espe-
c i a l l y after d r y i n g and h a n d l i n g . Context thin, soft, white and d u l l i n
pileus, shiny, w h i t i s h , cream-coloured to pale p i n k i s h b r o w n i n stipe.
S m e l l rather indistinct, not unpleasant, a bit sweetish. Taste indistinct.
Spore print probably white.
Spores i n side-view 6.0-9.5(-10.0) X 3.0-4.5(-5.0) p m , o n average
7.1- 8.2 X 3.5-4.3 u m , Q = 1.65-2.35, Q a v = 1.8-2.05, o b l o n g - a m y g -
d a l i f o r m to c y l i n d r i c a l - a m y g d a l i f o r m , without, o r w i t h a p i c a l p a p i l -
la; i n f. sericatellus 8.0-12.5 X 4.0-5.5 p m , o n average 9.2-10.7 X
4.2- 4.5 p m , Q = 1.75-2.6, Q a v = 2.1-2.35; o b l o n g to c y l i n d r i c a l w i t h
a m y g d a l i f o r m aspect, some more f u s i f o r m , w i t h the same k i n d o f
shape as spores o f Lepiota clypeolaria, dextrinoid, congophilous,
cyanophilous, metachromatic i n C r e s y l B l u e . B a s i d i a 15-34 X 6.0-
9.5 u m , predominantly 4-spored i n f. sericifer, 2-spored w i t h some 1-
spored b a s i d i a i n f. sericatellus. Lamella edge w i t h tufts o f
cheilocystidia. C h e i l o c y s t i d i a very variable i n shape and size, 17-90
X 6.5-22 p m ( i n c l u d i n g neck), l a g e n i f o r m w i t h broad body, slender
neck and s l i g h t l y w i d e n e d c a p i t u l u m , l a g e n i f o r m , clavate, n a r r o w l y
l a g e n i f o r m a n d slender, o r i r r e g u l a r l y shaped, c o l o u r l e s s , w h e n
l a g e n i f o r m neck t h i n - w a l l e d , and b o d y t h i c k - w a l l e d . P l e u r o c y s t i d i a
not observed. P i l e i p e l l i s a cutis w i t h bundles o f r a d i a l l y arranged
hyphae, inflated, 7.0-25 p m w i d e , w i t h rounded apices at the ends;
pigment pale y e l l o w - b r o w n and i n t r a c e l l u l a r i n the b i g g e r elements.
S t i p i t i p e l l i s a cutis o f c y l i n d r i c a l hyphae, 3.0-5.0 p m w i d e ; pale
F i g . 74. Leucoagaricus sericifer. b r o w n i n t r a c e l l u l a r p i g m e n t present i n u n d e r l y i n g w i d e r hyphae.
C l a m p - c o n n e c t i o n s absent.
S E L . I C O N . - M i g l . i n B o l l . G r u p p o m i c o l . G . B r e s . 4 0 (1): 21. 1997 H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic a n d
(as Sericeomyces sericifer); M i g l . & C o c c i a i n M i c o l . ital. 2 0 (2): p i . terrestrial on sandy, l o a m y and clayey soils, i n deciduous woods, rather
62. 1991 (as L. sericeus). rare, scattered throughout the country, w i t h the typical f o r m more c o m -
S E L . D E S C R . & F I G S . - H u i j s m a n i n M e d e d . N e d . m y c o l . Vereen. 28: mon than f. sericatellus. A u g . - O c t . Widespread i n E u r o p e .
44-46. 1943 (as Lepiota sericea); K e l d e r m a n , Parasolzw. Z u i d -
L i m b u r g : 146-149. 1994 (as L. sericatellus a n d L. sericeus resp.); Leucoagaricus sericifer i s variable i n terms o f spore size, a n d the
L o c q . i n F r i e s i a 5: 2 9 4 - 2 9 5 . 1 9 5 6 (as Lepiota sericifera); M a i . & Bert., shape a n d size o f the c h e i l o c y s t i d i a ( V e l l i n g a i n P e r s o o n i a 17: 477-
F l . C h a m p , sup. M a r o c 1: 150-151, f i g . 2 6 . 1970 (as Lepiota seri- 479. 2000).
catella); M i g l . & B i z z i i n M i c o l . ital. 24 (3): 81, 84, f i g . 7. 1995 (as K e l d e r m a n (Parasolzw. Z u i d - L i m b u r g : 149. 1994) reported that L.
Sericeomyces sericatellus); M i g l . & C o c c i a i n M i c o l . ital. 20 (2): 24- sericifer has abundant crystals o n the cystidia, but this c o u l d not be
28. 1991 (as L. sericeus). confirmed.
VERN. NAME - Zijdechampignonparasol, incl. Tweesporige The spores formed o n 1-spored basidia, w h i c h are often present i n
champignonparasol. f. sericatellus, are even longer than the sizes given above; lengths,up
to 15 u m have been found.
K E Y TO THE FORMAE Leucoagaricus medioflavoides M . B o n (in D o c . m y c o l . 6 (24): 4 4 .
1. B a s i d i a predominantly 4-spored; spores 6.0-9.5 X 3.0-4.5(-5.0) p m 1976) differs from L. sericifer i n the y e l l o w to sulphur coloured pileus
f. sericifer centre, and the s m a l l (up to 6.5 p m long) spores. T h i s species has been
1. B a s i d i a predominantly 2-spored; spores 8.0-12.5 X 4.0-5.5 u m recorded from D e n m a r k to Italy.
(longer f r o m 1-spored basidia) f. sericatellus
19. Leucoagaricus serenus (Fr.) B o n & B o i f f a r d i n B u l l , trimest. S o c .
Pileus 11-45 m m , bluntly c o n i c a l , conico-convex, expanding to plano- m y c o l . Fr. 90: 301. 1974. - F i g . 75.
convex, to almost applanate, always w i t h s m a l l u m b o , w i t h very pale Agaricus serenusFr., H y m e n o m y c . Eur.: 38: 1874; Lepiota serena
ochraceous cream, y e l l o w i s h centre, ( M u . 2.5 Y 8/4 to 10 Y R 8/4), (Fr.) Q u e l , i n B u l l . S o c . bot. France 26: 4 5 . 1880; Pseudobaeospora
w h i t e around centre, w i t h age tending to become very pale p i n k i s h serena (Fr.) L o c q . i n B u l l , trimest. S o c . m y c o l . F r . 6 8 : 169. 1952;
(7.5 Y R 8/4 to 5 Y R 8/3), felted at centre, around centre w i t h r a d i a l l y Sericeomyces serenus (Fr.) H e i n e m . i n B u l l . Jard. bot. natn. B e l g . 48:
arranged, shiny-satiny fibrils, more o r less squamulose-fibrillose; 403. 1978.
those fibrils (hazel-)brown w i t h age and o n d r y i n g ; m a r g i n irregular- E X C L . - Lepiota serena sensu J . L a n g e , F l . agar. dan. 1: p i . 1 1 B .
l y fringed w h e n y o u n g and exceeding lamellae, b e c o m i n g b r o w n b y 1935 ( = L. sericifer). - Lepiota serena sensu A . Pears, i n Trans. B r .
one day after p i c k i n g . L a m e l l a e , L = c. 5 5 - 7 0 , 1 = 0-3, very c r o w d - m y c o l . Soc. 32: 258. 1949 ( = L. pinguipes). - Leucoagaricus serenus
ed to moderately c r o w d e d , free, segmentiform to slightly ventricose, sensu M . B o n i n D o c . m y c o l . 11 (43): 5 2 . 1981; sensu Candusso i n
up to 3.5 m m w i d e , white to cream, pale p i n k - b r o w n (7.5 Y R 8/4) w i t h R i v . M i c o l . 33: 19-23. 1990; sensu C h r . L a n g e i n D o c . m y c o l . 25 (98-
H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic a n d

terrestrial i n h u m u s - r i c h , nutrient-rich sandy a n d l o a m y soils, i n
deciduous woods, i n the inner dune area o f H o l l a n d , but more wide-
spread i n southern L i m b u r g ; rare. Sept.-Oct. W i d e s p r e a d i n Europe,
and recorded f r o m many countries.
T h e name Leucoagaricus serenus has been used for several different

species (see under m i s a p p l i e d names). K u h n e r ' s interpretation o f the
species (in B u l l , trimest. S o c . m y c o l . F r . 52: 213-214. 1936) is f o l -
l o w e d here (see also V e l l i n g a i n P e r s o o n i a 17: 474-475. 2000).
Leucoagaricus serenus i n the present sense i s a white species, w h i c h
does not discolour o n d r y i n g , w i t h a d r y pileus covering, without a
really distinctly bulbous base, p r o v i d e d w i t h clavate c h e i l o c y s t i d i a
without o r w i t h some crystals o n the apex. A species w i t h narrow cys-
tidia w i t h crystals o n the apex is n o w distinguished as separate, a n d
described here under the name L. crystallifer. M o s e r (Rohrlinge
Blatterpilze, 5. A u f l . : 244. 1983) placed both species together under S.
F i g . 75. Leucoagaricus serenus.
serenus. B r e i t e n b a c h & K r a n z l i n ( P i l z e S c h w e i z 4: p i . 260, p. 2 2 2 .
1995) depicted the cheilocystidia o f S. serenus as fusiform-clavate, and
100): 251-253. 1995; Sericeomyces serenus sensu M . B o n , F l . m y c o l . often w i t h crystals, but very w i d e (up to 2 0 pm). T h e i r c o l l e c t i o n m a y
Eur. 3, Lepiotes: 85. 1993; sensu M i g l . & B i z z i i n M i c o l . ital. 24 (3): belong to a different species, close to L. crystallifer.
80-81. 1995; sensu R o d r i g u e z A r m a s e t a l . i n D o c . m y c o l . 18 (72): 67- T h e c h e i l o c y s t i d i a have a rather l o n g p e d i c e l , a n d are relatively
69. 1988 (in a l l cases L. crystallifer). thin-walled, w h i c h causes rupture o f the w a l l i n microscope slides.
S E L . D E S C R . & FIGS. - H u i j s m a n i n M e d e d . N e d . m y c o l . Vereen. 28:' Leucoagaricus erioderma ( M a l e n c . ) M . B o n , described f r o m
42-44. 1943 (as Lepiota serena); K e l d e r m a n , Parasolzw. Zuid*s M o r o c c o , has a distinctly squamulose pileus c o v e r i n g , a n d slightly
L i m b u r g : 144-145. 1994; K u h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 5 2 : narrower c h e i l o c y s t i d i a ( M a l e n c o n i n M a l e n c o n & Bertault, F l .
213-214. 1936 (as Lepiota serena); M a i . & Bert., F l . champ, sup. C h a m p , sup. M a r o c 1: 147-148, fig. 24. 1970, as Lepiota serena var.
M a r o c 1: 145-147, fig. 23. 1970 (as Lepiota serena); W u i l b a u t i n D o c . erioderma).
m y c o l . 17 (65): 44-45, figs. 5-8. 1986 (as Sericeomyces serenus). Leucoagaricus amylosporus (Malen$.) M . B o n may show a super-
V E R N . N A M E - Witte champignonparasol. ficial resemblance to L. serenus, but differs i n the differentiated hyphae
o f the pileus c o v e r i n g and the spores w h i c h are d i s c o l o u r i n g dark v i o -
P i l e u s 10-35 m m , w h e n y o u n g paraboloid, expanding to convex, let i n M e l z e r ' s Reagent ( M a l e n c o n i n M a l e n c o n & Bertault, F l .
plano-convex, o r applanate w i t h umbo, rather smooth a n d creamy, C h a m p , sup. M a r o c 1: 148-150, f i g . 25. 1970, as Lepiota amylospora).
cream-coloured, ochraceous cream to pale grey-brown at centre,
around centre r a d i a l l y f i b r i l l o s e to fibrillose-squamulose to slightly 20. Leucoagaricus crystallifer V e l l i n g a i n Persoonia 17: 475. 2000. -
sulcate i n outer marginal zone, shiny, pure white to pale cream, often F i g . 76.
w i t h white fme-flocculose edge. L a m e l l a e , L = 32-70,1 = ( 0 - ) l , m o d - M I S A P P L . - Leucoagaricus serenus sensu M . B o n i n D o c . m y c o l . 11
erately crowded, free, rounded near stipe, often remote f r o m stipe, (43): 52. 1981; sensu Candusso i n R i v . M i c o l . 33: 19-23. 1990; sensu
(sub)ventricose, up to 3.5 m m w i d e , white to pale cream w i t h age, w i t h Chr. L a n g e i n D o c . m y c o l . 25 (98-100): 251-253. 1995; Sericeomyces
white, rather even to flocculose edge. Stipe 25-85 X 1.5-3 m m , broad- serenus sensu M . B o n , F l . m y c o l . Eur. 3, L e p i o t e s : 8 5 . 1993; sensu
ening towards clavate base (up to 5 m m ) , h o l l o w , white to w h i t i s h M i g l . & B i z z i i n M i c o l . ital. 24 (3): 80-81. 1995; sensu R o d r i g u e z
lengthwise fibrillose over total length, i n lower half on slightly b r o w n - A r m a s et a l . i n D o c . m y c o l . 18 (72): 67-69. 1988.
ish background, sometimes finely hairy-pubescent or short-fibrillose, S E L . I C O N . - Candusso i n R i v . M i c o l . 33: 20. 1990.
i n general shiny. A n n u l u s ascending w i t h flaring margin, white cottony,
often rather loose around stipe. Context thin, white and shiny i n pileus,
white to pale cream and d u l l i n stipe. S m e l l o f cut basidiocarp like rub-
ber (Lepiota cristata), rarely slightly fruity, astringent o r indistinctly
fungoid. Taste not k n o w n . Spore print probably white.
Spores i n s i d e - v i e w 6.0-9.5 X 3.0-5.0 p m , o n average 6.7-7.9 X
3.8-4.2 m m , Q = 1.55-2.3, Q a v = 1.75-1.9, i n side-view a m y g d a l i -
form, more rarely w i t h rounded apex, some w i t h distinct apical p a p i l -
la, i n frontal view narrowly ovoid, dextrinoid, congophilous,
cyanophilous, metachromatic i n C r e s y l B l u e . B a s i d i a 16-29 X 6.5-9.5
pm, 4-spored, very rarely 2-spored. L a m e l l a edge heterogeneous to
x
totally sterile. C h e i l o c y s t i d i a 20-65 X 8.5-16 p m , clavate, rarely u t r i -
form-clavate, without crystals, rather thin-walled, w i t h greyish c o n -
tents, deep rooting. Pleurocystidia absent. Pileus covering a thin cutis
of narrow, c. 2.0-4.0 p m w i d e , c y l i n d r i c a l , colourless hyphae; hyphae
arranged i n adnate bundles and slightly ascending at the tips. H y p h a e
o f trama inflated, u p to 25 p m w i d e . S t i p i t i p e l l i s a cutis o f narrow
c y l i n d r i c a l , colourless hyphae, c. 2.0-5.0 p m w i d e , w i t h some irregu-
lar colourless hairs. Clamp-connections absent. F i g . 76. Leucoagaricus crystallifer.
S E L . D E S C R . & FIGS. - M i g l . & B i z z i i n M i c o l . ital. 24 (3): 80-81, fig. 21. Leucoagaricus menieri (Sacc.) S i n g , i n M y c o p a t h . M y c o l . appl.
5. 1995; R o d r i g u e z A r m a s et a l . i n D o c . m y c o l . 18 (72): 67-69. 1988. 34: 131. 1 9 6 8 . - F i g . 77.
Lepiota arenicola M e n i e r i n B u l l . S o c . m y c o l . Fr. 5: 174. 1890, non
P i l e u s 15-40 m m , campanulate, e x p a n d i n g to p l a n o - c o n v e x , a n d Lepiota arenicola Peck, 1888; Lepiota menieri S a c c , S y l l . F u n g . 9: 4.
applanate w i t h l o w u m b o , w i t h s l i g h t l y i n f l e x e d m a r g i n , w i t h cream 1891, n o n Lepiota menieri Q u e l . , 1896; Leucoagaricus arenicola
to pale ochre c o l o u r e d u m b o , m o r e intensely c o l o u r e d b y 2 days after (Menier) M . B o n i n B u l l , trimest. S o c . m y c o l . Fr. 86: 9 7 . 1970 (not
p i c k i n g , around u m b o w h i t e and r a d i a l l y short-fibrillose to s i l k y - f i b - valid); Leucoagaricus arenicola (Menier) B o n & Boiffard i n B u l l ,
rillose, sometimes slightly squamose, w i t h m a r g i n s l i g h t l y exceeding trimest. Soc. m y c o l . Fr. 88: 2 1 . 1 9 7 2 (not valid); Sericeomyces menieri
lamellae, rarely w i t h part o f annulus attached to it. L a m e l l a e , L = (Sacc.) C o n t u i n C r y p t o g . , M y c o l . 12: 4. 1991.
4 5 - 7 0 , 1 = 0-5, moderately c r o w d e d to v e r y c r o w d e d , free, and often E X C L . - Leucoagaricus menieri sensu S i n g , i n M y c o p a t h . M y c o l .
remote f r o m stipe, sometimes anastomosing, subventricose, 1-3.5 appl. 34: 131. 1968 ( = Leucoagaricus spec).
m m w i d e , cream, very pale beige, w i t h c o n c o l o r o u s to white f l o c c u - S E L . D E S C R . & FIGS. - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 142-
lose, rarely even, edge. Stipe 20-70 X 2.5-6 m m , broadening d o w n - 143. 1994; M e n i e r i n B u l l . S o c . m y c o l . Fr. 5: 174, p i . 18. 1890.
wards to often bulbous base (up to 8 m m ) , h o l l o w , w h i t e and shiny, V E R N . N A M E -Zandchampignonparasol.
lengthwise fibrillose, white-tomentose at basal part, b e c o m i n g y e l -
l o w o n d a m a g i n g . A n n u l u s ascending, sometimes w i t h a very distinct Pileus 20-60 m m , convex to applanate, without o r w i t h blunt w i d e
c u f f around stipe w i t h a straight l o w e r edge, and a fringed upper mar- umbo, when o l d w i t h uplifted, wavy margin, white to cream, somewhat
gin, and a narrow flaring part, white. Context white and d u l l i n pileus, ochraceous at centre, r a d i a l l y fine-fibrillose, satiny shiny; m a r g i n
w h i t e and shiny i n stipe. S m e l l indistinct, o f cut basidiocarp variable: fringed w i t h white fibrils. L a m e l l a e , L = c. 90-100, 1 = 1-5, very
f u n g o i d , unpleasant, astringent-fungoid, slightly rubberish (like crowded, crowded, free, segmentiform to slightly ventricose, 4-6 m m
Lepiota cristata). Taste i n d i s t i n c t o r absent. Spore print p r o b a b l y wide, white to cream-coloured w i t h white, fimbriate edge. Stipe 40-65
white. X 4-6 m m , widened towards base, and there abruptly and slightly mar-
Spores i n side-view 5.5-9.0 X 3.5-4.5(-5.0) um, o n average 6.3-7.7 ginately bulbous and up to 13 m m w i d e , h o l l o w , white, lengthwise
X 3.9-4.2 p m , Q = 1.45-2.1, Q a v = 1.65-1.9, v a r y i n g f r o m e l l i p s o i d innately fibrillose and shiny. A n n u l u s halfway or lower, ascending o r
to o b l o n g w i t h rounded apex to o b l o n g - a m y g d a l i f o r m without o r w i t h pendant, white. Context white and thick i n pileus, white shiny i n stipe,
apical p a p i l l a , o b l o n g - o v o i d , dextrinoid, congophilous, cyanophilous, slightly ochraceous i n base. S m e l l indistinct. Taste indistinct. Spore
metachromatic i n C r e s y l B l u e . B a s i d i a 17-28 X 6.5-8.5 u m , 4-spored. print whitish.
L a m e l l a edge sterile. C h e i l o c y s t i d i a i n tufts 23-51 X 5.5-13 p m , c y l i n - Spores 5.5-8.5(-9.0) X 4.0-5.0 m m , o n average 6.3-7.7 X 4.2-4.7
drical, cylindrical-fusiform, narrowly clavate, some w i t h a tendency to pm, Q = 1.3-2.0, Q a v = 1.4-1.85, i n side-view e l l i p s o i d - a m y g d a l i -
be utriform, colourless, w i t h (some without) s m a l l to b i g crystals o n f o r m or o b l o n g - a m y g d a l i f o r m w i t h acute apex, dextrinoid, c o n -
apex. Pleurocystidia not observed. Pileus covering a cutis w i t h bundles gophilous, cyanophilous, metachromatic i n C r e s y l B l u e , without germ
o f adnate r a d i a l l y arranged hyphae, 3.0-12 p m w i d e , w i t h adnate to pore. B a s i d i a 21-27 X 7.5-9.0 p m , 4-spored. L a m e l l a edge sterile.
slightly ascending terminal elements w i t h rounded apex, colourless, C h e i l o c y s t i d i a very variable, 20-70 X 6.0-18 p m , v a r y i n g f r o m clavate
rarely w i t h some pale b r o w n intracellular pigment. Stipitipellis a cutis to narrowly lageniform, misshapen, sometimes w i t h two apices, rather
of narrow, colourless, 2.0-4.0 p m w i d e c y l i n d r i c a l hyphae, rarely w i t h thin-walled and most w i t h s m a l l crystals o n the apex. P l e u r o c y s t i d i a
a loose c o v e r i n g o f wider, c y l i n d r i c a l a n d branched, colourless, absent. Pileus covering a cutis o f narrow c y l i n d r i c a l hyphae, 5.0-7.0
hyphae. Clamp-connections absent. p m wide, w i t h slightly w i d e r ascending terminal elements. C l a m p - c o n -
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic and nections absent.
terrestrial o n h u m u s - r i c h sandy and l o a m y soils, i n deciduous woods, H A B I T A T & DISTR. - Gregarious i n s m a l l groups, solitary, sapro-
i n the inner dune region, and i n southern L i m b u r g , rather rare. Sept.- trophic and terrestrial i n deciduous woods o n s o i l relatively p o o r i n
Oct. Widespread i n Europe.
Leucoagaricus crystallifer i s what several authors (see under misap-

p l i e d names) described as L. serenus. Leucoagaricus serenus itself has
broadly clavate a n d clavate c h e i l o c y s t i d i a without crystals. M o s e r
(Rohrlinge Blatterpilze, 5. A u f l . : 244. 1983) merged the two interpre-
tations, as he stated that 'sterile hairs are clavate o r c y l i n d r i c a l ' . T h e
w a l l s o f the c h e i l o c y s t i d i a i n L. crystallifer are sturdier than i n L.
serenus and stay intact i n microscope slides o f dried material.
A n o t h e r species w i t h crystals o n the c h e i l o c y s t i d i a is Sericeomyces
cinereopallidus C o n t u (in M i c o l . ital. 23 (2): 117. 1994) w i t h a pale
grey, v i s c i d pileus, and t h i c k - w a l l e d cystidia.
Leucoagaricus menieri (Sacc.) Sing., i s very close to L. crystallif-
er, but is i n general more sturdy, w i t h an abrupt bulbous base. T h e
cheilocystidia i n L. menieri are variably shaped, and often misshapen
i n the D u t c h material, w i t h s m a l l crystals.
Leucoagaricus subvolvatus ( M a i . & Bert.) M . B o n is another stur-
dy species, w i t h velar remnants at the bulbous base, and it has ellipsoid
spores without tendencies to amygdaliformity, and regularly shaped,
narrowly lageniform to c y l i n d r i c a l cheilocystidia, w i t h a tuft o f crys-
tals o n the apex. B o t h L. menieri and L. subvolvatus were described
f r o m sandy areas i n the mediterranean region. F i g . 77. Leucoagaricus menieri (habit d e l . P. K e l d e r m a n ) .
nutrients; i n the Netherlands o n l y k n o w n from a few places i n south- Sericeomyces singeri M . B o n ex C o n t u & Signorello (in Signorello
ern L i m b u r g . Sept.-Oct. K n o w n from sandy, dry habitats, along the & C o n t u i n M i c o l . Veget. medit. 13: 64. 1998) was described to a c c o m -
Mediterranean and southern A t l a n t i c coasts and from central Europe. modate Singer's interpretation o f an U r u g u a y a n collection o f L.
menieri, but a S a r d i n i a n gathering served as the type collection. T h i s
The above description is based on two collections from the same l o c a l - species was earlier i n v a l i d l y described as S. cylindrosporus C o n t u (in
ity i n the Netherlands. C r y p t o g . M y c o l . 12: 8. 1991), and is characterized by a slightly gelat-
Leucoagaricus menieri comes very close to L. crystallifer, and the i n i z e d pileus covering and subcylindrical spores. Whether it really is
differences between the two species are rather subtle; L. menieri being identical to Singer's c o l l e c t i o n s , as described i n M y c o p a t h . M y c o l .
the bigger, thicker-fleshed species, w i t h an abrupt bulbous base, w i t h appl. 34: 131. 1968, has still to be proven. Furthermore, Singer's spec-
lageniform cystidia. T h e cystidia i n the D u t c h material were atypical imens m a y represent L. menieri i n the European sense.
i n that they were misshapen and relatively thin-walled. D r i e d material A n o t h e r species i n this c o m p l e x is Leucoagaricus mairei M . B o n
of both L. crystallifer and L. menieri remains still cream-coloured, and (in D o c . m y c o l . 11 (43): 59. 1981), based o n M a i r e ' s interpretation o f
does not d i s c o l o u r b r o w n as i n L. sericifer. L. menieri. C h e i l o c y s t i d i a are absent i n this species.
Leucoagaricus subvolvatus ( M a i . & Bert.) M . B o n comes very close This w h o l e group o f species, w i t h a predominantly mediterranean
to the present concept o f L. menieri, but differs i n spores with rounded, distribution, is i n need o f revision.
not acute, apex, and regular, narrowly lageniform to cylindrical cheilo- Leucoagaricus menieri was described f r o m sandy habitats i n the
cystidia, w h i c h bear crystals o n the apex. A good description is given mediterranean region, and its occurrence i n southern L i m b u r g i n the
by M a l e n c o n & Bertault (in A c t a phytotax. bare. 8: 4 1 . 1971). Netherlands, is rather unexpected.
7. Lepiota (Pers.: Fr.) S.F. Gray
Agaricus sect. Lepiota Pers., Tent. Disp. meth. Fung.: 68. 1797; Agaricus tribus Lepiota Pers.: Fr., Syst. mycol. 1: 19.
1821; Lepiota (Pers.: Fr.) S.F. Gray, Nat. Arr. Br. PI. 1: 601. 1821. - Fusispora Fay. in Ann. Sc. nat., Bot. Ser. VII, 9:
351. 1889. - Lepiotula (Maire) Locq. ex Horak, Syn. Gen. Agar.: 337. 1968. - Echinoderma (Locq. ex M . Bon) M .
Bon in Doc. mycol. 21 (82): 61. 1991.
S E L E C T E D LITERATURE - M . Bon in Doc. mycol. 11 (43): 27-50. 1981; M . Bon, Fl. mycol. Eur. 3, Lepiotes: 44-82.
1993; Candusso & Lanzoni, Lepiota: 115-322. 1990; Kelderman, Parasolzw. Zuid-Limburg: 43-139. 1994; Knudsen
in Bot. Tidsskr. 75: 121-155. 1980 (sect. Echinatae); Vellinga & Huijser in Persoonia 15: 223-240. 1993 (species with
spurred spores); Vellinga & Huijser in Belg. J. Bot. 131: 191-210. ('1998') 1999 (species with hymeniform pileus
covering).
Basidiocarp pluteoid with persistent velum universale; pileus surface in most species squamulose; lamellae free; annu-
lus or annular zone, remnants of velum partiale, often present in mature specimens; spore print white to cream.
Spores smooth, in a few species, especially with spurred spores, minutely rough, hyaline, dextrinoid, rarely not dex-
trinoid, congophilous, cyanophilous, not metachromatic in Cresyl Blue, or with pink inner wall, with walls slightly or
not swelling in ammonia ( N H ( p and acetic acid (CH COOH( ), binucleate, in a few species uninucleate; cheilo-
3 aq 3 ag
cystidia present in most species; pleurocystidia not present in Netherlands' species; hymenophoral trama regular;
clamp-connections present in most species. Development bivelangiocarpic and pileostipitocarpic or stipitocarpic. -
Typus conservandus Agaricus colubrinus Pers. (= Lepiota clypeolaria (Bull.: Fr.) Kumm.).
HABITAT & DISTRIBUTION - Saprotrophic and terrestrial, often in nutrient-rich areas, in woods and in grasslands, cos-
mopolitan, but absent from arctic and high alpine regions.
The genus Lepiota differs from Cystolepiota Sing, in the structure of the velum universale; in Cystolepiota the universal
veil consists of loosely arranged globose or inflated elements; in most Lepiota-species the velum is made up of long,
cylindrical or clavate, erect or ascending elements; in one section globose elements are present, but these are connect-
ed with hyphae, with a gradual change from globose to longitudinal elements, and these chains are agglutinated into
pyramidal squamules. Whether the differences really warrant recognition of two genera has to be proved by further
research.
The genus Macrolepiota Sing, is not included within Lepiota, as proposed by Ballero & Contu (in Candollea 46:
480-481. 1991), because of the fundamental differences between the two genera in hymenophoral trama, spore wall
composition, and structure of the velum universale.
Lepiota lignicola P. Karst. is excluded from the genus Lepiota, and accepted, as Leucopholiota decorosa (Peck)
O.K. Miller et al. (in Mycologia 88: 138. 1996), within the family Tricholomataceae.
A simple infrageneric classification, in which six sections are recognized, is applied here, pending further research
on molecular and additional morphological characters. Candusso & Lanzoni (Lepiota: 738-740. 1990) gave the same
six sections, whereas Bon (Fl. mycol. Eur. 3, Lepiotes: 44-82.1993) gave a much more extensive division of the genus.
Many Lepiota-species, especially the group of species with ellipsoid spores and a pileus covering made up of elon-
gate elements only, contain amanitin, and are highly toxic, causing severe liver damage which can result in death.
The neutral term 'pileus covering' is used for all the covering layers of the pileus, not taking into account their ori-
gin (whether they are metableme, archibleme, teleobleme or pellis in the terminology of Clemencon, Anat.
Hymenomyc: 802-804. 1997).
The size and shape of the elements of the pileus covering have been observed just outside the central region or the
'calotte'.
109
1. Spores spurred, or with straight base (Fig. 78a) Key one

1. Spores not spurred, but ellipsoid, oblong, fusiform-amygdaliform or shaped as some Boletus-spores (with straight
abaxial side, convex adaxial side and suprahilar depression)
2. Spores shaped as some Boletus-spores (Fig. 78b) Key two
2. Spores either fusiform-amygdaliform, or (broadly) ellipsoid to oblong
3. Spores fusiform-amygdaliform with convex abaxial and convex adaxial sides (Fig. 78c) Key three
3. Spores ellipsoid, oblong, broadly ellipsoid (Fig. 78d)
4. Pileus surface smooth, closed or breaking open into smooth patches; covering made up of clavate to cylin-
drical elements, tightly packed, forming a hymeniderm (Fig. 79a) Key four
4. Pileus surface squamulose; pileus covering made up either of long elements, or of inflated to rounded ele-
ments in chains
5. Pileus set with acute, pyramidal squamules, made up of agglutinated chains of elements, from cylindri-
cal to inflated elements at base of squamules, gradually changing into ellipsoid to globose terminal ele-
ments (Fig. 79b) Key five
5. Pileus set with squamules, made up of elongate, cylindrical elements, without or with septa, which are
provided with or without clamp-connections (Fig. 79c, two types) Key six
v..
LEPIOTA (Vellinga) 111
ALTERNATIVE K E Y
1. Pileus surface set with acute, pyramidal to recurved squamules, made up of agglutinated chains of elements, from
cylindrical to inflated elements at base of squamules, gradually changing into ellipsoid to globose terminal elements
(Fig. 79b) Key five
1. Pileus surface smooth, closed or with patches and pileus covering a hymeniderm, or squamulose and breaking up
into squamules, made up of elongate elements
2. Pileus smooth and closed or breaking up into patches; pileus covering a hymeniderm (Fig. 79a) . . . . Key four
2. Pileus with squamules, made up of elongate, erect or adnate to ascending elements (Fig. 79c)
3. Spores ellipsoid, more rarely broadly ellipsoid or oblong, or fusiform to amygdaliform (i.e. with abaxial and
adaxial side convex)
4. Spores amygdaliform to fusiform (Fig. 78c) Key three
4. Spores ellipsoid to oblong, rarely broadly ellipsoid (Fig. 78d) Key six
3. Spores with broadened base, triangular in outline, spurred, or with straight abaxial and convex adaxial side and
suprahilar depression
5. Spores triangular to spurred (Fig. 78a) Key one
5. Spores shaped as some Boletus-spores (with straight abaxial and convex adaxial side, and suprahilar depres-
sion; 'penguin-shaped', Fig. 79b) Key two
K E Y ONE
Species with spurred spores
N.B. Spore width measured in middle of spores, in side-view
1. Pileus covering a hymeniderm
2. Smell fruity-sweet and rubber-like 28. L . cristata
2. Smell saponaceous Lepiota saponella
Bodin & Priou in in Doc. mycol. 23 (92): 62. 1994. Described from
southwestern France, not yet recorded from the Netherlands
1. Pileus covering made up adnate, ascending or erect long and slender elements
3. Pileus covering made up of adnate and ascending, articulate hyphae (Fig. 79c), with clamp-connections at the
septa, with intracellular brown pigment, very rarely lacking pigment 21. L . boudieri
3. Pileus covering made up of adnate to erect elements, unicellular or with septa without clamp-connections, with-
out or with pigment, which can be intracellular or parietal
4. Basidiocarp white to ochraceous cream 22. L . subalba
4. Basidiocarp grey-brown, brown, dark green, greenish etc., not white
5. Pileus predominantly with brown, orange-brown, red-brown or purplish brown colours
6. Pileus at centre (when young) dark red-brown, red-brown, orange-brown; context in stipe, and often also
pileus and stipe surfaces discolouring orange; pileus covering made up of erect, (non-)septate elements
and a layer of flexuous repent coloured hyphae 19. L . castanea
6. Pileus at centre dark brown, dark grey-brown; context in stipe not turning orange; pileus covering made
up of more or less erect elements only; a layer of repent coloured hyphae not present
7. Distinct girdles or bands of squamules present in lower half of stipe; pileus surface splitting up into
concentrical zones around centre; pileus (20-)30-55(-80) mm 20. L . cingulum
7. Distinct girdles or bands on stipe lacking, some scattered squamules on stipe often present; pileus sur-
face around centre irregularly breaking open; pileus 10-30(-40) mm 23. L . tomentella
5. Pileus predominantly with grey or (grey-)green colours (though context may discolour orange)
8. Pileus at centre green-black to blue-green, or blue-brown, with distinct squamules around centre; back-
ground turning orange; elements of pileus covering normally with up to 3 clamp-less septa, with intra-
cellular pigment, which is soluble in ammonia 27. L . grangei
8. Pileus at centre pinkish brown to dark grey (with or without greenish tinges), with granular, plush-like or
more clearly delimited squamules on background which turns orange or not; elements of pileus covering
without or with up to 2 clamp-less septa, with predominantly parietal pigment
9. Pileus background not turning orange with age or damaging
10. Pileus when young with pinkish brown or greyish brown tinges; elements of pileus covering with
up to 2 (-3), clamp-less septa 23. L . tomentella
10. Pileus when young very dark, often with olive or lilacinous tinges; elements of pileus covering nor-
mally without septa, sometimes in lower part l(-2) septate 25. L . griseovirens
9. Pileus background turning (slowly) orange with age or on damaging
11. Spores short (avl = 6.6-7.4 pm); elements of pileus covering short ((45-)70-180(-200) pm long)
and often irregular 26. L . poliochloodes
11. Spores long (avl = 7.0-9.6 pm); elements of pileus covering long ((55-)85-320(-400) pm long) and
straight
12. Pileus only when very young dark grey or olive, soon dull orange brown, dark brown or ochra-
ceous brown at centre, with granulate, rather thin-layered squamules around centre; cheilocys-
tidia broad (5.0-15(-18) pm wide) 24. L . pilodes
12. Pileus dark grey, black greyish olive-brown at centre, to pale grey-brown at margin, with wart-
like squamules around centre; cheilocystidia narrow (5.0-10.5 pm wide)
25. L . griseovirens
K E Y TWO
Species with 'penguin-shaped' spores (as in some Boletus-species, with straight abaxial side, and suprahilar depression)
1. Pileus covered in acute squamules (with terminal elements globose to ellipsoid)
2. Lamella edge dark; cheilocystidia with dark contents 39. L. hystrix
2. Lamella edge white or concolorous with sides; cheilocystidia not coloured
3. Basidiocarps big (pileus up to 115(-150) mm), with big hanging annulus, with squamules on underside; spores
6.0-9.0(-9.5) X 2.5-3.5(-4.0) um; cheilocystidia clavate to spheropedunculate 37. L . aspera
3. Basidiocarps medium (pileus up to 55 mm; stipe with annular zone, without hanging annulus; spores 5.0-6.5
X 2.5-3.0 um; cheilocystidia fusiform, narrowly clavate, cylindrical 38. L . perplexa
1. Pileus surface breaking up into smooth patches or into woolly, or felted squamules, which might be squarrose at cen-
tre; annulus absent, if present membranaceous, or only as a woolly zone on stipe
4. Pileus surface smooth, breaking up into patches around a distinct 'calotte'; stipe glabrous, devoid of squamules;
pileus surface made up of tightly packed narrowly clavate elements 28. L . cristata
4. Pileus surface squamulose to ragged; stipe woolly or tomentose in lower part; squamules made up of long, hairy
elements
5. Stipe woolly in lower part; spores (12.0-)13.5-25.0 X 4.0-6.0 pm 2. L . magnispora
5. Stipe glabrous to fibrillose, tomentose in lower part; spores (6.5-)7.0-9.0(-10.0) X (2.5-)3.0-3.5('-4.0) um
5. L. cortinarius
K E Y THREE
Species with fusiform to amygdaliform spores and a pileus covering made up of long elements
N.B. Spore length measured without hilar appendage
1. Spores 'penguin-shaped' with straight abaxial side, convex adaxial side and suprahilar depression (Fig. 78b)
2. Spores (12.0-)13.5-25.0 X 4.0-6.0 pm 2. L . magnispora
2. Spores (6.5-)7.0-9.0(-10.0) X (2.5-)3.0-3.5(-4.0) um 5. L. cortinarius
1. Spores fusiform-amygdaliform, with convex abaxial and adaxial sides (Fig. 78c)
3. Pileus without discrete squamules, white to (pale) brown at centre
4. Pileus white to pale brown at centre (colours in the range of Mu. 10 Y R ) ; spores 10.0-21.0(-23.0) X 5.0-7.0(-
8.0) pm, avw = 5.5-6.1 um 3. L . erminea
4. Pileus pale brown to brown at centre (colours in the range of Mu. 7.5 Y R ) ; spores on average narrower than in
L. erminea: 10.0-16.5 X 4.5-6.5 pm, avw — 4.8-5.5 pm 4. L . oreadiformis
3. Pileus with brown to pale brown squamules contrasting with background
5. Stipe in lower part discolouring orange with age and when touched; basidiocarps relatively big: pileus 50-90
mm; stipe 70-105 X 10-15 mm; spores 8.5-11.0(-12.5) X 5.0-6.0(-6.5) pm, Qav = 1.75-1.95
1
1. L . ignivolvata
5. Stipe not discolouring orange; basidiocarps small to big: pileus 20-100 mm; stipe 40-100 X 2-10 mm; spores
9.0-18.5 X 4.0-6.0(-6.5) pm, Qav = 1.9-2.85
6. Spores 11.0-18.5 X (4.0-)4.5-6.0(-6.5) pm, Qav = 2.4-2.85; stipe with annular zone, and lanate covering
below 1. L . clypeolaria
6. Spores 9.0-13.5 X 4.0-6.0(-6.5) pm, Qav = 1.9-2.1; stipe often with a lanate annular zone and in lower part
a brown ring, especially when young, with bands or squamules in lower part 6. L . subgracilis
K E Y FOUR
Species with a hymeniform pileus covering
1. Pileus surface smooth and glabrous, viscid when moist; pleurocystidia present, similar to cheilocystidia, conspicu-
ous Chamaemyces fracidus
1. Pileus surface dry, smooth or breaking up into patches; pleurocystidia absent; cheilocystidia present or absent
2. Basidiocarps with bright orange-red pileus and stipe; surface of pileus not broken up into squamules or patches
34. L . pyrochroa
2. Basidiocarps with whitish cream to dark brown or almost black colours on pileus, with whitish to reddish or lila-
cinous stipe; surface of pileus broken up into squamules or not (if surface closed, colour of pileus cream, pale brown)
3. Spores in.side-view spurred, triangular, with a distinct sac at base, or 'penguin-shaped' 28. L . cristata
3. Spores ellipsoid in side-view
4. Lamellae separated from stipe by a kind of collarium; basidiocarps big: pileus 25-70(-100) mm and orange-
brown to leather-brown; smell soapy-perfumy; spores dextrinoid, not or slightly metachromatic in Cresyl
Blue 33. L . ochraceofulva
4. Collarium-like structure absent; pileus in most species smaller, and with other colours; spores in most
species not distinctly dextrinoid, but metachromatic in Cresyl Blue, sometimes weakly dextrinoid and
weakly metachromatic
5. Annulus conspicuous with broadened rim, with coloured patches on it; pileus surface broken into patch-
es leaving a discrete disc at centre; colour of pileus varying from very pale brown to almost black, when
young often with lilacinous tinges 32. L . lilacea
5. Annulus, if present, membranaceous and/or fugacious, without coloured patches, or absent; pileus sur-
face broken up or not; colour either dark chocolate-brown, orange-brown or pale brown, creamy, not lila-
cinous when young
6. Cheilocystidia absent
7. Pileus surface breaking up into patches; remnants of partial veil adhering to pileus margin; spores
dextrinoid, but not strongly so, sometimes only after 10 minutes visible, weakly metachromatic in
Cresyl Blue, 4.5-6.5 X 3.0-4.0 um 29. L . apatelia
7. Pileus surface not breaking up or breaking up into small patches; ring ascending on stipe; spores
non-dextrinoid, but metachromatic in Cresyl Blue, 3.5-5.0(-5.5) X 2.5-3.5 um
30. L . cristatoides
6. Cheilocystidia present
8. Annulus present 35. L . psalion
8. Annulus absent
9. Pileus surface broken up into dark brown patches; some brown patches present at base of stipe;
spores 4.0-6.0(-7.0) X 2.5-3.5 um, Qav = 1.6-1.7 31. L . hymenoderma
9. Pileus surface creamy, pale brown or ochraceous, closed or broken up into patches
10. Pileus surface not breaking up into squamules or patches; spores 3.0-4.5 X 2.5-3.0(-3.5) pm
36. L . rufipes
10. Pileus surface squamulose, made up of clavate elements, giving rise to globose terminal ele-
ments; spores 6.0-8.0 X 3.5-4.5 pm L . cystophoroides
Joss. & L . Riousset in Bull. mens. Soc. linn. Lyon 41: 317. 1972. Described from south-
ern France, also recorded from northern France, not yet recorded from the Netherlands
K E Y FIVE
Species with acute squamules on pileus, made up agglutinated chains of elements, with ellipsoid to globose terminal
elements
1. Annulus membranaceous, hanging, provided with (brown) squamules; lamella edge white or concolorous with sides;
spores 6.0-9.0(-9.5) X 2.5-3.5(-4.0) pm, Q = 1.9-3.3, Qav = 2.2-2.7; basidiocarps big: pileus 50-115(-150) mm;
stipe 50-120 X 9-22 mm 37. L . aspera
1. Membranaceous (hanging) annulus absent (if such annulus present, clamp-connections absent); lamella edge white,
concolorous with sides or dark; spores 3.5-7.0 X 2.0-3.5 pm, Q = 1.35-2.6, Qav < 2.4; basidiocarps small to medi-
um: pileus up to 85 mm; stipe up to 90 X 14 mm
2. Lamella edge dark coloured
3. Pileus, especially at centre, densely set with small, blackish, squamules; spores 5.5-7.0 X 2 . 5 - 3 . 0 pm, Q = 2 . 0 -
2.6, Qav = 2.3; pigment in pileus elements, both intracellular and soluble in ammonia, and parietal
39. L . hystrix
3. Pileus surface strongly fibrillose-squamose, with long, up to 5 mm, brown to dark brown squamules; spores
4 . 0 - 6 . 0 X 2.0-3.5 pm, Q = 1.55-2.15, Qav = 1.6-1.85; pigment in pileus elements only parietal
40. L . calcicola
2. Lamella edge white or concolorous with sides
4. Membranaceous hanging annulus present; clamp-connections absent from all tissues 41. L . efibulis
4. Membranaceous and hanging annulus absent; an annular zone might be present; clamp-connections present in
all tissues (easily be seen at cylindrical elements on pileus)
5. Spores non-dextrinoid
6. Squamules on pileus made up of only one type of elements, viz. globose elements; terminal globose ele-
ments easily detached; cheilocystidia narrowly clavate to obovate, usually with cylindrical, moniliform
or branched, excrescence at apex • Cystolepiota moelleri
6. Squamules on pileus made up of globose terminal elements in chains, and coloured cylindrical elements;
terminal elements stuck to the chains; cheilocystidia narrowly clavate to clavate, without excrescence
46. L . boertmannii
5. Spores dextrinoid
7. Cheilocystidia absent or only some inconspicuous, basidiole-like structures present
8. Stipe more or less twice as long as pileus width (ratio of pileus width and stipe length 0 . 5 - 0 . 6 )
: 45. L . pseudoasperula
8. Stipe more or less as long as pileus width/ratio of pileus width and stipe length 0 . 8 - 1 . 0 )
9. Terminal elements of squamules on pileus 10-25(-45) X 10-21(-35) pm 44. L . carinii
9. Terminal elements of squamules on pileus 15-65 X 10-50 pm
10. Basidiocarp, though set with dark brown squamules, orange-brown; spines on stipe isolated,
not covered in wool, or absent 42. L . echinacea
10. Colours of basidiocarp sombre; spines of stipe covered in wool 43. L . jacobi
7. Cheilocystidia present and distinctly different from basidia and basidioles
11. Spores 5.0-6.5 X 2 . 5 - 3 . 0 pm, Q = 1.8-2.6, Qav = 2 . 0 5 - 2 . 3 5 , 'penguin-shaped' . . 38. L. perplexa
11. Spores 3 . 5 - 6 . 0 ( - 6 . 5 ) X 2.0-3.5 um, Q = 1.35-2.15, Qav = 1.6-1.85, ellipsoid
12. Apical elements of squamules on pileus ellipsoid to oblong 40. L . calcicola
12. Apical elements of squamules on pileus globose to ellipsoid
13. Apical elements of squamules on pileus ranging from small to big: 15-62 X 10-50 pm
42. L . echinacea
13. Apical elements of squamules on pileus small: 10-25(-45) X 10-21(-35) pm
44. L . carinii
K E Y SIX
Species with a pileus covering made up of long (erect or adnate) elements; spores ellipsoid to oblong
1. Pileus covering made up of only repent, septate or articulate elements with ascending terminal elements
2. Clamp-connections absent; pileus covering made up of long repent hyphae 18. L . fuscovinacea
2. Clamp-connections present
3. Pileus with brown colours; annulus absent, or only present as an annular zone on stipe; cheilocystidia present;
elements of pileus covering brown and thick-walled (walls up to 2 pm thick); in greenhouses; 16. L . rubella
3. Pileus white; annulus present; cheilocystidia absent; elements of pileus thin-walled; in woods
17. L . parvannulata
1. Pileus covering made up of often tufted, erect elements, occasionally septate; repent coloured hyphae may be pres-
ent as well / '
4. Only one type of elements, usually long erect elements, present in pileus covering
5. Basidiocarps with sulphur-yellow or olivaceous yellow tinges, especially in lamellae, context arid stipe; in hot-
houses . . . 15. L . elaiophylla
5. Basidiocarps with white, or pinkish to orange-brown colours; in natural habitats
6. Basidiocarps white; annulus present; cheilocystidia absent; hairs of pileus covering made up of long
clamped elements 17. L . parvannulata
6. Basidiocarps coloured; annulus only as woolly zone present; cheilocystidia present; hairs of pileus cover-
ing made up of non-septate elements
7. Elements of pileus covering long (70-400 pm long); spores (5.5-)6.0-7.5(-8.0) X 3.0-4.0(-4.5) pm
14. L . subincarnata
7. Elements of pileus covering relatively short (60-110(-140) pm); spores (7.0-)7.5-9.0(-10) X (4.5)5.0-5.5
pm L . helveola Bres.,
Fungi trident. 1:15. 1882. Recorded from Italy, France, and Germany,
but not yet known from the Netherlands
4. Pileus covering made up of long elements and short, clavate, narrowly clavate to cylindrical elements
8. Lamellae with distinct yellow tinges
9. Pileus covering made up of long elements, with short, clavate elements in between; cheilocystidia mostly
narrowly lageniform to narrowly uniform; in woods 11. L . xanthophylla
9. Pileus covering made up of long elements with occasionally short cylindrical elements in between; cheilo-
cystidia clavate; in hothouses 15. L . elaiophylla
8. Lamellae white to cream-coloured, occasionally pale grey with yellowish tinge; distinct yellow colours lack-
ing in basidiocarps
10. Basidiocarps with distinct olivaceous (brown) tinges 10. L. forquignonii
10. Basidiocarps either very dark, almost black, or pink to brown coloured, lacking olivaceous tinges
11. Annulus present, mostly as a distinct cuff around the stipe, with little squamulose patches on it
12. Basidiocarp relatively big and sturdy (pileus 30-75 mm; stipe 20-60(-110) X (5-)8-10(-15) mm),
growing in open sand on sanddunes; annulus thin; spores 9.0-11(-13) X 5.0-6.0 pm
L. brunneolilacea
Bon & Boiff. in Bull, trimest. Soc. mycol. Fr. 88: 18.1972. Recorded from the Atlantic and
Mediterranean coasts growing on white dunes amid Ammophila arenaria, from mid France south-
wards. To be expected in the dunes in the Netherlands.
12. Basidiocarps medium (pileus 15-45 mm; stipe 20-60 X 2-9 mm), growing in woods and copses;
annulus a distinct cuff set with little dark squamules; spores 6.0-10.0 X (3.5-)4.0-5.0 pm
13. Pileus covering broken up in discrete, dark patches around the dark calotte, made up of long
elements and a very dense layer of dark-pigmented clavate elements; repent coloured hyphae
on pileus surface absent 8. L . felina
13. Pileus covering squamulose, but lacking a distinct calotte and discrete squamules, varying in
colours from dark brown to pinkish brown; pileus covering made up of long elements and a loose
layer of narrowly clavate elements; repent coloured hyphae (mostly) present on pileus surface
9. L . pseudolilacea
11. Annulus absent; fugacious or woolly annular zone and bands of squamules on stipe might be present
14. Basidiocarps robust, with a relatively thick (5-9 mm wide) stipe, set with bands of squamules on
pinkish to brownish vinaceous background; cheilocystidia often septate; spores 7.0-10.0(-l 1.0) X
4.0-5.0(-6.0) pm, Qav — 1.7-1.9 3. L . brunneoincarnata
14. Basidiocarps slender, with a thin stipe (1-4 mm wide), with some incomplete bands or girdles of
small squamules; cheilocystidia not or rarely septate; spores either 4.5-7.5 X 3.0-4.5 pm with Qav
= 1.55-2.0 or 7.5-10.0 X 3.5-5.0 pm, Qav = 2.2
15. Cheilocystidia (narrowly) lageniform to (narrowly) utriform ('mummy-shaped'); spores 4.5-
6.5 X 3.0-4.0 pm; elements of pileus elements relatively long: 185-300 pm 12. L . echinella
15. A l l other combinations of characters: see discussion under L. echinella
Sect. Lepiota
Spores f u s i f o r m - a m y g d a l i f o r m w i t h convex abaxial and convex adax-
i a l sides, or w i t h a straight abaxial side, i n combination w i t h a pileus
covering made up o f l o n g c y l i n d r i c a l elements, often intermixed w i t h
short elements.
N . B . Spore length measured without hilar appendage.
1. Lepiota clypeolaria ( B u l l : Fr.) K u m m , Fiihr. P i l z k . : 137. 1871. -

F i g . 80.
Agaricus clypeolarius Bull., H e r b . F r a n c e : p i . 4 0 5 . 1789;
Agaricus clypeolarius B u l l . : Fr., Syst. m y c o l . 1: 21. 1 8 2 1 . - A g a r i c u s
colubrinus Pers., S y n . m e t h . F u n g . : 2 5 8 . 1801; Lepiota colubrina
(Pers.) S.F. G r a y , N a t . A r r . B r . P I . 1: 6 0 1 . 1821. -Lepiota ochraceo-
sulfurescens L o c q . ex M . B o n i n D o c . m y c o l . 16 (61): 4 6 . 1985. -
Lepiota clypeolaria var. minor J . L a n g e , F l . agar. dan. 1: 2 8 . 1935
(not v a l i d ) .
E X C L . - Agaricus clypeolarius sensu Sow., C o l . 111. E n g l . F u n g i 1:
p i . 14. 1796 ( = L. cristata); sensu Sow., C o l . 111. E n g l . F u n g i 2: p i . 171.
1798 ( = Leucocoprinus brebissonii); Lepiota clypeolaria sensu
B r e b i n a u d i n B u l l , trimest. S o c . m y c o l . Fr. 4 7 : 90. 1931 ( = Lepiota
s p e c ) ; Lepiota clypeolaria sensu R e a , B r i t . B a s i d i o m y c : 69. 1922 ( =
L. magnispora); Lepiota clypeolaria sensu Johnson & V i l g a l y s i n
M y c o l o g i a 90: 974-976. 1998 ( = L. magnispora). -Agaricus colubrh
nus sensu Krombh., N a t u r g . A b b . Beschr. essb. schadl. verd.
S c h w a m m e : p i . 1, figs. 10 & 11. 1831 ( = Macrolepiota spec);
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 220. 1995;
Candusso & L a n z o n i , L e p i o t a : p i . 15. 1990; D a h n c k e & Dahncke, 700
P i l z e : 313. 1979; J . L a n g e , F l . agar. dan. 1: p i . 1 1 D , H E . 1935 (as L.
clypeolaria and L. clypeolaria var. minor resp.).
S E L . D E S C R . & F I G S . - H o r a k , S y n . G e n . Agar.: 336-337. 1968 (as L.
colubrina); H . V S m i t h i n L l o y d i a 17: 314-315. 1954.
VERN. NAME - Bosparasolzwam.
Pileus 25-100 m m , when y o u n g campanulate, c o n i c a l w i t h small umbo,

expanding to plano-convex w i t h l o w broad umbo, at centre felted to
glabrous and pale b r o w n , pale/orange-like b r o w n ( M u . 10 Y R 6/3-8),
rarely more r e d (7.5 Y R 5-7/6)xOr pale buff, around centre w i t h c o n -
centrical zones o f more or less uplifted, discrete squamules more or less
w o o l l y , and concolorous to paler ochraceous b r o w n than disc, towards
margin more fibrillose, o n pale cream to white background, at margin
without or more rarely w i t h sulphur-yellow tinges, at outermost margin
w h e n y o u n g w i t h whitish floccose velar remnants, sometimes radially
sulcate at m a r g i n to 10 m m from margin. L a m e l l a e , L = 50-60,1 = 1-
3, moderately to very crowded, or moderately distant, free, not or slight-
l y remote from stipe, rarely anastomosing, subventricose or ventricose,
3-6 m m wide, white, cream or pale flesh-coloured cream, w i t h c o n c o l -
orous or white finely flocculose edge. Stipe 47-100 X 3-10 m m , taper-
ing upwards or cylindrical, w i t h bulbous base, fistulose, white to pale
cream-coloured, at apical part more or less glabrous or finely fibrillose-
striate, without real annulus, but w i t h annulus-like zone o f lanate or fib-
rillose squamules, below w i t h irregular bands o f lanate squamules or
w i t h incomplete girdles; squamules beige to pale brownish; not or y e l -
l o w i n g w h e n scratched, w i t h white m y c e l i u m cords at base. Context
white or pale cream and d u l l i n pileus, and white or y e l l o w and shiny
i n stipe. S m e l l strong, l i k e the rubber component o f the s m e l l o f L.
cristata, unpleasant, or l i k e the leaves o f Geranium robertianum, or
indistinct. Taste m i l d , or like smell and unpleasant. Spore print ' w h i t e ' .
Spores i n side-view 11.0-18.5 X (4.0-)4.5-6.0(-6.5) p m , Q = 2 . 1 -
3.4, Q a v = 2.4-2.8, i n side-view more or less a m y g d a l i f o r m w i t h c o n -
vex sides, some w i t h p a p i l l a - l i k e apex, w i t h or without suprahilar
depression, i n frontal v i e w more or less fusiform, t h i c k - w a l l e d , w i t h F i g . 80. Lepiota clypeolaria.
r
guttule or granular contents, dextrinoid, congophilous; not metachro- Lepiota clypeolaria var. pantherina G i l l e t , H y m e n o m y c e t e s : 62. 1874.
matic i n C r e s y l B l u e . B a s i d i a 26-43 X 8.0-13 u m , clavate, 4-spored, -Lepiota clypeolaria var. campanetta B a r l a i n B u l l . S o c . m y c o l . Fr. 2 :
rarely intermixed 4- and 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 116. 1886. - Lepiota ventriosospora var. fulva M . Bon in Bon &
12-33 X 5.0-15.5 p m , narrowly clavate, clavate, also c y l i n d r i c a l , Chevassut i n D o c . m y c o l . 4 (15): 2 7 . 1974. - Lepiota clypeolaria f.
broadly clavate or (sub)utriform, t h i n - w a l l e d and colourless. Pileus umbrinisquamosa H o n g o i n M e m . F a c . E d u c . S h i g a U n i v . Nat. S c i . 20:
covering made up o f erect elements, 150-c. 4 0 0 X 7.0-13 p m , mostly 51. 1970; Lepiota clypeolaria var. umbrinosquamosa (Hongo) M . B o n
longer than 2 0 0 p m , w i t h rounded apex, not o r very rarely septate, in D o c . m y c o l . 22 (88): 29. 1993. - Lepiota ventriosospora var. umbri-
slightly t h i c k - w a l l e d , c u r v e d i n l o w e r part, w i t h short elements i n norufescens M . B o n i n D o c . m y c o l . 22 (88): 46. 1993.
between, 15-60 X 6.0-10 p m ; pigment pale b r o w n and parietal. M I S A P P L . - Lepiota metulaespora sensu Bres., Iconogr. m y c o l . 1:
Stipitipellis a cutis made u p o f c y l i n d r i c a l , 4.0-15 p m w i d e hyphae, p i . 30. 1927; sensu Ktthn. & R o m a g n . , F l . anal. C h a m p , sup.: 4 0 2 .
w i t h y e l l o w , sometimes encrusted, parietal pigment. V e l u m partiale 1953. - Lepiota clypeolaria sensu R e a , B r i t . B a s i d i o m y c : 69. 1922.
made up o f short, often c. 2 0 p m long, 4.0-8.0 p m wide, c y l i n d r i c a l , S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 235. 1995;
colourless hyphae, w h i c h are often curved. B e l o w annular zone w i t h D a h n c k e & D a h n c k e , 7 0 0 P i l z e : 3 1 5 . 1979; R . P h i l l i p s , Paddest.
squamules, similar to those o n pileus. Clamp-connections present. S c h i m m . : 28. 1 9 8 1 ; R y m a n & H o l m a s e n , Svampar: 4 1 5 . 1984;
H A B I T A T & D I S T R . - Solitary o r i n s m a l l groups, saprotrophic and V e l l i n g a & Huijser i n C o o l i a 40: p i . 4. 1997.
terrestrial, m o s t l y i n beech-forests (Fagus) o n r i c h , often calcareous S E L . D E S C R . & F I G S . - B o n & Chevassut i n D o c . m y c o l . 4 (15): 27.
soils. Rare i n the Netherlands. Aug.-Oct., beginning of Nov. 1974 (as L. ventriosospora war. fulva); D . R e i d i n Trans. B r . m y c o l .
Widespread i n E u r o p e from the lowlands u p into the montane zone, Soc. 4 1 : 427-429, fig. 9, p i . 24 fig. 3. 1958.
and occurring also i n N o r t h A m e r i c a and A s i a . V E R N . N A M E - Geelbruine wolsteelparasolzwam.
Lepiota clypeolaria is often confused w i t h L. magnispora i n the field. Pileus 20-55 m m , hemispherical o r c o n i c a l w i t h obtuse apex w h e n
Spore shape i s the decisive character to distinguish between the two: young, expanding to broadly c o n i c a l or plano-convex w i t h broad h i g h
w i t h a curved abaxial side i n L. clypeolaria, and a straight abaxial side umbo, at centre felted and b r o w n ( M u . 7.5 Y R 4/8, 5/4, 7 . 5 - 1 0 Y R
i n L. magnispora. 5/6), around centre w i t h concentrical felted patches to fibrillose squa-
Lepiota ochraceosulfurescens L o c q . ex M . B o n (in D o c . m y c o l . 16 mules; squamules concolorous w i t h centre o n pale ochraceous buff to
(61): 46. 1985) was described as a species i n w h i c h the stipe discolours pale p i n k i s h ochraceous buff background; m a r g i n exceeding lamellae,
y e l l o w when scratched, and i n w h i c h stipe and pileus context are almost when young w i t h lanate-fibrillose y e l l o w to white remnants o f partial
confluent. B o t h are qualitative and variable characters, w h i c h cannot be v e i l . L a m e l l a e , L = 50-60,1 = l-3(-5), rather c r o w d e d to moderately
used o n species level. In other characters the two species do not differ. distant, free, subventricose to ventricose, up to 6 m m broad, cream to
pale b r o w n - y e l l o w (10 Y R 7-8/6), w i t h even, concolorous or slightly
2. Lepiota magnispora M u r r i l l i n M y c o l o g i a 4: 237. 1912. - F i g . 8 1 . paler edge. Stipe 40-110 X 3-10 m m , tapering upwards, not o r sub-
Lepiota fusispora C . H . K a u f f m . i n Papers M i c h . A c a d . S c i . , A r t s bulbous at base, up to 10 m m wide, hollow, at apex white to cream-
Letters 4 : 330. 1924, non Lepiota fusispora P. H e n n . , 1897. - Lepiota coloured and fibrillose (not pruinose), l o w e r d o w n w i t h more or less
ventriosospora D . R e i d i n Trans. B r . m y c o l . S o c . 4 1 : 427. 1958. - distinct lanate annular zone, b e l o w this zone w i t h y e l l o w i s h to w h i t i s h
F i g . 81. Lepiota magnispora.

