Fungi of Temperate Europe

The wheels
Thomas Læssøe & Jens H. Petersen
 Fungi of Temperate Europe is one of the most comprehensive my-
cological guides ever published. Featuring more than 7,000 photo-
graphs, this lavish two-volume set treats more than 2,800 species
of fungi across the region.

Including agarics, boletes, chanterelles and morels but also more obscure
groups such as cyphelloids, cup fungi, pyrenomycetous fungi and
hysterioids, this guide takes an unprecedentedly broad approach to
communicating fungal diversity. All species are illustrated with one or
more photographs and information is given on morphology, ecology
and distribution within temperate Europe. Furthermore, 1,500+ species
are discussed as potential look-alikes. The books are divided into eighty
“form groups,” each starting with an innovative identification wheel with
guiding photos, distinguishing characteristics and drawings of essential
microscopic features. Poisonous and edible species are colour coded
within the text.
Revealing the world of fungi in all its splendour, Fungi of Temperate
Europe is a must-have resource for any amateur or professional
mycologist.

To distribute the identification wheels as far as possible, we have

created this digital version for use on PC, tablet and smartphone. The
wheels are linked so you may jump forward from the table of contents
and the main wheels and back again with a home button.

The digital wheels can be downloaded from the MycoKey website (www.
mycokey.com). They may be printed for personal use and for education,
but must not be distributed commercially.

Thomas Læssøe og Jens H. Petersen

Contents
General identification wheels. . . . . . . . . . . . . . 6 (42)
Chanterelles and the like. . . . . . . . . . . . . . . . . 10 (46)
Identification wheels for groups of agarics. . 12 (56)
Pleurotoids. . . . . . . . . . . . . . . . . . . . . . . . . . . 16 (60)
Clitocyboids . . . . . . . . . . . . . . . . . . . . . . . . . . 18 (98)
Omphalinoids. . . . . . . . . . . . . . . . . . . . . . . . 20 (128)
Hygrocyboids . . . . . . . . . . . . . . . . . . . . . . . . 22 (142)
Mycenoids . . . . . . . . . . . . . . . . . . . . . . . . . . 24 (176)
Tricholomatoids. . . . . . . . . . . . . . . . . . . . . . . 26 (226)
Collybioids . . . . . . . . . . . . . . . . . . . . . . . . . . 28 (274)
Marasmioids. . . . . . . . . . . . . . . . . . . . . . . . . 30 (302)
Cystoderma and the like. . . . . . . . . . . . . . . . 32 (314)
Lepiotoids. . . . . . . . . . . . . . . . . . . . . . . . . . . 34 (320)
Chamaemyces and Limacella. . . . . . . . . . . . . 36 (348)
Amanitoids. . . . . . . . . . . . . . . . . . . . . . . . . . 38 (352)
Russula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 (372)
Lactarioids . . . . . . . . . . . . . . . . . . . . . . . . . . 42 (414)
Entolomatoids. . . . . . . . . . . . . . . . . . . . . . . . 44 (448)
Pluteoids. . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 (486)
Agaricus and Allopsalliota. . . . . . . . . . . . . . . 48 (500)
Coprinoids . . . . . . . . . . . . . . . . . . . . . . . . . . 50 (520)
Psathyrelloids . . . . . . . . . . . . . . . . . . . . . . . . 52 (546)
Hypholomatoids. . . . . . . . . . . . . . . . . . . . . . 54 (580)
Gomphidioids and Melanomphalia. . . . . . . . 56 (596)
Pholiotoids. . . . . . . . . . . . . . . . . . . . . . . . . . 58 (600)
Little brown mushrooms (LBMs) . . . . . . . . . . 60 (616)
Inocybe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 (652)
Hebeloma. . . . . . . . . . . . . . . . . . . . . . . . . . . 64 (680)
Cortinarius . . . . . . . . . . . . . . . . . . . . . . . . . . 66 (690)
Paxillus and the like. . . . . . . . . . . . . . . . . . . . 68 (756)
Boletes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 (760)
Polypores . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 (824)
Polypores with a stem. . . . . . . . . . . . . . . . . . 74 (826)
Clustered polypores . . . . . . . . . . . . . . . . . . . 76 (846)
Perennial polypores. . . . . . . . . . . . . . . . . . . . 78 (854)
Annual, capped polypores. . . . . . . . . . . . . . . 80 (886)
Annual, resupinate polypores . . . . . . . . . . . . 84 (932)
Rosette-fungi and the like. . . . . . . . . . . . . . . 86 (952)
Corticioids. . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 (960)
Capped corticioids . . . . . . . . . . . . . . . . . . . . 90 (962)
Merulioid corticioids. . . . . . . . . . . . . . . . . . . 92 (974)
Fragile corticioids . . . . . . . . . . . . . . . . . . . . . 94 (986)
Smooth, firm, ± whitish corticioids. . . . . . . 96 (1000)
Smooth, firm, coloured corticioids. . . . . . . . 98 (1012)
Spiny corticioids . . . . . . . . . . . . . . . . . . . . 100 (1036)
Hydnoids. . . . . . . . . . . . . . . . . . . . . . . . . . . 102 (1054)
Cyphelloids. . . . . . . . . . . . . . . . . . . . . . . . . 104 (1076)
Clavarioids. . . . . . . . . . . . . . . . . . . . . . . . . 106 (1092)
Dacrymycetales . . . . . . . . . . . . . . . . . . . . . 110 (1154)
Jelly fungi. . . . . . . . . . . . . . . . . . . . . . . . . . 112 (1162)
Rusts and smuts. . . . . . . . . . . . . . . . . . . . . 116 (1198)
Exobasidiales. . . . . . . . . . . . . . . . . . . . . . . 118 (1214)
Bird’s nest fungi. . . . . . . . . . . . . . . . . . . . . 120 (1216)
Stinkhorns . . . . . . . . . . . . . . . . . . . . . . . . . 122 (1222)
Puffballs and similar gasteroid fungi. . . . 124 (1228)
Truffles. . . . . . . . . . . . . . . . . . . . . . . . . . . . 126 (1256)
Operculate cup fungi. . . . . . . . . . . . . . . . . 130 (1278)
Morels and the like. . . . . . . . . . . . . . . . . . 132 (1280)
Operculate cup fungi with dull colours. . . 134 (1290)
Yellow, orange, red to pink cup fungi. . . . 136 (1328)
Inoperculate cup fungi . . . . . . . . . . . . . . . 138 (1348)
Earthtongues and the like. . . . . . . . . . . . . 140 (1350)
Large, gelatinous inoperculate cup fungi. . . . . 142 (1364)
Downy to hairy inoperculate cup fungi. . . . 144 (1372)
High, smooth to felty inoperculate cup fungi. 148 (1404)
Low, smooth to felty inoperculate cup fungi. . 150 (1426)
Erumpent inoperculate cup fungi . . . . . . . 154 (1468)
Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 (1484)
Calicioids and mycocalicioids . . . . . . . . . . 158 (1494)
Mildews and cleistothecial fungi. . . . . . . 160 (1500)
Pyrenomycetous fungi . . . . . . . . . . . . . . . . . . 162 (1510)
Hypocrealean pyreno­mycetous fungi and others. 164 (1512)
Hard, stromatic pyrenomycetous fungi . . . 166 (1546)
Non-stromatic, dark, unitunicate pyrenos. . 168 (1582)
Bitunicate, non-hysterioid pyrenos. . . . . . . 172 (1602)
Hysterioid, pyrenomycetous fungi. . . . . . . 174 (1614)
Laboulbeniales. . . . . . . . . . . . . . . . . . . . . . 176 (1620)
Taphrinales. . . . . . . . . . . . . . . . . . . . . . . . . 178 (1622)
Asexual fungi. . . . . . . . . . . . . . . . . . . . . . . 180 (1626)
Mycetozoa. . . . . . . . . . . . . . . . . . . . . . . . . 182 (1646)
Blue text are links to the wheels.
42 overview of all major form groups overview of form groups – with external spore production 43

rosette fungi and clavarioids

Form groups and fungal wheels the like page 1092 cyphelloids
page 952 page 1076
The fungi that are included in the two
corticioids
volumes of this publication have been page 960
organized into groups of morphologically
similar fungi, which we have called ‘form jelly fungi
groups’. With some experience, these page1162
groups can be recognized with the naked Dacrymycetales
eye or with a hand lens. It is important page 1154
to remember that form groups do not
necessarily reflect natural relationships, tongue-
since morphology only reflects such shaped or
with flattened clavate or
relationships to a certain degree. For branches; coralloid
example, fungi with polypore morphology hydnoids with hanging, tiny
smooth under/
page1054 discs or tubes
have evolved multiple times and, as upperside
± rubbery; basidia rusts and smuts
a result, many of the taxa concerned like tuning-forks; page 1198
flat or effused- spores often
are not closely related – even though
reflexed; smooth, septate
they may look similar. The section on warty, wrinkled or
polypores in this book therefore covers spiny
all the species with polypore morphology, often gelatinous;
spores repeating
regardless of their current classification.
There is one major exception to
this general rule. We have divided the basidiomycotes
fungi, irrespective of their macroscopic stem and
basidia
spines parasitize plants
appearance, into the two major one-celled,
phylogenetic groups: the Basidiomycota ‘normal’
without
and the Ascomycota. At the end of with fruitbodies
Volume 2, we have also included sections deviating
on asexual fungi, and organisms that basidia
fruitbodies

➡
belong in other kingdoms but resemble (when present) with

➡
external spore
true fungi, e.g. the slime moulds.

ycotes
production
The fungal wheels are the key feature and active spore discoid to clavate

ascom
(meiospore) release apothecia;
of this publication. We have adopted tubes firmly
asci with lids
operculate
attached discomycetes
a highly pragmatic approach when page 1278
preparing these wheels, so that, for polypores
page 824 with tubes that
example, even if you find yourself looking discoid, slit-
can be loosened
at the wheel that covers cyphelloids from the flesh like to clavate
(Basidiomycota) (p. 1076), you will apothecia;
asci without
find cross-references to similar discoid lids
fungi in the Ascomycota (p. 1372). For
this reason, prior knowledge of fungal fruitbodies with
internal spore thalli with
systematics is not a prerequisite. production, without
fruitbodies enclosed
The wheels on this and the following with gills
with or without algae and/or
page spread are organized based on how active spore release cyanobacteria
without
the sexual spores are produced. In order fruitbodies
asci on living inoperculate
boletes host tissue; on discomycetes
page 760 leaves and fruits page 1348
funnel-shaped, asexual
hymenophore
smooth or veined
hymenium

hymenium lichens
page 1484

⬅
chanterelles asexual
Fruitbodies with external spore production on agarics page 46 fungi Taphrina © Thomas Læssøe & Jens H. Petersen
see next page spread Fungi of Temperate Europe,
an exposed hymenium, see also page 31. page 56 page 1626 page 1622 Princeton University Press, 2019
 44 overview of form groups – with internal spore production overview of form groups – with internal spore production 45

Eurotiomycetes
to decide which wheel to use, you will first
page 1500
have to determine whether your chosen asexual fungi
Laboulbeniales powdery mildews page 1626
fungus produces spores from an external (Erysiphales)
page 1620
hymenium (see p. 31 and the illustration page 1500
p. 42 bottom left). The hymenium consists non-stromatic
pyrenomycetous fungi
of a palisade of basidia or asci, and page 1510
typically forms a smooth surface on the
fruitbody. Fruitbodies with internal spore
production do not display such a surface.
Instead, you may find small openings
(ostioles), through which the spores can externally on living
be ejected. Form groups with external insects and other on leaves and
arthropods stems; parasitic
production and active dispersal of the
sexual spores (meiospores) from hymenia
on plant litter,
are found on the previous page spread, cleistothecia-like feathers, hair
calicioids
(chasmothecia) page 1494
and groups with internal spore production etc.
are covered on this page spread. perithecia
On this page spread, the groups (incl. pseudothecia, cleistothecia conidia dry or in
thyriothecia slimy masses
with spores that are actively released are
& hysterothecia)
shown at the upper/left side; the groups almost closed
with fruitbodies that have passive spore fruitbodies with a without
round or slit-like fruitbodies
dispersal, or have a powdery or slimy inner opening very small with
spore mass, are represented in the lower/ dusty heads;
with active mainly on wood
right side of the wheel. For passive spore spore release and bark

tes
puffballs and the like
dispersal to take place an external force is stromatic

yco
pyrenomycetous fungi page1228

m
often required, e.g. rain drops or animal

o
page 1510

asc
activity.

➡ ➡
The wheels here and throughout this larger,
fruitbodies with
publication can be used in various ways. without with powdery
internal spore
active spore contents
By checking the images in the outer ‘rim’, production with or
perithecia without active spore (meiospore)
a quick overview can be obtained of the release

s
release

ycote
perithecia in a
variation in the fungal groups covered.
fleshy structure –

asco
Images in the pale blue sections (see, e.g.,

iom
a stroma; surface initially ± onion-

my
the wheel on page 46) represent fungi dotted with

sid
shaped, then

cot
ostioles

ba
that in some way recall species from the splitting star-like

es ➡
form group covered by the wheel, and
which are treated in other wheels in this ➡
➡
spores in
publication; the relevant page number ➡ mix small egg-like
is given for ease of cross-referencing. structures
(peridioles)
The inner part of the wheel shows earthstars
other features that could be considered, and the like
particularly microscopic characters such as page 1228
spore morphology (see, e.g., the wheel on bird’s nest fungi
page 60 and 176). page 1216
We recognize that the wheel approach foetid spore
to fungal identification will never reach mass; insect-
fruitbodies with external dispersed
the same level of accuracy as a traditional
spore production ± subterranean;
analytical key. The distinguishing features stinkhorns
and active spore release animal-dispersed
and the like
of some fungi are not always clear-cut, page 1222
and several wheels and species accounts
may need to be consulted in order to
reach a satisfactory conclusion. However,
we hope that the approach taken in this
publication will be easier to use than

⬅
trying to follow the often highly technical
keys found in other identification guides.
truffles
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe, page 1256
Princeton University Press, 2019 see previous page spread
46 chanterelles and the like
Chanterelles and the like
The form group have rather fleshy
fruitbodies with a ± central stem and
a deeply decurrent hymenophore that
may be completely smooth or consist
of ± branched veins to wrinkles. The
fruitbodies can be massive or hollow.
The core group is Craterellus and
Cantharellus. These genera belong
in Cantharellales and both form
ectomycorrhiza.
Craterellus
7 species, page 47
Cantharellus
5 species, page 50
fruitbody ± hollow; fruitbody
outer side smooth ± massive;
outer side
or veined
veined
chanterelles and the
like are ± funnel-shaped,
with a stem and a smooth
to veined hymenophore
Craterellus – hollow chanterelles 47
Gomphus clavatus, which – some omphalinoids have a veined
belongs in Gomphales, also forms hymenophore, but are smaller with
ectomycorrhiza, while Faerberia reduced stems (page 128).
carbonaria, which is situated in – funnel-shaped hygrocyboids and
Polyporales, is a decomposer. omphalinoids have decurrent gills
(pages 128 & 142).
Similar groups: – some mycenas (e.g. Hemimycena)
– clitocyboids have similar shapes to may look similar, but are small, fragile
the chanterelles, but have typical gills and white (page 176).
that may be forked (page 98).
Further reading: 50, 92, 231, 242.
Gomphus
1 species, page 54
Micro-drawings:
spores and
cystidia.
Approximate
Faerberia
species number
1 species, page 55
applies to
temperate
Europe. 7 mm
Chanterelles within the genus
clitocyboids
Craterellus have a smooth or veined
page 98 outer side where the hymenium is
fruitbody
situated. They also have a ± hollow
large and massive; stem.
outer side ± violet
and veined
•• Craterellus cornucopioides is
fruitbody greyish;
outer side veined;
a thin-fleshed, grey-black to brown,
on burnt ground rarely yellow-brown, trumpet-
shaped fungus with an undulating
omphalinoids and
hygrocyboids
margin and with an almost smooth,
pages 128 & 142 grey hymenium, which is strongly
decurrent. The stem is hollow; the cap
surface is felty and the smell is very
gills narrow, pleasant. The smooth, hyaline spores
simple or bifurcate measure (9–)10–13.5(–16) × (5.5–)6.5–
8.5(–9.2) µm. Forms ectomycorrhiza
Arrhenia
page 64
with mostly Fagus and Picea on better,
gills ± thick typically ± clay-rich, calcareous soils.
Craterellus cinereus ▷▷ has obvious
fruitbody wrinkles. The two species occasionally
mostly without stem; occur close together. A yellowish form
gills reduced to veins
has been separated as C. konradii
Hemimycena & (lower image).
small, thin and Delicatula Widespread and locally common,
whitish, gills mostly page 176
becoming scarcer towards the north;
reduced
mostly September–October.
similar
7 mm
groups small, thin and
brownish; Jan Vesterholt
Cotylidia © Thomas Læssøe & Jens H. Petersen
outer side page 958 Fungi of Temperate Europe,
smooth Princeton University Press, 2019
56 overview of pale-spored agaric groups 57
marasmioids
page 302
Agarics collybioids
I+ = may be amyloid page 274
The gilled mushrooms comprise a I+ (dex.) = may be tricholomatoids Cystoderma
very large group of macrofungi, with dextrinoid page 226 and Cystodermella
page 314 lepiotoids
more than 3,000 species in temperate page 320
Europe. They are characterized by
having gills under a cap, always held in mycenoids
page 176
the vertical postion. Most species also
have a stem. In the pleurotoid agarics
the stem is reduced or absent.

Spore colour small to medium, rather

tough; gills adnexed gills mostly adnexed;
The colour of spore-deposits is a to adnate ± tough, may revive
key character for dividing the gilled robust, after drying
medium-sized with grainy
mushrooms into manageable groups: surfaces
to large; gills
white, cream to yellow (this page ± emarginate (universal veil)
I+
spread) or brownish, rose-brown to
black (next page spread). In addition, a I+ I+
with free gills;
few species have green or lilac spore- slender, mostly small
and fragile; mostly with often with Limacella and
deposits (also next page spread). convex caps; often with partial veil Chamaemyces
amyloid spores page 348
Veil
In some agarics the gills develop freely with free gills; amanitas
with slimy universal page 352
– a naked development. Others have I+
spore-deposit veil
I+
a protecting tissue over the young white to cream (dex.)
gills – a partial veil. Some have the I+
(dex.)
entire fruitbody covered in an universal
veil. E.g. Amanita phalloides has a with thick,
wax-like with free or, rarely,
membranous version of both veil types: I+ adnexed gills and
gills
hygrocyboids agarics membranous
the partial veil is seen as a ring on the page 142 with ± soft-fleshed universal veil
stem at maturity, while the universal fruitbodies with a cap
veil is seen as a volva at the stem I+ and with gills below
base. In Cortinarius the two veil types small and
fragile, with spore-deposit
can be thread-like and in others, e.g. white to
decurrent gills with grainy
Gomphidius, they are present as slimy yellowish surfaces;

othlours
layers.

co
universal veil
ornament I+

er s
I+
Phaeolepiota

por
Gill attachment other page 314

e
similar
The way the gills attach to the stem is fungi
an important character for separating omphalinoids fragile;
page 128 fleshy, flesh with
species and genera. The gills may be with decurrent spherical cells;
free (not touching the stem), adnexed, gills without latex
adnate, emarginate or decurrent.

fragile; flesh with

spherical cells;
without or with with latex
free gills reduced or
eccentric stems

with decurrent wrinkles,

chanterelles, page 46

adnexed and broadly

adnate gills
clitocyboids
page 98
Lactarius and
Lactifluus

emarginate gills
⬅
page 414 Russula
pleurotoids page 372
page 60 leathery, broadly © Thomas Læssøe & Jens H. Petersen
short and deeply attached fruitbodies
decurrent gills next page spread Fungi of Temperate Europe,
polypores, page 824 Princeton University Press, 2019
58 overview of dark-spored agaric groups overview of dark-spored agaric groups 59

hypholomatoids gomphidioids and

psathyrelloids page 580 Melanomphalia I+ = may be amyloid
Microscopical characters page 546 page 596 I+ (dex.) = may be
Many species of agarics have cystidia dextrinoid
on the stem (caulocystidia), cap
surface (pileocystidia) or on the gills. pholiotoids
The gill cystidia are divided into those coprinoids page 600
page 520
in the hymenium (pleurocystidia) and
Agaricus and
those on the edges (cheilocystidia). Allopsalliota
The cystidia are very important in page 500
species identification. It it advisable
to use a dissection microscope when
gills adnexed to emarginate;
making preparations, to ensure that gills adnexed to
flesh rather tough; cap
the appropriate part of the gill is adnate, often
yellowish or slimy
marbled; gills
studied. flesh very decurrent
Agarics have a huge range of fragile
spore types: smooth, angular, to gills free and
deliquescent cap greasy-slimy
spiny, thin- to thick- or double- or scaly;
walled, with or without germ pore. Hebeloma
on wood
page 680
Some spores stain ± blue with I+
Melzer’s reagent (termed amyloid (dex.) small to large,
pluteoids gills free and ± smooth, with
or I+). When studying microscopical persistent
page 486 I+ brownish colours;
characters always check for amyloidity (dex.) mostly with radish-
I+
and surface ornaments (use oil (dex.) like smells; on soil
immersion). I+
spore-deposit dark (dex.)
brown to ± black
Relationships rather small, with
The agarics constitute a ‘form group’ fibrillose caps;
gills free spores grey-brown
– an artifical assemblage. The biggest and smells often Inocybe
monophyletic groups of agarics are spermatic; on soil page 652
the Agaricales, the Russulales and agarics spore-deposit
with ± soft-fleshed grey-brown to
the Hymenochaetales each of which spore­
fruitbodies with a cap
deposit red-brown Cortinarius
include many species that are not entolomatoids ± brownish-
with gills below
page 448 page 690
agarics, e.g. corticioids, clavarioids, rose small to large; veil
puffballs and polypores. thread-like, spores
I+
(dex.) rust-brown;
on soil

w
Other similar fungi:

-yello
en
gills adnexed,

gre
emarginate to spore-
– chantarelles do not have proper gills lilac to

nish
decurrent deposit
but, instead, sinuose, branching veins brown I+
white to

brow
(dex.)
or wrinkles (page 46). yellowish with adnexed,
emarginate to short,
– certain polypores have gill-like decurrent gills;
hymenophores, but these structures tricholomatoid, with mostly on soil
are normally very tough (page 824). emarginate gills and
sweetish smells
with central
stem and deeply
decurrent gills
with free gills

with grainy without or with

surface; reduced stem
universal veil and/or gills

Lepista
page 249
little brown
mushrooms
(LBMs)
page 616
Paxillus
and the like
page 756

⬅
lepiotoids
page 320
Phaeolepiota
page 314 © Thomas Læssøe & Jens H. Petersen
previous page spread pleurotoids Fungi of Temperate Europe,
page 60 Princeton University Press, 2019
60 pleurotoids Sarcomyxa pleurotoids 61
1 species, page 74
Hohenbuehelia
Resupinatus 12 species
Pleurotoids Arrhenia 3 species page 71
Neofavolus
5 pleurotoid species page 70 1 species
The pleurotoid agarics are either fan- Micro-drawings: Panus page 79
page 65 2 species
or tongue-shaped and lack or have an spores and cystidia.
page 75
excentric to lateral stem. Some forms Approximate species Chaetocalathus
are attached on the backside, with number applies to 1 species Neolentinus
temperate Europe page 67 2 species
the gills forming a radiating pattern. page 81
Some have the gills reduced to veins
or wrinkles, or even to a ± smooth rather large; Lentinus
small to large; ± tough; partially 2 species
hymenophore. Species with a smooth rather tough; rather large;
Cheimonophyllum small; rather yellowish page 80
hymenophore and no stem are treated 1 species rather tough;
mostly felty or greenish ± tough;
Lentinellus
base lilac, felty;
under the cyphelloid fungi (page 1076). page 67 fragile; ± felty 6 species
without veil with skeletal tough; gills with
Most species in this group are page 76
hyphae hyphal pegs;
decomposers and the majority grow smells of aniseed;
somewhat
on wood, even though some are tough; with skeletal
Panellus hyphae

us
also nematophagous. A few occur tomentose tough; gills

ino
5 species I+ serrated;
on herbaceous stems: Gloiocephala,

lat
page 68

sm
with skeletal

us
somewhat

all
Campanella, and some of the

– ge
o
hyphae

atin
tough;

to
Hohenbuehelia species, one species of

no

ge
without veil

el
tough; gills

lar
tg

lar

tg
Len­­tinellus, one Clitopilus, several

ge
serrated;

ela

– no
–g

o
with skeletal

t
Entoloma, some Deconica and a few Tectella

ll t
ino
somewhat Phyllotopsis

ela
1 species hyphae

ge
us;

sma
Crepidotus species.

tin
tough; 1 species

lar
page 69

sm

ou
without veil page 82
Pleurotoid agarics have different

all

s
I+ gill edge entire,
spore-deposit colours. The majority spore colour I+ yellow Pleurocybella
somewhat
are white- to cream-spored, but a tough; (I+)
1 species
large; gills page 83
smaller group is rose- or brown-spored with veil
smooth-edged,
and Pleurotus ostreatus has a greyish- with g
ills; sm simple, not
Gloiocephala all – not gel anastomosing
violet spore-deposit. The different 4 species witho atinou pleurotoids
small and somewhat ut or s
with at have no stem or a
spore colours reflects the fact that the page 66 tough; hymenophore ypical
gills
± excentric to lateral large, ± tough; gills
species concerned have completely smooth to veined;
stem. Decomposers of smooth-edged,
mostly hirsute from
different phylogenies. The spore colour wood or herbaceous anastomosing;
cystidia (use lens)
stems with skeletal hyphae
is used in the division of the groups in Pleurotus
the wheel. rather fragile; 6+ species
hymenophore page 84
Some species of pleurotoids are +/- spores longitudinallly
smooth to veined ridged; with a
remarkably tough, e.g. Panus and farinaceous smell
Neofavolus and also older fruitbodies
of Lentinellus and Pleurotus. Others, Arrhenia rather tough; other
5 pleurotoid species hymenophore similar
e.g. Schizophyllum, Hohenbuehelia page 64 reticulate I+ fungi spores angular Clitopilus
and Sarcomyxa, have gelatinous or in outline 1–4 pleurotoid
rubbery flesh. rather tough; species, page 88
Campanella hymenophore
1 species wrinkled large; without
Other similar fungi: page 63 tough; obvious
– the cyphelloid fungi have small with split remnants of veil;
‘gills’ thick-fleshed Entoloma
fruitbodies without gills and no or
without obvious ≈ 5 pleurotoid species
hardly any stem. They are typically page 88
remnants of veil;
attached on the upperside of the small to medium,
fruitbody (page 1076). with
small; with ± stemless and rather
without triangular thin-fleshed
– the clitocyboids have central stems, Plicatura remnants of
thread-like veil;
stem thin-fleshed
but may recall pleurotoids (page 98). 1 species and gills veil; small
page 63 cyphelloids
page 1076
Further reading: 11, 65, 66, 156, 181,
203, 223, 224, 245, 295.
tough and
stemless Tapinella
Crepidotus 2 species
with ≈ 20 species page 96
Schizophyllum
± central page 90
3 species
page 62 stem
Deconica
2 pleurotoids
page 95
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe, clitocyboids polypores Pleuroflammula
Princeton University Press, 2019 page 98 page 824 1 species, page 96
 Leucopaxillus Aspropaxillus
98 clitocyboids 5+ species 1 species clitocyboids 99
page 107 page 110
Lepista &
Clitocyboids Paralepista
13 species
Clitocyboid agarics have short to Pseudoclitopilus page 112
deeply decurrent gills. The cap is 2 species Omphalotus
Notholepista page 106 Bonomyces Ampulloclitocybe
umbonate, applanate or funnel-shaped 1 species 1 species 1 species
page 111 1 species
and the flesh is mostly obviously page 106 page 111 page 116
fibrillose but not very tough. Some
species form small fruitbodies, but
most are medium-sized to very large.
Most clitocyboids have white to cream
spore-deposits, but in some species
belonging to the genera Clitocybe and
Paralepista they are clay-pink.
Catathelasma large to very
Almost all clitocyboids are 1 species gills large; gills moderately Infundibulicybe
large; 10+ species
decomposers of litter or soil. Exceptions page 105 detachable detachable gills page 114
with pink, gills detachable yellow;
include Ossicaulis and Omphalotus detachable
gills
on wood distant;
that are lignicolous. Hygrophoropsis gills
± on burnt distinctly
and Aphroditeola may also grow on with pruinose, soil funnel-shaped,
remnants of wood. Catathelasma yellowish cap 1 species with
forms ectomycorrhiza with conifers. umbo
Rhizocybe
The generic division of the with a (I+) 3 species
I+ with soft,
clitocyboid agarics has gone through membranous page 118
I+ clavate stem Omphaliaster
large changes, with many species, veil 3+ species
Pseudoomphalina
I+ page 117
previously accepted in Leucopaxillus & Pseudolaccaria
2 species as a large with
and Clitocybe, now being assigned to a page 103 Omphalina with echinulate
number of new genera. amyloid spores
Pseudoclitocybe spores
2 species I+ I+
Other similar fungi: page 104
with striking
tall and slender clitocyboids
– omphalinoids may look rather with grey- I+ hyphal cords;
have ± decurrent
similar, but are mostly more fragile brown colours mostly in spring
gills and umbonate,
and smaller. Some have a biotrophic I+
applanate or funnel-
flesh shaped caps. Spore- whitish, smell
assosciation with mosses, etc., others deposit white to Ossicaulis
gelatinous farinaceous;
are decomposers (page 128). clay-pink on wood 2 species
– pleurotoids likewise have deeply page 119
Myxomphalia
decurrent gills but have excentric 1 species, page 103 flesh I+ stem whitish
gelatinous and gills brown-
stems. Many of these (e.g. Neolentinus
other grey
and Lentinellus) have distinctly tough similar fungi
gills whitish;
flesh (page 60). bifurcate, with
– entolomatoids may be shaped like Hohenbuehelia reddening characteristic
clitocyboids, but have brownish-rose, 1 species with central stem odours
(see page 73) remaining
verrucose, striate or angular spores gills bifurcate tough or with species with
and smell sweet excentric to lateral Atractosporocybe
(page 448). white, cream to
stem 1 species, page 119
clay-pink spore-
– Paxillus and others are shaped like deposits
clitocyboids but have brownish spore- Cantharellula ± orange with
1 species bifurcate gills
deposits (page 756).
page 102
with
chantarelle-
Litteratur: 8, 9, 10, 27, 110, 156, 174,
like wrinkles
181, 183, 340, 344. small and fragile or with
thick, fleshy gills brownish-rose,
brown or Leucocybe &
black spores Singerocybe
3 species, pages
119 & 120
Micro-drawings: Aphroditeola
spores first, 1 species
followed on the page 101
inside by the Clitocybe &
cystidia Clitopaxillus
Hygrophoropsis 30+ species
Approximate species page 121
3 species
number applies to
deep and short decurrent gills page 100
temperate Europe
© Thomas Læssøe & Jens H. Petersen chanterelles, pleurotoids, omphalinoids & entolomatoids & Paxillus
Fungi of Temperate Europe, page 46 page 60 hygrocyboids pages 448 & 756
Princeton University Press, 2019 pages 128 & 142
128 omphalinoids omphalinoids 129

Omphalinoids Arrhenia Micro-drawings:

Rickenella 23 species spores and gill
The omphalinoid fungi are a loosely Contumyces cystidia.
3 species page 135
defined form group of white- page 134 1 species
spored agarics with relatively small, page 139 Approximate species
number applies to
mostly dry, somewhat fragile and temperate Europe.
elegant fruitbodies and rather deeply
decurrent gills. Although the name Cantharellopsis Hydropus
1 species 1 omphalinoid species
suggests that all are umbilicate, this page 184
page 133
is far from the case. They can also
be convex, umbonate or applanate reddish, greenish, blueish,
greyish, brown or blackish; pink,
without an umbilicus. coloured; stem,
on soil or deciduous wood with cystidia;
cap and gills hairy
Some omphalinoid fungi are from cystidia
on soil
biotrophic, either as lichenized with Hodophilus
whitish, ± with spores
algae or as parasites on mosses pinkish-buff cap amyloid; cystidia 10+ species
(left side of the wheel). Others are unbranched; on soil page 164
centre
decomposers (saprotrophs) of litter, Arrhenia
23 species
herbs or wood (right side of the page 135
decom small, brown to
wheel). pose
ytes
yellowish; with
rs o I+
cap mostly
ryoph ns thick gills and small
Other similar fungi: with minute b oil spores; on soil
scales on or
– some hygrocyboids (Hygrocybe Omphalina

wo
and others) with decurrent gills look 16 species

od
page 139
distinctly omphalinoid. However, the

± brown or

um
hygrocyboid fungi have, in general, cap honey- white; on soil

agn
very slender basidia (6–9 times as Gyroflexus coloured; gills

on Sph
1 species whitish
long as wide) and live mostly as page 132
Omphalinoids
biotrophs amongst phanerogams have rather small and gills thick, yellow
(page 142). fragile fruitbodies or grey-rose;
– some mycenas with decurrent gills with decurrent, on soil or conifer Chromosera
Loreleia mostly thick gills and wood 5 species

on liverw
are also omphalinoid. Many mycenas 2 species cap convex, applanate or page 148
can be separataed by their amyloid page 133 orange-brown umbillicate caps
spores, but e.g. Phloeomana speirea gills

orts
and related species and the genus orange-yellow to
greenish-yellow; on
Hemimycena are inamyloid (page other conifer wood
176). cap on similar fungi
– the clitocyboids have predominantly pinkish-buff to lich
honey-coloured nse with shaggy
decurrent gills, but are mostly larger margin and
and tougher than the omphalinoids orange colours; on Chrysomphalina
lich hardwood 2 species
(page 98). eniz page 141
ed
– some species of entolomatoids also Blasiphalia
1 species yellow-orange
look similar, but these have brownish page 132
on Peltigera with orange
rose spore-deposits, and the spores with basal felt;
are ridged or angular (page 448). brownish-rose, with thick with gracile spores I+
with robust
angular/ridged fruitbodies gills fruitbodies; Haasiella
spores and long spores 2 species
Litteratur: 156, 181, 262. Arrhenia with a thallus at page 140
basidia mostly I+
23 species the stem base
page 135

I+/- I+

deeply decurrent gills Xeromphalina

Lichenomphalia page 185
© Thomas Læssøe & Jens H. Petersen 4 species
Fungi of Temperate Europe, entolomatoids clitocyboids hygrocyboids mycenoids
Princeton University Press, 2019 page 130 page 448 page 98 page 142 page 176
 142 hygrocyboids hygrocyboids 143

Hygrocyboids Micro-drawings:
The hygrocyboid agarics (waxcaps and spores.
Hygrocybe
others) are recognized by their thick, 35+ species
Neohygrocybe Approximate
3 species
wax-like and mostly rather distant gills, Gliophorus
page 150
page 160
species number
applies to
and many species have very vivid colours. 5+ species Chromosera temperate
Microscopically, most hygrocyboids have page 145 5 species Europe.
page 148
unusually long and slender basidia,
typically 6–9 times as long as wide. The
spore-deposits are whitish and the spores
smooth and inamyloid; as a general rule
cystidia are lacking.
Most hygrocyboids are thought to
be biotrophic, with a poorly understood
symbiosis with herbs. Using DNA-analysis
techniques, living hyphae have been Cuphophyllus
10 species
detected inside tissues and seeds of page 161
Plantago. Species of Hygrophorus small to large,
small, slimy to dry,
are also proven to be biotrotrophic, omphalinoid, smooth or scaly;
but they form ectomycorrhiza with a lubricous to dry with vivid colours
with decurrent large, ± greasy,
number of woody partners. One species gills with dull colours
(Hygrophorus exiguus ⋉) appears to be and adnexed gills
Gloioxanthomyces medium-
associated with Tricholoma mycorrhiza. 1 species sized, slimy,
Most hygrocyboid agarics have very page 149 with dry or
demanding habitat requirements. They slimy gill
edge
are particularly sensitive to fertilizers small to large,
and many favour habitats with a long greasy, mostly with
continuity. The hygrocyboids share small, slimy, dull colours and
with slimy ± decurrent gills
the same habitats and have similar thread on gill
preferences to a number of clavarioids, edge
entolomatoids and earthtongues; they
hygrocyboids
are all good indicators of sites of special
include white-spored
nature conservation value. small, dry, with
agarics with thick,
dull colours and
In most of Europe the typical habitat large, with wax-like gills and
short decurrent,
conical, rose- mostly slender, long
for hygrocyboids, except Hygrophorus coloured basidia; spores I-
rather thin gills
and Hodophilus, is old, unfertilized greasy cap
grassland – a habitat that has declined Camaro­phyl­lopsis
small, greasy, with
dramatically over the past 50 years. In 1 species
other dull colours and
other parts of the world, the hygrocyboid page 165
similar fungi decurrent, thick,
agarics are predominantly to be found in rather distant gills
forests with long continuity. Species of Porpolomopsis spore-deposit spore-deposit
Hodophilus tend to prefer thorny thickets 1 species whitish reddish; spores
page 144 angular
on clay soils.

rather large, ± slimy
Other similar fungi: with floccose stem
– omphalinoids may also have thick, apex; ectomycorrhizal
fleshy gills; many are parasites on I +/- Hodophilus
10+ species
mosses or are lichenized, but some are fruitbodies mostly ± funnel- mostly with normal thin gills;
page 164
shaped with decurrent gills; spores mostly I+; mostly with
saprotrophs (page 128). spores I- hymenial cystidia
– mycenoids generally have ‘typical’
gills and most have cystidia and amyloid
spores. They are all believed to be
saprotrophs (page 176).
– entolomatoids may be colourful, but
Hygrophorus
have angular, pinkish spores (page 448). 40+ species
page 166
Litteratur: 1, 2, 3, 43, 57, 156, 172,
178, 183.
© Thomas Læssøe & Jens H. Petersen omphalinoids mycenoids entolomatoids
Fungi of Temperate Europe, page 128 page 448
Princeton University Press, 2019 page 176
176 mycenoids mycenoids 177

Mycenoids Resinomycena
1 species Roridomyces Hemimycena
Mycenoids are small, rather fragile or Micro-drawings: 2 species
page 186 30+ species
somewhat tough agarics with white spores and cystidia. page 187 page 188 Mycena
Phloeomana
spore-deposits. Most species have a 150+ species
Approximate species 4 species
Atheniella and the like page 196
bell-shaped or convex cap, but may count applies to page 192
4 species
temperate Europe. Delicatula
become applanate or even somewhat 1 species pages 193, 194
funnel-shaped with age. The gill page 186
attachment comes in all forms, except
completely free. Smell and surface
Xeromphalina
features – dry versus slimy etc. – are 4+ species
important characters. In the field it page 185
is important to note whether the
downy from cap cuticle
fruitbodies have a distinctive odour, capitate cellular and
8 species
small and page 196
e.g. radish-like, like iodoform or cystidia dry; stem whitish; gills
nitrous, and whether parts of the gills vein- with with thick normal or greyish to
slime-coat whitish,
adhering reduced. brownish;
fruitbody are slimy. like; cap with
material spores I-
orange to rose-
a quickly coloured;
The mycenoid fungi often have disappearing
tough, spores I-
smooth, amyloid spores. Many species brownish- veil
I+ ra
have characteristic cystidia in the orange with ne
ge
Hydropus basal orange I- oi
d
hymenium or in other places. felt; gills I+ en en
a
5 species
8 species yc yc
The mycenoid fungi are mainly page 182 decurrent I- er
m
s
m page 198
th nu large;
decomposers and often occur on leaf o
e
ge
smell of radish
cap hairy or th
or needle debris, on dead wood or on innately fibrillose I+ I+ I-
the bark of living trees. A few species (cuticle a rectocutis
(e.g. Mycena galopus, page 205 ) may or a trichoderm) I+(-)
coloured
form mycorrhiza-like associations with
I+/- gill edge
ericaceous plants.
cap shiny or innately I+
Mycena is the most speciose fibrillose (cuticle
Fayodia
mycenoid genus in temperate Europe, 3 species a rectocutis); I+ mycenoids with
and the next is Hemimycena. The page 181 stem ± smooth
have small, rather latex
latter has inamyloid spores and the fragile, ± bell-
I- shaped fruitbodies 5 species
fruitbodies are small or very small and
cap shiny or innately and white spores page 203
whitish; some are without gills and
fibrillose (cuticle
form a link to the cyphelloid fungi stem
a rectocutis);
with
(page 1076). stem ± smooth I- basal disc

Other similar fungi: Gamundia

– omphalinoids and hygrocyboids 1 species cap cuticle a bluish colours or
with decurrent gills may look rather page 180 hymeniderm, surface similar on living bark
pruinose; white-spored
similar (pages 128 & 142). stem ± downy groups
– collybioids and marasmioids may very small, 9 species
pale or grey page 206
also look similar, but these are mostly
5 species
somewhat tougher or can revive after ± tough fragile, mostly page 209
desiccation (pages 274 & 302). with thick, wax- with
Mycenella decurrent gills
– similar entolomatoids have rose- 4 species like gills remaining
coloured spore-deposits (page 448). page 178 not desiccation greyish
desiccation to brown
– similar Panaeolus species have tolerant tolerant
mycenas
blackish spore-deposits (page 554).
– similar little brown mushrooms 6 species
(LBMs) have grey-brown to red-brown page 212
spore-deposits (page 616).

