PERSOONIA
Volume
Coprotus
17,
Part
4,
649-656
Jacques+R.
Unite
d'ecologie et
de
coprophilous species
new
reported
a
from five localities
of
group
It could
not be identified
to
as
first
was
Catholique
de
Louvain,
4-5 Place
Coprotus (Pezizales, Pyronemataceae)
of the Ardennes
any
of
Ascomycota)
Croix
Sud,
du
Louvain-la-Neuve, Belgium
identified
species mainly
Korf & Kimbr.
Coprotus
of the genus
of
new species
(Pezizales,
de Sloover
University
biogeographie,
B-1348
A
a
arduennensis,
coprophilous discomycetes
(2002)
by
the
of the 25
Within the
(Belgium).
presence
suggested by
it
of carotenoids within the
species known,
Korf
comprise mainly coprophilous
is described
genus,
to
paraphyses.
but is closest to C. ochraceus.
who later drafted the
(1954)
with
species
and
belongs
diagnosis
non-amyloid 8-spored asci,
smooth, hyaline ascospores and uncinate paraphyses i.e. species traditionally placed within
the old
& Korf
with
species
paraphyses
In the basic paper of
added
seven more
species.
of the 25 described
species,
al.,
1972).
et
et
here)
to
et
described
the best of our
al.
descriptions
of
more
discovered
new
a recent
taxa
of
are
study
Coprotus
For
species.
were
an
that their
up-to-date
combined, shortly
(1973), Jeng
et
al.
(1993)
has been made
study
admitting
427).
Kimbrough
often than not
For instance,
coprophilous
on
two
Krug (1977),
Wang (1994)
to assess
quite obviously
of which
&
and
the
validity
difficult to delimit
Pezizales in
could
not
be
Italy
assigned
species.
new
species
knowledge
for
hereafter described we
taxa
delimitations
specific
most of
extended the limits of the genus to
(1980), Gene
referred
confirmed that the informal groups outlined by
still hold but that
(1972)
(1972) eighteen species
Bell &
comprehensive
which
with the
dealing
adding
Kimbrough
al„ 2000) deals with nine
previously
Before
No
al.
Van Brummelen (1998:
(1971),
Gibson &
et
et
extracted from Rhyparobius Boud.,
Kimbrough
Thind
(Kimbrough
see
Moravec
keyed.
(1978),
al.
some
of the genus,
(Korf, 1958). Subsequently Kimbrough (1966),
Kimbrough
hooked and
strictly
are not
described and
(Doveri
and
(1967)
multispored asci,
circumscription
to
Boud.
heterogeneous Ascophanus
Kimbrough
the
species
combines sufficient differential
are
to
Kimbrough
hard
produced
Coprotus.
to
pin
et
This
al.
down
a
shown
within the genus
(1972)
on
key (not
comparative approach
the
of limited
grounds
involved. Nevertheless, the material described hereunder
diagnostic
characters
to
be
distinguished
as
a
separate spe-
cies.
The
dung
collected
Coprotus species
were
are
in
placed
was
among the last
mounted in distilled
water
moist chambers for several
discomycetes
in which all
to
appear.
Freshly
measurements were
weeks
knowing
collected
made.
Histological
have been studied from freehand sections. Cotton blue in lactic acid (CB) has
tested for its effects
artifact. Freehand
with
1)
on
light microscopy
This
key
ripe
drawings
with
ascospores with non-elastic walls
were
a
and comments on the
made for the illustrations.
Leitz
microscope using
species
involved
will be
a
35
developing
de
mm
published
Olympus
in the
near
been
bubbles
were
camera.
future.
details
only
Bary
Photomicrographs
that
specimens
made
650
PERSOONIA
Vol.
17,
Part
4,
2001
De
Sloover:
Coprotus arduennensis,
arduennensis J.R. De Sloover, spec.
Coprotus
discus aurantius
Apothecia discoidea, sessilia, dispersa;
trudentes
Superficies
ascos.
pulum ectale
textura
valde
16
cylindracei
late
pm,
ad
elongatae praesertim
vel
alba
cellulis valde
globulosa,
marginales
glabra,
externa
nov.
vel
basim,
lutea.
200
usque
inferne
cylindraceae omniquoque erectae, comparate crassiusculae,
ad
diam., raro
apicem
In fimo fero
J.
Typus:
leviter
incrassatae
porcino (Sus scrofa)
De
L
Sloover; isotypus
in calluneto
crescens,
A
Sacrawé,
from the Ardenne
Latin,
high. Receptacle
mm
shaped, light
or
first
at
above the disk. Disk flat,
in
paraphyses varying greatly
250-400 pm thick. Cortical
isodiametric
pacted
sessile
dry,
then
smooth with
oblong
or
width below the middle,
(but often with only
blue in Melzer's
clavate
across,
herb.