lanate floccules or rags, w i t h girdles o f b r o w n squamules o n this c o v -

ering; b a c k g r o u n d y e l l o w i s h - b r o w n i s h . Context white i n pileus,
cream-coloured w i t h reddish flush to y e l l o w i s h b r o w n i n stipe. S m e l l
rather strong and unpleasant, like L. cristata. Taste as smell, rarely m i l d
like Boletus edulis. Spore print white.
Spores i n side-view (12.0-)13.5-25.0 X 4.0-6.0 p m , Q = (2.6-)2.7-
4.8(-5.0), Q a v = 3.0-4.2, w i t h more or less straight abaxial side, usu-
a l l y strongly i n f l e x e d above h i l a r appendage ('penguin-shaped'), i n
frontal v i e w fusiform to c y l i n d r i c a l , t h i c k - w a l l e d , dextrinoid, c o n -
gophilous; not metachromatic i n C r e s y l B l u e . B a s i d i a 20-42(-52) X
7.0-13.5 u m , 4-spored, very rarely some 2-spored. L a m e l l a edge ster-
ile. C h e i l o c y s t i d i a 11-40 X 7.0-16 p m , clavate, narrowly clavate and
utriform, a few broadly clavate to spheropedunculate, thin-walled and
colourless. Elements o n pileus erect, (125-)170-400 X 9.0-16(-18) p m ,
rather t h i c k - w a l l e d , not, or rarely septate, i n between w i t h short nar-
r o w l y clavate to c y l i n d r i c a l elements, u p to 35-80 p m l o n g . Pigment
b r o w n , parietal i n lower part o f elements, also intracellular i n upper
part. S t i p i t i p e l l i s a cutis, made u p o f c y l i n d r i c a l , 4.0-8.0 p m w i d e ele-
ments w i t h pale y e l l o w - b r o w n w a l l s . V e l u m partiale made up o f curved
h y a l i n e , c y l i n d r i c a l , 6.0-12 u m w i d e hyphae, some w i t h encrusted
walls. Clamp-connections present.
H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial, i n
coniferous (Picea), m i x e d or deciduous woods, o n nutrient-rich often
calcareous soils. In the Netherlands more or less evenly distributed
over the country (dunes, province o f Flevoland, southern L i m b u r g ) , noj
c o m m o n . E n d o f A u g . - O c t . Widespread i n Europe and N o r t h A m e r i c a ,
f r o m the lowlands up into the montane and boreal zones.
F i g . 82. Lepiota erminea.
T h i s taxon was k n o w n as L. metulaespora i n European literature, but Pileus 20-50(-85) m m , w h e n y o u n g hemispherical, campanulate-con-
R e i d (in Trans. B r . m y c o l . S o c . 4 1 : 427. 1958) showed that that was a i c a l w i t h i n f l e x e d margin, expanding to plano-convex w i t h (broad)
different, tropical species, and described the European taxon as L. ven- u m b o w i t h deflexed margin, smooth at centre, slightly fibrillose-
triosospora ( i n Trans. B r . m y c o l . S o c . 4 1 : 4 2 7 . 1958). However, squamose around centre, at centre pale ochre or y e l l o w i s h b r o w n ( M u .
M u r r i l l described the species i n 1912 from western N o r t h A m e r i c a as 10 Y R 8/6, 8/4), rarely darker (10 Y R 6-7/6), at m a r g i n paler, pale
L. magnispora, a name w h i c h has hardly been i n use since (see cream to white, rarely very pale b r o w n (10 Y R 8/4). L a m e l l a e , L = 38-
V e l l i n g a i n M y c o t a x o n 76: 430-431. 2000 o n the synonymy). 50, 1 = 1-3, moderately distant to moderately crowded, free, ventri-
Descriptions o f L. clypeolaria from the 19th century often include cose, or subventricose, u p to 8 m m broad, w h i t e to pale cream w i t h
L. magnispora (e.g. Fries, M o n o g r . H y m e n o m y c . Sueciae (Lepiota): concolorous even o r f i n e l y flocculose edge. Stipe 15-60(-80) X 2-7
8-9. 1854). P o p u l a r m u s h r o o m guide books for the U . S . A . l i k e w i s e do mm, c y l i n d r i c a l , slightly broadened at base, v a r y i n g from thick and
not make a distinction between the t w o species and use the name L . short to l o n g and slender, h o l l o w , above annulus or annular zone white
clypeolaria for L. magnispora. T h e t w o species are rather s i m i l a r i n and pruinose, l o w e r d o w n white to w h i t i s h (rarely brownish) or y e l -
thi field, but i n general L. magnispora has darker and intenser colours l o w i s h w o o l l y - f i b r i l l o s e , often glabrescent w i t h age and rain, b r o w n -
than L. clypeolaria. T h e difference i n spore shape i s the most reliable ish at utmost base. A n n u l u s often present, hanging, w i t h b r o w n or
character to tell the species apart. y e l l o w i s h floccules. Context i n pileus white, y e l l o w i s h i n stipe to
B o n (in D o c . m y c o l . 4 (15): 27. 1974) distinguished two varieties, brownish, very rarely d i s c o l o u r i n g y e l l o w . S m e l l indistinct, indistinct-
v i z . L. ventriosospora var. ventriosospora w i t h y e l l o w v e i l remnants, ly fungoid, slightly r a n c i d or l i k e L. cristata, rarely raphanoid. Taste
g r o w i n g i n P/cea-forests, a n d var. fulva, more b r o w n i n appearance, m i l d , slightly unpleasant or rancid. Spore print white.
l a c k i n g the y e l l o w c o l o u r o n the v e i l , w i t h longer spores than var. ven- Spores i n side-view 10.0-21.0(-23.0) X 5.0-7.0(-8.0) p m , a v l X
triosospora, a n d g r o w i n g i n deciduous woods. B u t , as collections avw = 11.8-15.1 X 5.5-6.1 um, Q = 1.75-3.5, Q a v = 2.0-2.8, i n side-
f r o m deciduous w o o d s often have a y e l l o w v e i l , these t w o varieties v i e w more or less a m y g d a l i f o r m , o b l o n g to s u b c y l i n d r i c a l , w i t h or
are s y n o n y m i z e d . without suprahilar depression, i n frontal v i e w fusiform to s u b c y l i n d r i -
cal, dextrinoid and congophilous, not c o l o u r i n g i n C r e s y l B l u e , thick-
3. Lepiota erminea (Fr.: Fr.) K u m m . , Fiihr. P i l z k . : 1 3 6 . 1 8 7 1 . - F i g . 82. walled. B a s i d i a 23-47.5 X 7.5-13.5 p m , 4-spored. L a m e l l a edge sterile,
Agaricus ermineus Fr.: Fr., Syst. m y c o l . 1: 2 2 . 1821. - Lepiota but sometimes c h e i l o c y s t i d i a very inconspicuous. C h e i l o c y s t i d i a 11-
clypeolaria var. alba Bres., F u n g i trident. 1: 15. 1882; Lepiota alba 39 X 5.0-15(-27) p m , variable i n shape from one c o l l e c t i o n to anoth-
(Bres.) S a c c , S y l l . F u n g . 5: 37. 1887, non Lepiota alba B e e l i , 1932. er, usually narrowly clavate, but also c y l i n d r i c a l , utriform, or clavate,
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 213. 1995 (as thin-walled to s l i g h t l y t h i c k - w a l l e d a n d colourless. P i l e u s c o v e r i n g
L. alba); R y m a n & H o l m a s e n , Svampar: 414. 1984. made up o f more or less erect colourless or pale c o l o u r e d aseptate
S E L . D E S C R . & F I G S . - H 0 i l a n d i n B l y t t i a 35: 148. 1977 (as L. alba); hyphae, 40-400 X 5.0-12(-25) p m , mostly 200-300 X 6.0-12 p m ; these
H u i j s m a n i n M e d e d . N e d . m y c o l . Vereen. 28: 39-42. 1943; K u h n e r i n elements straight, often twisted at base, w i t h thickened w a l l s . A very
C r y p t o g . M y c o l . 4: 62-65. 1983 (as L. alba); T r i m b . i n D o c . m y c o l . 5 few short elements i n between sometimes present. Stipitipellis a cutis,
(20): 47-48. 1975 (as L. alba). made up o f cylindrical, 3.0-7.0 p m w i d e hyphae, w i t h encrusted, cream
V E R N . N A M E - Duinparasolzwam. walls. V e l u m partiale made up o f non-encrusted, often rather short (20-
30 p m long), 3.0-5.0 u m wide, hyphae. Clamp-connections present i n

all tissues.
H A B I T A T & DISTR. - Solitary to gregarious i n s m a l l groups, sapro-
trophic and terrestrial, i n grassy vegetation types, very rarely i n shrub
vegetation. I n the Netherlands almost completely restricted to the
coastal dunes and to the sandy beach areas o f the province o f Flevoland,
k n o w n from a very few localities i n l a n d (for a distribution map, see
A r n o l d s et al., O v e r z . Paddest. Nederland: 288. 1995); rather c o m m o n .
(June-)Aug.-late N o v . I n Europe widespread i n grasslands o n calcare-
ous soils up into the subalpine zone, probably also i n N o r t h A m e r i c a .
T w o names existed i n literature for a white species w i t h fusiform

spores: Lepiota erminea and L. alba and people have tried to fit the
collections they found into one o r the other, regarding L. erminea as a
species w i t h a raphanoid s m e l l and l o n g spores. H u i j s m a n (in M e d e d .
N e d . m y c o l . Vereen. 28: 39-42. 1943) described a c o l l e c t i o n f r o m the
Netherlands w i t h l o n g spores as L. erminea. However, the presence o f
numerous 2-spored basidia probably accounts for the extraordinary
length o f the spores i n h i s c o l l e c t i o n . T h e differences he described
between the two species do not warrant the distinction o f two species
and here they are considered one.
Specimens o f L. erminea w i t h rather b r o w n pileus can be distin-
F i g . 83. Lepiota oreadiformis.
guished f r o m L. oreadiformis by the w i d e r spores (average w i d t h i n L.
erminea > 5.5 u m , i n i . oreadiformis < 5.5 pm).
Lepiota ochraceodisca M . B o n (in D o c . m y c o l . 21 (81): 52. 1991) d o w n (below ringlike zone) more lanate-floccose o n fibrillose back-
comes very close and might be identical, as i t i s characterized b y an ground and sometimes i n basal part w i t h some b r o w n i s h squamules.
ochraceous-reddish disc a n d a more o r less reddish w o o l l y annulus. A n n u l u s rarely more developed than an arachnoid white zone, i f devel-
Recently it has been reduced to a forma o f L. alba ( M i g l i o z z i & C o c c i a oped, then hanging, white w i t h pale b r o w n i s h r i m . Context i n pileus
i n B o l l . G r u p p o m i c o l . G . B r e s . , n.S. 4 2 : 107. 1999). d u l l , w h i t i s h to pale brownish, i n stipe watery grey, cream, and (pink-
A variety w i t h relatively s m a l l and narrow spores, i.e. measuring ish) brownish i n lower part. S m e l l o f undamaged and o f cut basidio-
(8.5-)9.5-12(-13) X (3.0-)3.5-4.5 p m , w a s described as L. alba var. carps fungoid, unpleasant o f rubber, o r sweetish and pleasant. Taste
angustispora B i z z i & M i g l . (in M i g l . & B i z z i i n M i c o l . ital. 23 (3): 29. m i l d , fungoid, w i t h slightly bitter aftertaste. Spore print ' w h i t e ' .
1994). Spores i n side-view 10.0-16.5 X 4.5-6.5 p m , avl X avw = 11.5-14.4
X 4.8-5.5 um, Q = (1.85-)1.95-3.1, Q a v = (2.05-)2.25-2.7, i n side-view
4. Lepiota oreadiformis Velen., C e s k e H o u b y 2: 2 1 5 . 1 9 2 0 . - F i g . 83. fusiform to amygdaliform, w i t h or without confluent hilar appendage
Agaricus clypeolarius var. pratensis F r . , E p i c r i s i s : 15. 1838; and with or without suprahilar depression, i n frontal view subcylindrical
Lepiota pratensis (Fr.) B i g . & G u i l l . , F l . C h a m p , sup. France: 5 8 . 1 9 0 9 , to narrowly ovoid, dextrinoid, congophilous, not metachromatic i n
non Lepiota pratensis Speg., 1898. - Lepiota gracilis var. laevigata I . Cresyl B l u e , thick-walled. B a s i d i a 17-40 X 6.5-12 u m , 4-spored.
L a n g e i n D a n s k bot. A r k . 2 (3): 24. 1915; Lepiota laevigata (J. Lange) L a m e l l a edge sterile. C h e i l o c y s t i d i a 11-39 X 5.0-13 u m , narrowly
J. L a n g e i n D a n s k bot. A r k . 4 (4): 47. 1923; Lepiota oreadiformis var. clavate, narrowly utriform or more cylindrical, hyaline and slightly thick-
laevigata (J. L a n g e ) M . B o n i n D o c . m y c o l . 2 2 (88): 28. 1993. walled. Pleurocystidia not observed. Pileus covering made up o f more or
S E L . I C O N . - K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: A t l a s p i . less erect elements, non-septate, 60-220(-360) X 6.0-14 u m , cylindrical
71, f i g . 1. 1936 ( a s L . laevigata); R y m a n & H o l m a s e n , Svampar: 415. and attenuated into pedicel, often w i t h , but also found without, short,
1984 (as L. oreadiformis). narrowly clavate to short-cylindrical elements, 18-60 X 10-13 p m ; pale
S E L . D E S C R . & FIGS. - Babos i n A n n l s hist.-nat. M u s . natn. hung. 66: parietal b r o w n pigment present i n both short and l o n g elements.
70. 1974 (as Candusso & L a n z o n i , L e p i o t a : 179-182, fig. 3 1 . 1990; Stipitipellis a cutis o f 4.0-10 u m wide, cylindrical, cream-yellow hyphae.
K i i h n e r i n B u l l , trimest. Soc. m y c o l . Fr. 52: 231. 1936 (as L. laevigata). V e l u m partiale made up o f cylindrical, often curved, hyaline hyphae, c.
V E R N . N A M E - Gladde wolsteelparasolzwam. 6 p m wide. Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - Solitary o r i n s m a l l groups, saprotrophic and
Pileus 20-65 m m , w h e n y o u n g hemispherical to convex, later plano- terrestrial o n sandy to l o a m y soils ( m a i n l y sandy soils w i t h i n the
convex o r applanate w i t h low, broad u m b o , rather pale pinkish-ochra- Netherlands), o n road verges, i n grasslands, also i n deciduous woods.
ceous to dark y e l l o w - b r o w n at centre ( M u . 7.5 Y R 7/6, 4/6-5/8), and In the Netherlands very rare, scattered throughout the country, June-
paler around centre (7.5 Y R 7/6, 8/4-6), and paler, up to almost white, Sept. Rare and widespread i n E u r o p e .
at outermost margin, (very) finely plush-like, almost glabrous, to fine-
l y acute-squamulose at centre, adnate-fibrillose towards margin, w i t h Lepiota oreadiformis is taken i n a broad sense, including L. laevigata,
white w o o l l y o r c o b w e b b y m a r g i n and adhering v e l u m partiale i n originally described as a rather b i g species, with brownish pileus and a
y o u n g specimens; m a r g i n exceeding lamellae. L a m e l l a e , L = 40-60,1 slightly cottony-floccose stipe. Lepiota oreadiformis is generally con-
= 1-7, c r o w d e d to moderately distant, free, subventricose, up to 6 m m sidered a small, pale coloured rather smooth-stiped species. T h e present
wide, greyish white w h e n y o u n g , later more beige w i t h p i n k i s h sheen concept is the same as used by Candusso & L a n z o n i (Lepiota: 178-182.
(10 Y R 7/3), w i t h white, subflocculose edge. Stipe 32-70(-90) X 3-6(- 1990). It appears to be a variable taxon, both i n macroscopical charac-
8) m m , c y l i n d r i c a l w i t h s l i g h t l y broadened base, rarely narrowed at ters (size, development o f an annulus) and i n the presence o f short nar-
base, fistulose w i t h age, white, at apex lengthwise fibrillose, l o w e r rowly clavate elements between the long elements o f the pileus covering.
T h e name L. pratensis (Fr.) B i g . & G u i l l . cannot be used for this Lepiotes: 5 1 . 1993 (not v a l i d ) ; Lepiota cortinarius var. dryadicola
taxon, as i t is a later h o m o n y m o f a different species, described b y (Kuhner) M . B o n i n D o c . m y c o l . 23 (92): 48. 1994.
S p e g a z z i n i (in A n . M u s . nac. Hist. nat. B . A i r e s , Ser. II, 6: 92. 1898), VERN. NAME - Gordijnparasolzwam.
first as L. platensis, and subsequently, i n the Index and Errata, as L.

pratensis, since the name L. platensis was mistakenly used for two dif- K E Y TO THE VARIETIES
ferent species i n the o r i g i n a l publication. 1. Basidiocarps w i t h y e l l o w colours, without p i n k i s h b r o w n or dark
Lepiota pallida L o c q . ex M . B o n & Candusso (in Candusso & brown; pileus smooth, without distinct squamules or patches
L a n z o n i , L e p i o t a : 182. 1990) differs i n having septate pileus covering 5c. var. flava
elements. 1. Basidiocarps w i t h p i n k i s h b r o w n to dark b r o w n colours; pileus sur-
Lepiota granulopunctata L o c q . ex M . B o n (in D o c . m y c o l . 2 2 (88): face breaking up into squamules and patches
29. 1993) comes also very close. T h e m a i n characteristic seems to be 2. P i l e u s w i t h m e d i u m to pale b r o w n colours ( p i n k i s h orange-
the almost glabrous stipe, but whether this warrants recognition at brown, pale hazel-brown, reddish-ochraceous, isabella)
species level is unclear. Further study o f this group o f grassland species 5a. var. cortinarius
is badly needed. 2. Pileus w i t h dark b r o w n colours 5b. var. audreae
D a r k specimens o f L. erminea differ from L. oreadiformis i n the
more y e l l o w colours (in the range o f M u . 10 Y R ) , whereas L. praten- 5a. var. cortinarius
sis is characterized b y more red i n the pileus (in the range o f M u . 7.5 S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 19a. 1990; Cetto,
Y R ) . T h e average w i d t h o f the spores is an additional character to dis- F u n g h i Vero 3, E d . 3: p i . 863. 1982; J . L a n g e i n D a n s k bot. A r k . 2 (3):
tinguish the two species: 4.8-5.5 p m i n L. oreadiformis versus 5.5-6.1 p i . l b . 1915; J . L a n g e , F l . agar. dan. 1: p i . 1 0 B . 1935.
p m i n L. erminea. In both species the length (and also the length-width S E L . D E S C R . & FIGS. - H u i j s m a n i n Persoonia 1: 325, figs. 1-3. 1960;
ratio) o f the spores varies considerably. J. L a n g e i n Dansk bot. A r k . 2 (3): 2 5 . 1915; V e l l i n g a i n Persoonia 14:
Sometimes specimens o f L. oreadiformis resemble y o u n g speci- 410-412, fig. 2 . 1 9 9 2 ; G . A . de V r i e s i n C o o l i a 12:48-52, figs. 1-5.1966.
mens o f L. clypeolaria. Spore sizes and shapes o f the two species are
similar, but i n L. clypeolaria the surface o f the pileus splits u p i n a ' Pileus 40-80 m m , w h e n y o u n g broadly and obtusely c o n i c a l , rarely
distinct central part and discrete squamules around centre, whereas^ hemispherical, w i t h i n f l e x e d margin, expanding to conico-convex,
the surface i n L. oreadiformis is rather u n i f o r m l y c l o s e d and u n i - plano-convex or applanate, w i t h broad, l o w or h i g h umbo, often slight-
f o r m l y splitting u p into s m a l l patches, g i v i n g the surface an evenly l y undulating around centre, at centre tomentose, tomentose-felted,
coloured l o o k . around centre breaking up into concentrical zones o f patches or gran-
ulose to fibrillose-arachnoid squamules, at centre and when y o u n g i n
5. Lepiota cortinarius J . L a n g e i n D a n s k bot. A r k . 2 (3): 25. 1915. - squamules p i n k i s h orange-brown ( M u . 5 Y R 4-3/4, 4/6), later around
F i g . 84. centre squamules paler, pale hazel-brown to pale isabella (5-7.5 Y R Y R
Lepiota audreae ( D . R e i d ) M . B o n i n D o c . m y c o l . 11 (43): 3 5 . 5/8 to 7.5 Y R 7/6), o n cream)to"pale p i n k i s h background, when y o u n g
1981. - Lepiota dryadicola K u h n e r i n C r y p t o g . M y c o l . 4: 6 5 . 1983; w i t h white lanate-floccose velar remnants at margin, and m a r g i n
Lepiota audreae var. dryadicola (Kuhner) M . B o n , F l . m y c o l . Eur. 3, exceeding lamellae. L a m e l l a e , L = c. 100, 1 = ( l - ) 3 , moderately
crowded or crowded, free to remote from stipe, ventricose, subventri-
cose or segmentiform, up to 6 m m broad, white to cream, w i t h whitish,
flocculose edge. Stipe 48-90 X 3.5-12 m m , broadened towards base to
usually abrupt, up to 25 m m w i d e , bulb, h o l l o w w i t h age, whitish at
apex, c r e a m y - y e l l o w i s h , glabrous or innately fibrillose lengthwise,
lower d o w n w i t h lanate-floccose p i n k i s h b r o w n tomentum, w h i c h i s
appressed w i t h age; sometimes squamules as o n pileus present o n bulb.
Context thick, d u l l and white i n pileus, white i n stipe, w i t h y e l l o w -
brown to g o l d e n - y e l l o w tinges i n bulb. S m e l l strong, fruity, sweetish,
sometimes weak and reminiscent o f L. cristata. Taste unpleasant.
Spore print w h i t i s h to cream.
Spores (6.5-)7.0-9.0(-10.0) X (2.5-)3.0-3.5(-4.0) p m , Q = 2.1-2.9(-
3.1), Q a v = 2.3-2.7, i n side-view c y l i n d r i c a l , often w i t h slight suprahi-
lar depression ('penguin-shaped'), i n frontal v i e w c y l i n d r i c a l to
slightly o v o i d , dextrinoid, congophilous, not metachromatic i n C r e s y l
B l u e . B a s i d i a 18-29 X 5.5-9.0 p m , 4-spored, rarely also 2-spored.
L a m e l l a edge sterile. C h e i l o c y s t i d i a 15-50 X 6.0-13.5 p m , variable i n
shape and size: fusiform, broadly utriform, narrowly clavate or clavate,
colourless, though rarely w i t h y e l l o w i s h refringent contents, slightly
thick-walled. Pileus covering made up o f l o n g , adnate or ascending to
erect, not or rarely septate, elements, (50-)150-400(-800) X 10-20 p m ,
w i t h parietal b r o w n i s h pigment, slightly t h i c k - w a l l e d , o c c a s i o n a l l y
w i t h basal short, clavate elements, e.g. 30-35 X 14-18 u m . Stipitipellis
a cutis o f c y l i n d r i c a l , 3.0-10 u m w i d e hyphae, w i t h loose, irregular,
colourless, patent, c y l i n d r i c a l , c. 3.0 p m w i d e hyphae. C l a m p - c o n n e c -
tions present i n a l l tissues.
H A B I T A T & DISTR. - Solitary to gregarious, i n fairy rings, sometimes
F i g . 84. Lepiota cortinarius (a: var. audreae). fasciculate, saprotrophic and terrestrial o n rather nutrient-rich, often
calcareous, sandy to clayey soils, i n deciduous and m i x e d woods, city-

parks, and o n m i n e waste heaps. I n the Netherlands rare, m a i n l y i n the
new polders, scattered o n l o a m y localities i n the Pleistocene regions,
and i n southern L i m b u r g . S e p t . - O c t . ( - D e c ) . W i d e s p r e a d and rare i n
Europe, also k n o w n f r o m the eastern part o f the U . S . A .
5b. var. audreae D . R e i d i n F u n g . rar. Ic. c o l . 3: 8. 1968.

Lepiota audreae var. dryadicola ( K i i h n e r ) M . B o n , F l . m y c o l . Eur.
3, Lepiotes: 5 1 . 1993 (not valid); Lepiota cortinarius var. dryadicola
(Kiihner) M . B o n i n D o c . m y c o l . 23 (92): 4 8 . 1994.
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 19b. 1990; D . R e i d
i n F u n g . rar. Ic. c o l . 3: p i . 19a. 1968.
S E L . D E S C R . & FIGS. - W . B e y e r i n Z . P i l z k . 4 3 : 193-196. 1977 and
i n L i b r i bot. 5 : 2 1 1 . 1992; K e l d e r m a n i n C o o l i a 31: 1 6 - 1 7 , f i g . 3 . 1 9 8 8 ;
D . R e i d i n F u n g . rar. Ic. c o l . 3: 8-10. 1968.
C H A R A C T E R I S T I C S - P i l e u s 35-45(-60) m m , dark b r o w n and slightly

squarrose-felted at centre, around centre breaking up into discrete, c o n -
centrical, hazel-brown patches or squamules; stipe 25-50 X 6-7 m m ,
also w i t h rather dark b r o w n squamules.
M i c r o s c o p i c a l characters as i n the t y p i c a l variety.
H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial on nutri-
ent-rich soils; i n the Netherlands very rare, o n l y k n o w n from m i n e
F i g . 85. Lepiota subgracilus.
waste heaps i n southern L i m b u r g (Brunssum). Sept.-Oct. Rare but
widespread i n Europe, possibly more thermophilic than var. cortinarius.
Pileus 20-48 m m , campanulate, expanding to plano-convex, plano-
5c. var. flava B a s & V e l l i n g a i n V e l l i n g a i n Persoonia 14: 414. 1992. concave, w i t h broad l o w umbo, at centre dark red-brown, fulvous,
S E L . D E S C R . & FIGS. - B a s & V e l l i n g a i n V e l l i n g a i n Persoonia 14: greyish p i n k i s h b r o w n , paler towards margin, at centre densely set w i t h
414, figs. 3d-h. 1992. small fibrillose o r p y r a m i d a l squamules, not showing the underlying
context, towards m a r g i n w i t h fibrillose or more granular squamules,
C H A R A C T E R I S T I C S - P i l e u s ochraceous y e l l o w , smooth and glabrous, on white, ochraceous background, w i t h o r without vinaceous tinges,
but i n outer half w i t h granular dots, slightly darker than background; sometimes at m a r g i n w i t h a zone o f dark orange-brown squamules
stipe pale straw-yellow. ( M u . 7.5 Y R 6/6). L a m e l l a e , L = 4 5 , 1 = 1-3, moderately distant to
M i c r o s c o p i c a l characters as i n t y p i c a l variety, but elements o f crowded, free to narrowly adnexed, segmentiform to ventricose, up to
pileus covering w i t h y e l l o w w a l l s . 8 m m wide, creamy white, cream-coloured, very pale grey-brown w i t h
H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial o n clay age, w i t h finely serrate to almost even, white edge. Stipe 40-90 X 2-
in plantation o f deciduous trees. I n the Netherlands o n l y k n o w n from 6.5 m m , c y l i n d r i c a l o r slightly broadening at base, h o l l o w , cream, pale
one locality i n the province o f F l e v o l a n d (Dronten, Spijkbos), Oct. N o t brownish or pale ochraceous w i t h l i l a c i n o u s tinge, finely fibrillose,
k n o w n outside the Netherlands. w i t h i n lower half o r o n l y at base, some girdles o f squamules, c o n c o l -
orous w i t h those o n pileus. A n n u l u s usually present as white w o o l l y
6. Lepiota subgracilis K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 5 2 : remnants o f the partial veil, and also as a b r o w n ring, remnants o f the
231. 1 9 3 6 . - F i g . 8 5 . universal v e i l . Context white and d u l l i n pileus, i n stipe concolorous
Lepiota clypeolaria var. gracilis Q u e l , i n C . r. A s s . franc. A v . S c i . w i t h surface and shiny. S m e l l and taste not distinct, w h e n o l d reminis-
(Besancon, 1893) 2 2 : 4 8 5 . 1894 ( C h a m p . Jura Vosges S u p p l . 19); cent o f L. cristata. Spore print probably white.
Lepiota gracilis (Quel.) J . L a n g e i n D a n s k bot. A r k . 2 (3): 2 4 . 1915, Spores 9.0-13.5 X 4.0-6.0(-6.5) p m , Q = 1.8-2.4(-2.6), Q a v = 1.9-
n o n Lepiota gracilis Peck, 1899; Lepiota gracilis (Quel.) R e a , B r i t . 2.1, i n side-view a m y g d a l i f o r m to oblong, w i t h o r without suprahilar
B a s i d i o m y c : 70. 1922 (superfluous). - Lepiota subgracilis Wasser i n depression, i n frontal v i e w oblong, broadly fusiform or slightly ovoid,
U k r . bot. Z h . 3 5 : 518. 1978, non Lepiota subgracilis K i i h n e r , 1936; uniguttulate, t h i c k - w a l l e d , strongly dextrinoid, c o n g o p h i l o u s ; inner
Lepiota wasseri M . B o n i n D o c . m y c o l . 2 2 (88): 2 9 . 1993 (superflu- w a l l not metachromatic i n C r e s y l B l u e , w h i l e spore content colours
ous). - Lepiota kuehneriana L o c q . i n F r i e s i a 5: 296. 1956. blue. B a s i d i a (18-)21-35 X 7.5-10 p m , 4-spored, w i t h some 2- or even
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 2 3 0 . 1 9 9 5 (as 1-spored ones intermixed. L a m e l l a edge sterile, sometimes i n places
L. kuehneriana); Candusso & L a n z o n i , L e p i o t a : p i . 12. 1990 (as L. fertile. C h e i l o c y s t i d i a 15-43 X 6.5-11 u m , variable i n shape, c y l i n d r i -
kuehneriana); Hardtke & R o d e l i n M y k o l . M i t t b l . 36: 5. 1993; J . cal and subcapitate to narrowly clavate, or lageniform to utriform, nar-
L a n g e , F l . agar. dan. 1: p i . 11F. 1935 (as L. gracilis). r o w l y clavate, thin-walled and colourless. Pleurocystidia absent. Pileus
S E L . D E S C R . & FIGS. - B i z z i & M i g l . i n M i c o l . ital. 28 (2): 23-31. covering made up o f elongated elements, often united to squamules;
1999; Candusso & L a n z o n i , L e p i o t a : 159-162, fig. 2 5 ; 184-186, fig. elements 55-360 X 8.0-13 p m , c y l i n d r i c a l , slightly flexuous i n lower
33. 1990 (as L. kuehneriana and L. subgracilis resp.); E n d e r l e & part, non-septate, o r w i t h 1 septum, w i t h slightly thickened, b r o w n o r
Krieglst. i n Z . M y k o l . 5 5 : 97-98. 1989; Hardtke & R o d e l i n M y k o l . grey-brown w a l l s , sometimes also w i t h intracellular pigment, w i t h or
M i t t b l . 36: 1-3. 1993; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 66-67. without basal clavate elements, w i t h dark b r o w n parietal pigment.
1994; S v r c e k i n C e s k a M y k o l . 19: 47-49, fig. 5. 1965. Stipitipellis a cutis, made up o f c y l i n d r i c a l hyphae, 4.0-6.0 u m wide,
V E R N . N A M E - Slanke wolsteelparasolzwam. w i t h pale y e l l o w w a l l s ; elements o f squamules as those o n pileus.
Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - S o l i t a r y o r i n s m a l l groups, terrestrial a n d Sect. Ovisporae (J. Lange) Kuhner

saprotrophic, o n c l a y e y soils, often r i c h i n nutrients and/or c a l c i u m , i n
deciduous woods, i n the Netherlands very rare, k n o w n from southern Spores e l l i p s o i d to o b l o n g ; pileus covering made up o f l o n g , erect ele-
L i m b u r g (Valkenburg, Schaelsberg; C a d i e r en Keer, Riesenberg) a n d ments, w i t h o r without short clavate elements at the base o f these l o n g
f r o m one l o c a l i t y i n the river area (Barendrecht, H u y s ten D o n c k ) . elements.
Sept.-Oct. R a r e a n d scattered throughout E u r o p e , but absent from
alpine and arctic regions.
Lepiota subgracilis is a variable species, variable i n colours o f the Subsect. Felininae M . B o n

pileus (from very dark b r o w n to ochraceous brown), i n presence o f
velar remnants o n the stipe (as girdles, o r as two ringzones, one o f the Pileus covering made up o f l o n g , erect elements and short, clavate ele-
partial v e i l , one o f the universal veil), and i n presence o r absence o f a ments.
h y m e n i f o r m underlayer i n the pileus covering. It seems that the
h y m e n i f o r m underlayer i s present i n y o u n g specimens, whereas the 8. Lepiota felina (Pers.) P. Karst., R y s s L , F i n l . , S k a n d . H a l f 0 n s
elements lengthen and do not persist as a separate layer i n older spec- Hattsv.: 10. 1879. - F i g . 86.
imens. T h e variation i n colours and i n presence o f velar remnants was Agaricusfelinus Pers., S y n . meth. F u n g . : 261. 1801; Lepiota clype-
observed by K e l d e r m a n (Parasolzw. Z u i d - L i m b u r g : 67. 1994) i n seven olaria wax. felina (Pers.) G i l l e t , H y m e n o m y c e t e s : 62. 1874.
collections f r o m one locality. M I S A P P L . - Lepiota clypeolarioides sensu H u i j s m a n i n M e d e d .
Lepiota kuehneriana L o c q . , L. wasseri M . B o n , and probably also N e d . m y c o l . Vereen. 28: 25-27. 1943.
L. latispora (Wasser) M . B o n , b e l o n g to this same variable taxon. S E L . I C O N . - D a h n c k e , 1200 P i l z e : 519. 1993; Papeti & F o r t i i n
E x t e n s i v e descriptions o f the latter taxon are l a c k i n g . B o n (in D o c . B o l l . C i r c . m i c o l . G . C a r i n i 29-30: m i d d l e page. 1995.
m y c o l . 11 (43): 30, 32. 1981; and F l . m y c o l . Eur. 3, Lepiotes: 56-60. S E L . D E S C R . & FIGS. - Papeti & F o r t i i n B o l l . C i r c . m i c o l . G . C a r i n i
1993) placed L. latispora and L. kuehneriana i n subsect. Latisporinae 29-30: m i d d l e page. 1995.
M . B o n , characterized by spores w i t h length-width ratio < 2.5, where* VERN. NAME - Panterparasolzwam.
as L. subgracilis and L. wasseri were ranked i n subsect. Fusisporinae
(J. L a n g e ) M . B o n , characterized b y spores w i t h a length-width ratio Pileus 15-40 m m , hemispherical when young, expanding to plano-con-
> 2.5. However, the spore sizes given b y B o n scarcely differ for any vex, applanate,-a bit undulating, w i t h o r without umbo, at centre w i t h
o f these taxa, and the length-width ratio is less than 2.5 i n a l l cases. uplifted, plush-like, very dark b r o w n ( M u . 10 Y R 4-3/2, 7.5 Y R 3/2, 5
Y R 3/3), little tufts i n a distinct patch, very rarely more orange-brown
7. Lepiota ignivolvata Bousset & Joss, i n Joss, i n R i v . M i c o l . 33: 123. (7.5 Y R 3/4), around centre w i t h concentrical zones o f uplifted o r flat-
1990. tened, discrete patches, concolorous w i t h centre, o n a cream back-
Lepiota ignivolvata Bousset & Joss, i n Joss, i n B u l l , trimest. Soc. ground, towards m a r g i n the patches become more r a d i a l l y arranged,
m y c o l . Fr. 64: 5. 1948 (not valid); Lepiota ignivolvata Bousset & Joss, more c o b w e b b y squamules; margin a bit ragged, rarely w i t h annulus
i n Joss, i n B u l l . mens. S o c . l i n n . L y o n 4 1 : 5. 1972 (not valid); Lepiota remnants attached, and exceeding lamellae. L a m e l l a e , L = 30-45,1 =
ignivolvata Bousset & Joss, i n Joss, i n B u l l . mens. Soc. l i n n . L y o n 58: 1-3, moderately distant to moderately crowded, free, some anasto-
328. 1989 (not valid). mosing, (sub)ventricose, up to 5 m m w i d e , cream-coloured, sometimes
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 229. 1995; w i t h p i n k i s h sheen, w i t h white to concolorous eroded o r flocculose
Candusso & L a n z o n i , L e p i o t a : p i . 11. 1990; Courtec. & D u h e m , G u i d e edge. Stipe 32-60 X 2-5 m m , gradually w i d e n i n g towards base, h o l -
C h a m p . France Europe: p i . 6 6 8 . 1 9 9 4 ; Dahncke, 1200 Pilze: 5 1 3 . 1 9 9 3 . low, innately fibrillose lengthwise, w h i t i s h o r pale b r o w n i s h (7.5 Y R
S E L . D E S C R . & FIGS. - Joss, i n B u l l , trimest. Soc. m y c o l . Fr. 64: 7- 8/4) above annulus, b e l o w annulus p i n k i s h - b r o w n i s h (7.5 Y R 6/4) o r
8, fig. 1. 1948; S c h w o b e l i n Z . P i l z k . 32: 2-4, figs. 2 & 3. 1966. grey brownish, w i t h , rarely without, i n l o w e r h a l f some scattered gir-
i dles o f squamules, paler than those* o n pileus, w i t h white tomentum
C H A R A C T E R I S T I C S - P i l e u s 50-90 m m , w i t h glabrous b r o w n centre, and some white m y c e l i a l cords at base. A n n u l u s l i k e a cuff around
around centre w i t h adnate felted-tomentose, b r o w n i s h squamules o n
pale background; lamellae cream-coloured; stipe 70-100 X 10-15 m m ,
w i t h b r o w n narrow annulus, a n d belts o f floccules b e l o w annulus,
w h i t i s h cream, i n basal part d i s c o l o u r i n g orange w h e n touched, w i t h
age o r o n d r y i n g ; context white to creamy; smell strong, like the rub-
ber s m e l l o f L. cristata; taste unpleasant, l i k e rubber.
Spores 8.5-11.0(-12.5) X 5.0-6.0(-6.5) p m , Q = 1.75-1.9, Q a v =
1.75-1.95, a m y g d a l i f o r m w i t h rounded apex i n side-view; cheilocys-
tidia 17-37 X 9.0-17 m m , narrowly clavate to clavate; pileus covering
made u p o f l o n g (reaching over 500 pm) ascending to erect elements,
w i t h a rather regular layer o f short elements; pigment b r o w n and pari-
etal i n l o n g and short elements; clamp-connections present.
H A B I T A T & DISTR. - Solitary o r i n small groups, saprotrophicând
terrestrial, i n beech-forests (Fagus) o n calcareous soils. A c c o r d i n g to
Josserand ( i n B u l l , trimest. S o c . m y c o l . F r . 64: 8. 1948) i n Picea-
forests. K n o w n from Denmark, Switzerland, France, L u x e m b o u r g , and
Germany. Sept.-Oct.
F i g . 86. Lepiota felina.

stipe, rarely o n l y fibrillose-membranaceous, ragged at upper side, set

w i t h dark g r e y - b r o w n squamular patches, disappearing w i t h age.
Context white i n pileus, concolorous w i t h surface i n stipe. S m e l l sug-
gestive o f cedar w o o d , o f cut basidiocarps l i k e the rubber component
of the s m e l l o f L. cristata: astringent and unpleasant. Taste not k n o w n ,
probably unpleasant. Spore print white.
Spores 6.0-8.5 X (3.5-)4.0-5.0 p m , Q = 1.45-1.95, Q a v = 1.65-
1.75, e l l i p s o i d to o b l o n g , some i n side-view a m y g d a l i f o r m w i t h blunt
apex, w i t h abrupt hilar appendage, thick-walled, dextrinoid, though not
all spores reacting i n the same strong way, congophilous, not
metachromatic i n C r e s y l B l u e . B a s i d i a 21-35 X 6.5-9.0 p m , 4-spored,
intermixed w i t h a few 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a
21-53 X 4.5-10 p m , most often c y l i n d r i c a l or slightly irregularly c y l i n -
drical, w i t h acuminate apex, often w i t h septum, rarely w i t h capitate
apex, sometimes fusiform, colourless, w i t h s l i g h t l y thickened w a l l .
Pleurocystidia absent. P i l e u s covering made up o f agglutinated long,
non-septate, erect elements i n tufts, 110-415 X 9.0-17 p m , w i t h dark
b r o w n parietal pigment that decreases i n quantity towards apex, w i t h -
out septa, w i t h a very dense underlayer made up o f narrowly clavate to
clavate elements, up to 15 p m wide, w i t h dark b r o w n parietal and i n
lower parts sometimes encrusting dark b r o w n pigment; encrusting dark
b r o w n pigment also present i n l o w e r hyphae. Stipitipellis b e l o w annu- F i g . 87. Lepiota pseudolilacea.
lus, a cutis o f c y l i n d r i c a l , irregular, colourless hyphae, w i t h slightly

encrusted walls, 3.5-6.0 u m wide, w i t h some irregular b r o w n - w a l l e d squamules lacking, p i n k i s h greyish b r o w n to dark red-brown ( M u . 7.5
hyphae w i t h narrowly clavate terminal elements, c. 9.0-10 u m wide, Y R 4/3, 5/4; 5 Y R 4/4, 5-7.5 Y R 3/3), b r o w n w i t h some grey tinges
and squamules l i k e those o n pileus, w i t h clavate and l o n g elements. (7.5 Y R 5/4 plus 7.5 Y R 4/6), b r o w n (10 Y R 4/5) or, especially w h e n
Clamp-connections present i n a l l tissues. old, more orange-brown (7.5-10 Y R 4-6/6), rarely w i t h lilacinous
H A B I T A T & DISTR. - In s m a l l groups and saprotrophic o n nutrient- tinges, near m a r g i n s h o w i n g underlying w h i t i s h context, w i t h margin
r i c h , often calcareous soil, i n deciduous or coniferous woods, also o n exceeding lamellae, especially w h e n y o u n g . L a m e l l a e , L = 32-48,1 =
m i n e waste heaps. In the Netherlands rather rare, o c c u r r i n g i n the 1-3, moderately distant to crowded, free, but quite close to stipe, ven-
southern part o f L i m b u r g , the p r o v i n c e o f F l e v o l a n d and i n l o c a l l y tricose, subventricose, or segmentiform, up to 5 m m w i d e , cream w i t h
l o a m - r i c h places o f the Pleistocene regions. Sept.-Oct. Widespread and brownish tinge or w i t h p i n k i s h sheen (e.g. 10 Y R 8/2-3), w i t h white,
not c o m m o n throughout E u r o p e ; also recorded from N o r t h A m e r i c a . fimbriate or flocculose (lens) edge. Stipe 23-55 X 2-9 m m , cylindrical,
not or slightly broadened at base, h o l l o w , w i t h distinct annulus, and
Lepiota felina can be confused i n the field w i t h L. pseudolilacea, as when y o u n g above this annulus a white fugacious annulus, lengthwise
both are characterized b y the cuff-like annulus w i t h little dark patches fibrillose, cream(ish) (7.5 Y R 8/2) at apex, b r o w n i s h - p i n k i s h , p i n k i s h -
on it. Lepiota pseudolilacea can be distinguished o n account o f the lilacinous (7.5 Y R 6/4), to b r o w n i s h at base, i n lower part w i t h some
structure o f the pileus c o v e r i n g , w h i c h lacks the dense h y m e n i f o r m girdles o f rather w o o l l y squamules. A n n u l u s like a cuff around stipe,
underlayer o f L. felina, and is p r o v i d e d w i t h adnate coloured hyphae. white-cream and set w i t h a band o f squamules or patches concolorous
to those o n pileus, rarely w i t h a dark zone only, sometimes w i t h dark-
9. Lepiota pseudolilacea H u i j s m a n i n B u l l . mens. S o c . l i n n . L y o n 16: er upper r i m . Context white i n pileus, glassy above lamellae, shiny i n
182. 1 9 4 7 . - F i g . 87. stipe and concolorous w i t h surface. S m e l l o f basidiocarps w h e n cut,
Lepiotula pseudolilacea (Huijsman) Wasser i n N o v . Sist. v y s s h . fungoid-astringent, fruity-aromatic, even sweet-aniseed-like, or l i k e
n i z s h . Rast. 1975: 191. ( ' 1 9 7 5 ' ) 1976. - Lepiota pseudohelveola the rubber component o f the s m e l l o f L. cristata. Taste indistinct or as
K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 221-223. 1936 (not valid); smell. Spore print white.
Lepiota pseudohelveola K i i h n e r ex H o r a i n Trans. B r . m y c o l . S o c . 43: Spores 6.0-10.0 X 4.0-5.0 pm, Q = 1.4-2.1, Q a v = 1.6-1.8, i n side-
44. 1960. - Lepiota pseudohelveola var. sabulosa M . B o n in Doc. view ellipsoid, oblong, even subcylindrical, rarely w i t h straight adax-
m y c o l . 17 (67): 8. 1987. ial side or even w i t h slight suprahilar depression, e l l i p s o i d to oblong
E X C L . - Lepiota pseudohelveola sensu J . Favre, C h a m p , sup. Z o n e in frontal view, w i t h inner w a l l orange-brown i n M e l z e r ' s Reagent,
alpine: 157-158. 1955 ( = L.favrei). congophilous, cyanophilous, not metachromatic in Cresyl Blue.
S E L . I C O N . - Candusso i n R i v . M i c o l . 33: 17. 1990 (as var. sabu- B a s i d i a 15-33 X 6.0-9.0 u m , 4-spored, rarely 2-spored. L a m e l l a edge
losa); V e l l i n g a & Huijser i n C o o l i a 40: p i . 5. 1997 (as L. pseudohelve- sterile. C h e i l o c y s t i d i a 17-43 X 5.0-12 p m , mostly (irregularly) c y l i n -
ola). drical, but also n a r r o w l y utriform, clavate or narrowly clavate.
S E L . D E S C R . & FIGS. - D e r b s c h & Schmitt, A t l . P i l z e Saarlandes 2: Pleurocystidia absent. P i l e u s covering made up o f more or less erect
493. 1987; K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 221-223. 1936 l o n g elements i n tufts, 100-370 X 8-16 p m , sometimes o n l y up to 200
(both as L. pseudohelveola). pm long, w i t h short-clavate elements i n between, 13-60 X 10-15 p m ,
V E R N . N A M E - M a n c h e t p a r a s o l z w a m , i n c l . Valse l i l a parasolzwam. and a layer o f repent c y l i n d r i c a l coloured hyphae, 4.0-6.0 u m i n d i a m .
Pigment b r o w n , intracellular and parietal, also encrusting i n repent
Pileus 13-45 m m , w h e n y o u n g rounded-conical w i t h inflexed margin, hyphae, and pigment especially present i n the short elements.
expanding to plano-convex, applanate or plano-concave w i t h or w i t h - Stipitipellis a cutis o f colourless, c y l i n d r i c a l hyphae, 4.0-10 p m i n
out l o w u m b o , almost completely felted, p l u s h - l i k e at centre, towards diam. Squamules on stipe as those o n pileus. Clamp-connections pres-
margin more r a d i a l l y fibrillose, and the tufts a bit uplifted, but discrete ent i n a l l tissues.
H A B I T A T & DISTR. - Gregarious, rarely solitary, saprotrophic and ter- Pileus 18-42 m m , at first hemispherical, rounded c o n i c a l w i t h inflexed
restrial i n dune grasslands, and i n deciduous woods on calcareous soils, margin, expanding v i a campanulate to plano-convex or applanate w i t h
also o n m i n e waste heaps. In the Netherlands rather rare, i n southern umbo, w h e n y o u n g grey-brown squamulose, later at u m b o dark grey-
L i m b u r g and i n the coastal dunes. Sept.-Nov. Widespread i n Europe. b r o w n w i t h greenish or o l i v e tinge ( M u . 10 Y R 5/2, 5/3-4), velvety-
plushy, w i t h s m a l l tufts, around u m b o w i t h concentrical, small
Lepiota pseudolilacea and L. pseudohelveola are considered synony- plush-like, grey-brown (10 Y R 5/4 to 7/4) squamules o n pale to white
mous, as the differences i n spore shape and i n size o f the elements o f background, i n m a r g i n a l zone more r a d i a l l y fibrillose, w h e n y o u n g
the pileus c o v e r i n g o f the t w o species are s m a l l and there is a range w i t h white overhanging margin exceeding lamellae. L a m e l l a e , L = 4 0 -
i n spore shape a n d i n size o f the pileus elements. T h e better k n o w n 60,1 = 1-5, moderately crowded, free, ventricose or segmentiform, up
name L. pseudohelveola was v a l i d l y p u b l i s h e d o n a later date than L. to 6.5 m m w i d e , white at first, later cream-coloured, or greyish w i t h
pseudolilacea. y e l l o w i s h tinge (2.5 Y 8/4), sometimes w i t h p i n k i s h flush w h e n l o o k e d
Lepiota pseudohelveola var. sabulosa M . B o n (in D o c . m y c o l . 17 at from below, w i t h white floccose to eroded edge. Stipe 23-58 X 3-6
(67): 8. 1987) is identical i n most characters, though macroscopically m m , c y l i n d r i c a l or tapering towards apex, h o l l o w , white, lengthwise
distinct because o f its paler colours. innately creamy-fibrillose, i n lower 1/2-2/3 set w i t h w o o l l y white
In several Lepwto-species a w i d e range o f colours can be observed; v e l u m girdles, i n l o w e r part just above base, w i t h squamules, c o n c o l -
m i c r o s c o p i c a l characters and characters concerning presence and orous to those o n pileus (10 Y R 5-4/4), w i t h white m y c e l i a l cords.
shape o f a r i n g are considered to be more important than c o l o u r for Context white to cream i n pileus and stipe. S m e l l o f basidiocarps when
species delimitation. cut indistinct, fungoid, unpleasant: a bit a c i d , or fishy-rubber-like,
Lepiota pseudolilacea is a t o x i c species, containing amanitins sometimes reminiscent o f the s m e l l o f L. cristata. Taste not k n o w n .
(Gerault & G i r r e i n C . r. hebd. Seanc. A c a d . S c i . , Paris 2 8 0 , Ser. D : Spore print white.
2841-2843. 1966 ( a s L . pseudohelveola)). Spores (5.5-)6.0-8.0 X 3.5-5.0 p m , Q = 1.6-1.95, Q a v = 1.8-1.9,
Lepiota farinolens B o n & G . Riousset (in D o c . m y c o l . 22 (85): 6 5 . oblong i n side-view, sometimes w i t h the adaxial side flatter than the
1992), described from grassy localities i n southern France, has also an abaxial side, i n frontal v i e w oval-oblong, dextrinoid, congophilous, not
annulus, but this one is not set w i t h dark squamules, and is distinguished metachromatic i n C r e s y l B l u e . B a s i d i a 16-23 X 6.0-8.0 p m , 4-spored,
on account o f its pale p i n k i s h colours, and the farinaceous smell, whictf sometimes w i t h some 2-spored basidia intermixed. L a m e l l a edge ster-
is especially distinct after rubbing the basidiocarps. T h e spores o f L. ile. C h e i l o c y s t i d i a 15-32 X 3.5-8.0 u m , variable i n shape, almost c y l i n -
farinolens measure 7.5-9.5 X (4.0-)4.5-5.0 p m , Q = 1.6-2.0, and the drical to very n a r r o w l y utriform, slightly fusiform, narrowly clavate
elements o f the pileus covering are relatively short: 100-160(-180) p m etc., relatively s m a l l , and t h i n - w a l l e d . P l e u r o c y s t i d i a absent. Pileus
long, and a layer o f short clavate elements is lacking, p l a c i n g this covering made up o f erect to adnate, non-septate elements, 60-370 X
species i n the v i c i n i t y o f L. subincarnata and L. brunneoincarnata. 5.0-10 p m , rarely septate i n basal part, narrowed into base, w i t h pale
y e l l o w i s h to greyish-brownish parietal pigment, w i t h at base some nar-
10. Lepiota forquignonii Q u e l , i n C . r. A s s . franc. A v . S c i . ( B l o i s , r o w l y clavate to c y l i n d r i c a l short elements, 22-55 X 8.0-14.5 p m , also
1884) 13: 277. 1885 ( C h a m p . Jura Vosges S u p p l . 13). - F i g . 88. w i t h pale b r o w n i s h parietal pigment. S t i p i t i p e l l i s a cutis o f narrow,
Lepiota forquignonii var. coniferarum M . B o n i n D o c . m y c o l . 11 c y l i n d r i c a l hyphae, 3.0-5.0 p m w i d e ; partial v e i l consisting o f irregu-
(43): 4 1 . 1981. - Lepiota olivaceobrunnea P . D . Orton i n Trans. B r . lar, colourless hyphae, 3.0-5.0 p m w i d e , w i t h encrusting colourless
m y c o l . S o c . 9 1 : 561. 1988; Lepiota forquignonii var. olivaceobrunnea pigment; squamules o n stipe made up o f elements as o n pileus, but
(P.D. Orton) M . B o n i n D o c . m y c o l . 22 (88): 2 9 . 1993. w i t h a relatively h i g h number o f relatively short, narrowly clavate ele-
E X C L . - Lepiota forquignonii sensu B a r b i e r i n B u l l . mens. S o c . ments, c. 5 0 X 8.0 p m . Clamp-connections present i n a l l tissues.
l i n n . L y o n 3: 76-78. 1934 ( = L. grangei). H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial o n
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 30. 1990; Papeti & more or less calcareous soils, i n deciduous woods. In the Netherlands
F o r t i i n B o l l . C i r c . m i c o l . G . C a r i n i 29-30: m i d d l e page. 1995; D . R e i d rare, o n l y k n o w n f r o m several localities i n southern L i m b u r g (e.g.
i n F u n g . rar. Ic. c o l . 2: p i . 9 c & d . 1967. C a d i e r en Keer, Riesenberg and Orenberg; E l s l o o , Bunderbos), Sept.-
S E L . D E S C R . & FIGS. - H u i j s m a n i n M e d e d . N e d . m y c o l . Vereen. 28: Oct. Widespread, but apparently not c o m m o n i n the warmer regions o f
23-25, fig. 4. 1943; M a i . & Bert., F l . C h a m p , sup. M a r o c 1: 128-129. Europe (absent from D e n m a r k and N o r w a y , but present o n the S w e d i s h
1970. Island o f 01and).
VERN. NAME - Olijfparasolzwam.
Lepiota olivaceobrunnea P . D . O r t o n lacks p i n k i s h tinges and is less
grey than t y p i c a l L. forquignonii, but it is i n a l l other characters iden-
tical to L. forquignonii; reason for B o n to change its status into a vari-
ety o f L. forquignonii. H e r e it i s subsumed i n the s y n o n y m y o f L.
forquignonii.
Var. coniferarum differs from t y p i c a l L. forquignonii i n the slight-
ly bigger spores (up to 8.0 p m ) and the darker colours o f the basidio-
carps. Whether these differences warrant status o n varietal level is
questionable.
11. Lepiota xanthophylla P . D . O r t o n i n Trans. B r . m y c o l . S o c . 4 3 :