Further reading: 12, 15, 104, 156, 183,

206, 272, 273.

85+ species 15 species

© Thomas Læssøe & Jens H. Petersen omphalinoids
page 218 page 213
Fungi of Temperate Europe, marasmioids collybioids and hygrocyboids
Princeton University Press, 2019 page 302 page 274 pages 128 & 142
226 tricholomatoids tricholomatoids 227

Lepista
Gerhardtia
Tricholomatoids 1 species
13 species
Micro-drawings:
page 249
The tricholomatoid fungi are rather page 244 Melanoleuca spores first,
Tricholomella Dermoloma 20+ species Pseudoporpoloma thereafter cystidia.
fleshy agarics with ± solid stems, 1 species 3 species page 246 1 species cy. = cystide.
± emarginate gills and whitish page 244 page 245 page 253
Approximate
or, rarely, brownish-rose spore- species number
deposits. The genera Tricholoma and Calocybe applies to
10 species temperate Europe.
Catathelasma form ectomycorrhiza, page 240
while species of Squamanita are
parasites on other agarics. The Pseudotricholoma
remaining genera are decomposers cap
1 species, page 253
(saprotrophs), although Armillaria may ± vinaceous- cap cuticle elegant
kill trees and bushes before degrading brown; completely and mostly
fugaceous, on rich without fibres; umbonate;
the wood (necrotrophs). Hypsizygus stem ± Pogonoloma
thread-like to soil farinaceous with caulo-
2 species floccose; smell 1 species
The tricholomatoid fungi have membranous cystidia
page 239 cap cuticle sweetish cap conical, page 253
partial veil
rather few reliable characters for completely warts fibrillose; ±
I+ hygrocyboid
separating the genera. Besides the without
fibres; ± cap cuticle
spore morphology and the presense of farinaceous without
veils, identification very much depends I+/- fibres;
upon experience and gut feeling. It is, farinaceous
Floccularia
cap ± smooth; de
for example, relatively straightforward mostly high de
co
m 2 species
co po
for an experienced field mycologist to on hardwood m se cap margin page 252
po rs I+
se on strigose-hairy;
recognize the elegant melanoleucas Megacollybia rs
so smell soap-like
on il
as such, but it is very difficult to & Clitocybula w I+
cap radially oo
put into words the macroscopical 1 species d
cracking; mostly
page 238
differences between a Melanoleuca with thick I+ cap ± squarrose
hyphal cords; from remnants
and a Lyophyllum. on hardwood of veil

Other similar fungi: tricholomatoids

have rather large to cap smooth, soft-
– the large, brown hygrocyboids in cap scaly; gills large, fleshy fruitbodies fleshy, grey-brown
the genus Neohygrocybe are rather Tricholomopsis yellow; with ± emarginate
4 species on conifer wood gills and whitish, rarely
similar but have thick, wax-like gills
page 236 brownish-rose spore-
(page 160). deposits large; with
– collybioids may look very similar, with veil; cap veil and basal
Leucocortinarius
but are mostly more fragile with centre ± dark, I+ bulb
1 species
hollow stems, and typically have scaly; mostly on page 257
wood other
narrowly adnate gills (page 274). similar fungi
– species of Entoloma with roboust cy gills ± decurrent; Lyophyllum
+/- 15+ species
fruitbodies are very similar, but have cap veined; with membranous
Armillaria & double veil page 254
d

ec
pinkish, angular spores (page 478). Desarmillaria on deciduous I+ oo

to
w
wood

m
7 species s

yc
– Hebeloma and Cortinarius may
on

ric

o
page 232 j

rr
a
rs

ag

have the same stature, but brown

hi
+/-
se

z
with threadlike

al
po

on

gills emarginate;
spores (pages 680 & 690).
om

veil; cultivated; spores spores

ite

with or without
c

as

on deciduous whitish coloured

de

veil; on soil
r
pa

wood
Further reading: 28, 64, 88, 111,
Catathelasma
156, 157, 181, 183, 285, 342, 343, squarrose; page 105
Rhodotus on wood
345. 1 species mostly scaly;
page 231 with thick gills
lower stem; thick and spores angular, ± spores
on fungi wax-like gills normal brownish-rose verrucose, brown

Lentinula
1 species
page 230

Leucopholiota
1 species Tricholoma
types of page 230 67 species, page 258
© Thomas Læssøe & Jens H. Petersen Hebeloma &
emarginate gills Squamanita hygrocyboids collybioids entolomatoids Cortinarius
Fungi of Temperate Europe,
Princeton University Press, 2019 7 species, page 228 page 160 page 274 page 470 pages 680 & 690
274 collybioids collybioids 275

Macrocystidia Clitocybula
Callistosporium
Collybioids 2+ species
1 species and the likes
page 288 4 species
The collybioids are characterized by Micro-drawings: page 289 (see page 238)
convex or umbonate to applanate, silky- spores, followed on
the inside by cystidia.
fibrillose, greasy, hairy or slimy caps, Chl. = chlamydospores Gymnopus
Myochromella
adnexed to broadly adnate gills and † = dead. Tephrocybe 2+ species 30+ species
white spore-deposits. Rhodocollybia 1 species page 287 page 290
Approximate species Lyophyllum page 286
has a pale grey-rose deposit and count applies to 25+ species
Macrocystidia may have a ± brown temperate Europe. page 287
deposit. Most species are moderately Sagaranella sordid yellow-
Rhodocollybia
tough. The spores are, with a few 4 species 5+ species, page 296
stem brown;
exceptions, inert to iodine. page 287 floccose, dead spores
farinaceous have orange stem and gills with
The collybioids function deeply
or not; ‘drop’; on soil projecting cystidia;
rooting;
predominantly as decomposers. A strongly
on soil or wood smell of linseed; on
soil or wood chips cap surface
number of species have long, rooting farinaceous; radially cracking;
stems, which arise from buried Sphagnurus on soil and on soil on wood
1 species burnt †
substrates such as tree roots or buried ground gills distant
page 286
cones. Laccaria differs by being to crowded;
important ectomycorrhizals. cap greasy;
spores verrucose; on litter
In older literature many collybioid rather mycenoid;
species were included in the genera on soil gills ± crowded;
Lyophyllum, Collybia and Xerula. cap greasy; spores
in fairy rings on ± thick-walled and
However, molecular phylogenetic Sphagnum deposit pale grey-
studies have shown that these genera rose; on litter
Pseudobaeospora
are all strongly polyphyletic, and this has 14+ species
Xerula
± silky-fibrillose; I+/- 2 species
led to much ‘splitting’, as can be seen page 284 (dextrinoid) gills distant;
mostly with thick page 298
cap and stem
from the many unfamilar names. gills; mostly on
hairy;
calcareous soil I+/-
collybioids from roots
(dextrinoid) +/÷cy.
Other similar fungi: are rather tough,
on wood,
– mycenoids with narrowly adnate gills Hydropus have convex to
mostly from gills distant;
I+/- applanate caps, Paraxerula
may look very similar. However, they ≈ 8 species buried twigs
narrowly to broadly cap pruinose; 1 species
mostly have campanulate-umbonate page 182 adnate gills and, from roots page 298
caps, tend to be more fragile, and most on cones mostly, whitish
or wood; gills I+ spore-deposits
species have amyloid spores (page 176). very crowded
gills distant,
Baeospora mostly dark-edged;
– marasmioids are rather similar to the 2 species cap greasy;
collybioids in shape and habitat, but page 283 spores large;
on conifer
most can tolerate desiccation. The stem from roots
cones
Chl. other
is typically darker with a pale apex. They
similar fungi cap greasy-slimy;
are also somewhat tougher (page 302). stem dark,
Strobilurus on fungi;
– tricholomatiods are more fleshy and with conidia
tomentose;
3 species Hymenopellis
have ± emarginate gills (page 226). on wood
page 282 on rotten 1 species
– entolomatoids have brownish-rose fungi; two page 299
species cap slimy;
spore-deposits and angular spores (page stem with ring;
with brown
448). sclerotia with on wood
– LBMs (little brown mushrooms) may side
branches cap mostly
have collybioid shapes, but are brown- and black campanulate-
spored (page 616). sclerotia umbonate; gill
with thick, attachment variable tough and revive
Asterophora wax-like gills; after drying; gills
Further reading: 4, 13, 24, 25, 30, 156, 2 species ectomycorrhizal narrowly to broadly with
181, 183, 246, 348. page 281 adnate or with coloured
Flammulina
collarium spores
6 species
Collybia page 300
3 species
page 279

Dendrocollybia
1 species Mucidula
page 280 1 species
Laccaria entolomatoids page 301
adnexed to broadly © Thomas Læssøe & Jens H. Petersen 9 species
Fungi of Temperate Europe, mycenoids marasmioids and LBMs
adnate gills Princeton University Press, 2019 page 276 page 176 page 302 pages 448 & 616
302 marasmioids marasmioids with a garlic smell 303

Marasmioids Mycetinis is recognized by the smell

The marasmioid fungi have applanate
to convex, rather tough caps and
highly variable gill attachments,
including a collarium around the stem
(page 312). The stems are mostly dark,
except for the apex, and some species
have strong odours. The fruitbodies
revive after desiccation. Microscopically
many species have cystidia with finger-
like protuberances.
The marasmioid fungi can be
Rhizomarasmius
Mycetinis 2 species
3 species page 304
page 303
very difficult to distinguish from
the collybioids, especially the genus
Gymnopus (pages 290 & 306).
Likewise, Xeromphalina (page 185)
creates a bridge between marasmioids
and mycenoids.
Other similar fungi:
– collybioids usually have adnexed gills
and the stems are mostly not darker
than the caps (page 274).
Marasmiellus
11 species
page 305
– marasmioid species without stem or
with excentric stems are to be found
amongst the pleurotoids (page 60).
Further reading: 13, 156, 181, 202.
adnexed, narrowly adnate, broadly
adnate and decurrent gills
of stale garlic or old rubber-bands.
Mycetinis alliaceus is a large, tough,
strongly garlic-smelling marasmioid
with a very dark, finely felty-downy,
stiff stem. The gills are pale and
rather distant. The spores measure
7.5–11 × 6–8 µm. Almost exclusively
on trunks and buried twigs of Fagus.
Mycetinis querceus ▽ is browner
and occurs directly on leaf-litter.
Mycetinis alliaceus var. subtilis ⋉ is a
rather common dwarf form with a
pale, marginally striate cap.
Widespread and very common in
parts of temperate Europe, rare or
absent from others; May–January. 20 mm

Marasmius & Mycetinis querceus is a strongly

Cryptomarasmius garlic-smelling, large, pale marasmioid;
20+ species
page 308 the stem becomes felty towards
Crinipellis
2 species
the base. The spores measure
page 307 Gymnopus p.p. 7–10 × 4–5 µm. Occurs mostly on
30+ species Quercus leaf-litter late in the season
page 306
(often Q. rubra), but also on Fagus
leaf-litter.
Mycetinis alliaceus △ has darker
stems, grows on wood and, in
comparison to M. querceus, has
cheilocystidia. Mycetinis scorodonius ▽
odourless; gills has a reddish, smooth stem. Similar
smell of garlic; ± decurrent; species of Gymnopus (pages 290 &
on wood, herbs on leaves or odourless;
or leaves Pteridium gills adnexed to 306) are either odourless or have a
odourless;
adnate cap and stem cabbage-like smell.
strigose-felty; Widespread, but mostly rather
mostly on herbs
uncommon; mainly October–January. 15 mm
some with
cabbage-like Gloiocephala
page 66 Jean-Marc Moingeon
smells; gills ±
adnate Mycetinis scorodonius is a medium-
gills sized marasmioid with a shiny, reddish
adnexed or
joined to a stem, pale cap and a strong garlic
collarium around smell. Spores 7–10 × 3–5 µm. Occurs
the stem
in open, dry places, e.g. in dunes,
stem excentric, and can be found on e.g. stems of
± reduced; Artemisia campestris and similar
gills mostly
reduced coarse herbs, but occasionally also on
Xeromphalina wood, e.g. in Syringa hedgerows.
page 185 Crinipellis scabellus ▷▷ occurs in
yellow-orange
with basal felt; the same habitats but has projecting
I+ gills decurrent
hairs on the stem and is odourless.
Marasmius oreades ▷▷ is less red and
marasmioids
have adnexed, adnate has a different smell. Other Mycetinis
to decurrent gills or are species have ± felty stems, are larger
collariate and most are with reduced and more bound to woodland.
relatively dark-stemmed. stem rather similar, but Micro-drawings:
They are tough and not tolerant of spores outermost, Widespread and rather common;
desiccatiom 5 mm
tolerate desiccation other then cystidia. mainly July–November.
similar fungi
Species numbers © Thomas Læssøe & Jens H. Petersen
Chaetocalathus collybioids applies to tempe­ Fungi of Temperate Europe,
see the pleurotoids, page 60 page 274 rate Europe. Princeton University Press, 2019
314 Cystoderma and the like Phaeolepiota 315

Cystoderma and the like

This group is characterized by the
granulose surface of the cap and
stem that constitutes remnants of
a universal veil. In some species a
ring or ringzone is present on the
stem. The gills are adnexed, adnate
or emarginate and the spore-deposit
is either whitish or brownish-yellow.
The spores are amyloid or inamyloid
and smooth. The flesh may contain
asexual spores. Some species have
hymenial cystidia.
Other similar fungi:
– the lepiotoids have completely free
gills (page 320).
Left: young caps. Right: expanded caps
Further reading: 156, 181, 324. with remnants of veil at the margin.

Phaeolepiota
1 species 315

adnexed to broadly adnate or

Cystoderma emarginate gills
7 species
page 316 20 mm

Phaeolepiota aurea is a large,

fleshy, completely dry, orange-
yellow agaric with a large, flaring
to hanging ring, the upperside of
which is ± coloured by the brownish-
yellow spores. The entire fruitbody is
mealy-grainy. The gills are adnexed.
The odour may be somewhat like
cyanide or bitter almonds. The spores
very large;
smells of cyanide; Cystodermella are fusiform, smooth, and measure
spore-deposit 3 species 11–12 × 4.5–5.5 µm. Occurs on rich,
brownish-yellow page 318 often disturbed, soils, e.g. in fertilized
whitish, yellowish lawns or stands of nettles, typically in
to ± brown; troops and fairy rings.
spores amyloid 8 mm
Could possibly be confused with
the ± fibrillose Gymnopilus spectabilis
(page 602) but that species has
I+ ornamented spores and grows on
wood. It may also recall an enlarged
whitish, ±
brownish-red Cystoderma ▷▷ or Cystodermella ▷▷
to orange-red; but the spore-deposit colour is
spores inamyloid distinctive. In Japan it is parasitized by
a species of Squamanita (page 228),
which may reflect its close relationship
with Cystoderma, a common host for
this parasitic genus of agarics.
Widespread and locally common,
more or less absent from the boreal
Cystoderma and the like zone; mainly September–November.
have a grainy universal Micro-drawings:
veil, which is left as a spores.
coating and a ring zone.
Decomposers on soil or other Approximate species
rarely decayed wood similar fungi number applies to © Thomas Læssøe & Jens H. Petersen
with free gills
temperate Europe. Fungi of Temperate Europe,
lepiotoids, page 320 Princeton University Press, 2019
320 lepiotoids lepiotoids 321

Lepiotoids Leucocoprinus Leucocagaricus

The lepiotoids have whitish or greenish Echinoderma 5+ species 20+ species
9 species page 336 page 337
spore-deposits and free gills but the page 334
universal veil is less membranous in
comparison to the amanitoids, p. 352.
rather small; thin-
In some species the universal veil is fleshed; with ring, mostly
powdery or granulose, sometimes with ± plicate margin
in the form of small spines. Often
have complex, muff-like rings formed small to medium-
by parts of both the universal and sized; mostly with
well-defined, muff-
the partial veil. The spores are often Lepiota like ring
dextrinoid. 35+ species
page 326 medium-sized;
The lepiotoids are decomposers mostly with
and are mostly found on black, mull ring; cap with
soils, e.g. in damp, rich woodlands, pyramidal scales
along road-sides with Urtica, in scrub
I+ (dextrinoid),
and gardens, and on old compost. small; mostly without with a germ pore
Some species of Macrolepiota, ring, often with girdles; with a hyaline
with scaly, cracking cap cover
Chlorophyllum and Lepiota also grow
in open dry grasslands, including I+/-
(dextrinoid)
dunes. I+/- (dextrinoid),
with or without a germ
The smaller lepiotoids apparently pore; one species has a medium-sized to large;
need a specific climatic scenario in hyaline cover with well-defined,
spore colour muff-like, loose,
order to produce fruitbodies, and I+
(dextrinoid) mostly double ring
years may pass without significant
fruiting. When this does happens it is
often over in a week or two. Cystolepiota
11 species, page 324 small and
powdery; without lepiotoids
Other similar fungi: well-defined ring have free gills with whitish I+ (dextrinoid),
– the amanitoids are similar but have a I+/- or greenish spore-deposits, thick-walled, with a germ Macrolepiota
(dextrinoid) and non-membranous pore with a hyaline cover
membranous universal veil (page 352). 7 species
universal veils page 345
– Limacella and Chamaemyces may
look similar but have a greasy/slimy
cap surface (page 348).
– Cystoderma and others have
adnexed to emarginate gills (page other
I+/- (dextrinoid), thick-
walled, with an uncovered
314). similar fungi germ pore
– Floccularia has adnexed to
emarginate gills (page 252). small and
powdery; without medium-sized to large;
– Agaricus (page 500) has black well-defined ring; with well-defined,
spores. Young fruitbodies of e.g. A. gills red or green muff-like, loose,
mostly double ring
sylvicola, with immature spores, can Melanophyllum
be recognized by the sweet marzipan 2 species, page 322
smells. Reddning species may recall
species of Chlorophyllum, but these spore-deposit white; spore-deposit white; spore-
gills adnexed to gills free deposit
have more complex rings. emarginate black;
gills free
Further reading: 6, 85, 156, 183, 226, surface surface with membranous
332. granulose greasy/slimy universal veil
cap with
attached Chlorophyllum
scales 5 species
page 341

Micro-drawings:
spores.

Approximate species
number applies to
free gills temperate Europe.
© Thomas Læssøe & Jens H. Petersen
tricholomatoids Cystoderma and the like Limacella/Chamaemyces amanitoids Fungi of Temperate Europe,
page 252 page 314 page 348 page 352 Agaricus, page 500 Princeton University Press, 2019
348 Chamaemyces and Limacella Chamaemyces 349

Chamaemyces and Limacella

These genera are characterized by ±
free gills, white spore-deposits and the
absence of a membranous universal
veil, although some species may
have a slimy universal veil. They have
membranous or thread-like partial veils
left as a ring or ring zone on the stem.
The cap surfaces are smooth, do not
crack, and are mostly greasy to slimy.
Chamaemyces and Limacella are
decomposers, which occur mostly in
exceptionally mull-rich habitats in scrub
and forests. Most, perhaps all, species
are associated with calcareous soils.
Chamaemyces is closely related
to the lepiotoids (page 320), while
Limacella is relatively close to the
amanitoids (page 352).
Other similar fungi:
– the amanitoids have ± free gills and a
membranous universal veil that covers
the greasy cap (page 352).
– volvariellas have free gills, a
membranous universal veil seen as
a volva, and a brownish-rose spore-
deposit (page 487).
– the lepiotoids have free gills and
powdered, plicate, scaly or cracking,
dry, cap surfaces (page 320).
Further reading: 90, 156, 181.

free gills

Micro-drawings:
spores outermost,

Thomas Læssøe
then cystidia.

Approximate species
number applies to 10 mm
temperate Europe.
Chamaemyces
1 species Chamaemyces fracidus recalls
page 349 something between an Amanita and
a Limacella, but it is easily recognized
by the grainy-scaly covering below
the ring-zone and by the wet,
mostly guttulate, convex, pale buff
cap, which may have remnants
of a veil at the margin. The stem
cap greasy to dry; is also ± covered in yellow-brown
stem below the drops. The whitish gills are free to
ring-zone with almost free. The odour is strong, ±
yellow-brown
drops; spores gas-like or sour. Has clavate cheilo-
smooth and pleurocystidia. The spores are
smooth, inert to iodine, measure
4.5–5.5 × 2.5–4 µm, and lack a germ
pore. Occurs on calcareous soils in
Limacella deciduous forests (typically Fagus),
cap slimy to greasy; 6 species
stem below the ring page 350 mostly in sites with numerous other
zone dry or slimy; rare species of fungi.
spores smooth or without
This species also goes by the
finely verrucose membranous
universal veil; name Lepiotella irrorata. It may recall
with powdery or Echinoderma hystrix (page 335), but
I+/- with cracking cap
(dextrinoid) membranous
that species has pyramidal, brown
universal veil scales on the cap. Farther south a
darker type, Chamaemyces fracidus f.
pseudocastanea, can be found.
Limacella and Rather rare and local, absent from
Chamaemyces
have free gills, white the boreal zone; mainly July–October.
spore-deposits and
smooth, mostly greasy
to slimy caps other
similar fungi 5 mm

Martin Kříž
© Thomas Læssøe & Jens H. Petersen
Volvariella amanitas lepiotoids Fungi of Temperate Europe,
page 487 page 352 page 320 Princeton University Press, 2019
352 amanitoids Saproamanita 353

Amanitoids
The amanitoids are recognized by free,
or more rarely, adnexed gills, whitish
or somewhat greenish spore-deposits
and, almost always, a membranous
universal veil (velum), which, at
maturity, may be seen as a volva at
the base of the stem and/or as loose
patches on the cap. The spores may be
amyloid or inamyloid.
Most of the amanitoids form
ectomycorrhiza and are, therefore,
usually found in forests and parks. The
genus Saproamanita is saprotrophic.
Amanita is a species-rich genus
that includes some of the most
notorious poisonous mushrooms, not
least the ringless species (Amanita
sect. Vaginatae) which form a very
confusing group where specific
identification can be difficult.
Other similar fungi:
– the lepiotoids have either a powdery
universal veil, or it is absent (page 320).
– limacelloids are rather similar, but
have a slimy universal veil (page 348).
– volvariellas are separated by their
brownish-rose spores (page 486).
Further reading: 149, 156, 183, 218,
263, 281.
universal veil
partial veil

Saproamanita
3 species
page 353

25 mm
amanitas free to adnexed gills
with a ring Emilio Pini
20+ species Saproamanita are decomposing
page 355 Amanita-like agarics. They usually
occur in open land and mostly form
fairy rings.

Saproamanita vittadinii is a
magnificent, ± white to pale buff
amanitoid with highly projecting
amanitas scales on the cap and stem. The
with a partial veil without a ring
that mostly is left as
stem also has white to pale buff
(sect. Vaginatae)
a ring on the stem 20+ species, page 365 belts along almost its entire length;
(mostly central to it lacks a bulb but does have have a
southern Europe)
with a partial veil
complex ring. The gills are tinted buff
that mostly is left as or green. The spores are ellipsoid,
a ring on the stem amyloid and measure (9–)10–13

decomposers (–15) × (6.5–)7.5–10(–11) µm. Occurs
in fairy rings without connections to
ec
to
mycorrhizal trees or scrub.
m The mycorrhizal Amanita
yc
o r
solitaria ▷▷ is very similar, but has
rh

without partial
iza

veil or ring
echinulate scales on the lower part of
ls

the stem.
Mainly a southern species
lepiotoids
page 320
but reaches as far north as the
I+/- Chamaemyces/ Netherlands and central England;
Limacella autumn.
page 348

amanitoids

Emilio Pini
have free or adnexed
gills and a membranous
universal veil other
similar fungi © Thomas Læssøe & Jens H. Petersen
volvariellas Fungi of Temperate Europe,
page 486 Princeton University Press, 2019
372 russulas russulas 373

Russula More than 200 species in mild, pale-spored russulas

temperate Europe. page 386
Russula is a genus of fleshy agarics
with fragile flesh, and spores that have
amyloid ornaments. The spore-deposit is
white to orange-yellow.
A difficult genus. Among the most
important macroscopical characters
are the cap colour, the colour of the jade-russulas
spore-deposit, the taste and the smell. ≈ subgen. Heterophyllae
However, the cap colours can be variable, page 382 acrid, pale-spored russulas
since the pigments are water-soluble and page 390
easily washes out. The spore colour must taste mild; taste mild
be judged by obtaining a thick spore- cap with brown, to moderately acrid; with
deposit. The taste is fairly constant; some grey-rose, greyish or yellowish to reddish colours
greenish colours
species are burning hot like strong chilli
but for the milder species it is necessary
to chew on a suitably large sample for
taste
some time. There exist also truffle-like distinctly acrid
species of Russula (see page 1266).
cap ± buff
Russulas form ectomycorrhiza. Some fetid russulas
subgen. Ingratae to honey-yellow;
species may grow with many different margin sulcate-striate;
page 379 m
tree species, while others are ± limited to smell mostly unpleasant rea
oc
et ow
hit ell
partners of certain genera. It is therefore res
w
uff
-y
spo a l eb
important to note which ectomycorrhizal res
p
spo
trees and bushes are present in the with
lon
g gil
vicinity of a fruiting Russula. Mild-tasting with
sho
rt a
ls o
nly
nd l
russulas are considered edible but only a ong
gill
s
few are really choice edibles. russulas taste mild; smell of
are fragile agarics shellfish; stem browning
without milk (green with FeSO4)
Other similar fungi:
spores
– lactarioids have a similar fragile flesh pale b
uff-y ellow
spores
cap greyish to
and spores with similar ornaments, but brownish;
buff-
to or
ange-
yello
the flesh exudes a white or coloured latex flesh reddening
w
shrimp-russulas
when cut (page 414). or blackening subgen. Xerampelinae

ls
ea
m page 396

ng gil
– Paxillus and Gomphidius can have to
cr
llo
w
h ite -ye

ly
fragile flesh, but the spores are brown to w g e

and lo
s n

ls on
re ra
spo to o taste mild
black (pages 596 & 756). ite to moderately acrid
wh

ng gil
es

short
spor

Further reading: 76, 92, 151, 156, 165,

ith lo
with
191, 228, 276, 287, 288. charred russulas

w w
subgen. Compactae cap whitish, ± funnel-
low
page 376
yello
shaped; flesh not uff-yel
reddening ange-
or blackening
pale b

taste
to or

distinctly acrid
spores
buff-

A1 B1 C1
spores

A2 B2 C2

A3 B3 C3

Spore types (ornament stained with iodine reagents):

column A with isolated warts or spines. mild, dark-spored russulas
column B with somewhat connected warts or spines. funnel-russulas acrid, dark-spored russulas page 398
column C with reticulum (C1–C2) or crests (C3). ≈ subgen. Lactarioides page 408 © Thomas Læssøe & Jens H. Petersen
page 374 Fungi of Temperate Europe,
Princeton University Press, 2019
414 lactarioids lactarioids 415

fishy lactifluii
Lactarioids (Lactifluus p.p.)
page 416
The genera Lactarius and Lactifluus are lilac-staining lactarii
recognized by their fleshy fruitbodies page 434
with milk. They have the same fragile
flesh as the genus Russula, and similar foxy lactarii
spores with a very amyloid ornament page 428
of spines, reticulae or crests. Most
species also have large cystidia on the
gills. Some are mild-tasting but many
are very acrid.
The most important characters for yellow-staining lactarii
page 437
species identification are the colour
and stain of the milk, surface texture fleshy, with very
sticky lactarii ample and sticky
of the cap (including the degree of page 420 milk, and fishy
sliminess), overall colour, taste and smell
mycorrhizal partner. margin smooth;
milk and flesh
cap mostly pale,
All lactarioids are mycorrhizal with brown or lilac; brown milk on drying greyish-violet on
drying
trees and shrubs or, in the alpine zone, not sticky lk vio
g mi let
mi
with e.g. Dryas and Salix herbacea. yin lk
e on
The lactarioids are very closely gr d

or

ry
related to Russula, and in the

in g
g
in
tropics the genera are less easy to

ng
ch a
differentiate.

un
margin smooth;
With the exception of Lactarius

y el
milk at first white

te ,
cap mostly pale,

lo w
but quickly yellow

w hi
helvus, which smells like curry and is grey-brown or
on the gill flesh

i ng m
olive-brown,
slightly toxic, all mild-tasting European
greasy, sticky
lactarioids are edible. However, the to slimy

ilk
lactarioids
acrid lactarioids are toxic and some have fleshy fruitbodies
contain strong mutagens. with milk (latex) and
spores with an amyloid
cap ± felty, whitish;

æl k
ornament
Other similar fungi: fleecy lactifluii fruitbody large and

in g m
(Lactifluus p.p.) heavy
– some Mycena species have milk, milk at first white, pink-staining lactarii
page 417 page 440

den
but the stems are less than 2 mm later rose or only
on the gill flesh

re d
wide (page 203).
– in the subtropics/tropics other
agarics, e.g. Lactocollybia ⋉,
may have latex. i

lk
m
g e
an
Further reading: 92, 105, 152, 156, or
165, 228, 327. margin
milk rather sparse,
clear milk orange from the
bearded
start

with smell of curry

or stock cubes
A1 B1 C1

orange-milked
A2 B2 C2 lactarii
page 444

A3 B3 C3

Spore types (ornament stained with iodine-reagents):

Column A with isolated warts or spines.
Column B with somewhat connected warts or spines. More than 100 species bearded lactarii Lactarius helvus
Column C with reticulum or crests (C3). in temperate Europe. page 418 page 447
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
Princeton University Press, 2019
448 entolomatoids entolomatoids 449

floccose-scaly blue
Entolomatoids entolomas,
without clamps
This group includes the genus Entoloma Micro-drawings: page 459 floccose-scaly,
in a broad sense and the closely related spores and cystidia. pink or greenish
cy. = cystidia. entolomas floccose-scaly,
genera Clitopilus, Clitopilopsis, Clitocella, page 464 brownish
Rhodophana and Rhodocybe. They are Approximate species floccose-felty, entolomas
count applies to blue entolomas page 466
characte­rized by adnexed-adnate, emar­ with clamps
temperate Europe.
ginate to decurrent gills and ± brownish- page 457
rose spore-deposits. The spores are
either verrucose, striate-ridged or three-
dimentionally angular (faceted). An
impor­tant character of entolomatoids is omphalinoid
whether the pigment in the cap cuticle entolomas
is incrusted or intracellular. Species with page 455
clitocyboid
excentric or lateral stems are treated cap bluish; cap bluish; cap
entolomas
cap centre centre ± scaly;
under the pleurotoid fungi (page 88). page 454 cap rose or
scaly felty; stem mostly
Entoloma greenish;
Some species apparently have a stem mostly smooth
cap centre
200+ species coarsely cap
parasite-like association with plants page 454 ± scaly brownish;
fibrillose small,
in the Rosaceae, while other species slender, with cap centre ±
smooth stem mycen scaly
± smooth entolomas
around Entoloma rhodopolium are oid/c page 469
and decurrent olly

mycorrhizal. The ecology of many gills bio
id
species is still not known. oid small with ±
lin
All genera belong in the same family, fleshy-robust, with ha smooth,

p
fibrillose stem and greasy or

om
the Entolomataceae. decurrent gills shiny cap

id
ybo
itoc
Other similar fungi: Clitopilus

cl
10+ species gills decurrent;
– Pluteus, Volvariella & Volvopluteus with strong
page 453
have the same spore colour but free gills farinaceous cap whitish, hairy,
smell entolomatoids whitish entolomas
(page 486). and others
± hairy
+/-cy. page 475
– some species of Lepista have have ± brownish-
brownish-rose spores and tricholomatoid +cy. rose spore-deposits
gills decurrent and verrucose, striate-
fruitbodies. They are recognized by ridged or angular
small;
cap and
their minutely warty spores and a Clitopilopsis -cy. spores stem
characteristic sweetish smell (page 249). 1 species fibrillose-
page 452 gills decurrent; hairy
– some clitocyboids and the omphali­noid +cy.
cap ± cracking
Arrhenia discorosea have ± brownish-
-cy. +/-
rose, smooth spores (pages 98 & 138). cy.

tric
Clitocella rather large, cap

h
other

o
– the large collybioids in the genus 4+ species gills decurrent; with diverging

lom
page 452 cap ± cracking similar fungi
Rhodocollybia have somewhat brownish fibres; with unusual

ati
o
d
gills with brownish-rose cystidia
-rose, smooth spores (page 296). spore-deposits
fibrous to hairy, greenish
emarginate to brown entolomas
or short, page 476
rather large, with
Further reading: 45, 89, 156, 158, 181, decurrent
gills smooth, dry or ±
182, 204, 223, 224, 335, 336, 337, Rhodocybe emarginate greasy caps
338. 6+ species stem
page 451 excentric, gills gills gills
lateral or narrowly decurrent free
missing adnate
tricholomatoid,
Rhodophana fibrillose-hairy
3 species entolomas
page 450 page 478

adnexed, emarginate
pleurotoids Clitocybe & tricholomatoid, © Thomas Læssøe & Jens H. Petersen
and decurrent gills
page 88 Lepista Rhodocollybia Arrhenia Pluteus and others smooth entolomas Fungi of Temperate Europe,
page 249 page 296 p. 121 & 138 page 486 page 482 Princeton University Press, 2019
486 pluteoids Volvopluteus (cap smooth and shiny) 487

Pluteoids
Volvopluteus, Volvariella and Pluteus
are characterized by their free gills
and brownish-rose spore-deposits.
They are decomposers either of
wood or organic material in the soil.
One species, Volvariella surrecta, is a
parasite on fruitbodies of Clitocybe
nebularis.
Other similar fungi:
– Amanita has white spore-deposits
and the poisonous species have a
partial veil that is left as a ring on the
stem (page 352).
– Limacella is ± sticky and has white
spore-deposits and no volva at the base
(page 348).
– Leucoagaricus leucothites has pinkish
gills, white spores, and a ring but no
volva (page 339).
– entolomatoids have attached gills and
ridged or angular spores (page 448).
Further reading: 6, 89,140, 141, 156,
182.