289. 1960, non Lepiota xanthophylla S i n g . , 1973. - F i g . 89.
E X C L . - Lepiota xanthophylla sensu B e s l et a l . i n Z . M y k o l . 5 0 :
187-188. 1984; sensu Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 236.
1995; sensu M i g l . et a l . i n R i v . M i c o l . 32: 105-107. 1989; sensu M i g l .
F i g . 88. Lepiota forquignonii. & B i z z i i n M i c o l . ital. 26 (3): 57-58. 1997 (in a l l cases L. elaiophylla).
walls. Pleurocystidia absent. Pileus covering made up o f more or less

erect, long elements (more erect at pileus centre, more adnate towards
margin), (120-)150-400(-490) X 5.0-15 p m , flexuous i n lower part,
w i t h slightly thickened, y e l l o w - b r o w n w a l l s , b e c o m i n g thinner and
paler towards apex, also w i t h pigment soluble i n ammonia, intermixed
with pale coloured, narrowly clavate, clavate and spheropedunculate
elements, 22-80(-105) X (7.5-)10-25 u m , sometimes encrusted i n
lower part; lower l y i n g hyphae sometimes encrusted. S t i p i t i p e l l i s a
cutis o f colourless, c y l i n d r i c a l elements, 2.5-6.0 u m i n diameter, w i t h
velar hyphae i n lower part o f stipe, c y l i n d r i c a l , irregular and flexuous,
2.5-10 p m i n diameter, colourless, o r w i t h encrusting colourless p i g -
ments; squamules made up o f elements s i m i l a r to those o f pileus c o v -
ering, but shorter. Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - Gregarious and saprotrophic, terrestrial o n
humus-rich, nutrient-rich, clayey, probably calcareous soils, i n decid-
uous woods. I n the Netherlands very rare, o n l y k n o w n f r o m some
localities i n southern L i m b u r g (Valkenburg, Schaelsberg; C a d i e r en
Keer, Riesenberg and Orenberg; Margraten, Eijsderbos; Elsloo,
Bunderbos). Sept.-Oct. Scattered and rare i n temperate Europe, from
D e n m a r k southwards, also k n o w n from Japan.
F i g . 89. Lepiota xanthophylla.
The name Lepiota xanthophylla has been erroneously used for L. elaio-
M I S A P P L . - Lepiota citrophylla sensu B o u d . i n B u l l . S o c . m y c o l . Fr. phylla V e l l i n g a & Huijser, a toxic species g r o w i n g i n E u r o p e i n hot-
9: p i . 2, fig. 1. 1893; sensu R e a , B r i t . B a s i d i o m y c : 72. 1922. houses, w i t h more olive tinges i n the basidiocarps, a n d a different
S E L . I C O N . - Imaz. & H o n g o , C o l . 111. M u s h r . Japan 1: p i . 4 1 , fig. structure o f the pileus covering.
280. 1987; R a i d et a l . i n Svampe 26: 38. 1992; Tabares i n B o l e t s
Catalunya 10: p i . 480. 1991; V e l l i n g a & Huijser i n B o l l . G r u p p o m i c o l . 12. Lepiota echinella Q u e l . & B e r n , i n B u l l . S o c . m y c o l . Fr. 4: L I .
G . Bres., n.S. 4 0 (2-3): fig. 1. ('1997') 1998. 1888 (as Lepoita echinellus). - F i g . 90.
S E L . D E S C R . & FIGS. - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 124- Lepiota minuta I. L a n g e i n D a n s k bot. A r k . 4 (4): 4 8 . 1923, n o n
125. 1994; P . D . O r t o n i n Trans. B r . m y c o l . S o c . 4 3 : 289-290. 1960; Lepiota minuta V o g l . , 1896; Lepiota setulosa J . Lange, F l . agar. dan. 1:
R a i d et a l . i n Svampe 26: 38. 1992; D . R e i d i n F u n g rar. Ic. c o l . 3: 10- 3 4 . 1 9 3 5 . - Lepiota rhodorhiza (Romagn. & L o c q . ) L o c q . i n B u l l . mens.
12, f i g . 6. 1968; V e l l i n g a & Huijser i n B o l l . G r u p p o m i c o l . G . Bres., Soc. linn. L y o n 14: 98. 1945 (not valid); Lepiota rhodorhiza (Romagn.
n.S. 4 0 (2-3): 458-460, fig. 2. ( ' 1 9 9 7 ' ) 1998. & L o c q . ) ex P.D. Orton i n Trans. B r . m y c o l . Soc. 43: 285. 1960.
VERN. NAME - Geelplaatparasolzwam. E X C L . - Lepiota echinella sensu H u i j s m a n i n M e d e d . N e d . m y c o l .
Vereen. 28: 22. 1943 ( = Lepiota spec.)
Pileus 10-30(-40) m m , when y o u n g hemispherical, campanulate w i t h VERN. NAME - Piekhaarparasolzwam.
inflexed margin, expanding to plano-convex w i t h u m b o and undulat- A l t h o u g h numerous species i n the v i c i n i t y o f Lepiota echinella
ing l i m b , and deflexed margin, at centre b r o w n ( M u . 10 Y R 3/4) w i t h have been described, it is still possible to make a c o l l e c t i o n w h i c h does
pointed fibrillose squamules, around centre paler b r o w n (10 Y R 5/6) not fit i n any o f them. After an o v e r v i e w o f this species complex, t w o
with more adnate fibrils, at outermost margin revealing the y e l l o w c o n - varieties o f L. echinella are described.
text between them, w h e n y o u n g w i t h some y e l l o w i s h velar remnants There appears to be huge variation i n the m a i n characters used to
at m a r g i n . L a m e l l a e , L = c. 6 0 , 1 = 1-3, moderately c r o w d e d , dis- distinguish species i n the v i c i n i t y o f Lepiota echinella: spore size,
tinctly free, subventricose to 5 m m broad, y e l l o w (more green than 5 shape o f the cheilocystidia and the length and structure o f the elements
Y 8/6) w i t h pale uneven edge. Stipe 35-60 X 3-5 m m , slightly broad- of the pileus covering. Short characteristics o f the existing speciesare
ening downwards, h o l l o w , pale sulphur-yellow (slightly more greenish given i n the f o l l o w i n g overview, w h i c h i s a c o m p i l a t i o n o f literature
than 5 Y 8/6-8, resembling K . & W . 1 A 5 ) , at apex slightly fibrillose, on the existing names; the problems i n interpretation o f some names
b e l o w irregularly w o o l l y , i n l o w e r h a l f o r at base o n l y w i t h s m a l l , are outlined. B o n ( F l . m y c o l . Eur. 3, Lepiotes: 6 8 - 7 7 . 1 9 9 3 ) i n his treat-
incomplete bands o f b r o w n squamules o n heightened y e l l o w w o o l l y ment o f this group, emphasized c o l o u r o f the basidiocarps, w h i l e here
background, without real annulus, but w i t h a l o w r i m . Context y e l - priority is given to m i c r o s c o p i c a l characters.
l o w i s h cream i n pileus, i n stipe y e l l o w i s h to distinctly y e l l o w at base,
w h i t i s h fibrillose around cavity. S m e l l o f intact basidiocarps indistinct,
slightly astringent, w h e n basidiocarp cut strong, like L. cristata. Taste
not k n o w n . Spore print white (when fresh).
Spores (6.0-)6.5-9.0(-10,0) X (3.5-)4.0-5.0 u m , Q = (1.4-)1.6-2.0,
Qav = 1.7-1.9, oblong, often w i t h less convex adaxial than abaxial side
i n side-view, g i v i n g the spores a slight a m y g d a l i f o r m appearance,
oblong i n frontal view, dextrinoid, congophilous, not metachromatic i n
C r e s y l B l u e . B a s i d i a (17-)20-30 X (6.5-)7-9.5(-10,5) u m , mostly 4-
spored, but often also some (3-)2(-l) spored basidia present. L a m e l l a
edge sterile. C h e i l o c y s t i d i a (15-)20-35(-40) X (5.0-)6.5-12(-14.5) u m ,
often clustered, n a r r o w l y lageniform, n a r r o w l y utriform, narrowly
clavate to c y l i n d r i c a l , rarely clavate, hyaline w i t h slightly thickened F i g . 90. Lepiota echinella.
Lepiota lepida G u i n b . & B o d i n (in D o c . m y c o l . 23 (92): 6 1 . 1994) K e l d e r m a n (Parasolzw. Z u i d - L i m b u r g : 115. 1994) gave a description
comes v e r y c l o s e to L. echinella; the spores are slightly bigger (on of such a c o l l e c t i o n .
average 6.6-7.0 X 3.8-4.3 um), and the shape o f the c h e i l o c y s t i d i a is Lepiota speciosa (Trimb.) T r i m b . & A u g i a s (in T r i m b a c h i n D o c .
m o r e varied than i n L. echinella: not o n l y lageniform but also more m y c o l . 18 (72): 50. 1988), first described as a variety o f L. rhodorhiza
c y l i n d r i c a l c y s t i d i a occur; a septum might occur i n the pileus covering ( T r i m b a c h i n D o c . m y c o l . 5 (20): 4 5 . 1975), differs f r o m L. echinella
elements, w h i c h are l o n g and slender. Lepiota bickhamensis P.D. O r t o n i n the b i g basidiocarps (stipe up to 90 X 6 m m ) , w i t h copious v e i l o n
(in N o t e s R . bot. G d n E d i n b . 4 1 : 588. 1984) is probably identical w i t h the stipe, the b i g spores (6-7.5(-8) X 3-4.5 pm), and the clavate to
L. lepida, though it lacks septa i n the pileus c o v e r i n g elements. lageniform c h e i l o c y s t i d i a .
Basidiocarps have been found, fitting L. lepida as regards pileus cover-
ing elements and spore size, but with c h e i l o c y s t i d i a that are K E Y TO THE VARIETIES
predominantly clavate, and o n l y a few being widest i n the middle. 1. Stipe without any (red to vinaceous p i n k ) strigae at base
Lepiota clypeolarioides R e a (Brit. B a s i d i o m y c : 69. 1922) is 12a. var. echinella
m a c r o s c o p i c a l l y different f r o m a l l other species i n this group, resem- 1. Stipe red to vinaceous p i n k strigose at base . . . 12b. var. rhodorhiza
b l i n g L. clypeolaria, but w i t h e l l i p s o i d spores. A modern photograph
o f its var. armillata B o n et al. is p r o v i d e d by M i g l i o z z i & C h e r u b i n i 12a. var. echinella
(in B o l l . A s s . m i c o l . e c o l . R o m a n a 17: 27. 1989). T h i s species is sub- E X C L . - Lepiota setulosa sensu K u h n e r i n B u l l , trimest. Soc. m y c o l .
ject to various interpretations. K u h n e r (in B u l l , trimest. Soc. m y c o l . Fr. Fr. 52: 224-226. 1936 ( = var. rhodorhiza).
52: 228-229. 1936) described specimens f r o m A l g e r i a . H u i j s m a n (in S E L . I C O N . - M i g l . & C o c c i a i n M i c o l . ital. 19 (3): 9. 1990.
M e d e d . N e d . m y c o l . Vereen. 28: 25-27. 1943) used the name L. clype- S E L . D E S C R . & F I G S . - M i g l . & C o c c i a i n M i c o l . ital. 19 (3): 6-8, f i g .
olarioides for an a l c o h o l c o l l e c t i o n o f L. felina, and named K u h n e r ' s 2. 1990.
c o l l e c t i o n L. kuehneri. A s this was not a v a l i d description o f a new
species, H o r a (in Trans. B r . m y c o l . Soc. 43: 448. 1960) described L. P i l e u s 5-28 m m , convex to plano-convex, applanate w i t h l o w umbo,
kuehneri w i t h a L a t i n diagnosis and designated a type c o l l e c t i o n , sometimes slightly undulating around centre, at centre dark
w h i c h got lost. B o n (in D o c . m y c o l . 26 (102): 21-23. 1996) designate (red)brown ( M u . 7.5 Y R 3/2, 4/4), and p l u s h - l i k e w i t h little upright
ed Kuhner's c o l l e c t i o n (actually made by M a i r e ) as neotypc- tufts, around centre gradually breaking up into s m a l l granular patch-
Unfortunately, B o n ' s measurements o f the spores o f this neotype ((4.5- es, or s m a l l squamulose floccules, c o n c o l o r o u s w i t h centre, or paler
)5-6(-7) X (3-)4-4.5(-5) pm) differ considerably f r o m the present towards m a r g i n (e.g. 7.5 Y R 5/4), o n c r e a m to pale p i n k i s h back-
author's (6.0-7.5 X 3.5-4.0 p m , a v l X avw = 6.6 X 3.8 pm). K u h n e r ground. L a m e l l a e , L = 3 0 - 3 5 , 1 = 1-3, moderately distant to c r o w d -
(and H o r a for the B r i t i s h type) gave as spore sizes intermediate values: ed, free, segmentiform to ventricose, up to 2 mm wide,
5.5-7 X 3.2-4 p m . Lepiota kuehneri might w e l l be conspecific w i t h L. cream-coloured, rarely creamy white, w i t h white or concolorous erod-
lepida, but perhaps is better considered a nomen d u b i u m . ed to flocculose edge. Stipe 17-35 X 1-3 m m , c y l i n d r i c a l , sometimes
Lepiota coxheadii P.D. O r t o n (in Notes R . bot. G d n E d i n b . 4 1 : 589. s l i g h t l y w i d e n i n g towards subbulbous base, fistulose, w h i t i s h f i b r i l -
1984 (as L. coxheadi)) is characterized by spores 6.0-7.5 X 3.5-4.0(- lose i n upper part, l o w e r d o w n m o r e irregularly w o o l l y fibrillose and
4.5) p m , Q = 1.6-1.9, Q a v = 1.71, narrowly lageniform to c y l i n d r i c a l - more; p i n k i s h b r o w n , without real annulus, but at base sometimes w i t h
clavate c h e i l o c y s t i d i a , and i n contrast to L. echinella, L. lepida and L. some bands o f squamules, concolorous w i t h pileus. C o n t e x t white i n
bickhamensis, relatively short elements i n the pileus covering (80-135 pileus, i n stipe c o n c o l o r o u s w i t h surface and r e d d i s h i n l o w e r part.
X 10-14 p m ) . Specimens w i t h short spores (5.5-6.5 X 3.0-4.0 p m , avl S m e l l very variable, indistinct, fungoid, unpleasant or l i k e the s m e l l
X avw = 6.0 X 3.4 pm), and relatively short elements i n the pileus of L. cristata. Taste as s m e l l . Spore print p r o b a b l y white.
covering have been encountered i n the Netherlands, but whether they Spores 4.5-6.5 X 3.0-4.0 p m , Q = 1.4-2.0, Q a v = 1.55-1.8, e l l i p -
b e l o n g to L. coxheadii is not clear, as colours arid sizes o f the basidio- s o i d to o b l o n g i n side-view, i n maturity dextrinoid, c o n g o p h i l o u s ,
carps do not fit w i t h the concept o f L. coxheadii. cyanophilous, and not metachromatic i n C r e s y l B l u e . B a s i d i a 15-23 X
Lspiota favrei K u h n e r ex M . B o n (in D o c . m y c o l . 22 (88): 29. 6.0-8.0 pm, predominantly 4-spored. Lamella edge sterile.
1993) is a pale, s m a l l species w i t h spores 6-8.5 X 4-5 p m and C h e i l o c y s t i d i a 15-33 X 6-12 p m , w i t h 2.0-5.5(-6.5) p m w i d e neck,
' m u m m y - s h a p e d ' cheilocystidia, described f r o m alpine habitats o n cal- lageniform to utriform, w i t h short to l o n g neck, hyaline and slightly
careous s o i l (for a full description see Favre, C h a m p , sup. Z o n e alpine: t h i c k - w a l l e d . P l e u r o c y s t i d i a not observed. P i l e u s c o v e r i n g made up
157-158, f i g . 143. 1955 (as L. pseudohelveola)). erect, l o n g , c y l i n d r i c a l elements i n tufts, 185-300 X 8-12.5 p m and
Lepiota glareophylla F i l l i o n (in B u l l . F e d . m y c o l . D a u p h . Savoie short, narrowly clavate to c y l i n d r i c a l elements, 15-46 X 8.0-10 at base;
144: 7. 1997) is a species w i t h very s m a l l basidiocarps (pileus 8-12 repent c o l o u r e d hyphae absent. P i g m e n t especially situated i n l o w e r
mm), pale y e l l o w - b r o w n colours and darker b r o w n squamules, b i g part o f l o n g elements, and clavate elements, dark b r o w n , parietal,
spores (11-12(-12.5) X 6.0-7.0(-7.5) pm) and a pileus c o v e r i n g made encrusting sometimes i n l o w e r parts o f l o n g elements and u n d e r l y i n g
up o f relatively short and w i d e elements o f a l l lengths ((35-)50-120(- hyphae. Stipitipellis a cutis o f narrow, 2-4 p m w i d e c y l i n d r i c a l hyphae,
160) X 10-15(-20) pm). w i t h i n l o w e r part l o o s e l y attached, irregular hyphae w i t h b r o w n p a r i -
A n o t h e r s m a l l species w i t h a rather irregular pileus c o v e r i n g is L. etal pigment. C l a m p - c o n n e c t i o n s present i n a l l tissues.
locquinii M . B o n (in D o c . m y c o l . 16 (61): 20. 1985); the pileus meas- H A B I T A T & DISTR. - Solitary, or i n s m a l l groups, saprotrophic and
ures 1-2 c m , and the spores are smaller than i n L. glareophylla: 6.5-8.0 terrestrial i n deciduous w o o d s , o n humus-rich, nutrient-rich sandy to
X 3.5-4.5 p m ; the layers o f the pileus covering are not clearly sepa- clayey soils. In the Netherlands relatively rare, o c c u r r i n g i n the coastal
rated causing the irregular l o o k (the sketch given by B o n ( F l . m y c o l . dune area, southern L i m b u r g and some scattered localities, Sept.-Oct.
Eur. 3, L e p i o t e s : 69. 1993) is m i s l e a d i n g i n this respect). Specimens Scattered, rare and widespread i n E u r o p e .
w i t h such an irregular pileus c o v e r i n g have been encountered i n the \
Netherlands, but they were m u c h bigger and darker b r o w n or more Lepiota echinella var. echinella forms rather s m a l l basidiocarps, w h i c h
p i n k c o l o u r e d than L. locquinii i n the o r i g i n a l description, and the often grow solitary. T h u s , the chance o f o v e r l o o k i n g this taxon i n the
c h e i l o c y s t i d i a were l a g e n i f o r m (instead o f clavate to flexuous). field is rather great.
12b. var. rhodorhiza (P.D. Orton) Hardtke & R o d e l i n M y k o l . M i t t b l . ceous red-brown, and around centre w i t h concentrical zones o f squa-
35: 62. 1992. mules, slightly paler or concolorous w i t h centre ( M u . 7.5 Y R 3/2-5/2,
Lepiota setulosa var. rhodorhiza Romagn. & Locq. i n Locq. in 2.5 Y R 3/4-4/4, 3/6-4/6), rarely also w i t h centre cracked into squamules,
B u l l , trimest. S o c . m y c o l . Fr. 60: 38. 1944 (not valid). on whitish, cream to vinaceous p i n k or p i n k i s h buff background; squa-
M I S A P P L . - Lepiota setulosa sensu K i i h n e r i n B u l l , trimest. S o c . mules and centre tomentose, often made up o f small pyramids.
m y c o l . Fr. 52: 224-226. 1936. Lamellae, L = 45-55,1 = 1-3, rather to moderately crowded, free, ven-
S E L . I C O N . - L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l . G . Bres. tricose up to 7 m m wide, white to cream, when o l d often w i t h brownish
26: 109. 1983. red spots, with white, slightly eroded edge. Stipe 25-70 X 5-9 m m , cylin-
S E L . D E S C R . & FIGS. - K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 5 2 : drical or slightly broadened o r attenuate at base, fistulose, i n apical half
224-226. 1936 (as L. setulosa); M i g l . & C o c c i a i n M i c o l . ital. 19 (1): cream to pinkish cream to brownish-pinkish, lengthwise whitish fibril-
13-15, figs. 1 & 2. 1990 (as L. rhodorhiza); P . D . O r t o n i n Trans. B r . lose, i n lower half w i t h discrete squamules, forming incomplete girdles,
m y c o l . S o c . 4 3 : 285-286. 1960. concolorous with squamules on pileus on pinkish to brownish-vinaceous
pink background, w i t h white m y c e l i u m strands at base. Context white to
C H A R A C T E R I S T I C S - Differs from the typical variety i n the sometimes whitish i n pileus, y e l l o w i s h to glassy red-brown i n cortex o f stipe, i n
more robust basidiocarps (pileus up to 4 0 m m , stipe up to 65 X 5 m m ) , inner part o f stipe white and dull. S m e l l fruity-aromatic to more fungoid.
and the presence o f a vinaceous p i n k to red strigose tomentum at base Taste fungoid or unpleasant: rancid and astringent. Spore print 'white'.
of stipe. Spores i n side-view 7.0-10.0(-l 1.0) X 4.0-5.0(-6.0) p m , Q = (1.5-
M i c r o s c o p i c a l characters as i n the typical variety. )1.6-2.05(-2.15), Q a v = 1.7-1.9, oblong, slightly a m y g d a l i f o r m w i t h
H A B I T A T & DISTR. - Gregarious i n s m a l l groups, saprotrophic and straight adaxial side, i n frontal v i e w o b l o n g , t h i c k - w a l l e d , strongly
terrestrial o n humus-rich, mostly sandy soils, i n deciduous woods or dextrinoid, congophilous; w a l l not c o l o u r i n g i n C r e s y l B l u e . B a s i d i a
avenues w i t h deciduous trees. I n the Netherlands very rare, k n o w n 24-42 X 6.5-10.0 u m , predominantly 4-spored, also intermixed 2- and
from a few localities (for instance A r c h e m near O m m e n , and 4-spored (hence longer spores). L a m e l l a edge sterile. C h e i l o c y s t i d i a
H o l l a n d s e R a d i n g ) , Sept.-Oct. I n E u r o p e rare, and widespread. irregular, often septate, 14-37 X 4.0-10 p m , c y l i n d r i c a l , narrowly
clavate, or narrowly utriform. P l e u r o c y s t i d i a not observed. Pileus c o v -
ered w i t h elements i n squamules; elements 90-285 X 8.0-14 um, not
septate, sometimes w i t h attenuate apex w i t h thickened w a l l ; w i t h
Subsect. Helveolinae B o n & Boiffard brown parietal pigment; underlayer o f c y l i n d r i c a l twisting c y l i n d r i c a l
hyphae w i t h narrowly clavate terminal elements, w i t h b r o w n parietal
Pileus covering made up o f l o n g , erect elements, l a c k i n g short, clavate pigment. Stipitipellis a cutis o f c y l i n d r i c a l , non-coloured, 3.0-8.0 u m
elements. wide hyphae; elements o f squamules l i k e those o n pileus. C l a m p - c o n -
nections present i n a l l tissues.
13. Lepiota brunneoincarnata C h o d . & M a r t , i n B u l l . S o c . bot. H A B I T A T & DISTR. - Gregarious or more rarely solitary, terrestrial i n
Geneve 5: 222. 1889. - F i g . 9 1 . m i x e d deciduous woods, m a i n l y o n soils r i c h i n nutrients, w h i c h are
S E L . I C O N . - K o b l e r i n S c h w e i z . Z . P i l z k . 66: opp. p . 6. 1988; J . rather clayey or loamy. In the Netherlands rare and scattered (for a dis-
Lange, F l . agar. dan. 1: p i . 13F. 1935 (rather red-brown). tribution map, see A r n o l d s et a l . , O v e r z . Paddest. N e d e r l a n d : 2 8 8 .
S E L . D E S C R . & FIGS. - D e r b s c h & Schmitt, A t l . P i l z e Saarlandes 2: 1995); June-Oct., but mostly recorded i n the summer months.
490. 1987; K o b l e r i n S c h w e i z . Z . P i l z k . 66: 6. 1988; S c h u l z - W e d d i g e n Widespread i n Europe, but everywhere rare, absent f r o m the boreal,
in Z . M y k o l . 5 2 : 9 1 - 1 0 0 . 1986. (sub)alpine and arctic regions.
V E R N . N A M E - Gegordelde parasolzwam.
Lepiota brunneoincarnata is a very t o x i c species, causing, just as

Pileus 25-55 m m , when young conico-convex, w i d e l y campanulate or Amanita phalloides, severe damage to the liver, w h i c h c a n result i n
broadly obtusely conical, expanding to plano-concave or applanate, w i t h death (e.g. F u r i a et al. i n M i c o l . ital. 11 (3): 29-33. 1982; H e r m a n n i n
low broad u m b o o r plano-convex w i t h u m b o and deflexed margin, at M y k o l . M i t t b l . 3 0 : 73-76. 1987, a n d S c h u l z - W e d d i g e n i n Z . M y k o l .
centre dark greyish-brownish to nearly black or very dark to dark vina- 52: 91-100. 1986).
14. Lepiota subincarnata J . Lange, F l . agar. dan. 5: V. 1940. - Fig. 92.

Lepiota josserandii B o n & B o i f f . i n B u l l , trimest. S o c . m y c o l . Fr.
90: 2 8 9 . ( ' 1 9 7 4 ' ) 1975; Leucoagaricus josserandii ( B o n & Boiff.)
Raithelhuber i n M e t r o d i a n a 17: 7 5 . 1989; Lepiota subincarnata var.
josserandii ( B o n & Boiff.) G m i n d e r i n Beitr. K e n n t n . P i l z e Mitteleur.
12: 69. 1999. - Lepiota josserandii var. rosabrunnea Raithelhuber i n
M e t r o d i a n a 16: 42. 1988.
EXCL. - Lepiota subincarnata sensu Johnson & Vilgalys in
M y c o l o g i a 90: 974-976. 1998 ( = Lepiota spec).
M I S A P P L . - Lepiota helveola sensu Joss, i n B u l l , trimest. S o c .
m y c o l . Fr. 47: 65-66. 1931.
S E L . I C O N . - J . L a n g e , F l . agar. dan. 1: p i . 131. 1935; R . P h i l l i p s ,
Paddest. S c h i m m . : 29. 1981; R a i d et a l . i n Svampe 26: 39. 1992.
SEL. D E S C R . & F I G S . - M . B o n i n D o c . m y c o l . 2 0 (78): 4 8 , figs. 21-
L. 1990; D e r b s c h & Schmitt, A t l . P i l z e Saarlandes 2: 4 9 2 . 1987;
Enderle & Krieglst. i n Z . M y k o l . 55: 9 4 - 9 6 . 1 9 8 9 ; Esteve-Rav. & A l t e s
F i g . 9 1 . Lepiota brunneoincarnata. in B o l . Soc. micol. Madrid 14: 162-163, figs. 1-3. 1990 (as L.
ed i n lower part, w i t h parietal pigment. T h e longer elements present at

centre o f pileus, the smaller ones towards margin. U n d e r l a y e r w e l l -
developed, made u p o f twisted, c y l i n d r i c a l , repent elements, 4.0-5.0
u m wide, w i t h parietal, b r o w n pigment. Stipitipellis a cutis o f c y l i n -
drical elements, 3.0-6.0 p m i n diam., w i t h encrusting pale pigments;
squamules made up o f elements as o n pileus, but slightly shorter and
wider. Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - Gregarious i n small groups, saprotrophic and
terrestrial on humus- or nutrient-rich sandy to clayey soils, i n city-parks,
gardens, copses, deciduous or m i x e d woods and also o n m i n e waste
heaps, rather c o m m o n i n the Netherlands, A u g . - N o v . Widespread i n
Europe, also recorded from N o r t h A m e r i c a .
Lepiota subincarnata is, like L. brunneoincarnata and L. elaiophylla,

very toxic (Gerault & G i r r e i n C . r. hebd. Seanc. A c a d . S c i . , Paris, Ser.
D , 280: 2842. 1966), because o f the presence o f amanitins; i t also c o n -
tains amatoxins.
15. Lepiota elaiophylla V e l l i n g a & Huijser i n B o l l . G r u p p o m i c o l . G .

F i g . 92. Lepiota subincarnata. Bres., n.S. 4 0 (2-3): 462. ('1997') 1998.
M I S A P P L . - Lepiota xanthophylla sensu B e s l et a l . i n Z . M y k o l . 5 0 :
187-188. 1984; sensu Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 236.
josserandii); G m i n d e r i n Beitr. K e n n t n . P i l z e Mitteleur. 12: 68-70, figs. 1995; sensu M i g l . et a l . i n R i v . M i c o l . 32: 105-107. 1989; sensu M i g l .
6-10. 1999 (as L. subincarnata and L. subincarnata var. josserandii & B i z z i i n M i c o l . ital. 26 (3): 57-58. 1997.
resp.) K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 104-105. 1994; Kuhner*; S E L . I C O N . - Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 236. 1995;
in B u l l , trimest. S o c . m y c o l . Fr. 52: 223-224. 1936. M i g l . et al. i n R i v . M i c o l . 32: 106. 1989; M i g l . & B i z z i i n M i c o l . ital.
V E R N . N A M E - Vaalroze parasolzwam. 26 (3): p i . 3. 199.7 (all as L. xanthophylla); V e l l i n g a & Huijser i n B o l l .
G r u p p o m i c o l . G . Bres., n.S. 4 0 (2-3): fig. 3. ('1997') 1998.
Pileus 17-42(-60) m m , when young hemispherical w i t h inflexed margin, S E L . D E S C R . & FIGS. - V e l l i n g a & Huijser i n B o l l . G r u p p o m i c o l . G .
expanding to plano-convex or even plano-concave w i t h broad l o w umbo Bres., n.S. 4 0 (2-3): 462-463, fig. 4. ('1997') 1998.
and straight margin, at centre vinaceous pink, pinkish red, reddish brown VERN. NAME - Olijfplaatparasolzwam.
or grey-brown to pink-brown ( M u . 2.5 Y R 4/4, 5 - 7.5 Y R 4/4, 5/5, 5/4
or 7/4), paler towards margin (7.5 Y R 8-7/4, 7/6; 5 Y R 8/3-4, 7/6), floc- C H A R A C T E R I S T I C S - P i l e u s 14-33 m m , w i t h age expanding to plano-
culose or flocculose-squamulose at centre, towards margin either corti-^ convex w i t h or without l o w u m b o , w i t h central velutinous to subto-
pletely tomentose or split up i n small, appressed floccose, concentrically mentose dark orange-brown ( M u . 10 Y R 3/4) or reddish-ochraceous
or more irregularly arranged, patches or squamules o n a cream or pale b r o w n patch, around the patch breaking up into concentrical, s m a l l ,
p i n k i s h cream background, w h e n young w i t h white velar remnants at arachnoid-subtomentOjSe squamules slightly paler than at centre, o n
margin. L a m e l l a e , L = 40-80,1 = 0-3, moderately distant to crowded, pale olivaceous y e l l o w to pale sulphur-yellow background; lamellae, L
free, ventricose or subventricose, up to 7 m m broad, white to cream w i t h = c. 50,1 = 1-3, moderately distant to crowded, free, ventricose, olive-
age, w i t h concolorous to white, entire or finely granular edge. Stipe 15- yellow, greyish-olivaceous y e l l o w (5 Y 6/4; K . & W . 3 A 4 - 3 B 4 , later
55 X 1.5-7(-14) m m , cylindrical or slightly broadened at base, fistulose 3 C 5 - 4 C 5 ) , less grey towards edge; edge y e l l o w and slightly eroded; \
with age, i n upper half lengthwise innately striate, pale, cream or pink- stipe 32-40 X 2.5-3 m m , c y l i n d r i c a l , h o l l o w , i n upper part subglabrous
ish (5 Y R ' 8 / 3 ) , more or less halfway w i t h girdle of annulus, concolorous to lanate-fibrillose and olive-yellow, w i t h a b r o w n r i n g - l i k e zone, and
with pileus and lower d o w n provided w i t h more girdles or incomplete below this zone w i t h bands o f s m a l l b r o w n i s h squamules, as on pileus,
bands, made u p o f floccules o n a cream, fibrillose background, w i t h on more lanate covering; context pale sulphur-yellow to o l i v e - y e l l o w
white m y c e l i u m cords at base. Context i n pileus white, slightly p i n k brownish i n pileus (paler than 5 Y 8/4), darker o l i v e - y e l l o w (5 Y 6/4)
coloured under surface at centre, dull, i n stipe shiny and beige, beige- in stipe; s m e l l unpleasantly fungoid; spore print y e l l o w - c r e a m
p i n k or brownish i n cortex, i n inner part white and cottony. S m e l l always (Breitenbach & K r a n z l i n , P i l z e S c h w e i z : p i . 236. 1995).
with a fruity component and a fungoid or farinaceous component. Taste Spores 6.5-9.5 X 3.5-4.5 p m , Q = 1.75-2.25(-2.4), Q a v = 1.9-2.2,
unpleasant, rather astringent at back o f throat. Spore print white. oblong, s u b c y l i n d r i c a l i n side-view, dextrinoid, congophilous, not
Spores i n s i d e - v i e w (5.5-)6.0-7.5(-8.0) X 3.0-4.0(-4.5) u m , Q = metachromatic i n C r e s y l B l u e ; basidia 18-35 X 6.0-9.5 u m , 4-spored;
(1.45-)1.5-2.0(-2.3), Q a v = 1.65-1.85, o b l o n g or c y l i n d r i c a l w i t h c h e i l o c y s t i d i a 15-40(-48) X 5.0-12 p m , narrowly clavate to clavate,
rather straight sides, sometimes uniguttulate, dextrinoid, congophilous, disappearing w i t h age; pileus covering made up o f more or less erect
strongly cyanophilous, not metachromatic i n Cresyl Blue; hilar elements, 60-265 X 6.5-18 p m , w i t h b r o w n i s h parietal and intracellu-
appendage inconspicuous. B a s i d i a 17.5-30 X 5.0-8.5 p m , 4-spored, lar pigments, l a c k i n g a layer o f clavate, short elements; c l a m p - c o n -
rarely 4 - and 2-spored intermixed. L a m e l l a edge sterile. C h e i l o c y s t i d i a nections present i n a l l tissues.
usually abundant, rarely inconspicuous and not easily found, 11-37 X H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial,
5.0-10 u m , c y l i n d r i c a l , narrowly clavate, clavate, some septate, thin- until now o n l y k n o w n from greenhouses i n Europe together w i t h c u l -
or slightly t h i c k - w a l l e d , colourless. Pleurocystidia absent. Pileus c o v - tivated plants; i n the Netherlands k n o w n from pots w i t h Spathiphyllum
ered w i t h erect elements, 70-400 X 6.0-16 u m , sometimes not sur- i n a c o m m e r c i a l greenhouse (Uithoorn), and from a hothouse i n a
passing 200 um; elements c y l i n d r i c a l without attenuate apex or more botanical garden (Leiden). A u g . - O c t .
fusiform w i t h attenuate apex, narrower i n basal part, straight, or twist-
European authors called this taxon Lepiota xanthophylla (or L. citro- S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 25b. 1990.
phylla), but the former i s a species w i t h browner pileus, more y e l l o w S E L . D E S C R . & FIGS. - K e l d e r m a n i n C o o l i a 34: 52-53, fig. 2. 1991;
lamellae, and w i t h a layer o f clavate elements i n the pileus covering K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 218-219. 1936.
(see V e l l i n g a & Huijser i n B o l l . G r u p p o m i c o l . G . Bres., n.S. 4 0 (2-3): V E R N . N A M E - Witte dwergparasolzwam.
460-462. ('1997') 1998. for a discussion o n this species), and L. citro-
phylla has spurred spores (Pegler, A g a r i c F l . S r i L a n k a : 305. 1986). Pileus 5-25 m m , convex w i t h l o w u m b o , ochraceous w i t h reddish
Lepiota elaiophylla is h i g h l y toxic, and contains a-amanitin and B- tinge at centre, cream-coloured to white at m a r g i n , completely c o v -
amanitin ( B e s l et a l . i n Z . M y k o l . 50: 190. 1984). ered i n woolly-tomentose fibrils, w i t h some toothed v e l u m remnants
at margin. L a m e l l a e , L = 20-28, 1 = 1-3, c r o w d e d , free, subventri-
16. Lepiota rubella B r e s . i n P. H e n n . i n V e r h . bot. Ver. Prov. cose, white to slightly cream-coloured, w i t h even concolorous edge.
Brandenb. 3 1 : 149. 1890. Stipe 11-25 X 0.7-3 m m , c y l i n d r i c a l o r slightly broadened at base, fis-
Lepiota bettinae Dorfelt i n Z . M y k o l . 48: 246. 1982; Echinoderma tulose, white, w i t h annulus, b e l o w annulus w i t h some ochraceous fib-
bettinae (Dorfelt) M . B o n i n D o c . m y c o l . 2 2 (88): 28. 1993. rils o n white to cream background. A n n u l u s ascending, white, fragile
E X C L . - Lepiota rubella sensu M . B o n , F l . m y c o l . Eur. 3, Lepiotes: and membranaceous. Context white, i n basal part o f stipe b e c o m i n g
81. 1993 ( = Lepiota spec). b r o w n i s h w i t h age.
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 215. 1995 (as Spores 3.5-4.5 X 2.0-3.0 p m , Q a v = 1.7, non-dextrinoid, not c o n -
L . aff. bettinae); M i g l . & C o c c i a i n B o l l . A s s . m i c o l . e c o l . R o m a n a 18: gophilous, not metachromatic i n C r e s y l B l u e , uninucleate. B a s i d i a 14-
7. 1990 (as L. bettinae). 20 X 5.0-6.6 um, 4-spored. L a m e l l a edge fertile. C h e i l o c y s t i d i a absent.
S E L . D E S C R . & F I G S . - B a b o s i n A g a r i c a 6 (12): 204-205. 1985; Pleurocystidia absent. Pileus covering made up o f repent to ascending
C o n t u i n M i c o l . ital. 15 (3): 5 2 - 5 3 . 1 9 8 6 ; Dorfelt i n Z . M y k o l . 48: 245- hyphae w i t h terminal elements non- or up to 2-septate (without clamp-
251. 1982; M i g l . & C o c c i a i n B o l l . A s s . m i c o l . e c o l . R o m a n a 18: 5-11. connections at the septa), 170-300 X 5-6.5 p m , without pigment.
1990 (all as L. bettinae); V e l l i n g a et al. i n S y d o w i a 50: 272-277. 1998. Clamp-connections present i n a l l tissues.
V E R N . N A M E - R o d e dwergparasolzwam. H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres-
trial i n a nutrient-rich, relatively w a r m part o f a w o o d e d slope. I n the
C H A R A C T E R I S T I C S - P i l e u s 6-10 m m , rounded c o n i c a l w h e n y o u n g , Netherlands very rare, recorded o n l y once f r o m one locality i n south-
expanding to plano-concave, often w i t h s m a l l umbo, set with hairy l i l a - ern L i m b u r g (Valkenburg, Schaelsberg), Sept. K n o w n from only a very
cinous-purplish squamules, densely set at centre, more scattered near few finds throughout central and mediterranean Europe.
margin, o n whitish background, w i t h fimbriate margin; lamellae, L =
23-27,1 = 3(-7), rather crowded, free, i v o r y white and b e c o m i n g y e l - The above description has been based o n the selected descriptions cited.
l o w i s h , w i t h w h i t i s h edge; stipe 8-14 X 0.7-1 m m , tapering towards T h e spores o f this taxon are uninucleate, w h i l e most Lepiota-
apex, w h i t i s h at apex, p i n k i s h - l i l a c i n o u s to slightly b r o w n i s h at base, species have binucleate spores. T h e absence o f c h e m i c a l reaction o f
entirely pubescent, rarely w i t h floccose dark purplish annular zone, i n the spore w a l l i n M e l z e r ' s Reagent and C o n g o R e d is also a striking
lower part w i t h scattered minute p u r p l i s h b r o w n f l o c c u l i ; context character o f this species.
whitish i n pileus, brownish-lilacinous i n stipe; smell o f bruised basid- Lepiota nigrescentipes G . Riousset (in B o n & G . Riousset i n D o c .
iocarps a c i d spermatic. m y c o l . 22 (85): 67-68. 1992) comes very close to L. parvannulata, has
Spores 3.5-5.0 X 2.5-3.0 p m , Q = 1.6-1.8(-2.0), Q a v = 1.7, oblong also uninucleate spores w i t h the same k i n d o f c h e m i c a l reactions, but
i n side-view and i n frontal view; basidia 11-14 X 4.5-6.5 p m , 4-spored; differs i n the stipe w h i c h blackens w i t h age.
l a m e l l a edge sterile; c h e i l o c y s t i d i a 15-22 X 5.0-6.5 p m , narrowly Lepiota parvannulata resembles white Leucoagaricus-specits like
clavate; pleurocystidia absent; pileus c o v e r i n g made up o f very thick- L. serenus (Fr.) B o n & Boiffard and L. sericifer ( L o c q . ) V e l l i n g a , but
walled, b r o w n hyphae, 5-7 p m wide, w i t h w a l l s u p to 2 p m thick, with the presence o f numerous clamp-connections makes it u n l i k e l y that
terminal elements hardly w i d e n e d to distinctly clavate, u p to 17 p m this taxon belongs to that genus. Spores o f Leucoagaricus-species are
w i d e ; clamp-connections present i n a l l tissues. binucleate.
H A B I T A T & DISTR. - Gregarious i n b i g groups, saprotrophic and ter-
restrial, i n hothouses. K n o w n f r o m several (botanical) gardens i n
Europe, and there the w h o l e year through. T h e natural distribution area
is tropical A m e r i c a . Sect. Fuscovinaceae B o n & Candusso
F o r an elaborate discussion o n the s y n o n y m y o f Lepiota rubella and L. Clamp-connections absent; elements o f pileus covering articulate and
bettinae, see V e l l i n g a et al. (in S y d o w i a 50: 272-278. 1998). cylindrical, repent to ascending.
17. Lepiota parvannulata ( L a s c h : Fr.) G i l l e t , H y m e n o m y c e t e s : 66. 18. Lepiota fuscovinacea M 0 l l e r & I. L a n g e i n J . Lange, F l . agar. dan.
1 8 7 4 . - F i g . 93. 5 : V . 1 9 4 0 . - F i g . 94.
Agaricus parvannulatus L a s c h i n L i n n a e a 3: 156. 1828; Agaricus Lepiota fuscovinacea M 0 l l e r & J . L a n g e i n J . L a n g e , F l . agar. dan.
parvannulatus L a s c h : Fr., Syst. m y c o l . , Ind. gen.: 34. 1832. 1: 33. 1935 (not valid).
Enderle & Krieglst. i n Z . M y k o l . 55: between p. 64 and p. 65. 1989; I .
Lange, F l . agar. dan. 1: p i . 1 3 H . 1935; D . R e i d i n F u n g . rar. Ic. c o l . 2:
pi. 10a. 1967.
S E L . D E S C R . & FIGS. - K i i h n e r i n B u l l , trimest. Soc. m y c o l . Fr. 52:
0 00 00 0 220-221. 1936; D . R e i d i n F u n g . rar. Ic. c o l . 2: 8-9. 1967.

V E R N . N A M E - Purperbruine p a r a s o l z w a m .
F i g . 93. Lepiota parvannulata (habit del. P. K e l d e r m a n ) .

Lepiota fuscovinacea is unique w i t h i n the genus o n account o f the

c o m b i n a t i o n o f the f o l l o w i n g characters: absence o f clamp-connec-
tions, uninucleate spores, and a pileus c o v e r i n g made up o f repent
hyphae.
Sect. Stenosporae (J. Lange) Kuhner
Spores c y l i n d r i c a l w i t h a straight or rounded bottom, w h i c h i n most

cases is elongated into a distinct spur; pileus covering not hymeniform,
but made up o f adnate, ascending or erect l o n g and slender elements;
clamp-connections present.
N . B . Spore w i d t h measured i n m i d d l e o f spores.
19. Lepiota castanea Q u e l , i n C . r. A s s . franc. A v . S c i . (Reims, 1880)

9: 661. 1881 ( C h a m p . Jura Vosges S u p p l . 10). - F i g . 9 5 .
Lepiota ignicolor Bres., F u n g i trident. 2: 3. 1892; Lepiota ignipes
F i g . 94. Lepiota fuscovinacea.
L o c q . i n B u l l . mens. Soc. l i n n . L y o n 14: 59. 1945 (not valid); Lepiota
Pileus 17-42 m m , when y o u n g bluntly conical, expanding to more or ignipes L o c q . ex M . B o n i n D o c . m y c o l . 8 (30-31): 70. 1978.
less applanate or plano-convex w i t h broad prominent, w i d e u m b o to E X C L . - Lepiota castanea sensu Pilat i n A c t a M u s ! natn. Prag. 1 I B
plano-concave w i t h l o w u m b o and undulating margin, w i t h fibrillose- (2): 3-5. 1955 ( = probably L. boudieri).
lanate tomentose covering, i n centre w i t h uplifted squamules, towards M I S A P P L . - Lepiota fulvella sensu A l e s s i o i n M i c o l . ital. 21 (3): 16-
fnargin more appressed fibrillose, showing the background, sordid grfey- 20. 1992.
p i n k to dark purplish, purplish grey or dark reddish b r o w n porph^ra- SEL. I C O N . - M . B o n i n F u n g . rar. Ic. c o l . 11: p i . 84, fig. 2. 1979 (as
ceous ( M u . 7.5 - 10 Y R 3/2; K . & W . 6E5-4) nearly b l a c k i s h w i t h age L. ignicolor); Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 219, 228. 1995
at centre, towards margin slightly paler o n pale p i n k i s h background (10 (as L. castanea and L. ignicolor resp.); Candusso & L a n z o n i , L e p i o t a :
YR 7/3 - 6/4, 7 B 4 ) , w h e n y o u n g w i t h velar remnants at margin. p i . 18. 1990; K u h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 53: A t l a s p i . 74-
Lamellae, L = 50-80,1 = 1-3, very crowded, free, subventricose or w i t h 2 . 1 9 3 7 ; L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l . G . Bres. 26: 101.
straight edge, up to 4 m m broad, white-cream to cream-coloured, w i t h 1983; R . P h i l l i p s , Paddest. S c h i m m . : 29. 1981; R a i d et al. i n Svampe
age b r o w n spotted or sordid pale y e l l o w - b r o w n (10 Y R 8-7/4), w i t h 26: 39. 1992; D . R e i d i n F u n g . rar. Ic. c o l . 6: p i . 43b. 1972.
white fimbriate edge. Stipe 20-55(-95) X 3-11 m m , c y l i n d r i c a l or S E L . D E S C R . & FIGS. - Enderle & Krieglst. i n Z . M y k o l . 55: 53-55,
slightly broadened towards straight or subbulbous base, h o l l o w w i t h and 73-75 (as L. ignicolor). 1989; H o r a k i n B o l l . G r u p p o m i c o l . G .
age, i n apical part pale p i n k i s h cream to pale p i n k i s h o r sordid cream Bres. 26: 90-92. 1983 (as L. ignicolor); K e l d e r m a n , Parasolzw. Z u i d -
(10 Y R 8/4 - 7/3, 5 A 2 ) and innately fibrillose, rather glabrous, lower L i m b u r g : 78-79, 88-89, and 90-91. 1994 (resp. as L. castanea, L. igni-
d o w n w i t h or without distinct annular zone and i n lower half or 3/4 w i t h color, and L. ignipes); M a i . & Bert., F l . C h a m p , sup. M a r V 1: 130.
zones and bands o f tomentose squamules, concolorous w i t h covering 1970 (as L. ignipes); M i g l . & C l e r i c u z i o i n M i c o l . Veget. medit. 4: 36,
of pileus o n cream-coloured background, w i t h basal white tomentum, 38. 1989 (as L. ignicolor); D . R e i d i n F u n g . rar. Ic. c o l . 6: 11/14. 1972.
and white rhizoids. Context i n pileus white to off-white or w i t h p i n k i s h V E R N . N A M E - Kastanjeparasolzwam, incl. Vuurparasolzwam.
tinge, i n stipe at first also cream, discolouring i n cortex to grey-brown.

S m e l l fruity or sweetish fungoid, or earth-like and unpleasant, slightly
like the s m e l l o f L. cristata. Taste indistinct. Spore print white to cream.
» Spores (3.5-)4.0-6.5(-7.0) X 2.0-3.3(-3.5) p m i n side-view, Q =
(1.5-)1.75-2.3(-2.75), Q = 1.9-2.15, o b l o n g to c y l i n d r i c a l , a very few
slightly allantoid, dextrinoid, congophilous, not metachromatic i n
C r e s y l B l u e , often i n tetrads, uninucleate. B a s i d i a (12.5-)13.5-22 X
4.0-7.0 p m , 4-spored. C h e i l o c y s t i d i a 13-45 X 6-15 p m , abundant,
cylindrical, narrowly clavate or clavate, often uniseptate.
Pleurocystidia not observed. Pileus covering made up o f radial, loose-
l y arranged, appressed to ascending, inflated to c y l i n d r i c a l , septate
hyphae, 5.0-20 p m i n diam., slightly thick-walled, w i t h terminal ele-
ments w i t h rounded or acute apex; pigment brown, soluble i n ammo-
nia. Stipitipellis a cutis made up o f cylindrical, colourless to y e l l o w i s h ,
4.0-6.0 u m w i d e hyphae, w i t h i n lower part o f stipe a covering w i t h
elements l i k e those o f pileus covering. Clamp-connections absent.
H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic and
terrestrial o n l o a m y to c l a y e y soils, r i c h i n nutrients and humus, i n
deciduous and coniferous woods and plantations, also found o n m i n e
waste heaps. I n the Netherlands rare and mostly i n the western clayey-
peat regions, the river area and southern L i m b u r g , very rare elsewhere
(for a distribution map, see A r n o l d s et a l . , O v e r z . Paddest. Nederland:
288. 1995). A u g . - O c t . N o t c o m m o n i n the rest o f Europe. F i g . 95. Lepiota castanea.
Pileus 6-40 m m , campanulate or paraboloid w h e n y o u n g , expanding ly developed underlayer the underlying context shows more clearly.
to plano-convex, often w i t h (broad) umbo, when y o u n g equally dark Variants occurring in the dune area are strikingly slender and have a bril-
red-brown, red-brown, orange-brown (e.g. M u . 5 Y R 2.5/2, 5-7.5 Y R liant-white stipe; they also show more white on the pileus than the more
3/4, 5 Y R 4/6, c. 5 Y R 5-6/8 at centre and w h e n y o u n g , later and sturdy inland types. T h i s dune variant is probably B o n ' s L. ignipes.
towards m a r g i n 5 Y R 4/6, 7.5 Y R 4-5/6) and velvety-tomentose w i t h T h e spore sizes differ greatly between collections, from 7.0-9.0 p m
uplifted squamules, w i t h age breaking up i n smaller patches or squa- in one to 12-14 p m i n others. B u t the range o f the spore sizes is a c o n -
mules, often i n concentrical zones and towards margin more radially tinuum, without separable units. T h e small-spored collections tend to
fibrillose, pallescent towards margin, o n predominantly white or pale have less-septate elements i n the pileus covering. T h e number o f septa
background (7.5 Y R 7/8-8/6, 5 Y R 7/8), o r not or hardly showing the i n these pileus elements differs, not only between collections, but also
underlying context at a l l , w i t h age often orange d i s c o l o u r i n g , some- during the growth o f the pileus (the number increases w i t h growth) and
times w i t h some white velar remnants at margin, and margin exceed- the number o f septa is greater at the pileus centre than at the margin o f
ing lamellae, especially w h e n y o u n g . L a m e l l a e , L = 25-40,1 = 1-5, the pileus. The pigment is situated inside the walls o f these elements,
moderately distant to moderately crowded, free, ventricose, up to 6 m m and i n the walls o f the underlying elements, but i n addition to this pari-
w i d e , white at first, later cream, often w i t h orange sheen (e.g. 10 Y R etal pigment there is often some pigment that is soluble i n ammonia.
8/5) and w i t h age orange-spotted, w i t h white fme-flocculose edge. H o r a k (in B o l l . G r u p p o m i c o l . G . Bres. 26: 90-92. 1983) charac-
Stipe 22-65 X 1-6 m m , c y l i n d r i c a l , subbulbous at base, often curved terized L. ignicolor by its s m a l l spores, non-septate elements and vac-
in basal part, narrowly fistulose, at apex (1/4 to 1/3 o f length) cream o r uolar pigment. Unfortunately this combination is not as
pale p i n k i s h cream-coloured and lengthwise adnate-fibrillose, without, straightforward as it seems. Specimens w i t h s m a l l spores and up to 2
rarely w i t h , annulus, below this zone w i t h fibrillose to squamulose cov- septa i n their pileus elements do occur, and so do basidiocarps w i t h b i g
ering, concolorous w i t h squamules o n pileus o n s h i n i n g l y white to spores, and soluble pigment o r w i t h spores over 10 p m l o n g w i t h non-
slightly paler o r yellowish-reddish, p i n k i s h orange, pale red-brown or septate elements i n the pileus covering.
orange-brown (5 Y R 5/6, 6-5/8) background, often staining orange M i g l i o z z i & Z e c c h i n ( i n M i c o l . ital. 2 6 (2): 11-22. 1997) distin-
w h e n touched, sometimes orange-brown strigose at base, often w i t h guished three species i n this c o m p l e x : L. castanea Q u e l , sensu stricto,
white m y c e l i a l cords at base. A n n u l u s rarely present, s m a l l and w o o l - L. castanea sensu auct., and L. ignicolor Bres., differing i n colour,
ly. Context w h i t i s h cream i n pileus and i n stipe apex, l o w e r d o w n i n structure o f the pileus covering and spore size. In v i e w o f the transi-
stipe concolorous w i t h surface and often b e c o m i n g v i v i d orange w i t h tions i n spore size, the variability i n the number o f septa and the length
age. S m e l l fruity, l i k e the sweet component o f the L. cristata smell. of the pileus covering elements, their ideas are not f o l l o w e d here, and
Taste m i l d , fungoid. Spore print white. L. castanea is given i n a broad sense.
Spores i n side-view 7.0-14.0(-15.5) X 3.0-5.5 p m , Q = (1.85-)2.0- Lepiota cingulum is closely related to L. castanea and is often con-
3.0(-3.3), Q a v = 2.15-2.9, more o r less c y l i n d r i c a l , w i t h especially i n fused w i t h the latter i n the field; for differences see under L. cingulum.
the longer spores a distinct sac-like spur, w h i c h protrudes from abaxi- Lepiota andegavensis M o r n a n d ( i n D o c . m y c o l . 12 (48): 4 3 .
al side, rarely w i t h suprahilar depression, w i t h patent hilar appendage, ('1982') 1983) comes very close, but that species is characterized by
in frontal v i e w , ( s u b ) c y l i n d r i c a l , t h i c k - w a l l e d , uniguttulate o r not, very dark colours o n the pileus. J u d g i n g from B o n ' s description ( F l .
strongly dextrinoid, congophilous, not metachromatic i n C r e s y l B l u e . m y c o l . Eur. 3, Lepiotes: 54. 1993) the pileus covering resembles that
B a s i d i a 19-32 X 6-10 p m , 4-spored, rarely i n t e r m i x e d 2-spored. of L. boudieri, but that is not the case; the elements are septate and bear
L a m e l l a edge sterile. C h e i l o c y s t i d i a 19-41 X 5-10 p m , fusiform, nar- only a basal clamp-connection.
r o w l y utriform, narrowly clavate, o r c y l i n d r i c a l w i t h s u b c y l i n d r i c a l Lepiota castanea contains a l o w e r amount o f amanitins than the
apex, colourless, rarely thick-walled. Pleurocystidia absent. Pileus cov- species i n Lepiota sect. Ovisporae subsect. Helveolinae (Gerault &
ering made up o f erect c y l i n d r i c a l elements, 55-330 X 7-22 p m , w i t h - Girre i n C . r. hebd. Seanc. A c a d . S c i . , Paris, Ser. D , 280: 2842. 1966).
out o r w i t h u p to 3 septa, w i t h parietal b r o w n pigment and additionally
some pigment soluble i n a m m o n i a , w i t h a layer o f adnate, c y l i n d r i c a l , 20. Lepiota cingulum Kelderman i n Persoonia 15: 537. 1994. - F i g . 96.
flexuous hyphae, 2.0-5.0 p m w i d e , w i t h narrowly clavate to subcapi- M I S A P P L . - probably Lepiota fulvella sensu M . B o n i n D o c . m y c o l .
tate, terminal elements and b r o w n thickened w a l l s . Stipitipellis a cutis, 11 (43): 35-36. 1981; sensu M . B o n , F l . m y c o l . Eur. 3, Lepiotes: 5 1 .
made up o f colourless, c y l i n d r i c a l elements, 2.0-4.0 u m w i d e ; elements 1993.
i n squamules shorter, more irregular, more flexuous and more often S E L . I C O N . - H e i l m a n n - C l a u s e n i n Svampe 34: 33. 1996.
non-septate than those o n pileus; underlayer o f irregular, flexuous S E L . D E S C R . & FIGS. - K e l d e r m a n i n C o o l i a 31: 89-91, fig. 1. 1988
c y l i n d r i c a l hyphae present. Clamp-connections present i n a l l tissues. (as Lepiota s p e c ) ; K e l d e r m a n i n Persoonia 15: 537-539, fig. 1. 1994;
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres- K e l d e r m a n i n Parasolzw. Z u i d - L i m b u r g : 80-81. 1994.
trial i n deciduous, m i x e d o r coniferous woods, i n mossy dune scrub, VERN. NAME - Gordelsteelparasolzwam.
also o n m i n e waste heaps, o n sandy to c l a y e y soils, often r i c h i n
humus. In the Netherlands rather c o m m o n throughout the country, but Pileus 31-55(-80) m m , w h e n y o u n g hemispherical, expanding v i a
less c o m m o n i n districts characterized b y nutrient-poor sand; A u g . - m i d broadly campanulate to plano-convex, applanate, w i t h , rarely without,
Nov. Widespread i n E u r o p e and temperate parts o f A s i a , also i n N o r t h low to prominent, umbo, sometimes a bit undulating around centre, at
America. centre dark b r o w n ( M u . 5 Y R 3/4, 5-7.5 Y R 3/3, 7.5 Y R 4/6, 10 Y R 4-
3/4), sometimes more p i n k i s h b r o w n (5 Y R 4-3/2), felted-tomentose
Lepiota castanea i n the present sense is a variable species, both i n with small uplifted tufts, a bit p l u s h - l i k e (lens); around centre surface
macroscopical characters and i n microscopical characters like spore size splitting up into concentrical zones o f adnate to slightly uplifted patch-
and pileus covering. Some specimens show a more intense orange dis- es to squamules, those b e c o m i n g more adnate and radially arranged
coloration when touched or w i t h age than others. T h e aspect o f the towards margin, and b e c o m i n g paler b r o w n than centre o n cream,
pileus, completely b r o w n or showing white context, differs greatly, and occasionally p i n k i s h cream, background; m a r g i n exceeding lamellae.
depends on the development o f the felted underlayer; i f this layer is w e l l L a m e l l a e , L = 45-60,1 = 1-3, moderately crowded to moderately dis-
developed the pileus surface is totally covered; i n specimens w i t h a poor- tant, free, rarely almost free and narrowly adnate, ventricose, up to 6.5
Differences w i t h L. castanea are to be found i n the colours o f the

basidiocarps: more subdued b r o w n i n L. cingulum and more v i v i d
orange-brown i n L. castanea.
21. Lepiota boudieri Bres., F u n g i trident. 1: 4 3 . 1884, non Lepiota

boudieri Gueguen, ('1908') 1909. - F i g . 97.
Lepiota fulvella R e a i n Trans. B r . m y c o l . S o c . 6: 61. ('1917') 1918.
- Lepiota castaneopallida D a m b l o n & L a m b i n o n i n D a m b l o n et al. i n
L e j e u n i a 2 1 : 90. (' 1957') 1959. - Lepiota fulvella f. gracilis J . Lange,
F l . agar. dan. 1: 32. 1935 (not valid).
E X C L . - Lepiota fulvella sensu M . B o n i n D o c . m y c o l . 11 (43): 35-
36. 1981; M . B o n , F l . m y c o l . Eur. 3, L e p i o t e s : 5 1 . 1993 (see note);
sensu A l e s s i o i n M i c o l . ital. 21 (3): 16-20. 1992 ( = L. castanea).
M I S A P P L . - Lepiota acerina sensu M . B o n i n D o c . m y c o l . 11 (43):
36. 1981; sensu M . B o n , F l m y c o l . Eur. 3, Lepiotes: 52. 1993; proba-
bly Lepiota castanea sensu Pilat i n A c t a M u s . natn. Prag. 11B (2): 3-
5. 1955.
C h i a r i & F o r t i i n B o l l . C i r c . m i c o l . G . C a r i n i 29-30: centre o f issue.
1995; J . L a n g e , F l . agar. dan. 1: p i . 1 2 D , 12F. 1935 (as L. fulvella);
Krieglst. i n Z . M y k o l . 5 1 : opp. p. 132. 1 9 8 5 ( a s L . fulvella); Kuhner in
F i g . 96. Lepiota cingulum. B u l l , trimest. S o c . m y c o l . Fr. 53: A t l a s p i . 74-1. 1937 (as L. fulvella);
Patane i n S c h w e i z . Z . P i l z k . 68: 223. 1990 (as L. fulvella); R. Phillips,
mm wide, cream (2.5 Y 8/2) w i t h concolorous o r white, finely floccu-; Paddest. S c h i m m . : 29. 1981 (as L. fulvella).
lose (lens) to distinctly floccose edge. Stipe 28-58 X 4-10 m m , slight? S E L , D E S C R . & FIGS. - C h i a r i & F o r t i i n B o l l . C i r c . m i c o l . G . C a r i n i
ly broadening towards base, lengthwise innately fibrillose, hollow, at 29-30: 33-35. 1995; K e l d e r m a n i n C o o l i a 3 1 : 4 0 - 4 1 , f i g . 1. 1988;
apex w h i t i s h to cream w i t h p i n k i s h sheen to pale p i n k at base, i n lower K u h n e r i n B u l l - t r i m e s t . S o c . m y c o l . Fr. 52: 234-236. 1936 (as L. ful-
h a l f to 1/3 w i t h distinct girdles o f squamules, concolorous w i t h those vella); Patane i n S c h w e i z . Z . P i l z k . 68: 222-224. 1990 (as L. fulvella);
on pileus. Context w h i t i s h to creamy and d u l l i n pileus, i n stipe shiny V e l l i n g a & Huijser i n Persoonia 15: 237-239, fig. 6. 1993.
cream i n y o u n g specimens, later p i n k i s h - b r o w n i s h to red-brown o r V E R N . N A M E - Oranjebruine p a r a s o l z w a m .
b r o w n . S m e l l indistinct to strong, unpleasant fungoid. Taste fungoid.
Spore print white. Pileus 20-65 m m , w h e n y o u n g hemispherical o r (conico-)campanulate
Spores ( 7 . 5 - ) 8 . 5 - l l X 3.5-4.5(-5.0) p m , Q = 2.0-3.0, Q a v = 2.3- w i t h inflexed margin, expanding to convex, plano-convex, applanate,
1
2.55(-2.75), w i t h straight or outgrown spur, cylindrical to oblong, w i t h
rounded or more acute apex, i n frontal v i e w oval, c y l i n d r i c a l , strongly
dextrinoid, congophilous, not metachromatic i n C r e s y l B l u e . B a s i d i a
21-39 X 6.0-10 p m , 4-spored, o c c a s i o n a l l y 2-spored. L a m e l l a edge
sterile. C h e i l o c y s t i d i a abundant, 17.5-56 X 7.0-17 um, rarely m u c h
bigger: 48-90 X 10.5-24 p m , narrowly clavate, more rarely clavate,
c y l i n d r i c a l or narrowly utriform, slightly t h i c k - w a l l e d and colourless.
Pleurocystidia absent. Pileus covering made up o f erect to ascending,
sometimes very twisted, elements, 100-260 X 12-16 um, up to 4 0 0 p m
long, w i t h attenuate apex and narrowing into pedicel, without septa, or,
mere rarely, w i t h up to 2 septa i n lower part, w i t h d u l l b r o w n parietal
pigment; underlayer o f c y l i n d r i c a l b r o w n - w a l l e d , flexuous hyphae
present, w i t h narrowly clavate terminal elements. Stipitipellis a cutis,
made up o f c y l i n d r i c a l , 3.0-10 p m w i d e hyphae w i t h colourless
encrusting pigment; elements o f squamules as those o n pileus. C l a m p -
connections present i n a l l tissues.
H A B I T A T & D I S T R . - S o l i t a r y o r i n s m a l l to large groups, sapro-
trophic a n d terrestrial, o n c l a y e y soils, i n deciduous and coniferous
w o o d s i n the western part o f the Netherlands a n d i n southern
L i m b u r g , also o n m i n e waste heaps, very rare. A u g . - O c t . R a r e i n
E u r o p e , recorded f r o m the Netherlands, France, D e n m a r k , and Great
Britain.
Lepiota cingulum resembles L. brunneoincarnata i n habitus. B o t h

species have relatively sturdy basidiocarps w i t h distinct concentrical
zones o f squamules o n pileus a n d girdles o n the stipe. Y o u n g speci-
mens o f L. cingulum may have a purplish hue. T h e shape o f the spores
is the differentiating character: spurred i n L. cingulum, and ellipsoid i n
L. brunneoincarnata. F i g . 97. Lepiota boudieri.
w i t h deflexed margin, o r plano-concave, and often w i t h broad umbo, especially his variety subpurpurata M . B o n (in B o n & G . Riousset i n
finely tomentose-squamulose at centre, orange-brown, dark red-brown D o c . m y c o l . 22 (85): 63. 1992). These c o l o u r variants fit w e l l into the
or deep rusty b r o w n ( M u . 7.5 Y R 4/6, 5 Y R 4/6), towards m a r g i n present concept o f L. boudieri. A m e r i c a n collections o f L. acerina,
adnately, radially fibrillose-squamose, paler and more y e l l o w than at present i n the L e i d e n herbarium, proved to be L. boudieri.
centre (7.5 Y R 5-6/6, 7/8) not o r hardly showing the underlying pale Lepiota fulvella i n the sense o f A l e s s i o (in M i c o l . i t a l . 21 (3): 16-
cream, pale pinkish-ochraceous buff background (10 Y R 5-7/8, 7-8/6), 20. 1992) p r o b a b l y belongs to the c o m p l e x o f Lepiota castanea,
at m a r g i n , especially w h e n y o u n g , w i t h white velar remnants. because o f the slender basidiocarps, the spore size (spores 10.5-12
L a m e l l a e , L = 40-60, 1 = 1-5, c r o w d e d , moderately c r o w d e d , free, p m long) and the type o f elements o n the pileus (non-articulate, non-
straight to ventricose, up to 6 m m broad, white to pale cream, often septate).
w i t h reddish spots w i t h age, w i t h concolorous or slightly darker, floc- Lepiota fulvella i n the sense o f B o n (in D o c . m y c o l . 11 (43): 35-36.
culose edge. Stipe 30-70 X 3-8 m m , c y l i n d r i c a l and often gradually 1981 and F l . m y c o l . Eur. 3, Lepiotes: 5 1 . 1993) is probably Lepiota
broadened towards base, o r w i t h slightly bulbous base, h o l l o w w i t h cingulum. However, material from B o n ' s herbarium filed as L. fulvel-
age, at apex w h i t i s h to p i n k i s h , lengthwise adnately fibrillose, w h e n la proved to belong to several species.
y o u n g w i t h white flocculose annulus, disappearing w i t h age, o r w i t h T h e colours o f Lepiota boudieri are i n general a little bit more y e l -
orange-brown annular zone, w h i c h can be quite l o w o n stipe, below l o w i s h than those o f L. castanea.
this zone w i t h adnate to slightly squarrose orange-brown o r concolor-
ous w i t h pileus, fibrillose squamules o n b r o w n i s h flesh-pink, or p i n k 22. Lepiota subalba K i i h n e r ex P . D . O r t o n i n Trans. B r . m y c o l . Soc.
arachnoid-lanate background. Context white and d u l l i n pileus, shiny 43: 287. 1960. - F i g . 98.
i n stipe, cream and more or less concolorous w i t h surface at apex, to Lepiota subalba K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 233.
red-brown b e l o w and white at base. S m e l l varying from fungoid-earthy 1936 (not valid).
to sweetish-fruity, nutty etc. Taste m i l d , fungoid o r l i k e s m e l l . Spore M I S A P P L . - Lepiota albosericea sensu J . L a n g e , F l . agar. dan. 1: 32.
print white. 1935; Lepiota naucina sensu M o s . , R o h r l i n g e , Blatter-Bauchpilze, 2.
Spores i n side-view 7.0-10.0(-11.0) X 3.0-4.0(-4.5) p m , Q = (1.9- A u f l . : 135. 1955.
)2.1-2.9(-3.1), Q a v = (2.2-)2.35-2.7, w i t h truncate to spurred base, S E L . I C O N . - J . L a n g e , F l . agar. dan. 1: p i . 1 2 B . 1935 (as L.
c y l i n d r i c a l , narrowly triangular o r w i t h curved abaxial side, i n frontal albosericea).
v i e w c y l i n d r i c a l , t h i c k - w a l l e d , w i t h or without one guttule, w i t h patent S E L . D E S C R . & FIGS. - Enderle & Krieglst. i n Z . M y k o l . 55: 93-94.
hilar appendage, dextrinoid (orange-brown), congophilous, not 1989; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 96-97. 1994; K i i h n e r i n
metachromatic i n C r e s y l B l u e . B a s i d i a (15-)16-27.5 X 6.0-8.0 u m , B u l l , trimest. Soc. m y c o l . Fr. 52: 233-234. 1936; P . D . O r t o n i n Trans.
usually 4-spored, rarely i n t e r m i x e d w i t h some 2-spored basidia. Br. m y c o l . Soc. 43: 287-288. 1960; A . Pears, i n Trans. B r . m y c o l . S o c .
L a m e l l a edge sterile. C h e i l o c y s t i d i a 13-38 X 5.0-12 p m , subutriform, 35: 98. 1952.
clavate, narrowly clavate to o v o i d and pedicellate, colourless and not V E R N . N A M E - Bleke parasolzwam.
t h i c k - w a l l e d . P l e u r o c y s t i d i a absent. P i l e u s c o v e r i n g made up o f
hyphae i n adnate to ascending bundles, made up o f 2-5 articulate and Pileus 10-35 m m , w h e n y o u n g hemispherical, campanulate o r c o n i c a l
c l a m p e d elements, provided w i t h pigment; terminal elements 40-215 w i t h inflexed m a r g i n , expanding to plano-convex, applanate or even
X 8.0-24 u m , more or less c y l i n d r i c a l w i t h rounded apex, rarely inflat- plano-concave and then undulating, w i t h o r without broad (low) umbo,
ed, w i t h prominent vacuolar, b r o w n pigment, d i s s o l v i n g i n a m m o n i a at centre pale y e l l o w i s h , pale ochraceous o r creamy ( M u . 10 Y R 8/3,
and w i t h some parietal pigment. Stipitipellis a cutis o f n o n o r slightly
coloured, c y l i n d r i c a l , 3.0-14 p m w i d e hyphae, w i t h i n lower half o f
stipe the same k i n d o f elements as o n pileus. Clamp-connections pres-
ent i n a l l tissues.
H A B I T A T & DISTR. - I n s m a l l groups, gregarious, saprotrophic and
terrestrial o n h u m u s - r i c h , often nutrient-rich, loamy-clayey, rarely
sandy, soils, i n m i x e d or deciduous woods, copses o r city-parks, also
on m i n e waste heaps, rather rare i n the Netherlands, A u g . - O c t . ( b e g i n -
ning o f N o v . ) . Widespread i n the temperate regions o f Europe, proba-
bly also i n N o r t h A m e r i c a .
O c c a s i o n a l l y totally white basidiocarps are encountered. V e r y dark

specimens, b l a c k i s h b r o w n w i t h lilacinous tinges, are also rarely noted.
A s the m i c r o s c o p i c a l characters o f those t w o c o l o u r variants do not
deviate f r o m those o f n o r m a l Lepiota boudieri, (except for the absence
of pigments i n the former), no formal status is given to those variants.
F o r a discussion o f the s y n o n y m y o f L. boudieri and L. fulvella Rea,
see V e l l i n g a & Huijser (in Persoonia 15: 237-239. 1993).
Lepiota castaneopallida Damblon & Lambinon, a species
described f r o m B e l g i u m , i s characterized b y multiseptate hyphae o n
the pileus. Because o f this it i s considered a s y n o n y m o f L. boudieri.
T h e presence o r absence o f clamp-connections is not mentioned.
Lepiota acerina P e c k ( i n R e p . N . Y . St. M u s . nat. H i s t . 5 1 : 2 8 3 .
1898) is probably another s y n o n y m . However, S m i t h (in L l o y d i a 17:
317. 1954) stated that clamp-connections are absent. B o n ' s interpreta-
tion o f this taxon i s a more purplish variant than t y p i c a l L. boudieri, F i g . 98. Lepiota subalba.
8/4), towards m a r g i n cream to white, at centre felted, not broken up sensu K e l d e r m a n i n C o o l i a 3 1 : 43-44. 1988 ( = i n a l l cases L. pilodes);
into squamules, towards m a r g i n w i t h patches o f felted-floccose cover- sensu P e g l . & L e g o n i n M y c o l o g i s t 12: 37. 1998 ( = L. cingulum).
i n g o n a white background, towards margin more squamulose-fibril- S E L . I C O N . - J . L a n g e i n D a n s k bot. A r k . 4 (4): p i . I d . 1923; J .
lose, or completely sericeous-felted, when y o u n g w i t h velar remnants L a n g e , F l . agar. dan. 1: p i . 1 4 D . 1935; Z e c c h i n & M i g l . i n M i c o l . ital.
at margin (as covering), w i t h m a r g i n exceeding lamellae. L a m e l l a e , L 25 (2): p i . 1 & 2. 1996.
= 30-40, 1 = 1-3, rather distant to moderately crowded, free, some- S E L . D E S C R . & FIGS. - M . B o n i n B u l l , trimest. S o c . m y c o l . Fr. 92:
times some furcate, ventricose, subventricose or segmentiform, up to 324-326, fig. 6. 1976; P . D . O r t o n i n Trans. B r . m y c o l . S o c . 9 1 : 562.
3.5 m m broad, white, cream to brownish cream w i t h age, w i t h white 1988; V e l l i n g a & Huijser i n P e r s o o n i a 15: 224-226, fig. 1. 1993;
finely floccose edge (lens). Stipe 18-60 X 1.5-4.5 m m , c y l i n d r i c a l or Z e c c h i n & M i g l . i n M i c o l . ital. 25 (2): 35-37, figs. 1 & 2. 1996.
slightly broadened at base, or more rarely, tapering towards base, fis- V E R N . N A M E - Viltparasolzwam.
tulose w i t h age, cream, w h i t i s h at apex, and there lengthwise innately
fibrillose or glabrous, l o w e r d o w n more p i n k i s h b r o w n or p i n k Pileus 10-28(-34) m m , w h e n y o u n g p a r a b o l o i d to campanulate w i t h
coloured, w i t h r i n g - l i k e zone o f w h i t i s h lanate-floccose material, more inflexed margin, expanding to convex, plano-convex or slightly plano-
b e l o w w i t h floccose or lanate girdles; squamules or f l o c c i w i t h y e l - concave w i t h l o w broad umbo, at centre dark grey-brown, ochre
lowish-ochraceous or y e l l o w brownish tinges, vanishing w i t h age and b r o w n i s h or grey-brown w i t h p i n k i s h tinge ( M u . 7.5 Y R 3/4, 5 Y R 3-
rain; white m y c e l i u m strands at base. Context i n pileus w h i t i s h and 5/3-4, K . & W . 7 - 8 E 4 , 7 F 6 - 5 , 5-6D6), paler towards m a r g i n (7.5 Y R 5-
d u l l , i n stipe at apex white to pale pink, or pale p i n k i s h b r o w n to rather 7/4), felted at centre, around centre flocculose-felted or broken up i n
dark reddish b r o w n at base. S m e l l sweetish, like the sweet component more or less distinct appressed or slightly uplifted felted squamules;
o f the s m e l l o f L. cristata, fungoid, faintly or strongly like the smell o f margin w i t h some white velar remnants w h e n young. L a m e l l a e , L =
L. cristata, or fishy-fruity. Taste bitterish. Spore print probably white. 28-44,1 = 1-3, rather crowded or crowded, free, (sub)ventricose, up to
Spores i n side-view (6.5-)7.5-10.0(-10.5) X 2.5-3.5(-4.0) um, Q = 5 m m broad, w h i t i s h at first, later cream or pale beige w i t h slight p i n k -
(2.05-)2.15-3.0(-3.3), Q a v = 2.35-2.85, w i t h spurred, or truncate ish reflex, w i t h w h i t i s h finely flocculose edge. Stipe 15-50 X 2-4.5(-
base, rarely w i t h confluent base, i n frontal v i e w c y l i n d r i c a l , unigut- 6.5) m m , c y l i n d r i c a l and usually w i t h broadened base, subfistulose,
tulate, often i n groups (tetrads), d e x t r i n o i d , c o n g o p h i l o u s ; w a l l not' w h e n y o u n g w h i t i s h , cream-brown, finely fibrillose, later from base
c o l o u r i n g i n C r e s y l B l u e . B a s i d i a 15-34 X 5.0-8.5 p m , 4-spored# upwards reddish b r o w n , pale b r o w n rusty, i n lower 1/2 to 2/3 w i t h scat-
L a m e l l a edge sterile. C h e i l o c y s t i d i a 15-42 X 6.0-10 p m , u s u a l l y tered squamules and girdles, like covering o n pileus, without distinct
c y l i n d r i c a l , w i t h subcapitate apex or slightly s w o l l e n at centre, rarely annulus, w h e n y o u n g sometimes w i t h a white fugacious ring-zone,
narrowly clavate, not coloured, with slightly thickened wall. w i t h white m y c e l i u m cords at.base. Context i n pileus whitish cream,
P l e u r o c y s t i d i a absent. P i l e u s covering made up o f erect, irregular, nar- in stipe white at first, later b r o w n i s h or pale brown. S m e l l indistinct to
r o w l y fusiform to c y l i n d r i c a l elements, 30-210 X 6.0-21 p m , w i t h slightly sweetish. Taste slightly bitter. Spore print white.
rounded apex, narrowed into pedicel, slightly thick-walled, colourless, Spores i n side-view (6.5-)7.0-9.5(-10.5) X 3.0-4.0(-4.5) p m , Q =
w i t h basal clamp-connection. B a s a l layer o f short elements not pres- 1.7-2.7, Q a v = 2.1-2.4, w i t h straight or outgrown broad spur, some-
ent. S t i p i t i p e l l i s a cutis o f c y l i n d r i c a l elements 3.0-8.0 p m w i d e ; ele- times allantoid or phaseoliform, usually rather c y l i n d r i c a l , some atten-
ments o f squamules as those o n pileus, but u s u a l l y shorter (e.g. uate towards base, i n frontal v i e w s u b c y l i n d r i c a l , dextrinoid (pale
36-115 X 12-20 pm) and c y l i n d r i c a l to n a r r o w l y fusiform. C l a m p - b r o w n or orange-brown), c o n g o p h j l e u s ^ w a l l not coloured i n C r e s y l
connections present i n a l l tissues. B l u e . B a s i d i a (16.5-)20-32 X / 6 . 0 - 9 . 5 p m , 4-spored, rarely (3-)2-
H A B I T A T & DISTR. - Solitary to gregarious i n s m a l l groups, sapro- spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a (17-)20-40(-48) X 5.0-12
trophic and terrestrial i n deciduous woods, scrub etc., rarely i n conif- um, c y l i n d r i c a l , narrowly clavate, or rarely narrowly utriform, slightly
erous forests, o n sandy to l o a m y soils r i c h i n humus, widespread i n the t h i c k - w a l l e d , not c o l o u r e d . P l e u r o c y s t i d i a absent. P i l e u s c o v e r i n g
Netherlands, often o n the inner coastal dunes. E n d o f July-Oct. K n o w n made up o f c y l i n d r i c a l erect elements, (35-)45-320(-440) X 6.0-19 p m ,
f r o m the temperate zones o f E u r o p e ; distribution outside E u r o p e usually narrowed into a pedicel, without or w i t h up to 2(-3) clampless
unknown. septa, w i t h b r o w n i s h , slightly thickened w a l l s , c l a m p e d at base.
Stipitipellis a cutis o f c y l i n d r i c a l hyphae, 2.0-8.0 p m i n d i a m . , w i t h
23. Lepiota tomentella J. L a n g e i n D a n s k bot. A r k . 4 (4): 48. 1923. - some patent colourless hyphae; squamules o n stipe made up o f b r o w n -
F i g . 99. w a l l e d elements, as o n pileus, but shorter, up to 160 p m long. C l a m p -
E X C L . - Lepiota tomentella sensu Candusso & L a n z o n i , L e p i o t a : connections present i n a l l tissues.
226-228. 1990; sensu M . B o n i n D o c . m y c o l . 11 (43): 36-37. 1981; H A B I T A T & D I S T R . - Gregarious or m o r e rarely solitary, sapro-
trophic and terrestrial i n deciduous w o o d s o n r i c h loamy-sandy soils,
often r i c h i n l i m e , i n the Netherlands very rare, o n l y i n southern
L i m b u r g ( C a d i e r en K e e r , Orenberg and R i e s e n b e r g ; V a l k e n b u r g ,
Schaelsberg and St. Jansbosch). E n d o f A u g . - O c t . K n o w n f r o m
B e l g i u m , France, D e n m a r k , Germany, and E n g l a n d , everywhere rare
except i n E n g l a n d .
24. Lepiota pilodes V e l l i n g a & Huijser i n Persoonia 15: 226. 1993. -

F i g . 100.
M I S A P P L . - Lepiota tomentella sensu Candusso & L a n z o n i , L e p i o t a :
226-228. 1990; sensu M . B o n i n D o c . m y c o l . 11 (43): 36-37. 1981;
sensu K e l d e r m a n i n C o o l i a 31: 43-44. 1988.
S E L . I C O N . - Z e c c h i n & M i g l . i n M i c o l . ital. 25 (2): p i . 3. 1996.
F i g . 99. Lepiota tomentella. 42. 1990 (as L. tomentella); V e l l i n g a & Huijser i n Persoonia 15: 226-
squamules o n stipe made up o f elements as those o n pileus but 25-160

X 8.0-23 um. Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - Gregarious i n s m a l l groups, often together w i t h
L. tomentella and other Lepiota-species, terrestrial and saprotrophic i n
deciduous woods o n r i c h l o a m y soils, also o n m i n e waste heaps. In the
Netherlands o n l y k n o w n f r o m some localities i n southern L i m b u r g ;
Sept.-Oct. K n o w n from France, G e r m a n y and Italy.
This species comes very close to Lepiota tomentella and was not dis-
tinguished from i t i n the past. Characteristics o f this species are the
orange to ochraceous discoloration o f the pileus surface, the broad
cheilocystidia and the non-septate elements o f the pileus covering.
25. Lepiota griseovirens M a i r e i n B u l l , trimest. S o c . m y c o l . Fr. 4 4 :

37. 1 9 2 8 . - F i g . 101.
Lepiota griseovirens ssp. obscura L o c q . i n B u l l . mens. Soc. l i n n .
L y o n 14: 6 1 - 6 2 . 1 9 4 5 (not valid); Lepiota griseovirens var. obscura M .
B o n i n D o c . m y c o l . 6 (24): 4 4 . 1976; Lepiota obscura (Locq.) Babos
F i g . 100. Lepiota pilodes.
i n A n n l s hist.-nat. M u s . natn. hung. 5 0 : 89. 1958 (not valid); Lepiota
obscura ( M . B o n ) M . B o n i n D o c . m y c o l . 23 (91): 33. 1993. -Lepiota
229, fig. 2. 1993; Z e c c h i n & M i g l . i n M i c o l . ital. 25 (2): 37-41, figs. 3 grangei f. brunneoolivacea Pilat i n A c t a M u s . natn. Prag. 1 I B (2): 9.
& 4. 1996. 1955.
V E R N . N A M E - Valse viltparasolzwam. E X C L . - Lepiota griseovirens sensu D . R e i d i n F u n g . rar. Ic. c o l . 6:
14-16. 1972; sensu M . B o n , F l . m y c o l . Eur. 3, L e p i o t e s : 5 5 . 1993;
Pileus 13-34 m m , expanding f r o m campanulate to convex or plano- Lepiota griseovirens var. griseovirens sensu M . B o n i n D o c . m y c o l . 11
convex, with or without broad u m b o , when y o u n g w i t h m a r g i n exceed- (43): 38. 1981 (in a l l cases L. poliochloodes).
ing lamellae, at centre d u l l orange b r o w n , dark b r o w n or ochraceous M I S A P P L . - Lepiota pseudofelina i n the sense o f many authors, e.g.
b r o w n ( M u . 2.5 Y R 4/4, 10 Y R 3-4/4), towards m a r g i n fading to pale M . B o n , F l . m y c o l . Eur. 3, Lepiotes: 5 5 . 1993; Candusso & L a n z o n i ,
brown-beige (e.g. 10 Y R 7/6), at first at outermost m a r g i n very pale, L e p i o t a : 222-225, 22b. 1990; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g L
w h e n very y o u n g and undamaged w i t h dark fuligineous colours some- 94-95. 1994; Chr. L a n g e i n S v a m p e 3 1 : 4 2 . 1995; M i g l . & C o c c i a i n
times w i t h a hue o f green or olive, d i s c o l o u r i n g orange w h e n touched B o l l . A s s . m i c o l . ecol. R o m a n a 19: 2 0 . 1990.
and w i t h age, c o m p l e t e l y covered w i t h a relatively thin c o v e r i n g o f S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 226. 1995;
small plush-like squamules, at centre closed and velutinous-tomentose, Candusso & L a n z o n i , L e p i o t a : p i . 22a, 22b. 1990 (as L. griseovirens
towards m a r g i n more fibrillose to arachnoid w i t h context showing i n and L. pseudofelina resp.); Chr. L a n g e i n S v a m p e 3 1 : 4 2 . 1995 (as L.
between squamules. L a m e l l a e , L = 31-35,1 = 1-3, moderately distant pseudofelina); L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l . G . B r e s .
to rather crowded, free, not or slightly cream-coloured, w i t h age more 26: 104. 1983; M i g l . & C o c c i a i n B o l l . A s s . m i c o l . ecol. R o m a n a 19:
orange b r o w n i s h tinged or w i t h orange-brown spots, w i t h w h i t i s h or 20. 1990 (as L. pseudofelina); R a i d et a l . i n Svampe 26: 3 8 . 1992.
white, even to flocculose edge. Stipe 15-50 X 2-4 m m , c y l i n d r i c a l with S E L . D E S C R . & FIGS. - F A . v.d. B e r g h i n C o o l i a 23: 54-56. 1980 (as
broadened to bulbous base, fistulose, cream to pale p i n k i s h cream at L. grangei); Candusso & L a n z o n i , L e p i o t a : 218-220, 223-225, figs. 4 0
base, d i s c o l o u r i n g orange to reddish b r o w n especially at base, length- & 4 1 . 1990 (as L. griseovirens var. griseovirens and L. pseudofelina
w i s e innately fibrillose, i n upper h a l f more or less pruinose, lower w i t h resp.); Courtecuisse i n B u l l , semest. S o c . m y c o l . N o r d 4 2 : 8-9, figs.
faint girdles a n d scattered squamules (rarely without), concolorous 12-14.1988; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 94-95. 1994 (as L.
w i t h pileus, w i t h white m y c e l i u m cords at base. Context i n pileus white pseudofelina); K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 5 2 : 236-237.
and dull, i n stipe creamy, especially i n cortex discolouring orange, w i t h
age i n l o w e r part reddish b r o w n (5 Y R 4/4). S m e l l faint, sweetish,
fruity-fungoid or farinaceous, not l i k e L. cristata. Taste not k n o w n .
Spore print white.
Spores i n side-view 7.0-10.0(-11.5) X 3.0-4.0(-4.5) p m , Q = 1.9-
2.8(-3.0), Q a v = 2.2-2.6, c y l i n d r i c a l , s u b c y l i n d r i c a l , w i t h rounded
apex or tapering towards apex, without or w i t h lateral spur, w i t h round-
ed or spur-like base, i n frontal v i e w s u b c y l i n d r i c a l or o v o i d , w i t h dis-
tinct patent hilar appendage, dextrinoid, c o n g o p h i l o u s ; w a l l not
c o l o u r i n g i n C r e s y l B l u e . B a s i d i a (19-)22-34(-38) X 6.5-9.5 u m , 4 -
spored, also some 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a (16-
)20-45(-50) X 5.0-15(-18) u m , n a r r o w l y clavate, clavate, narrowly
u n i f o r m to more or less c y l i n d r i c a l , not coloured. P l e u r o c y s t i d i a
absent. C o v e r i n g o f pileus made up o f erect c y l i n d r i c a l to slightly
fusiform elements, 60-280(-330) X (6.5-)8.0-21 u m , narrowed into
pedicel, without (very rarely w i t h one) clampless septa, w i t h some
basal clavate elements i n between, w i t h parietal and intracellular
b r o w n pigment, w h e n fresh also w i t h grey intracellular pigment.
Stipitipellis a cutis o f narrow 2.0-6.5 p m w i d e c y l i n d r i c a l hyphae; F i g . 101. Lepiota griseovirens.
1936 (as L. pseudofelina); L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l . ering elements, i n d i c a t i n g that this is most p r o b a b l y a y o u n g L .
G . B r e s . 26: 103-107, f i g . 2. 1983; M i g l . & C o c c i a i n B o l l . A s s . m i c o l . grangei.
ecol. R o m a n a 19: 17-19. 1990 (as L. pseudofelina); A . Pears, i n Trans. The name L. griseovirens has been interpreted by some authors i n the
B r . m y c o l . S o c . 29: 192. 1946 (as L. pseudofelina); Romagn. & Locq. same way as is presented here; others regard it as the grey-green species
i n B u l l , trimest. S o c . m y c o l . Fr. 60: 54-55. 1944; V e l l i n g a & Huijser i n with small spores, and short elements i n the pileus covering. F o r an
Persoonia 15: 231-234, fig. 4. 1993. extensive discussion o n the history o f the name, see Vellinga & Huijser
V E R N . N A M E - Grijsgroene parasolzwam. (in Persoonia 15: 234. 1993). B o n ( F l . m y c o l . Eur. 3, Lepiotes: 55.1993)
used L. griseovirens for L. poliochloodes, and both L. pseudofelina and
Pileus 12-32 m m , at first conico-campanulate, expanding to plano-con- L. obscura for L. griseovirens i n the interpretation given here.
vex o r slightly plano-concave w i t h broad umbo, when y o u n g w i t h
w h i t i s h m a r g i n exceeding lamellae, at centre w i t h a closed velvety- 26. Lepiota poliochloodes V e l l i n g a & H u i j s e r i n Persoonia 15: 229.
tomentose covering w i t h s m a l l tufts (plush-like), dark grey to b l a c k o r 1 9 9 3 . - F i g . 102.
greyish o l i v e - b r o w n , sometimes w i t h a hue o f green, blue or violet, M I S A P P L . - Lepiota griseovirens sensu M . B o n , F l . m y c o l . Eur. 3,
towards m a r g i n cracked into small wart-like squamules, paler than at Lepiotes: 55. 1993; sensu D . R e i d i n F u n g . rar. Ic. c o l . 6: 14-16. 1972;
centre to pale olive-beige b r o w n i s h o r grey-brown ( M u . 2.5 Y R 3/2, 5 Lepiota griseovirens var. griseovirens sensu M . B o n i n D o c . m y c o l . 11
Y 2.5/1 to 2.5 Y 7/6, 2.5 Y 6/2, 10 Y R 6/4) o n a pale cream to isabella (43): 38. 1981.
background, slightly pallescent and more brownish w i t h age and some- S E L . I C O N . - K n u d s e n & Vesterholt, Truede Storsvampe D a n m a r k :
times d i s c o l o u r i n g y e l l o w - b r o w n , orange-brown o r salmon-brown at 33. 1990 (as L. griseovirens); D . R e i d i n F u n g . rar. Ic. c o l . 6: p i . 43c &
margin. L a m e l l a e , L = 36-41,1 = 1-5, crowded, free, subventricose o r d, 1972 (as L. griseovirens).
ventricose and rounded near stipe, 2-4 m m broad, white to cream (10 S E L . D E S C R . & FIGS. - D . R e i d i n F u n g . rar. Ic. c o l . 6: 14-16. 1972
Y R 8/2), w h e n y o u n g sometimes w i t h greyish tinge, turning orange (as L. griseovirens).
b r o w n i s h or w i t h orange-brown spots w h e n old, w i t h white finely floc- V E R N . N A M E - Grauwgroene parasolzwam.
culose edge. Stipe 20-60 X 2-5 m m , c y l i n d r i c a l and slightly broadened
at base, fistulose, i n upper half pale, w h i t i s h to greyish, lengthwise fib- Pileus 10-31 m m , w h e n y o u n g spherical to ellipsoid, expanding to c o n -
rillose, without real annulus, but especially when y o u n g w i t h an annus vex, campanulate, finally applanate w i t h l o w obtuse u m b o , at first with
lar zone, i n the l o w e r half covered w i t h bands or many scattered a closed relatively thin covering, green-grey, grey-olive, grey-brown,
grey-black to grey-brown squamules as o n pileus; background turning light o l i v e - b r o w n ( M u . 5 Y 6-5/3-4, 1 0 Y R 5-2/3, 2 . 5 Y R 6-5/2-4),
orange b r o w n i s h at base; w i t h white m y c e l i u m cords. Context i n pileus towards margin splitting up into very s m a l l fibrillose squamules with
white, w i t h glassy line above lamellae, i n stipe cortex pale y e l l o w or age; underlying context d i s c o l o u r i n g , p i n k - y e l l o w to light orange-
brownish, at base o f stipe turning orange-brown w i t h age, inner part o f brown ( 1 0 Y R 8-7/6-8, 2 . 5 Y 8-7/6-8); margin w h e n y o u n g fringed by
stipe white cottony. S m e l l i n y o u n g and undamaged specimens fruity, velar remnants, exceeding lamellae. L a m e l l a e , L = 36-42,1 = l-3(-7),
w h e n damaged o r w i t h age more l i k e L. cristata, l a c k i n g the rubber moderately crowded, free, (sub)ventricose, up to 4 m m broad, round-
component, astringent. Taste rather strong, unpleasant, resembling ed near stipe, cream-coloured, w i t h white somewhat flocculose edge,
smell. Spore print white. discolouring light orange-brown o n h a n d l i n g and w i t h age. Stipe 20-
Spores i n side-view (6.0-)6.5-9.5(-11.0) X (3.0-)3.5-4.5 p m , Q = 55 X 2-4 m m , slightly broadening towards base, w h e n y o u n g white
1.6-2.5(-2.7), Q a v = 1.8-2.4, o b l o n g to c y l i n d r i c a l , w i t h truncate base, and lengthwise fibrillose, w i t h scattered squamules a n d girdles l i k e
the bigger the more distinctly spurred, i n frontal v i e w ± c y l i n d r i c a l , covering o n pileus, at base discolouring orange when touched and with
w i t h thickened w a l l and patent hilar appendage, strongly dextrinoid, age. Context i n pileus white, i n stipe white at first, i n l o w e r parts later
congophilous; w a l l i n C r e s y l B l u e not colouring. B a s i d i a 1 7 - 3 1 X 5 . 5 - on, m a i n l y i n the cortex, brawn-orange to red-brown. S m e l l faint,
9.0 p m , 4-spored, often also 2-spored. L a m e l l a edge sterile w i t h tufts slightly sweetish o r unpleasant, not l i k e L. cristata. Taste u n k n o w n .
o f cheilocystidia. C h e i l o c y s t i d i a (16-)20-42(-50) X 5.0-10.5 um, usu- Spore print white.
a l l y n a r r o w l y utriform, also o b o v o i d to narrowly clavate to nearly Spores i n side-view 6.0-8.0(-9.0) X (3.0-)3.5-4.5 p m , Q = 1.4-2.3,
c y l i n d r i c a l . P l e u r o c y s t i d i a absent. Pileus covering made up o f tightly Qav = 1.7-2.0, usually o b l o n g to c y l i n d r i c a l , w i t h rounded o r truncate
packed, erect rather straight elements, (55-)85-320(-400) X 6.5-20(- base, more rarely w i t h distinct spur, showing lateral hilar appendage,
24) p m , c y l i n d r i c a l o r fusiform, often w i t h attenuate apex, rarely fur- i n frontal v i e w more e l l i p s o i d , dextrinoid, c o n g o p h i l o u s ; w a l l not
cate, w i t h g r e y i s h - b r o w n i s h parietal a n d i n basal hyphae encrusting coloured i n C r e s y l B l u e . B a s i d i a 20-32 X 6.0-9.0 u m , 4-spored, some
pigment, also w i t h some vague coloured granules i n H 0 , sometimes 2 (3-)2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a (12-)16-35(-40) X
in l o w e r part w i t h l ( - 2 ) clampless septa. Stipitipellis a cutis o f c y l i n - 4 . 0 - 1 0 ( - l 1) u m , variable i n shape, l o n g and slender, ± cylindrical, short
drical, not c o l o u r e d hyphae 2.5-8 p m i n diameter; elements o f squa-
mules l i k e those o n pileus but slightly shorter 35-240 X 6.5-22 p m .
Clamp-connections present i n a l l tissues.
H A B I T A T & D I S T R . - Gregarious i n s m a l l groups, terrestrial a n d
saprotrophic i n deciduous woods o n sandy to l o a m y soils r i c h i n
humus and nutrients; k n o w n i n the Netherlands from several localities
in the inner coastal dunes, the river area and the western clay-areas,
and f r o m southern L i m b u r g . I n the Netherlands the most c o m m o n
species o f the dark, greenish taxa. E n d o f A u g . - N o v . Rare i n Europe.
M o s t E u r o p e a n authors c a l l e d this species Lepiota pseudofelina J.

L a n g e . H o w e v e r , L a n g e ' s description and figure ( F l . agar. dan. 1: p i .
12C. 1935) depict a slender s p e c i m e n w i t h a r a p i d l y d i s c o l o u r i n g
pileus, l o n g spores, and w i t h intracellular pigment i n the p i l e u s c o v - F i g . 102. Lepiota poliochloodes.
clavate, narrowly utriform, often w i t h basal broad element, colourless S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 225. 1995;
w i t h fine granular contents w h e n fresh. P l e u r o c y s t i d i a not observed. Candusso & L a n z o n i , L e p i o t a : p i . 2 1 . 1990; Enderle i n Z . M y k o l . 5 1 :
C o v e r i n g o n pileus rather irregular and made up o f elements (45-)70- between p. 16 and 17. 1985; J . L a n g e , F l . agar. dan. 1: p i . 1 0 A . 1935;
180(-200) X 6.5-20(-26) p m , often broadened at m i d d l e and narrowed L a n z o n i i n B o l l . G r u p p o m i c o l . G . Bres. 29: 85. 1986; L o c q . i n B u l l ,
into pedicel, w i t h 0-1 (-2) clampless septa, w i t h short clavate elements trimest. Soc. m y c o l . Fr. 60: p i . 2, fig. 3. 1944; R a i d et a l . i n Svampe
in between; pigment b r o w n , parietal, i n l o w e r parts also intracellular, 26: 38. 1992.
in basal parts o f terminal elements encrusting. Stipitipellis a cutis o f S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 213-216.
narrow hyphae, 1.5-6.5 u m broad; elements o f squamules and girdles 1990; Enderle i n Z . M y k o l . 5 1 : 19-22. 1985; H e r i n k i n C e s k a M y k o l .
as those o n pileus o r shorter 25-120(-180) X 9.0-25(-33) u m . C l a m p - 16: 228-234. 1962; K e l d e r m a n i n C o o l i a 3 1 : 91-92, f i g . 2. 1988;
connections present i n a l l tissues. In fresh material a l l elements are full L a n z o n i i n B o l l . G r u p p o m i c o l . G . B r e s . 29: 83-87, fig. 2. 1986; L o c q .
of little granules o r o i l drops; sometimes these granules are also v i s i - in B u l l , trimest. Soc. m y c o l . Fr. 60: 41-42. 1944; V e l l i n g a & Huijser i n
ble i n exsiccata. Persoonia 15: 235-237, fig. 5. 1993.
H A B I T A T & D I S T R . - Gregarious i n s m a l l groups, terrestrial and V E R N . N A M E - Groenschubbige parasolzwam.
saprotrophic i n deciduous w o o d s o n nutrient-rich soils, l o a m y o r
sandy, but also o n m i n e waste heaps; very rare i n the Netherlands, only Pileus 13-40 m m , w h e n y o u n g campanulate o r hemispherical w i t h
k n o w n f r o m V a l k e n b u r g (Schaelsberg) a n d K e r k r a d e (former m i n e inflexed margin, expanding to plano-convex w i t h u m b o , at centre
Laura-Julia); O c t . K n o w n f r o m D e n m a r k , France and Great B r i t a i n , green-black, dark grey-green, grey-blue o r blue-green, then green-
apparently very rare. brown o r blue-brown to b r o w n , velvety-tomentose w i t h s m a l l erect
squamules, around centre w i t h concolorous o r slightly paler small to
T h i s species i s k n o w n i n European literature as Lepiota griseovirens very s m a l l uplifted o r erect squamules o n pale cream to pale b r o w n
sensu D . R e i d (in F u n g . rar. Ic. c o l . 6: 14-16. 1972). B u t the name L. background, w i t h age often d i s c o l o u r i n g orange-brown, more glabrous
griseovirens is best applied to the species formerly k n o w n under the and squamules less conspicuous; margin irregular and w i t h some velar
name L. pseudofelina J . L a n g e . See notes o n L. griseovirens and for an remnants especially when y o u n g . L a m e l l a e , L = c. 30,1 = 1-3, m o d -
elaborate discussion see V e l l i n g a & H u i j s e r ( i n Persoonia 15: 234. erately to rather crowded, free, ventricose o r broadly ventricose, up to
1993). 6 m m broad, when y o u n g w h i t i s h to cream, w h e n o l d w i t h orange-
Lepiota poliochloodes differs f r o m L. griseovirens i n the lighter brown spots, w i t h white finely floccose edge. Stipe 25-60 X 3-6 m m ,
colours o f the basidiocarp, the smaller spores and the shorter elements c y l i n d r i c a l o r slightly broadening towards base or w i t h subbulbous
of the pileus covering. It is m u c h rarer than L. griseovirens. base, fistulose w i t h age, at apex glabrous and w h i t i s h o r pale p i n k i s h ,
lower d o w n more orange (5 Y R 5/8) o r orange-brown, especially o n
27. Lepiota grangei (Eyre) K i i h n e r i n B u l l . mens. S o c . l i n n . L y o n 3: handling, i n lower half w i t h some scattered girdles o f squamules, as
79. 1934. - F i g . 103. on pileus, b l a c k i s h green-blue, blue-green o r grey-green, without dis-
Schulzeria grangei E y r e i n A . L . S m . & R e a i n Trans. B r . m y c o l . tinct annulus. Context i n pileus white, i n stipe concolorous w i t h sur-
Soc. 2: 3 7 . 1903; Hiatula grangei (Eyre) W . G . S m . , Synopsis B r i t . face. S m e l l unpleasant, rubber-like, musty-stuffy o r l i k e L. cristata.
B a s i d i o m y c : 2 7 . 1 9 0 8 ; Lepiotula grangei (Eyre) H o r a k i n N . Z . J . B o t . Taste 'mild-aromatic'. Spore print white.
18: 184. 1980. - Lepiota ochraceocyanea K i i h n e r i n B u l l . mens. S o c . Spores i n side-view (7.5-)8.0-13.0(-14.5) X 3.5-4.5(-5.0) p m , Q =
l i n n . L y o n 3: 4 3 . 1934. 1.9-3.0(-3.2), Q a v = 2.2-2.9, w i t h truncate to distinctly spurred base,
M I S A P P L . - Lepiota forquignonii sensu B a r b i e r i n B u l l . mens. Soc. in frontal v i e w c y l i n d r i c a l o r subcylindrical, not or uniguttulate, slight-
l i n n . L y o n 3: 76-78. 1934. ly thick-walled, w i t h distinct patent hilar appendage, immediately dex-
E X C L . - Lepiota grangei sensu F A . v.d. B e r g h i n C o o l i a 23: 54-56. trinoid, congophilous; w a l l not c o l o u r e d i n C r e s y l B l u e . B a s i d i a
1980 ( = L. griseovirens); sensu H o r a k i n N . Z . J . B o t . 18: 184-185, (20-)22-35 X 6.5-11.5 p m , 4-spored, rarely some 2-spored. L a m e l l a
fig. 2. 1980 ( = Lepiota spec). edge sterile, made up o f cheilocystidia, (16-)20-45(-55) X 5.0-13(-15)
um, ± c y l i n d r i c a l , narrowly utriform or narrowly clavate, colourless,
often o n broad basal element or septate i n lower part. Pleurocystidia
absent. Pileus covering made up o f more o r less erect elements, (30-
)50-300(-400) X (6.5-)8.0-20(-24) p m , c y l i n d r i c a l w i t h rounded apex,
attenuate towards base, septate, w i t h up to 5 clampless septa, rarely not
septate, w i t h thickened w a l l s and w i t h basal clamp-connection; p i g -
ment brownish parietal and intracellular, w h e n fresh blue-green intra-
cellular, i n dried material i n granules, v i s i b l e i n H 0 . S t i p i t i p e l l i s a
2
cutis o f narrow 2.0-7.0 p m w i d e , c y l i n d r i c a l , colourless hyphae; ele-

ments o f squamules and girdles as those on pileus but less septate, 35-
310 X 8.0-21 p m . Clamp-connections present i n a l l tissues.
H A B I T A T & DISTR. - Gregarious, i n s m a l l groups, terrestrial a n d
saprotrophic i n deciduous woods, especially i n Fagus-woods o n ± cal-
careous, loamy o r sandy soils and soils r i c h i n humus, but also on mine
waste heaps under Salix and Betula. I n the Netherlands very rare ('s-
Graveland; C a d i e r en Keer, Riesenberg; Kerkrade, m i n e L a u r a - J u l i a ;
Valkenburg, Schaelsberg); more c o m m o n i n south-eastern B e l g i u m ,
western Germany (Eifel) etc. E n d o f Sept.-Oct. K n o w n f r o m Eurasia,
and also recorded f r o m A r g e n t i n a and N e w Zealand, but these reports
may concern closely related species.
Fig. 103. Lepiota grangei.
138 FLORA AGARICTNA NEERLANDICA
Sect. Lilaceae M . B o n sometimes pale b r o w n i s h buff; the c o l o u r s fading w i t h age and

weather; b a c k g r o u n d white to cream-coloured, r a d i a l l y adnate-fibril-
Pileus c o v e r i n g a h y m e n i d e r m , made u p tightly packed clavate, nar- lose; remnants o f annulus sometimes attached to the m a r g i n .
r o w l y clavate to c y l i n d r i c a l elements; spores subglobose to ellipsoid or L a m e l l a e , L = 4 0 - 6 0 , 1 = 1-3, moderately c r o w d e d to c r o w d e d , free,
spurred. ventricose u p to 4 m m broad, w h i t e to cream, rarely furcate, w i t h
white, even to f i n e l y f l o c c u l o s e edge. S t i p e 15-80 X 2-8.5 m m ,
28. Lepiota cristata (Bolt.: Fr.) K u m m . , Fiihr. P i l z k . : 1 3 7 . 1 8 7 1 . - F i g . c y l i n d r i c a l or s l i g h t l y broadened or even bulbous at base, fistulose
104. w i t h age, at apex w h i t i s h a n d at base p i n k i s h or s o r d i d b r o w n i s h ,
Agaricus cristatus B o l t . , H i s t . F u n g . H a l i f a x 1: 7. 1788, n o n innately s i l v e r y fibrillose, w i t h or without annulus, at base often w i t h
Agaricus cristatus S c o p . , 1774; Agaricus cristatus B o l t . : Fr., Syst. c l u m p o f s o i l particles, o r w i t h w h i t e m y c e l i u m strands. A n n u l u s
m y c o l . 1: 2 2 . 1821; Lepiotula cristata (Bolt.: Fr.) H o r a k , S y n . G e n . loosely attached, straight or ascending, w h i t i s h o n upper side, b r o w n -
Agar.: 3 3 8 . 1968. - Lepiota cristata var. felinoides M . B o n in Doc. ish or w i t h s m a l l b r o w n squamules o n l o w e r side. C o n t e x t i n pileus
m y c o l . 11 (43): 34. 1981; Lepiota felinoides ( M . B o n ) P . D . Orton i n d u l l a n d w h i t e , i n stipe c o n c o l o r o u s w i t h p i n k i s h surface. Smell
Notes R . bot. G d n E d i n b . 4 1 : 591. 1984, non Lepiota felinoides Peck, strong, sweet a n d w i t h a rubber component, very rarely without any
1900; Lepiota subfelinoides B o n & O r t o n i n P.D. O r t o n i n D o c . m y c o l . s m e l l at a l l . Taste as s m e l l , unpleasant. Spore print c o l o u r white to
14 (56): 56. ( ' 1 9 8 4 ' ) 1985. - Lepiota cristata var. exannulata M .Bon cream, but l e m o n - y e l l o w a c c o r d i n g to B r e i t e n b a c h & K r a n z l i n ( P i l z e
i n D o c . m y c o l . 11 (43): 34. 1981. - Lepiota cristata var. pallidior S c h w e i z 4 : 197. 1995).
B o u d . ex M . B o n i n D o c . m y c o l . 11 (43): 34. 1981. Spores i n side-view (5.0-)5.5-8.5(-10.0) X 2.5-3.5 p m , Q = 1.8-
E X C L . - Lepiota cristata sensu A r o r a , M u s h r . demyst., E d . 2: 306. 2.9, Q a v = 2.0-2.5, truncate and narrowly triangular, rarely w i t h dis-
1986 ( = L. casteneidisca); sensu Sundb. i n M y c o t a x o n 34: 240-242. tinct, saccate spur, c y l i n d r i c a l i n frontal view, slightly t h i c k - w a l l e d ,
1989 ( = L. castaneidisca and L. cristata). dextrinoid, but not strongly so, congophilous, not metachromatic i n
S E L . I C O N . - A r n o l d s & V e e r k a m p , G i d s Paddest. Meetnet: 7 3 . C r e s y l B l u e . B a s i d i a 14-28 X (4.0-)5.0-7.5 p m , 4-spored, rarely inter-
1999; Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 221, 222. 1995 (as L . m i x e d 4 - and 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 17-55 X
cristata and L. cristata var. pallidior resp.); Candusso & L a n z o n i , 7.0-18 p m , clavate, broadly clavate, spheropedunculate or n a r r o w l y
L e p i o t a : p i . 2 0 . 1990; L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l ^ clavate, slightly t h i c k - w a l l e d , colourless. Pleurocystidia absent. Pileus
B r e s a d o l a 26: 105. 1983; Lavorato i n S c h w e i z . Z . P i l z k . 66: 127. 1988 covering a h y m e n i d e r m , made up o f erect elements, 16-70 X 5.0-18
(as L. cristata wax. felinoides); R . P h i l l i p s , Paddest. S c h i m m . : 29. 1981; pm, c y l i n d r i c a l - a n d often "subcapitate to n a r r o w l y clavate, c l o s e l y
R y m a n & H o l m a s e n , Svampar: 413. 1984. packed, slightly t h i c k - w a l l e d a n d pigment b r o w n and parietal.
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 207-210, fig. Stipitipellis a cutis made up o f c y l i n d r i c a l , 4.0-12 p m w i d e , colourless
37. 1990; H o r a k , S y n . G e n . Agar.: 338-339. 1968 (as Lepiotula crista- hyphae. Clamp-connections present i n a l l tissues.
ta); K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 82-83. 1994. H A B I T A T & D I S T R . - Gregarious, sometimes i n b i g groups, rarely
VERN. NAME - Stinkparasolzwam. solitary, saprotrophic and terrestrial, o n humus-rich, often nutrient-rich
soil, i n a w i d e variety o f habitats, from open road-sides to coniferous
P i l e u s 10-75 m m , w h e n y o u n g campanulate, conico-campanulate, woods, often i n ruderal and disturbed places; i n a l l parts o f the
e x p a n d i n g to p l a n o - c o n v e x or applanate w i t h or without l o w u m b o , Netherlands c o m m o n (for a distribution map, see A r n o l d s et a l . , O v e r z .
at centre w i t h a c l o s e d , smooth, c o v e r i n g , often c a l l e d a 'calotte' and Paddest. Nederland: 288. 1995); the most c o m m o n Lep/ota-species i n
around this centre w i t h rather broad c o n c e n t r i c a l squamules, some- the Netherlands. J u l y - N o v. Widespread i n a l l temperate and mediter-
times w i t h s m a l l , m o r e granular, squamules; most t y p i c a l l y the ranean parts o f the Northern Hemisphere.
c o l o u r o f the calotte a n d squamules i s orange-brown to reddish
b r o w n ( M u . 7.5 Y R 4-5/6), sometimes they are very dark b r o w n , Lepiota saponella B o d i n & P r i o u , f r o m the west coast o f France,
comes very close, but differs m a c r o s c o p i c a l l y i n the smaller squa-
mules o n the pileus, <and the strong s m e l l o f soap; m i c r o s c o p i c a l l y the
broader, more triangular-shaped spores are a g o o d differentiating
character ( B o d i n ifej^riou i n B u l l , trimest. S o c . m y c o l . Fr. 110: 125-
137. 1994). y
Lepiota apatelia and L. hymenoderma resemble L. cristata closely,
but can be easily distinguished o n account o f the e l l i p s o i d spores.
V e l l i n g a & Huijser (in B e l g . J . B o t . 131: 198-199. ('1998') 1999)
showed that L. cristata is more closely related to species w i t h e l l i p s o i d
spores and a h y m e n i f o r m pileus covering than to species w i t h spurred
spores and l o n g elements i n the pileus covering.
29. Lepiota apatelia V e l l i n g a & H u i j s e r i n B e l g . J . B o t . 131: 196.