Volvariella
Volvopluteus 9 species
1 species page 488
Pluteus free gills
page 487
30 species, page 491

15 mm

Volvopluteus gloiocephalus is a
plutei with fibrous caps large agaric with a smooth, greasy
page 491 to slimy, greyish cap surface, a
long, greyish stem and a rather
large, white volva (the remnants
of the universal veil) at the base.
The spore-deposit is brownish-rose.
Cheilocystidia clavate. The spores
are smooth and measure 10.5–
cap smooth and
shiny
17.5 × 6.5–9 µm. Mostly found in
cap fibrillose plutei with hairy caps disturbed places, including stubble
to hairy page 493 fields, compost heaps, bales of 7 mm
damp straw, etc. It can occur by the
thousands in stubble and in newly
sown fields late in the season.
The amanitoids (page 352)
usually have the same greasy-smooth
cap surface but they are generally
cy. cy. ectomycorrhizal and have white
cap cuticle cy.
radially fibrillose, spores; the most similar species,
sometimes with Amanita phalloides (page 359),
hairs; with hooked has a ring on the stem. Species of
cystidia
Volvariella ▷▷ are dry to somewhat
with universal veil/ cap cuticle hairy; sticky on the cap and always
volva at the stem without hooked distinctly radially fibrillose. Species of
base cystidia
plutei with cellular caps Pluteus ▷▷ lack a universal veil and
without universal page 496
veil/volva at the thus a volva.
cy.
stem base Widespread and common,
decreasing towards the north; May–
pluteoids
have free gills and December.
brownish-rose spore- cap cuticle cellular
deposits; with or without (possibly including
universal veil (volvas) scattered hairs); without
hooked cystidia © Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
Princeton University Press, 2019
500 Agaricus and Allopsalliota Agaricus and Allopsalliota 501

Agaricus and Allopsalliota royals Micro-drawings:

Agaricus
4+ species spores and
The genera Agaricus and Allopsalliota 40+ species, page 503
page 503 cheilocystidia.
are characterized by free, permanent,
non-dissolving gills and dark Approximate species
Allopsalliota number applies to
chocolate-brown spore-deposits. 1 species, page 502 temperate Europe.
Ink caps (Coprinus, Coprinopsis and
Coprinellus, page 520) also have ± free
gills and ± black spores, but their gills
normally dissolve with maturiy.
Molecular phylogenetic studies
have led to descriptions of a
large number of Agaricus species
that cannot be recognized by
morphological charac­ters. Using dwarfs
ring thick, double; 4+ species, page 506
traditional morphology, only 40+ fruitbodies rather large;
broadly defined species can be keyed spores 5–12 µm long
out in temperate Europe.
Agaricus and Allopsalliota are ring slender;
fruitbodies small; yellow stainers
decomposers that often occur in fairy spores 4.5–6.5 µm long 3+ species, page 508
rings.
Many species are edible but slowly yellowing;
the slowly yellowing species (the smell of aniseed or
almonds
royals) may contain high levels of
cadmium, and the yellow stainers can
cause stomach upsets. In addition, yellows, then
reddens and
most species contain potentially blackens; spore-
carcinogenic hydrazines (agaritin). end thick-walled Agaricus and
Species of Agaricus should therefore Allopsalliota
have free gills that immediately yellowing;
only be consumed in fairly small are pale grey or pink smell of carbol
(ink, phenols)
amounts and then only when cooked. gille adnate; at first but quickly
spore-deposits become dark brown
chocolate-brown
Other similar fungi: other
– Stropharia coronilla recalls a small, similar
yellowish Agaricus but has narrowly fungi
± brown or red-staining;
adnate gills (page 590). smell faint, sour or
Psathyrella Stropharia
– annulate species of Psathyrella have page 546 page 590 unpleasant
narrowly adnate gills (page 546). gills free;
spore-deposits
– similar lepiotoids have white spore- whitish
deposits (page 320).
– the very poisonous Amanita virosa
has a membranous universal veil and
a white spore-deposit, as do other
species of Amanita (page 357).

Further reading: 91, 147, 148, 156,

182, 226, 236, 237.

red stainers
18+ species, page 510

© Thomas Læssøe & Jens H. Petersen

amanitoids lepiotoids Fungi of Temperate Europe,
free gills
page 352 page 320 Princeton University Press, 2019
520 coprinoids coprinoids 521

Coprinopsis nivea group

Coprinoids page 532
Micro-drawings:
spores first,
The coprinoids include the genera Coprinus, Coprinopsis thereafter cystidia.
Coprinellus, Coprinopsis and Parasola, of 70+ species Coprinopsis urticicola group ve. = hyphae of the
page 523 page 530 universal veil.
which there are are more than 150 species
in temperate Europe. They are recognized small to medium; Approximate species
Coprinellus micaceus group,
by the black spores and the almost always cap powdery from page 537 number applies to
Coprinopsis lagopus group globose veil cells temperate Europe.
free gills. Many coprinoids also have a
page 526
unique maturation process. The spores small;
mature from the gill edges inwards and, cap with small
Coprinellus
flake-like remnants
as soon as the spores are released, that 50+ species
of veil; mostly on
page 537
part of the gill disintegrates and becomes herbs
cap ± orange-
liquid, and so the process continues. This brown, with
allows for very closely spaced gills, since thread-like and Coprinellus disseminatus group
the risk of spores being caught between spherical remnants page 540
of veil
them is reduced. Nevertheless, many
spores are still caught and produce the inky
liquid associated with the vernacular name
ink cap. The coprinoids can be grouped cap ± orange-brown,
hairy from projecting
according to the appearance of the surface cystidia (pielocystidia)
of the cap and veil, e.g. whether the veil is
grainy or thread-like. These characters are with thread-like
borderline between macro and micro, and a veil

sis

us
Coprinopsis

op
good hand lens is required. The coprinoids ve.

ell
rin
atramentarius group
may produce many different spore types,

rin
Cop
page 523

Cop
from smooth and ellipsoid to lemon-shaped,
heart-shaped or truncate and they may large, dark;
cap centre with
have prominent warts. They always have large flake-like coprinoids
a germ pore and occasionally a loosening, remnants of veil have free gills Coprinellus
colourless outer wall. All have cystidia. that dissolve with
age; the spores are Parasola
The phylogenetic and morphological black
differences between psathyrelloid (page Parasola
13+ species
546) and coprinoid fungi are not page 544
straightforward. Some members of
other
e.g. Coprinopsis do not have gills that similar fungi
dissolve and are treated among the
psathyrelloids. ps
is
no
All coprinoids are decomposers and ri
op us
C in
some coprophilous species have a very pr cap smooth, mostly
Co ± plicate; gills very
short life-cycle, fruiting just a few weeks free and very distant
after spore germination. All fruitbodies
are very short-lived. At least some of
the coprilous species are able to kill
competing mycelia in the substrate. spores
dark brown to black

Other similar fungi:

large, whitish; spores
– Psathyrella and the like mostly have with thread-like brown
adnate gills that never dissolve during cap scales;
maturation (page 546). stem with ring
– Bolbitius are slimy-capped and have
brown spore-deposits (page 618).
Coprinus
Further reading: 156, 182, 207, 208, 227, 2 species
page 522
290, 322.

Psathyrella little brown

© Thomas Læssøe & Jens H. Petersen and the like mushrooms (LBMs)
free gills Fungi of Temperate Europe, page 546 page 616
Princeton University Press, 2019
546 psathyrelloids psathyrelloids 547
© Thomas Læssøe & Jens H. Petersen
scaly red-margined Fungi of Temperate Europe,
Psathyrelloids Psathyrella (& psathyrelloids psathyrelloids psathyrelloids with
Princeton University Press, 2019
Coprinopsis p.p.) page 562 page 564 abundant veil
A group of very fragile agarics, most 75+ species
page 566
page 562
with black to dark brown, rarely more
grey-brown, spore-deposits, mostly dry remaining Psathyrella
caps with dull brownish colours, and Cystoagaricus species
2 species page 574
adnexed, adnate to shortly decurrent page 561
gills. A few species have prominent
stem rings but most have veil remnants
on the cap or, rarely, no veil at all. It
includes a number of genera formerly
treated in Psathyrella: Cystoagaricus,
Homophron, Kaufmannia, Lacrymaria with red line
cap dark-scaly; above the gill
and Typhrasa, as well as the ± classic with a ring edge
genera Panaeolus, Psilocybe, Deconica cap fibrillose- with distinct
and part of Parasola. scaly; cystidia veil covering
Kauffmania
Microscopically, most species have 1 species with or without
green drops in
spores with germ pores. In rare cases page 560
ammonia
the spores are ornamented (Lacrymaria,
large with
Panaeolus p.p.). Shape and contents of smooth cap and
the cystidia (cheilo- and pleurocystidia) sparse veil; spores
rather pale,
are important for species identification. without a germ in ammonia cap without
Ecologically, vary from lignicolous scales; gill edge
pore
not red;
over soil to dung-inhabiting, and one veil sparse
species is a parasite on Pluteus.
cap with
thread-like veil;
Other similar fungi: cystidia with
Typhrasa yellowish, psathyrelloids fleshy; veil
– coprinoids have ± dissolving gills have black to dark absent; spores
2 species oily inclusions
(page 520). page 560 brown spores, strikingly pale
– other black-spored agarics, e.g. fragile fruitbodies and cystidia
with non-free gills crowned and
Stropharia, Leratiomyces and Hypho­ and brown to thick-walled
loma), have mostly more vivid colours cap
greyish colours
and often slimy caps (page 580). fibrillose; gills
– Inocybe has grey-brown spore- marbled, edge hair
with drops
deposits (page 652). veil Homophron
absent; 2 species
– Agrocybe has somewhat paler, grey- other
page 578
similar fungi cap and stem
brown spore-deposits and mostly sticky base with
cap surfaces (page 625). brown, thick-
stem rigid; walled hairs
cap smooth; (hand lens)
Further reading: 86, 154, 155, 156, 171, required!)
gills marbled,
182, 225, 370. Lacrymaria some with Parasola
3 species drops 1 psathyrelloid species
page 558 page 579

with ± black with paler brown

stem base mostly spore-deposits spore-deposits
bluing; gills
± marbled

stem not mostly with

gills dissolving
bluing; gills sticky cap or
with age
± marbled yellowish colours
Panaeolus
12+ species Micro-drawings:
page 554 spores outside,
cystidia inside.

Approximate species
adnexed, adnate and
number applies to
decurrent gills
temperate Europe.

Psilocybe Deconica Hypholoma

6+ species 15+ species and the like coprinoids LBMs Inocybe
page 552 page 548 page 580 page 520 page 616 page 652
580 hypholomatoids hypholomatoids 581

Hypholomatoids © Thomas Læssøe & Jens H. Petersen Micro-drawings:

Fungi of Temperate Europe, spores and cystidia.
Princeton University Press, 2019
This group includes mostly fairly robust
agarics with dark chocolate-brown Approximate species
number applies to
to black spore-deposits and adnate, temperate Europe.
emarginate or slightly decurrent Stropharia
gills. The caps are mostly slimy and Leratiomyces 9 species
5 species page 590
the colours a rather vivid yellowish, page 588
greenish or orange-brown. With the
exception of Hypholoma, the gills are ±
marbled due to uneven maturation.
Microscopically, many of the species Protostropharia
have dark spores with a germ pore 4 species stem mostly
page 587 with small woolly-scaly
and many have cystidia with yellow cap-scales in
contents (chrysocystidia). However, slime
Hemistropharia
similar cystidia are also common in 1 species
the brown-spored genus Pholiota page 595
(page 608).
All species are decomposers. The
majority decay wood or wood chips,
but a few are decomposers of herbs or
Sphagnum. cap and stem slimy;
on dung
Hypholoma
Other similar fungi: 15 species
– psathyrelloids also have blackish page 582
spores but the fragile fruitbodies are with ring or ring zone; veil
mostly abundant;
less brightly coloured and the caps are gills marbled
mostly dry (page 546). gills with drops;
– gomphidioids are slimy with black veil prominent,
spores and deeply decurrent gills cottony
(page 596). hypholomatoids
are somewhat tough,
– pholiotoids may be both slimy and small to medium-sized,
yellow but have paler, grey-brown without dark-spored agarics
spores (page 600). ring; with mostly adnate gills
gills not and slimy or yellowish
– the group of little brown mushrooms caps other similar fungi
marbled
(LBMs) is separated by their paler
brown spore colours (page 616).

Further reading: 156, 169, 181, 225. spores ± black spores ± brown

mostly with
yellowish to
reddish colours

adnexed, adnate, emarginate

and shortly decurrent gills

psathyrelloids gomphidioids pholiotoids little brown mushrooms (LBMs)

page 546 page 596 page 600 page 616
596 gomphidioids and Melanomphalia Chroogomphus 57
597

Gomphidioids and
Melanomphalia fusiform, typical boletoid spores. The Other similar fungi:
genus Melanomphalia has no veils and – Paxillus and Phylloporus are similar
This group includes agarics with dark belongs to the Agaricales; it has finely but have brown spore-deposits (page
brown to black spore-deposits and dotted-verrucose spores. 756).
deeply decurrent gills. The two genera The species of gomphidioids are – Tapinella has a brown spore-deposit
of gomphidioids, Chroogomphus probably parasites on ectomycorrhizal and a ± excentric or missing stem and
and Gom­­phidius, have a ± slimy fungi with conifers (boletes and grows on wood (page 96).
universal veil that covers the entire Rhizopogon); the single species in
young fruitbody. They are related to Melanomphalia is presumed to be a Further reading: 156, 181, 289.
the boletes and have long, somewhat decomposer.

Chroogomphus
1 species
page 597
deeply decurrent gills

Micro-drawings:
Gomphidius spores to the left,
3 species cystidia to the right
page 598 within each sector.

Approximate species
number applies to
temperate Europe.

7 mm

large and fleshy; Chroogomphus rutilus s.l. is a

brownish-rose; sturdy, fleshy, red-brown, somewhat
with threadlike slimy to dry, almost black-spored
large and fleshy;
veil
rose or greyish; agaric with deeply decurrent gills. The
with slimy veil
stem bears an indistinct ring zone
from the ephemeral partial veil, and
the flesh has wine-red tinges, but is
Melanomphalia yellower towards the base. The gills
I+ 1 species are fimbriate from cystidia. The spores
(dex.)
page 599 are smooth, ± fusiform, ± dextrinoid
and measure 15–22 × 5.5–7 µm.
slender with dry and
minutely scaly cap; Always found with Pinus, but probably
without veil parasitic on species of Suillus (p. 796)
that have Pinus as a mycorrhizal
partner, e.g. S. granulatus (page 800).
Gomphidius glutinosus ▷▷ is more
slimy and grey. Recent studies indicate
that there are eight European species in
the Chroogomphus rutilus complex. In
the strict sense, C. rutilus should have
gomphidioids and rather crowded gills (upper picture);
Melanomphalia other species have more distant gills.
have deeply decurrent The distribution of the eight species
gills and dark brown to
black spore-deposits is, as yet, unclear. As a whole they
other with brown
are widespread, rather common to
with brown
similar fungi spore-deposits spore-deposits occasional; June–November. 10 mm 2 mm
and ± central and ± excentric
stem or missing stem © Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
paxilloids and the like, page 756 Tapinella, page 96 Princeton University Press, 2019
600 pholiotoids – ± lignicolous agarics with brown spores pholiotoids – ± lignicolous agarics with brown spores 601

Kuehneromyces

 Pholiotoids © Thomas Læssøe & Jens H. Petersen 2 species
Fungi of Temperate Europe, page 606 Micro-drawings:
Princeton University Press, 2019 Galerina
spores outwards
This group includes wood-inhabiting page 638
followed by
agarics with brown (grey-, ochre-, red- Deconica cystidia.
to orange-brown) spore-deposits. The page 548
Approximate species
principal genera are Hemipholiota, Flammula number applies to
Kuehneromyces, Pholiota and 2 species temperate Europe.
page 605
Gymnopilus. Many of the species
included are rather fleshy, with either Meottomyces
1 species
slimy or scaly cap surfaces. A number page 605
of similar, but mainly terrerestrial, stem with ring and
genera are treated under the group stem with ring, brown scales along
‘little brown mushrooms’ (LBMs – page silvery fibrillose length with greasy,
with white smooth cap;
616) and the dividing line between Hemipholiota
remnants of no pleurocystidia
2 species
these two groups is not well defined. veil; gills broadly page 607
All species in the group are decom­ adnate
posers, and almost all degrade wood.
with grey-brown, very fleshy and
smooth, sticky- large; spore-
Other similar fungi: Flammulaster slimy cap with white deposit grey-
– some ‘little brown mushrooms’ page 748 remnants of veil; on brown; universal
leaf-litter veil thick and
(LBMs) grow on wood, but at most cottony
have only faintly scaly caps (page 616).
– some lignicolous brown-spored
agarics are pleurotoid with excentric, with brown to
lateral or missing stems. See the orange colours;
cap cuticle grainy, spores mostly mostly slimy or Pholiota
pleurotoids (page 60). without radial with germ pore pointed-scaly, ± 22+ species
– lignicolous species with brownish- structure yellowish or ± orange page 608
rose spore-deposits and free gills can pholiotoids
are lignicolous with
be found under the pluteoids (p. 486). brown spores and spores with
– scaly or slimy species with dark greasy/slimy or scaly germ pore and
(see more genera with
± olive-brown; cap caps; some have chryso­cystidia
chocolate-brown to black spore- chrysocystidia under
Simocybe cuticle pruinose, chrysocystidia hypholomatoids, p. 580)
deposits are found under the grainy to hairy,
5+ species spores
hypholomatoids (page 580). page 647 without radial without
structure germ pore
Further reading: 112, 156, 181, 225.
other similar fungi
I+
(dex.)
small, dry,
with erect scales

spore-deposits spore-deposits spore-deposits

± black brownish-rose ± brown

Phaeomarasmius with bitter

2 species taste
page 604

with broadly
adnate gills and a
adnexed to adnate, emarginate or short well-defined ring hypholomatoids
decurrent gills page 580

Gymnopilus Tubaria
15+ species page 644
page 602
little brown
mushrooms
pluteoids pleurotoids (LBMs)
page 486 page 60 page 616
616 LBMs LBMs 617

Conocybe
Little brown mushrooms 60+ species Micro-drawings:
page 631 spores outwards
This group includes mostly small, Naucoria followed by cystidia.
predominantly brownish agarics, e.g. 21 species Dex. = dextrinoid.
Macrocystidia page 628 Galerina
Tubaria, Naucoria, Galerina, Conocybe, 1 species Pholiotina 55+ species Approximate species
Pholiotina, Simocybe, Agrocybe and page 288 30+ species page 638 number applies to
Flammulaster. The label LBM that is page 634 temperate Europe.
used for this group of fungi was coined
many years ago. The more vividly
coloured Bolbitius species are also
included here, and also the more fleshy
Agrocybe, Cyclocybe and Phaeocollybia.
Simocybe adnexed gills Tubaria
The group overlaps somewhat with the 7 species mycorrhizal; and ± pruinose 10+ species
stem mostly adnexed gills and
pholiotids (page 600). page 647 stem from pruinose stem apex;
page 644
with remnants cystidia
The majority of the species are Cyclocybe of veil mostly with veil,
decomposers but Descolia and Naucoria 2 species mostly as a ring gill attachment
stem and gill variable; stem rarely
are ectomycorrhizal and Phaeocollybia is page 624 Phaeogalera
sides downy; pruinose but often
smell rancid 1+ species
possibly a root parasite. cap
I+ with remnants page 646
(dex.)
mostly with of veil gills broadly
greenish shades; adnate; stem with
Other similar fungi: I+ remnants of veil,
cap cuticle ± (dex.)
– psathyrelloids and hypholomatoids spores
pruinose rarely as ring spore-deposit
grey-brown, dark grey-brown; with
have darker, purple-brown to black
germ pore absent remnants of veil
spores (pages 546 & 580). or tiny; cap cuticle Mythicomyces
– pholiotoids are mostly wood- cellular; 1 species
with a ring spore-deposit grey- page 646
inhabiting, larger and have slimy or reasy to dry; smooth
cap g or r
ugu brown; without veil;
scaly caps (page 600). l stem rather tough, dark
spores grey-brown,

ose

– Inocybe has mostly coarsely fibrillose LBMs
towards base
Agrocybe with germ pore; cap I+
cap surfaces and spermatic or strong 11 species cuticle cellular; with are the brown-spored
agarics that remain pale grey spore- Homophron
fruity odours (page 652). or without a ring

page 625

slimy-greasy
once the pholiotoids, deposit, without veil; 2 species
– Hebeloma has mostly earthy, fruity or Inocybe, Hebeloma, stem white, fragile page 578
rooting and with
flower-like odours (page 680). Cortinarius and the

grainy
Phaeocollybia pointed, greasy-slimy

cap ±
paxilloids have been
– Cortinarius has mostly larger 6 species cap; probably parasitic cap pruinose to

to h
excluded
fruitbodies with red-brown spore- page 622 grainy, mostly with

airy
greenish shades;
deposits and threadlike partial veils I+ without veil
stem with
(dex.)
(page 690). remnants of veil;
spore-deposit grey- other
I+ cap ± floccose-
brown (dex.) similar fungi
Further reading: 102, 103, 113, 113, Phaeonematoloma
grainy from
Simocybe
remnants of veil
121, 122, 123, 156, 181, 182, 201, 1 species 7 species
227, 296, 331, 341. page 620 page 647
with ring; cap margin
mycorrhizal with often covered
Nothofagus by long, rigid
hairs
gills mostly
broadly adnate

gills free to
adnexed dark brown to ± brown spore-
black spore-deposits deposits

Flammulaster
10 species
page 648
Descolea
1 species Ripartites
page 621 1-5 species
free, adnexed to adnate, emarginate or
short decurrent gills page 651
Deconica
17 species
page 548

© Thomas Læssøe & Jens H. Petersen

Bolbitius hypholomatoids psathyrelloids Inocybe Cortinarius Hebeloma pholiotoids Fungi of Temperate Europe,
6+ species, page 618 page 580 page 546 page 652 page 690 page 680 page 600 Princeton University Press, 2019
652 Inocybe – mycorrhizal agarics with grey-brown spore-deposits and ± fibrillose caps inocybes with nauseating smells of marzipan, honey or perfume 653

Inocybe
This genus includes ecto­mycorrhizal
agarics with fibrillose to scaly caps,
grey-brown spore-deposits and mostly
conspicuous cystidia. Many species
have spermatic or, more rarely, flower-
or fruit-like, strong odours.
Inocybes may have a thread-like
universal veil and a similar partial
veil and cystidia above the veil zone.
Species without partial veil are typically
downy from cystidia along the entire
lengh of the stem. Young fruitbodies
with untouched stems are important
when assessing these characters. The
genus can be divided into subgenera
based on cystidia characters.
Many species are poisonous and
contain, among other compounds,
muscarin – none of the species should
be consumed.
Other similar fungi:
– Cortinarius has more rust-brown
spore-deposits (page 690).
– LBMs have different odours or more
rust-brown spores (page 616)
Further reading: 156, 173, 184, 315.

More than 185 species

in temperate Europe.

Micro-drawings:
spores outwards
followed by cystidia.

emarginate gills

page 659
page 653

page 656 8 mm
page 662

with nauseatingly
sweetish smells of smell mostly
marzipan, honey neither sweetish
or perfume nor spermatic;
flesh green or
reddening
Inocybe
has emarginate gills, grey-
brown spore-deposits
and often spermatic or
sweetish smells; all are
ectomycorrhizal
†? Inocybe corydalina is a large, on top. The spores are amygdaliform have distinct brown scales. Inocybe
pale Inocybe with greenish, appressed and measure 7.5–9.5 × 5–6 µm. Occurs erinaceomorpha ▽, sometimes treated
fibrous scales mostly towards the on ± calcareous soils or clay, mostly as a variety of I. corydalina, differs in
the umbonate centre. The smell is with Fagus and other deciduous trees, having a dark brown cap centre and
sweetish, perfume-like, nauseating. rarely with conifers. mostly dark brown fibrous, adpressed
The flesh is ± reddening. The stem is Other large inocybes with sweetish scales.
pale, but darkens ± towards the base, odours, e.g. I. fraudans ▷▷ and Widespread, occasional, absent
sometimes greenish. The pleurocystidia I. bongardii ▷▷, are not green at the from the boreal zone; mainly July–
without partial are thick-walled (to 2 µm) with crystals cap centre, redden less obviously, or October.
veil; stem with page 665
cystidia to the
base
smell mostly †? Inocybe erinaceomorpha is a
spermatic; rather fleshy, cinnamon-brown, scaly-
flesh not green with partial
or reddening; veil; stem fibrous Inocybe with a nauseatingly
with evident with cystidia strong smell of cider. The stem is
basal bulb only on the ± cylindrical, at first white, later
upper
part tinged in the cap colour. The flesh can
smell mostly
spermatic;
redden slightly. The pleurocystidia are
not green or ± clavate with up to 2 µm wide walls.
reddening; Spores, smooth, amygdaliform and
nodulose without evident
spore basal bulb measure 7.5–9.5 × 5–6 µm. Occurs on
clay soils with Fagus and Quercus.
Inocybe coryda­lina △ has a grey-
green cap centre. Other somewhat
similar species have flesh that reddens
strongly or thin-walled pleurocystidia.
Widespread, occasional to rather
rare, absent from the boreal zone;
August–October. 20 mm
little brown
other Thomas Læssøe
mushrooms
similar fungi © Thomas Læssøe & Jens H. Petersen
(LBMs) Cortinarius Fungi of Temperate Europe,
page 616 page 690 Princeton University Press, 2019
680 Hebeloma – mycorrhizal agarics with grey-brown spore-deposits and ± radish to sweet smells hebelomas with membranous ring and marcipan smell 681

Hebeloma
This genus includes mycorrhizal agarics
with ± grey-brown spore-depopsits,
mostly smooth, greasy to slimy caps,
emarginate gills and fruity, flower-
like, radish-like to earth-like odours.
The spores are ± verrucose and may
be ± dextrinoid and have a ± loosening
outer wall; the gill edge is covered with
cystidia which, in some cases, produce
a clear liquid that traps the dark spores.
Other similar fungi:
– Cortinarius may look similar, but has
more rust-brown spores (page 690).
– the remaining group of brown
spored mushrooms (LBMs) has different
odours and mostly more rust-brown
spores (page 616).
Further reading: 29, 156, 184.

More than 55 species in emarginate gills

temperate Europe.

Micro-drawings:
spores first followed
by cystidia.

Approximate species
number applies to
temperate Europe.

8 mm
Hebeloma Hebeloma
radicosum mesophaeum group
page 681 page 682 Hebeloma sacchariolens group
page 684 Hebeloma radicosum is a very
distinctive, large Hebeloma with
a deeply rooting stem, a striking
Hebeloma marzipan- or bitter almond-like
crustuliniforme group
page 685 smell and a floccose ring. The cap
is dull yellowish brown to pinkish
buff with scaly remnants of veil. The
spores measure 8–10 × 4.5–5.5 µm.
Occurs in deciduous forests, mostly
with Fagus, appearing from latrines
or middens made by moles or other
small forest-dwelling mammals; a
so-called ‘ammonia fungus’.
May recall some pholiotoids,
with membranous
ring; deeply particularly Hemipholiota populnea
rooting; with thread-like (page 607), but the smell, its soil-
smell of partial veil; smell inhabiting nature with a very deeply
marzipan of radish and soil
naked; rooting stem and ornamented spores
with strong, are good separating characters.
sweetish,
perfumed smells Widespread and fairly common
in the nemoral zone but becoming
naked; scarcer towards the north and
smell of radish,
earth, fruit or absent from the boreal zone; July–
cacao; often with November.
drops on the gill
may be edges
dextrinoid

Hebeloma
has emarginate gills,
± grey-brown spore-
deposits and ± radish- 20 mm
like or sweet smells; other
the species are similar fungi
© Thomas Læssøe & Jens H. Petersen
ectomycorrhizal LBMs, Cortinarius Fungi of Temperate Europe,
page 616 page 690 Princeton University Press, 2019
690 Cortinarius – mycorrhizal agarics with rust-brown spores and veil Cortinarius – mycorrhizal agarics with rust-brown spores and veil 691

phlegmacia
with a marginate bulb;
Cortinarius phlegmacia
young gills olive-brown,
with a marginate bulb;
Cortinarius is a genus of ectomycor­ brown to greyish dermocybes
young gills yellow to
page 718 (section Dermocybe)
rhizal agarics with ± emargi­nate yellow-green
page 722
page 714
gills, rusty-brown, verrucose, mostly
dextrinoid spores and well-defined large, dry,
± colourful cortinarias
veils. The partial veil is mostly thread- (subgenus Cortinarius p.p.)
like, while the universal veil leaves a page 728
phlegmacia
woolly, thread-like or slimy covering on with a marginate
the lower stem and on the cap. Only bulb; young gills
bluish
very rarely with cheilocystidia (of a page 708
simple type), and never pleurocystidia.
The genus is enormous and
difficult to deal with. New species
are constantly being described and
new synonymies published. Safe
identification is in many cases not small, brightly
possible without sequencing. Important coloured, but never
macroscopical characters include the with bluish to
violet gills
type of veil (slimy or thread-like), the
gill and flesh colour when young, the stem with a
overall shape, the smell, and chemical marginate bulb

colour reactions with KOH. large, with intense

cap slimy, orange-brown, telamonias
stem dry above cap dry, not yellowish or violet (subgenus Telamonia)
Other similar fungi: the base hygrophanous, colours page 732
mostly intensely
– Inocybe has fibrillose to coarsely coloured
fibrillose caps and grey-brown spores.
species of
The spores are smooth, nodulose Cortinarius
stem without a
or star-shaped, and all species have marginate bulb are mostly rather
small
cystidia (page 652). large and fleshy with
vivid rusty-brown dry to greasy,
– Hebeloma likewise has grey-brown spores and a thread- mostly
spores and smells mostly earth- or like partial veil hygrophanous,
radish-like. The spores are verrucose phlegmacia mostly with dull
colours small telamonias
and all species have prominent cystidia (≈ subgenus medium-sized page 732
Phlegmacium) to large
on the gill edge (page 680). page 698
cap and
stem slimy similar
– little brown mushrooms (LBMs) are large telamonias
brown-spored,
page 739
typically decomposers, and generally soil-inhabiting
fungi medium-sized
have cystidia (page 616).
to large; with a
– Leucocortinarius bulbiger recalls a membranous ring
white-spored phlegmacia (page 257).
myxacia
(subgenus Myxacium)
Further reading: 46, 69, 124, 125, 126, page 692
127, 156, 184, 305, 323.

More than 500 species

in temperate Europe.

universal veil spores

partial veil emarginate gills

universal veil

Cortinarius caperatus
and the like
page 754

© Thomas Læssøe & Jens H. Petersen

LBMs Hebeloma Inocybe Fungi of Temperate Europe,
page 616 page 680 page 652 Princeton University Press, 2019
756 Paxillus – brown-spored agarics with deeply decurrent gills and ± central stem brown-spored agarics with deeply decurrent gills and ± central stem 757

Paxillus and the like

This group includes agarics with
a ± brown spore-deposit, deeply
decurrent gills and a ± central stem.
Both of the genera included, Paxillus
and Phylloporus, are ectomycorrhizal
and belong phylogenetically to the
Boletales.
Other similar fungi:
– gomphidioids have dark brown to
black spore-deposits (page 596).
– the brown-spored genus Tapinella
has ± excentric, lateral or missing
stems (pleurotoids page 96).
– Lepista & Paralepista are more
smooth-capped and have whitish to
brownish-rose spore-deposits and
verrucose spores (page 112).
Further reading: 97, 133, 333.

Micro-drawings:
spores outwards
then cystidia.

Approximate species
number applies to
temperate Europe.

Paxillus
5+ species
page 757 deeply decurrent gills

12 mm

Phylloporus
1 species Paxillus is a genus of ectomycor­
page 759 rhizal, brown-spored agarics with
fairly crowded, decurrent gills that
can be loosened from the cap flesh.
felty-hairy, inrolled The colour of a fresh spore-deposit is
margin; gills brown when margin ± straight; important for identification (although
bruised gills thick and yellow, this can change over time). Repeated
not bruising brown Ripartites
consumption of paxilloids may cause
see page 651
a life-threatening autoimmune
reaction.
fruitbodies very
pale, with ± strigose †† Paxillus involutus is a relatively
margin; gills not
staining brown slender, rather pale Paxillus with a
downy, inrolled, wrinkled margin. The
stem is ± smooth; lacks conspicuous
reddish spots or yellow bands. The
gills are pale straw-yellow to yellowish-
brown and stain brown when bruised;
they can be loosened from the pale
cap flesh. The spore-deposit is yellow-
brown to grey-olive. The smooth
spores measure 7–11 × 5–6.5 µm.
Mostly mycorrhizal with Picea and
Paxillus
and the like
Betula, and usually on poor acidic soils.
are ectomycorrhizals Paxillus filamentosus ▷▷ is more
with deeply decurrent yellow-fleshed and occurs with Alnus.
gills, ± central stem
and and ± brown with dark
Other species are shorter and thicker-
spore-deposits brown to black stemmed.
spore-deposits with brown 7 mm
spore-deposits
Widespread and very common;
and central
stem and ± excentric June–November.
other © Thomas Læssøe & Jens H. Petersen
or missing stem
similar fungi Fungi of Temperate Europe,
gomphidioids and the like, page 596 Tapinella, page 96 Princeton University Press, 2019
760 boletes boletes 761
Hemileccinum Buchwaldoboletus Xerocomus
1 species 2 species, page 783 4 species, page 784 Hortiboletus &
page 781 Micro-drawings:
Boletes Rheubarbariboletus
spores.
4 species
The boletes is a group of fungi that form Cyanoboletus page 786 Xerocomellus
1 species Approximate
rather soft-fleshed fruitbodies that have 6+ species
Leccinellum page 781 species number
page 790
tubes rather than gills below the cap 2 species applies to
& Neoboletus p.p. temperate Europe.
and a sturdy central stem. The tubes are page 780 Pseudoboletus
generally not joined firmly to each other, 1 species
or to the cap flesh, from which the page 793
entire tube layer can easily be loosened. tubes tubes not cap mostly Imleria
iodoform ± decurrent decurrent red; spore 1 species
The spores form inside the tubes and smell at base cap mostly page 794
Q 2.3–2.4
are mostly brown. Many boletes change immediately
brownish; Gyrodon
colour when bruised or sectioned. These Q 2.5–2.9 1 species
Aureoboletus bluing
page 795
colour changes, the colour of the pores 2 species cap dry, with parasitic on
page 779 thick felt Scleroderma
(the visible part of the tubes) and the tubes pale yellow,
cap surface structure are very important cap not felty immediately bluing;
cap dry, smooth mostly with conifers
characters. or slimy to finely felty;
Imperator fibrillose to do stem less than
The boletes constitute a monophy­ smooth, tted-
3 species stem stem min 15 mm wide
letic entity and are classified in the page 778 tapering ute
ly s
order Boletales. The order also includes or not cal tubes decurrent and
y
immediately bluing;
some false truffles (e.g. Rhizopogon), stem long with Alnus

sc tem
some corticioids (e.g. Serpula) and some and slender

aly
s
with a coarse Aureoboletus
species with gills (e.g. Paxillus and the reticulum 2 species
gomphidioids). cap
tubes page 795
mild; bright yellow,
Almost all boletes are ectomycor­ immediately greasy-slimy
not bluing; cap

lum
rhizal with deciduous trees or conifers. bluing colour of the pores when wet
slime sour

reticu
The genus and sometimes the species of Butyriboletus
the host is important for identification,

stem with
6 species
mild
especially in the genera Leccinum and page 774 boletes
are soft-fleshed stem with or
Suillus. A few species are parasitic: without ring
fungi with soft,
Pseudoboletus parasitize Scleroderma; ± separable zone; with
Buch­waldo­boletus is probably parasitic bitter taste tubes below conifers

ooth
Caloboletus Suillus p.p.
a cap
on Phaeolus; and Chalciporus perhaps

stem sm
3 species flesh not 16+ species
parasitizes Amanita muscaria. page 772 bluing
smoo
page 796

stemth
There are many choice edibles mild; stem
without ring zone;

ulum
chambered
among the boletes, but also a few bitter, reti
flesh stem flesh yellow

retic
witulum
inedible species (e.g. Tylopilus felleus, moderately
c
h

ith
Gyroporus mild; bluing; base

rw
Caloboletus radicans and C. calopus) reticulum not reddish

o
3 species

th
and most species of Rubroboletus and other Chalciporus &
ret
wi culu
mostly pale all parts

oo
page 782
th m
i

sm
similar immediately Rubinoboletus
Imperator are moderately poisonous. st fungi ly 3 species
sca em sca bluing; flesh
page 812
bitter; ly in base ±
Other similar fungi: reticulum tubes all parts reddish
oth
– polypores with a stem (e.g. Albatrellus dark stem decurrent; stem smo immediately
flesh ± tough scaly stem bluing; base
long
and Boletellus) may look very similar. and not reddish
Boletus Rubroboletus
They have tubes that are firmly joined – 4 species scaly
5 species
both with each other and the flesh. They page 769 cap cap and stem page 804
also often have decurrent tubes and and stem ± smooth
coarsely
tough flesh – see page 826. woolly-scaly

cap slimy Suillellus

Further reading: 71, 150, 156, 170, 194, 3 species
205, 228. cap dry
page 808

Tylopilus
tubes 1 species
page 768

Suillus p.p. Neoboletus

Leccinum Imperator
16+ species 2 species
pores 12+ species 3 species
Strobilomyces Porphyrellus page 796 page 810
page 762 page 807
polypores with stem 1 species 1 species © Thomas Læssøe & Jens H. Petersen
page 826 page 813 page 813 Fungi of Temperate Europe,
Princeton University Press, 2019
824 groups of polypores groups of polypores 825

Polypores
The polypores is a form group that is
characterized by a tubular hymenophore
where the spores are formed on a
hymenium inside the downwards-
pointing tubes. The tubes are normally
inseparable from each other and from perennial
the cap flesh. The pores (tube mouths) polypores
may be spherical, angular, stretched, page 854

labyrinthine or gill-like. The number of clustered polypores

pores per millimetre is an important page 846
character, best measured with a trans­ fruitbodies
lucent ruler using a lens. The pore edges perennial;
hard or tough
are sterile. Fruitbodies can be completely
appresed to the substrate or have ± well-
annual, capped and
developed caps, and rarely also stems;
adnate polypores
they may be soft and annual or tough page 886
to hard, and in some cases perennial.
Perennial fruitbodies may be recognized
fruitbodies
by typically having several tube layers
annual,
and broad growth zones on the caps. polypores composite,
All polypores belong to Basidio­ with a stem clustered
page 826
mycota and have one-celled basidia. The
only exception is Aporpium (page 1186), fruitbodies annual, with
which belongs to the jelly fungi and caps but possibly partly
resupinate (reflexed)
has longitudinally divided basidia. Most
species with tough or hard fruitbodies polypores
form the spores
have a complicated hyphal structure, in tubes on the
which, besides the normal generative underside of the
hyphae, consists of branched binding fruitbodies annual, fruitbody. Mostly
tongue-shaped or wood decayers.
hyphae and/or skeletal hyphae (page
with a stem fruitbodies
35). Fruitbodies with two hyphal types annual,
are termed dimitic, and those with three tough or soft
are referred to as trimitic.
Almost all polypores are decomposers
and typically occur on wood. They
produce a brown rot or a white rot (page other
14). A few species are ecto­mycorrhizal, similar fungi
e.g. Albatrellus & Scutiger (page 828)
and Coltricia (page 843).