('1998') 1 9 9 9 . - F i g . 105.
M I S A P P L . - Lepiota cristatoides sensu B i z i o et al. i n R i v . M i c o l . 36:
226-227. 1993; sensu A . H a u s k n . i n Enderle & Krieglst. i n Z . M y k o l .
55: 63-64. 1989; sensu Huijser & V e l l i n g a i n A r n o l d s et a l , O v e r z .
Paddest. N e d e r l a n d : 290-291. 1995; sensu K e l d e r m a n , Parasolzw.
Z u i d - L i m b u r g : 132-133. 1994; sensu V e l l i n g a & Huijser i n C o o l i a 40:
p i . 3. 1997.
S E L . I C O N . - B i z i o et al. i n R i v . M i c o l . 36: 226. 1993 (as L. crista-
F i g . 104. Lepiota cristata. toides); V e l l i n g a & Huijser i n C o o l i a 40: p i . 3 . 1 9 9 7 (as L. cristatoides).
H A B I T A T & DISTR. - Gregarious, often i n b i g groups, saprotrophic

and terrestrial, i n deciduous woods, also o n m i n e waste heaps. In the
Netherlands very rare, k n o w n from two localities i n southern L i m b u r g
(Brunssum, m i n e H e n d r i k ; M a r g r a t e n , Eysderbos), Sept.-Oct. Very
rare i n Europe, k n o w n also from A u s t r i a , Germany, and Italy.
Lepiota apatelia m a c r o s c o p i c a l l y resembles L. cristata very closely.

The shape o f the spores o f course is the easiest character to separate the
two species: spurred i n L. cristata and e l l i p s o i d i n L. apatelia.
Furthermore, the annulus i n L. apatelia c l i n g s to the pileus margin,
w h i l e i n L. cristata it can often be found o n the stipe, and the smell o f
L. apatelia lacks i n general the rubber component, so characteristic o f
the smell o f L. cristata.
Another closely related species is L. cristatoides, characterized b y
a smoother and pinker pileus, a r i n g that clings to the stipe, the smell
of L. cristata, and the slightly smaller, non-dextrinoid spores.
30. Lepiota cristatoides E i n h . i n Ber. bayer. bot. G e s . 44: 3 1 . 1 9 7 3 -

F i g . 106.
F i g . 105. Lepiota apatelia. E X C L . - Lepiota cristatoides sensu B i z i o et a l . i n R i v . M i c o l . 36:
226-227. 1993; sensu A . H a u s k n . i n Enderle & Krieglst. i n Z . M y k o l .
S E L . D E S C R . & F I G S . - B i z i o et a l . i n R i v . M i c o l . 36: 226-227. 1993 55: 63-64. 1989; sensu K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 132-
(as L. cristatoides); K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 132-133. 133. 1994; sensu V e l l i n g a & Huijser i n C o o l i a 40: p i . 3. 1997 (in a l l
1994 (as L. cristatoides). cases L. apatelia).
V E R N . N A M E - Valse stinkparasolzwam. MISAPPL. - Lepiota lilacea var. pallida f. 'zonder cystiden'
K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 137. 1994. - Lepiota rufipes f.
Pileus 10-40(-60) m m , campanulate w i t h inflexed margin when young, phaeophylla sensu B i z i o et a l . i n Riv. M i c o l . 36: 240-241. 1993.
expanding to plano-convex, applanate, or w i d e l y conical with undulat- S E L . I C O N . - B i z i o et a l . i n Riv. M i c o l . 36: 240. 1993 (as L. rufipes
ing l i m b , w i t h or without blunt umbo, w i t h uplifted margin i n o l d spec- f. phaeophylla); B o n & G . Riousset i n D o c . m y c o l . 22 (85): 64-65, p i .
imens, orange-brown to y e l l o w brownish ( M u . 7.5 Y R 6/6-8, 5-6/6, 10 I B . 1992; M o s . & J u l . , Farbatl. B a s i d i o m y c . I l l L e p i o t a 9. 1990.
Y R 7/8) and glabrous at centre, around centre breaking up into broad S E L . D E S C R . & FIGS. - E i n h . i n Ber. bayer. bot. G e s . 44: 31-32. 1973;
concentrically arranged, adnate patches, fading i n colour towards mar- K e l d e r m a n i n C o o l i a 35: 79-82, fig. 2. 1992 (as L. lilacea var. pallida);
gin, rarely breaking up i n smaller, granular patches, paler than central K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 136-137. 1994 (as L. lilacea
part, o n a pale cream to w h i t i s h , fibrillose-silky background, when var. pallida f. 'zonder cystiden'); Laber et a l . i n Siidwestd.
young w i t h white velar remnants at margin. L a m e l l a e , L = 30-45,1 = Pilzrundschau 31 (1): 4-5. 1995.
1-3, rather c r o w d e d to moderately distant, sometimes anastomosing, V E R N . N A M E - K l e i n e stinkparasolzwam.
free, ventricose o r segmentiform, cream w i t h pale cream-orange-pink

tinge when l o o k e d into, w i t h even o r eroded concolorous edge. Stipe Pileus 9-30 m m , convex, plano-convex to applanate w i t h l o w umbo,
25-60 X 2-5 m m , c y l i n d r i c a l and i n some specimens widened at base, brown or p i n k i s h orange ( M u . 7.5 Y R 6-7/6; K . & W . 6 B - C 3 - 5 ) at cen-
hollow, w h e n y o u n g shiny cream-coloured, lengthwise adnately fibril- tre, around centre breaking up into s m a l l concentrical patches, often
lose, later from base upwards discolouring purple vinaceous red (c. 2.5 only i n marginal zone, m a k i n g a fairly cream impression, glabrous at
Y R 3/4), w i t h vanishing w h i t i s h b l o o m , w i t h white basal tomentum, centre, not rugulose o r micaceous. L a m e l l a e , L = 30-40,1 = 1, m o d -
and white m y c e l i a l cords. A n n u l u s white, membranaceous, often c l i n g -
ing to pileus margin, sometimes o n stipe, easily removed. Context white
to cream i n pileus, brownish to reddish i n stipe, concolorous w i t h sur-
face. S m e l l strong, sweet, also when basidiocarp crushed, rarely with
the rubber component o f the smell o f L. cristata, sometimes a bit astrin-
gent. Taste like smell. Spore print white.
Spores 4.5-6.5 X 3.0-4.0 u m , Q = 1.4-1.8(-2.0), Q a v = 1.4-1.65,
e l l i p s o i d to o b l o n g i n side-view, often w i t h straight abaxial side, ellip-
soid to o b l o n g i n frontal view, c o l o u r i n g orange-brown i n M e l z e r ' s
Reagent, but not strongly so, and often after 5-10 minutes, not or faint-
ly metachromatic i n C r e s y l B l u e . B a s i d i a 15-27(-32) X 5.0-8.0 p m , 4-
spored, rarely some 2-spored present. Lamella edge fertile;
c h e i l o c y s t i d i a absent. P l e u r o c y s t i d i a absent. Pileus c o v e r i n g a euhy-
m e n i d e r m w i t h tendencies to a transition between a h y m e n i d e r m and
an epithelium, made up o f narrowly clavate to spheropedunculate ter-
0QQ00
m i n a l elements, 15-45(-60) X 10-20(-25) u m , w i t h slightly thickened
b r o w n walls, some elements w i t h b r o w n intracellular pigment; under-
l y i n g trama hyphae w i t h colourless encrusting pigment. Stipitipellis a
cutis o f c y l i n d r i c a l , colourless hyphae, 3.5-4.5 p m i n diameter. C l a m p -
connections present i n a l l tissues. F i g . 106. Lepiota cristatoides.
erately c r o w d e d , free, ventricose up to 2 m m broad, cream-coloured Pileus 5-32 m m , hemispherical when young, expanding to plano-con-
(10 Y R 8/3) w i t h concolorous o r white, even to eroded, edge. Stipe 15- vex or almost applanate, w i t h o r without (low) umbo, at centre o n
50 X 1-3.0 m m , c y l i n d r i c a l , fistulose, whitish or cream, (sub)fibrillose u m b o smooth, chestnut-brown, dark b r o w n ( M u . 10 Y R 3/2-3, 5-7.5
and shiny at apex, more distinctly fibrillose i n lower part on p i n k i s h to Y R 3/4), around u m b o split u p into s m a l l rather granulose, radially
vinaceous b a c k g r o u n d (7.5 Y R 8-7/6). A n n u l u s fugacious and fibril- arranged patches, slightly paler than centre (10 Y R 3/4, 4/3) towards
lose-membranaceous, or distinct as a tube around stipe and m e m - margin fading to pale b r o w n (7.5 Y R 6/6; 10 Y R 4/6 a n d paler), o n
branaceous, rarely absent. Context w h i t i s h cream i n pileus, i n stipe cream-coloured r a d i a l l y fibrillose, s i l k y background; m a r g i n not
w i t h p i n k i s h tinges, especially at base. S m e l l , especially o f cut basid- exceeding lamellae. L a m e l l a e , L = 25-36, 1 = 1-3, rather distant to
iocarps, l i k e the rubber component o f the smell o f L. cristata, some- moderately crowded, free, slightly ventricose to ventricose, up to 6 m m
times w i t h spermatic component o r like chlorine. Taste not k n o w n . wide, cream w i t h greyish-brownish tinge (paler than 7.5 Y R 7/4. 10
Spore print off white. Y R 7/3), w i t h even, concolorous o r w h i t i s h flocculose edge. Stipe 23-
Spores 3.5-5.0(-5.5) X 2.5-3.5 p m , Q = 1.3-1.7(-1.85), Q a v = 1.4- 4 0 X 1.5-3 m m , c y l i n d r i c a l , narrowly fistulose, cream at apex, cream
1.65, e l l i p s o i d to o b l o n g , not dextrinoid, hardly c o l o u r i n g i n C o n g o w h i t i s h at base, slightly d i s c o l o u r i n g orange-like at base, covered i n
R e d , metachromatic i n C r e s y l B l u e , rather thin-walled. B a s i d i a 16-25 adnate w h i t i s h fibrils, without annulus, i n l o w e r 1/3 o f stipe length
X 4.0-7.0 p m , 4-spored, rarely intermixed w i t h 2-spored. L a m e l l a edge either w i t h faint girdles o r w i t h scattered b r o w n squamulose floccules.
fertile; c h e i l o c y s t i d i a absent. Pleurocystidia absent. Pileus covering a Context d u l l and w h i t i s h o r cream i n pileus, glassy o r silky, white i n
h y m e n i d e r m o r irregular h y m e n i d e r m w i t h several layers o f elements; stipe and faintly orange (7.5 Y R 7/5) i n base w h e n o l d . S m e l l strong,
terminal elements clavate to almost ellipsoid-globose, (10-)15-50(-55) w i t h both the fruity and the rubber component o f the s m e l l o f L. crista-
X 8-25(-35) p m , without or w i t h y e l l o w i s h parietal pigment, w i t h scat- ta. Taste unpleasant, like smell. Spore print probably white.
tered some narrow elements up to 5.0 p m w i d e w i t h intracellular b r o w n Spores i n side-view 4.0-6.0(-7.0) X 2.5-3.5 p m , Q = 1.4-1.9(-2.0),
pigment; pigment encrusting i n l o w e r layers. S t i p i t i p e l l i s a cutis o f Qav = 1.6-1.7, ellipsoid, oblong to cylindrical, slightly o v o i d i n frontal
c y l i n d r i c a l , colourless hyphae, 2.0-4.0 p m i n diameter. C l a m p - c o n - view, not or i n some stages slightly c o l o u r i n g pale orange-red i n
nections present i n a l l tissues studied. M e l z e r ' s Reagent o r L u g o l , pale p i n k i n C o n g o R e d , metachromatic i n
H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial, espe- • C r e s y l B l u e . B a s i d i a 13-24 X 5.5-7.0 um, 4-spored, rarely 2-spored.
c i a l l y i n nutrient-rich places, for instance o n the verge o f sunken roads, j L a m e l l a edge sterile. C h e i l o c y s t i d i a 11.5-37 X 6.0-10.5 p m , clavate
or i n the margins o f woods. In the Netherlands very rare, k n o w n from with rounded apex, slightly thick-walled, refractive, both i n fresh and i n
several localities i n southern L i m b u r g . Sept.-Oct. A l s o recorded from dried material. Pleurocystidia absent. Pileus covering a hymeniderm,
other European countries (France, Germany and Italy). made up o f narrowly clavate to c y l i n d r i c a l elements 12-63 X 8-13(-25)
pm, w i t h slightly thickened pale b r o w n walls, interspersed w i t h some
Lepiota cristatoides differs f r o m L. rufipes i n the presence o f an annu- narrow, up to 5 p m w i d e , c y l i n d r i c a l elements w i t h intracellular p i g -
lus, absence o f c h e i l o c y s t i d i a and the relatively longer spores (average ment, covered b y a hyaline or brown-coloured gelatinous layer, 10-12
length 4.3-4.8 p m , and Q a v = 1.4). p m thick; underlying elements w i t h b r o w n parietal to encrusting p i g -
Lepiota mediorosea C o n t u (in B o l . S o c . m i c o l . M a d r i d 2 3 : 4 4 . ment. Stipitipellis a cutis, made up o f c y l i n d r i c a l , colourless hyphae,
1998), described f r o m S a r d i n i a , comes very close, but differs i n the 2.0-4.0 p m wide, w i t h encrusting hyaline pigment. Elements i n floc-
smaller spores, v i z . 3.0-4.2 X 2.0-2.8 p m . cules as those o n pileus. Clamp-connections present i n a l l tissues.
See under L. rufipes and i n V e l l i n g a & Huijser (in B e l g . J . B o t . 131: H A B I T A T & DISTR. - Gregarious, saprotrophic and terrestrial, o n
199-201. ( ' 1 9 9 8 ' ) 1999) for more comments o n this group o f species. clayey to loamy soil, r i c h i n nutrients and often o n calcareous bedrock,
in deciduous and coniferous woods and plantations. Very rare i n the
31. Lepiota hymenoderma D . R e i d i n F u n g . rar. Ic. c o l . 1: 24. 1966. Netherlands, o n l y k n o w n from several localities i n southern L i m b u r g
- F i g . 107. (e.g. C a d i e r en Keer, Orenberg; E l s l o o , Bunderbos), Sept.-Oct.
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 35a. 1990; M i g l . & Widespread but rare i n E u r o p e ; the localities i n the Netherlands are
C o c c i a i n M i c o l . ital. 19 (1): p i . 55. 1990; D . R e i d i n F u n g . rar. Ic. c o l . probablv^meTmost northern i n m a i n l a n d Europe.
1: p i . 6e. 1966; W i n t e r h o f f & B o n i n C a r o l i n e a 52: p i . l a . 1994.
S E L . D E S C R . & FIGS. - Huijser & K e l d e r m a n i n C o o l i a 34: 80-83, R e i d (in F u n g . rar. Ic. c o l . 1: 2 4 . 1 9 6 6 ) stated that the spores o f Lepiota
fig. 1. 1991; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 134-135. 1994; D . hymenoderma are dextrinoid. T h i s is not confirmed by observations o n
R e i d i n F u n g . rar. Ic. c o l . 1: 24-25, figs. 6a-c. 1966; Winterhoff & B o n the Netherlands' material. H e r e the spores do c o l o u r slightly i n
in C a r o l i n e a 52: 6-7, fig. 1. 1994. M e l z e r ' s Reagent, but o n l y some spores, and the reaction is difficult to
V E R N . N A M E - R i n g l o z e stinkparasolzwam. observe. Candusso & L a n z o n i ' s ( L e p i o t a : 306. 1990) observations
agree w i t h the present description i n this respect, as these authors
called the spores slightly dextrinoid.
Lepiota hymenoderma can be distinguished from s m a l l specimens
of L. cristata i n the field b y the presence o f some squamules i n the
lower part o f the stipe and the absence o f an annulus.
32. Lepiota lilacea Bres., F u n g i trident. 2: 3. 1892. - F i g . 108.

Lepiotula lilacea (Bres.) Wasser i n N o v . Sist. vyssh. n i z s h . Rast.
1975: 191. ( ' 1 9 7 5 ' ) 1976. - Lepiota lilacea var. pallida M . B o n et al.
in D o c . m y c o l . 19 (76): 7 1 . 1989. - Lepiota lilacea f. micropholoides
M i g l . & C o c c i a i n M i c o l . ital. 21 (2): 40. 1992.
S E L . I C O N . - A n g e l i n Bolets Catalunya 14: p i . 681. 1995; Candusso
& L a n z o n i , L e p i o t a : p i . 36. 1990; C h r i s p i j n , C h a m p . Jordaan: 6 9 .
F i g . 107. Lepiota hymenoderma. 1999; R a i d & H e i l m a n n - C l a u s e n i n Svampe 28: 48. 1993.
Netherlands rare, k n o w n from several localities i n the western part o f

the country (province o f N o o r d - H o l l a n d ) and i n southern L i m b u r g ;
Sept.-Oct. N o t a c o m m o n species i n E u r o p e , but recorded f r o m
D e n m a r k i n the north to Italy i n the south; also i n N o r t h A m e r i c a .
The above macroscopical description is based o n notes accompanying

collections and the selected descriptions as cited above.
Lepiota lilacea is a variable species, concerning size and colours o f
the basidiocarps. It is characterized b y its annulus, and m i c r o s c o p i c a l
characters (spores, cystidia and pileus covering). D a r k variants resem-
ble L. felina i n structure o f the annulus and the discrete central part and
squamules o f the pileus covering, but are easily distinguished o n
account o f the structure o f the pileus covering and spore size.
33. Lepiota ochraceofulva P . D . O r t o n i n Trans. B r . m y c o l . Soc. 4 3 :

284. 1960. - F i g . 109.
Lepiota cookei H o r a i n Trans. B r . m y c o l . S o c . 4 3 : 446. 1960. -
Lepiota ochraceofulva var. huijsmanii M . B o n i n D o c . m y c o l . 22 (88):
30. 1993.
S E L . I C O N . - Enderle & Krieglst. i n Z . M y k o l . 55: opp. p. 96. 1989;
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 309-310, fig. R a i d & H e i l m a n n - C l a u s e n i n Svampe 2 8 : f i g . 1. 1993; D . R e i d i n
63. 1990; K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 214-215. 1936; F u n g . rar. Ic. c o l . 1: p i . 6b. 1966; R o m a g n . i n B u l l , trimest. S o c . m y c o l .
R a i d & H e i l m a n n - C l a u s e n i n Svampe 28: 50-52, fig. 6. 1993. Fr. 83: A t l a s p i . 172. 1967.
V E R N . N A M E - L i l a parasolzwam. S E L . D E S C R . & FIGS. - H u i j s m a n i n Persoonia 2: 364, figs. 10-15.
1962; P . D . Orton i n Trans. B r . m y c o l . S o c . 4 3 : 284-285. 1960; R a i d &
Pileus 12-35 m m , w h e n y o u n g campanulate-hemispherical, convex to H e i l m a n n - C l a u s e n i n Svampe 2 8 : 49-50, f i g . 4. 1993; R o m a g n . i n
applanate w i t h umbo, w i t h glabrous and closed umbo, surface around B u l l , trimest. Soc. m y c o l . Fr. 83: A t l a s p i . 172. 1967; Stridvall & A .
u m b o splitting up into squarrose, concentrical squamules w i t h uplift- Stridv. i n Goteborgs S v a m p e k l . Arsskr. 1981: 124-127. 1981.
ed tips, i n s m a l l specimens w i t h more adnate patches, very variable i n V E R N . N A M E - Okerbruine parasolzwam.
colour, f r o m very pale p i n k - b r o w n , v i a p i n k - b r o w n , b r o w n to dark

b r o w n ( K . & W . 8 F 6 at centre, 7 D 6 at squamules, 7 F 7 , M u . 7.5 Y R
3/4, 7.5-10 Y R 5/4), w h e n y o u n g i n most cases w i t h distinctly l i l a -
cinous sheen, o n an almost white background i n pale specimens, to
rose-tinged radially fibrillose background i n m e d i u m - c o l o u r e d speci-
mens, w i t h velar fringes at margin. L a m e l l a e , L = c. 3 5 , 1 = 0 - l ( - 3 ) ,
moderately crowded, free, ventricose, u p to 4 m m wide, pale cream,
w i t h p i n k i s h reflex, w i t h white fine-flocculose edge. Stipe 16-55 X
1.5-3 m m , c y l i n d r i c a l or broadening at apex, fistulose, at apex white to
isabella-buff, l o w e r d o w n more p i n k i s h to p i n k o r l i l a c i n o u s , shiny
lengthwise fibrillose. A n n u l u s l i k e a cuff around stipe, w i t h underside
squamulose and concolorous w i t h pileus centre, w i t h reflexed m a r g i n
and upper side cream-coloured. Context white-cream i n pileus, cream
i n stipe w i t h p i n k i s h streaks. S m e l l o f cut specimens sweetish, like the
sweet component o f the s m e l l o f L. cristata, aromatic. Taste not
k n o w n . Spore print c o l o u r not recorded.
Spores 4.0-6.0(-7.0) X 2.5-4.0 p m , Q = 1.3-1.7(-1.8), Q a v = 1.4-
1.55, ellipsoid, often uniguttulate, not distinctly thick-walled, hardly
congophilous, non-dextrinoid, metachromatic i n C r e s y l B l u e , w i t h con-
spicuous hilar appendage i n spores o f 1- and 2-spored basidia. B a s i d i a
18-26 X 5.5-7.5 p m , 4-spored; sometimes 1- and 2-spored basidia pre-
dominant. L a m e l l a edge sterile. Cheilocystidia 17-41 X 7.0-14 pm, nar-
rowly clavate, some clavate, a bit t h i c k - w a l l e d and colourless.
Pleurocystidia absent. Pileus covering a hymeniderm, made up o f (15-
)25-72(-85) X 8.5-21(-25) p m , cylindrical to narrowly clavate, rarely
clavate to globose elements; pigment only present i n some elements, and
there intracellular and brown, i n the very dark basidiocarps encrusting
the lower parts o f the terminal elements, i n paler specimens only encrust-
ing the hyphal walls i n the underlying layer. Stipitipellis a cutis, made up
of cylindrical elements, 2.0-4.0 p m i n diameter, w i t h finely encrusting
colourless pigment. Clamp-connections abundant i n a l l tissues.
terrestrial o n nutrient-rich soils, often i n p a r k - l i k e situations, i n the F i g . 109. Lepiota ochraceofulva.
Pileus 28-70(-100) m m , hemispherical w i t h inflexed margin, expand- Lepiota ochraceofulva is a t o x i c species as it contains amanitins
ing to plano-convex or applanate, w i t h or without umbo, w i t h smooth (Gerault & G i r r e i n C . r. hebd. Seanc. A c a d . S c i . , Paris, Ser. D , 280:
u m b o o r centre a n d s m a l l , broad, adnate squamules i n concentrical 2842. 1966).
zones; centre a n d squamules orange-brown ( M u . 7.5(-5) Y R 4/6) or
paler leather-brown, o n a w h i t i s h to cream or pale y e l l o w - b r o w n to 34. Lepiota pyrochroa M a l e n c . i n M a i . & Bert., F l . C h a m p , sup.
pale orange radially fibrillose background (up to 7.5-10 Y R 7/6), when M a r o c 1: 137. 1 9 7 0 . - F i g . 110.
y o u n g w i t h margin inflexed and exceeding lamellae. L a m e l l a e , L = c. S E L . I C O N . - B i z i o et a l . i n Riv. M i c o l . 36: 235. 1993; M a i . & Bert.,
50-100,1 = 3, crowded, free, separated from the stipe by a k i n d o f c o l - F l . C h a m p , sup. M a r o c 1: p i . 5. 1970.
l a r i u m , segmentiform up to 10 m m broad, cream-coloured, pale y e l l o w S E L . D E S C R . & FIGS. - B i z i o et a l . i n R i v . M i c o l . 36: 235-237. 1993;
(10 Y R 8/4) to pale orange-brown w i t h age (10 Y R 8-7/6), w i t h white, B o n & G . Riousset i n D o c . m y c o l . 2 2 (85): 69. 1992; M a i . & B e r t , F l .
even to flocculose edge. Stipe 50-70(-100) X 5-10 m m , enlarged at C h a m p , sup. M a r o c 1: 136-137, fig. 2 1 . 1970; G . Riousset & l o s s , i n
base up to 18 m m wide, c y l i n d r i c a l , h o l l o w , whitish or pale y e l l o w - B u l l . mens. Soc. l i n n . L y o n 4 5 : 198-200, fig. 1. 1976.
brown, discolouring y e l l o w i s h , orange or orange-brown when handled, V E R N . N A M E - Oranje p a r a s o l z w a m .
w i t h bands or girdles o f orange-brown squamules o n lower part o f stipe
or o n bulbous base only, w i t h vague annular zone, when y o u n g w i t h Pileus 9-17 m m , plano-convex, applanate and undulating around cen-
distinct annulus made o f partial v e i l , and w i t h white m y c e l i a l cords at tre, w i t h l o w umbo, orange ( M u . 5 Y R 5/6 at centre, 5 Y R 6/6 around
base. Context rather thick, w h i t i s h and d u l l i n pileus, whitish and shiny centre), w h e n o l d b r o w n i s h orange (7.5 Y R 6/6 at centre, 7.5 Y R 7/6
i n stipe. S m e l l w h e n basidiocarps are cut unpleasant, vaguely sugges- around centre), rugulose especially w h e n o l d , and especially around
tive o f L. cristata, sweetish-soapy o r perfumy. Taste m i l d , pleasant. centre, micaceous (lens). L a m e l l a e , L = 2 0 , 1 = 1-3, moderately
Spore print probably white. crowded, free, not far remote f r o m stipe, ventricose, up to 2.5 m m
Spores i n s i d e - v i e w 6.0-8.5(-10.0) X (3.5-)4.0-5.0 p m , Q = 1.5- wide, pale pinkish-orangelike (10 Y R 8/6), contrasting w i t h pileus and
1.9(-2.05), Q a v = 1.65-1.8, o b l o n g to c y l i n d r i c a l , some s l i g h t l y stipe colours, when o l d more brownish, w i t h even, concolorous edge,
o v o i d ; i n frontal v i e w slightly o v o i d , dextrinoid, c o n g o p h i l o u s , not w i t h age orange-spotted. Stipe 15-20 X 0.7-1,5 m m , c y l i n d r i c a l or
or s l i g h t l y metachromatic i n C r e s y l B l u e . B a s i d i a 23-31 X 6.0-8.5 slightly w i d e n e d at apex, orange-red (2.5 R 4-5/6), shiny w i t h some
pm, 4-spored, rarely 1- o r 2-spored. Lamella edge sterile. longitudinal innate fibrils i n lower part, w i t h narrow, white m y c e l i a l
Cheilocystidia 15-36(-51) X 4.0-8.5 pm, cylindrical, narrowly cords at base. Context orange a n d thin. S m e l l indistinct. Taste not
clavate, n a r r o w l y u t r i f o r m , sometimes w i t h very l o n g p e d i c e l , i n k n o w n . Spore print probably white.
some basidiocarps for the m a i n part slender: c y l i n d r i c a l to n a r r o w l y Spores 3.5-4.0(-4.5) X 2.5-3.0 p m , Q = 1.25-1.45, Q a v = 1.35,
clavate, h y a l i n e a n d s l i g h t l y t h i c k - w a l l e d . P l e u r o c y s t i d i a absent. ellipsoid i n side-view and i n frontal view, non-dextrinoid, not or hardly
P i l e u s c o v e r i n g a h y m e n i d e r m , made u p o f clavate, n a r r o w l y clavate congophilous, metachromatic i n C r e s y l B l u e , binucleate, seemingly
or i r r e g u l a r l y u t r i f o r m elements, 2 3 - 5 0 X 8.0-11 p m , w | t h b r o w n rough i n M e l z e r ' s Reagent, but smooth, w h e n observed w i t h Scanning
parietal pigment. Lower cylindrical elements sometimes with E l e c t r o n M i c r o s c o p e . B a s i d i a 14-15 X 5.0-5.5 p m , 4-spored. L a m e l l a
encrusting, b r o w n pigment. S t i p i t i p e l l i s a cutis, made up o f c y l i n d r i - edge fertile. C h e i l o c y s t i d i a not observed (but see comments).
cal, colourless, 3.0-5.0 p m w i d e hyphae. C l a m p - c o n n e c t i o n s present Pleurocystidia not observed. Pileus covering a h y m e n i d e r m , made up
i n a l l tissues. o f clavate to sphero-pedunculate elements, 22-50 X 14-28 um, w i t h
H A B I T A T & DISTR. - Solitary, i n s m a l l groups to gregarious, sapro- thickened, colourless w a l l s ; some elements w i t h intracellular b r o w n
trophic and terrestrial o n humus-rich and nutrient-rich l o a m y soil, o n pigment. Stipitipellis a cutis o f c y l i n d r i c a l , colourless hyphae, 3.0-6.0
the edge o f deciduous woods o r i n Fagus-woods, outside the p m i n diameter. Clamp-connections present i n a l l tissues.
Netherlands rather frequently encountered i n Juniperus-vegetation. In H A B I T A T & DISTR. - Solitary or i n s m a l l groups, saprotrophic a n d
the Netherlands k n o w n f r o m t w o localities i n southern Limburg terrestrial o n humus-rich and nutrient-rich soil, i n deciduous w o o d o n
(Elsloo, Bunderbos; Stokhem), f r o m the province o f F l e v o l a n d calcareous bedrock. I n the Netherlands k n o w n from one l o c a l i t y i n
(Noordoostpolder, U r k e r b o s ) , a n d f r o m R i j s w i j k (province o f Z u i d - southern L i m b u r g ( E l s l o o , Bunderbos). Sept. O r i g i n a l l y described
H o l l a n d ) , Sept.-Oct. Rare, but widespread i n Europe, k n o w n from from M o r o c c o a n d also recorded f r o m Cedrus-woods i n southern
S w e d e n southwards. France, Sardinia, and from deciduous forests i n Germany.
B o n ( F l . m y c o l . Eur. 3, Lepiotes: 77-78.1993) placed Lepiota ochrace- C h e i l o c y s t i d i a were absent i n the D u t c h material o f Lepiota pyrochroa,
ofulva i n subsect. Paralilaceae o f sect. Felinae, a subsection w h i c h is though they have been reported for the M o r o c c a n and F r e n c h c o l l e c -
characterized b y a dominant h y m e n i f o r m underlayer and rare or bare- tions (see selected descriptions). T h e spores i n the D u t c h c o l l e c t i o n
l y emerging l o n g elements. However, this type o f element is absent i n were considerably smaller than as given by M a l e n c o n (in M a l e n c o n &
L. ochraceofulva. Bertault, F l . C h a m p , sup. M a r o c 1: 136-137. 1970).
H o r a (in Trans. B r . m y c o l . S o c . 4 3 : 446. 1960) described L. cookei, Singer (Agar. m o d . Taxon., E d . 4: 494. 1986) mentions this species
w h i c h is identical to L. ochraceofulva, described by Orton i n the same (erroneously c a l l e d L. polychroa) as possibly belonging to the genus
issue o f the same j o u r n a l . B o n ( F l . m y c o l . Eur. 3, L e p i o t e s : 77-78.
1993) observed emerging elements i n the pileus covering o f L. cookei,
and kept the species separate.
Lepiota ochraceofulva var. huijsmanii M . B o n was distinguished
because o f a fishy s m e l l a n d the i n c o n s p i c u o u s c h e i l o c y s t i d i a . It
appears that c h e i l o c y s t i d i a are always abundant and quite obvious; a
different s m e l l i s not considered to warrant distinction o f a variety
(see H u i j s m a n i n Persoonia 2: 364. 1962; B o n i n D o c . m y c o l . 2 2 (88):
30. 1993; a n d V e l l i n g a & H u i j s e r i n B e l g . J . B o t . 131: 202-203.
( ' 1 9 9 8 ' ) 1999). F i g . 110. Lepiota pyrochroa.
Smithiomyces, a genus characterized b y a v e i l made u p o f very thin

hyphae w i t h embedded globose elements. H o w e v e r , a l l characters
point to a close relationships w i t h the species around L. lilacea.
35. Lepiota psalion Huijser & V e l l i n g a i n V e l l i n g a & Huijser i n B e l g .

J. B o t . 131: 203. ( ' 1 9 9 8 ' ) 1999.
M I S A P P L . - Lepiota rufipes sensu Krieglst. i n Beitr. K e n n t n . P i l z e
Mitteleur. 7: 69-71. 1991.
S E L . I C O N . - A . H a u s k n . i n K r i e g l s t . i n Beitr. K e n n t n . P i l z e
Mitteleur. 7: opp. p. 64. 1991 (as L. rufipes).
S E L . D E S C R . & FIGS. - K r i e g l s t . i n Beitr. K e n n t n . P i l z e Mitteleur. 7: F i g . 111. Lepiota rufipes.
69-70, fig. 2. 1991 (as L. rufipes); V e l l i n g a & Huijser i n B e l g . J . B o t .
131: 203-204, fig. 4. ( ' 1 9 9 8 ' ) 1999; W i n t e r h o f f & B o n i n C a r o l i n e a times with p i n k i s h tinge, especially at centre, ( M u . 10-7.5 Y R 7/6, 7.5
52: 8. 1994 (as L. rufipes f. phaeophylla). Y R 7-8/4 at centre, 10-7.5 Y R 8/4, 8/2 around umbo), slightly m i c a -
ceous (lens) or d u l l and glabrous, very slightly rugulose around umbo,
CHARACTERISTICS - P i l e u s 7-22 m m at first, later expanding to sometimes w i t h some white, arachnoid v e l u m fibrils at margin.
applanate c o n i c a l , w i t h inconspicuous u m b o , w i t h i n f l e x e d m a r g i n L a m e l l a e , L = c. 25-40, 1 = 1-5, moderately c r o w d e d to crowded,
w h e n y o u n g w i t h some adhering velar remnants, w h e n y o u n g at cen- slightly ventricose up to 4 m m w i d e , cream o r creamy (2.5 Y - 1 0 Y R
tre p i n k i s h to pale b r o w n , sometimes almost white, paler towards mar- 8/4), darkening w i t h age, w i t h white eroded or fimbriate edge. Stipe
gin, cream to pale flesh-coloured, smooth at centre, w i t h age breaking 13-32 X 1-2 m m , c y l i n d r i c a l , narrowly fistulose o r solid, pale cream
open around centre into patches; lamellae free, crowded, relatively i n upper part, lower d o w n w i t h w h i t i s h b l o o m o n vinaceous red back-
broad, w h e n y o u n g creamy white, cream flesh-coloured, soon darker, ground, often w i t h white appressed fibrils i n lower part, mostly w i t h -
w i t h w h i t i s h , slightly eroded edge; stipe 12-22(-37) X 1-3 m m , at apex out any trace o f annulus, sometimes w i t h adnate w h i t i s h remnants.
whitish, cream-white to slightly incarnate, towards base p i n k i s h , pink- Context pale o r sordid cream and dull i n pileus, shiny i n stipe and espe-
b r o w n to vinaceous b r o w n , almost smooth, s l i g h t l y fibrillose, often c i a l l y i n lower part reddish b r o w n (7.5 Y R 6/4-6). S m e l l indistinct or
w i t h white, tomentose, ascending, often incomplete annulus, white- w i t h faint smell l i k e L. cristata. Taste not k n o w n . Spore print pale y e l -
tomentose at base, often w i t h white m y c e l i a l cords; context w h i t i s h i n l o w i s h cream (c. 10 Y R 8/4).
pileus, i n lower part o f stipe p i n k b r o w n i s h to vinaceous b r o w n ; smell Spores 3.0-4.5 X 2.5-3.0(-3.5) p m , Q = 1.2-1.5(-1.6), Q a v = 1.25-
indistinct, like L. cristata. 1.45, broadly e l l i p s o i d to ellipsoid, uniguttulate, thin-walled, non-dex-
Spores 3.0-4.5 X 2.5-3.5 p m , Q = 1.2-1.6, Q a v = 1.25-1.45, broad- trinoid, seemingly r o u g h i n M e l z e r ' s Reagent, but smooth w h e n
ly e l l i p s o i d , e l l i p s o i d , non-dextrinoid; basidia 15-22 X 4.0-6.0 um, 4 - observed w i t h Scanning E l e c t r o n M i c r o s c o p e , very pale p i n k i n C o n g o
spored, some 2-spored; c h e i l o c y s t i d i a (13-)20-40 X 5.5-10.5 p m , Red, metachromatic i n C r e s y l B l u e . B a s i d i a 11-22 X 4.0-6.5 p m , 4 -
narrowly clavate, or clavate; pileus c o v e r i n g a h y m e n i d e r m w i t h tran- spored, rarely 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 14-34 X
sitions to an epithelium, w i t h up to 3(-4) elements o n top o f each other; 3.5-11 um, c y l i n d r i c a l and flexuous, often o n top o f each other, nar-
terminal element 18-55(-72) X 4-16(-20) u m , narrowly clavate, r o w l y clavate, relatively thin-walled, colourless. P l e u r o c y s t i d i a absent.
clavate, or o b p y r i f o r m ; clamp-connections present i n a l l tissues. Pileus covering a h y m e n i d e r m , made up o f tightly packed, narrowly
H A B I T A T & DISTR. - Gregarious and terrestrial i n h u m u s - r i c h decid- clavate to broadly clavate elements, 15-48 X 8-24 p m ; pigment brown,
uous woods, June-Oct. N o t with certainty recorded f r o m the parietal i n underlying hyphae. Stipitipellis a cutis o f c y l i n d r i c a l , 2.0-
Netherlands, k n o w n from A u s t r i a , Germany, and Hungary. 4.0 p m w i d e hyphae, w i t h b r o w n w a l l s (especially i n l o w e r part o f
stipe), w i t h loose l y i n g colourless, c y l i n d r i c a l velar hyphae. C l a m p -
T h i s taxon has been confused w i t h L. rufipes, but differs i n the pres- connections present i n a l l tissues.
ence o f an annulus, the fact that the pileus covering splits open i n L. H A B I T A T & DISTR. - Solitary or i n small groups, saprotrophic and
psalion, the u n i f o r m l y n a r r o w l y clavate c h e i l o c y s t i d i a and the b i n u - terrestrial on nutrient-rich, l o a m y o r sandy, humus-rich soils, i n decid-
cleate spores. uous woods and city-parks. In the Netherlands rare, occurring i n south-
ern L i m b u r g , i n the dune area and very rarely i n c l a y e y areas i n the
36. Lepiota rufipes M o r g . i n J . M y c o l . 12: 156. 1906. - F i g . 111. western part o f the country, l u l y - O c t . Rare i n Europe, not recorded i n
Lepiota rufipes f. phaeophylla M . B o n i n B o n & G . Riousset i n the northern countries, probably reaching the northernmost l i m i t o f its
D o c . m y c o l . 2 2 (85): 69. 1992. distribution area i n continental Europe i n the Netherlands. A l s o k n o w n
E X C L . - Lepiota rufipes sensu K r i e g l s t . i n Beitr. K e n n t n . P i l z e from N o r t h A m e r i c a .
Mitteleur. 7: 69-71. 1991 ( = L. psalion). - Lepiota rufipes f. phaeo-
phylla sensu B i z i o et a l . i n Riv. M i c o l . 36: 240-241. 1993 ( = L. crista- T h e concept o f Lepiota rufipes, as described above, is mostly based o n
toides); Lepiota rufipes f. phaeophylla sensu W i n t e r h o f f & B o n i n the interpretation o f K i i h n e r & M a i r e (in B u l l . S o c . H i s t . nat. A f r . N .
C a r o l i n e a 52: 8. 1994 ( = L. psalion). 28: 108-109. 1937) o f L. rufipes M o r g . M o r g a n ' s description ( i n J .
S E L . I C O N . - B i z i o et a l . i n Riv. M i c o l . 36: 239. 1993; M y c o l . 12: 156. 1906) is rather vague, characterizing a s m a l l pale
S E L . D E S C R . & FIGS. - B i z i o et a l . i n R i v . M i c o l . 36: 239-240. 1993; mushroom, without squamules, w i t h a reddening stipe and an evanes-
Joss, i n B u l l , trimest. S o c . m y c o l . F r . 7 1 : 65-67, f i g . 1. 1955; cent ring. Lepiota rufipes i n the present sense lacks a r i n g , although
K e l d e r m a n i n C o o l i a 35: 76-79, fig. 1. 1992. some velar remnants m a y be present.
V E R N . N A M E - K a l e parasolzwam. Lepiota rufipes f. phaeophylla M . B o n was described as a taxon
w i t h rather dark lamellae, a n d variable c h e i l o c y s t i d i a . I n later inter-
Pileus 7.5-18 m m , w h e n y o u n g c o n i c o - h e m i s p h e r i c a l w i t h inflexed pretations o f this forma (e.g. B i z i o et al. i n R i v . M i c o l . 36: 240-241.
margin, expanding to plano-convex o r applanate w i t h , rarely without, 1993) the cheilocystidia are said to be rare to almost absent, and a r i n g
u m b o , w h e n y o u n g w h i t i s h cream, later more ochre coloured, some- to be present. T h e original description and material clearly depict L.
rufipes s.str., whereas the later interpretations agree either w i t h Lepiota S E L . D E S C R . & F I G S . - Candusso & L a n z o n i , L e p i o t a : 127-128, fig.
cristatoides o r w i t h L. psalion. 17. 1990; H o r a k i n S y d o w i a 33: 118-119. ( ' 1 9 8 0 ' ) 1981; K e l d e r m a n ,
Several authors described L. rufipes as h a v i n g a distinct annulus. Parasolzw. Z u i d - L i m b u r g : 44-45. 1994; K n u d s e n i n B o t . Tidsskr. 7 5 :
T h i s taxon differs f r o m L. rufipes s.str. not o n l y i n the presence o f this 146-147, figs. 1 L , 2 A , 31 & 20. 1980; K u h n e r i n B u l l , trimest. S o c .
annulus, but also i n the looser structure o f the pileus covering, the pres- m y c o l . Fr. 52: 210-211. 1936 (as L. acutesquamosa var. furcata).
ence o f two n u c l e i i n the spores (versus one i n L. rufipes s.str.) and the V E R N . N A M E - Spitsschubbige parasolzwam.
absence o f o i l - l i k e substances i n the m i c r o s c o p i c a l structures; further-
more the c h e i l o c y s t i d i a are less variable i n shape than i n L. rufipes. P i l e u s 50-115(-150) m m , h e m i s p h e r i c a l w i t h i n f l e x e d m a r g i n w h e n
T h i s taxon has been described as L. psalion Huijser & V e l l i n g a ( i n y o u n g , expanding v i a b r o a d l y b l u n t l y c o n i c a l to applanate o r slightly
V e l l i n g a & Huijser i n B e l g . J . B o t . 131: 203. ('1998') 1999). plano-concave w i t h broad l o w umbo, w h e n y o u n g completely covered
Differences between L. rufipes and L. cristatoides are the presence w i t h b i g p y r a m i d a l warts o n a c l o s e d b a c k g r o u n d , later at centre
i n the former o f cheilocystidia, the rounder spores, the absence o f an closely set w i t h warts, towards m a r g i n w i t h more repent spines w i t h
annulus, and the non-squamulose pileus. recurved tips, w i t h r a d i a l fibrils around those spines o r their place o f
Lepiota neophana var. europaea B i z i o & M i g l . and its f. papillata attachment, easily removable, often, but not always, s h o w i n g the
M i g l . & Perrone differ f r o m L. rufipes i n the absence o f cheilocystidia, underlying b a c k g r o u n d i n between; centre and spines b r o w n , p i n k i s h
the scarcity o f clamp-connections, and the longer spores ( B i z i o et a l . orange-brown, orange-brown, rarely p i n k i s h y e l l o w i s h b r o w n ( M u .
i n R i v . M i c o l . 36: 228-234. 1993; see also V e l l i n g a & Huijser i n B e l g . 7.5 Y R 4-5/4-6, 7/6; 5 Y R 5/6, 6/6), often spines darker than back-
J . B o t . 131: 205. ( ' 1 9 9 8 ' ) 1999). ground and w i t h dark tips, very rarely colourless (see notes); w h e n
Lepiota rufipes might be confused w i t h specimens o f Cystolepiota y o u n g w i t h white c o b w e b b y velar remnants at margin. L a m e l l a e , L =
seminuda after heavy rain. T h e presence o f c h e i l o c y s t i d i a a n d the c. 200, number o f l a m e l l u l a e difficult to assess, very crowded, free,
structure o f the pileus covering are the best m i c r o s c o p i c a l characters often furcate either near stipe o r i n outer part, segmentiform or slight-
to distinguish L. rufipes from C. seminuda. l y ventricose, up to 8.5 m m w i d e , usually rather narrow, creamy white,
w i t h age w i t h orange spots, w i t h white, flocculose edge. Stipe 50-120
!
X 9-22 m m , c y l i n d r i c a l often w i t h bulbous o r turnip-shaped base, h o l -
s'low, annulate, above annulus cream-coloured, lengthwise fibrillose,
Sect. Echinatae Fay. b e l o w annulus b r o w n i s h , o r p i n k i s h pale b r o w n (c. 7.5 Y R 7/4) fibril-
lose o n white background, w i t h i n l o w e r 1/3 to 1/4 girdles o f b r o w n
Pileus covered w i t h acute squamules, made u p o f globose to e l l i p s o i d floccose to acute, dark-tipped squamules as o n pileus, w i t h white
(oblong) elements i n agglutinated chains, transient to l o n g coloured m y c e l i u m cords at base. A n n u l u s pendant, membranaceous, often dis-
hyphae o n pileus surface. Spores dextrinoid, rarely not reacting w i t h appearing as it is not p r o p e r l y attached to stipe, w i t h white felted
M e l z e r ' s Reagent; clamp-connections present, rarely absent. upper side confluent w i t h upper part o f stipe, a n d pale b r o w n i s h
underside beset w i t h dark b r o w n acute squamules, as o n pileus.
37.Lepiotaaspera(Pers.:Fr.)Quel.,Enchir.Fung.: 6 . 1 8 8 6 . - F i g . 112. Context thick, white and d u l l i n pileus, shiny and cream to pale y e l -
Agaricus asper Pers. i n H o f f m . , A b b . S c h w a m m e 3: p i . 2 1 . 1793; l o w i s h - p i n k i s h i n stipe. S m e l l w h e n cut and fresh very strong, l i k e the
Amanita aspera (Pers.) Pers., Observ. m y c o l . 2: 38. 1799; Agaricus rubber component o f the L. cristata-oAom, i n some cases also w i t h
asper (Pers.) Pers.: Fr., Syst. m y c o l . 1: 18. 1821; Cystolepiota aspera the sweet component o f that s m e l l . Taste as s m e l l , slightly bitterish,
(Pers.: Fr.) K n u d s e n i n B o t . Tidsskr. 73: 129. 1978; Echinoderma aspe- unpleasant. Spore print white.
rum (Pers.: Fr.) M . B o n i n D o c . m y c o l . 21 (82): 62. 1991. -Agaricus Spores 6.0-9X)(-9.5) X 2.5-3.5(-4.0) p m , Q = 1.9-3.3, Q a v = 2.2-
acutesquamosus W e i n m . i n Hornsch., S y l l . PI. nov. Ratisb. 1: 7 0 . 1 8 2 2 ; 2.7, i n side-view c y l i n d r i c a l , slightly triangular w i t h widest part near
Lepiota acutesquamosa (Weinm.) K u m m . , Fuhr. P i l z k . : 136. 1871; hilar appendage, some w i t h convex adaxial side, some w i t h slight hilar
Cystolepiota acutesquamosa (Weinm.) M . B o n i n D o c . m y c o l . 7 (27- depression i n side-view ('penguin-shaped'), i n frontal v i e w c y l i n d r i c a l
28): 11. 1977; Echinoderma acutesquamosum (Weinm.) M . B o n i n to slightly o v o i d , dextrinoid, red, but sometimes hardly, i n C o n g o R e d ,
D o c . m y c o l . 2 2 (88): 28. 1993; Lepiota friesii var. acutesquamosa not metachromatic i n C r e s y l B l u e . B a s i d i a (12-)15-26 X 5.5-9.0 p m ,
(Weinm.) Q u e l , i n M e m . S o c . E m u l . M o n t b e l i a r d , Ser. II, 5: 72. 1872 4-spored, very rarely 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a
( C h a m p . Jura Vosges 1); Lepiota aspera var. acutesquamosa (Weinm.) abundant, 15.5-45 X 7 . 0 ^ p m , clavate to spheropedunculate, rarely
S i n g , i n Persoonia 2: 9. 1961; Agaricus friesii L a s c h i n L i n n a e a 3: 155. narrowly clavate, slightly t h i c k - w a l l e d , hyaline, rarely y e l l o w i s h .
1828; Agaricus friesii L a s c h : Fr., Syst. m y c o l . , Ind. gen.: 2 1 . 1832; Pleurocystidia absent. Elenients o f pileus covering agglutinated into
Lepiota friesii (Lasch: Fr.) Q u e l , i n M e m . Soc. E m u l . M o n t b e l i a r d , Ser. warts; terminal elements o f chains 30-57 X 15-45 um, globose to e l l i p -
II, 5: 72. 1872 ( C h a m p . Jura Vosges 1); Cystolepiota friesii (Lasch: Fr.) soid or clavate, w i t h o r without b r o w n w a l l s . B a s a l elements c y l i n d r i -
M . B o n i n D o c . m y c o l . 7 (27-28): 12. 1977; Echinoderma friesii cal to slightly inflated, 2.0-12 u m wide, b r o w n - w a l l e d (not encrusted),
( L a s c h : Fr.) M . B o n i n D o c . m y c o l . 22 (88): 2 8 . 1993. - Lepiota w i t h clamp-connections. Stipitipellis a cutis o f c y l i n d r i c a l , 3.0-5.0 u m
acutesquamosa var. furcata K u h n e r i n B u l l , trimest. Soc. m y c o l . F r . w i d e , hyphae, colourless, b e l o w annulus also b r o w n hyphae present,
52: 210. 1936. - Lepiota acutesquamosa f. gigantea Pilat in Acta M u s . w i t h elements i n spines and warts as those o n pileus. C l a m p - c o n n e c -
natn. Prag. 11B (2): 3. 1955; Echinoderma acutesquamosum f. gigan- tions abundant i n a l l tissues.
teum (Pilat) M . B o n i n D o c . m y c o l . 2 2 (88): 28. 1993. H A B I T A T & DISTR. - Gregarious, rarely solitary, saprotrophic a n d
E X C L . - Lepiota acutesquamosa var. typica K u h n e r i n B u l l . ' trimest. terrestrial o n h u m u s - r i c h and/or nutrient-rich sandy to clayey soils,
Soc. m y c o l . F r . 5 2 : 209. 1936; Lepiota acutesquamosa sensu M o s . , often i n slightly disturbed places like gardens and around badger dens,
R o h r l i n g e Blatterpilze, 4. A u f l . : 238. 1978 (in both cases L. perplexa). i n deciduous, m i x e d , and coniferous woods, city-parks, and orchards.
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 214. 1995; A l s o recorded f r o m greenhouses. W i t h L. cristata and Cystolepiota
Candusso & L a n z o n i , L e p i o t a : p i . 7. 1990; Dahncke, 1200 P i l z e : 514, seminuda f o r m i n g the trio o f most c o m m o n Lepiota-species i n the
515. 1993; J . L a n g e , F l . agar. dan. 1: p i . 10F. 1935 (as L. Netherlands, occurring rather c o m m o n l y throughout the country. A u g . -
acutesquamosa); R . P h i l l i p s , Paddest. S c h i m m . : 27. 1981. N o v . Widespread i n E u r o p e and the rest o f the temperate parts o f the
N o r t h e r n H e m i s p h e r e a n d everywhere scattered to moderately c o m - 38. Lepiota perplexa K n u d s e n i n B o t . Tidsskr. 75: 137. 1980. - F i g .
mon, o n l y l a c k i n g north o f the arctic c i r c l e and i n alpine habitats. 113.
Cystolepiota perplexa (Knudsen) M . B o n i n D o c . m y c o l . 15 (60):
Lepiota aspera i s very variable i n terms o f pileus covering; the spines 38. 1985; Echinoderma perplexum (Knudsen) M . B o n i n D o c . m y c o l .
can be b i g and standing out o n a w h i t i s h background, but also s m a l l 21 (82): 63. 1991.
spines o n a b r o w n background do occur. T h e c o l o u r i n g is not constant M I S A P P L . - Lepiota acutesquamosa var. typica sensu K i i h n e r i n
either. O c c a s i o n a l l y totally white or sordid-white specimens are B u l l , trimest. S o c . m y c o l . Fr. 5 2 : 209. 1936; Lepiota acutesquamosa
encountered. A l s o w h i t i s h specimens w i t h pale b r o w n tips o f the squa- sensu M o s . , R o h r l i n g e Blatterpilze, 4. A u f l . : 238. 1978.
mules are met w i t h (see for example B a l k e i n C o o l i a 13: 11-12. 1966). S E L . D E S C R . & F I G S . - Candusso & L a n z o n i , L e p i o t a : 148-150, fig.
R a r e l y p i n k - b r o w n colours occur, reminiscent o f those o f L. fuscov- 22. 1990; E n d e r l e & K r i e g l s t . i n Z . M y k o l . 5 5 : 84-85. 1989;
inacea. I n a l l these cases, m i c r o s c o p i c a l characters, and the other K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 54-55. 1994; K n u d s e n i n B o t .
macroscopical features (annulus, size, number o f lamellae etc.) leave Tidsskr. 75: 137-139, figs. I I , 2 E , 3 L , 12 & 13. 1980.
no doubt that these variants still belong to L. aspera. VERN. NAME - Egelparasolzwam.

deciduous woods or city-parks o n humus-rich, nutrient-rich soils, rare,
m a i n l y i n the western c l a y region, the river area and i n southern
L i m b u r g . A u g . - O c t . Widespread but rare i n temperate parts o f Europe.
39. Lepiota hystrix M 0 l l e r & J . L a n g e i n J . L a n g e , F l . agar. dan. 5: V .

1940.
Lepiota hystrix M 0 l l e r & J . L a n g e i n J . L a n g e , F l . agar. dan. 1: 26.
1935 (not valid); Cystolepiota hystrix ( M 0 l l e r & J . L a n g e ) K n u d s e n i n
Bot. Tidsskr. 73: 128. 1978; Echinoderma hystrix ( M 0 l l e r & J . Lange)
M . B o n i n D o c . m y c o l . 21 (82): 63. 1991.
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 227. 1995; J .
Lange, F l . agar. dan. 1: p i . 10E. 1935; R . P h i l l i p s , Paddest. S c h i m m . :
26. 1981; S t r i d v a l l i n Jordstjarnan 12 (2): opp. p. 1. 1991.
S E L . D E S C R . & FIGS. - K n u d s e n i n B o t . Tidsskr. 75: 143-144, figs.
I K , 2F, 3F, 17 & 18. 1980.
C H A R A C T E R I S T I C S - Pileus 30-70 m m , c o n i c a l to convex w i t h a broad

umbo, tomentose, greyish orange or b r o w n , covered w i t h pointed, c o n -
i c a l , rather persistent squamules, b l a c k i s h and very c r o w d e d at centre,
more scattered towards m a r g i n and paler w i t h b l a c k i s h tip, towards
margin surrounded w i t h arachnoid fibrils showing the whitish context
F i g . 113. Lepiota perplexa.
in between; lamellae w h i t i s h w i t h dark l a m e l l a edge; stipe 40-80 X 4-
8 m m , c y l i n d r i c a l w i t h annulus, b e l o w annulus w i t h pointed squa-
Pileus 28-55 m m , w h e n y o u n g hemispherical w i t h inflexed margin, jgiules; annulus membranaceous, w i t h w o o l l y squamules o n underside
expanding to p i a n o - c o n i c a l , plano-convex and eventually applanate and when fresh e x u d i n g b l a c k i s h drops; smell unpleasant, reminiscent
w i t h low, broad u m b o , at centre densely set w i t h irregular to c o n i c a l o f the smell o f L. cristata.
warts up to 1 m m h i g h , around centre w i t h warts i n concentrical zones Spores 5.5-7.0 X 2.5-3.0 p m , Q = 2.0-2.6, Q a v = 2.3, c y l i n d r i c a l
or more scattered, w i t h radial, arachnoid fibrils around base, on tomen- or subcylindrical w i t h convex abaxial side i n side-view, i n frontal v i e w
tose-felted background, p i n k i s h b r o w n ( M u . 7.5 Y R 5/5, 5/6) at centre narrowly o v o i d ; l a m e l l a edge sterile, w i t h parallel hyphae w h i c h give
and p i n k i s h buff (7.5 Y R 7/6) at margin, sometimes darker ( K . & W . rise to clusters o f t h i c k - w a l l e d cheilocystidia; cheilocystidia 17-40 X
6 C 4 - 6 D 4 ) w i t h concolorous to slightly darker warts (e.g. 7.5 Y R 4/6); 6-10 p m , narrowly clavate, sometimes slightly capitate or c y l i n d r i c a l ,
warts easily rubbed and washed off; w h e n y o u n g w i t h remnants o f par- h y a l i n e at first, then w i t h b r o w n w a l l s , finally w i t h dark contents;
tial v e i l at margin, and margin exceeding lamellae. L a m e l l a e , L = 55- pileus covering w i t h warts, made up o f chains o f irregularly shaped and
70, 1 = 1-3, c r o w d e d , free and remote from stipe, not furcate, sized elements elements, up to 110 X 30 p m , w i t h intracellular dark
subventricose, ventricose, rather broad (up to 6 m m wide), sordid brown pigment w h i c h is soluble i n ammonia, and w i t h b r o w n w a l l s ;
cream, sometimes w i t h p i n k i s h sheen, w i t h white, subgranular edge. clamp-connections present.
Stipe 35-70 X 5-11 m m , c y l i n d r i c a l w i t h (sub)bulbous, up to 15 m m H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres-
wide, base, fistulose, at utmost apex w h i t i s h to cream, below annular trial i n deciduous woods o n nutrient-rich soils. N o t yet recorded i n the
zone darkening towards p i n k i s h b r o w n at base, i n lower 1/3 or less w i t h Netherlands and rare i n E u r o p e .
darker reddish b r o w n , slightly lanate, fibrils (7.5 Y R 6-5/4-6) length-
wise and at base w i t h distinct warts, concolorous or slightly darker than T h e above description is based o n descriptions i n the literature and o n
those o n pileus. Context w h i t i s h i n pileus, glassy cream or very pale observations o n two collections f r o m the same l o c a l i t y i n northern
b r o w n i n stipe and white around tube. S m e l l strong i n cut basidiocarps, France. Lepiota hystrix is easily recognized b y the c o m b i n a t i o n o f the
like the s m e l l o f L. cristata. Taste unpleasant, somewhat acrid-astrin- dark, almost b l a c k squamules o n the pileus, and the dark l a m e l l a edge.
gent. Spore print probably white. Lepiota calcicola o c c a s i o n a l l y has a dark l a m e l l a edge, but i n that
Spores 5.0-6.5 X 2.5-3.0 p m , Q = 1.8-2.6, Q a v = 2.05-2.35, i n species the elements o f the warts are regularly shaped, e l l i p s o i d to
side-view o b l o n g to c y l i n d r i c a l , some w i t h slightly convex, abaxial o b l o n g (not globose), and the pigment is situated i n the w a l l s o f the
side, some w i t h suprahilar depression ('penguin-shaped'), i n frontal elements. K n u d s e n (in B o t . Tidsskr. 7 5 : 144. 1980) stated that i n L.
view o b l o n g to c y l i n d r i c a l , dextrinoid, metachromatic i n C r e s y l B l u e , hystrix the elements o f the warts were t y p i c a l l y 20-30 u m and globose;
but not very distinctly so. B a s i d i a 12.5-23 X 4.0-7.0 p m , 4-spored, however, i n the specimens studied the elements were very variable,
very rarely 2-spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 24-40(-50) both i n shape and size. T h e shape was very irregular, and only a few
X 7.0-10(-13) p m , fusiform, narrowly clavate, c y l i n d r i c a l , often w i t h elements were globose.
constriction, slightly t h i c k - w a l l e d and colourless. P l e u r o c y s t i d i a
absent. Pileus covering made up o f c y l i n d r i c a l , b r o w n - w a l l e d , 4.0-16 40. Lepiota calcicola K n u d s e n i n B o t . Tidsskr. 75: 140. 1980. - F i g .
p m w i d e hyphae, g i v i n g rise to agglutinated chains o f more or less glo- 114.
bose elements, unified to warts; terminal elements o f warts 16-60 X Cystolepiota calcicola (Knudsen) B o n & Courtec. i n D o c . m y c o l .
12-57 p m , t h i c k - w a l l e d and brown. Stipitipellis at apex o f stipe a cutis 18 (69): 38. 1987; Echinoderma calcicola (Knudsen) M . B o n i n D o c .
of c y l i n d r i c a l , 3.0-6.0 p m w i d e , y e l l o w - b r o w n hyphae. Stipe covering m y c o l . 21 (82): 62. 1991.
made up o f irregularly flexuous, c y l i n d r i c a l hyphae, b r o w n and thick- M I S A P P L . - Lepiota hispida sensu J . L a n g e , F l . agar. dan. 1: 26.
walled, w i t h elements as i n squamules o n pileus. Clamp-connections 1935; Cystolepiota hispida sensu M . B o n i n D o c . m y c o l . 11 (43): 28.
present i n a l l tissues. 1981.
w i d e , when y o u n g white, later more cream to y e l l o w i s h , w i t h c o n c o l -

orous, white, or rarely dark, even, pruinose to irregular edge. Stipe 30-
90 X 4-14 m m , c y l i n d r i c a l , o r slightly attenuate at apex a n d slightly
widened towards subbulbous, 8-13 m m w i d e , base, fistulose w i t h age,
at apex white when y o u n g , brunnescent w i t h age, and fibrillose, below
annular zone completely covered w i t h dark b r o w n fibrils, w i t h zones
or girdles o f acute squamules as o n pileus, sometimes more w i d e -
spread, showing more o f the u n d e r l y i n g context, and m a k i n g a paler
impression than pileus, sometimes very dark b r o w n at base. Context
thick, d u l l and white i n pileus, w h i t i s h and shiny i n stipe. S m e l l l i k e
s o i l or cellars; when basidiocarp cut, s m e l l distinctly l i k e L. cristata
or fruity-resinaceous, and not reminiscent o f L. cristata. Taste m i l d ,
fungoid. Spore print ' w h i t e ' .
Spores 4.0-6.0 X 2.0-3.5 p m , Q = 1.4-2.2, Q a v = 1.6-1.85, o b l o n g
to subcylindrical to c y l i n d r i c a l , i n frontal v i e w sometimes slightly
o v o i d , dextrinoid, c o n g o p h i l o u s , not metachromatic i n C r e s y l B l u e .
Basidia 15.5-22 X 5.0-7.0 u m , 4-spored. L a m e l l a edge sterile.
Cheilocystidia i n fascicles, 18-35 X 4.0-10 p m , narrowly clavate, some
c y l i n d r i c a l , some narrowly utriform, hyaline, very rarely w i t h dark
brown contents. Pleurocystidia absent. Elements o f pileus covering i n
agglutinated chains; a p i c a l elements (20-)30-70(-100) X 15-40 u m ,
ellipsoid to oblong, a very few globose, w i t h thickened, b r o w n w a l l s ;
intermediate elements more elongate-fusoid; basal elements, 2.0-6.0
p m wide, c y l i n d r i c a l , w i t h b r o w n w a l l s . Stipitipellis a cutis o f c y l i n -
drical, b r o w n - w a l l e d , 3.0-5.0 p m w i d e , y e l l o w - b r o w n hyphae, w i t h
b e l o w annular zone, elements i n squamules, as o n pileus, o r longer.
Clamp-connections abundant i n a l l tissues.
H A B I T A T & DISTR. - I n s m a l l groups, gregarious, saprotrophic and
terrestrial o n humus-rich soils and o n calcareous bedrock, i n decidu-
ous woods. Very rare i n the Netherlands, o n l y k n o w n f r o m several
places i n southern L i m b u r g ; A u g . - N o v . Widespread and rare i n Europe.
41. Lepiota efibulis K n u d s e n i n B o t . Tidsskr. 7 5 : 151. 1980.

Echinoderma efibule (Knudsen) M . B o n i n D o c . m y c o l . 21 (82):
62. 1991.
S E L . D E S C R . & F I G S . - K n u d s e n i n B o t . T i d s s k r . 7 5 : 151. 1980.
VERN. NAME - Kasstekelparasolzwam.
F i g . 114. Lepiota calcicola. C H A R A C T E R I S T I C S - Pileus 20-55 m m , w h e n y o u n g truncate-conical,

expanding to plano-convex w i t h broad, prominent umbo, w h e n young
completely covered w i t h adjacent, acute warts, later at u m b o covered
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 217. 1995; with small up to 1 m m h i g h warts, and around u m b o w i t h concentrical
H e i l m a n n - C l a u s e n i n S v a m p e 34: 3 3 . 1996; K a s p a r e k i n M i t t b l . uplifted acute squamules, dark red-brown o n u m b o and warts ( M u . 7.5
A r b e i t s g e m . P i l z k . N i e d e r r h e i n 11: after p. 102. 1993. Y R 3/4) o n paler background (7.5 Y R 5/5 to 10 Y R 8/6 at margin);
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 132-134, fig. cream context not or hardly showing at umbo; lamellae moderately dis-
18. 1990; Kasparek i n M i t t b l . A r b e i t s g e m . P i l z k . Niederrhein 11: 7 8 - tant, free, w i t h up to 1 l a m e l l u l a , ventricose, not furcate, creamy floc-
83. 1993; K e l d e r m a n , Parasolzw. Zuid-Limburg: 46-47. 1994; culose edge; stipe 35-55 X 3-5 m m , c y l i n d r i c a l , fistulose w i t h age,
K n u d s e n i n B o t . Tidsskr. 7 5 : 140, figs. 1 H , 2 G , 3 B , 15. 1980; w i t h white m y c e l i u m cords at base, w i t h distinct annulus 1/3 from
L o h m e y e r et a l . i n Z . M y k o l . 59: 205-206. 1993. apex, above annulus p i n k i s h (5 Y R 8/3), lengthwise fibrillose, below
V E R N . N A M E - Kalkstekelparasolzwam. annulus purplish red-brown (5 Y R 4/4, 7.5 Y R 6/4) w i t h i n lower 1/3
zones o f blunt floccose squamules; annulus w i t h red-brown squamu-
P i l e u s 26-85 m m , h e m i s p h e r i c a l , convex o r r o u n d e d - c o n i c a l w i t h lose warts o n upper part, less b i g and less acute than o n pileus; context
inflexed m a r g i n at first, expanding to applanate or plano-concave w i t h white and d u l l i n pileus, i n stipe concolorous w i t h upper outer part,
u m b o and undulating, deflexed m a r g i n , strongly fibrillose-squamose, shiny white around tube; smell unpleasant, somewhat suggestive o f the
at centre w i t h large, p y r a m i d a l , slender squamules up to 5 m m h i g h smell o f L. cristata; taste none.
and 1.5 m m at base, towards m a r g i n w i t h more appressed fibrillose Spores 4.5-5.5 X 2.5-3.0(-3.5) p m , Q = 1.55-1.95, Q a v = 1.7,
squamules; u n d e r l y i n g context hardly s h o w i n g ; rather equally b r o w n oblong, dextrinoid, metachromatic i n C r e s y l B l u e ; b a s i d i a 4-spored;
or dark b r o w n ( K . & W . 7 E 4 - 5 ; M u . 5 Y R 4/3-2), sometimes more lamella edge sterile; c h e i l o c y s t i d i a 18-32 X 8.0-12 p m , clavate; pileus
vinaceous b r o w n ( 8 D 3 - 4 ) ; squamules not r e m o v a b l e ; m a r g i n w i t h warts made up o f chains o f globose to o v o i d elements, 24-40 X 18-32
some overhanging v e l u m remnants; w h e n y o u n g w i t h a cortina o f pm, w i t h brown, thick w a l l s ; warts o n stipe made up o f more irregu-
white and b r o w n fibrils between stipe and pileus m a r g i n . L a m e l l a e , L larly shaped elements than o n pileus; clamp-connections absent i n a l l
= 70-80,1 = 3-6, c r o w d e d to very c r o w d e d , free, ventricose, 5-8 m m tissues.
H A B I T A T & D I S T R . - Gregarious, saprotrophic and terrestrial o n at base; at base w i t h w h i t e m y c e l i u m cords. C o n t e x t white, d u l l and
h u m u s - r i c h soil, very rare, o n l y k n o w n f r o m a non-heated greenhouse f i r m i n p i l e u s , i n stipe c r e a m to b r o w n i s h i n l o w e r part, shiny. S m e l l
in A m s t e r d a m . Sept. A l s o recorded f r o m several greenhouses i n Great sweetish fruity l i k e L. cristata, e s p e c i a l l y o f cut basidiocarps. Taste
Britain. l i k e taste o f L. cristata: rubber-like and unpleasant. Spore print prob-
ably white.
T h i s species i s aberrant w i t h respect to the other representatives o f the Spores ( 3 . 5 - ) 4 . 0 - 5 . 5 ( - 6 . 5 ) X ( 2 . 0 - ) 2 . 5 - 3 . 5 u m , Q = 1.4-2.0(-2.15),
section i n E u r o p e o n account o f the absence o f clamp-connections. Qav = 1 . 6 - 1 . 8 , oblong or s u b c y l i n d r i c a l i n side-view, similar or slight-
ly o v o i d i n frontal view, dextrinoid, congophilous, and not metachro-
42. Lepiota echinacea J . L a n g e , F l . agar. dan. 5: V . 1940. - F i g . 115. matic i n C r e s y l B l u e . B a s i d i a 1 5 - 2 6 X 4 . 5 - 7 . 0 u m , 4-spored, rarely
Lepiota echinacea J . L a n g e , F l . agar. dan. 1: 26. 1935 (not valid); 2-spored; o l d basidia i n some cases t h i c k - w a l l e d . L a m e l l a edge het-
Cystolepiota echinacea (J. L a n g e ) K n u d s e n i n B o t . Tidsskr. 7 3 : 127. erogeneous or sterile. C h e i l o c y s t i d i a absent or present i n tufts; 8 . 0 - 3 2
1978; Echinoderma echinaceum (J. Lange) M . B o n i n D o c . m y c o l . 21 X 4 - 1 1 . 5 p m , clavate, narrowly clavate and narrowly utriform, slight-
(82): 63. 1991. l y t h i c k - w a l l e d , colourless. P l e u r o c y s t i d i a absent. Pileus c o v e r i n g
S E L . I C O N . - Enderle & K r i e g l s t . i n Z . M y k o l . 55: opp. p. 64. 1989; made up o f agglutinated chains o f e l l i p s o i d to rounded elements; ter-
J. L a n g e , F l . agar. dan. 1: p i . 1 0 D . 1935; V e l l i n g a & Huijser i n C o o l i a m i n a l elements o f chains o f squamules 1 5 - 6 2 X 1 0 - 5 0 p m , e l l i p s o i d ,
40: p i . 2. 1997. globose, w i t h thick, b r o w n w a l l s . Stipitipellis a cutis made up o f c y l i n -
S E L . D E S C R . & FIGS. - K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 50-51. drical hyphae, 4 . 0 - 1 0 u m w i d e , w i t h pale b r o w n , sometimes encrust-
1994; K n u d s e n i n B o t . Tidsskr. 7 5 : 135, 137, figs. 1 G , 3 J , 9 & 10. ed, walls, w i t h squamules as o n pileus, elements o f these squamules
1980; K i i h n e r i n B u l l , trimest. S o c . m y c o l . Fr. 52: 211-212. 1936. often more c y l i n d r i c a l - e l l i p s o i d than o n pileus, w i t h b r o w n w a l l s .
V E R N . N A M E - Fijnschubbige parasolzwam. Clamp-connections present i n a l l tissues.
H A B I T A T & D I S T R . - Gregarious i n s m a l l groups, rarely i n fairy
P i l e u s 18-50(-60) m m , w h e n y o u n g b r o a d l y h e m i s p h e r i c a l or broad- rings, saprotrophic and terrestrial i n deciduous woods, city-parks, mine
ly conical-hemispherical with inflexed margin, expanding to waste heaps etc. o n humus-rich, sandy or loamy-clayey, nutrient-rich,
applanate, w i t h straight, i n f l e x e d or deflexed m a r g i n , c o m p l e t e l y set ; soils; rather rare, absent f r o m the regions w i t h nutrient-poor sand as
w i t h spines, those at centre fine and almost c y l i n d r i c a l to p y r a m i d a l , . main component o f the s o i l . M i d A u g . - m i d N o v . W i d e s p r e a d i n
0.5-1.5 m m h i g h , around centre often i n concentrical zones and more Europe, but nowhere very c o m m o n .
adnate, r e c u r v e d and fibrillose; c o l o u r o f spines dark b r o w n (at least
at apex) o n a p i n k i s h b r o w n , p i n k i s h orange-brown to o c c a s i o n a l l y The squamules o n pileus and stipe show a remarkable variability, from
c r e a m b a c k g r o u n d ( M u . 7.5 Y R 7/4-6, 6/4, 6-8/5-6; K . & W . 4 A 3 , fine and short to more p y r a m i d a l ; K n u d s e n (in B o t . Tidsskr. 7 5 : 1 3 5 .
5 B 4 ) ; b a c k g r o u n d f e l t e d - f i b r i l l o s e , e s p e c i a l l y at centre i n patches 1 9 8 0 ) distinguished three types o f velar coverings o n pileus and stipe,
around squamules; spines sometimes washed o f f and l e a v i n g i n d i s - but he also discovered transitions between the different types. O n
tinct, pale scars; m a r g i n e x c e e d i n g lamellae. L a m e l l a e , L = 4 0 - 7 0 , 1 account o f these transitions it was not possible to distinguish taxa o n
= 1-5, c r o w d e d to moderately distant, free, ventricose to subventri- this character; n o r was there a correlation between macromorphology
cose, up to 9 m m w i d e , white to c r e a m or pinkish-creamy, w i t h even and the presence or absence o f cheilocystidia. T h i s leaves Lepiota echi-
to very fine-flocculose white edge. Stipe 22-62 X 4 - 1 2 m m , c y l i n - nacea as a variable species.
d r i c a l , often s l i g h t l y broadened at utmost base, fistulose, i n upper
half c r e a m to pale b r o w n i s h or p i n k i s h c r e a m and shiny to w o o l l y fib- 43. Lepiota jacobi V e l l i n g a & K n u d s e n i n V e l l i n g a i n Persoonia 1 4 :
r i l l o s e , i n l o w e r half, b e l o w indistinct annular zone, pale p i n k - b r o w n 407. 1992.-Fig. 116. S
to orange-brown (e.g. 7.5 Y R 7/6) tomentose, w i t h distinct and acute, Echinoderma jacobi ( V e l l i n g a & Knudsen) R a i d et a l . i n Svampe
though often less acute than o n pileus, or floccose, dark red-brown or 2 6 : 3 6 . 1 9 9 2 (not v a l i d ; as E. jacobii); Lepiota langei K n u d s e n i n B o t .
orange-brown squamules or warts, often i n bands, rather densely set Tidsskr. 7 5 : 1 3 0 . 1 9 8 0 , non Lepiota langei L o c q . , 1 9 4 5 .
M I S A P P L . - Lepiota eriophora sensu auct. eur.; Cystolepiota erio-
phora sensu K n u d s e n i n B o t . Tidsskr. 7 3 : 1 2 7 . 1 9 7 8 ; Echinoderma
eriophorum sensu M . B o n i n D o c . m y c o l . 2 1 ( 8 2 ) : 6 3 . 1 9 9 1 .
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4 : p i . 2 3 1 . 1 9 9 5 (as
L. langei); J . L a n g e , F l . agar. dan. 1: p i . 1 2 H . 1 9 3 5 (as L. echinella var.
eriophora); R a i d et a l . i n Svampe 2 6 : 3 5 . 1 9 9 2 (as Echinoderma jaco-
bii).
F i g . 115. Lepiota echinacea. F i g . 116. Lepiota jacobi.

S E L . D E S C R . & F I G S . - Candusso & L a n z o n i , L e p i o t a : 144-146, fig.

21. 1990 (as L. langei); E n d e r l e & K r i e g l s t . i n Z . M y k o l . 55: 78. 1989 Q
n
(as L. langei); K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 52-53. 1994;
K n u d s e n i n B o t . Tidsskr. 75: 130-131, figs. I B , 3 K & 6. 1980 (as L .
langei); Schatzle & O t t m a n n i n U l m e r P i l z f l . 3: 63-64. 1992 (as L.
0
langei).
V E R N . N A M E - WoUige stekelparasolzwam.
Pileus 19-51 m m , w h e n y o u n g hemispherical w i t h inflexed margin,

expanding to plano-convex or applanate w i t h low, broad umbo, and
undulating marginal zone, completely covered w i t h small discrete acute
spines, up to 1 m m high, higher than broad, towards margin spines c o n -
F i g . 117. Lepiota carinii.
centrically arranged, slightly bigger and more adnate and fibrillose,
around squamules, except at centre, showing the underlying pale sordid
brown, beige to buff ( M u . 10 Y R 8-7/4) background; squamules dark Echinoderma carinii (Bres.) M . B o n i n D o c . m y c o l . 21 (82): 6 3 .
b r o w n to b l a c k - b r o w n (10 Y R 2/2, 3/3-4); w h e n y o u n g w i t h velar rem- 1991.
nants at margin. L a m e l l a e , L = 35-50,1 = 1-3, moderately crowded to E X C L . - Lepiota carinii sensu Breitenb. & K r a n z l . , P i l z e S c h w e i z
moderately distant, free, subventricose, 4-6 m m w i d e , cream, or cream 4: 194-195. 1995 ( = Lepiota spec).
w i t h pale orange tinge, w i t h concolorous, even to white, flocculose M I S A P P L . - Lepiota echinella var. asperula sensu J. L a n g e , F l . agar,
edge. Stipe 20-60 X 3-6 m m , cylindrical, sometimes slightly broadened dan. 1: p i . 10C. 1935; Lepiota eriophora var. asperula sensu M a i . &
towards base, hollow, w i t h annular zone halfway, above this zone white- Bert., F l . C h a m p , sup. M a r o c 1: 126-127. 1970.
cream w h e n young to pale p i n k i s h orange-tinged, or pale brown, fibril- S E L . I C O N . - Bres., Iconogr. m y c o l . 12: p i . 5 9 8 . ( ' 1 9 2 9 ' ) 1930
lose-flocculose (10 Y R 8/5, 7.5 Y R 7/6-8/5), below annular zone w i t h G a l l i n a r i & T o m a s i i n B o l l . C i r c . m i c o l . G . C a r i n i 15: opp. p . 18. 1988
girdles and zones o f woolly, b r o w n (7.5 Y R 5-4/4) squamules o n paler J. Lange, F l . agar. dan. 1: p i . 10C. 1935 (as L. echinella var. asperula)
background. Context whitish and d u l l i n pileus, whitish around tube i n Partacini i n Riv. M i c o l . 38: 283. 1995.
stipe, and concolorous w i t h surface i n outer part o f stipe, shiny. S m e l l S E L . D E S C R . & FIGS. - G a l l i n a r i & Tomasi i n B o l l . C i r c . m i c o l . G .
unpleasant, earth-like and sweet, a bit like the s m e l l o f L. cristata. Taste C a r i n i 15: 15. 1988; K n u d s e n i n Bot. Tidsskr. 75: 131,132, figs. I D , 3 C
unpleasant, slightly bitterish raphanoid. Spore print cream (Breitenbach & 8.1980; M a i . & Bert., F l . C h a m p , sup. M a r o c 1: 126-127.1970 (as L.
& K r a n z l i n , P i l z e S c h w e i z 4: p i . 231. 1995). eriophora var. asperula); M i g l . & C o c c i a i n M i c o l . ital. 21 (2): 35-40.
Spores 3.5-5.5 X 2.0-3.0 um, Q = 1.45-2.05, Q a v = 1.65-1.75, 1992; Partacini i n Riv. M i c o l . 38: 282-284. 1995; Schatzle & Ottmann
e l l i p s o i d to oblong, often w i t h parallel sides i n side-view, the same o r in U l m e r P i l z f l . 3: 65-67. 1992; U l j e i n C o o l i a 27: 12-16. 1984.
slightly o v o i d i n frontal view, dextrinoid, not metachromatic i n C r e s y l V E R N . N A M E - Bruingele stekelparasolzwam.
B l u e , often i n tetrads. B a s i d i a 13.5-22 X 4.0-6.0 p m , narrowly clavate,
a few c y l i n d r i c a l , 4-spored. L a m e l l a edge fertile. C h e i l o c y s t i d i a Pileus 15-45 m m , when y o u n g hemispherical, then broad-conical w i t h
absent. Pleurocystidia absent. Elements o f pileus covering organized inflexed margin, expanding to plano-convex or convex, w i t h undulat-
in squamules, made up o f more or less regular chains, w i t h terminal ing m a r g i n and broad umbo, completely beset w i t h acute or blunt,
elements globose, p y r i f o r m o r e l l i p s o i d , 20-60 X 15-48 p m , w i t h orange-brown to dark b r o w n squamules (darker than M u . 7.5 Y R 4/6;
b r o w n , t h i c k w a l l s ; intermediate elements b r o w n - w a l l e d , inflated to 7.5 Y R 4/4), c. 0.5 m m h i g h and broad, o n a y e l l o w - b r o w n , pale
cylindrical, 5.0-20 p m w i d e ; basal elements c y l i n d r i c a l to inflated, 4.0- orange-brown to (pale) b r o w n background (5 Y R 5/8, 7.5 Y R 7/6, 8/6
15 p m wide, w i t h b r o w n , not encrusted w a l l s . S t i p i t i p e l l i s a cutis, or paler), when y o u n g w i t h overhanging margin set w i t h w h i t i s h w o o l -
lower part made up o f c y l i n d r i c a l to inflated, 5.0-20 p m w i d e b r o w n ly v e l u m . L a m e l l a e , L = 45-55,1 = 1-3, rather crowded, free, ventri-
hyphae, w i t h encrusted pigment, w i t h elements i n squamules as o n cose, subventricose o r segmentiform, 3-5 m m broad, white w h e n
pileus, w i t h b r o w n not-encrusted walls, and u p to 110 X 45 p m . young, cream and brown-spotted w i t h age, w i t h w h i t i s h , even or f i m -
Clamp-connections present i n a l l tissues. briate (lens), concolorous edge. Stipe (15-)25-60 X 3.5-7(-8) m m ,
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres- c y l i n d r i c a l o r slightly w i d e n e d at base, c u r v e d near base, h o l l o w , i n
trial o n humus-rich o r nutrient-rich l o a m y to clayey soil, i n city-parks, upper 1/4-1/3 part white and fibrillose-striate, i n lower part w i t h gir-
gardens, deciduous and m i x e d w o o d s , also o n m i n e waste heaps. dles o f orange-brown, acute squamules, w i t h white m y c e l i a l cords at
Rather rare, mostly occurring i n southern L i m b u r g and H o l l a n d . June- base. A n n u l u s w o o l l y and evanescent. Context white and d u l l i n pileus,
Oct. Widespread but not c o m m o n i n Europe. in stipe white to pale creamy and shiny, w i t h age b r o w n i s h at base.
S m e l l indistinct to strong and like the s m e l l o f L. cristata. Taste not
T h i s species has been k n o w n as Lepiota eriophora A t k . , but this, k n o w n . Spore print c o l o u r white.
according to K n u d s e n (in B o t . Tidsskr. 75: 131. 1980), is a different Spores (3.0-)3.5-4.5(-5.0) X 2.0-3.0 um, Q = 1.35-1.8, Q a v = 1.6-
species, not occurring i n E u r o p e . B o n ( i n D o c . m y c o l . 21 (82): 6 3 . 1.65, ellipsoid to oblong, dextrinoid, not or w e a k l y metachromatic i n
1991) d i d not agree w i t h K n u d s e n ' s point o f v i e w and c a l l e d the C r e s y l B l u e , often i n tetrads. B a s i d i a 13.5-23 X 4.0-6.5 p m , 4-spored.
E u r o p e a n species E. eriophorum. Unfortunately K n u d s e n ( i n B o t . L a m e l l a edge fertile o r w i t h some scattered c h e i l o c y s t i d i o i d elements
Tidsskr. 75: 130. 1980) chose a new name already v a l i d l y published i n or rarely completely sterile. C h e i l o c y s t i d i a 16-33 X 5.0-8.5 p m ,
Lepiota, v i z . L. langei L o c q . , for the E u r o p e a n interpretation o f L . clavate, rarely c y l i n d r i c a l , h y a l i n e and t h i n - w a l l e d . P l e u r o c y s t i d i a
eriophora, so again a name change was necessary: L. jacobi Vellinga absent. Squamules o n pileus made u p o f agglutinated chains o f glo-
& K n u d s e n (in V e l l i n g a i n Persoonia 14: 4 0 7 . 1992). bose to e l l i p s o i d elements, w i t h the terminal elements more globose
and bigger than the penultimate elements; terminal elements 10-25(-
44. Lepiota carinii Bres., Iconogr. m y c o l . 12: p i . 598. ('1929') 1930. 45) X 10-21(-35) u m w i t h y e l l o w - b r o w n w a l l s ; intermediate hyphae
-Fig. 117.
w i t h inflated elements, 16-25 X 4-6 u m , w i t h b r o w n w a l l s ; elements broad, cream w i t h pale s a l m o n sheen, w i t h even, c o n c o l o r o u s edge.
of pileus c o v e r i n g made up o f non-coloured, c y l i n d r i c a l hyphae, c. 4.0 Stipe 20-45 X 1.5-3 m m , c y l i n d r i c a l o r s l i g h t l y broadened at base,
p m w i d e . S t i p i t i p e l l i s a cutis made up o f c y l i n d r i c a l , non-coloured h o l l o w w i t h age, c r e a m c o l o u r e d a n d almost glabrous at apex, i n
hyphae, 2.0-5.0 p m i n diam.; squamules o n stipe made u p o f chains o f l o w e r 3/4 w i t h s l i g h t l y w o o l l y girdles o f squamules, more o r less
elements, comparable i n shape and size to those on pileus. C l a m p - c o n - c o n c o l o r o u s w i t h squamules o n pileus (5 Y R 5/4-6) o n c r e a m o r
nections present i n a l l tissues. slightly p i n k i s h b a c k g r o u n d . C o n t e x t i n pileus w h i t e - c r e a m and d u l l ,
H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres- i n stipe shiny c r e a m at apex to p i n k - b r o w n at base. S m e l l o f cut
trial o n the outskirts o f deciduous woods and i n city-parks on nutrient- basidiocarps strong, unpleasant, mealy-turpentine-like a n d s l i g h t l y
rich, clayey o r l o a m y soils. V e r y rare i n the Netherlands, k n o w n from sweetish. Taste unpleasant, l i k e the taste o f L. cristata. Spore print
several places i n southern L i m b u r g and i n the western clay area. Sept.- probably white.
beginning o f N o v . Rare i n Europe, k n o w n northwards into Denmark, Spores (3.5-)4.0-5.0(-6.0) X 2.0-3.0 p m , Q = (1.4-)1.55-2.1(-2.35),
perhaps more c o m m o n i n Italy, and recorded from M o r o c c o . Qav = 1.65-1.9, o b l o n g to c y l i n d r i c a l i n side-view and i n frontal view,
often i n clusters, dextrinoid, (weakly) metachromatic i n C r e s y l B l u e .
Lepiota sinopica comes very close, but it seems to lack distinct squa- B a s i d i a 14-24 X 4.5-6.5 p m , c y l i n d r i c a l to narrowly clavate, 4-spored,
mules o n the stipe (see R o m a g n e s i i n B u l l . Soc. nat. O y o n n a x 10-11: a few 2-spored. L a m e l l a edge fertile. C h e i l o c y s t i d i a and pleurocystidia
91-92. 1957 ( C o m p l . F l . anal. 7)). Lepiota sinopica is characterized by absent. Squamules o n pileus made up o f chains o f elongate to more
orange-brown colours, and a velvety pileus w i t h small squamules. T h i s ellipsoid to globose terminal elements; terminal elements (15-)30-55
species, however, i s rather enigmatic a n d the finds o f i t since X (15-)26-48 p m , w i t h y e l l o w - b r o w n w a l l s . B a s a l elements c y l i n d r i -
R o m a g n e s i ' s description are dubious (e.g. Braiotto i n B o l l . A s s . m i c o l . cal, 3.0-6.0 p m w i d e , w i t h b r o w n w a l l s . Stipitipellis a cutis o f c y l i n -
ecol. R o m a n a 25: 28-33. 1992). I n the latter the p i n k colours and the drical, 2.0-6.0 u m w i d e elements; c o v e r i n g made up o f elements
s m a l l m e a l y squamules indicate a possible similarity o f the collection resembling those o n pileus, often w i t h spheropedunculate elements.
depicted w i t h Cystolepiota moelleri. Clamp-connections abundant i n a l l tissues.
H A B I T A T & DISTR. - Solitary o r gregarious, saprotrophic and terres-
45. Lepiota pseudoasperula (Knudsen) K n u d s e n i n B o t . Tidsskr. 75: trial o n humus-rich, clayey or l o a m y soils, i n deciduous woods and o n
128. 1 9 8 0 . - F i g . 118. a-anine waste heap. Very rare i n the Netherlands, k n o w n from several
Cystolepiota pseudoasperula K n u d s e n i n B o t . Tidsskr. 7 3 : 125. places i n southern L i m b u r g , and from the province o f Flevoland. E n d
1978; Echinoderma pseudoasperulum (Knudsen) M . B o n i n D o c . of J u l y - O c t . V e r y rase a n d scattered i n E u r o p e , f r o m N o r w a y i n the
m y c o l . 21 (82): 63. 1991. north, to the north o f Italy i n the south.
E X C L . - Lepiota pseudoasperula sensu Enderle & K r i e g l s t . i n Z .
Mykol. 5 5 : 86. 1989 ( = Cystolepiota moelleri); Lepiota aff. Lepiota pseudoasperula i s often mistaken for Cystolepiota moelleri
pseudoasperula sensu Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: 204-205. (and v i c e versa) i n the field. T h e m i c r o s c o p i c a l data, v i z . cheilocys-
1995 ( = probably L. carinii) tidia absent i n L. pseudoasperula, present i n C. moelleri, and spores
M I S A P P L . - Lepiota eriophora sensu D . R e i d i n Trans. B r . m y c o l . dextrinoid i n L. pseudoasperula and non-dextrinoid i n C. moelleri, are
Soc. 4 1 : 4 2 6 - 4 2 7 . 1958. decisive, and easy to check.
S E L . I C O N . - R a i d et a l . i n Svampe 26: 35. 1992.
S E L . D E S C R . & FIGS. - Candusso & L a n z o n i , L e p i o t a : 151-152, fig. 46. Lepiota boertmannii K n u d s e n i n B o t . Tidsskr. 75: 150. 1980. -
23. 1990; Courtecuisse i n Beitr. K e n n t n . P i l z e Mitteleur. 3: 314-316, F i g . 119.
fig. 2. 1987; Huijser & K e l d e r m a n i n C o o l i a 34: 83-86, fig. 2. 1991; Echinoderma boertmannii (Knudsen) M . B o n i n D o c . m y c o l . 21
K n u d s e n i n B o t . Tidsskr. 75: 128, figs. I C , 3 G & 4. 1980. (82): 62. 1991.
V E R N . N A M E - K l e i n e stekelparasolzwam. S E L . I C O N . - R a i d et a l . i n Svampe 26: 35. 1992.
S E L . D E S C R . & FIGS. - K n u d s e n i n B o t . Tidsskr. 75: 150-151, figs.
P i l e u s 10-27 m m , w h e n very y o u n g h e m i s p h e r i c a l , later more c o n i - IF, 2 D & 3 D . 1980.
cal w i t h blunt apex, expanding to plano-convex w i t h l o w umbo, beset VERN. NAME - Buitenbeenstekelparasolzwam.
w i t h p i n k i s h b r o w n to dark r e d d i s h b r o w n ( M u . 5 Y R 3/3, 4/4-6),
pointed squamules up to 1 m m h i g h and 1 m m broad, close together Pileus 20-40 m m , rounded c o n i c a l w i t h inflexed m a r g i n to plano-con-
at centre, more spaced out towards m a r g i n , o n pale, r a d i a l l y short- vex w i t h o r without broad umbo, set w i t h c o n i c a l , p i n k - b r o w n ( K . &
fibrillose b a c k g r o u n d (7.5 Y R 7-8/4-6); squamules easily removable; W . 7 E F 6 ) o r dark b r o w n ( M u . 5-7.5 Y R 4/4) squamules up to 0.75 m m
m a r g i n e x c e e d i n g l a m e l l a e . L a m e l l a e , L = 25-45, 1 = 0-3, moder- h i g h o n felted cream-pink o r dark isabella (7.5 Y R 7/6) background.
ately c r o w d e d , free, segmentiform to s l i g h t l y ventricose up to 2 m m L a m e l l a e , L = c. 6 0 , 1 = 3, moderately c r o w d e d to crowded, free, u p
to 3 m m broad, w h i t i s h but rather y e l l o w i s h w h e n l o o k e d into ( 3 A 2 -
3), w i t h white flocculose edge. Stipe 30-40 X 3-4(-9) m m , c y l i n d r i c a l
and slightly broadened at base, w i t h w h i t i s h to p i n k - b r o w n annular
zone, above this zone smooth, b e l o w this zone w o o l l y and w i t h w o o l -
ly girdles o f squamules as o n pileus (10 Y R 4/4), w i t h l o n g i t u d i n a l
b r o w n fibrils over w h o l e length. Context white i n pileus a n d stipe.
S m e l l more o r less like L. cristata, or not distinct. Taste not k n o w n .
Spore print white.
Spores 4.0-6.0(-6.5) X 2.5-3.0 p m , Q = 1.6-2.3, Q a v = 1.85-2.0,
Q
oblong to c y l i n d r i c a l i n side-view, i n frontal v i e w more o v o i d , non-
DO a m y l o i d , not d e x t r i n o i d (not c o l o u r i n g i n L u g o l , nor i n M e l z e r ' s

Reagent), pale p i n k i n C o n g o R e d , metachromatic i n C r e s y l B l u e .
F i g . 118. Lepiota pseudoasperula. B a s i d i a 14-24 X 5.0-6.5 p m , 4-spored, some 2-spored, or even 1-
cylindrical, c. 30-40 X 12-20 p m ; basal hyphae cylindrical, 4.0-6.0 p m ,

w i t h encrusting b r o w n pigment. S t i p i t i p e l l i s w i t h squamules as o n
pileus, but elements slightly smaller and more irregular. C l a m p - c o n -
terrestrial o n nutrient- and humus-rich soils i n deciduous woods. Very
rare i n the Netherlands, k n o w n from two localities i n southern L i m b u r g
(Elsloo, Bunderbos; B e m e l e n ) ; Aug.-Sept. Very rare i n Europe, record-
ed from Germany and D e n m a r k .
The macroscopical description is based o n the o r i g i n a l description by

K n u d s e n (in B o t . Tidsskr. 75: 150-151. 1980) and o n notes accompa-
F i g . 119. Lepiota boertmanii.
n y i n g the Netherlands' collections.
T h e photograph published by R a i d et a l . (in Svampe 26: 35. 1992)
spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a 15-31 X 5.5-10 um, nar- depicts a specimen o f this species w i t h a different k i n d o f pileus c o v -
r o w l y clavate to clavate, slightly t h i c k - w a l l e d , hyaline. Pleurocystidia ering than the D u t c h specimens. In the Netherlands' collections, the
absent. Pileus covering consisting o f squamules, made up o f aggluti- pilei were covered w i t h a tomentose layer, set w i t h s m a l l , pointed squa-
nated elements i n chains, 12-35(-47) X 10-30(-40) u m , globose to mules. It is entirely possible that this species, just l i k e Lepiota aspera
ellipsoid, w i t h b r o w n , thickened w a l l s ; intermediate hyphae inflated- and L. echinacea, shows a w i d e variation i n pileus covering.
8. Chamaemyces Batt. ex Earle
ELSE C . VELLINGA
Chamaemyces Batt. ex Earle in Bull. N.Y. bot. Gdn 5: 446. 1909; Lepiota subgen. Lepiotella E.J. Gilb., Genre Amanita:
159. 1918; Lepiotella (E.J. Gilb.) Konr. in Schweiz. Z. Pilzk. 12: 177. 1934, non Lepiotella J. Rick, 1938. - Drosella
Maire in Kiihn, & Maire in Bull, trimest. Soc. mycol. Fr. 50: 15. 1934 (not valid).
SELECTED LITERATURE - Migl. & Zecchin in Micol. ital. 22 (3): 142-156. 1993.
Basidiocarp collybioid; pileus rather fleshy, viscid; lamellae free or almost free and then emarginate; spore print pale
cream.
Spores hyaline, without germ pore, binucleate, non-amyloid, not dextrinoid; complete wall metachromatic in Cresyl
Blue; hymenophoral trama regular; cystidia present; covering of pileus a hymeniderm; clamp-connections present;
development monovelangiocarpic and stipitocarpic; universal veil with radiating hyphae at the outside present on pileus
only (Reijnders in Persoonia 8: 318. 1975). - Holotype species: Armillaria fracida (Fr.) Gillet.
HABITAT & DISTRIBUTION - Saprotrophic, terrestrial and on very decayed wood. One species known from Europe and
possibly two other species in South America.
Chamaemyces fracidus (Fr.) D o n k i n B e i h . N o v a H e d w i g i a 5: 4 8 .

1962. - F i g . 120.
Agaricus fracidus Fr., E p i c r i s i s : 25. 1838; Armillaria fracida (Fr.)
G i l l e t , H y m e n o m y c e t e s : 7 7 . 1874; Drosella fracida (Fr.) S i n g , i n
L i l l o a 22: 446. ('1949') 1951 (not valid). - Lepiota irrorata Q u e l , i n
C . r. A s s . franc. A v . S c i . ( L a R o c h e l l e , 1882) 11: 387. 1883 ( C h a m p .
Jura Vosges S u p p l . 11); Lepiotella irrorata (Quel.) K o n r . i n S c h w e i z .
Z . P i l z k . 12: 177. 1934; Drosella irrorata (Quel.) K i i h n e r & M a i r e i n
Bull, trimest. S o c . m y c o l . F r . 5 0 : 15. 1934 ( n o m . invalid). -
Chamaemyces fracidus var. pseudocastaneus B o n & Boiff. i n B u l l ,
trimest. S o c . m y c o l . Fr. 90: 303. ('1974') 1975; Chamaemyces pseudo-
castaneus ( B o n & B o i f f . ) C o n t u i n B o l . S o c . B r o t . , Ser. II, 6 3 : 384.
1990; Lepiotella irrorata f. pseudocastanea ( B o n & Boiff.) M i g l . &
Z e c c h i n i n M i c o l . ital. 22 (2): 5 1 . 1993. - Lepiotella irrorata f.
passerinii M i g l . & Z e c c h i n i n M i c o l . ital. 22 (2): 51. 1993. - Agaricus
demisannulus Fr., E p i c r i s i s : 19. 1838; Lepiota demisannula (Fr.) S a c c ,
S y l l . F u n g . 5: 6 9 . 1887; Drosella demisannula (Fr.) L o c q . i n B u l l ,
mens. S o c . l i n n . L y o n 2 3 : 5 4 . 1954 (not valid); Chamaemyces
demisannulus (Fr.) M o s . , R o h r l i n g e Blatterpilze, 3. A u f l . : 194. 1967.
D a h n c k e , 1200 P i l z e : 5 0 9 . 1993; Laess0e i n S v a m p e 4 1 : 5 6 . 2 0 0 0 ;
M i g l . & Z e c c h i n i n M i c o l . ital. 2 2 (3): 146-147. 1993 (as Lepiotella
irrorata, i n c l . f. passerinii and f. pseudocastanea); Pepetti et a l . i n
B o l l . C i r c . m i c o l . G . C a r i n i 39: 15. 1999 (as Lepiotella irrorata); Pat.
i n B u l l , trimest. S o c . m y c o l . Fr. 102: A t l a s p i . 2A5. 1986 (as Lepiota
irrorata). \
S E L . D E S C R . & F I G S . - A n o n , i n B u l l , trimest. S o c m y c o l . Fr. 102:
A t l a s p i . 245. 1986 (as Lepiota fracida); Breitenb. & K r a n z l . , P i l z e
S c h w e i z 4 : 182. 1995; K i i h n e r i n B u l l , trimest. Soc. m y c o l . F r . 52:
2 0 2 - 2 0 4 . 1 9 3 6 (as Lepiota irrorata); M i g l . & Z e c c h i n i n M i c o l . ital. 22
(3): 144-150, figs. 1-3. 1993 (as Lepiotella irrorata, i n c l . f. passerinii
and f. pseudocastanea); Reijnders i n Persoonia 8: 309-311; 313-315;
pi. 48, 4 9 A , B . 1975 (on the development o f the basidiocarps).
VERN. NAME - Druppelparasolzwam. F i g . 120. Chamaemyces fracidus.
152
CHAMAEMYCES (Vellinga) 153
Pileus 37-65 m m , when y o u n g more or less hemispherical, expanding derm, covered w i t h or without gelatinous layer up to 10 p m thick,
to applanate w i t h low, broad u m b o or plano-convex w i t h applanate made up o f clavate and narrowly clavate elements, 24-60 X 8.5-19 pm,
centre, ochre-yellow or c r e a m y - p i n k i s h at centre to very pale y e l l o w at with or without more c y l i n d r i c a l elements, 20-50 X 4.0-8.5 p m , w i t h
margin, w h e n moist a bit v i s c i d , d i s c o l o u r i n g w h e n handled, smooth, brown refractive pigment; walls o f elements metachromatic i n C r e s y l
but i n marginal part slightly radially rugose, w i t h fibrillose velar rem- B l u e . Stipitipellis a cutis, made up o f c y l i n d r i c a l , colourless hyphae,
nants between margin and stipe. L a m e l l a e , L = c. 4 0 , 1 = c. 3, m o d - 2.0-4.0 p m w i d e , w i t h i n l o w e r part o f stipe scattered clusters o f
erately c r o w d e d to crowded, free or narrowly adnate, ventricose, up to brown-coloured, irregular, sometimes ascending hyphae, up to 50 X
6 m m broad, white, pale beige or cream w i t h p i n k i s h sheen, w i t h even, 5.0 um. Clamp-connections present i n a l l tissues, very conspicuous.
concolorous edge. Stipe 40-65 X 10-12 m m , narrowing towards base, H A B I T A T & DISTR. - In s m a l l groups, probably saprotrophic, terres-
fistulose, white, smooth and glabrous at apex, b e l o w annular zone w i t h trial i n avenues and woods o n nutrient-rich and calcareous sandy and
small brownish floccules o n w h i t i s h to ochre-brown background, when l o a m y soils, also k n o w n f r o m chalk-grasslands. In the Netherlands
y o u n g w i t h droplets. A n n u l u s present and ascending, or only as a velar rare, especially occurring i n the river area, the dune area and southern
zone. Context white and d u l l i n pileus, shiny and cream i n stipe. S m e l l L i m b u r g , d e c l i n i n g . Sept.-Oct. W i d e s p r e a d i n Europe, but absent or
l i k e L. cristata, or sweet and more sourish w h e n basidiocarp cut. Taste very rare i n the northern countries.
not k n o w n . Spore print pale cream.
Spores (4.0-)4.5-5.5(-6.0) X 2.5-4.0 p m , Q = (1.25-)1.35-1.75, D o n k (in B e i h . N o v a H e d w i g i a 5: 48-49. 1962) has been f o l l o w e d for
Qav = 1.5-1.55, e l l i p s o i d , though some o b l o n g , sporadically broadly the nomenclature o f this species.
e l l i p s o i d i n side-view, i n frontal v i e w more o v o i d , non-dextrinoid, not Chamaemyces demisannula is, according to L o c q u i n (in B u l l ,
congophilous, metachromatic i n C r e s y l B l u e , cyanophilous. B a s i d i a mens. Soc. l i n n . L y o n 23: 54. 1954), a different species, characterized
15.5-30 X 5.0-8.5 p m , 4-spored. L a m e l l a edge heterogeneous. by a hardly developed gelatinous layer o n the pileus and scarce cys-
C h e i l o c y s t i d i a abundant, occasionally very rare, s i m i l a r to pleurocystidia. However, these characters are not correlated; basidiocarps w i t h
tidia. Pleurocystidia abundant, 36-73 X 11.5-21 p m , oblong-utriform, a l o w number o f cystidia may have a thick gelatinous layer, so the two
rounded-fusiform or more clavate, w i t h at least i n dry material refrac- species are considered synonymous.
tive, evenly distributed or at base a bit granular, content, sometimes D a r k variants o f Ch. fracidus are recognized as forma pseudocas-
w i t h crystal floes at apex; cystidia occasionally very rare and then h y a - tanea ( B o n & Boiff.) M i g l . & Z e c c h i n , but this taxon has not yet been
line; walls metachromatic i n C r e s y l B l u e . Pileus covering a h y m e n i - encountered i n the Netherlands.
9. Cystolepiota Sing.
E L S E C. VELLINGA
Cystolepiota Sing, in Sing. & Digilio in Lilloa 25: 281. ('1951') 1952. - Pulverolepiota M . Bon in Doc. mycol. 22 (88):
30. 1993. - Lepiota subgen. Sphaerocystae Wasser in Ukr. bot. Zh. 35: 517. 1978.
SELECTED LITERATURE - M . Bon in Doc. mycol. 11 (43): 23-26. 1981; M . Bon, Fl. mycol. Eur. 3, Lepiotes: 38-42.
1993; Candusso & Lanzoni, Lepiota: 77-114. 1990; Kelderman, Parasolzw. Zuid-Limburg: 29-41. 1994; Vellinga &
Huijser in Persoonia 16: 513-526. 1998; Zecchin in Riv. Micol. 43: 147-165. 2000.
Basidiocarp pluteoid, rather small; pileus covered in granules from velum universale; velum partiale leaving an annu-
lar zone on stipe; lamellae free; spore print white to pale yellow or pale lilac.
Spores hyaline, with smooth or slightly rough wall, without a germ pore, uninucleate, in a few species binucleate,
often in tetrads, non-amyloid nor dextrinoid, rarely (faintly) dextrinoid, in most species with inner wall pink in Cresyl
Blue, congophilous, with walls not swelling in ammonia (NH - )) and acetic acid ( C H C O O H ( ) ; cheilo- and pleu-
3( aq 3 aq
rocystidia present or absent, or only cheilocystidia present; hymenophoral trama regular; pileus covering made up of
more or less loosely arranged globose to oblong or elongate, inflated elements; clamp-connections present, in some
species absent. Development bivelangiocarpic and pileostipitocarpic. - Type species: Cystolepiota constricta Sing.
HABITAT & DISTRIBUTION - Saprotrophic, terrestrial in woods, world-wide, but very rare in alpine and arctic regions.
Cystolepiota is closely related to the genus Melanophyllum Velen. and to Lepiota sect. Echinatae Fay. It differs from
Melanophyllum mainly in the coloration of the spores, and from Lepiota sect. Echinatae by the structure of the veil:
only thin hyphae and globose (or inflated), relatively loosely arranged elements in Cystolepiota, versus thin hyphae,
gradually transient into globose to ellipsoid elements, agglutinated to form pyramidal squamules in Lepiota sect.
Echinatae. Furthermore, in most Cystolepiota species the spores are uninucleate, whereas they are binucleate in Lepiota
sect. Echinatae. For an extensive comparison of the two taxa see also Knudsen (in Bot. Tidsskr. 75: 123-127. 1980).
K E Y TO T H E SPECIES
1. Basidiocarps with distinct lilac or violaceous tinges and a strong smell of coal gas, or with yellow (sulphur-yellow
or yellowish brown) tinges in all parts; spores dextrinoid (though the reaction may be slow)
2. Basidiocarps with distinct lilac or violaceous tinges and a strong smell of indole (coal gas); spores (6.5-)7.0-9.0(-
10.0) pm long 6. C . bucknallii
2. Basidiocarps yellow (sulphur-yellow or yellowish brown), smell indistinct; spores 3.5-5.5(-6.0) pm long
7. C . icterina
(if pileus margin/sulcate, clamp-connections absent, and spores 5.0-8.5 X 4.0-7.0 pm, see Leucocoprinus
' straminellus)
1. Basidiocarps with white,\greyish, orange or pink to brownish colours; spores not reacting in Melzer's Reagent (if
slowly and weakly dextrinoid, then elements of pileus covering elongate)
3. Elements of pileus covering elongate, oblong, inflated; clamp-connections absent; cystidia absent
8. C . pulverulenta
(if spores with germ pore, and cheilocystidia present, see Leucocoprinus)
3. Elements of pileus covering globose; clamp-connections present; cystidia present or absent
4. Cheilocystidia absent; pileus white, rarely pale pink or yellowish, not discolouring with age or when damaged
5. C . seminuda
4. Cheilocystidia present; pileus either white and discolouring orange-brown, or with other colours
5. Basidiocarps white, cream or greyish, discolouring (orange-)brown with age or on bruising
154
CYSTOLEPIOTA (Vellinga) 155
6. Cheilocystidia and pleurocystidia with yellow contents and exudates 1. C . cystidiosa

6. Cheilocystidia (and pleurocystidia) without yellow contents and exudates
7. Spores 4.0-6.0(-6.5) X 2.0-3.0(-3.5) pm, Q = (1.5-)1.6-2.1(-2.3), Qav = 1.8-2.0; pleurocystidia pres-
ent, especially close to lamella edge 2. C . hetieri
7. Spores 5.0-6.5(-7.0) X 2.0-3.0 pm, Q = (1.8-)2.0-2.6, Qav = 2.1-2.4; pleurocystidia absent
3. C . adulterina
5. Basidiocarps with greyish, pale lilac-brown, cream or pink(-brown) covering, not discolouring orange-
brown when damaged or with age
8. Pileus covered with small pink-coloured granulose warts, often resembling members of Lepiota sect.
Echinatae; cheilocystidia narrowly clavate to obovate, usually with cylindrical, moniliform or branched
excrescence at apex; pleurocystidia absent 4. C . moelleri
8. Pileus covered with a uniform, thick velar layer, or with pinkish grey-brown to greyish yellow, pyrami-
dal warts; pleurocystidia present or absent, if present with distinct yellow contents
9. Cheilocystidia and pleurocystidia with yellow contents or exudate; spores 3.5-5.5(-6.0) X 2.0-3.0 pm,
Q = 1.5-2.2(-2.4), Qav = 1.7-2.1; elements of pileus covering (10-)20-70(-95) pm
1. C . cystidiosa
9. Cheilocystidia without yellow contents or exudate; pleurocystidia absent; spores 5.0-6.5(-7.0) X 2.0-
3.0 pm, Q = (1.8-)2.0-2.6, Qav = 2.1-2.4; elements of pileus covering 15-45 pm
3. C . adulterina
Sect. Cystolepiota S E L . D E S C R . & FIGS. - H u i j s m a n i n S c h w e i z . Z . P i l z k . 39: 5 1 . 1961;