Other similar fungi: annual, resupinate

(without caps but
– the boletes are always soft-fleshed,
sometimes with small
and their tubes are not well-joined and projections)
can be loosened from the cap flesh;
most are mycorrhizal (page 760). merulioid annual, completely
– merulioid corticioids have a ± folded corticioids resupinate (flat)
hymenophore where also the edges are page 974 polypores
page 932
covered with hymenium (page 974).
– cyphelloids are small, cup-shaped
Basidiomycota. They may occur in dense
groups and mimic a polypore (page
1076).

Further reading: 17, 37, 161, 163, 193,

219, 220, 235, 282, 283, 308, 309, boletes
339, 356, 366, 367. page 760
© Thomas Læssøe & Jens H. Petersen
cyphelloids Fungi of Temperate Europe,
page 1076 Princeton University Press, 2019
 Cerioporus
826 polypores with central or lateral stem, or with tongue-shaped fruitbodies 2 species &
polypores with central or lateral stem, or with tongue-shaped fruitbodies 827
Picipes
4 species Lentinus
Polypores with a stem page 836 4+ species Neofavolus
Polyporus page 838 1 species
This form group includes somewhat 1 species page 839
boletoid fruitbodies and more tongue- & Cerioporus
shaped fruitbodies, that may have a 2 species Jahnoporus
Piptoporus page 834 1 species
short lateral stem. All form annual 1 species page 833
fruitbodies. In one genus, Fistulina, page 833
the tubes are not joined. Most species
produce hyaline, smooth spores, but a Ganoderma
7 species, 3 with a stem
few have brown, ornamented spores. page 840
Prominent brown setae are found in Buglossoporus
1 species
one genus. page 832 Podofomes
The group includes both 1–7 pores/mm; 1 species
mostly with
stem black with rudimentary page 842
ectomycorrhizals, e.g. Albatrellus, 4–9 pores/mm;
cap smooth or with age; cap stem and 1–2
Scutiger and Coltricia, decomposers hairy ± smooth pores/mm; on felty-hairy;
on soil, e.g. Xanthoporus, and wood coarse scales; deciduous wood from buried
pores large, conifer wood
decayers that produce a white or 1–2 mm
brown rot. with lacquer-like
with papery
cap crust that can
crust;
be melted
Other similar fungi: on Betula
Boletopsis Pelloporus
– boletes with decurrent tubes are flesh pale pinkish-
3 species durng summer, 3 species, 1 with a stem
soft-fleshed, and usually have tubes wood decayers, white rot buff; with a black

rot rs
page 842

wn ye
page 831 mostly low

bro d deca
zone below the cap
that are easy to loosen from the cap down on old felt; on Abies

woo
flesh. Gyrodon has firmly attached Quercus
tubes, but these stain blue when flesh colour
cap and stem
bruised (pages 760 & 795). stem greyish brown felty;
– in clustered polypores the individual to black; with flesh with duplex
1–4 pores/ structure
caps may also have a stem (page 846). seta
mm
– broadly adnate polypores on top Scutiger
3 species
of stumps and trunks may develop page 828
polypores
a contracted base and thus recall

m y co rr hiz al
e ct o m y c or rh i
stem mostly Coltricia
with a stem
polypores with a stem, but usually yellowish; with concentric 5 species
pores 0.5–2 mm zones and thin page 843
there will be more normal fruitbodies large felty covering

z al
in the same fructification (pages 854
& 886).

rot s,
w n ay e r
bro d e c
tubes and
See also the overview and the refe­ other

od
stem whitish; o
ec il-inh

wo
similar fungi

s
rences to further reading on page 824. with 2–5 o m ab i brown-felty with

d
pores/mm po ting ro t labyrinthine pores
ser it e
s w h


small with
dentate-floccose
pores and phenolic
smell with red juice; tubes
Albatrellus in mature fruitbodies
10+ species not joined; mostly
page 829 on Quercus
fasciculate small and thin-
yellow; polypores Phaeolus
stemmed; with
in nitrogen page 846 (see page 853)
large pores
enriched habitats

Sistotrema
1 species with a stem
(see page 1075)

Micro-drawings:
spores, cystidia and
setae. Fistulina
Favolaschia 1 species
Approximate species 1 species page 845
number applies to Xanthoporus page 845
temperate Europe. 1 species boletes
page 828 (Buchwaldoboletus, unusally formed, normally
© Thomas Læssøe & Jens H. Petersen
Gyrodon and others) broadly adnate fruitbodies Fungi of Temperate Europe,
page 760 pages 854 & 886 Princeton University Press, 2019
846 clustered polypores (polypores with numerous caps from a common base) clustered polypores 847

Clustered polypores Meripilus Micro-drawings:

1 species spores outwards
This group includes species with page 850 then cystidia.
Bondarzewia cy. = cystidia.
numerous caps from a narrow, 1 species
Grifola cl. = clamps
common base or from a thick, short 1 species page 851
stem. All species in the group form page 849 Approximate
species number
annual, rather soft-fleshed fruitbodies applies to
and are typically decomposing Abortiporus
temperate
1 species
parasites on trees. Europe.
page 852
A number of these species
are prized edibles, e.g. Grifola,
Cladomeris umbellatus and
Laetiporus, and less so Meripilus
and Fistulina. Phaeolus is used for
dyeing wool. Laetipours is known to
cause mild poisoning when cooked
insufficiently. Cladomeris
- cl. I+
1 species
Other similar fungi: page 849
cy. - cl.
– polypores with stems are similar,
but only have one or a few caps
Buglossoporus
(page 826). (see page 2)
– perennial, broadly adnate polypores flesh colour
may form numerous, layered caps, ite
rot
wh
but do not have a common stem or cy. rot
wn
bro
constricted base (pages 854 & 886).
– Sparassis and similar fungi lack
pores (page 952). clustered
polypores
See also the overview and the refe­ form numerous caps
from a common cy.
rences to further reading on page 824. base
t
white ro
rot - cl.
brown

cy.

other
similar fungi

- cl.

broadly adnate
polypores
pages 854 & 886 Phaeolus
1 species
page 853

Fistulina
(see page 845)
Laetiporus
2+ species
page 848

polypores
with a stem Sparassis and similar fungi © Thomas Læssøe & Jens H. Petersen
page 826 page 952 Fungi of Temperate Europe,
Princeton University Press, 2019
 Fomitopsis
Laricifomes
1 species
654 perennial polypores 1 species
page 867
perennial polypores 855
page 866

Daedalea
Perennial polypores Perenniporia 1 species
5 species page 865
All polypores where the fruitbodies page 863
Gloeophyllum Fomes
continue to produce new tube layers 4 species
Rhodofomes 1 species
in successive years belong here. The page 868 page 870
1 species,
growth periods can often be seen as page 866
Rigidoporus
raised concentric zones on the caps. 6+ species Osmoporus
The group includes most polypores page 861 1 species
page 868
with hard fruitbodies. In general, the
fruitbodies have one growth period
high,
per year, but Fomes fomentarius has hoof-shaped;
with
two growth periods: a strong one large, flesh very pores flesh pale
bitter pink yellow in
in the spring and a weaker one the labyrinthine young parts with strong, Ganoderma
autumn; this creates two zones of pores
spores mostly sweetish smell 7 species
growth per year. dextrinoid and pores mostly page 871
Some species are borderline thick-walled gill-like,
Daedaleopsis stretched

br
perennial. For example, species of

w
(see page spore-deposit white;

ow rot
hi

n
Trametes, Cerrena and Antrodia may,

te
910) fruitbody
with cystidia;

ro
I+ with mycelial core

t
during a mild spring, continue to flesh mostly
t
grow and thus produce two-layered ± pale brown ro
t
n
w ro
fruitbodies. However, these ro te
with b hi
w
fruitbodies typically rot away during Funalia reddening,
- cl. Phellinus
radially spore-deposit red-
the summer. (see page flesh colour 10+ species
stretched brown; mainly on
Most perennial polypores have 916) page 874
pores deciduous trees
a complicated hyphal system with
both generative hyphae with septa, with deep mostly with hoof-
and branched binding hyphae and/or felt and large shaped, usually cracked
pores caps; on deciduous
thick-walled skeletal hyphae without perennial
- cl. trees
septa (page 35). polypores
The species can be divided into with pale grey, – pileate or
mostly with hoof- Fomitiporia
labyrinthine resupinate; on
white- and brown-rotters; although - cl. shaped, cracked caps; 4 species
pores wood.
Cerrena on deciduous trees or page 879
the two types may look surprisingly conifers
1 species
similar, they are not closely related. page 860 - cl.
with radially
stretched pores mostly resupinate or
Other similar fungi: other with sharp-edged
similar caps; mostly on
– similar but annual species of deciduous trees
with small fungi
polypore are divided into two groups: Trametes - cl. - cl.
spores and crystal- setae
those with caps (pileate) and those (see page 918) covered - cl.
without (resupinate) (pages 886 & hyphae I-/I+
932). - cl. with caps or
with a darker resupinate;
margin; often - cl.
with black lines in the
See also the overview and the refe­ high on Quercus flesh; on conifers Fuscoporia
rences to further reading on page Skeletocutis or Corylus 5 species
(see page 902) ± hairy and page 880
824.
high on Salix with large,
or Quercus ± labyrinthine pores;
on conifers
with small caps;
on roots and mostly on Salix Phellopilus
stumps; mostly on annual, resupinate polypores 1 species
Dichomitus conifers page 932 page 884
3 species with dark line in
page 859 the flesh; mostly
on Ribes or
Micro-drawings: Euonymus
Porodaedalea
spores outwards,
3 species
followed on the
Phellinopsis page 883
inside by cystidia
1 species
and setae.
page 885
cl. = clamps. Haploporus
2 species
Approximate species page 858
number applies to
temperate Europe. Heterobasidion Phylloporia © Thomas Læssøe & Jens H. Petersen
3 species, page 856 annual, pileate polypores 1 species, page 884 Fungi of Temperate Europe,
page 886 Princeton University Press, 2019
886 annual, capped polypores Tyromyces &
Datronia annual, capped polypores 887
2 species
Loweomyces
page 911
6 species Abortiporus
Spongipellis
Annual, capped polypores 3 species
page 908 (see page 852)

This group includes the reflexed page 906 Daedaleopsis Bjerkandera

Odoria & Aurantiporus 2 species 2 species
and capped polypores that have 2 species page 910 page 912
fruitbodies which rot away after one pages 904 & 905
season. The species form either soft
or tough, but never hard, fruitbodies.
All capped and stemless species with
soft fruitbodies belong here. Climacocystis
1 species
Continued on next page spread . . . page 903 Trichaptum
narrow; with medium-wide;
medium-wide; dark back side; 4 species
pores labyrinthine
mostly on on deciduous page 914
and reddening;
wide; pores deciduous wood on deciduous
mostly labyrin­thine; wood wide; pores oblong
wood
Micro-drawings: on deciduous and reddening; on
spores on the wood deciduous wood
outside followed by wide; soft
cystidia. flesh reddish with
medium-wide; with
cl. = clamps. Skeletocutis KOH; on deciduous Cerrena
grey pores or brown
13+ species wood (see page 860)
line; on deciduous
Approximate species page 901 wood
wide; upper side
number applies to deeply tufted- narrow; with
temperate Europe. Gloeoporus narrow; hairy; on conifers violet colours and
1 species with crystals on irregular pores
page 900 the hyphae; on flesh white, cream,
deciduous and pale pinkish-buff or
conifer wood pale blue medium-wide; Dichomitus
narrow; with pale grey, (see page 859)
pores pinkish; labyrinthine pores
rubbery; on deci­duous
wood narrow; with darker
Diplomitoporus margin; mostly
annual, capped high on Quercus or
2 species narrow with rather large
polypores Corylus
page 900 pores; on conifers
with broadly
+/- adnate, soft- to Funalia
narrow; cl. medium-wide; very 2 species
t tough-fleshed caps; felty and with large
extremely tough-fleshed white ro page 916
rot
on wood pores
brown
Antrodiella
17+ species
narrow; medium-wide; with
page 898
tough- to soft-fleshed yellowish colours and
small pores

medium-wide; with
medium-wide; large pores that
soft-fleshed split into teeth
I+ similar Frantisekia
groups 1 species
Antrodia, Cartilosoma narrow to wide; upper page 917
& Neoantrodia side densely zonate,
15+ species wide;
mostly hairy; pores small
page 896 tounge-shaped;
- cl. to large, rounded to
on Quercus Trametopsis
stretched
1 species
page 917
medium-wide;
reddening;
capped
on conifers, medium-wide; soft-
corticioids
boreal fleshed and page 962
reddening;
on conifers, boreal

Trametes
9 species
Postia
page 918
15+ species
Buglossoporus
page 891
(see page 832)

⬅
Amylocystis
1 species
page 890 Leptoporus
© Thomas Læssøe & Jens H. Petersen 1 species capped, perennial polypores flesh orange to dark brown
Fungi of Temperate Europe, see next page spread
Princeton University Press, 2019 page 890 page 854
888 annual, capped polypores, continued annual, capped polypores, continued 889

Daedaleopsis
Cerrena 2 species
Annual, capped polypores Funalia
(see page 860) page 910
Hapalopilus
2 species 2 species
Bjerkandera
continued from previous page spread . . . page 916 page 924 Ischnoderma
2 species
page 912 2 species
page 925
Some annual polypores may
consist solely of generative
hyphae and are termed
monomitic; such fruitbodies are Pelloporus
typically soft and easy to break. 3 species
When either skeletal hyphae or page 926

binding hyphae are also present pores oblong and

with pale grey, reddening; base stains
the fruitbodies are termed dimitic; laby­rinthine on deciduous violet; on with dark crust below Mensularia
if all three hyphal types are pores; on wood deciduous the cap felt; pores ± 3 species
very felty deciduous wood light-reflecting; page 926
present they are termed trimitic. and with wood on deciduous and
Dimitic and trimitic annual large pores; conifer wood
on deciduous flesh with
fruitbodies are typically tough and duplex structure;
wood
leathery. (See page 35 for more pores ± light-
on hyphal systems) with grey pores reflecting; on
or brown line; conifer wood
on deciduous
Other similar fungi: wood pores ± light-
– capped, broadly adnate, perennial reflecting; caps Inonotus
setae 5 species
flesh dark brown - cl. rather narrow,
polypores over time form several ± smooth; on page 928
layers of tubes. They also often deciduous wood
orange, thick and
have raised zones on the surface soft-fleshed;
- cl.
(page 854). Aurantiporus
on Quercus
pores ± light-
– clustered polypores are annual 4 species reflecting;
and form numerous caps from a page 923 annual, capped caps wide, deeply
polypores tomentose; on
common stem (page 846). - cl.

flesh orange
with broadly deciduous wood
– capped corticioids do not have adnate, soft- to
pores; a hand lens is required (page orange tough-fleshed caps;
and tough; on wood - cl. pores ± light-
962). on deciduous reflecting; caps
wood e ro
t very large, ± smooth,
whit flesh without mycelial
rot
See also the overview and the refe­ bro
wn
core; on Quercus
- cl.
rences to further reading on page - cl.
824. Pycnoporus
w
1 species - cl. br
hi
t pores ± light-
page 922 ow e ro
n t reflecting; flesh with
with splitting ro
t mycelial core; on
pores; mostly on
deciduous wood
conifers
similar Pseudoinonotus
fungi 1 species
pores ± light- page 930
reflecting,
labyrinthine

not light-reflecting;
pores gill-like
capped corticioids
Pycnoporellus page 962 Inocutis
2 species 2 species
page 922 page 931

Micro-drawings:
spores outermost,
followed by cystidia
or setae.
cl. = clamps.

Approximate species

⬅
Phaeolus
number applies to (see page 853)
temperate Europe. Gloeophyllum
flesh white, cream, pale (see page 869)
capped, perennial polypores © Thomas Læssøe & Jens H. Petersen
pinkish-buff or pale blue
page 854 Fungi of Temperate Europe,
see previous page spread
Princeton University Press, 2019
 Sidera Physisporinus Rigidoporus & Emmia
932
84 annual, resupinate (flat) polypores 2 species 4 species 6 species annual, resupinate (flat) polypores 933
page 943 page 944 page 945 Porothelium
Cinereomyces (see page 1084)
1 species Ceriporia & Meruliopsis p.p.
Annual, resupinate polypores page 943 7+ species
page 946
This group includes polypores Antrodiella
with annual, flat fruitbodies 17+ species
page 942 Gloeoporus
that completely lack caps. Some 2 species
species may, however, form small page 948
protuberances (false caps) without Niemelaea
Skeletocutis mostly perennial,
an evident cap surface. Although discolours when 2 species
13+ species with free- with crystal-
bruised; page 949
some species can be recognized in page 941 composed
floating with spherical tipped cystidia of confluent
the field, it is a very difficult group with skeletal star- spores and spherical
hyphae that spores cups; dimitic
crystals with pastel
to identify, and in most cases correct dissolve
extremely colours or white;
identification requires microscopy. Xylodon tough; in KOH soft-fleshed
with greenish Ceriporiopsis
Many species have tough, long- 3 species with small tint; waxy, 5+ species
lasting fruitbodies that persist long with pores mostly spores soft-fleshed page 949
page 940 with very
after spore production has ceased. pale brown, large
small spores and
Sterile material is easily sampled pores; mostly with pored, ± soft-
but cannot be identified. It may be a crystals on the loose fleshed
hyphae star-
good idea to check if the specimen is Sistotrema with irregular crystals - cl. Porpomyces
able to deposit spores on a coverslip 3–4 species pores and with pastel 2 species
+/- colours, ± soft- page 950
with pores several types of cl. cy.
or microscope slide. Keep the page 939 cystidia crystal- fleshed
- cl.
specimen moist in order to enhance covered - cl.
hypha
the chance of obtaining a good cottony; whitish, with
spore-deposit. Trechispora with 4–, 6- or hyphal cords,
All species are wood-degrading 3 species 8-spored basidia ± soft-fleshed Junghuhnia
& Butyrea
and most produce a white rot. with pores cy. white rot 2+ species
page 938 basidium
cottony; vinaceous-brown with page 950
Other similar fungi: with echinulate +/- pale margin; with
spores cl. crystal-tipped cystidia
– annual reflexed (polypores with a annual, resupinate
polypores
resupinate decurrent lower part) and Anomoloma cy. Aporpium
form flat fruitbodies
capped polypores (page 886). 2 species cottony; that completely lack (see page 1186)
page 939 I+ basidium on deciduous wood;
– perennial, resupinate polypores (page with amyloid caps. On wood or basidia with
spores rarely on herbs. - cl. longitudinal walls
854). seta
– merulioid corticioids (corticioids with clamydo­

br
I+ spore wn

o
wrinkled-veined hymenophore) may soft; with ro t brown-fleshed;
basidium
look similar but the ‘pores’ are rounded amyloid spores with setae;
cy. other - cl. spores yellow Inonotus
and have fertile pore mouths – in true similar fungi 6+ resupinate
polypores the pore mouths are sterile species
(page 974). Anomoporia soft; with page 951
3 species chlamydospores cy.
– gregarious cyphelloids may recall orange-brown;
page 937 with rounded, fertile
resupinate polypores (page 1076). pores
soft-fleshed;
See also the overview and the refe­ Postia rose; on conifer yellow, ± labyrinthine; Meruliopsis
rences to further reading on page 824. 4+ resupinate wood with ornamented (see page 977)
species, spores and
- cl. cyanophilous basidia
page 936
± crisp; contents
dimitic

crisp; mostly
Rhodonia disc-shaped;
1 species dimitic ± crisp, with cyphelloids
pores Lindtneria
page 936 hyphal cords; orange; page 1076
disintegrating, 1 species with ‘pores’
dimitic ± crisp; with thick-walled (see page 980)
dimitic almost as
Micro-drawings: cystidia
spines
spores first,
followed on the
inside by cystidia.
cl. = clamps.
cy. = cystidia.

Approximate species Amyloporia Cartilosoma

number applies to 4+ species (see page 897)
temperate Europe. page 935
Fibroporia
1+ species perennial (incl. resupinate) and © Thomas Læssøe & Jens H. Petersen
Flavidoporia Pycnoporellus Auriporia merulioid corticioids annual, capped polypores Fungi of Temperate Europe,
page 936 1+ species, page 934 2 species, page 934 1 species, page 934 page 974 pages 854 & 886 Princeton University Press, 2019
952 rosette-fungi and the like rosette-fungi and the like 953

Rosette-fungi and the like

This form group comprises species Micro-drawings:
that, although not closely related, spores first,
followed by cystidia Cotylidia
are collectively known as ‘stipitate on the inside. 3 species
stereoids’. All the speices in this cl. = clamps. page 958
group produce fan- to rosette- Thelephora Stereopsis
Approximate species 9+ species
shaped or folded fruitbodies with a number applies to page 956
1 species
page 959
± smooth hymenium on the under/ temperate Europe.
outer side. The upper/inner side is
mostly sterile. The fruitbodies are medium-sized; with brown
typically rather tough and range from flesh, brown hyphae and
brown, verrucose spores medium-sized to
very large in Sparassis to small in Podoscypha small; with white
Muscinupta. Species of Thelephora 1 species flesh and numerous,
page 955 ± projecting cystidia
form ectomycorrhiza, while the
remaining species are presumed to be
decomposers or moss parasites.

Other similar fungi:

small; with white
– the clustered polypores may form flesh, yellow
large, rosette-shaped fruitbodies colours and very
but they always have pores on the small spores
Stereum
underside (page 846). (see page 962)
+/- cl.
– capped stereoids are tough, mostly - cl.
broadly attached to the substrate, and
not stipitate (page 962).
with - cl.
– pleurotoids with reduced gills (e.g. medium-sized;
skeletal with tough flesh
Gloiocephala and Arrhenia) form soft- hyphae and thick-walled
fleshed fruitbodies that usually have a stipitate stereoids
photo Lukas Large I+ cystidia
form fan- to rosette-
veined hymenophore (page 60). +cl. shaped or folded - cl.
– cyphelloids do not normally form large; with numerous fruitbodies with a
rosettes; with gloeocystidia hymenium on the - cl.
stems (page 1076). ± smooth lower/
and skeletal hyphae
– flattened, clavate species of outer side
Ascomycota are soft-fleshed with very large; with numerous small and
folds; without skeletal spatula-shaped;
a hymenium on both sides and the
hyphae with projecting
spores are formed within asci (page cystidia;
1350). other on mosses
similar fungi

Further reading: 92, 142,

+/- cl.
156, 163, 264.
Muscinupta
± tough ± fragile and soft-fleshed 1 species
page 959

Sparassis
2 species
page 954

earthtongues and others

page 1350
© Thomas Læssøe & Jens H. Petersen
capped corticioids clustered polypores pleurotoids without gills cyphelloids Fungi of Temperate Europe,
page 962 page 846 page 60 page 1076 Princeton University Press, 2019
960 corticioids corticioids 961

Corticioids corticioids
with veins or folds
This form group includes species with smooth, cottony
page 974
resupinate (flat) or reflexed (flat with a or peeling, fragile
narrow protruding cap) fruitbodies and corticioids
page 986
with a smooth, warty, spiny, toothed
or veined hymenophore. Species with
divided basidia or with spores that smooth, firm,
germinate with a replicate spore are whitish to greyish
corticioids corticioids
shown on this wheel, but are treated with spines or teeth page 1000
with the resupinate jelly fungi (page page 1036
1162).
Some distinctive species can be
told by macroscopical means. Colour,
cottony or flaky,
shape (mainly of the hymenophore) fragile
and ecology are the most important
characters. The majority of the species,
however, require microscopical study in
order to reach an identification.
Phylogenetically the species are
distributed among a series of orders
that include fungi with completely
without cap;
different morphologies (boletes, hymenophore gelatinous, waxy
agarics, polypores, etc.). folded or or leathery;
Most corticioids are decomposers, veined whitish to
and the majority are white-rotters. greyish
Species in the Boletales (e.g. Serpula
(page 984) and Coniophora (page
1034) produce a brown rot, and corticioids
most members of the Thelephorales, form ± flat fruitbodies without cap;
without cap; that are found mainly on hymenophore
e.g. Tomentella (page 998), are the underside or sides smooth or warty
hymenophore with smooth, firm, coloured
ectomycorrhizal. A handful of genera spines or teeth of dead wood or on corticioids
in other orders are also mycorrhizal. herbaceous stems page 1012
The corticioid fungi is a large group
gelatinous, waxy
with about 500 species in temperate or leathery;
Europe. Many rare or inconspicuous coloured (not
species are not included here, but whitish to greyish)
other
representatives of most of the genera similar fungi
that occur in the area are included.
with cap and
capped corticioids a flattened
Other similar fungi: (see page 962) (resupinate) part
– the rosette fungi have spathulate, (reflexed)

cauliflower to rosette-shaped
fruitibodies, with or without a ± well
developed stem (page 952).
– thick-fleshed, resupinate hydnoids
are treated on page 1054.
– resupinate polypores with thick-fleshed hydnoids
labyrinthine pores have sterile pore page 1054
margins (page 932).
– plant parasites on living stems and
resupinate
leaves, see Exobasidiales (page 1214) polypores
and Taphrinales (page 1622).

Further reading: 36, 50, 67, 74, 75, 81,

99, 159, 163, 339.

rosette-fungi Exobasidiales Taphrinales © Thomas Læssøe & Jens H. Petersen

page 932 Fungi of Temperate Europe,
page 952 page 1214 page 1622 Princeton University Press, 2019
962 capped corticioids with smooth hymenophore Porostereum capped corticioids with smooth hymenophore 963
1 species Thelephora
page 967 (see page 956)
Laurilia
Laxitextum
Capped corticioids 1 species,
1 species Micro-drawings:
page 967 spores first,
This form group includes capped page 966 followed on the
inside by cystidia.
(reflexed/stereoid) corticioids with Veluticeps cl. = clamps.
a smooth hymenophore. Capped Plicatura 2 species
1 species page 967 Approximate
genera with a wrinkled or echinulate page 966
Cystostereum species number
hymenophore are shown in the blue applies to
1 species
part of the wheel and are treated Cylindrobasidium page 968 temperate Europe.
under merulioid corticioids (page 974) (see page 1014) felt-like; with felt-like; flesh
and echinulate corticioids (page 1036). brown flesh; brown; on soil,
felt-like; with deciduous
Many species in this group have stems and wood perennial;
brown flesh; wood on Picea
caps with colours, hairs and other deciduous perennial; dimitic
cottony; wood Chondrostereum
useful characters. With experience with skeletal
wrinkled with 1 species
I+ hyphae; brown rot
quite a few can be recognized without Aleurocystidiellum age; deciduous page 968
in conifer wood
2 species wood
the use of a microscope. All capped page 965
annual; I+ Stereum
corticioids may start out without a cap, perennial; smell of
cracking; coconut cake; 7 species
so always look for older fruitbodies. deciduous page 970
J+ conifer wood
Phylogenetically the species are wood
distributed in a number of groups that small,
Peniophora but thick
are not closely related, including orders 22 species; ± purple;
and
dominated by corticioid, poroid or only a few with cap I+ deciduous wood
perennial;
page 965
lamellate species. deciduous
Most capped corticioids are wood
decomposers and form a white rot. with mainly perennial;
loosening ± reddening, ±
Serpula (page 985), Pseudomerulius hairy; deciduous and
margin;
(page 981) and Veluticeps (page 967) enophore ± smoot I+ conifer wood Amylostereum
Dendrocorticium
deciduous +/- cl. hym h 3 species
are brown-rotters and species in the wood
page 969
5 species
genus Thelephora (page 956) form page 964
ectomycorrhiza. - cl. I+ not reddening;
margin capped
conifer wood
loosens; corticioids
deciduous with caps from
Other similar fungi: wood the upper part
dendro-
– annual, capped polypores always hyphidia of resupinate - cl. I+
have proper pores on the underside of fruitbodies perennial,
sp seta
iny cracking;
the caps (check with a hand lens) (page Terana ed
(see page 1024) indigo-blue; wrinkl Quercus
886). deciduous
– rosette-fungi either have a stem or wood seta Xylobolus
- cl.
1 species
narrow towards the base (page 952). very stiff; with page 970
projecting setae
(strong lens!);
See also the overview and the refe­ rubber-like; Quercus
rences to further reading on page 960. deciduous
wood stiff and elastic;
Cytidia I+ - cl. with projecting
+/-
1 species cl. setae (strong lens!); Hymenochaete
page 964 deciduous wood 5 species
covered in long, page 973
rigid hairs
margin ±
+/- loosening
cl. I+
mostly brownish-
other similar rose, with pale large and
fungi margin rubber-like Hymenochaetopsis
whitish-grey 2 species
with small caps
page 973
mainly in houses bright orange
Gloiodon
(see page 1060 Byssomerulius
(see page 977)

Merulius
Steccherinum (see page 978)
(see page 1949)

annual, capped polypores rosette-fungi Plicatura

Irpex © Thomas Læssøe & Jens H. Petersen
page 886 page 960 Serpula Pseudomerulius (see page 63) Fungi of Temperate Europe,
(see page 1048)
(see page 985) (see page 981) Princeton University Press, 2019
974 merulioid corticioids – veined-wrinkled hymenophore merulioid corticioids – veined-wrinkled hymenophore 975

Phlebia
Merulioid corticioids 20+ species
page 978 Micro-drawings:
This form group includes species with spores first,
Merulius followed on the
resupinate and reflexed fruitbodies 1 species inside by cystidia.
that have a wrinkled or veined page 978 cl. = clamps,
hymenophore. All such species were Meruliopsis Lindtneria bas.= basidium.
once included in the genus Merulius. 1 species 5 species
page 977 page 980 Approximate
Some merulioid corticioids are very species number
distinctive and can easily be recognized Pseudomerulius applies to
2 species, page 981 temperate Europe.
without the aid of a microscope, while
others, e.g. species in Leucogyrophana,
Serpula & Pseudomerulius are very rubber-like, resupinate;
difficult to separate without using rubber-like, brownish-lilac, greyish to orange
micro-characters. The majority are capped;
cottony; yellowish
± flesh-pink
white-rotters, but species related to Byssomerulius
to yellow-orange;
basidia cyanophilous
the dry rot fungus Serpula lacrymans 3 species
page 977 rubber-like;
produce brown rots. Hydnomerulius
± capped;
1 species
brownish-rose to
mostly capped; page 981
coral-red
Other similar fungi: yellow-orange
– the wrinkled hymenophore grades with well-defined
into the spiny corticioids (page 1036). margin
– polypores with labyrinthine - cl.
hymenophore (page 932) have sterile whitish to bas.
pore margins, whereas the species ± pale brown
covered here have fertile margins. medallion- labyrithine
white rot clamps dentate
Ceraceomyces - cl.
See also the overview and the refe­ 5 species
rences to further reading on page 960. page 976 Leucogyrophana
merulioid 5 species
corticioids page 982
whitish to form flat or reflexed ± yellow-
fruitbodies with a brown
± pale brown orange with
veined-wrinkled rot
woolly, whitish
hymenophore margin
(underside)

smell of other distinctly wrinkled;

naphthalene similar fungi greyish-violet to
(moth balls) brownish with
woolly, whitish
margin
wrinkled hymenophore

I+
small with
at most somewhat
loosening
wrinkled; yellowish-
margin,
brown with woolly,
Merulicium gradually a I+
resupinate whitish margin Serpula
1 species small agaric
page 976 spiny corticioids polypores 3 species,
page 984
cottony; caps
develop over time
hymenophore with radiating wrinkles

Plicatura crispa Coniophora

(see page 63) (see page 1034)

Plicatura nivea
(see page 966)
veined hymenophore
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
page 1036 page 932 Princeton University Press, 2019
986 fragile corticioids with smooth to warty hymenophore Gyrophanopsis Hyphoderma
fragile corticioids with smooth to warty hymenophore 987
1 species (see page 1004) Peniophorella
page 994 (see page 1006)
Litschauerella
Fragile corticioids 2 species
Micro-drawings:
Bulbillomyces spores first,
This rather poorly defined form group Thanatephorus page 994 followed on the
1 species
(see page 1178) page 995 Subulicystidium inside by cystidia
includes resupinate corticioids where and basidia (bas.).
1 species
the fruitbody is relatively loosely page 995 cl. = clamps.
attached to the substrate. They may be
Approximate
either cobweb-like, cotton-like or flaky Botryobasidium species number
and fragile. Species with thin, waxy or 15 species Lyomyces applies to
gelatinous fruitbodies are not included page 992 anonymous pale (see page 1007) temperate Europe.
downy corticioids; with anonymous pale
here, but amongst the firm, white or from corticioids; with
± projecting
coloured corticioids with a ± smooth downy cystidia cystidia ± projecting with globose,
from cystidia cystidia
hymenophore. smooth asexual
spores state; in very wet Tubulicrinis
Species in this group are best Athelia
places (see page 1007)
11 species repeating with globose,
identified by using a compound
page 991 hairy asexual
microscope but macro-morphology state
may provide useful clues. Since all 4–8-spored; thin and white,
mostly with an often with
the species included have a loose asxeual state amyloid cystidia
hyphal structure it is easy to prepare Membranomyces
flat preparations for use under the 3 species
microscope – and as the hyphal page 996
Leptosporomyces flaky, with aberrant
structure can be seen clearly, they are 4 species whitish cystidia
page 991
much easier to work with than the
- cl.
waxy or gelatinous species. basidia with
flaring sterigmata, Sistotrema
flaky with +/-
cl. 2–4-spored 26 species
Other similar fungi: brownish- page 996
– whitish to pale grey corticioids with yellow hyphal
cords +/-
firm, tougher, waxy structures (page cl.
Piloderma basidia with
1000). 3 species white-
rotter
s
flaring sterigmata,
page 990 ectomyc bas. 4–8-spored
– coloured corticioids with firm, orrhiz
al
bas.
+/- cl.
flaky with fragile corticioids Trechispora
tougher, waxy structures (page 1012). +/- cl. 24 species
yellow form flat (resupinate) ampullate swelling mostly with page 997
hyphal cords fruitbodies with a smooth or
See also the overview and the refe­ warty hymenophore ampullate hyphae
and crystals
rences to further reading on page 960.
+/-
cl. +/-
cl.
flaky, sp. blue downy from cystidia;
blue-green in KOH with skeletal hyphae
other +/- Odontia
+/- +(-) cl.
+/- cl. similar fungi cl. 2+ species
cl. page 1053
wh
ec ite
to
my
-ro
tt
flaky, co
rr
er
s with bluish or
Byssocorticium hiz
greenish colours
5 species, page 990 brown to al
sordid yellow +/-
cl.
I+ firm, whitish
(dextrioid) to pale grey Amaurodon
corticioids 4 species
pale yellow or
firm, page 997
on soil and brown
coloured
litter corticioids mostly
-rotters

Tretomyces ± coloured
1 species
page 989 on soil and litter
brown

mostly in Tomentellopsis
Amphinema 3 species
3 species very wet
places page 998
page 989

Cobweb-like Loosely constructed

fruitbodies of fruitbodies of
Botryobasidium with Tomentella with Tylospora
8-spored basidia in 4-spored basidia. 2 species
close up. page 988 Tomentella
page 1012 30+ species
Jaapia
© Thomas Læssøe & Jens H. Petersen 2 species page 998
Fungi of Temperate Europe, page 1000
Princeton University Press, 2019 page 988
1000 smooth, firm, whitish to greyish corticioids Hypochnicium Tubulicrinis smooth, firm, whitish to greyish corticioids 1001
14 species Lyomyces
20+ species, page 1007
Peniophorella page 1005 5+ species, page 1006 Hyphodontia
10+ species 5+ species
Smooth, firm, ± whitish corticioids page 1006 pages 1008
This form group includes smooth, firm, Hyphoderma & Kurtia Scopuloides
15+ species 2 species
relatively well-attached, resupinate Radulomyces
page 1004 page 1008
3 species
corticioids with whitish or pale greyish page 1009
hymenophores. The texture of the
fruitbodies may be cheesy, waxy or Bulbillomyces
leathery. (see page 995) Phlebiopsis
2 species
Most species in this group, also page 1009
colloquially called ‘white paint’, can with conspicuous often on
only be recognized with the aid of a cystidia Sambucus with
with
microscope and suitable reagents. conspicuous dental drill-like
waxy;
Dendrothele cystidia cystidia
Basidia attached to the side, rather downy from
8 species mostly with projecting
than at the base, are termed pleural. page 1003 conspicuous cystidia
cystidia waxy; Gloeohypochnicium
with phenolic 1 species
Other similar fungi: smells page 1010
with globose
– the distinction between this form ‘bulbils’; in very wet
habitats - cl. stearin-like
group and the fragile, white
and very large
corticioids is not clear-cut (page Aphano­
986). basidium
3+ species on living
– dark grey and pinkish-buff page 1002 Vuilleminia
bark with strong 4 species
species are treated amongst - cl.
fruity smell page 1010
the attached, coloured
corticioids (page 1012).
basidia
See also the overview and the pleural dendro- erumpent
Xenasma hyphidia through
refe­rences to further reading 17+ species bark
on page 960. page 1002
bas.
attached, +/- cl.
white-rott s
basidia ers
brown-ro
tter
pleural, spore whitish to
greyish corticioids bas. with extremely
ornament visible
when stained bas. with flat fruitbodies long basidia;
with a smooth or a brown-rotter Chaeto­
bas. +/- cl.
warty hymenophore white-ro dermella
tters
1 species
bas. +/- page 1011
cl. I+
sterigmata - cl. - cl.
+/- cl. with
inflated
other gloeocystidia
- cl.
similar fungi
bas.
I+
bas.

Tulasnella basidia with rather small,

(see page 1176) longitudinal fragile corticioids woody and
bas. I+ Gloeocystidiellum
walls perennial 5 species
page 1011

I+
basidia with woody and
longitudinal perennial
- cl.
walls
firm, coloured
Sebacina corticioids
(see page 1173) basidia in woody and Aleurocystidiellum
rows perennial 1 species
basidia with page 965
septa and
Micro-drawings: ‘pouch’ Xylobolus
spores first, (see page 970)
followed on the
inside by cystidia.
Exidiopsis
cl. = clamps.
(see page 1172)
bas. = basidia

Approximate species Stereum

number applies to Basidiodendron (see page 970)
temperate Europe. (see page 1175)

Helicogloea © Thomas Læssøe & Jens H. Petersen

Fungi of Temperate Europe,
(see page 1166) page 986 page 1012 Princeton University Press, 2019
1012 ± smooth, firm, coloured corticioids Corticium & Terana
± smooth, firm, coloured corticioids 1013
Marchandiomyces Vuilleminia Peniophora
1 species 4 species 22+ species
5 species page 1024
page 1023 page 1025 page 1020
Smooth, firm, coloured Tubulicrinis
Aleurodiscus
Cytidia 11 species
corticioids (see page 1007) (see page 964) page 1026
Phlebia
This form group includes resupinate 30+ species Xenasmatella
corticioids with buff, brown, dark page 1019 17+ species
grey or vividly coloured, smooth to page 1018
warty hymenophores. The texture
of the fruitbodies varies from
gelatinous, cheesy, waxy to leathery.
erumpent Scytinostroma
Some of the more brightly indigo-blue through waxy, drought- 5 species
± pink
coloured species can be identified in bark resistant page 1027
cystidia discoid,
the field, but the more anonymous gelatinous
Crustoderma thick-walled Conferticium
require microscopic examination.
1 species often discoid; 2 species
It is difficult to make suitable thread-like;
page 1018 spores very page 1028
basidia pleural
preparations of the waxy species. large
The preparations are hard to thick and +/- perennial;
cl. smell of
make sufficiently thin and the Dacryobolus waxy
naphthalene
tissues may be filled with lots 2 species
page 1017 +/-
of ‘goo’ that makes it difficult thick and cy. Megalocystidium
waxy; I+ 2 species
fo spores
to make out the details. brown-rotter fo rm
rm ing very finely page 1028
in I+
g aw verrucose
a +/-
br hite
Other similar fungi: ow r +/- bas. cl. - cl.
thick and n ot cl.
– species with completely
ro
t
+/-
Phanerochaete & leathery; cl. confluent,
+/- I+
white or pale grey fruitbodies Rhizochaete brown-rotters cl. thick, small Gloiothele
11 species fruitbodies 1 species
(page 1000). dendro-
page 1016 page 1029
– species with fragile, cobweb- hyphidia I+
like or flaky fruitbodies (page often with
hyphal cords +/- - cl. with milk and
986). cl. bitter taste
I+
– resupinate jelly fungi (with Candelabro­ ± smooth,
chaete firm, coloured - cl.
divided basidia) (page 1162). Vesiculomyces
3 species mostly with corticioids 1 species
- cl. - cl. often
page 1015 fimbriate with a smooth or page 1029
- cl. fimbriate at
See also the overview and the margin warty hymenophore
margin
refe­rences to further reading
on page 960. cracking upon +/- I+ leathery and
drying, often cl. perennial;
with narrow bas. hyphae often Stereum
Cylindrobasidium - cl.
1 species cap - cl. reddening & Xylobolus
- cl. 7 species
page 1014 I+
+/- (see page 970)
mostly whitish, cl.
bas. leathery and
± brownish
+/- perennial
with age bas. cl. I+
+/-
cy. - cl. I+/-
brown with brown with Amylostereum

fo
boat-shaped +/- cl. asterosetae 3 species

rm
spores

in
page 1030

g
t
Hyphoderma

a
e ro

w
and the like brown with

hi
al

sterigmata
whit

te
dichohyphae
rrhiz

(see page 1004) inflated

ro
with

t
ing a
myco

Micro-drawings: soft, waxy; projecting

Luellia setae (strong
form

spores first, basidia with Asterostroma

ecto

3 species longitudinal walls lens!)

followed on 2 species
page 1015 brown-rotters
the inside by basidia page 1030
brown to dark grey, rather loose
cystidia. helicoid-curved
very tough textured
cy. = cystidia
cl. = clamps.
bas. = basidia
Dichostereum &
I+ = amyloid
Vararia
7 species
Approximate Tulasnella page 1031
species number (see page 1176
applies to Hymenochaete &
temperate Sebacina Hymenochaetopsis
Europe. (see page 1173) 6 species
Coniophora page 1032
7 species © Thomas Læssøe & Jens H. Petersen
Helicobasidium Pseudotomentella Tomentella Fungi of Temperate Europe,
(see page 1168) 5+ species, page 1035 (see page 998) page 1034
Princeton University Press, 2019
1036 spiny corticioids spiny corticioids 1037
Lyomyces Resinicium
Basidioradulum
Hyphodontia & Xylodon 6+ species 3 species, page 1046
Spiny corticioids 15+ species page 1045 1 species
Micro-drawings:
page 1047
pages 1044 & 1045 spores first,
This form group includes resupinate Peniophora followed on the
and reflexed, thin-fleshed cortici­ (see page 1020) inside by cystidia.
oids with a dentate to spiny Dentipellis Irpex cl. = clamps.
1 species bas. = basidia
hymenophore. A few species page 1043
1 species
I+ = amyloid
page 1048
can be identified in the field, but hy. = hyphae
Sistotrema
in general the most important 26 species
Approximate
characters are the appearance page 1042
species number
of the spores and basidia, and in effused, somewhat applies to
with short temperate Europe.
some cases the types of cystidia spines
waxy with coarse
effused, with ‘teeth’
and hyphal structures, all of which Kavinia short spines coarsely warty-
require microscopy. 1 species Steccherinum
spiny, waxy
page 1041 effused, with 7 species
long spines page 1049
Other similar fungi:
effused, with ± effused-reflexed
– the hydnoids are more thick- Mycoacia
long spines; 3+ species
fleshed and often have a cap and basidia page 1019
stem (page 1054). 6–8-spored
Hydnocristella mostly brownish-
– pendent clavarioids form ‘clubs’ 1 species green with I+
rose with pale
directly on the substrate, rather than page 1040 FeSO4; margin
spines on +/-
in connection with a subiculum or subiculum cl.
the flesh (page 1092). Radulomyces
– the merulioid corticioids have a ± waxy 2 species
green with page 1051
veined hymenophore, which may FeSO4;
partially split into rounded ‘teeth’ Trechispora spines on
subiculum bas.
(page 974). 24 species
page 1039 waxy;
– the pores of annual, resupinate with phenolic
polypores may, with age, split into smell
cottony; with Amaurodon
‘teeth’ (page 932). ampullate spiny corticioids 6 species
hyphae form flat or effused- page 1052
reflexed fruitbodies spore blue
See also the overview and the refe­ a with spiny/ in KOH blue-green
rences to further reading on page Irpicodon dentate lower side whit
bro e-rot
I+ (hymenophore) wn- t
rot ers asteroseta
960. 1 species pendent ec
to ters
page 1039 my
clusters; w
hi
co
rr - cl.
on Pinus te
-ro hiz
al with brownish Asterodon
tt
er
flesh and brown 1 species
s hyphae
other page 1052
somewhat similar fungi hy.
waxy;
basidia with bas. + spores without flesh, but labyrinthine-
cyanophilous cyanophilous
with ± isolated dentate; brown-
drops spines rotter
somewhat
waxy; bas.
Hydnomerulius
basidia with with brown (see page 981)
Lindtneria cyanophilous hyphae
(see page 980) drops

on Carex or ferns
thick-fleshed pale brown
and spiny with brown
somewhat hyphae Tomentella
gelatinous 30+ species
Cristinia page 1052
4 species tough
page 1038 clavarioids & jelly fungi
page 1092 & 1162

veined Odontia
Epithele 2+ species
2 species page 1053
page 1038

Stypella
(see page 1171)
spines (round in section) ‘teeth’ (flattened) Heteroradulum © Thomas Læssøe & Jens H. Petersen
(see page 1174) Fungi of Temperate Europe,
hydnoids, page 1054 merulioid corticioids, page 974 Princeton University Press, 2019
1054 hydnoids (with spiny hymenophore, rather thick-fleshed or with stems) hydnoids 1055

Gloiodon Auriscalpium Bankera

Hydnoids 1 species 1 species 2 species
page 1060 page 1060 page 1061
The hydnoids are defined here as Hericium Phellodon
4 species 5 species
either stemmed or thick-fleshed page 1062
page 1058
fungi with a spiny or dentate
hymenophore. Fungi with thin, Hydnellum
14+ species
resupinate, spiny fruitbodies are page 1064
treated under the form group
corticioids (page 960).
The hydnoids constitute a
form group of species that, for
example, belong to the orders large and rather
Thelephorales, Cantharellales and crisp; bulb-
shaped, with
Russulales. caps or heavily
branched tough;
with brown,
Other similar fungi: on cones crisp;
broadly adnate
– spiny species with flat small caps smell of curry
(resupinate) fruitbodies and thin
tough;
flesh are included under the smell of curry
corticioid fungi (page 1036).
– species with isolated spines that I+
lack a common flesh are treated I+
tough;
under the clavarioids (page 1092). on without smell of
I+ cones; curry Sarcodon
with 15+ species
Further reading: 50, 142, 163, 221, stem page 1068
229, 346.
Spongipellis ± crisp;
(see page 906) without smell of
hydnoids curry
on wood; form hoof- on soil;
without shaped, branched with stem
stem, or stemmed
some are fruitbodies with
branched a spiny underside crisp;
Micro-drawings: tough and with (hymenophore) cap smooth and
spores and cystidia ± flat teeth pale or apricot-
orange
Approximate species
number applies to other
temperate Europe. similar fungi

thin-fleshed small; with tooth-

like spines to
Climacodon labyrinthine pores
1 species Hydnum
page 1056 11 species
small; with very page 1072
small, crowded
huge, tough spines
with small caps

Sistotrema
1 species with stem
page 1075
isolated, hanging
spines
Sarcodontia
1 species
page 1057
thick, yellow and Mycorrhaphium
± resupinate; 1 species
mainly on Malus page 1075

© Thomas Læssøe & Jens H. Petersen

spiny corticioids clavarioids Fungi of Temperate Europe,
page 1036 page 1092 Princeton University Press, 2019
1076 cyphelloids Woldmaria Ceriporia cyphelloids 1077
Henningsomyces & 1 species 7+ species
Rectipilus page 1083 (see page 946) Porothelium
4+ species Resupinatus
Cyphelloids page 1082
1 species
3+ cyphelloid species
Micro-drawings:
page 1084 spores first,
This form group includes page 1084 then surface
Cyphella hairs.
basidiomycotes with small cup-shaped, 2 species
bell-shaped or tubular fruitbodies page 1081 Approximate
on various substrates. The largest species number
Schizophyllum applies to
fruitbodies are up to 10 mm wide 2 species temperate
(Schizophyllum amplum), while the page 1081 Europe.
Chromocyphella
smallest are only around 0.2 mm wide 1 species
orange-brown, polypore, but page 1085
(Pseudolasiobolus minutissimus). The tubular; initially cup-shaped numerous cups
whitish,
fruitbodies are generally positioned tubular; on Matteuccia on a subiculum on a subiculum
Cellypha rather large,
with the hymenium pointing down- 1 species
on wood with fimbriate small, often
brown, bell- margin
wards, allowing the spores to fall freely page 1079 shaped; on Abies extremely
crowded Crepidotus
when released. and deciduous
1 cyphelloid species
wood fruitbodies
Most cyphelloids are reduced large, with brown page 1085
small, with pale
agarics, i.e. members of Agaricales hymenium; brown hymenium;
that have become gill-less during the on Salix and moss parasite
Populus
process of evolution. Some polypores, Clitopilus
rather small; small, whitish,
e.g. species of Ceriporia (page 946), 1 cyphelloid soon flat, with
species mostly with veins
may initially recall cyphelloids but, with and drops; dry, tinged hymenium;
(see page 88) on bark and litter Episphaeria
age, the cups join to form ‘normal’ on herbs, rarely
1 species
polypore fruitbodies.The numerous wood
page 1086
separated tubes in Fistulina hepatica very small, whitish
(page 845) and the unusual ‘gills’ of medium-sized, with tinged
with rose spores; hymenium;
Schizophyllum commune (page 62) on Fraxinus
on miscellaneous
can be interpreted as many cyphelloid substrates
fruitbodies joined on a common fleshy
small; with projecting
meta-structure. These species are also hairs; on herbs and Lachnella
nested within the Agaricales. twigs, tolerates 2 species
medium-sized; cyphelloids
dessication page 1086
hanging bell- form cup-shaped,
shaped; mainly bell-shaped to tubular,
Other similar fungi: hanging fruitbodies
Calyptella on herbs
– many species of e.g. Hyaloscyph- +3 species
on wood, herbaceous extremely small; with
aceae may look similar, but the stems, leaf-litter and projecting hairs; on
page 1080
living mosses wood and herbs
apothecia can point in all directions
and they tend to look neater and less medium-sized,
white; moss Pseudolasiobolus
irregularly hairy. They form spores in small; with long 1 species , page 1087
parasites
asci (page 1372). other
projecting hairs,
crystal-tipped;
– some rosette-fungi recall large similar fungi
on litter
cyphelloids, but cyphelloids are never
Rimbachia rather large,
long-stemmed (page 952). 3 species grey to white;
– pleurotoids are typically larger, with page 1078 parasite on small; hairs with
with asci
more well-developed gills (page 60). mosses; clamps flagella-lke tips; on Seticyphella
absent with herbs, wood etc. 2 species
basidia page 1088
Further reading: 5, 6, 40, 156, 186. large,
ear-shaped, small; hairs with
wrinkled lower glabrous, swollen
Arrhenia side tips; on ferns
1 cyphelloid
species small; with brown
(see page 64) hairs; gregarious
Flagelloscypha
or in clusters on
small; with brown 8+ species
wood and bark
hairs; on wood and page 1089
herbs
Cephaloscypha
Campanella 1 species
1 species page 1088
(see page 63)

Merismodes
2+ species
page 1090
Maireina © Thomas Læssøe & Jens H. Petersen
rosette-fungi pleurotoids inoperculate cup fungi 2+ species Fungi of Temperate Europe,
(page 952) (page 60) (page 1348) page 1091 Princeton University Press, 2019
1092 clavarioids clavarioids 1093
Pterula
4+ species
page 1100
Clavarioids Multiclavula © Thomas Læssøe & Jens H. Petersen Micro-drawings:
3 species Fungi of Temperate Europe, spores first,
Clavarioids includes Basidiomycota page 1114 Princeton University Press, 2019 then basidia
with normal, one-celled basidia and (bas.) and
thread-like, awl-shaped, clavate or Pistillina cystidia (cy.).
branched fruitbodies. Most clavate 1 species
Typhula page 1099 Approximate
fruitbodies are negatively geotropic 35+ species species number
(they grow/point upwards), but some page 1102 applies to
Clavicorona
temperate
species with very small fruitbodies 1 species
Europe.
page 1130
are ageotropic (the orientation is
random) and species in the genus
Mucronella ▷▷ are positively geotropic
(they point downwards).
The clavate fruitbody is one of the
simplest, most basic types of fungal Physalacria orientation Artomyces
orientation random
2 species random; very 2 species
fruitbody, and clavarioids have arisen page 1099 small; with
or upwards; small
page 1130
and ± pointed; with
numerous times through the evolution sclerotium; with skeletal hyphae upright orientation;
of the fungal kingdom. Clavate marginal hairs; medium-sized;
on Molinia lichenized
fruitbodies occur in Ascomycota (e.g.
orientation random upright orientation;
earthtongues), within the rust fungi, medium-sized; white;
or upwards; small
the jelly fungi and in numerous places to medium; ± with with widened, almost
amongst the remaining Basidiomycota sclerotium cup-shaped tips Lentaria
with one-celled basidia, e.g. amongst 3 species
page 1132
the Agaricales. orientation
Hirticlavula random; very small; marginal
The group of clavarioids with hair upright orientation;
1 species cystidia in the medium-sized; brown,
one-celled basidia includes about 20 page 1098 hymenium; e.g. on hairs with widened tips
genera, with around 200 species in Cryptomeria
temperate Europe.
Many small clavarioids live as I+
decomposers of plant material, orientation ± upright
e.g. on stems of herbs or leaves. random; orientation;
Some species of Typhula ▷▷ may extremely small; cy.
medium-sized;
with upturned green with
parasitize plants. Amongst the larger hairs; on bark FeSO4; on wood
clavarioids the genera Ramaria ▷▷,
Ceratellopsis hair
Clavariadelphus ▷▷ and Thele­ 3+ species, small,
phora ▷▷ are ecto­mycor­ page 1097 thin, unbranched
or slightly branched
rhizals. orientation clavarioids
Like the hygrocyboid random or – mostly on wood or
downwards; stems
fungi (page 142), some
very small; larger, more than 2 mm thick
or all species of Clavaria, ± pointed or highly branched clavarioids
Clavulinopsis and – on wood, stems or soil,

⬅
Ramariopsis ▷▷ may live see next page spread
other
in a not yet understood I+ similar fungi
symbiosis with herbs (page
with with
18). These species are mostly downturned basidia asci
very demanding with regard spines
to habitat, and, among
other things, are sensitive to fertilizers, bas.
preferring habitats with a long
continuity. They often share habitat
with a number of rare hygrocyboids,
entolomatoids, earthtongues – e.g.
in unimproved grasslands or in damp Mucronella
forests on mull soils – and they are 3+ species
page 1096
particularly good indicator species of
especially biodiverse fungus habitats.

Continued on next page spread . . .

club-shaped to tongue-
Calocera Eocronartium Phleogena Gymnosporangium Acrospermum shaped Ascomycota
(page 1159) (page 1184) (page 1246) (page 1208) (page 1523) (page 1350)
1094 clavarioids clavarioids 1095

 Clavarioids Ramariopsis
6+ species Micro-drawings:
continued from previous page spread . . . page 1121 Pterula spores first,
4+ species Clavulina then basidia (bas.)
Clavulinopsis page 1100 6+ species Artomyces and cystidia (cy.).
Other similar fungi: 15+ species page 1128 2 species
page 1125 page 1130 Approximate species
– species of Calocera look similar but number applies to
have rubbery textures and tuning temperate Europe.
fork-shaped basidia (page 1159).
– greyish-white species of clavarioids
with septate basidia are treated with Clavaria
the jelly fungi (page 1164). 20+ species
page 1116
– orange-yellow, gelatinous clavarioids Lentaria
with teliospores that germinate with with skeletal 3 species
hyphae; page 1132
septate basidia belong to the rust on debris
basidia 2-spored;
sterigmata bull-horn
fungi (page 1208). basidia with
shaped; on soil
– Ascomycota with club-shaped normal clamps;
on soil with flaring tips;
apothecia are included with the earth basidia with on deciduous
tongues and others (see page 1350). normal clamps; wood
– Acrospermum is a small, dark, on soil
clavate Ascomycota on herbaceous
stems with internal asci (see page
green with
1523). basidia without or FeSO4; on
with open clamps; bas.
– Ascomycota with perithecia conifer debris
on soil
immersed in a clavate stroma are
included with the pyrenomycetous Clavariadelphus
fungi (see page 1510). 5 species
Alloclavaria
– for small, club-shaped fruitbodies 1 species with cystidia; page 1112
green with FeSO4;
with internal spore production and page 1115 on soil or needle bas. large and fleshy;
I+
debris on soil
passive, dusty spore dispersal, see
Phloeogena, asexual fungi and myze­
to­zoa (pages 1246, 1626 and 1646). Macrotyphula cy.
2+ species larger, more than
page 1110 2 mm thick or heavily
Further reading: 31, 32, 38, 44, 63, 92, green with
branched clavarioids FeSO4; spores
100, 142, 143, 230, 232, 243, 244. late season; on – on wood, stems cinnamon-buff;
twigs and leaf- or soil on litter
litter
cyanophilous
ornament

Phaeoclavulina
other green with 15 species
similar fungi FeSO4; spores page 1134
cinnamon-buff;
small, thin, unbranched on wood or soil
or slightly branched
clavarioids – mainly on
with with
wood or stems, basidia asci Ramaria

⬅
see previous page spread 60+ species
page 1138
asci formed asci formed
externally in perithecia

flesh dark brown;

on soil; mycorrhizal

club-shaped to stromatic
tongue-shaped pyrenomycetous
© Thomas Læssøe & Jens H. Petersen Calocera Tremellodendropsis Ascomycota fungi Thelephora
Fungi of Temperate Europe, 9+ species
Princeton University Press, 2019 (page 1159) (page 1197) (page 1350) (page 1510)
page 1152
1154 Dacrymycetales – with tuning fork-shaped basidia and budding spores Dacrymyces 1155

Dacrymycetales Dacrymyces form small, cushion-

The Dacrymycetales is a natural
group characterized by tuning fork-
shaped basidia, mostly septate spores
that bud off microconidia, and by
the predominantly orange-yellow,
gelatinous to rubbery fruitbodies.
Mature spores are best studied from
deposits on slides. Some species form
simple asexual spores (arthrospores)
from structures that are similar to the
fruitbodies, but softer.
Dacrymyces
13 species
page 1155
The individual genera are poorly
delimited in relation to each other,
and the largest genus, Dacrymyces, is
not monophyletic.
All species are wood-decayers and
form a brown rot. Some species, e.g.
Dacrymyces stillatus, can cause serious
damage to timber, including wooden
window frames.
Further reading: 50, 142, 163, 265.
Ditiola
2 species
page 1157
Femsjonia
1 species
page 1157
shaped to turbinate fruitbodies
that may aggregate in resupinate
structures. The spores have up to
nine septa. All are wood-degraders
and brown-rotters.
Dacrymyces stillatus often occurs
in both a sexual and an asexual state
on the substrate. The asexual state
is reddish-orange and soft, and can
Development of basidia (left); types of basidia easily be squashed (lower image),
(right). while the sexual state is adnexed,
Micro-drawings: translucently yellow and firmer, more
spores (outer rubbery-gelatinous. The hyphae lack
part of wheel) clamps. The basidia are tuning fork-
and spores
budding off shaped. The thick-walled spores are
microconidia flattened on one side, at maturity 2 mm
(inner part).
have three thick septa and measure
Approximate 12–14 × 3.5–4 µm; they bud off
species number small, globose conidia from each
applies to
temperate cell. Occurs on moist, rotten wood of
Europe. coniferous and deciduous trees.
May be difficult to separate from
D. lacrymalis ▽ but that species lacks
the prominent asexual state and has
somewhat thinner-walled spores.
Widespread and very common; all
year.

overmature
spore with Guepiniopsis
microconidia 4 species
page 1158
2 mm

Dacrymyces lacrymalis is bright

yellow, trans­lucent, wrinkled,
narrowly attached, medium-sized,
Dacrymycetales Dacrymyces that does not have a
have tuning fork- proper stem. The spores have three
shaped basidia,
spores that bud somewhat thickened septa and
off microconidia measure 12–14 × 3.5–4 µm. Occurs
and gelatinous-
mostly on decomposed wood from
rubbery, ± yellowish
fruitbodies deciduous trees, more rarely on
Calocera conifer wood.
5 species
with with page 1159 The similar Dacrymcyes stillatus △
asci other basidia is usually accompanied by a reddish
similar fungi
asexual state.
Widespread and common; all
year.

1 mm

© Thomas Læssøe & Jens H. Petersen

Orbilia clavarioids jelly fungi Fungi of Temperate Europe,
page 1464 page 1092 page 1162 Princeton University Press, 2019
1162 jelly fungi (fungi with aberrant basidia and repeating spores) Exidiopsis Sebacina jelly fungi (fungi with aberrant basidia and repeating spores) 1163
12 species 5 species
page 1172 page 1173
Protodontia & Stypella Protohydnum Basidiodendron
Jelly fungi 9+ species 1 species 9 species, page 1174 Micro-drawings:
page 1171 spores first,
page 1169 Heteroradulum
This form group includes species 3 species
Tulasnella thereafter basidia.
29 species cl. = clamps.
that mostly have septate, long, page 1175
page 1176
sterigmate basidia and spores Approximate species
that can usually produce a copy number applies to
temperate Europe.
of themselves (repeating to form
secondary spores). Many species Efibulobasidium
1 species
are gelatinous and have basidia page 1169
imbedded in a protective gel. tough and
The sterigmata find their way thin and cartilaginous; with ±
fragile; on on wood, soil repeating
from within the fruitbody to the wood
with spines; and herbs basidia; on
surface, where the spores are on wood wood with teeth; Scotomyces
Xenolachne 1 species
formed. 1 species
on wood
confluent small above page 1179
The fruitbodies are highly page 1181 fruitbodies; on ground

pta
variable in shape and colour, wood

ia
se
sid
from flat (resupinate), verrucose with inflated

th

ba
on sterigmata;

wi
or spiny, to large, folded, brain-

ed
stems of on wood

ia
ell
sid
like, clavate, branched or herbs

e-c
ba
on
cap-like. One group of jelly fungi
does not even produce obvious basidia Rhizoctonia
in with dark 7 species
fruitbodies, but occurs either Tremella and the like Hymenoscyphus
40 species contents; page 1178
as parasites inside other fungal and Cudoniella on wood
page 1192 hymenia
fruitbodies or forms almost
invisible films on, for example, - cl. basidia short,
+/-
wood. on wood with cl. not forming a
other fungi - cl. true hymenium;
As defined above (aberrant
or directly on herbs and
basidia and repeating spores), Sirobasidium on fungal wood Digitatispora
the jelly fungi is a form group 3 species fruitbodies
+/-
1 species
page 1168 cl. page 1180
that also includes fruitbodies - cl.
that are not jelly-like, and basidia in chains;
on wood with
phylogenetically they usually pyrenomycetous spore on driftwood
jelly fungi at sea shores
belong to the subphylum fungi
with small or simple
Pucciniomycotina and the orders basidia with longitudinal
septa
fruitbodies
Auriculariales, Cantharellales, basidia with septa
Helicogloea
Sebasinales and Tremellales, basidia with Syzygospora
2+ species on other
all of which are within the page 1166 probasidial sac; & Hetero­
on deciduous +/- fungi,
subphylum Agaricomycotina cl. cephalacria
including

⬅
wood 8 species
(page 13). jelly fungi
lichens
page 1196
Jelly fungi are decomposers other with larger or
similar fungi more complex
or parasites on plants or other basidia with on
probasidial sac; fruitbodies, e-
on
fungi. A group of species are next page spread
ce
ll
on wood ed
ba Granulobasidium
part of the so-called ‘orchid with tuning sid
fork-shaped ia vellereum and on
mycorrhiza’, where orchids basidia mouldy insects
parasitize the fungi involved. on stems of Filobasidiella
herbs and 2+ species
Jelly fungi includes more with normal, page 1181
Saccosoma deciduous
than 50 genera, with more wood one-celled
2+ species basidia
than 200 species in temperate page 1167
Europe. In this section, 30 on wood with
genera are covered by the two other fungi
wheels, including the most
on
showy species. The majority of Lasiosphaeria
the genera not treated in this ovina
Helicobasidium
publication form resupinate, 2 species
mostly invisible fruitbodies page 1168

Achroomyces
continued on next page spread . . . 7 species
page 1167
© Thomas Læssøe & Jens H. Petersen corticioids
Fungi of Temperate Europe, Krieglsteinera page 960
Princeton University Press, 2019 1 species , page 1166 Dacrymycetales, page 1154
1164 jelly fungi (fungi with aberrant basidia and repeating spores) jelly fungi (fungi with aberrant basidia and repeating spores) 1165

Aporpium
 Jelly fungi 2 species
page 1186
Ditangium
2 species
Micro-drawings:
spores first,
continued from previous page spread . . .
page 1187 Myxarium then basidia.
10+ species cl. = clamps.
Pseudohydnum
page 1170
or are parasites within other 1 species
page 1185 Approximate species
fungi or plants, e.g. the genera number applies to
temperate Europe.
Bourdotigloea, Endoperplexa,
Hauerslevia, Herpobasidium, Exidia
10+ species
Heteroacanthella, Insolibasidium, page 1187
Kriegeria, Naohidea, Occultifur,
Oliveonia, Phragmoxenidium,
on cherries and
Pseudostypella, Renatobasidium, other species of Tremella, Phaeotremella
on wood with white
Serendipita, Spiculogloea and Prunus & Naematelia
from inclusions; on
Trimorphomyces. deciduous 40 species
wood
trees page 1191

Other similar fungi: Tremiscus

on conifer wood turbinate to expanded
– species of Dacrymycetales are 1 species
and brain-like;
page 1185
also gelatinous but are mostly on wood
yellow and have characteristic, - cl.
tuning fork-shaped basidia. The
spores are mostly septate and do mainly on sawdust
not repeat. However, conidia are and wood chips
+/-
often produced directly from the cl.
spores (page 1154).
– resupinate jelly fungi can be very ba
sid
ia
difficult to tell apart from corticioid ba
sid
wi
th
ia lo
fungi; those, however, have wi
th
sep
ng
itu
din
ta al
one-celled basidia and spores that Auricularia sep
ta
do not repeat (page 960). 2 species
page 1182 cushion-shaped
– Aporpium may be confused with to ± brain-like,
rubbery tough; veined; mostly on
resupinate polypores (page 932). on wood wood on/in other
– Pseudohydnum may be confused fungi
with hydnoids, but hydnoids are jelly fungi
with large or rather
not gelatinous (page 1054). complex fruitbodies with basidia that
– Tremellodendropsis and Eocro­ bud; on sclerotia
basid
nartium may be confused with parasitic on - cl. ia w
ith l
ong on herbs and
bryophytes itudin
clavarioids (page 1092). one-
al se
pta leaves
celle
d ba
– some Ascomycota, e.g. Asco­ sidia Tetragoniomyces
1 species

⬅
tremella, Ombrophila and Bulgaria, Eocronartium jelly fungi
1 species page 1195
superficially recall jelly fungi, but with small other
page 1184 or simple
they form spores in asci (page with internal similar fungi
spore production; fruitbodies, deform fruitbodies
1364). on exposed wood previous page of other fungi
Phleogena spread
(see page 1246) with normal,
Further reading: 50, 70, 100, 142, with one-celled
tuning basidia
163, 176, 189, 190, 192, 222, fork-shaped
268, 269, 270, 271, 306, 307. basidia
on soil, mostly in Syzygospora &
open land Heterocephalacria
with
asci 8 species
page 1196

Tremellodendropsis
2 species
page 1197

gelatinous inoper­ © Thomas Læssøe & Jens H. Petersen

culate cup fungi Dacrymycetales hydnoids clavarioids Fungi of Temperate Europe,
page 1364 page 1154 page 1054 page 1092 Princeton University Press, 2019
1198 rusts and smuts 0 Puccinia
rusts and smuts 1199
188+ species
page 1204
Rusts and smuts
Uromyces
Tranzschelia
4 species I Gymnosporangium
This group includes a number of Micro-drawings: 66+ species page 1210 6+ species
biotrophic plant parasites that do not teliospores (III) and page 1203 page 1208
form proper fruitbodies; all belong to
basidia (bas.). I III
the Basidiomycota. There are about 0: spermogonia
I
I: aecidia Chrysomyxa
30 genera of rusts and 20 of smuts in
temperate Europe; the aim here is only
II: uredia
Cronartium
3 species 5+ species I Triphragmium
III: telia
to provide an overview of this group.
page 1202 page 1202
III 2 species
page 1210
Approximate species
Rusts have a particularly
complicated life-cycle that may involve
number applies to
temperate Europe.
I 0–I on 0–I ± brown,
up to five types of spore, and in many numerous
hosts; 0–I on numerous mostly on
Ranunculaceae;
I
cases a change (shift) in host. The II–III mostly hosts; II–III mostly 0–I on Pyrus, Phragmidium
spore types can be summarized as III 0–I on Picea;
II ± orange, III brownish, on brownish, on
numerous hosts
II–III brown, on
Ranunculaceae or
Crataegus or
Sorbus;
11 species
± white, yellow numerous page 1211
follows. Type 0 (spermogonia with hosts
Rosaceae II–III orange, cells
to red, columnar, with two pores;
spermatia): a basidiospore (Type IV)
germinates on a suitable host and an
both mostly on
Rhododendron
III
bas.
gelatinous; on II+III
Juniperus
internal mycelium develops from which
Melampsora 0–I on Pinus;
III III
0–I on Filipendula;
I
tiny, flask-shaped spermogonia (n) are 14. species II ± orange, cushion- II–III brownish, on
formed in spots on the host’s surface. page 1201 shaped, III ± brownish, III Filipendula
II

te iosp
bas.