K r e i s e l i n W i s s . Z . Ernst M o r i t z A r n d t - U n i v . G r e i f s w a l d 16: 237-238.
Spores not dextrinoid, n o r a m y l o i d ; clamp-connections present; velar 1967 (as Lepiota lycoperdoides); D . R e i d i n F u n g . rar. Ic. c o l . 2: 9-10,
elements globose. fig 3. 1967 (as Lepiota luteicystidiata); A . H . S m i t h i n Papers M i c h .
A c a d . S c i . , A r t s Letters 27: 58-60, p i . 1, 2. ( ' 1 9 4 1 ' ) 1942; H . V . S m i t h
1. Cystolepiota cystidiosa ( A . H . S m i t h ) M . B o n i n D o c . m y c o l . 11 i n L l o y d i a 17: 318-319. 1954; V e l l i n g a & Huijser i n Persoonia 16: 518-
(43): 2 6 . 1 9 8 1 . - F i g . 121. 521, figs. 2b-d, 3b-c. 1998; Winterhoff & B o n i n C a r o l i n e a 52: 6. 1994;
Lepiota cystidiosa A . H . S m i t h i n Papers M i c h . A c a d . S c i . , A r t s Z e c c h i n i n R i v . M i c o l . 4 3 : 158-159. 2000.
Letters 27: 58 ('1941') 1 9 4 2 . - L e p i o t a luteicystidiata D . Reid in Fung, V E R N . N A M E - G e e l c e l l i g e poederparasol.
rar. Ic. c o l . 2: 9. 1967; Cystolepiota luteicystidiata (D. Reid) M . B o n

i n D o c . m y c o l . 6 (24): 4 3 . 1976. - Lepiota lycoperdoides Kreisel i n Pileus 20-40(-70) m m , w h e n y o u n g spherical to h e m i s p h e r i c a l w i t h
Wiss. Z . Ernst M o r i t z A r n d t - U n i v . G r e i f s w a l d 16: 2 3 8 . 1967; inflexed margin, expanding to applanate-campanulate, plano-convex
Cystolepiota luteicystidiata var. lycoperdoides (Kreisel) M . B o n i n w i t h o r without broad umbo, covered w i t h floccose, granular p y r a m i -
D o c . m y c o l . 11 (43): 2 6 . 1981. dal warts, up to several m i l l i m e t e r s h i g h , v a r y i n g i n c o l o u r from
M I S A P P L . - Lepiota rufescens sensu H u i j s m a n i n M e d e d . N e d . w h i t i s h when very y o u n g , to c r e a m d i s c o l o u r i n g p i n k i s h , to p i n k i s h
m y c o l . Vereen. 28: 47-48. 1943. grey-brown, p i n k i s h - b r o w n i s h ( M u . 7.5 Y R 6 / 4 , 1 0 Y R 4/3-7.5 Y R 5/4,
S E L . I C O N . - L o n a t i i n B o l l . A s s . m i c o l . e c o l . R o m a n a 12: 15. 10 Y R 7/6, 10 Y R 8/4 to 7.5 Y R 7/6), often staying paler at margin,
('1987') 1988 (as C . luteicystidiata); M i g l . et a l . i n R i v . M i c o l . 32: 103. where covering is thinner, showing the w h i t i s h context; w h e n warts are
1989 (as C. luteicystidiata); D . R e i d i n F u n g . rar. Ic. c o l . 2: p i . 10b. removed, a w h i t i s h 'scar' remains; m a r g i n exceeding lamellae.
1967 (as Lepiota luteicystidiata, dark specimens); Z e c c h i n i n R i v . L a m e l l a e , L = 30-45,1 = 1-3, moderately crowded to crowded, free,
M i c o l . 4 3 : 159. 2000. segmentiform to slightly ventricose, u p to 4.5 m m w i d e , cream
coloured, and brown-spotted w i t h age, w i t h even o r flocculose, white
to concolorous edge, w h i c h is also brown-spotted w i t h age. Stipe 15-
70 X 2.5-5 m m , c y l i n d r i c a l , sometimes w i d e n e d at apex, occasionally
curved i n lower part, fistulose, at apex w h i t i s h o r cream, finely pubes-
cent, glabrescent w i t h age, i n l o w e r half o r 3/4 o f length, b e l o w an
annular zone w h i c h is more or less distinct, granular floccose o r w i t h
floccose patches, concolorous w i t h pileus, often d i s c o l o u r i n g (e.g.
p i n k i s h b r o w n , 7.5 Y R 6/6) w h e n touched. Context white o r w h i t i s h
cream and d u l l i n pileus, shiny and cream i n stipe, b r o w n i s h o r vina-
ceous downwards. S m e l l fungoid or i e p i o t o i d ' , often l i k e Lepiota
cristata, but not always. Taste a b i t unpleasant, musty fungoid, w i t h
unpleasant lingering aftertaste. Spore print cream.
Spores 3.5-5.5(-6.0) X 2.0-3.0 p m , Q = 1.5-2.2(-2.4), Q a v = 1.7-
2.1, ellipsoid to c y l i n d r i c a l w i t h parallel sides, sometimes i n side-view
slightly w i d e n e d at base, often i n tetrads, colourless, w i t h slightly
thickened w a l l , non-dextrinoid, n o n - a m y l o i d , cyanophilous, and inner
w a l l p i n k i n C r e s y l B l u e . B a s i d i a 12.5-20 X 4.5-6.5 p m , 4-spored.
F i g . 121. Cystolepiota cystidiosa. L a m e l l a edge sterile, i n fresh specimens covered i n a y e l l o w exudate.
C h e i l o c y s t i d i a 17-40(-50) X 6.5-12 u m , cylindrical-fusoid, narrowly S E L . D E S C R . & FIGS. - H e r i n k i n C e s k a M y k o l . 15: 226-233, figs. 6-

clavate, rather variable i n shape, without o r w i t h apical excrescence, 8. 1961; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 36-37. 1994; K u h n e r
varying from a s m a l l capitulum, 3.0-4.0 X 3.0 p m , to a l o n g m o n i l i - i n B u l l , trimest. S o c . m y c o l . Fr. 52: 205-206. 1936; F. M 0 l l e r i n F r i e s i a
f o r m neck, u p to 4 5 p m l o n g ; o c c a s i o n a l l y w h o l e c y s t i d i u m m o n i l i - 6: 25. ('1957-1958') 1959; V e l l i n g a & Huijser i n Persoonia 16: 5 2 1 -
form; often t h i c k - w a l l e d , w i t h y e l l o w , often granular, contents. 524, figs. 2e & 3d. 1998.
Pleurocystidia abundant, s i m i l a r to cheilocystidia, but without o r w i t h V E R N . N A M E - V l e k k e n d e poederparasol.
s h o r t - m o n i l i f o r m neck; at a magnification o f 4 0 0 X c y s t i d i a clearly

v i s i b l e as y e l l o w dots, evenly distributed over l a m e l l a surface. Pileus Pileus 10-22(-30) m m , w h e n young hemispherical, expanding to plano-
covering a velar epithelium, made up o f elements (10-)20-70(-95) p m convex w i t h umbo, withXjyelar remnants at margin, at first white to
i n diam., globose to slightly subglobose, thin-walled o r slightly thick- whitish cream, rarely greyish, soon discolouring to orange-brown i n
w a l l e d , often w i t h slightly b r o w n i s h w a l l s ; i n fresh specimens w i t h patches or totally discoloured ( M u . 5 Y R 5/6-8), granulose to more or
very s m a l l y e l l o w droplets or particles. Stipitipellis a cutis made up o f less squamulose, when y o u n g often w i t h l o w pyramidal warts on w o o l -
c y l i n d r i c a l narrow hyphae, 1.5-5.0 p m i n diam.; below covered i n ele- ly background; warts at centre rather crowded. Lamellae, L = 35-60,1
ments as o n pileus, and often more irregular i n shape (pyriform, e l l i p - = 3, moderately crowded, free, segmentiform to subventricose, up to 3
soid etc.). C l a m p - c o n n e c t i o n s present i n a l l tissues. m m broad, white to cream coloured, w i t h age discolouring, especially at
H A B I T A T & D I S T R . - G r e g a r i o u s , sometimes i n b i g flocks, sapro- edge, to orange-brown or rarely to pale buff (7.5 Y R 8/4) when touched,
trophic and terrestrial i n deciduous woods o n nutrient-rich soils, and with white, concolorous even edge. Stipe 15-40 (50) X 2-4 m m , c y l i n -
i n greenhouses. K n o w n i n the Netherlands f r o m several localities i n drical, fistulose, w i t h o r without w o o l l y ring-like zone, whitish cream,
southern L i m b u r g . A u g . - O c t . i n the w i l d , throughout the year i n rarely greyish, at apex pruinose, below ring-like zone granulose to floc-
greenhouses. A l s o f o u n d i n several regions o f Germany, and i n N o r t h culose, often rather q u i c k l y discolouring orange-brown or red-brown,
America. especially when touched, w i t h white basal tomentum. Context whitish
i n pileus, i n stipe apex cream coloured, and orange-brown (7.5 Y R 4/6)
Cystolepiota cystidiosa is to be recognized b y the y e l l o w contents o f at base, white around cavity. S m e l l variably recorded, unpleasant sweet-
cystidia o n the sides and the edge o f the lamellae. It is easily confused 4sh-fungoid, suggestive o f the smell of Lepiota cristata, or not at a l l like
with C. hetieri, but that species lacks the y e l l o w contents i n the cystidia cristata. Taste not recorded. Spore print colour not k n o w n .
and is white, d i s c o l o u r i n g orange-red. Spores 4.0-6.0(-6.5) X 2.0-3.0(-3.5) p m , Q = (1.5-)1.6-2.1(-2.3),
F o r an extensive discussion o n s y n o n y m y see V e l l i n g a & Huijser Qav = 1.8-2.0, oblong, some (sub-)cylindrical, some slightly broad-
(in Persoonia 16: 520-521. 1998). ened at basal part, colourless, w i t h slightly thickened w a l l , often i n
tetrads, non-dextrinoid, n o n - a m y l o i d , w i t h cyanophilous w a l l ; inner
2. Cystolepiota hetieri (Boud.) S i n g , i n B e i h . S y d o w i a 7: 67. 1973. - w a l l p i n k i n C r e s y l B l u e . B a s i d i a 15-25 X 5.0-7.0 u m , 4-spored.
F i g . 122. C h e i l o c y s t i d i a usually abundant, 14-32 X 7.0-12 u m , w i t h up to 15 p m
Lepiota hetieri B o u d . i n B u l l . S o c . m y c o l . F r . 18: 137. 1902; l o n g abrupt, capitulate a n d c y l i n d r i c a l or m o n i l i f o r m excrescent at
Cystoderma hetieri (Boud.) S i n g , i n S c h w e i z . Z . P i l z k . 17: 53. 1939. apex, clavate o r e l l i p s o i d , t h i n - w a l l e d to slightly t h i c k - w a l l e d espe-
-Agaricus granulosus var. rufescens B . & B r . i n A n n . M a g . nat. H i s t . c i a l l y i n apical part, usually colourless, some w i t h b r o w n i s h granular
Ser. V , 7: 124. 1881 ( N o t i c . B r i t . F u n g i 1834); Lepiota rufescens ( B . & contents. P l e u r o c y s t i d i a present, especially near l a m e l l a edge, some-
Br.) J . L a n g e i n D a n s k bot. A r k . 9 (6): 6 5 . 1 9 3 8 , non Lepiota rufescens times rather rare, s i m i l a r to cheilocystidia. H y m e n i u m at l a m e l l a edge
M o r g a n , 1906; Lepiota langei L o c q . i n B u l l . mens. Soc. l i n n . L y o n 14: b r o w n coloured. Pileus covered w i t h a velar epithelium, made u p o f
87. 1945, n o n Lepiota langei K n u d s e n , 1981; Cystolepiota langei globose to ellipsoid and spheropedunculate elements, 20-60(-80) u m i n
( L o c q . ) M . B o n i n D o c . m y c o l . 2 2 (88): 27. 1993. diameter, slightly t h i c k - w a l l e d , w i t h , rarely without, b r o w n i s h parietal
E X C L . - Lepiota hetieri sensu J . L a n g e , F l . agar. dan. 1: 35-36, p i . pigment. Stipitipellis at apex a cutis o f c y l i n d r i c a l elements, 4-10 u m
14J. 1935 ( = C . adulterina); Cystolepiota hetieri sensu Breitenb. & in diam., usually colourless, sometimes red-brown coloured, slightly
K r a n z l . , P i l z e S c h w e i z 4: p i . 210. 1995 ( = C. adulterina or C. cys- thick-walled, lower d o w n w i t h chains o f globose elements, 20-60 p m
tidiosa). - Lepiota rufescens sensu H u i j s m a n i n M e d e d . N e d . m y c o l . in d i a m . , slightly t h i c k - w a l l e d a n d b r o w n c o l o u r e d . C l a m p - c o n n e c -
Vereen. 28: 46-51. 1943 ( = C. cystidiosa); sensu K u h n e r & R o m a g n . , tions present i n a l l tissues.
F l . anal. C h a m p , sup.: 396. 1953 ( = Cystolepiota spec). H A B I T A T & DISTR. - Saprotrophic a n d terrestrial, gregarious i n
S E L . I C O N . - B o u d . i n B u l l . S o c . m y c o l . Fr. 18: p i . 6, fig. 1. 1902; deciduous woods o n clayey, l o a m y soils, r i c h i n humus, often r i c h i n
J. L a n g e , F l . agar. dan. 1: p i . 141. 1935; R a i d et al. i n Svampe 26: 34. l i m e ; also i n greenhouses. In the Netherlands widespread, but not c o m -
1992; Z e c c h i n i n R i v . M i c o l . 4 3 : 156. 2000. mon. O c c u r r i n g from the end o f A u g . - O c t . ( - N o v . ) , but i n greenhouses
throughout the year. K n o w n f r o m temperate regions o f Europe.
Cystolepiota hetieri c a n be confused w i t h C. pulverulenta. T h e latter

is characterized b y the volatile, flocculose v e i l , a n d the o b l o n g ele-
ments o f w h i c h i t i s made up, the absence o f cystidia and c l a m p - c o n -
nections, and the slightly rough spores.
F o r the differences w i t h respect to C. adulterina, see the notes
accompanying the description o f that species.
Cystolepiota cystidiosa differs i n the presence o f y e l l o w contents o f
the cystidia, and the abundance o f pleurocystidia.
Three Cystolepiota-species have been tested for the presence o f
amanitins: C. seminuda, C. bucknallii a n d C. hetieri (as Lepiota
rufescens) (Gerault & G i r r e i n C . r. hebd. Seanc. A c a d . S c i . , Paris, Ser.
F i g . 122. Cystolepiota hetieri. D , 280: 2842. 1966). O n l y the last was found to be t o x i c .
3. Cystolepiota adulterina (F. M 0 l l e r ) M . B o n i n D o c . m y c o l . 7 (27-

28): 54. 1977.
Lepiota adulterina F. M 0 l l e r i n F r i e s i a 6: 23. ('1957-1958') 1959.
- Cystolepiota adulterina f. reidii M . B o n i n D o c . m y c o l . 11 (43): 2 5 .
1981; Cystolepiota adulterina var. reidii ( M . B o n ) M . B o n i n D o c .
m y c o l . 22 (88): 27. 1993. - Cystolepiota subadulterina M . B o n in Doc.
m y c o l . 6 (24): 4 3 . 1976; Cystolepiota adulterina var. subadulterina
( M . B o n ) M . B o n i n D o c . m y c o l . 2 2 (88): 27. 1993.
M I S A P P L . - Cystolepiota hetieri sensu J . L a n g e , F l . agar. dan. 1: 35-
36, p i . 14J. 1935.
E X C L . - Cystolepiota adulterina sensu K e l d e r m a n i n C o o l i a 31: 15.
1988 ( = C. cystidiosa). - Cystolepiota adulterina f. reidii sensu M .
B o n i n D o c . m y c o l . 11 (43): 2 5 . 1981; sensu L a n z o n i & Z e c c h i n i n F i g . 123. Cystolepiota moelleri.
R i v . M i c o l . 31 (3-4): 104. 1988; sensu Candusso & L a n z o n i , L e p i o t a :
88-90. 1990 (in a l l cases C. moelleri). M I S A P P L . - Cystolepiota adulterina f. reidii sensu M . B o n i n D o c .
S E L . I C O N . - J . Lange, F l . agar. dan. 1: p i . 14J. 1935 (as C. hetieri); m y c o l . 11 (43): 25. 1981; sensu L a n z o n i & Z e c c h i n i n R i v . M i c o l . 31
R a i d et a l . i n Svampe 26: 34. 1992 (dry specimens); R y m a n & (3-4): 104. 1988; sensu Candusso & L a n z o n i , L e p i o t a : 88-90. 1990;
Holmasen, Svampar: 4 1 1 . 1 9 8 4 ; Z e c c h i n i n R i v . M i c o l . 4 3 : 1 5 9 0 . 2000. Lepiota pseudoasperula sensu Enderle & Krieglst. i n Z . M y k o l . 55: 86.
S E L . D E S C R . & FIGS. - F. M 0 l l e r i n F r i e s i a 6: 22-23, figs. a-d. ('1957- 1989.
1958') 1959; D . R e i d i n F u n g . rar. Ic. c o l . 6: 10-11, figs. 18a & b . 1972; S E L . I C O N . - Enderle & Krieglst. i n Z . M y k o l . 55: opp. p . 96. 1989
V e l l i n g a & H u i j s e r i n Persoonia 16: 515-518, figs. 2 a & 3a. 1998; (as Lepiota pseudoasperula); R . P h i l l i p s , Paddest. S c h i m m . : 30. 1981
Z e c c h i n i n R i v . M i c o l . 4 3 : 148-160. 2000. (as Lepiota rosea); Z e c c h i n i n R i v . M i c o l . 4 3 : 153 & 155. 2000 (as C.
rosea).
C H A R A C T E R I S T I C S - P i l e u s 15-30(-50) m m , conico-convex, applanate S E L . D E S C R . & FIGS. - K e l d e r m a n i n C o o l i a 31: 12-14, fig. 1. 1988;
to convex, w h e n y o u n g covered w i t h a thick cream whitish, to beige P.D. Orton i n Trans. B r . m y c o l . S o c . 43: 287. 1960 (as Lepiota rosea);
layer, later w i t h cream-ochre to greyish y e l l o w flocculose warts; mar- V e l l i n g a & Huijser i n P e r s o o n i a 16: 524-525, figs. 2 f & 3e. 1998;
g i n w i t h velar remnants. L a m e l l a e crowded, free, sordid cream. Stipe Z e c c h i n i n Riv. M i c o l . 4 3 : 153-155. 2000 (as C. rosea).
30-60 X 3-7 m m , c y l i n d r i c a l or tapering downwards, fistulose, at apex V E R N . N A M E - R o z e poederparasol.
whitish, lower d o w n covered w i t h floccules, concolorous w i t h pileus,

b r o w n d i s c o l o u r i n g at base; s m e l l and taste not k n o w n . Pileus 8-40 m m , w h e n y o u n g hemispherical w i t h i n f l e x e d margin,
Spores 5.0-6.5(-7.0) X 2.0-3.0 p m , Q = (1.8-)2.0-2.6, Q a v = 2.1- expanding to plano-convex w i t h shallow central depression, o r w i t h
2.4, c y l i n d r i c a l to slightly broadened i n basal part, non-dextrinoid, w i t h applanate centre, at centre densely set w i t h l o w acute and (sub-)pyram-
p i n k inner w a l l i n C r e s y l B l u e ; basidia 14.5-23 X 5.0-9.0 p m , (2-)4- idal p i n k or p i n k i s h red-brown squamules or warts ( M u . 2.5 Y R 3/5, 3-
spored; c h e i l o c y s t i d i a abundant, 17-32 X 5.5-9.0 u m , lageniform w i t h 2.5/2, 2.5-5 Y R 4/4), at margin w i t h more widespread squamules, o n a
abrupt, c y l i n d r i c a l to slightly m o n i l i f o r m , flexuous, 1.5-4.0 p m wide very pale pinkish-brownish background (7.5 Y R 7/6), w i t h velar rem-
neck, rarely without neck or o n l y w i t h short excrescence, colourless.; nants at margin when y o u n g . L a m e l l a e , L = 30-40, 1 = 1-3, moder-
pleurocystidia absent; pileus covering a velar epithelium made up o f ately crowded to crowded, free, segmentiform to ventricose, up to 3.5
globose elements, 13-46 p m i n d i a m . , t h i c k - w a l l e d a n d slightly m m wide, w h i t i s h cream, greyish cream w i t h age, w i t h concolorous to
encrusted, colourless; clamp-connections present i n a l l tissues. p i n k i s h even to flocculose (lens!) edge. Stipe 15-40(-70) X 2-5 m m ,
H A B I T A T & DISTR. - I n s m a l l groups, saprotrophic and terrestrial, cylindrical or slightly broadening towards base, curved at base, fistu-
often o n m u l l , i n deciduous woods, often o n calcareous soils. S e p t - lose, at apex p i n k i s h cream a n d pruinose at fibrillose background,
lower d o w n w i t h scattered p i n k o r reddish b r o w n flocculose o r lanate
Oct. N o t yet k n o w n from the Netherlands, rare and scattered i n Europe,
bands o f warts (2.5 Y R 6/4) o n pale p i n k to vinaceous red background.
from southern S c a n d i n a v i a southwards, not yet recorded from
Context i n pileus white to cream, white to pale p i n k i s h i n stipe. S m e l l
Mediterranean countries.
faint and sweetish, pleasant o r l i k e Lepiota cristata. Taste indistinct,
fungoid. Spore print ' w h i t e ' .
Cystolepiota adulterina i s often confused w i t h C. hetieri, but easily
distinguished from that species b y the narrower and longer spores, the Spores 4.0-5.5(-6.0) X 2.5-3.0 p m , Q = (1.5-)1.6-2.2(-2.3), Q a v =
absence o f pleurocystidia and the different shape o f the cheilocystidia 1.7-1.9(-2.0), oblong to c y l i n d r i c a l , some slightly phaseoliform i n side-
(see M 0 l l e r i n F r i e s i a 6: 20-25. ('1957-1958') 1959; and V e l l i n g a & view, thin-walled and colourless, non-dextrinoid, w i t h p i n k inner w a l l
Huijser i n Persoonia 16: 516-518. 1998 for discussions o f the charac- in C r e s y l B l u e , often i n tetrads. B a s i d i a 16-26 X (3.5-)5.0-7.5 p m , 4-
ters). T h e latter authors also extensively treat the s y n o n y m y o f the spored. L a m e l l a edge sterile. C h e i l o c y s t i d i a abundant, 15-35(-39) X
species. 7.0-15 u m , n a r r o w l y clavate to obovate, u s u a l l y w i t h c y l i n d r i c a l ,
T h o u g h this species has been recorded f r o m adjacent regions i n m o n i l i f o r m o r branched, excrescence at apex, u p to 2 2 p m l o n g ,
G e r m a n y and B e l g i u m , and is also k n o w n to occur i n Great B r i t a i n , it colourless and thin-walled. Pleurocystidia not present. Squamules o n
has not yet been found i n the Netherlands. pileus made u p o f globose to e l l i p s o i d elements, 15-70 u m i n diam.,
slightly thick-walled, w i t h b r o w n parietal pigment. Stipitipellis at apex
4. Cystolepiota moelleri K n u d s e n i n B o t . Tidsskr. 73: 134. 1978. - of stipe a cutis o f c y l i n d r i c a l to inflated, 4.0-8.0 p m w i d e , not coloured
elements, lower d o w n w i t h chains o f globose to e l l i p s o i d elements, 30-
F i g . 123.
60 p m i n diam., brown-coloured and slightly thick-walled. C l a m p - c o n -
Lepiota rosea R e a i n Trans. B r . m y c o l . S o c . 6: 61-62. ('1917')
1918; Cystolepiota rosea (Rea) M . B o n i n D o c . m y c o l . 6 (24): 4 3 .
1976, non Cystolepiota rosea Sing., 1969; Cystolepiota rosella M o s . , H A B I T A T & DISTR. - Gregarious or solitary, saprotrophic and terres-
R o h r l i n g e Blatterpilze, 4. A u f l . : 236. 1978 (superfluous). trial i n m i x e d deciduous woods o n loamy, calcareous o r nutrient-rich
soils; i n the Netherlands rare, mostly i n southern L i m b u r g , A u g . - O c t . c y l i n d r i c a l , fistulose, cream to pale lemon-yellow, w i t h age and when
Widespread and rather rare i n temperate parts o f Europe. touched i n lower half and especially at base purplish or vinaceous pink,
very rarely not d i s c o l o u r i n g at a l l , minutely pubescent at apex, when
Bon ( F l . m y c o l . E u r . 3, L e p i o t e s : 4 0 . 1993) cited this taxon as y o u n g w h i t i s h pruinose, glabrescent w i t h age. Context thin, concolor-
Cystolepiota rosea (Rea) S i n g . However, Singer ( i n B e i h . N o v a ous w i t h surfaces. S m e l l not distinct to fruity-fungoid. Taste indistinct.
Hedwigia 2 9 : 168. 1969) v a l i d l y described the n e w species Spore print white.
Cystolepiota rosea, a species i n its o w n right, differing from C. moel- Spores 3.5-5.0(-5.5) X 2.0-3.0 p m , Q = 1.35-2.1(-2.2), Q a v =
leri i n the absence o f cheilocystidia and i n the rather smooth surface 1.45-1.95, e l l i p s o i d to c y l i n d r i c a l , rather thin-walled, w i t h distinct or
of the pileus. indistinct hilar appendage, non-dextrinoid, w i t h inner w a l l p i n k i n
Cystolepiota moelleri macroscopically resembles Lepiota C r e s y l B l u e , s l o w l y p i n k i n C o n g o R e d . B a s i d i a 11-19 X 4.5-6.5 p m ,
pseudoasperula, but i s m i c r o s c o p i c a l l y quite distinct, because o f the 4-spored. C h e i l o - a n d p l e u r o c y s t i d i a absent. P i l e u s covered w i t h a
non-dextrinoid spores and the clavate cheilocystidia that are often pro- velar epithelium, made up o f a layer o f rather thin-walled to slightly
v i d e d w i t h an apical excrescence. thick-walled globose, spheropedunculate o r e l l i p s o i d elements, 15-40
p m i n diameter; inner elements thin-walled and compressed, colour-
5. Cystolepiota seminuda (Lasch) M . B o n i n D o c . m y c o l . 6 (24): 43. less; outer elements w i t h encrusting, colourless pigment. Stipitipellis
1976. - F i g . 124. a cutis made up o f c y l i n d r i c a l , 4.0-10 p m w i d e hyphae w i t h some glo-
Agaricus seminudus L a s c h i n L i n n a e a 3: 157. 1828; Lepiota semi- bose elements as o n pileus. Clamp-connections present.
nuda (Lasch) K u m m . , Fiihr. P i l z k . : 1 3 6 . 1 8 7 1 ; Cystoderma seminudum H A B I T A T & D I S T R . - Gregarious, rarely solitary, saprotrophic and
(Lasch) Fay. i n A n n l s S c i . nat., B o t . , Ser. V I I , 9: 351. 1889; Lepiota terrestrial o n soils r i c h i n humus a n d nutrients, i n deciduous a n d i n
sistrata var. seminuda (Lasch) Q u e l , i n M e m . Soc. E m u l . M o n t b e l i a r d , coniferous woods, not u n c o m m o n , A u g . - O c t . K n o w n from temperate
Ser. II, 5: 2 3 1 . 1872 ( C h a m p . Jura Vosges 1). - Lepiota sororia regions o f the Northern Hemisphere.
H u i j s m a n i n Persoonia 1: 326. 1960; Cystolepiota sororia (Huijsman)
S i n g , i n B e i h . S y d o w i a 7: 67. 1973. - Lepiota seminuda f. minima J . Cystolepiota seminuda is very variable i n size and colours o f the basid-
L a n g e , F l . agar. dan. 1: 36. 1935 (not valid); Lepiota sistrata f. mini- iocarps. T h e spores m a y be smooth, or r o u g h as observed by Scanning
ma (J. L a n g e ) B a b o s i n A n n l s hist.-nat. M u s . natn. hung. 50: 91. 1958. «Clectron M i c r o s c o p e (see Keller, A t l . B a s i d i o m y c : p i . 284. 1997 (as
M I S A P P L . - Lepiota sistrata and Cystolepiota sistrata sensu auct. eur. C. sistrata) for a picture o f rough spores).
E X C L . - Lepiota sistrata sensu H e r i n k i n C e s k a M y k o l . 15: 217. T h e name C. sistrata (Fr.: Fr.) Sing, ex B o n & B e l l u has been used
1961 ( = C . pulverulenta). for this species, but is here considered a nomen d u b i u m , as Fries (Syst.
S E L . I C O N . - Breitenb. & K r a n z l . , P i l z e S c h w e i z 4: p i . 211. 1995; m y c o l . 1: 24. 1821) described a rather b i g species, and various inter-
Candusso & L a n z o n i , L e p i o t a : p i . 2a. 1990 (as C. sistrata); Dahncke, pretations o f this name exist i n literature (see V e l l i n g a i n Persoonia 13:
1200 P i l z e : 512. 1993 (as C. sistrata); J . L a n g e , F l . agar. dan. 1: p i . 323. 1987).
14A, 1 4 B . 1935; L a n z o n i & Candusso i n B o l l . G r u p p o m i c o l . G . Bres.
2 6 : 1 1 6 . 1983 (as C. sistrata); R . P h i l l i p s , Paddest. S c h i m m . : 3 0 . 1 9 8 1 ;
Z e c c h i n i n R i v . M i c o l . 4 3 : 152. 2000.
S E L . D E S C R . & F I G S . - A t k . i n M e m . N . Y . bot. G d n 6: 2 1 6 - 2 2 1 , p i . Sect. Pseudoamyloideae Sing. & Clem.
22, 25 & 26. 1916 (development); B a b o s i n A n n l s hist.-nat. M u s . natn.
hung. 5 0 : 9 1 . 1958 (as Lepiota sistrata f. minima); Candusso & Spores dextrinoid; clamp-connections present; velar elements globose.
L a n z o n i , L e p i o t a : 80-81, f i g . 7. 1990; H u i j s m a n i n Persoonia 1: 326-
327. 1960 (as Lepiota sororia); K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 6. Cystolepiota bucknallii ( B . & Br.) Sing. & Clem, i n Nova
32-33. 1994; V e l l i n g a i n Persoonia 13: 321-325, figs. 1-4. 1987; H e d w i g i a 23: 338. ( ' 1 9 7 2 ' ) 1973. - F i g . 125.
Z e c c h i n i n R i v . M i c o l . 43: 150-151. 2000. Agaricus bucknallii B . & B r . i n A n n . M a g . nat. H i s t . Ser. V , 7: 124.
V E R N . N A M E - K l e i n e poederparasol. 1881 (as A . bucknalli) ( N o t i c . B r . F u n g i 1836); Lepiota bucknallii (B.
& Br.) S a c c , S y l l . F u n g . 5: 50. 1887. - Lepiota seminuda var. lilacina
Pileus 3-20(-30) m m , hemispherical o r obtusely c o n i c a l w h e n y o u n g , Q u e l , i n B u l l . S o c . bot. Fr. 23: 325. 1877 ( C h a m p . Jura Vosges S u p p l .
w i t h inflexed m a r g i n and w i t h velar remnants attached, expanding to 4); Lepiota lilacina (Quel.) B o u d . i n B u l l . Soc. m y c o l . Fr. 9: 6. 1893;
plano-convex w i t h or without l o w umbo, white, white w i t h cream to Cystolepiota bucknallii var. lilacina (Quel.) M . B o n i n D o c . m y c o l . 7
y e l l o w i s h tinge at centre o r White w i t h p i n k i s h tinge at centre, rarely (27-28): 21. 1977 (not valid).
w i t h b r o w n i s h centre, w h e n y o u n g w i t h densely floccose-verrucose S E L . I C O N . - Breitenb. & K r a n z l , P i l z e S c h w e i z 4: p i . 209. 1995;
covering, later o n granulose to pruinose-farinose o r even glabrous Candusso & L a n z o n i , L e p i o t a : p i . 6b. 1990; B . H a n f f i n P i l z f l .
when very o l d or after heavy rain. L a m e l l a e , L = 30-40,1 = 1-3, rather Nordwestoberfrankens 1-4: p i . 8 (023). 1982; K u h n e r i n B u l l , trimest.
crowded, free or nearly free, ventricose to subventricose, 2-3 m m wide, Soc. m y c o l . Fr. 52: A t l a s p i . 72, fig. 2. 1936; Lavorato i n S c h w e i z . Z .
white, yellowish-creamy, o r w i t h pale l e m o n - y e l l o w tinge, w i t h even P i l z k . 67: 207. 1989; R . P h i l l i p s , Paddest. S c h i m m . : 30. 1981; R a i d et
to finely flocculose, concolorous edge. Stipe 15-50(-70) X 1-3 m m , al. i n Svampe 26: 34. 1992; Z e c c h i n i n R i v . M i c o l . 4 3 : 161. 2000.
15. 1990; K e l d e r m a n , Parasolzw. Z u i d - L i m b u r g : 30-31. 1994; K u h n e r
i n B u l l , trimest. S o c . m y c o l . F r . 5 2 : 204-205. 1936; L a v o r a t o i n
S c h w e i z . Z . P i l z k . 6 7 : 204-208. 1989; S i n g . & C l e m , i n N o v a
H e d w i g i a 23: 338-339. ('1972') 1973.
V E R N . N A M E - V i o l e t s t e l i g e poederparasol.
OQQQ@D©00 Pileus 8-25 m m , w h e n y o u n g hemispherical or bluntly c o n i c a l w i t h

F i g . 124. Cystolepiota seminuda. inflexed margin, expanding to (conico-)convex, w i t h broad u m b o o r
S E L . D E S C R . & FIGS. - K n u d s e n i n B o t . Tidsskr. 73: 131, figs. 3-4,

13 (upper left row). 1978; L o n a t i i n B o l l . A s s . m i c o l . ecol. R o m a n a 12:
12-14. ('1987') 1988.
C H A R A C T E R I S T I C S - Pileus 15-30 m m , obtusely c o n i c a l to convex w i t h

low, broad umbo, w i t h undulating marginal zone, sulphur-yellow at
first, pallescent to y e l l o w i s h o r pale buff, or b r o w n i s h yellow, finely
floccose-farinose, almost glabrous at margin; lamellae rather crowded,
free, cream-coloured; stipe 30-55 X 2-3 m m , c y l i n d r i c a l , concolorous
with pileus, brownish at base, floccose-farinaceous at first, glabrescent
with age; context thin and concolorous w i t h surfaces; s m e l l and taste
not distinct; spore print pale cream.
Spores 3 . 5 - 5 . 5 ( ^ 0 ) X 2.5-3.0 u m , o b l o n g , o v o i d , dextrinoid;
basidia 4-spored; cheilocystidia scattered, 30-45 X 5.0-8.0 u m , irreg-
ularly sinuous, narrowly clavate or fusiform; p l e u r o c y s t i d i a absent;
elements o f velar covering globose to subglobose, 20-30(-40) p m i n
diam., w i t h y e l l o w i s h vacuolar pigment; clamp-connections present i n
F i g . 125. Cystolepiota bucknallii. all tissues.
deciduous woods. N o t w i t h certainty k n o w n f r o m the Netherlands,
broadly campanulate, cream to white at m a r g i n , usually w i t h v i o l a - probably found once i n southern L i m b u r g ; very rare i n Europe, only
ceous o r l i l a c tinge (e.g. K . & W . 15D4, 14D3), when o l d w i t h ochra- k n o w n from a few finds i n D e n m a r k , S w e d e n , B e l g i u m and Italy.
ceous o r y e l l o w i s h tinge at centre, completely finely micaceous
granulose, w h e n y o u n g w i t h some velar remnants at margin. L a m e l l a e , Unfortunately no material o f the Netherlands find o f Cystolepiota icte-
L = 25-40, 1 = ( l - ) 3 - 5 , moderately distant to moderately crowded, rina has been conserved; the characteristics above are c o m p i l e d from
free, ventricose, white to cream c o l o u r e d w i t h age, w i t h white even the descriptions cited.
edge. Stipe 22-80 X 1.5-4 m m , c y l i n d r i c a l o r broadening upwards,
sometimes w i t h subbulbous base, fistulose, white at apex, darkening
to violet, deep violaceous o r discoloured to dark b r o w n at base, (14F4,
1 5 D 4 - F 5 ) , pruinose to flocculose at apex, finely granulose to glabrous Sect. Pulverolepiota ( M . Bon) Vellinga
l o w e r d o w n , w h e n y o u n g w i t h some floccules o f annulus. Context
white i n pileus; i n stipe concolorous w i t h surface, but sometimes dark- Spores very s l o w l y b e c o m i n g red-brown i n M e l z e r ' s Reagent o r not
er i n base. S m e l l strong, l i k e indole (coal gas, l i k e Tricholoma sul- c o l o u r i n g at a l l , rough, and binucleate; velar elements elongate and
phureum). Taste never tried. Sporeprint white. inflated.
Spores i n side v i e w (6.5-)7.0-9.0(-10.0) X 2.5-3.5(-4.0) p m , Q =
(2.3-)2.4-3.1(-3.3), Q a v = 2.6-2.95, c y l i n d r i c a l , for the greater part 8. Cystolepiota pulverulenta (Huijsman) V e l l i n g a i n Persoonia 14:
w i t h truncate base, some even slightly spurred, not coloured, slightly 407. 1992. - F i g . 126.
thick-walled, slowly c o l o u r i n g pale orange-brown i n Melzer's Lepiota pulverulenta H u i j s m a n i n Persoonia 1: 3 2 8 . 1960;
Reagent, p i n k i n C o n g o R e d , often i n tetrads; inner w a l l p i n k i n Leucoagaricuspulverulentus (Huijsman) M . B o n i n D o c . m y c o l . 8 (30-
C r e s y l B l u e . B a s i d i a 18-35 X 6-8 p m , 4-spored. C h e i l o - and pleuro- 31): 70. 1978; Leucoagaricus pulverulentus (Huijsman) M o s . ,
cystidia absent. P i l e u s c o v e r i n g a loose velar e p i t h e l i u m made up o f Rohrlinge Blatterpilze, 4. Aufl.: 246. 1978 (superfluous);
globose to e l l i p s o i d , some spheropedunculate elements, 10-35 u m i n Pulverolepiota pulverulenta (Huijsman) M . B o n i n D o c . m y c o l . 2 2
d i a m . , colourless, some w i t h diffuse very pale b r o w n intracellular (88): 30. 1993. - Leucoagaricus pulverulentus f. minimus M . B o n et al.
pigment, slightly t h i c k - w a l l e d , w i t h some incrustations (not evanes- in D o c . m y c o l . 19 (75): 54. 1989; Pulverolepiota pulverulenta f. mini-
cent i n h y d r o c h l o r i c acid). S t i p i t i p e l l i s a cutis w i t h loosely arranged ma ( M . B o n et al.) M . B o n i n D o c . m y c o l . 22 (88): 30. 1993;
or attached globose and spheropedunculate velar elements, u p to 2 0 Cystolepiota pulverulenta f. minima ( M . B o n et al.) C h i u s a i n R i v .
p m i n d i a m . , l i k e those o n pileus. C l a m p - c o n n e c t i o n s present, very M i c o l . 4 1 : 152. 1998. - Lepiota hetieriana L o c q . i n B u l l . mens. Soc.
o c c a s i o n a l l y absent. l i n n . L y o n 14: 94. 1945 (not valid); Cystolepiota hetieriana (Locq.)
H A B I T A T & D I S T R . - Gregarious, rarely solitary, saprotrophic and Bresinsky i n Bresinsky & Haas i n B e i h . Z . P i l z k . 1: 46. 1976 (not valid).
terrestrial i n deciduous woods o f Fagus, Corylus, Carpinus on cal- M I S A P P L . - Lepiota sistrata sensu H e r i n k i n C e s k a M y k o l . 1 5 : 2 1 7 .
careous l o a m y soils r i c h i n nutrients. I n the Netherlands m o s t l y i n 1961.
southern L i m b u r g and k n o w n from a very few places i n the Pleistocene S E L . I C O N . - Partacini i n R i v . M i c o l . 30 (3-4): front cover. 1987; D .
areas (Dorst, and i n the province o f Drenthe), w h i c h are l o c a l l y r i c h i n R e i d i n F u n g . rar. Ic. c o l . 2: p i . 9e, 9f. 1967; V e l l i n g a & Huijser i n
l o a m , o r enriched w i t h shells. A u g . - O c t . I n temperate regions o f C o o l i a 40: p i . 1. 1997.
Europe, and also k n o w n from N o r t h A m e r i c a . S E L . D E S C R . & FIGS. - H u i j s m a n i n P e r s o o n i a 1: 328-329. 1960;
Kelderman, Parasolzw. Z u i d - L i m b u r g : 34-35. 1994; Partacini i n R i v .
7. Cystolepiota icterina K n u d s e n i n B o t . Tidsskr. 7 3 : 130. 1978. M i c o l . 30: 132-133. 1987; D . R e i d i n F u n g . rar. Ic. c o l . 2: 6-7. 1967;
Lepiota icterina F. M 0 l l e r i n F r i e s i a 7: 453. ('1965') 1967 (nom. V e l l i n g a i n Persoonia 14: 407-408, fig. 1. 1992.
nud.). VERN. NAME - Kegelpoederparasol.
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . 6 1 . 1990; L o n a t i i n

B o l l . A s s . m i c o l . e c o l . R o m a n a 12: 14. ( ' 1 9 8 7 ' ) 1988; R y m a n & Pileus 11-50 m m , when y o u n g c o n i c a l w i t h rounded apex or conico-
H o l m a s e n , Svampar: 412. 1984. convex, w i t h inflexed m a r g i n , expanding to plano-convex o r piano-
uous, fistulose, at apex cream-coloured and glabrous to slightly pru-

inose; rest o f stipe covered w i t h pale cream f l o c c i , l i k e those o n pileus,
and d i s c o l o u r i n g l i k e f l o c c i on pileus. Context w h i t i s h or creamy i n
pileus and stipe, and d i s c o l o u r i n g to orange-brown, especially i n base
o f stipe, when disposed to air. S m e l l indistinct, unpleasant. Taste not
k n o w n . Spore print white.
Spores i n side-view (3.5-)4.0-5.5(-6.0) X (2.0-)2.5-3.5 p m , Q =
(1.3-)1.4-2.25(-2.35), Q a v = 1.55-2.0, e l l i p s o i d to oblong w i t h paral-
l e l to convex sides, i n frontal v i e w o b l o n g , often i n tetrads, w i t h slight-
ly rough w a l l , s l o w l y or not at a l l dextrinoid, w i t h p i n k inner w a l l i n
C r e s y l B l u e , binucleate. B a s i d i a 13-25 X 4.0-7.5 p m , 4-spored.
C h e i l o - and pleurocystidia absent. Elements o f velar, epithelial cover-
ing o n pileus and stipe 60-110 X 9.0-15 p m , elongate, inflated w i t h
rounded apices, sometimes branched, catenulate, w i t h encrusting
colourless or pale b r o w n i s h substance. Stipitipellis a cutis o f c y l i n d r i -
cal, 4.0-6.0 p m w i d e , hyphae, w i t h y e l l o w , not encrusting pigment.
Clamp-connections absent.
H A B I T A T & DISTR. - In s m a l l groups, saprotrophic and terrestrial i n
deciduous woods o n rather nutrient-rich, l o a m y and moist soils. In the
Netherlands k n o w n from several localities i n the province o f L i m b u r g
(Cadier en Keer, Heerlen, L i n n e , and Valkenburg), very rare elsewhere
(Wassenaar). A u g . - O c t . T h i s species m i g h t reach the northern l i m i t o f
itsÊuropean distribution area i n the Netherlands and Germany.
T n e reasons to i n c l u d e this species i n Cystolepiota are given b y

V e l l i n g a (in Persoonia 14: 4 0 9 - 4 1 0 . 1992), and are supported b y
molecular data.
F i g . 126. Cystolepiota pulverulenta.
Cystolepiota pulverulenta has, i n contrast to the species i n sections
Cystolepiota and Pseudoamyloideae, binucleate spores, and no c l a m p -
concave, w i t h or without obtuse u m b o or papilla, w i t h straight margin, connections.
w h i t i s h or cream-coloured at first, later o n and when touched p i n k i s h Cystolepiota pseudogranulosa ( B . & Br.) Pegl., a tropical species
b r o w n ( M u . 7.5 Y R 5/6), p i n k - b r o w n (7.5 Y R 7/6) or rusty b r o w n to k n o w n from S r i L a n k a , East A f r i c a , Central and S o u t h A m e r i c a comes
dark hazel-brown (7.5 Y R 4/4-6/6, 10 Y R 4/4), covered w i t h a thick, very close. T h e pileus covering o f that species is also made up o f inflat-
loose, flocculose layer, u n i f o r m l y thick or w i t h warts, w i t h overhang- ed, oblong elements and the spores are rough as w e l l , but the hyphae
ing floccules or w i t h a r i m at margin. L a m e l l a e , L = 30-35,1 = 1-7, are provided w i t h numerous, though small, clamp-connections (Pegler,
moderately c r o w d e d to crowded, free and remote from stipe, thin, pers. c o m m . , Febr. 1996), and the spores are said to be strongly dex-
slightly ventricose, 2-4.5 m m broad, cream-coloured, pale cream to trinoid (Dennis i n K e w B u l l . 7: 486. 1952), though personal observa-
cream-yellow, sometimes slightly p i n k i s h , w i t h even, concolorous or tions contradict this.
paler edge. Stipe 30-65 X 2-5.5 m m , c y l i n d r i c a l or slightly tapering T h e spores o f C. pulverulenta, as seen w i t h S c a n n i n g E l e c t r o n
towards base, sometime^ w i t h up to 8 m m wide, subbulbous base, flex- M i c r o s c o p e , have discrete, rounded warts.
10. Melanophyllum Velen.
ELSE C. VELLINGA
Agaricus subgen. Melanophyllum (Velen.) Pilat in Acta Mus. natn. Prag. 7B (1): 26. 1951; Melanophyllum Velen.,
Ceske Houby: 569. 1921. - Chlorospora Mass. in Kew Bull. 1898: 136. 1898, non Chlorospora Speg., 1891;
Glaucospora Rea, Brit. Basidiomyc: 62. 1922. - Chlorosperma Murrill in Mycologia 14: 96. 1922.
SELECTED LITERATURE - Candusso & Lanzoni, Lepiota: 68-76. 1990; Sing., Agar. mod. Taxon., Ed. 4: 491-492.1986.
Basidiocarp pluteoid with persistent velum universale, made up of globose elements; velum partiale an evanescent
annulus; lamellae free, brightly coloured; spore print when fresh green or blue-green, in one species on drying turning
into brown and finally to black.
Spores ellipsoid to oblong, finely verrucose, non-amyloid, non-dextrinoid, metachromatic in Cresyl Blue, and walls
not swelling in ammonia ( N H ) and acetic acid ( C H C O O H ) ; cheilocystidia present and inconspicuous or absent;
3(aq) 3 (aq)
hymenophoral trama subregular; clamp-connections present. Development of basidiocarp bivelangiocarpic and

pileostipitocarpic. - Type species: Melanophyllum canali Velen.
HABITAT & DISTRIBUTION - Saprotrophic and terrestrial; cosmopolitan.
The genus Melanophyllum is very closely related to Cystolepiota Sing., differing mainly in the pigmentation of the
spores. Because of the coloured spores it was often placed close to Agaricus L.: Fr. in tribus Agariceae (e.g. Singer,
Agar. mod. Taxon., Ed. 4: 482-483. 1986), but that is not supported by molecular, nor by morphological, data.
K E Y TO THE SPECIES
1. B a s i d i o c a r p dark b r o w n , d i s c o l o u r i n g b l a c k i n exsiccates; lamellae reddish coloured 1. M . haematospermum

1. B a s i d i o c a r p white to pale y e l l o w - b r o w n , not changing into black i n exsiccates; lamellae greenish blue 2. M . eyrei
1. Melanophyllum haematospermum ( B u l l . : Fr.) K r e i s e l i n R e p r i u m S E L . D E S C R . & FIGS. - H o r a k , S y n . G e n . A g a r . : 379-380. 1968 (as

nov. Spec. R e g n i veg. 95: 700. 1984. - F i g . 127. M. echinatum); Krieglst. i n Siidwestd. P i l z r u n d s c h a u 17 (2): 1-3. 1981
Agaricus haematospermus B u l l . , Herb. France: p i . 595. 1793; (as M. echinatum); L a z z a r i & B e l l u , A t l . iconogr.: 2 0 6 - 2 0 7 . 1 9 8 5 ; Pegl.
Agaricus haematospermus B u l l . : Fr., Syst. m y c o l . 1: 282. 1821; Lepiota & B r a n d i n M y c o l o g i s t 11: 180. 1997; Wasser, Tr. A g a r i c e a e Soviet
haematosperma ( B u l l : Fr.) Q u e l , i n B u l l . Soc. m y c o l . Fr. 9: 6. 1893; U n i o n : 11-13. 1989.
Psalliota haematosperma ( B u l l . : Fr.) L u n d . & Nannf. i n F u n g i exs. suec. V E R N . N A M E - Verkleurzwammetje.
no. 341. 1935. -Agaricus echinatus Fr.: Fr., Syst. m y c o l . 1: 286. 1821;
Psalliota echinata (Fr.: Fr.) K u m m . , Fiihr. Pilzk.: 7 3 . 1 8 7 1 ; Pratella echi-
nata (Fr.: Fr.) Gillet, Hymenomycetes: 5 6 5 . 1 8 7 4 ; Lepiota echinata (Fr.:
Fr.) Quel., Enchir. Fung.: 8.1886; Inocybe echinata (Fr.: Fr.) S a c c , S y l l .
F u n g . 5: 773. 1887; Naucoria echinata (Fr.: Fr.) Schroet. i n C o h n ,
K r y p t . - F l . Schlesien 1: 6 0 7 . 1 8 8 9 ; Cystoderma echinatum (Fr.: Fr.) Sing,
i n A n n l s M y c o l . 34: 338. 1936; Melanophyllum echinatum (Fr.: Fr.)
Sing, i n L i l l o a 22: 436. ('1949') 1951. - Melanophyllum canali Velen.,
C e s k e H o u b y 3: 5 6 9 . 1921. - Lepiota haematosperma f. gracilis J.
Lange, F l . agar. dan. 1: 37. 1935 (not valid).
EXCL. - Melanophyllum haematospermum sensu Mitchell &
B r e s i n s k y i n M y c o l o g i a 9 1 : 811-819. 1999 ( = Leucoagaricus spec).
S E L . I C O N . - B o u d . , Ic. m y c o l . 1: p i . 12. 1905 (as Lepiota
haematosperma); Candusso & Lanzoni, L e p i o t a : p i . l a . 1990;
Liitjeharms i n F l . batava 28: p i . 2203. 1934 (as Lepiota echinata);
K r i e g l s t . i n Siidwestd. P i l z r u n d s c h a u 17 (2): 2. 1981 (as M. echina-
tum); L a z z a r i & B e l l u , A t l . iconogr.: 2 0 7 . 1985; P e g l . & B r a n d i n
M y c o l o g i s t 11: 180. 1997. F i g . 127. Melanophyllum haematospermum.
161
Pileus 6 - 3 5 m m , at first convex or bluntly c o n i c a l , expanding to

applanate w i t h or without l o w broad umbo, at centre dark grey-brown
( M u . 10 Y R 3/2 - 4/3), towards margin paler b r o w n on pinkish-reddish
background, o n d r y i n g pallescent to pale p i n k - b r o w n , completely c o v -
ered w i t h granules i n heaps, the biggest at centre, after heavy rainfall
completely glabrous; m a r g i n w h e n very y o u n g w i t h triangular velar
remnants then flocculose-fimbriate, exceeding lamellae. L a m e l l a e , L
= 27-40, 1 = 1-7, moderately crowded to crowded, free, straight to
ventricose, up to 3.5 m m broad, y o u n g red then purplish p i n k to red-
b r o w n or vinaceous r e d (2.5 Y R 3/4 - 5 Y R 3/3) w i t h even and c o n -
colorous edge. Stipe 10-50 X 1-3.5 m m , cylindrical, fistulose w i t h age,
when y o u n g vinaceous p i n k and w i t h fugacious annulus, later reddish
or purple-brown (2.5 - 7.5 Y R 3/2) lengthwise fibrillose w i t h granu-
lose c o v e r i n g as o n pileus. Context i n pileus pale cream to p i n k i s h
white, i n stipe concolorous w i t h surface, i n inner parts paler. S m e l l F i g . 128. Melanophyllum eyrei.
strong, l i k e the s m e l l o f Lepiota cristata or L. aspera, earth-like. Taste
suggestive o f L. cristata. Spore print w h e n fresh green ( K . & W . 2 8 D 4 )
turning into vinaceous red, o n d r y i n g into black. S E L . D E S C R . & FIGS. - B a b o s i n A n n l s hist.-nat. M u s . natn. hung.
Exsiccate black.
50- 87. 1958; H o r a k , S y n . G e n . Agar.: 658-660. 1968 (as Glaucospora
Spores (4.5-)5.0-6.5(-7.0) X 2.5-4.0 p m i n side view, Q = (1.4- eyrei).
)1.5-2.0(-2.3), Q a v = 1.55-1.95, o b l o n g , a few c y l i n d r i c a l , slightly VERN. NAME - Groenplaatzwammetje.
thick-walled, pale greenish-brownish i n ammonia, slightly verrucose
all over, n o n - a m y l o i d , non-dextrinoid, w i t h p i n k inner w a l l i n C r e s y l Pileus 6.5-25 m m , p a r a b o l o i d o r conico-campanulate when y o u n g ,
B l u e . B a s i d i a 14-25 X 5.0-7.0 p m , 4-spored, some 2-spored. C y s t i d i a expanding to applanate, often w i t h l o w broad umbo, very young almost
not observed, but see notes. H y m e n o p h o r a l trama w i t h b r o w n parietal White o r cream, then y e l l o w - b r o w n or pale p i n k i s h b r o w n ( c . M u . 10
pigment. P i l e u s covered w i t h v e l u m universale, forming an epithelium Y R 8/4) at centre, pallescent towards margin (10 Y R 8/6), a n d there
made up o f mostly globose a n d some e l l i p s o i d o r spheropedunculate sometimes slightly b l u i s h , completely finely granulose (lens!), when
to p y r i f o r m elements, 10-55 p m i n d i a m . , t h i c k - w a l l e d w i t h b r o w n y o u n g w i t h triangular velar remnants at margin. L a m e l l a e , L = 25-30,
intracellular pigment; terminal elements also w i t h some encrusting 1 = l-3(-6), moderately crowded, free, ventricose, up to 3 m m broad,
pigment; elements i n heaps. S t i p i t i p e l l i s a cutis o f c y l i n d r i c a l thick- strikingly greyish blue-green or blue-green. Stipe 20-55 X 1-3 m m ,
w a l l e d hyphae, 4.0-20 p m wide, w i t h b r o w n intracellular and parietal c y l i n d r i c a l , fistulose, at apex cream and finely pubescent or more o r
pigment, w i t h some l o o s e l y attached globose velar elements. C l a m p - less glabrous, lower d o w n pale p i n k - b r o w n , concolorous w i t h pileus
connections present. and granulose, b r o w n i n g w h e n touched. Context white to cream i n
H A B I T A T & D I S T R . - Gregarious, saprotrophic a n d terrestrial i n pileus. S m e l l indistinct, slightly resembling the s m e l l o f Lepiota crista-
deciduous forests o n c l a y o r l o a m , o n compost heaps, i n gardens, ta. Taste not k n o w n . Spore print b l u i s h green.
parks, and hothouses. I n the Netherlands not u n c o m m o n , throughout Spores (3.5-)4.0-5.5(-6.0) X 2.5-3.5 p m , Q = (1.2-)1.4-1.8(-2.0),
the country (for a distribution map see N a u t a & V e l l i n g a , A t l . N e d e r l . Qav = 1.5-1.7, e l l i p s o i d , oblong, some more c y l i n d r i c a l , o r amygdal-
Paddest.: 205. 1995); ( J u l y - ) A u g . - O c t . ( - N o v ) , i n hothouses throughout i f o r m w i t h rounded apex, slightly t h i c k - w a l l e d , w i t h b l u i s h w a l l i n
the year. C o s m o p o l i t a n . a m m o n i a , verrucose, n o n - a m y l o i d , non-dextrinoid, cyanophilous,
metachromatic i n C r e s y l B l u e . B a s i d i a 17-23 X 5.0-7.0 p m , 4-spored
T h e use o f the epithet 'haematospermum' instead o f 'echinatum' has or m i x e d 2- and 4-spored. C y s t i d i a not observed. H y m e n o p h o r a l trama
been discussed b y L a n z o n i & B e l l u ( A t l . iconogr.: 207. 1985). Pegler not coloured. Pileus covered w i t h v e l u m universale, c o m p o s e d o f glo-
(in K e w B u l l . add. Series 12: 316. 1986) gave an extensive list o f syn- bose and spheropedunculate elements, 15-45(-55) u m , slightly thick-
onyms o f Melanophyllum haematospermum. walled, w i t h pale y e l l o w - b r o w n parietal pigment. Stipitipellis a cutis o f
C h e i l o c y s t i d i a were not observed i n the D u t c h material, though c y l i n d r i c a l 4.0-10 p m w i d e colourless hyphae covered w i t h elements
Bon ( i n D o c . m y c o l . 11 (43): 2 3 . 1981) a n d Pegler & B r a n d ( i n as o n pileus. Clamp-connections present i n a l l tissues.
M y c o l o g i s t 11: 180. 1997) stated them to be present and inconspicu- H A B I T A T & DISTR. - Solitary to gregarious, saprotrophic and terres-
ous, rare, and clavate. trial i n m i x e d deciduous forests ( w i t h Fagus) o n h u m u s - r i c h soil, r i c h
The spores are ornamented w i t h more or less isolated warts, as can in nutrients, probably also calcareous and loamy. U p to n o w k n o w n
be seen w i t h S c a n n i n g E l e c t r o n M i c r o s c o p e . from several localities i n the southern part o f the Netherlands
O n d r y i n g the c o l o u r o f the spore print changes from green, v i a (Valkenburg, Schaelsberg; C a d i e r en Keer, Riesenberg and Orenberg),
dingy b r o w n to black ( M e l z e r i n C e s k a M y k o l . 8: 82-87. 1954). not recorded f r o m B e l g i u m . Sept.-Oct. W i d e s p r e a d a n d very rare i n
temperate Europe.
2. Melanophyllum eyrei (Mass.) S i n g , i n L i l l o a 22: 436. ( ' 1 9 4 9 ' )
1 9 5 1 . - F i g . 128. Urbonas (in Urbonas et a l . , K o n s p . agar. G r i b . L S S R , L S S R , E S S R :
Schulzeria eyrei M a s s . i n G r e v i l l e a 22: 38. 1894; Chlorospora eyrei 54. 1974) described Melanophyllum eyrei var. macrosporum, with
(Mass.) M a s s . i n K e w B u l l . 1898: 136. 1898; Lepiota eyrei (Mass.) J . spores 5-7.5 X 2-2.5 p m : bigger and slightly narrower than those o f the
L a n g e , F l . agar. dan. 1: 36. 1935. typical variety.
S E L . I C O N . - Candusso & L a n z o n i , L e p i o t a : p i . l b . 1990; J . L a n g e , T h e spores are ornamented w i t h isolated rounded warts, as seen
F l . agar. dan. 1: p i . 1 3 B . 1935; L a n z o n i & Z e c c h i n i n R i v . M i c o l . 3 1 : w i t h Scanning E l e c t r o n M i c r o s c o p e .
100. 1988; R . P h i l l i p s , Paddest. S c h i m m . : 30. 1981.
Index
Illustrations are i n d i c a t e d b y a n asteriks (*) a d d e d to the p a g e number.
abruptibulba, M a c r o l e p i o t a 72 L e u c o a g a r i c u s m a r r i a g e i 98 b e n e s i i , P s a l l i o t a 41 caepestipes, L e p i o t a 77
abruptibulba, Psalliota 47 amylosporus, Leucoagaricus bemardii, Agaricus 33* caepestipes, L e u c o c o p r i n u s
abruptibulbus, A g a r i c u s 47 106 b e r n a r d i i f o r m i s , A g a r i c u s 34 77, 7 8 *
acerina, L e p i o t a 132 andegavensis, L e p i o t a 131 bettinae, E c h i n o d e r m a 129 caeruleoviolaceus var. o f
acutesquamosa, Cystolepiota a n g u s t i s p o r a , var. o f L e p i o t a bettinae, L e p i o t a 129 L e u c o a g a r i c u s i o n i d i c o l o r 99
144 a l b a 118 biberi, Agaricus 32 caeruleoviolaceus,
a c u t e s q u a m o s a , L e p i o t a 144, apatelia, L e p i o t a 138, 1 3 9 * b i c k h a m e n s i s , L e p i o t a 126 Leucoagaricus 99
145 a r e n i c o l a , A g a r i c u s 35 biornata, L e p i o t a 9 3 c a e s p i t i p e s , A g a r i c u s 37
a c u t e s q u a m o s a , var. o f L e p i o t a a r e n i c o l a , L e p i o t a 107 biornatus, L e u c o a g a r i c u s 93 c a l c i c o l a , C y s t o l e p i o t a 146,
aspera 144 arenicola, Leucoagaricus 107 birnbaumii, A g a r i c u s 80 147*
a c u t e s q u a m o s a , var. o f L e p i o t a a r e n i c o l a , P s a l l i o t a 35 birnbaumii, Leucocoprinus c a l c i c o l a , L e p i o t a 146
f r i e s i i 144 a r e n o p h i l u s , A g a r i c u s 35 80* c a l d a r i o r u m , L e u c o c o p r i n u s 93
acutesquamosum, a r m i l l a t a , var. o f L e p i o t a b i s p o r a , P s a l l i o t a 35 calongei, Agaricus 32
Echinoderma 144 clypeolarioides 126 bisporus, A g a r i c u s 3 5 * c a m p a n e t t a , var. o f L e p i o t a
a c u t e s q u a m o s u s , A g a r i c u s 144 arvensis, A g a r i c u s 4 3 * bitorquis, A g a r i c u s 3 2 * c l y p e o l a r i a 117
adulterina, C y s t o l e p i o t a 157 arvensis, P s a l l i o t a 4 3 bitorquis, Psalliota 32 c a m p e s t r i s var. o f P s a l l i o t a
adulterina, L e p i o t a 157 asper, A g a r i c u s 144 boertmannii, E c h i n o d e r m a 150 squamulosa 29
aestivalis, A g a r i c u s 3 0 , 31 aspera, A m a n i t a 144 b o e r t m a n n i i , L e p i o t a 148, campestris, A g a r i c u s 28*, 30
aestivalis, P s a l l i o t a 3 0 aspera, C y s t o l e p i o t a 144 151* c a m p e s t r i s , P s a l l i o t a 28
affinis, M a c r o l e p i o t a 7 0 aspera, L e p i o t a 144, 145*, 151 b o h e m i c a , L e p i o t a 71 c a m p e s t r i s , var. o f A g a r i c u s
Agaricus 23 a s p e r u l a , var. o f L e p i o t a b o h e m i c a , M a c r o l e p i o t a 71 campestris 28*, 29
a l b a , L e p i o t a 118 e c h i n e l l a 149 b o h e m i c a , var. o f camporum, A n n u l a r i a 74
alba, var. o f L e p i o t a c l y p e o l a r - asperula, var. o f L e p i o t a e r i o - M a c r o l e p i o t a r a c h o d e s 71 camporum, L e p i o t a 74
i a 118 p h o r a 149 bohusii, A g a r i c u s 37, 38* c a n a l i , M e l a n o p h y l l u m 161
albertii, A g a r i c u s 49 asperum, Echinoderma 144 b o u d i e r i , L e p i o t a 8 3 , 130, carinii, Echinoderma 149
a l b i d a , f o r m a o f A g a r i c u s hort- a t r o b r u n n e a var. of. 132* carinii, L e p i o t a 149*
ensis 34 Macrolepiota mastoidea 70 b r e b i s s o n i i , L e p i o t a 81 carneifolia, L e p i o t a 89
a l b i d a , f o r m a o f P s a l l i o t a hort- audreae, L e p i o t a 120 brebissonii, Leucocoprinus carneifolia, Leucoagaricus 89
ensis 3 4 audreae, var. o f L e p t i o a c o r t i - 81* carneifolius, Leucocoprinus 89
a l b o s a n g u i n e u s , A g a r i c u s 41 n a r i u s 120, 121 bresadolae, L e p i o t a , 9 2 , 93 carneifolius,var. o f L e p i o t a
a l b o s e r i c e a , L e p i o t a 133 augusta, P r a t e l l a 4 4 bresadolae, L e u c o a g a r i c u s 8 5 , l e u c o t h i t e s 89
albosericeus, A g a r i c u s 30 augusta, P s a l l i o t a 4 4 93 c a r o l i , A g a r i c u s 41
albus, f o r m a o f A g a r i c u s per- augustus, A g a r i c u s 4 4 * bresadolae, L e u c o c o p r i n u s 93 c a r o l i , var. o f A g a r i c u s squa-
rarus 4 5 aurantiovergens, bresadolanus, A g a r i c u s 6 0 * m u l i f e r u s 41
albus, var. o f L e u c o c o p r i n u s Leucoagaricus 98 brunneocingulatus, castanea, L e p i o t a 1 3 0 *
straminellus 83 aurea, L e p i o t a 8 0 Leucoagaricus 101 c a s t a n e i d i s c a , L e p i o t a 138
Allopsalliota 6 2 brunneoincarnata, L e p i o t a c a s t a n e o p a l l i d a , L e p i o t a 132,
altipes, A g a r i c u s 3 0 * b a d h a m i i , A g a r i c u s 95 124, 127*, 128, 132 133
altipes, P s a l l i o t a 3 0 badhamii, Lepiota 71, 93, 95 brunneola, Agaricus 40 cepaestipes, L e p i o t a 91
americanus, Leucocoprinus badhamii, L e p i o t o p h y l l u m 95 brunneoolivacea, forma o f cepaestipes, L e u c o c o p r i n u s
85,92 badhamii, Leucoagaricus 93, L e p i o t a g r a n g e i 135 78*
americanus, A g a r i c u s 92 9 5 * , 97 brunnescens, A g a r i c u s 35 C h a m a e m y c e s 152
americanus, L e p i o t a 92 badhamii, Leucocoprinus 93, brunnescens, P s a l l i o t a 35 Chlorophyllum 74
americanus, Leucoagaricus 95 b u c k n a l l i i , A g a r i c u s 158 Chlorosperma 74
92*, 96 b a r s s i i , L e p i o t a 88 b u c k n a l l i i , C y s t o l e p i o t a 158, Chlorospora 74
a m e t h y s t i n a , P s a l l i o t a 5 0 , 51 b a r s s i i , L e u c o a g a r i c u s 85, 8 8 * 159* cinerascens, Leucoagaricus 90,
a m m o v i r e s c e n s , var. o f benesii, Agaricus 41, 4 2 * b u c k n a l l i i , L e p i o t a 159 91
163
164 INDEX
cinereolilacinus, d e m i s a n n u l a , D r o s e l l a 152 eriophorum, Echinoderma fracidus, C h a m a e m y c e s 152*,