te
lio
The spermatia may then transfer nuclei columnar, both on III

l
sp res
herbs or Ribes

or
to suitable compatible hyphae on

o
I–II yellowish,

es ± un
III III brownish,

lo
other spermogonia. Type I (aecidia

ng tem
on members
with aecidiospores): the dikaryotic

st
0–I on trees and

s
of Rosaceae –

em ed
hyphae that result from fertilization herbs;

me
without host shift

m
II–III yellow or III

d
with spermatia may then produce ± III
± brownish, mostly III
discoid aecidia in which dikaryotic on Salix or Populus
(n+n) aecidiospores are formed. Type II es
inial
(uredia with uredospores): in host- I III rusts and smuts are rust
s – pucc
otry
ales
transforms anthers to
purple-brown spore
shifting species the aecidiospores infect 0–I on conifers; micr
ob
parasites on plants powder
II–III orange to
a second host and a new dikaryotic ± brownish, on
where they form Microbotryum
pustules with spores smuts – ustilaginales 30+ species
mycelium develops from which uredia deciduous trees and
of various types page 1212
and uredospores are formed (n+n). herbs
Type III (telia with teliospores): a Type II Pucciniastrum transforms flowers and
III
mycelium finally forms telia with thick- 10 species seeds to a brownish-
page 1201 black spore powder
walled teliospores (n+n) that typically
I–II yellow, III whitish,
overwinter. The following spring the I on Rubus – without
other
similar fungi
teliospores germinate with 4-celled host shift III
basidia after fusion of the two nuclei transforms flowers
and a subsequent meiosis. Type IV: of Carex and a few Ustilago
other Cyperaceae to a 15+ species
basidiospores (n) are then produced brownish black spore
0–I on Anemone; page 1213
and can reinfect the first host. II pale brown, III pale III powder covered by a
Smuts and Microbotryales have Kuehneola red, on members of grey film
1 species Rosaceae
much simpler life-cycles, infecting just
one host and finally producing spores
page 1200 I other kingdoms
than the fungi
Ascomycota Basidiomycota transforms flowers,
leaves and stems
externally or internally. They can, for 0–I whitish, on of numerous plants
Abies; II–III whitish, to brownish-black
example, transform the inner parts of spore powder
on ferns
a flower to spores or may break out
Anthracoidea
through stems, roots, seeds or leaves. 20+ species
Both rusts and smuts are serious
Ochropsora III page 1212
pathogens on crop plants. 1 species
page 1200
Other similar fungi:
– numerous fungi and fungoid
organisms live as parasites on plants, Urocystis
some of which may form structures 35+ species
Milesina page 1212
that resemble those from rusts and Further reading: 9+ species
smuts, see images in the blue section. 78, 80, 164, 325, 330. page 1200
© Thomas Læssøe & Jens H. Petersen
other kingdoms powdery mildews (Erysiphales) Taphrinales Exobasidiales Fungi of Temperate Europe,
page 1645 page 1500 page 1622 page 1214 Princeton University Press, 2019
1214 Exobasidiales
Exobasidiales
The order Exobasidiales has just
one genus of biotrophic parasites in
temperate Europe. It partly deforms
shoots and leaves of the host, which is
also stained red, and forms a whitish
covering of hymenia that have long,
one-celled basidia on some surfaces.
Some species form infections only in
a few leaves, while others produce
systemic infections. The species in
Taphrina and others powdery mildews (Erysiphales)
page 1622
3 mm
temperate Europe occur on shrubs and
dwarf shrubs of the Ericaceae.
Farther south the related parasite
Laurobasidium lauri ⋉ produces horn-
like protuberances on Laurus trunks.
Other similar fungi:
– the ascomycote Taphrina occurs
predominantly on trees and may form
witch’s brooms (page 1622).
– the powdery mildews (Erysiphales)
other kingdoms
page 1500 page 1645
are more mealy and, with age, form
small, spherical fruitbodies with asci
(page 1500).
– some organisms that resemble fungi,
e.g. Albugo, have neither basidia nor
asci (page 1645).
Further reading: 80, 100, 164, 209.
Exobasidiales
is parasitic on species of
Ericaceae that become
deformed (galled) and
red-stained
other
similar fungi
basidium
with
Exobasidium spores
25+ species;
page 1214
Exobasidium oxycocci occurs
within Vaccinium oxycoccos and
V. microcarpum. The annual shoot
of the host stretches and becomes
pale, and finally is completely
covered by erumpent basidia. The
curved, 1–3-septate spores measure
12–15 × 3–3.5 µm.
Exobasidium rostrupii ⋉ occurs
on the same host but only causes
symptoms on the leaves, which
acquire bright red spots on the upper
side and have spore production
underneath.
Widespread and probably rather
common; summer–autumn.
Exobasidiales 1215
Exobasidium vaccinii occurs on
leaves of Vaccinium vitis-idaea. The
leaves discolour red and become thick,
and spores are formed on basidia on
the mealy white lower sides. May also
attack flower buds and young shoots,
which become misshapen. The curved,
1–3(–7)-septate spores measure
11–19 × 2–4 µm; germinates with
microconidia.
Exobasidum splendidum ⋉ and
E. juelianum ▽ occur on the same host
but form larger, systemic infections
that include both leaves and stems.
Widespread and very common
where the host occurs; summer–
autumn.
8 mm
Thomas Læssøe
Exobasidium juelianum is systemic
and discolours the shoots of Vaccinium
vitis-idaea bright red; it also causes
dwarf growth. The spore-producing
pale tissue is seen mostly under
the leaves. The spores are either
one-celled, or have one septum, and
measure 9–14 × 2–4 µm.
Exobasidium splendidum ⋉, on the
same host, is an even red­der, more
northern, 2-spored species with curved,
0–1-septate spores that measure
20–27 × 6–11.5 µm; E. vaccinii △
causes localized infections of single
leaves, but not of whole shoots.
Widespread, common in the
hemiboreal–boreal zones, rarer farther
south; summer–autumn. 5 mm
Exobasidium uvae-ursi infects
plants of Arctostaphylos uva-ursi.
The parasitized plants typically
produce dense, annual shoots with
numerous dark red leaves and
stems. The hymenium is formed on
the underside of the leaves. The
curved, 1–3-septate spores measure
15–22 × 5–6 µm; germinates with
microconidia.
Exobasidium sydowianum ⋉
occurs on the same host but the
infection is limited to just tiny areas
on the leaves.
Widespread in the hemiboreal–
boreal zones, occasional, rare in the
nemoral zones; summer–autumn.
15 mm
Julia Kruse
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
Princeton University Press, 2019
1216 bird's nest fungi Sphaerobolus 1217

Bird’s nest fungi Sphaerobolus stellatus is a unique,

Bird’s nest fungi belong to the All species in the group are – Stictis may superficially recall approximately 2 mm wide gasteroid
Basidiomycota, and produce basidia decomposers and occur typically on Sphaerobolus but lacks peridioles and fungus that opens in a star-like fashion,
and spores within special egg-like large herbaceous stems, wood or forms the spores in asci in a gelatinous after which a ± orange-brown ball (a
structures called peridioles. In dung. hymenium (page 1472). peridiole) is shot several metres away.
Sphaerobolus the single peridiole is During the opening an orange colour is
shot actively away from the fruitbody. Other similar fungi: Further reading: 50, 92, 156, 241. seen (upper image), and after shooting
In other genera the small peridioles are – some operculate discomycetes a pale projecting ‘balloon’ is seen
held within cup-shaped structures, and may superficially resemble bird’s nest (middle image). Spores and basidia
are ‘splashed out’ by heavy raindrops. fungi, but they do not have peridioles are held within the peridiole. The
Traditionally, bird’s nest fungi are and the spores are borne in asci in a hyaline, thick-walled spores measure
included in the gasteroid form group. hymenium inside the cup (page 1278). 6–10 × 5.5–6.5 µm. Occurs on litter,
very decomposed wood, and old dung,
mainly in open habitats.
Distinctive and unlikely to be
Micro-drawings: misidentified. Only one other European
spores. genus, Pilobolus (page 1642), has the
Sphaerobolus ability to propel a spore ball several 800 µm
Approximate
1 species species number metres, but the mechanism used by
page 1217 applies to
Mycocalia the two genera is quite different. In
temperate
4 species Europe. Sphaerobolus the propelling force
page 1218 is created by tensions between six
different wall layers. At maturity, one
of the layers under the peridiole swells,
and the resulting tension results in an
inversion of the upper layer and the
Nidularia
1 species
peridiole is shot away. The inverted
page 1218 layer is the pale ‘balloon’. Pilobolus
propel the spore ball by creating an
internal pressure within the head that
with active dispersal eventually ‘explodes’ and delivers the
of peridiole from with passive
dispersal of necessary force.
a star-shaped
fruitbody peridioles; Widespread and common; mainly
fruitbody June–December.
without lid, less
than 2 mm with passive
dispersal of
peridioles; Crucibulum
fruitbody without 1 species 1½ mm
lid, 4–7 mm page 1219
Jan Vesterholt

with passive
dispersal of white
peridioles;
fruitbody with lid

with passive dispersal

of grey to black
peridioles;
fruitbody with lid Cyathus
bird’s nest fungi
3 species
form the spores in
page 1220
peridioles inside
the fruitbody that
either cracks open
or opens with a
special lid

4 mm
other
operculate Jan Vesterholt
similar fungi © Thomas Læssøe & Jens H. Petersen
discomycetes Stictis and others Fungi of Temperate Europe,
page 1278 page 1472 Princeton University Press, 2019
1222 stinkhorns unbranched stinkhorns 1223

Stinkhorns Species of Mutinus are small

The stinkhorns and their allies
(Phallales) is a monophyletic group
of fungi that form spores internally
in a bulb-shaped ‘witch’s egg’. At
maturity, the egg opens to reveal
a slimy, dark spore mass. At the
same time the fruitbodies emit a
nauseating smell of carrion or faeces
that attracts flies and other insects,
which then disperse the spores. The
Phallus
2 species
page 1224
spore mass also contains sugars as a
reward for the insects. All species live
as decomposers and are traditionally
included in the gasteroid form group.
The insect-aided dispersal limits
the potential distance that spores
can travel compared to normal wind-
disseminated spores. A consequence,
it has been possible to follow the
relatively slow spread of some of the
Mutinus
3 species
page 1223
introduced species in this order, e.g. stinkhorns with a not very clearly
Clathrus archeri. delimited fertile area on top of the
stem. Has red and orange colours.
Other similar fungi:
– some basidiomycote truffles form Mutinus caninus is a very slender
structures similar to witch’s eggs, but stinkhorn with orange colours under
these stay underground (page 1256). an olive-black spore mass, as well
as part way down the stem. The
Further reading: 50, 92, 156, 241. white witch’s egg is elongated and
has orange colours when sectioned.
It is not as foul-smelling as Phallus
impudicus ▷▷. The smooth, dark
Micro-drawings:
spores.
spores measure 4.5–6.5 × 1.8–3 µm.
Occurs on wood chips, on and
Approximate species around old stumps, and in similar
number applies to
temperate Europe. places, in parks, deciduous forests
and with conifers.
Mutinus ravenelii ▽ has red
colours under the spore mass and
normally also down the stem; it
mainly occurs in gardens. Mutinus
elegans ⋉ is very pointed, has vivid
orange colours on the stem and
the spore mass is even less well
delimited.
Widespread and common; mainly
June–January.
Clathrus
2 species
page 1226

large;
unbranched;
white to small;
purplish unbranched;
red to orange 5 mm

Mutinus ravenelii is a slender

large; branched stinkhorn that tapers towards the
from the base;
top, does not have a well-delimited
red to orange
‘head’, and has red colours under the
olive-black spore mass; it also usually
has red tinges down the stem. The
large, branched witch’s egg is slender and white,
stinkhorns from the top of and has red colours when sectioned.
form spores inside a stem; red
a slimy spore mass, Not as foul-smelling as Phallus
which, at maturity, Aseroë impudicus ▷▷. The dark, sooty
emits a smell that 1 species spores measure 5–7 × 1.8–2.5 µm.
attracts insects rather large; page 1227
Sometimes occurs in large troops,
branched from
the top of a typically on compost and disturbed
other stem; pale soil in gardens and parks.
similar fungi orange
Mutinus caninus △ is ± orange
under the spore mass.
Rare and meteoric in its
Lysurus occurrence – probably introduced 10 mm
1 species from North America; June–October. Jacques Landry
page 1227 © Thomas Læssøe & Jens H. Petersen
basidiomycote truffles Fungi of Temperate Europe,
page 1256 Princeton University Press, 2019
1228 puffballs and the like puffballs and the like 1229

Puffballs and similar Micro-drawings:

Queletia
gasteroid fungi 1 species spores first,
page 1243 then capillitium
This form group includes (ca.) and basidia
basidiomycotes where the spores (bas.).
are produced inside the fruitbody Phleogena Tulostoma
Mycenastrum 1 species 19+ species Approximate
and form a powder at maturity. 1 species page 1246 page 1244 species number
The species may either be ball- page 1243 applies to
temperate
shaped, with or without a stem, or Europe.
have a star-like appearance when Langermannia
mature. The latter may open and 1 species
page 1242
close depending on moisture – they Lycoperdon & prominent stem Battarraea
are hygroscopic. The surface ‘skin’ Apioperdon part and scaly small, 1 species
large with thick, 1–3 mm wide; Chlorophyllum
25+ species surface; spore page 1246
is termed a peridium. Some species page 1237
leathery, splitting
mass brownish- on wood; agaricoides
without stem; peridium; in dry, 1 species
have a sterile elongation of the stem nitrophilous places
orange basidia septate
large, more than page 1247
within the spore-containing ball – a Calvatia 8 cm wide; the small ‘puffball’
columella. Others may have a short, 2 species whole surface on a long,
± scaly stem brown, dusty cap
wide stem, which can be massive or page 1241 breaks down
on a long stem;
have spongy flesh. mainly
when young has
universal veil
The spores are hydrophobic with spongy stem;
and dispersed by the splash from opens with a pore bas.
or rots away from
raindrops. Almost all species have the top
dark, thick-walled spores – probably cap with dusty,
ca. ca. brown gill
an adaptation to prolonged exposure with massive stem; remnants on
to sun and wind. The spores may spore mass brown a short stem;
or violet; rots away ca. reddens Montagnea
be mixed with elastic, branched,
from the top ca. 1 species
thick-walled hyphae – a capillitium. page 1247
A spore mass with all elements it is cap with dusty, black
ca. gill remnants on a long
termed a gleba. without sterile stem,
stem; when young has
or stem small and puffballs
Traditionally, all the species massive; with pore ca. and the like universal veil
included here were placed in the Bovista in the top form fruitbodies
11+ species with an inner, dusty
artificial class Gasteromycetes, ca.
spore mass
page 1235
along with the bird´s nest fungi, opens outer peridium
stinkhorns and basidiomycote with a splits into rays;
pore on the with columella;
truffles. Phylogenetically the hygroscopic
underside and
majority belong within the agarics rotates spore or not
(Agaricales). Battarraea, Montagnea
and Chlorophyllum agaricoides are other
similar fungi
intermediate types that in some Disciseda
1–5 mm in diameter;
3 species outer peridium
respects look similar to closely related on marine
splits into rays;
page 1234
agarics, but the gills are transformed driftwood
with columellae Geastrum
to a dusty spore mass at maturity. and many poroid 30+ species
Nia openings page 1248
Such forms are also called secotioids 1 species thick peridium;
after the genus Secotium, where page 1234 spore mass black
outer peridium
they were formerly placed. and marbled
splitting into Myriostoma
spore mass ± marbled rays; 1 species
Other similar fungi: in internal without columella; page 1254
chambers hygroscopic
– truffles have tuber-like fruitbodies
that develop below ground (page
1256).
– cleistothecial fungi are asco­
mycotes with tiny, 0.2–3 mm wide,
closed fruitbodies (page 1500).

Further reading: 92, 134, 135, 136,

137, 138, 142, 156, 162, 241, 320, Astraeus
365. 2 species
page 1254
Scleroderma Pisolithus
7+ species 3+ species © Thomas Læssøe & Jens H. Petersen
page 1231 page 1230 Fungi of Temperate Europe,
truffles (page 1256) cleistothecial fungi (page 1500) Princeton University Press, 2019
1256 basidiomycote, zygomycote and glomeromycote truffles basidiomycote, zygomycote and glomeromycote truffles 1257
Melanogaster
7 species
page 1264
Truffles Micro-drawings:
spores.
Truffles constitute a form group where
Octaviania
the tuber-like fruitbodies are formed Rhizopogon 2+ species Chamonixia Approximate species
± underground (hypogaeus) and 12 species page 1263 1 species number applies to
page 1262 page 1263 temperate Europe.
spore dispersal is passive. This strategy
has arisen multiple times during the
evolution of the fungal kingdom, both
amongst the Ascomycota, Basidiomycota
and Zygomycota and within the Hymenogaster
Glomales (A-mycorrhiza-forming fungi), 30+ species
where truffle-like asexual structures can page 1265
be found. The fungal wheels shown
here have the basidiomycote and other
non-ascomycote truffles on the first
page spread and the ascomycotes (the Sclerogaster
3 species Macowanites
‘true truffles’) on the second. page 1263 1 species
Almost all truffles are ectomycor- page 1266
rhizal. They are dispersed by animals,
and various mammals are attracted by
the pungent odours. Boar, deer, rodents,
etc. dig the fruitbodies out of the soil
and eat them. Some rodents hoard
truffles. Spores from some of the species
tolerate passage through the gut. The
odours vary from species to species, and Stephanospora
1 species
some may resemble pheromones. It is page 1266
probably compounds of the latter nature
that make true truffles an exclusive
ingredient in fine cuisine.
The spores are formed internally and
the mechanism for active release has
been lost. Within the basidiomycote Gautieria
truffles this means that the sterigmata 4 species
page 1261 basidiomycote and
on the basidia and the apiculus on the zygomycote truffles
spores are absent or have changed, are typically rubber-
and that the spores are typically more like, foam-like or
slimy inside – rarely ×¼
symmetrical than in basidiomycotes with folded/veined as most
active dispersal. Within the ascomycotes ascomycote truffles
the cylindrical ascus has typically ×¼
evolved into a ± balloon-like shape
without special structures, but in some Endogone
species the cylindrical shape has been 2 species
other
page 1267
maintained. The ascospores tend to be similar fungi ascomycote truffles
extremely large (20–50 µm). have asci and are
mostly folded/veined
Hysterangium internally
10+ species
page 1261 with
basidia

Phallogaster
Glomus
1 species
3+ truffle-like
page 1260
species
page 1267
False truffle basidia: left, a 2-spored

⬅
basidium from Hymenogaster; right, a Geastrum
6-spored basidium without sterigmata from
Hysterangium. 30+ species, © Thomas Læssøe & Jens H. Petersen
one basidiomycote truffle puffballs and the next page spread Fungi of Temperate Europe,
page 1260 like page 1228 Princeton University Press, 2019
1258 ascomycote truffles ascomycote truffles 1259

How to find truffles (truffling) Hydnotrya Pachyphloeus Micro-drawings:

5 species spores.
Some truffles may form fruitbodies at page 1270
3 species
Balsamia page 1271
soil level with the top clearly visible; 2 species Approximate species
this is often the case with, for example, page 1271 number applies to
temperate Europe.
species of Rhizopogon and Choiromyces.
However, as the majority form completely
immersed fruitbodies in litter or rotten
wood, or in soil down to 10–15 cm,
sometimes deeper, they have to be
Hydnobolites
dug out (e.g. with a small hand rake). 1 species
Successful truffling requires a certain ‘feel’ page 1271
for the ecology of the species. One simple Hydnocystis
1 species
prerequisite is the presence of suitable page 1269
mycorrhizal partners, mainly trees and
shrubs, e.g. Corylus, Tilia, Fagus, Quercus
or ectomycorrhizal conifers. Relatively Choiromyces
light, mull soils can often be rewarding 1 species
page 1272
to rake through, mainly in places with a
warm microclimate. Rake the surface and
keep an eye out for any tuberous object
(alas, many will be fungal primordia,
pebbles, old nuts, etc.). Some truffles
form potato-sized fruitbodies, but many
are small, down to the size of a pea. If
you think you have found a truffle, cut
it through with a sharp blade and check
the internal structure with a hand lens; if Paurocotylis
veined or spongy you are in luck. 1 species
A shortcut can be taken by looking page 1269
out for animal scrapes, e.g. from deer or ascomycote truffles
squirrels – or, even better, train a dog! are typically folded/
veined internally,
except the more
Other similar fungi: uniform Elaphomyces
– witch's eggs may look similar but do not that eventually
grow underground (page 1222). become powdery
– puffballs and the like typically become
dusty at maturity and are normally not
other
immersed in soil (page 1228).
truffles
– cleistothecial fungi produce tiny are typically other
fruitbodies with tiny asci (page 1500). rubber-like, similar
foam-like or fungi
Geopora
9+ species but slimy inside
Further reading: 7, 98, 163, 197, 238, 239, only 3 truffles
240, 303, 317, 318. page 1268 Tuber
with
basidia 14 species
with asci;
fruitbodies page 1273
without tiny
basidia
and asci

Genea
4+ species
page 1268 photo: Flemming Rune

Elaphomyces

⬅
Types of truffle asci: left, a balloon-shaped 10+ species
tytpe from Tuber aestivum; right, a cylindrical
ascus from Hydnocystis. page 1276
© Thomas Læssøe & Jens H. Petersen
basidiomycote truffles Endogone/Glomus cleistothecial fungi Fungi of Temperate Europe,
previous page spread previous page spread page 1500 Princeton University Press, 2019
1278 operculate cup fungi and morels operculate cup fungi and morels 1279

© Thomas Læssøe & Jens H. Petersen

Operculate cup fungi Fungi of Temperate Europe,
Princeton University Press, 2019
The operculate cup fungi are charac­ operculate cup fungi with
terized by a small lid (an operculum) at operculate cup fungi with yellow, orange, red to
white, greyish, violet, brown pinkish hymenium
the top of the asci, and all operculate page 1328
to black hymenium
species are classified in the order page 1290
Pezizales. The open fruitbodies
(apothecia, see page 30) have the
hymenium on the surface/inner side of
the cap or cup. They are often rather
large and may be ± cup-shaped, either
have or lack a stem or, as in the morels,
be more folded or pitted.
The hymenium is a mixture of asci
and paraphyses. The spores are mostly
large and may be ornamented with
e.g. a raised reticulum or with warts
or spines. The ends of the spores may
have appendages.
Some species are ectomycorrhizal,
while others are decomposers of dead
organic material, e.g. dung, wood or
debris. A third group is parasitic, mainly
on bryophytes and hepatics.
The dung-inhabiting (coprophilous)
species often produce very small
fruitbodies compared to the ecto­
mycor­rhizal species. The smallest
fruitbodies are found in the genera operculate
Ascozonus, Coprotus and Thelebolus cup fungi
have open fruit­
and can be less than 0.5 mm in bodies (apothecia)
diameter. The species in these three operculate cup fungi with and asci with a lid
a folded, wrinkled or (an operculum)
genera also have aberrant asci, where pitted cap on a stem
the top cracks instead of opening with page 1280
a lid, and are classified in an order of other
their own, the Thelebolales. similar fungi

Other similar fungi:

– inoperculate cup fungi have asci
with an internal plug-like structure
or no structure at all. They are mostly
smaller, typically 0.5–5 mm in diameter,
and mainly occur on dead herbaceous
stems, fallen leaves or dead wood.
Club-shaped inoperculate cup fungi
(page 1348) may resemble small morels
(page 1280) and often occur on soil.

Amyloid asci with asci without

lid in a Peziza operculum,
before and after but often
spore discharge. with an inoperculate
inner apical cup fungi
apparatus page 1348
1280 morels and the like Morchella 1281

Morels and the like Morchella is a genus of large,

All species belong to the operculate hymenophore is often highly folded, Other similar fungi: operculate cup fungi with a pitted
cup fungi, i.e. Ascomycota, and have resulting in the hymenium having a – some species in the group and hollow head on a hollow stem.
an exposed hymenium and asci that large surface area. ‘earthtongues and the like’ may look The spores are large and smooth.
open with a lid (an operculum). All Morchella may form mycorrhiza- similar to tiny morels. These, however, The genus probably includes both
have inamyloid asci. They produce like associations with plants, e.g. with have inoperculate asci (page 1350). mycorrhizals and saprotrophs.
the largest fruitbodies (apothecia) Rosaceae, Ulmus or herbs. Gyromitra
within the ascomycotes – species of is saprotrophic, while Helvella is See also the main wheel, page 1278. •• Morchella esculenta is a large,
Morchella ▷ and Gyromitra ▷▷ can ectomycorrhizal. at maturity ± buff Morchella with a
reach more than 20 cm in height. The Further reading: 101, 180, 266, 301 328. hymenophore of large, rather regular
honeycomb-like cells. The hollow top
may be conical or almost globose, and
Morchella the stem length and thickness is very
20+ species variable. It has a pleasant, spicy odour.
page 1281 The smooth spores measure 18–22
(–27) × 12.5–14 µm, and they have
Micro-drawings: numerous external drops at the poles
spores and asci.
but none internally. Occurs on rich, ±
Approximate calcareous soils – often with Ulmus,
species number but also found with e.g. Fraxinus and
applies to
temperate Populus; mycorrhizal.
Verpa Europe. Morchella esculenta is part of
2 species difficult species complex. Morchella
page 1284
vulgaris ▽ is darker with more irregular
hymenophoral cells. Morchella
americana ⋉, found in central Europe,
was probably recently introduced from
North America. Morchella steppicola ⋉
occurs in the steppes of Eastern
Gyromitra
with one large 12+ species Europe.
cavity that unites page 1285 Widespread, occasional; April–June.
stem and fertile cavity only formed 15 mm
part in the stem that
is attached to the
cap in a narrow •• Morchella vulgaris is a relatively
point large to large, rather dark Morchella
with a hymenophore that has a rather
irregular honeycomb-like pattern of
cells with thick walls. The smooth
spores measure 18–22(–27) × 
both stem and cap
folded and veined 12.5–14 µm and have numerous
and not completely external drops at the poles, but none
hollow; hymenium inside. Occurs on ± calcareous soil
yellow-brown to
ascus dark red-brown in deciduous forests, parks, gardens
and dunes, may occur with a range of
partner trees, e.g. Crataegus, Malus,
morels Fraxinus and Sorbus; mycorrhizal.
and the like
form ± folded fertile
Morchella americana ⋉, which was
heads on a stem. apparently introduced to Europe from
The asci open by a North America, is very similar and
lid (operculum) stem grooved
or smooth; probably requires sequencing in order
hymenium to confirm identification. Morchella
other ± veined/folded,
white to dark esculenta △ is typically paler and the
similar fungi
grey-brown cells in the hymenophore are more
regular.
Widespread, occasional; mainly
April–June. 10 mm
small; asci Helvella
inoperculate 35+ species
page 1288 © Thomas Læssøe & Jens H. Petersen
earthtongues and others Fungi of Temperate Europe,
page 1350 Princeton University Press, 2019
1290 operculate cup fungi with white, grey, brown, violet to black colours operculate cup fungi with white, grey, brown, violet to black colours 1291
Sphaerosporella
Tricharina, Trichophaea Humaria
Geopora 1 species Tarzetta &
Operculate cup fungi with 8 species
& Trichophaeopsis
page 1307 2 species
Sepultariella Micro-drawings:
20+ species page 1308
dull colours page 1304
page 1305 6+ species spores first,
Otidea page 1310 followed by
Apothecial ascomycotes with disc- 22 species Geopyxis
paraphyses (p), asci
to cup-shaped apothecia, with or page 1302 4 species
(asci) and hairs (h).
page 1311
without stems, and with whitish,
Approximate species
greyish, brownish, violet, or black
count applies to
hymenia. The asci have a lid (an Leucoscypha temperate Europe.
2 species
operculum) or, in Thelebolus and small; rather large;
Wynnella page 1309
Ascozonus, a cracking top. small; with pointed with pointed,
1 species with pointed brown hairs;
Helvella brown hairs; margin pale,
page 1301 brown hairs; autumn
35+ species autumn dentate; on margin pale Arpinia
partly under spring to 2 species
Other similar fungi: page 1298 rich soil and felty;
ground; with autumn page 1309
– operculate cup fungi with on burnt
brown hairs;
ground
yellow, orange, red to pink ± split down spring to small; white;
one side; autumn with pointed Pseudombrophila
colours are treated in a separate ± ear-shaped; hairs 20+ species
wheel (page 1328). soft-fleshed page 1311
narrowly
ear-shaped; h fasciculate
See also the general tough-fleshed h
wheel, page 1278.
margin smooth
Desmazierella or felty; mostly Smardaea
typically with stem;
Further reading: 34, 49, 2 species p on dung 3+ species
outer side ± felty
51, 52, 53, 73, 101, 108, page 1298 page 1312
to hairy; spring to
109, 118, 233, 301, autumn p h
± purple
310, 329, 349, 358. with brown
pointed hairs in Marcelleina
hymenium and 5+ species
Pseudoplectania margin; spring h
small, bluish to page 1312
4 species greyish-violet
page 1296 h+p
outer side with Plicaria
± black felt; operculate 4 species
winter-spring to cup fungi asci page 1322
h asci i-
autumn with whitish, amyloid
asci i+
Plectania p greyish, violet,
outer side with brownish or
1 species
rust-brown felt; ± black colours asci amyloid;
page 1297
spring outside with Plicariella
h conical warts 2 species

asci
page 1322

asci
large and

i+
i-
gelatinous; asci amyloid;
winter-spring on wood in
Sarcosoma I+ water Adelphella
1 species 1 species
page 1295 large; with stellate page 1322
asci amyloid;
opening; grey-
on damp wood
brown hymenium;
winter-spring
large; with stellate
asci
opening; ± greyish-
amyloid Pachyella
Urnula violet hymenium;
spring 3+ species
2 species page 1323
page 1294 large; with asci on dung;
hyphal cords; cracking asci amyloid;
± all year spores in ‘packets’
very large and
veined; nitrous; mostly on
spring dung;
large; asci amyloid
on wood; mostly on
spring on dung; dung;
Sarcosphaera on dung;
asci large asci amyloid
1 species asci large
page 1294 Saccobolus
15+ species
page 1325
Peziza s.l.
Ascobolus 50+ species
Rhizina page 1313
1 species 30+ species
page 1293 page 1324
Thecotheus
Disciotis 7+ species © Thomas Læssøe & Jens H. Petersen
Gyromitra Thelebolus Ascozonus Fungi of Temperate Europe,
1 species, page 1293 12+ species, page 1292 page 1326
2+ species, page 1367 3+ species, page 1326 Princeton University Press, 2019
1328 operculate cup fungi with yellow, orange, red to pink colours operculate cup fungi with yellow, orange, red to pink colours 1329
Lamprospora
Octospora
40+ species
Yellow, orange, red to pink 45+ species Micro-drawings:
Ramsbottomia page 1338
page 1339 Neottiella spores first,
operculate cup fungi 4 species 5 species followed by
page 1337 page 1337 paraphyses (p), asci
Fungi with cup- to goblet-shaped or Pyronema Anthracobia and hairs (h).
2 species 5 species
flattened apothecia, yellow, orange, page 1336 page 1340 Approximate species
red or pinkish hymenium, and asci with Miladina number applies to
a lid (an operculum) or, in the genus 1 species temperate Europe.
page 1336
Coprotus, with a slit-like opening Melastiza
mechanism. All genera, apart from Pulvinula 5+ species
5+ species very small; very small; page 1341
Iodophanus and Peziza, have inamyloid page 1336 small; on moss parasites; moss parasites;
asci. mineral soil; autumn–winter autumn–winter small; with stem;
autumn moss parasites;
small; in troops on
autumn–winter
burnt ground; small; on burnt
Other similar fungi: spring–autumn ground; mostly
– operculate cup fungi with with appressed
whitish, greyish, brownish, small; on deciduous dark hairs;
autumn Spooneromyces
violet or black hymenia are wood in streams; medium; with small, 3 species
autumn p
found in the wheel on page dark hairs; mostly page 1340
on rich, disturbed
1290. soil; summer–
small; on mineral p autumn
See also the general or burnt soil;
p p Scutellinia
summer–autumn
wheel, page 1278. Aleuria p rather small; on 25+ species
5+ species p h p needle debris; with page 1342
page 1335 dark, pointed hairs;
Further reading: 33, 34, 39, small to large; autumn
49, 73, 101, 119, 198, mostly on rich, p
disturbed soil;
199, 200, 233, 292, 359, autumn p
rather small; with
360, 361, 363, 364. h p
dark, pointed,
Sowerbyella projecting hairs;
medium to large; p h
6+ species spring–autumn
yellow with h
page 1334 p Parascutellinia
distinct stem;
autumn p operculate h 3 species
cup fungi page 1343
p small; with
with yellow,
orange, red to h pointed hairs;
pink colours autumn
medium to large; p h
± ear-shaped; h
autumn p p
rather large with
brown, pointed,
Otidea p projecting hairs;
medium; with
22 species autumn
greenish-/bluish-
page 1332 staining outer side; ascus p
I+
spring small; with Humaria
pointed hairs; (see page 1308)
rather small, yellow- p p ascus mostly on dung;
orange; on Abies I+ all year-round
and Cupressaceae; asci
winter-spring cracking h
Caloscypha small; in clusters
1 species where deer urinate;
page 1332 small with long, autumn–spring
branched stem p
(clustered);
winter-spring medium; asci
p amyloid; mostly
autumn
large with short very small; pale Cheilymenia
stem; winter-spring pinkish; on dung and 10+ species
very small; with page 1344
Pithya very small; on dung; pale hairs; on dung; plant litter; all year
2 species Microstoma all year all year Byssonectria
page 1331 1 species 2 species
page 1331 page 1345

Peziza
page 1318

Sarcoscypha Iodophanus
2+ species © Thomas Læssøe & Jens H. Petersen
3 species Coprotus Lasiobolus Fungi of Temperate Europe,
page 1330 page 1347
10+ species, page 1347 7 species, page 1346 Princeton University Press, 2019
1348 inoperculate cup fungi inoperculate cup fungi 1349

Inoperculate cup fungi high or long-stemmed, non-gelatinous,

low, smooth to felty
inoperculate cup fungi
smooth to felty inoperculate cup fungi
The inoperculate cup fungi is a form (high = more than 2 mm above the
(low = less than 2 mm
above the substrate, page 1426)
group of ascomycotes with open fruit- substrate, page 1404)
bodies (apothecia) where spore release
is mostly controlled by an inner, cork-
like structure at the ascus tip. The name
refers to the lack of an operculum (a lid
on the top).
The apothecia are mostly small (less
than 2 mm wide) and cushion- or disc-
shaped to cupulate, and numerous
species occur on wood or herbaceous
stems. A few genera produce larger or
± club-shaped apothecia. Species in the
Sclerotiniaceae produce apothecia from
overwintered sclerotia; many of these
are plant parasites. erumpent inoperculate cup fungi
(opening by a lid, with teeth or slit-like,
The hymenium is in most cases page 1468)
made of asci and paraphyses. The
spores are typically hyaline, small and
smooth.
This form group is very large with smooth,
cupulate,
nearly 300 genera and perhaps around smooth, discoid,
mostly long-
sessile to short-
1,000 species in temperate Europe. The stemmed;
stemmed,
not or hardly
generic delimitation will undoubtedly gelatinous or
with gelatinous
undergo many changes in the coming downy to hairy not
inoperculate cup fungi projecting
years, and many more species will be (page 1372) hairs; not
described. Here we have included 139 gelatinous
inoperculate slit-like or with lid
genera with 256 species. cup fungi or teeth
large, gelatinous
inoperculate cup fungi (open fruitbodies
and asci
Other similar fungi: (page 1364) mainly large,
without lid)
– the operculate cup fungi have asci gelatinous

that open with a lid (an operculum).