Leucoagaricus 91 d e m i s a n n u l a , L e p i o t a 152 147, 148 153
cinereopallidus, Sericeomyces d e m i s a m i u l u s , A g a r i c u s 152 erminea, Lepiota 118* f r i e s i i , A g a r i c u s 144
107 demisannulus, Chamaemyces e r m i n e u s , A g a r i c u s 118 f r i e s i i , C y s t o l e p i o t a 144
c i n g u l u m , L e p i o t a 132, 1 3 3 * , 152, 153 erubescens, var. o f L e p i o t a friesii, Echinoderma 144
134 d e n s e s q u a m o s a , L e p i o t a 91 b a d h a m i i 96 f r i e s i i , L e p i o t a 144
citrifolius, f.of Leucoagaricus d e n s i f o l i a , L e p i o t a 89 esculenta, L e p i o t a 7 4 fuligineobrunneus, forma o f
m e l a n o t r i c h u s var. f u l i g i - densifolius, Leucoagaricus 89 esculentum, C h l o r o p h y l l u m 74 Leucoagaricus melan-
n e o b r u n n e u s 100 denudata, L e p i o t a 83 essettei, A g a r i c u s 4 7 o t r i c h u s 100
c i t r o p h y l l a , L e p i o t a 125 denudatus, A g a r i c u s 83 e x a n n u l a t a , var. o f L e p i o t a fuligineodiffractus,
c l e l a n d i i , M a c r o l e p i o t a 65 denudatus, L e u c o c o p r i n u s 83 cristata 138 Leucocoprinus 100
c l y p e o l a r i a , L e p i o t a 116*, 117 depauperata, P s a l l i o t a 4 2 excellens, A g a r i c u s 49, 50 fuligineosquarrosa,
clypeolarioides, L e p i o t a depauperatus, A g a r i c u s 4 2 * , excellens, Psalliota 49, 50 Macrolepiota 70
122,126 43 e x c e l l e n s , subsp. o f A g a r i c u s fuliginobrunneus, var. o f
c l y p e o l a r i u s , A g a r i c u s 109 devoniensis, A g a r i c u s 3 5 * macrosporus 49, 50 Leucoagaricus melan-
c o c c i n e o b a s a l i s var. of. d e y l i i , A g a r i c u s 4 2 , 43 e x c e l l e n s , var. o f A g a r i c u s o t r i c h u s 100
Macrolepiota mastoidea 70 diversisporus, var. o f albertii 49, 50 f u l i g i n o s a , L e u c o c o p r i n u s 67
collina, Psalliota 49 Leucocoprinus croceoveluti- e x c e l l e n s , var. o f A g a r i c u s fuliginosa, M a c r o l e p i o t a 67,
c o l u b r i n a , L e p i o t a 109 nus 97 macrosporus 49, 50 68*
c o l u b r i n u s , A g a r i c u s 109 D r o s e l l a 152 excoriata, L e p i o t a 7 0 f u l i g i n o s a , var. o f ••
c o m t u l u s , A g a r i c u s 5 2 * , 53 d r y a d i c o l a , L e p i o t a 120 excoriata, M a c r o l e p i o t a 69, 7 0 * M a c r o l e p i o t a p r o c e r a 67
c o n g o l e n s e , var. o f d r y a d i c o l a , var. o f L e p i o t a excoriatum, L e p i o t o p h y l l u m f u l i g i n o s a , var. o f L e p i o t a p r o -
C h l o r o p h y l l u m molybdites audreae 120, 121 70 c e r a 67
74 d r y a d i c o l a , var. o f L e p i o t a excoriatus L e u c o a g a r i c u s 70 f u l v a , var. o f L e p i o t a v e n -
congolensis, A g a r i c u s 74 cortinarius 120, 121 excoriatus, A g a r i c u s 70 t r i o s o s p o r a 117, 118
c o n i f e r a r u m , var. o f d u l c i d u l u s , A g a r i c u s 50. 5 1 * e y r e i , C h l o r o s p o r a 162 f u l v e l l a , L e p i o t a 130, 131, 132
f o r q u i g n o n i i , L e p i o t a 124 e y r e i , L e p i o t a 162 - furcata, var. o f L e p i t o a
c o o k e i , L e p i o t a 141 e c h i n a c e a , C y s t o l e p i o t a 148 eyrei, M e l a n o p h y l l u m 162* a c u t e s q u a m o s a 144
coprinoides, L e p i o t a 80 e c h i n a c e a , L e p i o t a 148* e y r e i , S c h u l z e r i a 162 f u s c o f i b r i l l o s a , P s a l l i o t a 41
cortinarius, L e p i o t a 120* echinaceum, Echinoderma 148 fuscofibrillosus, A g a r i c u s 4 1 *
c o x h e a d i i , L e p i o t a 126 echinata, I n o c y b e 161 farinolens, L e p i o t a 124 fuscosquamatus var. o f
cretacea, L e p i o t a 78 echinata, L e p i o t a 161 favrei, L e p i o t a 123, 126 A g a r i c u s s y l v a t i c u s 39
cretaceus, A g a r i c u s 78 echinata, N a u c o r i a 161 f e l i n a , L e p i o t a 122*, 141 f u s c o v i n a c e a , L e p i o t a 129,
cretaceus, L e u c o a g a r i c u s 7 8 , echinata, P r a t e l l a 161 f e l i n a , var. o f L e p i o t a c l y p e o - 130*
79* echinata, P s a l l i o t a 161 l a r i a 122 F u s i s p o r a 109
cretata, L e p i o t a 78 echinatum, Cystoderma 161 f e l i n o i d e s , var. o f L e p i o t a fusispora, L e p i o t a 117
cretatus, L e u c o c o p r i n u s 78 echinatum, M e l a n o p h y l l u m cristata 138
cristata, L e p i o t a 7 7 , 138*, 161 felinus, A g a r i c u s 122 gaillardii, Leucoagaricus 102
139, 140 echinatus , A g a r i c u s 161 fissurata, P s a l l i o t a 4 3 gauguei, Leucoagaricus 99,
cristatoides, L e p i o t a 138, e c h i n e l l a , L e p i o t a 125*, 126 fissuratus, A g a r i c u s 4 3 104*
139*, 140 e c h i n e l l a , var. o f L e p i o t a f l a v a , var. o f L e p i o t a c o r t i n a r - gauguei, Sericeomyces 104
cristatus, A g a r i c u s 138 e c h i n e l l a 126 ius 120, 121 geesterani, A g a r i c u s 6 2
croceovelutinus, e c h i n e l l u s , L e p i o t a 125, 126 flavescens, A g a r i c u s 4 7 , 5 4 geesterani, A l l o p s a l l i o t a 6 2 ,
Leucoagaricus 97* Echinoderma 109 flavescens, P r a t e l l a 4 7 63*
crystallifer, Leucoagaricus edulis, A g a r i c u s 32 flavescens, P s a l l i o t a 4 7 , 54 gennadii, Agaricus 34*
106*, 107, 108 e d u l i s , var. o f A g a r i c u s flavotactus, var. o f A g a r i c u s g e n n a d i i , C h i t o n i a 34
c u p r e o b r u n n e a , P s a l l i o t a 31 c a m p e s t r i s 32 aestivalis 3 0 georginae, A g a r i c u s 9 6
c u p r e o b r u n n e a , var. o f efibulis, E c h i n o d e r m a 147 flavotactus, var. o f P s a l l i o t a georginae, L e p i o t a 96
Psalliota campestris 28*, 29 e f i b u l i s , L e p i o t a 147 aestivalis 3 0 georginae, L e u c o a g a r i c u s 96*,
cupreobrunneus, A g a r i c u s 3 1 * e l a i o p h y l l a , L e p i o t a 124, 128 floccipes, A g a r i c u s 29 97
cupreus, A g a r i c u s 82, 9 2 e l v e n s i s , A g a r i c u s 37 f l o c c i p e s , var. o f A g a r i c u s gigantea, f o r m a o f L e p i o t a
c y g n e a , C y s t o l e p i o t a 82 E n d o p t y c h u m 21 campestris 29 a c u t e s q u a m o s a 144
cygnea, L e p i o t a 82 equestris, var. o f A g a r i c u s flocculosa, Psalliota 30 giganteum, forma o f
cygnea, Pseudobaeospora 82 campestris 28, 29 f l o s - s u l p h u r i s , A g a r i c u s 80 E c h i n o d e r m a acutesquamo-
cygneus, Leucocoprinus 82* equestris, var. o f P s a l l i o t a flos-sulphuris, L e p i o t a 80 s u m 144
cygneus, Sericeomyces 82 campestris, 28, 29 flos-sulphuris, Leucocoprinus glabridiscus, Leucoagaricus
cylindrosporus, Sericeomyces e r i o d e r m a var. o f L e p i o t a sere- 80 102
108 n a 106 f l o s - s u l p h u r i s , var. o f L e p i o t a g l a r e o p h y l l a , L e p i o t a 126
cystidiosa, Cystolepiota 155* erioderma, Leucoagaricus 106 caepestipes, 8 0 Glaucospora 161
c y s t i d i o s a , L e p i o t a 155 e r i o p h i o r a , C y s t o l e p i o t a 147, forquignonii, L e p i o t a 124* gracilenta, L e p i o t a 6 8 , 6 9
Cystolepiota 154 148 fracida, D r o s e l l a 152 gracilenta, M a c r o l e p i o t a 6 8 ,
e r i o p h o r a , L e p i o t a 147 fracidus, A g a r i c u s 152 69
INDEX 165
gracilentus, A g a r i c u s 68 hystrix, Echinoderma 146 latispora, L e p i o t a 122 m a c r o l e p i s , var. o f A g a r i c u s

gracilis, forma of Lepiota h y s t r i x , L e p i o t a 146 l e p i d a , L e p i o t a 126 arvensis 2 6
h a e m a t o s p e r m a 161 Lepiota 109 m a c r o r h i z a , L e p i o t a 88
gracilis, forma of Lepiota ful- ianthina, L e p i o t a 80 L e p i o t e l l a 152 macrorhizus, Leucoagaricus
v e l l a 132 ianthinophaeus, Leucoagaricus lepiotoides, var. o f A g a r i c u s 88
g r a c i l i s , L e p i o t a 121 99 xanthodermus 54 macrorhizus, Leucocoprinus
g r a c i l i s , var. o f L e p i o t a c l y p e - ianthinosquamulosus, L e p i o t u l a 109 88
o l a r i a 121 Leucoagaricus 99 Leucoagaricus 85 macrospora, Psalliota 49
g r a n g e i , H i a t u l a 137 ianthinus, A g a r i c u s 80 Leucocoprinus 7 6 m a c r o s p o r a , subsp. o f
g r a n g e i , L e p i o t a 124, 1 3 7 * i a n t h i n u s , L e u c o c o p r i n u s 80, leucothites, A g a r i c u s 89 P s a l l i o t a arvensis 4 9
g r a n g e i , L e p i o t u l a 137 81* leucothites, L e p i o t a 89 macrosporoides, Agaricus 50
g r a n g e i , S c h u l z e r i a 137 i c t e r i n a , C y s t o l e p i o t a 159 leucothites, L e u c o a g a r i c u s 7 8 , macrosporus, Agaricus 49
g r a n u l o p u n c t a t a , L e p i o t a 120 i c t e r i n a , L e p i o t a 159 79, 8 9 * macrosporus, forma o f
grisea, P s a l l i o t a 5 4 idae-fragum, Leucoagaricus leucothites,var. o f L e u c o c o p r i n u s caepestipes
grisea, var. o f P s a l l i o t a x a n t h o - 103 Leucoagaricus carneifolius 77
dermus 54 i g n i c o l o r , L e p i o t a 130, 131 89 m a c r o s p o r u s , var. o f A g a r i c u s
griseodiscus, Leucocoprinus i g n i p e s , L e p i o t a 130 l e u c o t r i c h a , P s a l l i o t a 43 lutosus 5 1 , 5 2
100 i g n i v o l v a t a , L e p i o t a 122 leucotrichus, A g a r i c u s 4 3 m a g n i s p o r a , L e p i o t a 117*,
griseovirens, L e p i o t a 135*, i m p u d i c a , Psalliota 39 l i g n i c o l a , L e p i o t a 109 118
136, 137 impudicus, Agaricus 39* l i g n i c o l a , L e u c o p h o l i o t a 109 mairei, Leucoagaricus 108
g r i s e o v i r e n s , var. o f L e p i o t a i m r e h i , var. o f A . m a s k a e 58 lilacea, Lepiota m a l e o l e n s , A g a r i c u s 33
griseovirens 135 infida, Psalliota 60 l i l a c e a , L e p i o t a 139, 140, 1 4 1 * m a r g i n a t a , var. o f L e p i o t a
griseus, var. o f A g a r i c u s x a n - infidus, A g a r i c u s 60 l i l a c i n a , L e p i o t a 158 m o l y b d i t e s 74
thodermus 54 i n f i d u s , var. o f A g a r i c u s l i l a c i n a , var. o f L e p i o t a s e m i - m a r r i a g e i , L e p i o t a 98
guadelupensis, A g a r i c u s 74 romagnesii 60 n u d a 158 m a r r i a g e i , L e u c o a g a r i c u s 98,
g u e g u e n i i , L e p i o t a 83 i n g r a t a , P s a l l i o t a 33 l i l a c i n a , var. o f C y s t o l e p i o t a 99*
ingratus, A g a r i c u s 33 b u c k n a l l i i 158 maskae, A g a r i c u s 58, 59
h a e m a t o s p e r m a , L e p i o t a 161 ionidicolor, Leucoagaricus 99* l i l a c i n o g r a n u l o s a var. o f Mastocephalus 76
h a e m a t o s p e r m a , P s a l l i o t a 161 irrorata, D r o s e l l a 152 H i a t u l a cepaestipes 8 0 m a s t o i d e a , L e p i o t a 68
haematospermum, irrorata, L e p i o t a 152 lilacinogranulosa, L e p i o t a 80 mastoidea, M a c r o l e p i o t a 68,
M e l a n o p h y l l u m 161* irrorata, L e p i o t e l l a 152 lilacinogranulosus, 69*
haematospermus, Agaricus Leucocoprinus 80 mastoidea, subsp.of Lepiota
161 jacobii, Echinoderma 147 litoralis, A g a r i c u s 58*, 59 e x c o r i a t a 68
h a e m o r r h o i d a r i u s , A g a r i c u s 38 jacobi, L e p i o t a 148* l i t o r a l i s , P s a l l i o t a 58 mastoideum, Lepiotophyllum
h e i m i i , M a c r o l e p i o t a 71 jodoformicus, Agaricus 54 littoralis, L e p i o t a 102 68
heinemannii, Leucocoprinus j o s s e r a n d i i , L e p i o t a 127 littoralis, L e u c o a g a r i c u s 102 m a s t o i d e u s , A g a r i c u s 68, 6 9
83,84 josserandii, Leucoagaricus 127 lividonitidus, Agaricus 32 medioflavoides, Leucoagaricus
h e l v e o l a , L e p i o t a 127 j o s s e r a n d i i , var. o f L e p i o t a l o c q u i n i i , L e p i o t a 126 105
hetieri, Cystolepiota 156* s u b i n c a r n a t a 127, 128 l o n g i c a u d u s , A g a r i c u s 31 mediofusca, Psalliota 40
h e t i e r i , L e p i o t a 156 jubilaei, Leucoagaricus 98 lutea, L e p i o t a 8 0 mediofuscus, A g a r i c u s 40
hetieriana, C y s t o l e p i o t a 159 luteicystidiata, C y s t o l e p i o t a m e d i o f u s c u s , var. o f A g a r i c u s
hetieriana, L e p i o t a 159 koelerionensis, A g a r i c u s 39, 155 langei 40
h i s p i d a , C y s t o l e p i o t a 146, 159 40 luteicystidiata, L e p i o t a 155 m e d i o r o s e a , L e p i o t a 140
h i s p i d a , L e p i o t a 146 k o e l e r i o n e n s i s , var. o f luteolorufescens, A g a r i c u s 5 9 , m e d i t e r r a n e a , var. o f
h o l o s e r i c e u s , A g a r i c u s 89 Psalliota variegata 39 60 Macrolepiota procera 76
holosericeus, Leucoagaricus k o n r a d i i , L e p i o t a 67 l u t e o l o s p e r m a , var. o f M e l a n o p h y l l u m 161
89 k o n r a d i i , M a c r o l e p i o t a 67 Chlorophyllum molybdites m e l a n o t r i c h a , L e p i o t a 100
holosericeus, Leucocoprinus k o n r a d i i , var. o f L e p i o t a e x c o - 74 melanotrichus, Leucoagaricus
89 riata 69 l u t e o m a c u l a t a , P s a l l i o t a 51 84, 8 5 , 100*
holospilotus, Leucocoprinus k u e h n e r i , L e p i o t a 126 luteomaculatus, A g a r i c u s 5 1 , melanotrichus, Leucocoprinus
94 k u e h n e r i a n a , L e p i o t a 121, 122 52* 100
hortensis, A g a r i c u s 71 kuehnerianus, A g a r i c u s 49 luteus, A g a r i c u s 8 0 meleagris, A g a r i c u s 56, 94
hortensis, P s a l l i o t a 71 luteus, L e u c o c o p r i n u s 8 0 m e l e a g r i s , G y m n o p u s 94
hortensis, var. o f L e p i o t a l a e v i g a t a , var. o f L e p i o t a gra- lutosus, A g a r i c u s 51 meleagris, Hiatula 94
r a c h o d e s 71 c i l i s 119 l y c o p e r d o i d e s , L e p i o t a 155 meleagris, L e p i o t a 94
hortensis, var. o f A g a r i c u s laevigata, var. o f L e p i o t a ore- l y c o p e r d o i d e s , var. o f m e l e a g r i s , L e u c o a g a r i c u s 85,
c a m p e s t r i s 71 adiformis 119 Cystolepiota luteicystidiata 93, 94*
h u i j s m a n i i , var. o f L e p i o t a langei, Agaricus 4 0 * 155 meleagris, Leucocoprinus 94
o c h r a c e o f u l v a 1 4 1 , 142 l a n g e i , L e p i o t a 147 meleagris, Psalliota 94
h y m e n o d e r m a , L e p i o t a 138, langei, Psalliota 40 m a c r o c a r p a , P s a l l i o t a 48 m e l e a g r i s , var. o f A g a r i c u s
140* lanipes, A g a r i c u s 5 9 * macrocarpus, A g a r i c u s 4 8 * moelleri 56
h y s t r i x , C y s t o l e p i o t a 146 lanipes, Psalliota 59 Macrolepiota 6 4 m e l e a g r o i d e s , L e p i o t a 95
166 INDEX
m e n i e r i , L e p i o t a 107 olgae, L e p i o t a 89 * l i l a c e a 139 p s e u d o l i l a c e a , L e p i o t a 123*,

menieri, Leucoagaricus 107*, olgae, L e u c o a g a r i c u s 89 phaeophylla, forma o f Lepiota 124
108 olgae, L e u c o c o p r i n u s 89 rufipes 143 p s e u d o l i l a c e a , L e p i o t u l a 123
menieri, Sericeomyces 107 o l i v a c e o b r u n n e a , L e p i o t a 124 p h a l l o i d e s , A m a n i t a 127 pseudolutosus, A g a r i c u s 52
m e t u l a e s p o r a , L e p i o t a 117, o l i v a c e o b r u n n e a , var. o f pilatianus, A g a r i c u s 5 5 * , 56 p s e u d o o l i v a s c e n s , var. o f
118 L e p i o t a f o r q u i g n o n i i 123 pilatianus, L e u c o a g a r i c u s 9 7 , M a c r o l e p i o t a p r o c e r a 67
micropholoides, forma of o l i v a s c e n s , M a c r o l e p i o t a 67 98* pseudopratensis, A g a r i c u s 5 6 *
L e p i o t a l i l a c e a 140 olivieri, forma o f Macrolepiota pilatianus, L e u c o c o p r i n u s 97 pseudopratensis, P s a l l i o t a 56
microsporias, ssp. o f A g a r i c u s rachodes 73 pilatianus, var. o f A g a r i c u s pseudoumbrella, Agaricus 46,
g e n n a d i i 34 o l i v i e r i , L e p i o t a r a c h o d e s var. x a n t h o d e r m u s 55 47*
m i n i m a , f o r m a o f L e p i o t a sis- o l i v i e r i 73 pilatianus, var. o f p s e u d o v i l l a t i c u s , A g a r i c u s 37
trata 158 olivieri, Macrolepiota 71, 72, Leucocoprinus badhamii 97 pudicus, Leucoagaricus 90
minimus, forma of Cysto- 73*, 95 p i l o d e s , L e p i o t a 134, 1 3 5 * p u e l l a r i s , L e p i o t a 91
l e p i o t a pulverulenta 159 o l i v i e r i , var. o f M a c r o l e p i o t a p i n g u i p e s , var. o f L e u c o - p u e l l a r i s , L e u c o c o p r i n u s 91
minimus, forma of Leuco- r a c h o d e s 73 agaricus m a c r o r h i z u s 88 p u e l l a r i s , M a c r o l e p i o t a 91
a g a r i c u s p u l v e r u l e n t u s 159 oreadiformis, L e p i o t a 119* p i n g u i p e s , L e p i o t a 88 p u e l l a r i s , var. o f A g a r i c u s
m i n o r , var. o f L e p i o t a c l y p e o - osecanus, A g a r i c u s 4 3 , 4 5 , 4 6 * pinguipes, Leucoagaricus 85, r a c h o d e s 91
l a r i a 116 otsuensis, L e u c o c o p r i n u s 81 88, 105 p u e l l a r i s , var. o f
m i n u t a , L e p i o t a 125 p l a c o m y c e s , A g a r i c u s 5 6 , 57 L e p i o o p h y l l u m r a c h o d e s 91
moelleri, A g a r i c u s 56, 57* p a l l i d a , L e p i o t a 120 platensis, L e p i o t a 120 p u e l l a r i s , var. o f L e p i o t a p r o -
moelleri, Cystolepiota 150* p a l l i d a , var. o f L e p i o t a l i l a c e a p o l i o c h l o o d e s , L e p i o t a 136*, c e r a 91
moellerianus, Agaricus 29* 139, 140 137 p u e l l a r i s , var. o f L e p i o t a
molybdites, Leucocoprinus 74 p a l l i d i o r , var. o f L e p i o t a porphyhizon, Agaricus 53, 54* r a c h o d e s 91
molybdites, A g a r i c u s 74 cristata 138 p o r p h y r e a , P s a l l i o t a 31 P u l v e r o l e p i o t a 154
molybdites, C h l o r o p h y l l u m 74 p a l l i d u s , var. o f A g a r i c u s s y l - * p o r p h y r e u s A g a r i c u s 31 pulverulenta, Cystolepiota
molybdites, M a c r o l e p i o t a 74 vaticus 39 porphyrocephalus, Agaricus 159, 1 6 0 *
molybdites, Mastocephalus 74 p a l l i d u s , f. L e u c o a g a r i c u s pur- 31* p u l v e r u l e n t a , L e p i o t a 159
morganii, Agaricus 74 p u r e o r i m o s u s 103 praeclaresquamosus, A g a r i c u s pulverulenta, Pulverolepiota
morganii, C h l o r o p h y l l u m 74 pampeanus, A g a r i c u s 29, 3 0 * 56 159
morganii, L e p i o t a 74 p a n t h e r i n a , var. o f L e p i o t a P r a t e l l a 23 pulverulentus, Leucoagaricus
morganii, Mastocephalus 74 c l y p e o l a r i a 117 pratensis, L e p i o t a 119, 120 159
moseri, Leucoagaricus 90 p a r k e n s i s , var. o f A g a r i c u s pratensis, var. o f A g a r i c u s p u r p u r a s c e n s , A g a r i c u s 53
n i v e s c e n s 45 c l y p e o l a r i u s 119 p u r p u r a s c e n s , P s a l l i o t a 53
n a u c i n a , L e p i o t a 133 p a r k e n s i s , var. o f P s a l l i o t a p r o c e r a , L e p i o t a 65 p u r p u r a s c e n s , var. o f A g a r i c u s
naucinus, A g a r i c u s 89 n i v e s c e n s 45 procera, M a c r o l e p i o t a 65, 66* arvensis 53
naucinus, Leucoagaricus 89 p a r v a n n u l a t a , L e p i o t a 129* p r o c e r u m , L e p i o t o p h y l l u m 65 purpurella, Psalliota 50
n i g r e s c e n t i p e s , L e p i o t a 129 p a r v a n n u l a t u s , A g a r i c u s 129 procerus, Mastocephalus 65 purpurellus, A g a r i c u s 50
niveolutescens, A g a r i c u s 52 passerinii, forma o f prominens, Macrolepiota 70 purpureolilacinus,
nivescens, A g a r i c u s 45 C h a m a e m y c e s pseudocasta- psalion, L e p i o t a l 4 3 L e u c o a g a r i c u s 103*, 104
nivescens, Psalliota 45 neus 152 P s a l l i o t a 23 purpureorimosus,
niveus, var. o f A g a r i c u s p e q u i n i i , A g a r i c u s 34 psammophila, Macrolepiota 70 L e u c o a g a r i c u s 103, 104
p s e u d o p r a t e n s i s 56 p e r d i n i c u s , A g a r i c u s 57 pseudoasperula, Cystolepiota pyrochroa, L e p i o t a 142*
n y m p h a r u m , A g a r i c u s 91 p e r d i n i c u s , var. o f A g a r i c u s 150
n y m p h a r u m , L e p i o t a 91 m e l e a g r i s 57 pseudoasperula, L e p i o t a 150* rachodes, A g a r i c u s 71
nympharum, Leucoagaricus p e r m i x t a , L e p i o t a 67 pseudoasperulum, r a c h o d e s , L e p i o t o p h y l l u m 71
91*, 92 p e r m i x t a , M a c r o l e p i o t a 66, 67 Echinoderma 150 rachodes, L e u c o c o p r i n u s 71
n y m p h a r u m , M a c r o l e p i o t a 91 p e r m i x t a , var. o f M a c r o l e p i o t a Pseudobaeospora 82 rachodes, M a c r o l e p i o t a 7 1 *
p r o c e r a 67 pseudocastaneus, rachodes, var. o f A g a r i c u s p r o -
o b s c u r a , L e p i o t a 135,1 3 6 p e r m i x t u s , L e u c o c o p r i n u s 67 C h a m a e m y c e s 152 cerus 71
o b s c u r a , ssp. o f L e p i o t a peronata, P s a l l i o t a 4 4 pseudocastaneus, var. o f radicatus, (3 o f A g a r i c u s
griseovirens 135 peronatus, A g a r i c u s 4 4 C h a m a e m y c e s fracidus 152 campestris 60
obscurata, var. o f P s a l l i o t a p e r p l e x a , C y s t o l e p i o t a 145 pseudocinerascens, radicatus, A g a r i c u s 6 0
m e l e a g r i s 57 p e r p l e x a , L e p i o t a 144, 1 4 6 * Leucoagaricus 88 reai, A g a r i c u s 3 9
obscuratus, var. o f A g a r i c u s perplexum, Echinoderma 145 p s e u d o c i n e r a s c e n s , var. o f reidii, forma o f Cystolepiota
x a n t h o d e r m u s 5 6 , 57 perrara, P s a l l i o t a 4 4 Leucoagaricus macrorhizus adulterina 157
o c h r a c e o c y a n e a , L e p i o t a 137 perrarus, A g a r i c u s 4 4 , 4 5 88 r h a c o d e s , L e p i o t a 71
o c h r a c e o d i s c a , L e p i o t a 118 perrarus, var. o f A g a r i c u s pseudocretaceus, A g a r i c u s 5 4 rhacodes, L e p i o t o p h y l l u m 71
ochraceofulva, L e p i o t a 141*, augustus 4 4 p s e u d o f e l i n a , L e p i o t a 135, r h o d o r h i z a , L e p i o t a 125
142 phaeodisca, Macrolepiota 70 136, 137 r h o d o r h i z a , var. o f L e p i o t a
ochraceosulfurescens, L e p i o t a phaeolepidotus, A g a r i c u s 57* p s e u d o h e l v e o l a , L e p i o t a 123, s e t u l o s a 127
116, 117 p h a e o l e p i d o t u s , P s a l l i o t a 57 124 r h o d o r h i z a , var. o f L e p t i o a
ochrospora, Lepiota 74 phaeophylla, forma of Lepiota pseudolicmophora, L e p i o t a 80 e c h i n e l l a 126, 127
INDEX 167
r h o d o s p e r m a , L e p i o t a 67 serenus, A g a r i c u s 105 subadulterina, C y s t o l e p i o t a u m b r e l l o i d e s , var. o f A g a r i c u s

r h o d o s p e r m a , M a c r o l e p i o t a 67 serenus, L e u c o a g a r i c u s 105, 157 arvensis 4 6
Rhozites 90 106*, 107, 129 subalba, L e p i o t a 89, 1 3 3 * umbrinisquamosa, forma of
r i c k e n i i , L e p i o t a 69 serenus, S e r i c e o m y c e s 105 subcretaceus, L e u c o a g a r i c u s L e p i o t a c l y p e o l a r i a 117
r i c k e n i i , M a c r o l e p i o t a 69 sericata, L e p i o t a 104 89 u m b r i n i s q u a m o s a , var. o f
r o b y n s i a n u s , A g a r i c u s 34 sericatella, L e p i o t a 104 subfelinoides, L e p i o t a 138 L e p i o t a c l y p e o l a r i a 117
rollanii, Agaricus 34 sericatellus f o r m a o f subfloccosa, f o r m a o f P s a l l i o t a u m b r i n o r a f e s c e n s , var. o f
romagnesii, Agaricus 60 L e u c o a g a r i c u s sericifer 104, hortensis 36 L e p i o t a v e n t r i o s o s p o r a 117
rorulentus, forma o f 105 subfloccosa, P s a l l i o t a 36 urinascens, A g a r i c u s 4 9 *
L e u c o c o p r i n u s caepestipes sericatellus, Leucoagaricus subfloccosus, A g a r i c u s 3 6 * urinascens, Psalliota 49
78 104 subglobisporus, var. o f u r i n a s c e n s , var. o f A g a r i c u s
r o s a b r u n n e a , var. o f L e p i o t a sericatellus, S e r i c e o m y c e s 104 Leucocoprinus lilacinogran- urinascens 49
josserandii 127 sericatellus, var. o f u l o s u s 81
rosea, C y s t o l e p i o t a 157 S e r i c e o m y c e s sericatus 104 subgracilis, L e p i o t a 1 2 1 * v a l i d u s , var. o f A g a r i c u s
rosea, L e p i o t a 157 sericatus, S e r i c e o m y c e s 104 subincarnata, L e p i o t a 124, bitorquis 32
r o s e l l a , C y s t o l e p i o t a 157 sericea, L e p i o t a 104 127, 128* v a l i d u s , var. o f P s a l l i o t a
r u b e l l a , L e p i o t a 129 sericea, var. o f L e p i o t a cristata sublittoralis, Leucoagaricus bitorquis 32
rubens, L e p i o t a 97 104 1 0 1 * , 102 v a p o r a r i a , P s a l l i o t a 37
rubescens, var. o f Sericeomyces 85 sublittoralis, L e p i o t a 101 v a p o r a r i u s , A g a r i c u s 37
M a c r o l e p i o t a e x c o r i a t a 71 sericeus, L e u c o a g a r i c u s 104 sublittoralis, Leucocoprinus vaporarius, ? o f A g a r i c u s
rubrotinctus, Leucoagaricus sericeus, S e r i c e o m y c e s 104 101 c a m p e s t r i s 37
85, 103 sericifer, L e u c o a g a r i c u s 104, subolivaceus, Leucoagaricus variegans, A g a r i c u s 39
rufescens, L e p i o t a 155 105* 102 variegata, P s a l l i o t a 39
m f e s c e n s , var. o f A g a r i c u s sericifer, S e r i c e o m y c e s 104 subperonata, f o r m a o f variegatus, A g a r i c u s 39
g r a n u l o s u s 155 sericifera, L e p i o t a 104 P s a l l i o t a hortensis 37 venenata, M a c r o l e p i o t a 7 2 , 7 3
rufipes, L e p i o t a 140, 1 4 3 * sericifera, P s e u d o b a e o s p o r a subperonata, P s a l l i o t a 37 veneris, A g a r i c u s 3 0
rufotegulis, A g a r i c u s 6 0 , 6 1 * 104 subperonatus, A g a r i c u s 3 7 * veneris, var. o f A g a r i c u s aesti-
r u f o v e l u t i n a , L e p i o t a 9 5 , 97 setulosa, L e p i o t a 125 subprocera, L e p i o t a 71 valis 30
r u s s i o p h y l l u s , A g a r i c u s 53 singeri, Sericeomyces 108 subpurpurata, var. o f L e p i o t a veneris, var. o f P s a l l i o t a aesti-
s i n o p i c a , L e p i o t a 150 b o u d i e r i 133 valis 30
sabulosa, var. o f L e p i o t a s l o v e n i c u s , A g a r i c u s 34 subrubens, var. o f v e n t r i o s o s p o r a , L e p i o t a 117,
p s e u d o h e l v e o l a 123, 124 sororia, L e p i o t a 158 L e u c o c o p r i n u s p i l a t i a n u s 98 118
salmoneophyllus, s p e c i o s a , L e p i o t a 126 subrufescens, P s a l l i o t a 4 4 v e r e c u n d a , var. o f P s a l l i o t a
Leucoagaricus 97 spissa, P s a l l i o t a 58 subsquarrosa, M a c r o l e p i o t a lanipes 59
s a l m o n e o p h y l l u s , var. o f s p i s s i c a u l i s , A g a r i c u s 59 substramineus, A g a r i c u s 4 9 v e r e c u n d u s , var. o f A g a r i c u s
L e u c o a g a r i c u s pilatianus 9 7 , s q u a m u l i f e r a , P s a l l i o t a 41 subtomentosus, A g a r i c u s 71 l a n i p e s 59
98 s q u a m u l i f e r u s , A g a r i c u s 41 subtomentosus, var. o f v i n o s o b r u n n e u s , A g a r i c u s 38
s a n g u i f l u a , L e p i o t a 95 s q u a m u l o s u s , var. o f A g a r i c u s A g a r i c u s p r o c e r u s 71 v i n o s o b r u n n e u s , var. o f
s a n g u i n a r i u s , A g a r i c u s 38 c a m p e s t r i s 2 8 , 29 subvolvatus, Leucoagaricus A g a r i c u s s y l v a t i c u s 38
sanguinescens, var. o f L e p i o t a squarrosipes, var. o f A g a r i c u s 107, 108 v o l v a t u s , A g a r i c u s 34
r u f o v e l u t i n a 95 nivescens 45 s y l v a t i c a , P s a l l i o t a 38
s a p o n e l l a , L e p i o t a 138 squarrosipes, var. o f A g a r i c u s sylvaticus, A g a r i c u s 38, 39*, w a s s e r i , A g a r i c u s 31
saturata, var. o f P s a l l i o t a s y l - osecanus 4 5 , 4 6 * 60 w a s s e r i , L e p i o t a 121
v a t i c a 38 squarrosus, var. o f A g a r i c u s sylvicola, Agaricus 47* w i c h a n s k i i , L e p i o t a 102
saturatus, var. o f A g a r i c u s s y l - s y l v i c o l a 4 7 * , 48 sylvicola, Psalliota 47 wichanskii, Leucoagaricus
vaticus 38 straminea, Psalliota 49 s y l v i c o l a , var. o f A g a r i c u s 101, 102*, 103
s e m i n u d a , C y s t o l e p i o t a 158* s t r a m i n e l l a , L e p i o t a 4 9 , 83 campestris 47
s e m i n u d a , L e p i o t a 158 s t r a m i n e l l a , var. o f L e p i o t a x a n t h o d e r m a , P s a l l i o t a 54
s e m i n u d a , var. L e p i o t a sistrata cepaestipes 83 tener, L e p i o t a 100 x a n t h o d e r m u s , A g a r i c u s 54,
158 s t r a m i n e l l u s , A g a r i c u s 83 tener, L e u c o a g a r i c u s 100, 55*
s e m i n u d u s , A g a r i c u s 158 straminellus, Leucocoprinus 101* xantholepis, A g a r i c u s 5 3 *
semota, Psalliota 50 83* terricolor, var. o f A g a r i c u s x a n t h o l e p i s , P s a l l i o t a 53
semotus, A g a r i c u s 5 0 stramineosquamulosus, m e l e a g r i s 56, 57 x a n t h o p h y l l a , L e p i o t a 124,
septentrionalis, var. o f Agaricus 49 terricolor, var. o f A g a r i c u s 1 2 5 * , 128, 129
Leucoagaricus melan- stramineus, A g a r i c u s 4 9 praeclaresquamosus 56
o t r i c h u s 100 stramineus, A g a r i c u s 5 0 tomentella, L e p i o t a 134*
serena, L e p i o t a 104, 105 s t r a m i n e u s , var. o f A g a r i c u s t y p i c a , var. o f L e p i o t a
serena, P s e u d o b a e o s p o r a 105 macrosporus 49 a c u t e s q u a m o s a 144, 145
Index to the generic names in volumes 1-5
V o l u m e n u m b e r s are g i v e n i n R o m a n n u m e r a l s ; a c c e p t e d n a m e s are p r i n t e d i n b o l d .
A c a n t h o c y s t i s III: 158 F l a m m u l i n a III: 170 Macrolepiota V : 6 4 P s a m m o s p o r a I V : 153

A e r u g i n o s p o r a II: 71 F l o c c u l a r i a III: 34 Marasmiellus III: 123 Pseudobaeospora III: 132
A g a r i c o c h a e t e III: 158 F u s i s p o r a V : 109 M a r a s m i u s III: 136 Pseudoclitocybe III: 9 2
A g a r i c u s V : 23 Mastocephalus V : 76 P s e u d o h y g r o c y b e II: 71
A l b o l e p t o n i a I: 85 G a l e r o p s i n a I V : 28 Megacollybia I V : 172 Pseudoomphalina III: 93
Allopsalliota V : 6 2 G a m u n d i a III: 155 M e g a t r i c h o l o m a I V : 107 Psilocybe I V : 28
A n d r o s a c e u s III: 136 G e o p e t a l u m II: 30,111: 158 M e l a l e u c a I V : 153 P u l v e r o l e p i o t a V : 154
A r m i l l a r i a III: 34 G e o p h i l a I V : 28 Melanoleuca I V : 153
A r r m l l a r i e l l a III: 34 G l a u c o s p o r a V : 161 M e l a n o p h y l l u m V : 161 Resupinatus III: 166
A r r h e n i a III: 39 G l i o p h o r u s II: 71 M e l a n o t u s I V : 28 R h o d o c o l l y b i a III: 106
A s p r o p a x i l l u s III: 76 G l o i o c e p h a l a III: 136 M i c r o c o l l y b i a III: 106 Rhodocybe I: 7 7
A s t e r o t u s III: 166 G l u t i n a s t e r I V : 107 M i c r o m p h a l e III: 129 R h o d o p h y l l u s I: 85
G o d f r i n i a l l : 71 M o n o m y c e s I V : 107 R h o d o s p o r u s I: 8 2
Baeospora I V : 165 G y m n o p u s III: 106 M u c i d u l a I V : 177 Rhodotus III: 175
B o t r y i d i n a III: 89 G y r o p h i l a I V : 107 Mycenella I V : 173 Rickenella III: 157
M y c e n i t i s III: 136 R i m b a c h i a III: 134
Callistosporium III: 104 Haasiella III: 63 M y x o c o l l y b i a III: 170 Ripartites III: 9 4
Camarophyllopsis II: 111 H e l i o m y c e s III: 136 M y x o m p h a l i a III: 156 R u s s u l i o p s i s III: 9 6
C a m a r o p h y l l u s II: 7 1 , 115 H e m i p h o l i o t a I V : 80
C a m p a n e l l a III: 104 H e x a j u g a I: 8 2 N a e m a t o l o m a I V : 28 S a r c o m y x a III: 168
C a n t h a r e l l u l a III: 41 H i r n e o l a I: 7 7 N o l a n e a I: 85 S c l e r o s t i l b u m III: 106
Catathelasma III: 176 H o d o p h i l u s II: 111 S c o r t e u s III: 136
C h a m a e m y c e s V : 152 Hohenbuehelia III: 158 Omphaliaster III: 78 S c y t i n o t o p s i s III: 166
Chlorophyllum V : 74 H y d r o p u s I V : 166 O m p h a l i n a III: 78 S c y t i n o t u s III: 168
C h l o r o s p e r m a V : 161 H y g r o c y b e II: 71 Omphalotus III: 88 S e r i c e o m y c e s V : 85
C h l o r o s p o r a V : 161 Hygrophoropsis III: 64 O r c e l l a I: 82 S e t u l i p e s III: 136
C l a u d o p u s I: 85 H y g r o p h o r u s II: 115 Ossicaulis III: 131 S i n g e r e l l a III: 4 2
C l i t o c y b e III: 42 L i m a c i u m l l : 115 O u d e m a n s i a I V : 177 S p h a e r o c e p h a l u s I V : 107
C l i t o p i l o p s i s I: 7 7 H y g r o t r a m a II: 111 Oudemansiella I V : 177 S t a c h y o m p h a l i a III: 155
Clitopilus I: 8 2 H y p h o l o m a I V : 28 S t e r c o p h i l a I V : 28
C o l l y b i a III: 106 Panellus III: 168 S t e r e o p h y l l u s III: 63
C o l l y b i d i u m III: 170 Kuehneromyces I V : 80 P a x i l l o p s i s I: 82 Strobilurus I V : 178
C o r i s c i u m III: 89 P h a e o n e m a t o l o m a I V : 80 S t r o p h a r i a I V : 28
C o r n i o l a III: 3 9 L a c c a r i a III: 9 6 Phaeotellus III: 78 S t r o p h o l o m a I V : 28
C o r t i n e l l u s I V : 107 L e n t i d i o p s i s II: 2 0 Pholiota I V : 8 0
Crinipellis III: 135 L e n t i n u l a II: 25 P h o l i o t e l l a I V : 28 T r i c h o l o m a I V : 107
C u p h o p h y l l u s II: 71 Lentinus II: 25 Phyllotopsis II: 24 Tricholomopsis I V : 151
Cyphellostereum III: 6 3 L e p i o t a V : 109 P h y l l o t u s III: 166 T r i c h o l o s p o r i u m I V : 107
Cystolepiota V : 154 L e p i o t e l l a V : 152 Phytoconis III: 89 T r i g o n i p e s III: 4 2
D e c o n i c a I V : 28 L e p i o t u l a V : 109 P l e u r o p u s I: 82
D e l i t e s c o r I V : 28 L e p i s t a III: 6 7 P l e u r o t o p s i s III: 158 U r c e o l u s III: 158
D e r m o l o m a III: 3 0 L e p t o g l o s s u m III: 3 9 Pleurotus II: 2 0 U r o s p o r e l l i n a III: 168
D i c t y o l u s III: 39 L e p t o n i a I: 85 P l u t e o s p o r a I: 7 7
L e p t o t u s III: 39 PluteusII: 31 V o l v a II: 5 6
E c c i l i a I: 85 Leucoagaricus V : 85 P o l y m a r a s m i u s III: 136 Volvariella II: 5 7
E c h i n o d e r m a V : 109 Leucocoprinus V : 76 P o r p o l o m a I V : 149 V o l v a r i o p s i s II: 56
E n t o l o m a I: 85 Leucopaxillus III: 7 6 P o u z a r e l l a I: 85 V o l v a r i u s II: 56
F a e r b e r i a II: 3 0 P o u z a r o m y c e s I: 85
F a y o d i a III: 153 M a c r o c y s t i d i a III: 174 P r a t e l l a V : 23 X e r u l a I V : 181
F l a m m u l a I V : 80 M a c r o c y s t i s III: 174 P s a l l i o t a V : 23
169

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FLORA AGARICINA NEERLANDICA

Critical monographs on families of agarics and boleti occurring

A.A. B A L K E M A P U B L I S H E R S LISSE / ABINGDON / EXTON (PA) / TOKYO

Typesetting: Red Barn Publishing, Skeagh, Skibbereen, Ireland.

Index to the generic names i n volumes 1-5 169

Scope, methods and presentation

Introductory chapters Illustrations

N a m e s o f pteridophytes and phanerogams mentioned are i n accor- D r . T h . W . K u y p e r , A g r i c u l t u r a l University, Section S o i l B i o l o g y

LIST OF ABBREVIATIONS IN TEXT AND DRAWINGS

ad int. - ad interim nom.conserv. - nomen conservandum

lamellae (only used w h e n confusion w i t h c h e i l o c h r y s o c y s t i d i a deliquescent - (lamellae and/or basidiocarp) b e c o m i n g l i q u i d after

lichenized - l i v i n g i n s y m b i o t i c association w i t h an alga, f o r m i n g a widest part above ( V o l . 1, F i g . 34.32).

Abbreviations of authors' names in this volume

A . & S. - J . B . v o n A l b e r t i n i & L . D . von S c h w e i n i z Bres. - G . Bresadola

Eyre - W.L.W. Eyre C . H . K a u f f m . - C . H . Kauffman

M o s . & Jul. - M . M . Moser & W . R B . Jiilich J. Schaeff. - J . Schaeffer

a. Books tif des champignons superieurs de l a zone subalpine du Pare

Naturgetreue A b b i l d u n g e n und Beschreibungen der essbaren, R y m a n & H o l m a s e n , Svampar - Svampar.

Ber. naturf. Ges. Augsburg - Bericht der naturforschenden Cryptog. M y c o l . - Cryptogamie, M y c o l o g i e .

Mycologist - The Mycologist; Mycologist. Rev. catal. M i c o l . - R e v i s t a Catalana d i M i c o l o g i a .

1. Pileus covered with dark (grey-)brown to purple-brown fibrils or squamules

3. Spores > 6.5 mm long; average spore length 6.9-8.3 u,m

V e i l c o m p o s e d o f hyphae w i t h usually c y l i n d r i c a l , 3.0-8.0 |i.m w i d e

Pileus not discolouring, sometimes y e l l o w i n g ; v e i l usually present as

w o u l d c a l l A. bisporus. T h e interpretation o f L a n g e is generally f o l - Pileus 45-85(-95) m m , at first convex, expanding to plano-convex to

Pileus 4 5 - 8 5 ( - l 15) m m , at first convex to hemispherical, expanding to

F i g . 4. Agaricus pampeanus. (habit X A).

w h e n y o u n g , tapering towards base, sometimes c y l i n d r i c a l , straight,

P i l e u s (25-)40-60(-110) m m , at first convex w i t h flattened centre, 6. Agaricus porphyrocephalus F. M l l e r i n F r i e s i a 4: 204. 1952. - F i g .

F i g . 6. Agaricus cupreobrunneus. (habit X A). F i g . 7. Agaricus porphyrocephalus. (habit X 'A).

Agaricus lividonitidus (F. M 0 l l e r ) P i l a t i s very close, and perhaps a

F i g . 8. Agaricus bitorquis. (habit X 'A).

lus smooth, b e l o w annulus fibrillose, sometimes w i t h velar sock form-

ma. Stipitipellis a cutis o f unbranched, irregularly arranged, 2.5-5.5

Pileus (35-)60-120(-130) m m , at first hemispherical or conico-convex,

c o m m o n , widespread. M a y - N o v . C o m m o n and widespread i n Europe.

The cultivated variant w i t h the almost white pileus is sometimes dis-

11. Agaricus devoniensis P.D. Orton i n Trans. B r . m y c o l . S o c . 4 3 : 173.

X (9-) 12-20 m m , annulate, c y l i n d r i c a l to subclavate, slightly attenuat-

appressed radially fibrillose, w i t h m u c h adhering sand. L a m e l l a e , L =

H A B I T A T & DISTR. - U s u a l l y gregarious, saprotrophic, terrestrial on,

T h e differences between Agaricus pseudovillaticus (= Psalliota vapo-

14. Agaricus bohusii M . B o n i n D o c . m y c o l . 13(49): 56. 1981. - F i g .

slightly curved, deeply rooting w i t h pseudorhiza, stuffed to narrowly

F i g . 15. Agaricus bohusii. (habit X A).

appearing q u i c k l y , i n base o f stipe rarely y e l l o w i s h b r o w n . S m e l l like

The identity o f the taxa Agaricus sylvaticus var. pallidus (F. M 0 l l e r ) F.

16. Agaricus impudicus (Rea) Pilat, K l i c urc. H u b hfib. bedl.: 403.

er p i n k i s h b r o w n (5 Y R 6-7/4,7.5 Y R 6/4), finally b l a c k i s h brown, with

hyphae w i t h c y l i n d r i c a l , sometimes up to 14 m m inflated elements,

T h e above description is also based o n a c o l l e c t i o n from Denmark.

P i l e u s 4 5 - 1 0 5 m m , at first h e m i s p h e r i c a l to convex, e x p a n d i n g to Pileus 50-60 m m , convex, w i t h inflexed margin, slightly exceeding

F i g . 22. Agaricus arvensis. (habit X A).

F i g . 23. Agaricus augustus. (habit X A).

S E L . D E S C R . & F I G S . - H e i n e m . i n B u l l , trimest. S o c . m y c o l . Fr. 81: very pale-coloured variant w h i c h was p r o v i s i o n a l l y named by M o s e r

elements, w i t h very pale intracellular pigment, with scattered slightly

Agaricus nivescens and A. osecanus are considered conspecific, for an

23b. var. squarrosipes ( B o n & C o l l i n ) N a u t a i n Persoonia 17: 460.

C H A R A C T E R I S T I C S - Differing from the typical variety i n the covering

24. Agaricus pseudoumbrella B o h u s i n M i k o l . K o z l e m . 34: 26. 1995.