The fruitbodies are mostly larger and other
many species occur on soil or wood. similar fungi

A group of species on dung have very

small apothecia (page 1278).
– the apothecial lichens may have club- to without asci or
tongue- basidia
fruitbodies that can look very similar shaped
to species in this form goup, but
most have obvious thalli with algae or
cyanobacteria (page 1484).
with with
– cyphelloids are small basidiomycotes asci basidia
with mainly downturned, cupulate or
ear-shaped fruitbodies (page 1076).

asci with a: simple

top; b: cylindrical
apical apparatus
and simple base;
c: cylindrical apical
apparatus and earthtongues and the like asexual fungi
crozier at the (page 1350) (page 1626)
base; d: thick-
walled top. a b c d © Thomas Læssøe & Jens H. Petersen
operculate cup fungi apothecial lichens cyphelloids Fungi of Temperate Europe,
(pages 1278) (page 1484) (page 1076) Princeton University Press, 2019
1350 tongue- to club-shaped inoperculate cup fungi tongue- to club-shaped inoperculate cup fungi 1351

Spathularia
3 species
Earthtongues and the like page 1358 Micro-drawings:
Leotia spores first, followed
This form group includes the species
2 species by paraphyses (p),
of inoperculate cup fungi that have page 1360 Cudonia ascus tops (at),
Microglossum 2 species amyloid (I+),
clavate, tongue-shaped, spatulate or 10+ species page 1359 inamyloid (I-), and
morel-like fruitbodies. page 1356 Mitrula
hairs (h).
Almost all species in the group 1 species
page 1361
have asci with an internal apparatus Approximate species
Thuemenidium number applies to
near the tip that is blown out and 1 species temperate Europe.
yellowish fertile part
inverted during spore ejection. This page 1355 to olive, with helmet-like
flattened; in forests fertile part; on litter
apparatus often stains blue in iodine. rubbery;
in forests in forests top yellow, somewhat
The genus Neo­lecta (page 1363) gelatinous; on leaves Bryoglossum
reddish, olive- 1 species
deviates by having asci that split brown to green; in clean fresh water;
spring–summer page 1361
open (no apparatus) and by the lack grasslands and at (I-)
forests
of paraphyses among the asci.
Many species are decomposers pale yellow with
but earthtongues, for example, at (I+) at (I-) ± free lower margin;
Sarcoleotia dark brown; spores on mosses; ± boreal
possibly form biotrophic associations large, hyaline;
1 species
with plants in the same way as the grasslands p
page 1355
Cudoniella
hygrocyboid fungi (Hygrocybe and (page 1370)
others) are thought to do (page 18). at (I-) greyish white to
p
at (I+/-) cream, gelatinous,
stem brown, scaly; ± nail-shaped;
Other similar fungi: p on wood, stems,
spores large, p
at (I+) leaves, etc.
– clavarioids may have very similar hyaline; on sand in p
heathlands p
fruitbodies but are basidiomycotes,
i.e. the spores form on basidia (page p
Heyderia
1092).
at (I-) 2 species
– jelly fungi and Calocera may be p tiny, with buff cap; page 1362
clavate in shape but are tougher and black; spores large, on needles
Sabuloglossum translucent; with earthtongues p
have septate or tuning fork-shaped Empetrum and the like
1 species at (I+)
basidia (page 1154 & 1162). page 1355 (inoperculate cup
photo Thomas Kehlet fungi with clavate p
– stromatic pyrenomycetous fungi at (I+) p fruitbodies)
such as, for example, Cordyceps, Episclerotium
x½
2 species
Trichoderma and Xylaria, have the stem slimy; spores large,
p page 1362
0–7-septate, hyaline h
flesh filled with perithecia (see page to brown; paraphyses
30) and the openings can be seen ± straight; grasslands h
small, fertile

x½
and forests part orange-
as small dots on the surface (page p other at (I+)
brown;
similar fungi
1510). p on fungal
– long-stemmed inoperculate cup Glutinoglossum sclerotia
x½ p
5+ species x½ at (I-)
fungi have a rather well-defined disc
page 1354
with hymenium (page 1404).
spores large, with veined head;
p from sclerotia
multiseptate,
See also the general wheel, brown; with setose formed on herbs
page 1348. black hairs; Scleromitrula
grasslands and with with asci and 5 species
forests at (I+) basidia dotted with tough-gelatinous; page 1363
Further reading: 14, 21, 49, 101, perithecia on twigs in clean
stem hairy; spores
166, 167, 168, 277. large, multiseptate, p fresh water; spores
Trichoglossum hyaline; thread-like
3+ species e.g. with Littorella in spores large, yellow and rather
page 1354 lakes 0–multiseptate, almost fragile; without
hyaline to brown; paraphyses; asci
grasslands, forests, etc. split open;
on soil, boreal

Hemileucoglossum Vibrissea
2+ species 1 club-shaped species
page 1353 page 1360

clavarioids
Neolecta
(page 1092)
Geoglossum 1 species
10+ species jelly fungi page 1363 © Thomas Læssøe & Jens H. Petersen
page 1352 and Calocera stromatic pyrenomycetous Fungi of Temperate Europe,
(pages 1154 & 1162 fungi (page 1510) Princeton University Press, 2019
1364 large, gelatinous inoperculate cup fungi large, gelatinous inoperculate cup fungi 1365

Ionomidotus
(page 1407)
Large, gelatinous Bulgariella
inoperculate cup fungi Holwaya 1 species
(page 1412) page 1370
A form group with often quite large,
Cudoniella
discoid, turbinate or brain-like, highly 4+ species
gelatinous apothecia. The spore page 1370
release is controlled by an internal,
Bulgaria
cork-like structure in the ascus 1 species
top. The species in this group are page 1371
predominantly wood-decayers.
medium to large,
greenish-yellow to black;
Other similar fungi: medium-sized, stains dark in KOH medium-sized,
– grades into the group of small, discoid, black; with (ionomidotic reaction); on black; only slightly
conspicuous asexual deciduous wood gelatinous; on
gelatinous, inoperculate cup fungi. Vibrissea
state; on wood, deciduous wood
7 species
It can be difficult to determine Ascocoryne mainly Tilia (page 1455)
whether or not a tiny cup fungus 7+ species medium-sized, large, purple
page 1368 to black; highly
has gelatinous matter in the tissues, turbinate to discoid,
whitish to pale gelatinous (like jellied
and it is important to check the candy); on deciduous
brown;
wheels for both the short- and long- on wood or stems wood
stemmed inoperculate cup fungi
(pages 1404 & 1426).
– gelatinous, tongue-shaped species small and only
are included on page 1350. moderately
medium to large,
gelatinous; with
– gelatinous, operculate cup fungi turbinate to
thread-like spores; on
discoid, brown to
are included on page 1290. violet; often with
deciduous wood in
fresh water
– jelly fungi have basidia, see page conspicuous asexual
1162. However, they can superficially state; on wood

resemble species of Ascotremella and large, gelatinous

Ombrophila and microscopy may be inoperculate
cup fungi small and at
required. p most moderately
with brain-shaped,
large and brain-like; turbinate to discoid gelatinous; with
See also the general wheel, page violet-brown; spores apothecia small spores and
with indistinct stripes swollen paraphyses;
1348. in cotton blue; on wood or stems
on deciduous wood
other
Further reading: 21, 49. similar fungi

Orbilia
(page 1464)
medium-sized,
Ascotremella discoid to
1 species turbinate, pale with asci with basidia
page 1366 violet-brown;
on cones, wood or
stems

asci asci
inoperculate operculate

Micro-drawings:
spores outermost,
followed by
paraphyses (p). Ombrophila
12+ species
Approximate species page 1366
count applies to
temperate Europe. © Thomas Læssøe & Jens H. Petersen
short- and long-stemmed inoperculate cup fungi earthtongues and the like operculate cup fungi jelly fungi Fungi of Temperate Europe,
(pages1404 & 1426) (page 1350) (page 1290) (page 1162) Princeton University Press, 2019
1372 downy to hairy inoperculate cup fungi downy to hairy inoperculate cup fungi 1373

Brunnipila
11 species , page 1380
Downy to hairy inoperculate Micro-drawings:
Cejpia
cup fungi Venturiocistella
(page 1435) spores first, followed
4 species by hairs (h).
Lachnellula Dennisiodiscus
This form group of small inoperculate 5 species with brown
page 1379
(page 1451)
cup fungi has the outer side, and hairs Approximate species
page 1378 number applies to
especially the margin, covered by temperate Europe.
projecting hairs. They are mainly
decomposers and occur predominantly
on herbaceous stems, fallen leaves,
cones and wood.
The group contains many genera, Neodasyscypha sessile to short-
1 species ± sessile; stemmed; sessile with downy grey
and the appearance of the hairs dark-haired
page 1378 pale brown; hymenium and dark
(the shape, size, colour, crystal- with two hair on herbs and leaves basal hairs; on Poaceae,
covering, guttulation, etc.) is a very types; on and Cyperaceae.
wood, leaves sessile with greenish- Perrotia
important distinguishing character. ± sessile to short- and herbs grey hymenium and dark 2 species
stemmed; with brown orange hairs; on Poaceae page 1382
Some of the more distinctive species and Carex
outer side and yellow
can be recognized in the field, but hymenium; on conifer
identification mostly requires the use wood ± sessile;
of a microscope. with orange-yellow
± sessile; to orange hairs; on
The majority of the species are Lasiobelonium with brownish- deciduous wood or
classified within the Hyaloscy­pha­ceae, 6+ species yellow to yellow h Poaceae
page 1377 colours; on
and many have long paraphyses with
deciduous wood
lanceolate or tapering tips that project
above the asci (see figure below). h ± sessile; with
h Albotricha
white or coral 6 species
Other similar fungi: h hairs; on herbs page 1382
± sessile; h or deciduous
– cyphelloid fungi (page 1076) have brown-haired; h wood
small, hanging fruitbodies. They are on deciduous wood h
basidiomycotes and can be recognized
h h
by their basidia and spore morphology ± hairy
– the spores having a small appendage inoperculate ± sessile on
(apiculus) where they were attached to cup fungi subiculum;
h – species with h
the sterigmata. They are often not as sessile; mostly with
brown-haired; dark hairs yellowish to
elegant as the hairy inoperculate cup on herbs orange hairs;
fungi and the hairs are more disorderly. mostly on wood
– some other inoperculate cup fungi other
have hairs but they are short and Trichopezizella h similar fungi species with Arachnopeziza
10 species

⬅
adpressed, making the apothecia look 5+ species pale hairs,
sessile; page 1383
page 1376 see next page spread
felty rather than hairy (a hand lens or mostly with
dissecting microscope is required) – see yellowish hairs;
on herbs and
the general wheel to the inoperculate wood
cup fungi, page 1348.
with with
Further reading: 21, 22, 23, 49, 78, 101, asci basidia
115, 116, 160, 253, 319.

Trichopeziza
6+ species
page 1392

A typical hymenium in the

genus Lachnum with asci
and lanceolate, projecting
paraphyses.
sessile to short-stemmed, high, sessile to long-stemmed, © Thomas Læssøe & Jens H. Petersen
smooth to felty smooth to felty cyphelloids Fungi of Temperate Europe,
(page 1426) (page 1404) (page 1076) Princeton University Press, 2019
1374 downy to hairy inoperculate cup fungi Calycina downy to hairy inoperculate cup fungi 1375
15+ species
page 1389 Hyphodiscus
Calycellina Psilachnum 12+ species
15+ species 10 species page 1390 Cistella
page 1387 page 1388 30+ species
page 1391

Hamatocantoscypha Cyathicula
8 species (page 1425)
page 1386

Trichopeziza
6+ species
± sessile; very short- ± sessile; very small page 1392
± sessile; haired; on wood,
very short-haired; and short-haired;
cones and herbs mostly on
mostly on ferns
± sessile; very and grasses other fungi ± sessile; very small
short-haired, hairs and short-haired;
± hooked; mostly on mostly on herbs
wood
Hyaloscypha h
± sessile; with short, teeth of thin-walled
15 species p
hooked hairs; (+VB) hyphae; mostly on
page 1385
mostly on wood herbs
h

± sessile; rather
short-haired; p
± greyish translucent; (+VB) sessile; ± cream; Amicodisca
h on herbs and wood 3 species
mostly on wood p
h page 1394
(+VB) h
h
Dematioscypha ± sessile amongst p
3 species black conidiophores; h
(+VB)
page 1384 greyish translucent; ± sessile; yellow-
mostly on wood h haired; on wood in
wet habitats
asex ± hairy
h h
barrel-shaped; inoperculate Proliferodiscus
with very short cup fungi 2 species
h
glassy hairs; – species with h page 1395
h ± sessile; felty;
on herbs and pale hairs
on wood,
wood h
drought-
Olla resistant
h
6 species h h
page 1384
barrel-shaped with h
projecting, glassy
Urceolella hairs; on herbs
11 species sessile to short-
page 1384 stemmed; hymenium
h yellow; on conifer
h
wood Lachnellula
species with 12 species with
on subiculum;
dark hairs, white hairs
on Quercus wood
± short-stemmed; page 1396

⬅
see previous page
spread hymenium yellow;
on deciduous wood,
on leaves, herbs 
Eriopezia subiculum; on etc.
1 species various substrates ± short- to long-
page 1383 stemmed; whitish to
yellowish; on wood
and leaves Capitotricha
± short-stemmed;
± short- to long-stemmed; hymenium white; on 4 species
hymenium white or coloured; wood, leaves, herbs  etc. page 1397
on wood, leaves, herbs  etc.
Micro-drawings:
spores first,
followed by hairs
(h), paraphyses (p)
refractive vacuolar
bodies (VB) and
asexual state (asex) Arachnopeziza Incrucipulum
10 species 4+ species
Approximate species page 1383 page 1398
number applies to
temperate Europe.
Dasyscyphella © Thomas Læssøe & Jens H. Petersen
Lachnum 10+ species, page 1399 Fungi of Temperate Europe,
35+ species, page 1399 Princeton University Press, 2019
1404 high or long-stemmed, smooth to felty inoperculate cup fungi Sclerotinia high or long-stemmed, smooth to felty inoperculate cup fungi 1405
13 species
Dumontinia page 1416 Monilinia
2 species 16 species
High or long-stemmed, page 1415 page 1416 Myriosclerotinia Micro-drawings:
smooth to felty inoperculate 9 species spores, followed on
page 1417
cup fungi Ciboria
the inside by asci (a),
conidia (con).
A form group of non-gelatinous to 15+ species
page 1414 Approximate species
slightly gelatinous, distinctly long- Lambertella Gloeotinia
number applies to
stemmed inoperculate cup fungi or, if 5 species 3 species
temperate Europe.
page 1413 page 1417
short-stemmed, growing to 2 mm or
more above the substrate. The outer Tatraea
(page 1444)
side is smooth to felty but does not
from sclerotia,
have projecting hyphal hairs. A few from mostly on grasses mostly from
sclerified sclerified fruits
species have teeth at the margin of the rhizomes
and herbs from sclerotia, on
Piceomphale on leaves,stems, Juncus, Luzula or
apothecia made up of aggregations of 1 species seeds, catkins, Cyperaceae
hyphae. Some species form apothecia page 1413 wood, acorns,
cupules, galls, etc. on seed from Cudoniella
from sclerotia. Juncus, Carex or 3+ species
Some species are decomposers, dotted from dark Poaceae page 1418
e.g. of wood or herbaceous stems, spores; mostly on pale to dark
while others – members of the sclerified leaves grey-brown;
and fruits large spored;
Sclerotiniaceae – parasitize plants.
on deciduous
Holwaya wood
1 species black;
Other similar fungi: on cones
page 1412 mostly rather
– downy to hairy inoperculate cup in spring
tough; Antinoa
fungi are covered by ± projecting hairs. mainly on 5+ species
Many species also have lanceolate, deciduous wood page 1418
rather tough;
± projecting paraphyses (page 1372). black; with
– small, short-stemmed to sessile, distinct asexual
state; on small with
smooth to felty inoperculate cup fungi tough stem;
deciduous
arise to less than 2 mm above the wood on needles
high, ± smooth, and cones
substrate (page 1426). non-gelatinous
fragile to tough; inoperculate Chloroscypha
See also the general wheel, with yellow- cup fungi greenish-yellow; 4 species
green colours; (long-stemmed or arise on needles of page 1419
page 1348. 2 mm or more above
on deciduous Thuja, Cupres­
wood the substrate) sus and Juniperus
Further reading: 19, 21, (Juniperus)
Chlorencoelia
49, 59, 72, 78, 95, 101, 1 species,
fragile to tough;
252, 267, 354, 362. page 1412
with green brown or
colours; mostly on other greenish-yellow;
deciduous wood similar mostly from
Chlorociboria and herbs fungi sclerified plant
4 species material
page 1409
tough; erumpent mostly whitish
a
in clusters; on con to yellowish; on
deciduous wood in wood or various
spring herbaceous
debris
whitish to
tough; dissolves buff; flesh
dark pigment in usually slightly Rutstroemia
KOH; on deciduous gelatinous; mostly 22+ species
wood on herbs page 1419
tough;
erumpent in
clusters on pink; on mosses
wood and Equisetum

Encoelia & Sclerencoelia Hymenoscyphus

10 species 40+ species
page 1408 page 1421
Cyathicula
Ionomidotus & Tympanis 25+ species
Diplocarpa 12+ species small, ± smooth ± hairy inoperculate page 1424
3 species, page 1406 page 1406 inoperculate cup cup fungi Roseodiscus © Thomas Læssøe & Jens H. Petersen
fungi, (page 1426) (page 1372) 4 species , page 1423 Fungi of Temperate Europe,
Princeton University Press, 2019
 Niptera Heterosphaeria
1426 low, ± sessile, smooth to felty inoperculate cup fungi 5+ species 5+ species Godronia low, ± sessile, smooth to felty inoperculate cup fungi 1427
page 1441 16 species
page 1441 page 1442
Pyrenopeziza
& Pirottea Tatraea
Low, smooth to felty 50+ species 1 species
inoperculate cup fungi page 1438 page 1444

This group includes sessile to very Dermea

Mollisia 6 species
short-stemmed, less than 2 mm
50+ species page 1444
high, inoperculate cup fungi with Cejpia page 1436
an outer side and margin that is 1 species
smooth to felty or rarely toothed, page 1435
but which never has projecting,
hyphal hairs. Large genera in the small, smooth, initially closed, Bulgariella
brown; on later cracking; ± spherical;
group include Mollisia and Orbilia. mostly (page 1370)
Poaceae, on herbaceous mostly on twigs
erumpent;
Coronellaria Cyperaceae stems from deciduous grey-brown;
spherical VBs;
3+ species and Juncaceae trees large-spored; Rhizodiscina
Other similar fungi: ± greyish; on on leaves and
on deciduous brownish-
page 1435 1 species
– long-stemmed or high (growing many substrates; herbaceous wood black; page 1445
paraphyses with stems
to 2 mm or more above the mostly on twigs
oblong VBs of deciduous
substrate), ± smooth inoperculate wood
downy; rather large;
cup fungi (page 1404).
on Molinia rather gelatinous;
– downy to hairy inoperculate on deciduous
cup fungi may be sessile to long- Polydesmia wood
Patellaria
stemmed but have ± projecting 3+ species downy; 1 species
page 1434 on Cyperaceae, black; mostly with page 1445
hairs. Many species also Juncaceae and p p small teeth at
have lanceolate, ± projecting Typha margin; on wood
paraphyses (page 1372). con
– erumpent inoperculate cup downy;
fungi are typically ± immersed on old stromatic black;
pyrenomycetous ± smooth;
in the substrate from where the p on dry wood
Pezoloma fungi h Sarea
apothecia emerge by splitting 4+ species 2 species
the substrate or by pushing a lid page 1431 page 1445
p with many
aside. The fruitbodies can often low, smooth to a
whitish with long teeth small, spherical
close again during dry spells and felty inoperculate
at margin; on wet litter spores; on
cup fungi a
reopen when wetted (page 1468). or Marchantia
(species with conifer resin
teeth whitish, greyish to
a
See also the general wheel, black hymenium)
very small, whitish with reddish Patinellaria
page 1348. with long teeth at p 1 species
subiculum;
margin; on leaves on wood page 1446
Further reading: 20, other
species with vividly
a similar fungi

⬅
21, 49, 78, 79, 153, white, short- coloured hymenia,
177, 210, 211, 212, stemmed; on see next page spread
Echinula
3 species herbaceous stems
213, 214, 216, 286,
page 1430
291, 334.
on soil, litter a
or liverworts or
Continued on next Calycina mosses
page spread . . . 8+ species
page 1433 asci flat-topped;
spores with
internal body

on living
Sphagnum
on living
Sphagnum resupinate; mostly
under Pinus bark
Micro-drawings:
spores first, followed
by paraphyses Orbilia
(p) possibly with 40+ species
refractive vacuolar page 1464
Hymenoscyphus &
bodies (VBs), conidia
Phaeohelotium
(con) and asci (a).
45+ species
page 1433
Approximate species
number applies to
temperate Europe.
long-stemmed or high, ± hairy erumpent © Thomas Læssøe & Jens H. Petersen
Discinella Trizodia
Ascocorticium ± smooth inoperculate inoperculate cup fungi inoperculate cup fungi Fungi of Temperate Europe,
5+ species, page 1432 1 species, page 1432
1 species, page 1430 cup fungi (page 1404) (page 1372) (page 1468) Princeton University Press, 2019
1428 small, ± sessile, smooth to felty inoperculate cup fungi Dennisiodiscus Catinella small, ± sessile, smooth to felty inoperculate cup fungi 1429
1 species 1 species
page 1451 page 1450 Godronia
Velutarina
16 species
1+ species Micropeziza
page 1442
Claussenomyces page 1451 2+ species
Chloroscypha page 1452
9+ species 5+ species
page 1449 page 1450 Pseudohelotium
3 species
page 1452

Mniaecia Bryoscyphus
3+ species 7 species
page 1448 page 1453
Chlorociboria
rather large; Vibrissea
(page 1409) rather large
7 species
rather large; with orange felty; on large, ± spherical;
rotten trunks of very small, page 1455
felty; on dry hairs; on mostly on twigs
yellow- stems and monocots deciduous trees of deciduous erumpent
green; on twigs trees on Poaceae, pale pinkish-buff;
greenish Cryptomeria Cyperaceae and mostly on conifer
to black; Juncaceae debris
small; on wood and
resin pale pinkish-buff;
blue-green;
on mosses and
on liverworts
p liverworts
Gorgoniceps
6 species (+VBs) spores thread-like;
small; turquoise;
page 1448 somewhat gelatinous; Podophacidium
on herbs or wood
on deciduous wood in 1 species
Graddonia p fresh water page 1455
1 species blue-green; (+VBs)
page 1447 on wood, cones, erumpent from Cenangium
herbs etc. brown flesh; on 3 species
needle debris page 1454

brownish-rose with
dark flesh;
on wood in erumpent; on
p
fresh water low, conifers
smooth to felty
inoperculate
Discinella brownish-rose; cup fungi
immersed in black
5+ species on soil (species with
asexual state; on
page 1447 or mosses vividly coloured Strossmayeria
deciduous wood
hymenium) p 1+ species
and Rubus
page 1456
brownish-rose; paraphyses with
erumpent with teeth; a oblong VBs; on wet
Diplonaevia on herbs deciduous wood,
15 species Phragmites,  etc.
page 1447

wine-red to black; yellowish; on Mollisia &

mostly on red species with deciduous wood, Phaeangella
subiculum; on wood whitish, greyish con bamboos and 50+ species
to black hymenia, reeds page 1457
a

⬅
see previous page p
spread
wine-red; apricot-yellow;
Patinellaria without subiculum; a
on wood, leaves
1 species on wood and soil
page 1446

± cushion-shaped;
with numerous mostly on stems of
erumpent; mostly small, spherical Urtica
asci with rounded asci flat-topped; spores; on Bisporella
clusterered;
tips; spores without spores with conifer resins & Calycina
on deciduous and
internal body internal body 12+ species
coniferous wood
page 1458
Hymenoscyphus
Micro-drawings: 40+ species
spores first, followed Patellariopsis page 1460
by paraphyses 2 species
(p) possibly with page 1446
refractive vacuolar
Calloria
bodies (VBs), conidia
3 species
(con) and asci (a).
page 1461
Approximate species Sarea
number applies to 2 species
temperate Europe. Hyalorbilia Pezicula page 1461 © Thomas Læssøe & Jens H. Petersen
5+ species, page 1467 Orbilia Fungi of Temperate Europe,
12 species, page 1462
40+ species, page 1464 Princeton University Press, 2019
1468 erumpent inoperculate cup fungi (with a lid, teeth or slit-like opening) erumpent inoperculate cup fungi (with a lid, teeth or slit-like opening) 1469
Ostropa Robergea
Stictis 1 species, page 1473
13 species 1 species, page 1473
Erumpent inoperculate cup page 1472 Trochila
fungi Phacidium
4 species
page 1474
A form group of inoperculate cup 15 species
page 1471 Ascodichaena
fungi that develop fruitbodies Coccomyces
Durella 1 species
(apothecia) within the substrate 9+ species
page 1475
page 1471 6 species
(typically in stems, leaves or wood). page 1474
At maturity the apothecia breaks
through the surface (erumpent). They
Karstenia
may be disc-shaped or elongated and 8+ species
with pale with transverse deep inside
are often surrounded by torn parts page 1470
slit; on deciduous
teeth; deciduous wood
of host tissue, which is sometimes on wood wood (seen in section) opening by a lid;
evident as distinct teeth or ‘lips’. The dentate with dark and stems on evergreen
outer side; mostly
teeth may involve both host and on leaves and
leaves
fungus tissue. Many species have a dentate with in deciduous
needles Propolis
Cryptodiscus dark, possibly wood which
black, almost carbonous outer side. 3 species
3 species black outer side; colours green
page 1476
page 1470 on wood and
leaves asexual state on
Other similar fungi: Fagus and Quercus;
pale to greyish;
– grades into the sessile, smooth on wood and stems apothecia mostly on
inoperculate cup fungi (page VB Quercus
of herbs
1426).
pale, mostly elongated, pale; Colpoma
– some large, inoperculate cup erumpent on
Pezicula somewhat 4 species
fungi are also erumpent, but (page 1462) elongated; deciduous and page 1480
form higher fruitbodies (page on deciduous coniferous trees
1404). wood
con at
– hysteroids have a slit-like p
erumpent; mostly elongated with
opening but are hard and the in clusters; ‘lips’; erumpent on
hymenium is rarely exposed on deciduous and p twigs
(page 1614). coniferous wood
erumpent Hypoderma
Diplonaevia 6 species
inoperculate p
(page 1447 page 1477
See also the general wheel, page cup fungi elongated with ‘lips’;
brownish-rose; (with lids, teeth or on stems of herbs,
1348. erumpent with teeth; slit-like opening) Rubus or leaves of
on herbs deciduous trees
p
Further reading: 18, 21, 49, 78,
139, 215, 297, 298. very small,
elongated with ‘lips’;
erumpent; on
p on stems, twigs,
Micropeziza grasses, sedges other cones, needles and
(page 1452 and rushes similar fungi leaves

mostly erumpent; a
with spherical VB; con somewhat elongated;
on leaves and stems buff with black outer
of herbs side; on bark of Lophodermium
conifers 22+ species
page 1478
erumpent; on hardly elongated;
Pyrenopeziza wood ± greyish with
(page 1438)
tough; black outer side;
erumpent on twigs
tough; with sinuous
in crowded cracks in a
clusters; on erumpent in
clusters; on less than 2 mm or more hard, mostly stroma; on
wood leaves of trees Pseudographis
deciduous 2 mm high high closed 2 species
wood; spring and shrubs
Micro-drawings: page 1482
spores first, followed Cenangium
by conidia (con), (page 1454)
paraphyses (p),
refractive vacuolar
bodies (VB), asci (a)
and ascus tops (at) Pseudophacidium
3 species
Approximate species Tympanis page 1482
number applies to (page 1406) Rhytisma
temperate Europe. 6 species
low, ± smooth high, ± smooth hysterioids
page 1483
inoperculate inoperculate (page 1614) © Thomas Læssøe & Jens H. Petersen
Encoelia & Sclerencoelia cup fungi cup fungi Fungi of Temperate Europe,
(page 1408) (page 1426) (page 1404) Princeton University Press, 2019
1484 lichens lichens 1485
crustose apothecial lichens
page 1492
Lichens foliose lichens (underside © Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
well delimited)
Lichenized fungi live in a mutualistic page 1490 Princeton University Press, 2019
sym­biosis with green algae and/or
cyano­bacteria that are able to perform
erect, simple to branched
photosynthesis. The algae/cyanobacteria (fruticose) lichens
(called the photobiont) provide the fungus page 1486
with sugars and the fungus affords shelter calicioid lichens
for the photobiont in the form of a ± well- page 1494
developed thallus (see page 19). Most
species are perennial and slow growing.
Lecanoralean lichens are by far the
largest group of lichens, with more than
1,500 species in temperate Europe. They
are traditionally studied by lichenologists
and are only briefly introduced on the
following pages. When sexual, the lecano­
ralean lichens produce apothecia on a
flattened, clavate or branched thallus. They
Ascomycota Ascomycota
have thick-walled asci with amyloid walls. with leaf- with completely
The calicioid lichens constitute an shaped thallus adhering thallus
aberrant group that form tiny fruitbodies and open and open
fruitbodies fruitbodies
with a dusty spore mass on the top of a (apothecia) (apothecia)
pyrenolichens
small stem (page 1494). when present
pages 1545 & 1615
Ascomycota with
Pyrenolichens form perithecioid fruit­ Ascomycota with stipitate fruitbodies
bodies and belong to the bitunicate, erect thalli and and passive spore
pyreno­mycetous fungi, pages 1615 and open fruitbodies dispersal
Ascomy (apothecia) when
1545. cota
present
Multiclavula Basidiom
Basidiolichens are lichenized basidio­ ycota Ascomycota
page 1114 lichens with ± closed
mycotes. They may be agarics, clavarioids
(fungi in symbiosis (perithecioid)
or (in warmer climates) corticioids (pages with algae and/or fruitbodies
130 & 1114). Basidiomycota cyanobacteria)
forming clavate
The most frequent photobiont is the fruitbodies
green alga Trebouxia, more rarely e.g. the
yellowish-green alga Trente­pohlia. Nostoc,
similar, non-lichenized
a genus of cyanobacteria, can also act or groups
co-act as the photobiont, and can benefit mycocalicioids have
the symbiosis by fixing nitrogen from the Basidiomycota active spore release from
atmosphere. with gills fruitbodies recalling those with ± closed fruitbodies and
(agarics) with apothecia of calicioids active spore release
Identification of lichens normally
involves the use of a series of chemical
characters but these are not covered here. Lichenomphalia
page 130
Other similar fungi:
– the inoperculate cup fungi have similar,
small, discoid fruitbodies but usually much
simpler asci (page 1348).
– Trizodia has a not fully understood
symbiosis with Sphagnum and
cyanobacteria (page 1432).
– mycocalicioids recall the calicioids but are
not lichenized (page 1497)
– ‘bitunicates’ and hysterioids may recall
crustose (crust-like) lichens (page 1602 &
1614).

Further reading: 302, 316, 355. Lecanoralean asci with thick-

walled tips.
inoperculate cup fungi Trizodia mycocalicioids non-lichenized ‘bitunicates’
(page 1348) (page 1432) (page 1497) (page 1602 & 1614)
1494 calicioids and mycocalicioids calicioids 1495

Calicioids and mycocalicioids algae such as Trebouxia, Stichococcus, Other similar fungi: Calicium form very small, stipitate
Calicioids and mycocalicioid fungi Dictyochloropsis and Trentepolia; the – Phleogena has larger fruitbodies; it fruitbodies with a widened head,
are two groups of distantly related mycocalicioid fungi are decomposers. is a basidiomycote with transversely the top of which is powdery from
ascomycotes with very similar, tiny, Many species can be found on divided basidia (page 1246). a thick, dark spore mass. Thalli are
pin-shaped fruitbodies. In some old, sun-exposed wood, but the small – asexual fungi may form very similar either superficial or partly immersed
species the spores are deposited in fruitbodies are easily overlooked; they structures (page 1626). in the substrate. Asci dissolve quickly;
a powdery column on the top and are best found by using the sky as a – many slime moulds may look similar spores 2-celled, thick-walled and
passively dispersed. The calicioids live background when inspecting potential but they are often larger and may have usually ornamented. Lichenized with
in a mutualistic symbiosis with green substrates. spores in a thread-like capillitium (page green algae belonging to the genus
1646). Trebouxia. Typically on sun-exposed
bark and wood.
Calicium Further reading: 302, 316, 326, 355.
13+ species Chaenotheca
page 1495 Calicium viride is a calicioid with a
17+ species
page 1496 finely grainy, yellow-green thallus.
Microcalicium The fruitbodies have rather long,
3 species Asci (dissolve at
page 1496 maturity in the black stems and a convex, black
calicioids). spore mass on the top. The lower
Chaenothecopsis
16+ species side of the head may be ± brown
page 1497
and finely powdery. Asci clavate.
Spores dark brown, 2-celled, 400 µm
have a ± regular to irregularly
spiralling pattern, and measure
Mycocalicium 12–14 × 6–7 µm. The algal partner
1 species
mostly with a tall page 1498
(photobiont) is a Trebouxia. Occurs
mostly with
column of spores;
yellowish or green mostly on dry bark and wood, both
asci dissolve quickly; deciduous and coniferous; mostly in
colours; spores mostly with a
spores 2-celled
mostly one-celled tall column of a sterile version. It is rather pollution
spores; spores
multicelled
tolerant.
on dead wood;
Calicium salicinum ▽ usually
spores mostly has an immersed thallus, browner
Phaeocalicium
one-celled fruitbodies, cylindrical asci and
8 species
page 1499 smaller spores; C. glaucellum ▷▷ has
on dead wood; small, dark and very open fruitbodies
spores one-celled Widespread, common to
occasional; all year. 5 mm

Calicium salicinum is a calicioid

on living twigs; with an almost invisible, ± immersed
lichens or parasite
Stenocybe
s
mature spores
2 species thallus, fruitbodies with a brown,
spores a 2-celled
page 1499 almost globose head topped with
± ornamented;
asci mostly a black spore mass, and stems that
are black towards the base. Asci
de

dissolve
co

quickly cylindrical. The spores are 2-celled,

m

on wood or
pos

spores smooth black, have a spiralling or irregularly

living twigs;
e rs

to finely ver­ru­
cose; asci ± mature spores cracking pattern and measure
persistent 4-celled 8–11 × 3.5–4.5 µm. The photobiont is
calicioids and
mycocalicioids a Trebouxia. Occurs on dry bark and
form spores in asci wood.
on the top of a
small ‘pin’ Calicium viride △ has a superficial,
yellow-green thallus, clavate asci,
larger spores and less brown powder
on the outside of the head.
other Widespread, common to
similar fungi
Micro-drawings: occasional; all year.
spores and ascus
tops (a). 300 µm

Approximate species
number applies to © Thomas Læssøe & Jens H. Petersen
Phleogena asexual fungi slime moulds/Myzetozoa Fungi of Temperate Europe,
temperate Europe. (page 1246) (page 1626) (page 1646) Princeton University Press, 2019
 Albertiniella
1500 mildews and cleistothecial fungi 1 species
mildews and cleistothecial fungi 1501
page 1505
Kernia
Mildews and cleistothecial 2+ species
fungi page 1506
Orbicula
Two form groups of ascomycotes with 1 species
page 1505
spherical fruitbodies are included here: Amylocarpus
1 species
the mildews (Erysiphales), which crack page 1505 Myxotrichum
open at maturity and disperse spores 10+ species
actively; and the cleistothecial fungi, page 1506
which do not open at maturity but decay Aspergillus
and disperse the spores passively. In both 100+ species
groups the fruitbodies are very small and page 1507
small, black cleistothecia
may act as dispersal units. To this end, smooth, brown with plate-like surface; triangular, black and
many have a surface with branched or ‘balls’ with yellow on dead Ganoderma shiny with hairs at
interior; the ‘corners’;
hooked appendages that may attach to on rotten litter and on faeces with black, hooked
passing animals, etc. Blumeria bird droppings hairs or ‘barbed-
1 species wire’-like hyphae;
The mildews are biotrophic plant page 1504 small, smooth, on straw and other
parasites that form whitish coverings of yellowish organic substrates
mycelia on stems, leaves and twigs, from cleistothecia; on wet
wood on seaside a
which haustoria can penetrate the living beaches ± smooth, pale Arthroderma
plant tissues (see page 16). The mealy J± per cleistothecia; on 10+ species
hyphal covering also produces asexual, a various plant litter, page 1507
a food items, etc.
wind-dis­seminated conidia. As leaves get a
de
older (typically in the autumn), mildews bio
co
mp
os
tr
often switch to producing tiny fruitbodies with simple op
hi
er
s
appendages; c
(chasmothecia) that overwinter and crack pa
r as a
on grasses ite
open when new leaves are available for so
n per
Podosphaera le
av hairy, pale
infection. 30+ species es,
et a cleistothecia;
c.
The cleistothecial fungi are decom­ page 1504 on hair, e.g. in
a
po­sers of organic material. The genera carnivore dung
Aphanoascus, Arthroderma, Gymnoascus only one ascus mildew and a
and Onygena (Eurotio­mycetidae) are per fruitbody; cleistothecial
specialists in degrading keratin (hair, appendages fungi
hyphae-like; on with ± spherical,
horn, etc.). Many have prominent asexual shoots and leaves a tiny fruitbodies and
states, but they may also produce tiny of herbs active or passive hairy, yellow
dispersal
spherical fruitbodies (cleistothecia, see cleistothecia;
a on hair, e.g. in
page 30) with ± globose asci; the spores carnivore dung
are released passively as the ascus walls a
and fruitbodies dissolve/decay.

other
Other similar fungi: with simple similar fungi per Gymnoascus
– pyrenomycetous fungi may have an or branched
appendages; on ± smooth, greyish 10+ species
almost invisible opening (ostiole), see cleistothecia; on page 1508
shoots and leaves of a a
page 1510. deciduous trees and hair, and carnivore
dung.
– truffles are ± spherical and decay at herbs
maturity but are much larger, see page
1256.
a
relatively large, to seveal
Further reading: 35, 47, 68, 72, 73, 78, mm high, ± stemmed,
79, 80, 164, 188. with sticky side and greyish cleistothecia; on
stilt-like appendages; feathers, horns and hoofs.
on shoots and leaves
of deciduous trees
Micro-drawings: Aphanoascus
outermost asci (a) and 5+ species
sometimes spores followed page 1508
by peridial appendages. Three
inset images depict peridia, Erysiphae
as seen in a compound 40+ species
microscope (per). page 1502 Onygena
2 species
Approximate species number Phyllactinia page 1508
applies to temperate Europe. 10+ species truffles pyrenomycetous fungi © Thomas Læssøe & Jens H. Petersen
page 1502 (see page 1256) (see page 1510) Fungi of Temperate Europe,
Princeton University Press, 2019
1510 pyrenomycetous fungi pyrenomycetous fungi 1511

Pyrenomycetous fungi non-stromatic, unitunicate

This is a form group of fungi with pyrenomycetous fungi non-hysterioid, bitunicate
page 1582 pyrenomycetous fungi

asci double-walled, spore release mostly via an inner sac

small, mostly spherical, flask-
page 1602
shaped or sac-like fruitbodies with
an opening (ostiole) from where the
spores, in most cases, are shot out.
The fruitbodies are here all termed
‘perithecia’ (see more on page 30),
although a more precise terminology
is often used by mycologists that
reflects ontogeny and phylogeny,
whereby the fruitbodies of

asci single-walled
ascomycotes with double-walled asci
(right-hand side of the wheel) may
be termed pseudothecia, thyriothecia
and hysterothecia.
In some species the perithecia hysterioid
are ± imbedded in a uniting flesh, a pyrenomycetous fungi
perithecia (pseudothecia, page 1614
stroma (pl. stromata), which may be perithecia thyrothecia) superficial or
flattened (applanate), cushion-shaped with ± circular immersed with ± circular
(pulvinate) or club-shaped (clavate). opening; to oval opening;
asci single- asci double-walled
Many species are decomposers, walled
decaying dead wood, herbs, dung,
etc, while others are parasites on
plants, mosses, insects, etc. Some perithecia ± black
kill insects and spiders and some are and in a stroma f ree or in
ecia ho
(surface dotted r i th st
lichenized. Many live inside living pe ti
perithecium with ostioles)
ri th cia ± bla
e
plant tissues (endophytic) and only pe ck

ss
ue

fruit when the plant dies. perithecia (hysterothecia)

ic
stromat
The pyrenomycetous fungi pyrenomycetous superficial and ± compressed;
fungi opening ± slit-like;
is a very large group with more (forming perithecial
asci double-walled
than 2,500 species in temperate

fruitbodies)
Europe. Parallel evolution has led
to similar-looking structures in perithecia and
stromata white
many ascomycote orders and only or ± brightly other
very pyrenomycetous fungi can hard, stromatic coloured and similar fungi
pyrenomycetous fungi rather soft
be identified without microscopy. page 1546
Experience and specialized literature
are required for detailed studies of
pyrenomycetous fungi, and there are
also good internet fora available (for fruitbodies perithecioid; on living at first closed, later opening to club-shaped, without dotted surface;
insects, millipedes, etc. reveal the hymenium hymenium exposed
references, see the following wheels).

Other similar fungi:

– Laboulbeniomycetes has very small,
peritecioid fruitbodies on living
insects, millipedes, etc. (page 1620).
– some erumpent inoperculate cup
fungi may be completely closed, but
the hymenium is revealed in wet
weather (page 1468). Laboulbeniales
– fungi with clavate, non-composite (page 1620)
fruitbodies may look similar, but
the hymenium is on the outside,
completely exposed (pages 1092 &
clavarioids and tongue-shaped
1350). inoperculate cup fungi
hypocrealean pyrenomycetous (page 1092 & 1350)
fungi and others erumpent inoperculate cup fungi © Thomas Læssøe & Jens H. Petersen
page 1512 (page 1468) Fungi of Temperate Europe,
Princeton University Press, 2019
 Trichoderma
1512 hypocrealean pyrenomycetous fungi and others 50+ species Protocrea
Arachnocrea
hypocrealean pyrenomycetous fungi and others 1513
page 1528 1 species
page 1533 1 species
Hypocreopsis page 1533 Nectria
Hypocrealean pyreno­ 2 species 3 species
page 1534
mycetous fungi and others page 1526

A form group of fungi with small, Neonectria

mostly globose to flask-shaped peri­ 8+ species Cosmospora, Dialonectria
page 1535
the­cia (page 30). The perithecia are & Macroconia
typically relatively soft (leathery) and Selinia 16+ species
1–2 species page 1536
often brightly coloured or whitish; Tubeufia & page 1527
Acanthophiobolus
they may be free, sit on a soft, often 5 species
brightly coloured stroma or immersed page 1524 effused,
± free or
cushion- or
in a stroma (page 30). The dots on club-shaped;
immersed; ± free; spores
the surface of the stromata represent spores split into split into two; on
spores split free to
stroma cushion to two; on fungi leaves and twigs
the ostioles (perithecial openings). into two; stromatic; on
finger-shaped; on fungi,
The spores are typically hyaline, or living and dead
on Hymeno­ wood, etc. wood
nearly so, and vary from multicelled, chaetopsis
± free; on living
thread-like (some split into part- stroma immersed/
and dead wood
cushion-shaped; Flammocladiella
spores) to one-celled; 2-celled spores spores large with splitting & Thyronectria
are common and may split into two at slime coat; on spores 11+ species
maturity. dung free; on fungi page 1539
Acrospermum
Many species are parasitic on 3 species free; with thick-
plants, animals or fungi. Species page 1523 walled asci; on wood, a
of Hypomyces may replace parts herbs and fungi
free to stromatic; Hydropisphaera
of the living fruitbodies of larger on wood and 3 species
fungi with perithecia. Cordyceps pyrenomycetous page 1537
fungi
and Ophiocordyceps are predatory club-shaped; mostly
fungi that kill insects and spiders, Neobarya on herbaceous stems
later producing stromata from the 7+ species free; on wood
carcasses. Epichloë and Claviceps live page 1522 and fungi
at
as endophytes in grasses and other hypocrealean Lasionectria
almost free,
monocots; they are well known due to mostly olive-green fungi 4 species
at
their toxic alkaloids (ergotism is caused perithecia; have ± soft, free; on wood, page 1538
on fungi white or brightly stems of herbs
by Claviceps). coloured and lichens
perithecia
Other similar fungi:
stroma tubular;
– hard, ± black pyrenomycetous fungi endophytes in grasses spores
free or immersed;
are more carbonous (page 1546).
Epicloë on fungi and
– clavarioid basidiomycotes and 10+ species other myzetozoa
tongue-shaped inoperculate cup fungi, page 1521 similar fungi
stroma club-shaped,
have external hymenia, and thus no from sclerotia;
parasitic on grasses at
dots from immersed perithecia (pages a stromatic or rarely
and Eleocharis
1092 & 1350). ± free; mostly on
splitting
Claviceps spores fleshy fungi
3 species Nectriopsis
See also the general wheel, page 1520 stroma club-shaped;
spores split into 8+ species
page 1510. part-spores; on free; with thick page 1540
Elaphomyces ascus walls; on
stroma club- spores formed spores formed
wood, fungi
shaped; with inside perithecia externally on
Further reading: 48, 55, 60, 96, 106, (incl. lichens) or
splitting spores; club-shaped
107, 128, 129, 131, 132, 175, 248, stroma lichenized
on insects fruitbodies
274, 275, 278, 279, 280, 284, 302, flattened; with
splitting spores;
311. on spiders
Tolypocladium
4 species
Micro-drawings: page 1518
spores first, followed
on the inside by
asci (a) and ascus Hypomyces &
tops (at). Cordyceps & Sphaerostilbella
Ophiocordyceps Thelocarpon 38+ species
Approximate species 11 species 12+ species page 1541
number applies to page 1515 page 1545
temperate Europe.
hard, ± black tongue-shaped
Torrubiella pyrenomycetous fungi inoperculate cup fungi clavarioids © Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
3 species, page 1514 see groups page 1546 (page 1350) (page 1092) Princeton University Press, 2019
1546 hard, stromatic pyrenomycetous fungi hard, stromatic pyrenomycetous fungi 1547
Cryptosphaeria Eutypa
3 species 8+ species, page 1569 Eutypella
page 1568 10+ species, page 1571 Micro-drawings:
Hard, stromatic Quaternaria spores, asci (a) and
pyrenomycetous fungi Kretzschmaria
Lopadostoma 2 species ascus tops (at);
7 species page 1571 perithecia = p;
A form group of fungi with black, 1 species page 1566 dehiscent in KOH (KOH).
page 1563
mostly rather carbonized perithecia
Approximate species
that are ± immersed in a stroma (plural Diatrype number applies to
stromata), and which may be club-, 6 species temperate Europe.
page 1572
nail- or cushion-shaped or quite flat.
The stroma may be well defined, as in
most genera on the left hand side of Nemania, Entoleuca
& Euepixylon
the wheel, or composed of a mixture 15 species, Diatrypella
of tissue from the host plant and the page 1564 8+ species
page 1574
fungus, as in many species on the right perithecia
hand side of the wheel. immersed; immersed with
in wood/ Peroneutypa
asci expanded
All species are decomposers, but ± cylindrical, bark stroma;
erumpent p. mostly 1+ species
from bark in clusters of page 1574
some may also kill the host, e.g. I+(–); in in wood/bark
large, black and four; erumpent

as
Kretzschmaria deusta and Eutypa Biscogniauxia wood/bark

as
from bark

ci
crumbly; asexual at

ci
10 species erumpent

± ta lindric
spinosa. Some species may survive as state prominent;

±c
page 1562 from bark
on wood

per
y
endophytes in living tissues and only ostioles
asci

in
umbilicate or
become active when the host dies.

g
multi-spored;
conical; brown

al
The majority of the hard, stromatic erumpent from
to black; apical
bark Diaporthe &
pyrenomycetous fungi are found in apparatus tall;
Diaportella
on wood long necked;
three families: Xylariaceae has brown, 35+ species
a erumpent from page 1575
± asymmetrical ellipsoid spores, bark
erumpent; ostioles
mostly with a germ slit; Diatrypaceae Jackrogersella ± conical; ± black; tac
hed
3 species i at ing
asc
has paler allantoid spores and on wood il o o sen
page 1561 asc
asci with a long tail-like base; mostly with long,
projecting necks; spores
and Diaporthaceae has hyaline, ostiole conical, brown
2-celled; erumpent from
1–2-celled spores and loosening to black; apical a
apparatus low; stromatic, dark bark
KOH pyrenomycetous
asci. The first two families mostly on wood
fungi have spores one-celled;
have an amyloid ascus apparatus. Hypoxylon
ostioles umbilicate, black, rather hard
22+ species disc-like;
The genera Daldinia, Hypoxylon, page 1556 with ± vivid KOH perithecia in a erumpent from
and Jackrogersella have diagnostic pigments; apical stroma bark
apparatus low; at
pigments that can be dissolved on wood Cytospora
a a a
with KOH; scrape the surface onto a disc-like; 15+ species
erumpent from page 1576
blotting paper and add a drop of rather large, with
zonate stroma in bark
10% KOH (corrosive) – the pigment
section; on wood
will be drawn into the paper (see
disc-like;
pages 1557–1560). Daldinia erumpent from
rather large, x1/2 as
9 species as
ci bark
± orange with lo
page 1554 ci os
± 
Other similar fungi: watery interior; x at
en
in
1/2 ta g
– hypocrealean pyrenomycetous on wood at at broadly ched Amphiporthe
fungi have a softer texture and at erumpent 3+ species
stroma with a at
broadly from bark page 1577
paler or brighter colours (page flat or round erumpent from
1512). head on a stem;
p. ± spherical;
p. mostly bark
on dung stroma club- oblong;
– the non-stromatic pyreno­myce­ shaped or with double-walled
in wood
asci
asci

asci; on wood

asci singl
tous fungi and the bitunicate branched;
sing

on wood or litter
dou

pyrenomycetous fungi have ± free

le-w
b

perithecia (pages 1582 & 1602).

l e

e-wa
Melanconis
-
alle
wal

8+ species
lled
d
led

See also the general wheel, page 1577

page 1510. Entonaema
1 species Melanamphora
page 1555 1 species
Further reading: 61, 78, 82, 83, 93, page 1578
130, 179, 250, 251, 254, 255, 256,
Sillia
313, 314, 353. Poronia & 1 species
Podosordaria page 1578
4 species
page 1552 Xylaria
Camarops © Thomas Læssøe & Jens H. Petersen
12+ species Cucurbitaria Fungi of Temperate Europe,
page 1548 25+ species, page 1581 7 species, page 1579 Princeton University Press, 2019
1582 non-stromatic, unitunicate, dark pyrenomycetous fungi non-stromatic, unitunicate, dark pyrenomycetous fungi 1583
Barbatosphaeria
7 species, page 1588
Calosphaeria
Non-stromatic, unitunicate, 5+ species Micro-drawings:
dark pyrenomycetous fungi Natantiella page 1588 Melanospora spores first, followed
1 species 7+ species on the inside by
Species in this form group have more Lentomitella page 1587 page 1589 asci (a).
or less solitary, perithecia and the asci 7 species
page 1587 Approximate species
have single walls (unitunicate). They are number applies to
generally ± brown or black but may be temperate Europe.
covered in white or more colourful hairs.
The perithecia may be seated on the Pseudohalonectria
3 species
surface of the substrate or immersed and page 1590
almost invisible, with only the ostiolar Ophiognomonia necks ± hairy;
10+ species asci with asci loosening; necks ± smooth;
necks appearing at or above substrate asci non-loosening;
page 1587 thin, tail- perithecia between
level. like base, bark and wood perithecia between
The majority of the species are loosening; bark and wood
± hairy with thin
on wood
decomposers of leaves, herbaceous asci non-loosening; necks and spores
stems, wood or dung – a few are spores longitudinally in a drop; on other
ribbed, hyaline, fungi
parasitic.
1–3-celled; on wood Ceratosphaeria
The non-stromatic pyrenomycetous 1 species
with long necks,
fungi form a heterogeneous group of permanently immersed in wet
page 1590
fungi with representatives from a range Gnomonia immersed; thin- wood; spores
10+ species necked;
of orders, of which Sordariales is the multicelled
page 1586 in leaves of
most prominent. Many of these species Betulaceae and
produce large, ± brown spores. Fagaceae

a asci attached;
Other similar fungi: thin-necked, loosening
a a spores hyaline,
a multicelled;
­– the bitunicate pyrenomycetous fungi asci and narrow spores
in a drop; on wood
have thick, double-walled asci. The a
a
superficial on a Sordaria
spores are mainly released when the leaves and wood of 12+ species
outer wall ruptures, allowing an inner Betulaceae page 1595
a
sac to stretch out (page 1602). long-necked, a
spores brown
– hypocrealean pyrenomycetous fungi non-stromatic with slime coat,
can also be non-stromatic, but the with thin necks and pyrenomycetous a one-celled, with
spores in a drop; fungi germ pore;
perithecia are mostly brightly coloured a asci single-walled
on other fungi, on dung
and rather soft (page 1512). mostly insect
– the dark stromatic pyrenomycetous parasites
short-necked or
fungi grade into the non-stromatic. neckless, non-stromatic
For species with immersed perithecia it other

⬅
pyrenomycetous fungi,
Syspastospora similar fungi
can be especially difficult to determine see next page spread
1 species with very thin
whether or not a stroma is present (page page 1586 necks and spores
in a drop at the tip;
1546). on wood and old
polypores
See also the general wheel, page 1510.
asci single-walled asci double-walled
Further reading: 16, 54, 58, 62, 73, 78, (unitunicate) (bitunicate)
185, 195, 196, 247, 257, 258, 259, 260,
261, 304, 350, 353.
Sporothrix
10+? species
page 1586

Asci with single walls (unitunicate). hypocrealean pyrenomycetous fungi and others stromatic bitunicate hysterioid and
© Thomas Læssøe & Jens H. Petersen
(mostly soft, mostly coloured) pyrenomycetous fungi pyrenomycetous fungi Fungi of Temperate Europe,
(page 1512) (page 1546) (pages 1602 & 1614) Princeton University Press, 2019
1584 non-stromatic, unitunicate, dark pyrenomycetous fungi Bombardia non-stromatic, unitunicate, dark pyrenomycetous fungi 1585
1 species
Schizothecium page 1594 Sordaria
8+ species, page 1594 12+ species
page 1595
Podospora
25+ species Echinosphaeria
page 1593 4+ species
page 1595 Helminthosphaeria
12+ species
page 1596
Immersiella
2+ species
page 1593

1–2 mm high, thick Chaetomium

smooth and pear- 20+ species
hairs in tufts; and rubber-like;
shaped with a hairs brown, page 1595
downy-hairy, not on dung on wood
short neck; thick-walled,
tufted-scaly; on dung septate; spores
on dung boomerang-
Ruzenia
shaped; on wood with dark setae;
1 species
± felty-hairy; spores brown, with
page 1593
in/on wood small germ pores;
mostly on other
Lasiosphaeria
fungi
10+ species ± smooth; Chaetosphaerella
page 1592 mostly in dense 2 species
spores collected in
groups; spores page 1596
a column; asci soon
boomerang-shaped;
dissolve; on litter and
on wood
dung

smooth or white-
to yellow-haired, on black hyphal mat
non-setose; spores h with production of
S-shaped; on wood or conidia hyphae; on
Lasiosphaeris herb stems wood/fungi
a Acanthonitschkea
2 species h 2 species
page 1592 con page 1597
a short-necked
or neckless, black setose;
with thick-walled, h ± cupulate when
non-stromatic
black hairs; on wood pyrenomycetous h dry; on wood and
h other fungi
fungi
a
asci single-walled

± smooth;
Strattonia ± cupulate when Nitschkia
h
5 species on burnt soil or dung dry; on wood and 6+ species
page 1591 other fungi page 1598
a

mostly hairy; smooth but typically

spores with sticky a amongst dark hair-
appendages; long-necked, like asexual states;
on dung, herbs and non-stromatic on wood
wood pyrenomycetous

⬅
fungi,
Arnium see previous page spread Chaetosphaeria
with black setae; mulberry-like;
8+ species spores with germ slit; on wood 15+ species
page 1591 on wood, dung etc. page 1598
confluent dark
spots; on living plant
perithecia large; hyphal with shield on tissue
mat ± prominent, on top; on dung
wood or herbs
Bertia
2 species
Micro-drawings: Coniochaeta
page 1598
spores first, followed 15+ species
on the inside by asci page 1591
(a), hairs (h) and
conidia (con).
Phyllachora
Approximate species
10+ species
number applies to
page 1599
temperate Europe.
Hypocopra © Thomas Læssøe & Jens H. Petersen
Rosellinia 10+ species, page 1599 Fungi of Temperate Europe,
12+ species, page 1600 Princeton University Press, 2019
1602 non-hysterioid, bitunicate pyrenomycetous fungi non-hysterioid, bitunicate pyrenomycetous fungi 1603
Melanomma
Capronia
4 species, page 1610
20+ species, page 1611
Non-hysterioid, bitunicate Leptospora
Cucurbitaria Micro-drawings:
pyrenomycetous fungi Lophiostoma 1 species
(see page 1581) spores first,
15+ species page 1609 followed on
Species in this form group have thick- page 1609 Venturia the inside by
25+ species asci (a).
walled asci with several functional wall page 1612
layers. Most species have small, free Approximate
perithecia (also termed pseudothecia) or species count
Pleospora applies to
even smaller, circular, flattened perithecia 40+ species temperate
(termed thyriothecia – page 1604). page 1608 Europe.
The young asci are typically very thick-
perithecia small,
walled, especially the upper parts, where crowded; perithecia small;
long-necked;
a characteristic bulge often can be seen on wood on plants and fungi
on herbaceous mostly stromatic;
(see arrow below). The spores are ejected ostiole stems mostly below bark
compressed; Trichodelitschia
from an inner sac that stretches out from Sporormiella on wood and 4 species
the outer sac once the ascus tip ruptures 30+ species herbaceous stems very small with page 1611
page 1608 setae; on leaves
– a so-called jack-in-the-box mechanism. ostiole mostly
Most species in this form group are somewhat Pseudotrichia
flattened; mostly 3 species
plant parasites, or decomposers of wood, page 1612
below the surface
herbs or dung. of herbaceous small with setae;
stems on dung
Other similar fungi:
Ophiobolus mostly
– the ± black, non-stromatic immersed; on
15+ species
unitunicate pyrenomycetous fungi are page 1608 dung
with yellowish or
similar but the asci are single-walled orange hairs; on
(page 1582). wood of deciduous
trees Tubeufia
– the hysterioid pyrenomycetous fungi pear-shaped to
(see page
conical; mostly a a
have similar asci (and phylogenetically a a 1524)
below the surface
belong to the same class), but have of herbaceous a
a a
stems
compressed, superficial fruitbodies one part-spore bitunicate, small, ± soft and
that open by a narrow slit (page pyrenomycetous brown to yellowish;
pear-shaped to fungi on wood and fungi
1614). conical; mostly with, ± black
below the perithecia and a
surface of stems thick-walled asci
See the wheel page 1510.
of herbs very small, soft
Leptosphaeria and yellow;
a
Further reading: 61, 73, 77, 78, 75+ species on wood, fungi
page 1607 large, thick-walled,
84, 234, 299, 300, 302, 312, ± semiglobose; (incl. lichens) etc.,
a some species
347, 353, 357. mostly below bark other
a lichenized
of deciduous trees similar fungi

black, in blue-
small, convex green thallus; on
pseudothecia; on Thelocarpon
bark; (see page 1545)
Massaria living herbs, fallen ostioles rounded;
Asci in most bitunicate pyrenomycetous lichenized
15+ species leaves, etc. asci single-walled
fungi have a characteristic thick-walled tip page 1606
with a bulge, here in Cucurbitaria.
small, flattened, black, in black
circular, with slit-like ostioles;
asci double- thallus; on rocks;
central ostiole; lichenized
on living herbs walled
tiny, flattened,
circular; Pyrenula
Mycosphaerella on leaves, stems 10+ species
100+ species and lichens tiny, flat-topped; page 1613
page 1605 on pyreno­
mycetous fungi

Asterina
1 species Hydropunctaria
page 1605 8 species
page 1613
Asci with bitunicate walls – from left Leptosphaeria (one
mature and one with the inner sac ejected), Cucurbitaria, Lichenopeltella
Sporormiella, Venturia, Massaria & Asterina. 20+ species, page 1604 © Thomas Læssøe & Jens H. Petersen
Pachythyrium non-stromatic hysterioid, pyrenomycetous fungi Fungi of Temperate Europe,
1 species, page 1604 pyrenomycetous fungi, page 1582 page 1614 Princeton University Press, 2019
1614 hysterioid pyrenomycetous fungi low, elongate, hysterioid pyrenomycetous fungi 1615

Hysterioid axe-like. All have double-walled asci Other similar fungi: Morenoina pteridicola is a black,
pyrenomycetous fungi (bitunicate) that typically release the – inoperculate cup fungi with slit- bitunicate pyrenomycetous fungus
This group of fungi includes species spores after an inner sac has stretched like openings reveal the hymenium in with oblong, somewhat branched,
with mostly superficial perithecia that through the outer sac (jack-in-the box- wet weather. They have single-walled very small, ± flattened fruitbodies
have distinct, ± compressed ostioles release). (unitunicate) asci (page 1468). that have slit-like ostioles. The margin
and a slit-like opening (hysterothecia). All species are decomposers of – some bitunicate pyrenomycetous has brown, radiating, ± branching
The fruitbody shape is highly variable: wood, cones, herbs, etc. They mostly fungi (e.g. Lophiostoma) have flattened hyphae. Asci double-walled, measure
many are elongate, others clam- or occur in exposed, dry habitats, e.g. on ostioles, but the fruitbodies are 18–27 × 9–14 µm. The spores are
bark of ± sun-exposed twigs. immersed in the substrate (page 1602). 2-celled, not completely symmetrical,
smooth, somewhat brownish when
Further reading: 41, 42, 368. old, measure 9–13 × 3–4 µm. Occurs
Morenoina Rhopographus on petioles of Dryopteris, Pteridium
1 species 1+ species Hysterographium and probably other ferns.
page 1615 page 1615 2 species
page 1616 There are apparently no other
Hysterobrevium Micro-drawings: similar species on these substrates.
2+ species spores first, Possibly widespread, but very rarely
page 1615 Gloniopsis followed on the
2 species reported, most likely overlooked;
inside by asci.
page 1616 most of the year. 200 µm
Approximate
species number Rhopographus filicinus is a
applies to tem­pe­
Hysterium rate Europe. bitunicate pyrenomycetous fungus
2 species that forms elongated-oblong,
page 1617 black, ± confluent fruitbodies
very small, ± cylindrical;
lip-shaped; with slit-like ostioles along the
sinuous; on on Pteridium length of the hosts’ ‘stems’. Asci
on bark
ferns of deciduous long, ± sinuous; double-walled, 8-spored, measure
trees on bark
of deciduous 70–86 × 20–25 µm, I-. The somewhat
lip-shaped; Acrogenospora
trees
on bark 2 species bent, yellow-brown, 4–8-celled
of deciduous page 1617 spores measure 27–35 × 7–8 µm.
trees Occurs on petioles of Pteridium.
± lip-shaped; Scirrhia aspidiorum ⋉ forms grey
on bark
of deciduous fruitbodies/stromata and hyaline
trees 2-celled spores.
Actidium
high, lip- to clam- 3 species
Widespread, very common; all
shaped; on bark page 1619 year.
of deciduous and
coniferous trees
4 mm 200 µm

± clam-shaped; Hysterographium fraxini has

on bark
of Juniperus
Mytilinidion elongated, black, longitudinally
6+ species
page 1618
furrowed, half-open fruitbodies
(hysterothecia) that may branch
± clam-shaped; slightly. Asci double-walled, I-. Spores
on conifers incl. yellow-brown, muriform-septate
hysterioid Juniperus
pyrenomycetous with a slightly constricted middle
fungi septum and many transverse and
with compressed, longitudinal septa; they measure
slit-like ostioles Lophium
clam-shaped; 40–48 × 16–20 µm, are mostly broadly
2 species
on bark, wood
page 1618 rounded and have a slime coat. Mainly
and cones of
other similar fungi conifers occurs on Fraxinus bark.
Hysterographium flexuosum ⋉
has somewhat longer, more pointed
axe-­shaped; spores; H. elongatum ⋉ apparently
on bark of has 1 or no longitudinal septa, but is
deciduous trees Glyphium perhaps just a synonym of H. fraxini.
2 species
Widespread, occasional; all year. 400 µm
page 1619

inoperculate cup fungi with immersed bitunicate © Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
slit-like opening (page 1468) pyrenomycetous fungi (page 1602) Princeton University Press, 2019
1620 Laboulbeniales
Laboulbeniales
The Laboulbeniales are tiny, 0.03–2 mm
high, ascomycotes that grow on
living insects, millipedes and other
arthropods. Each species has one or
several hosts and some are always
situated on specific areas of the host,
e.g. the antenna. The entire fungus
(thallus) includes a dark ‘foot’, a stem,
a perithecium (with asci and 2-celled
spores with a slime coat), antheridia,
which form spermatia, and finally
hypocrealean pyrenomycetous fungi,
(page 1512)
1 mm
appendages
antheridium
spores
perithecium
50 µm
sterile appendages. During the sexual
process spermatia are transferred from
the antheridia to a trichogyne. After
this the perithecium develops.
The Laboulbeniales do not have
a true mycelium, and apparently do
not enter the host cavity, but get their
nutrients only from the exoskeleton.
Large aggregations of fruitbodies may
weaken some hosts.
The Laboulbeniales belong in their
own class, Laboulbeniomycetes, with
other parasites
on arthropods
asexual fungi
(page 1626)
stem
foot cell
Jimmie Høier
50+ genera in temperate Europe. Only
a few examples are included here – all
from beetles.
Other similar fungi:
– insect parasites are also found
among the hypocrealean fungi (page
1512), and within the asexual fungi
(pages 1630, 1635 & 1643), but none
resemble Laboulbeniales.
Further reading: 114, 117, 187, 352.
the Laboulbeniales
are tiny fungi on living
arthropods, e.g. beetles
and flies
Laboulbenia is the largest genus
of the Laboulbeniales. It has a short
foot cell and lateral appendages.
Laboulbenia argutoris is an
approximately 275 µm long
Laboulbenia with a dark olive-brown,
perithecium about 125 µm long. The
outer, simple appendages may have
up to 9 cells. Spores asymmetrically
2-celled, hyaline, and have a thick
slime coat that enables it to stick to
the host animal. Occurs on beetles in
the genus Pterostichus.
At least four species of
Laboulbenia occur on this type of
beetle. Laboulbenia pseudomasei ⋉
and L. flagellata ▷ have more
complex outer appendages than
those in L. argutoris. Laboulbenia
kajanensis ⋉ is very similar to L.
argutoris but has small differences in
cell dimensions.
Distribution and frequency not
well known, but probably rather
common; all year.
Laboulbeniales 1621
Laboulbenia flagellata is a ± olive-
green Laboulbenia with a thallus
about 400 µm long. The perithecia
appear from three large basal cells
that arise from the dark attachment
cell. The appendages appear from
a 2-celled extension of the basal
cells. Occurs on a number of carabid
beetle genera and species.
A very variable and rather difficult
species to identify, with identification
further complicated by the long list
of hosts.
Probably widespread and
common; all year.
50 µm
Jimmie Høier
Monoicomyces fragilis is a
laboulbenialean fungus with thalli
approximately 200 µm long and
140 µm wide. Each thallus divides
above cell two, and one perithecium
develops on each side. Two long
appendages with brown basal
pigmentation arise from just above
the lower pigmented cell. Spores
hyaline, 2-celled, and have a slime
coat. Occurs on the rove beetles
Ocalea picata and Oxypoda opaca.
The distinctive split thallus makes
this species easy to identify.
Distribution and frequency poorly
known; all year.
50 µm
Jimmie Høier
Rhachomyces furcatus is a
complex laboulbenialean fungus,
up to 500 µm long with many
dark appendages and one or two,
± centrally positioned, approximately
250 µm long perithecia. The cell size
gradually increases up to the cell
that supports the perithecium, and
decreases again above it. The spores
are 2-celled, hyaline, and have a
slime coat. Occurs on rove beetles in
the genus Othius.
A relatively easy species to
recognize, provided the host animal
is taken into account.
Distribution and frequency poorly
known; all year.
80 µm
Jimmie Høier
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
Princeton University Press, 2019
1622 Taphrinales
Taphrinales
Species in the order Taphrinales are all
biotrophic plant parasites that never
form proper fruitbodies, but mostly
produce a layer of asci within the host
tissue. The hosts may be galled in
various ways, from small leaf blisters to
huge witch’s brooms in the crown of
infected trees.
All belong in Taphrinomycotina
– one of three subphyla within the
Exobasidium Erysiphales
(page 1214) (page 1500)
phylum Ascomycota. The other
two subphyla are the yeasts in
Saccharomycotina and the often
fruitbody-forming Pezizomycotina
(page 12).
Other similar fungi:
– the basidiomycotes in Exobasidium
parasitize species of Ericaceae (page
1214).
– Erysiphales are more mealy from
other
similar groups
non-fungal
(page 1645)
surface production of conidia and form
small spherical fruitbodies with asci
(page 1500).
– some non-fungal organisms, Albugo
for example, may look similar but do
not have basidia or asci (page 1645).
Further reading: 78, 80, 87, 164.
Taphrinales
parasitizes plants
– often visible as galled
tissues
Protomyces
Taphrina page 1625
page 1622
Taphrina is a large genus of plant
parasites. All species deform the
infected tissue in more or less
spectacular ways: from curly or
blistered leaves, through deforming
fruits, to inducing ‘witch’s brooms’.
The small asci are formed in a
palisade directly on the surface of
the host tissue. The spores mostly
produce yeast stages within the asci.
When sporulating the infected tissue
may have a pruinose appearance.
Taphrina betulina causes the
formation of Magpie nest-like witch’s
brooms. The asci are formed in the
leaves of the host, which appear
from the brooms; they measure
25–70 × 10–25 µm and are I-. Spores
measure 4–6.5 × 2.5–5 µm. Occurs in
the crown of living Betula trees.
Other organisms may also
cause the formation of witch’s
brooms in Betula, so in reality
reliable identification requires
microscopy. Taphrina betulae ⋉
forms discoloured, swollen spots
on the leaves of Betula rather than
conspicuous galls.
Widespread, very common; all
year (as brooms).
Taphrina 1623
Taphrina wiesneri causes a very
spectacular formation of drooping
witch’s brooms. The asci are formed
on discoloured, somewhat bumpy
leaves; they measure 17–35 × 15 µm
and are I-. The spores measure
3.5–9 × 3–6 µm and form yeast-like
cells. Occurs on both cultivated and
wild species of cherry (Prunus cerasus
and P. avium).
In order to ensure a correct
identification, infected leaves require
microscopical investigation.
Mainly nemoral–hemiboreal,
rather common; all year (as brooms).
Taphrina carpini forms conspicuous
witch’s brooms. Asci occur on the
pale leaves of the host that appear
as long shoots from the brooms;
they measure 20–30 × 7–15 µm. The
spores measure 3.5–5 × 3–4.5 µm.
Found on Carpinus, including in
mature hedgerows.
Microscopical investigation is
recommended as other organisms
can induce the formation of witch’s
brooms in Carpinus.
Nemoral, rather common; all year
(brooms).
Tom Smidth
Taphrina farlowii deforms leaves,
shoots and fruits of the host with
yellowish to reddish blisters and
swollen galls. The asci are formed
on the galls and measure 20–30 × 
8–9 µm, the basal cell measures
8–9 × 15–25 µm. The spores are
± spherical to broadly egg-shaped
and measure 4–6 × 4–5 µm. Affects
Prunus serotina, a North American
species that is widely planted and
now spreading.
The very similar T. deformans ⋉
deforms the leaves of other Prunus
species, including P. amygdalus and
P. persica.
Nemoral, probably common;
mainly June–October. 5 mm
© Thomas Læssøe & Jens H. Petersen
Fungi of Temperate Europe,
Princeton University Press, 2019
1626 asexual fungi (hyphomycetous, coelomycetous, mucoralean, entomophthoralean fungi) and Oophyceae asexual fungi 1627

hae
hyphoid (hyphomycetous)

ngia
hyp
Asexual fungi asexual fungi)

o ra
page 1635

sed
l sp
Many fungi have asexual states where

xpo

ica
ne
propagules are produced following mitotis

her
no
rather than meiosis. These mitospores

± sp
tio
(typically termed conidia) are ± genetically

duc

in
res
identical to the mycelium that produced

pro
spo
them. Contrary to this, meiospores (in this

idia

gio
con
publication termed spores) are formed cushion-shaped or bulbil-forming

ran
asexual fungi
in, for example, asci, or on basidia, by a

spo
page 1632
sexual process that involves nuclear fusion
and subsequent meiosis, see pages 11 &
22.
Many species produce both sexual and
asexual states. Where the two states occur
together both states are usually included
in the description of the sexual state.
Traditionally, asexual and sexual states
were given separate scientific names. For
example, Aspergillus glaucus (page 1507) Mucorales
page 1640
was the name for an asexual state, while
Eurotium herbariorum was the name ran
gia
al spo
for the sexual state of the same species. eric les)
s in
± sph hth
ora
ore mop
However, in 2011, the code that governs ran
gio s p
lly
(En t o
hyphoid; mostly spo rna
the naming of fungi was changed so exte
dusty med
ia for
that one name is used, the first described conid

taking precedence irrespective of the state cushion-shaped, ± pin-shaped

to which it was attached. Thus, Aspergillus greasy-slimy
or;dusty Entomophtorales
glaucus became the valid name for all
page 1643
states of this fungus. club-shaped (synnematoid)
asexual
Asexual fungi constitute a pool of asexual fungi
states
page 1629 club-shaped; waxy;
thousands of ‘species’. These are not a greasy-slimy or mostly on insects
main theme of this publication, and on the dusty
following pages only a few characteristic
examples are given.
Asexual propagation is found in almost atypical;
all fungal groups. However, the more at first closed, hyphae without
other septa
spectacular examples are found mainly perithecioid or
similar groups
within the ascomycotes and zygomycotes, hysterioid
and less so within the basidiomycotes.
co
nid
ia
fo
Other similar fungi: rm
ed
ex
– species of mildew form asexual spores ter
na
lly
from a mealy covering on living leaves, but (En
to
mo
later form tiny, spherical fruitbodies with ph
th
or
al
internal asci (page 1500). Oo es)
ae ph
ph fungi fungi yc
hy ea
– rusts and smuts have a series of asexual sed with with e
po asci basidia
spore states, see page 1198. ex ll
y
on na
– cobweb-like corticioids may look similar ct
io
n
in
t er
u n
to asexual moulds but form meiospores pr
od
u ct
io
a od
from basidia (page 986). di
ni pr
co ia
id
– Myzetozoa (slime moulds) may look co
n protozoan
similar to asexual fungal moulds (page
1646). hyphoid, non-fungal species
coelomycetous fungi (Oophyceae)
page 1628 (page 1645)
Further reading: 78, 94, 144, 145, 146,
293, 294, 321, 351, 369.

© Thomas Læssøe & Jens H. Petersen

mildews rusts and smuts corticioids Myzetozoa Fungi of Temperate Europe,
(page 1500) (page 1198) (page 986) (page 1646) Princeton University Press, 2019
1646 Mycetozoa Mycetozoa with external spores 1647

Mycetozoa sporulating stage mostly has internal fruitbodies but transversely divided
Mycetozoa (slime moulds) do not spore production and the spores can basidia.
belong in the fungal kingdom but be mixed with a hyphoid capillitium. – small clavarioids have basidia and
within the Protozoa, However, their There are approximately 48 genera in are not dusty at maturity (page 1092).
spore-producing structures may temperate Europe; a few examples are
resemble those from fungi. They included here to show the variation. Further reading: 120, 217, 249.
ingest organic particles in their mobile
amoeboid stages and can be found Other similar fungi:
in many environments, particularly on – asexual fungi may look similar (page
bark and rotten wood. The amoeboid 1626).
stages range from tiny to huge. The – Phleogena (page 1246) has dusty

Ceratiomyxa
page 1647 cushion-shaped
(aethalia-forming)
Mycetozoa
page 1648
A plasmodium of Badhamia utricularis.
ty

composite, cushion-shaped
maturi

ty

(pseudoaethalia-forming)
maturi

Mycetozoa
4 mm
ery at

page 1650
sty at
al; wat

Ceratiomyxa fruticulosa is a white

al; du

to somewhat yellowish slime mould

extern

intern

with external spore production

(inset image). It forms large, dense
spores

oblong to sinuous
spores

(plasmodiocarp-forming) areas of long-branched fruitbodies

Mycetozoa that ± dissolve when touched. The
page 1652
egg-shaped to broadly ellipsoid,
mostly hyaline, smooth spores are
large, cushion- positioned on a small stem and
when mature still shaped, dusty
soft and watery measure 8–15 × 6–10 µm. Occurs on
large, rotten wood, both deciduous and 1 mm
composite,
cushion-shaped, coniferous.
dusty The variety Ceratiomyxa fruticu­
losa var. porioides (lower image)
sinuous, almost looks like a resupinate
net-like, polypore but is much softer and
dusty
disintegrates when touched.
narrowly adnate to ± stemmed
(sporocarp-forming) Mucronella (page 1096) and
Mycetozoa Mycetozoa Ceratellopsis (page 1097) are firmer
are protozoa small, narrowly page 1653
with fungus-like when touched and have basidia.
adnate to
fruitbodies ± stipitate, dusty Ceratiomyxa is not closely related to
the other groups of slime moulds.
Widespread, very common; May–
October, peaking during the summer.
other similar fungi

4 mm

Thomas Læssøe
© Thomas Læssøe & Jens H. Petersen
asexual fungi Phleogena clavarioids Fungi of Temperate Europe,
(page 1626) (page 1246) (page 1092) Princeton University Press, 2019