3
to
up
globular
200 pm
cells
(150—)160(—185)
some
of the spores
its
causing
x
drying. Hymenium
near
uniseriate or
the base, of
closely
Habitat
tiny (up
On
—
wet Vaccinietum
herb. J. De
BELGIUM:
Sloover; isotype
wild boar
michaudii
dung in peaty
walls
hyaline
m),
on
deer
a
maximum
pm, rounded above,
furaceus Pers.,
Fig.
1.
birch-oak
simple
or
and
one
d.
paraphyses
paraphysis;
00C9, holotype;
with
with
branched below the
f.
b.
with
d & f: De Sloover
stercoreus
median
section
on
ascospores
99C111).
—
long,
pm
vacuoles.
large
585
Fond
(alt.
440
deer
on
m),
dung
in
a
on
&
et
dung in
wet
640
m),
al.,
Ascobolus
afforested
des Cavaliers
Br.) Boud.,
and
c.
lower flank
refringent guttules;
showing
de
markers:
Bary
raised
a
&
c
=
50
μm;
b
ascospores
CB.
=
360
fur-
De Sloover 00C114.
a-c,
100
μm;
e:
ectal
external
two asci
in
De
d-f
margin
and
excipulum with
e.
bubbles in
site,
(alt.
Ascobolus
through ripe apothecium:
flank;
wild boar
(alt.
through ripe apothecium: hymenium
the lower
Scale
on
deer
spruce
Tode, 20.VIII.2000, J.
section
m),
Holzwarche
Coprotus leucopocillum Kimbrough
granular content
with
long
(holotype of Coprotus arduennensis,
Ascozonus woolhopensis (B.
median
not
tip (10
to
(60-)80(-100)
and
Liege Province, Biillingen,
99C111; Biron,
elongated cells
ascus
guttules
Province, Oignies-en-Thierache, Trieu
Longitudinal
hymenium;
with bullate cells and
oil
J. De Sloover 00C9
Crouan and Thelebolus
a.
the upper cell
-667); Petit-Thiers, Grand
Sloover
wood
arduennensis.
(left)
elongated cells;
flanking
in
A. albidus
Coprotus
of bullate cells
excipulum
J. De
8-spored
the wall
rarely 16-spored),
heathland.
wet
grassland
the
appearance. Asci
rough bumpy
Luxemburg Province, Vielsalm, A Sacrawd (alt.
De Sloover 00C139. Namur
dung
plasmatic
De Sloover 00C106.
Molinia
near
rarely biseriate, ellipsoid, hyaline, smooth,
septate),
more
orange
L 998.171
Boud., 04.VII.1999,
15.X.2000,J.
pm)
heathland, 23.111.2000,
Molinietum, 15.VII.2000, J.
on
1
of wild boar in
dung
Specimens examined.
dung in
to
(textura
across
and downwards from
or
com-
into the lower and then the
cells 30-40 pm
(10—)12.8(—16)
fully developed
saucer-
rough margin
and
slightly protruding
with thin
long
(6.5—)6.8(—7.5) pm. Paraphyses
x
4-septate (exceptionally
or
numerous
De
mm across,
upper cell, strictly cylindrical and straight, rather thick, 6-9 pm wide from base
with
J.
the
spreading
gently tapering upwards
reagent. Ascospores
(12.5—) 14.5(—15.5)
pm),
pm
orange upon
250-300 pm thick
cells 10-45 pm
made of inflated
cylindrical
or
6-9
ramosae,
finally high
a narrow
bright orange, roughened by
globulosa
angularis),
and
expanding
subglobular-subangular
margin. Margin
raro
obconical base, 0.5-1.5
length, remaining deep
excipulum
globose
on an
cells
cylindrical
et
sphagnis (Vaccinietum).
cum
upper flank made of isodiametric,
to
10-
x
6.5-7.5
x
parvis aurantiisque guttulis,
septatae
Belgium, 23.111.2000, (holotypus:
subglobular,
whitish when
turning
orange
slightly raised
asci
12.5-15.5
country.
Apothecia discoid, superficial, scattered,
0.5-0.7
150-185
octospori
998.171-667).
from
Etymology:
Exci-
cellulae
marginem;
(10 pm) obtusaeque.
00C9, Vielsalm,
Sloover
Asci
Ascosporae
multis
cum
inferne
ad
ob pro-
prominens.
parum
praecipue
longae.
pm
leviter attenuati.
pm, ellipsoideae, uniseriales, hyalinae, levigatae. Paraphyses
diametro,scabriusculus
mm
Margo alba,
10-45 pm diam.
inflatis,
cylindrico-clavati,
0.5-1.5
pallide
1, 2
Figs.
—
651
Nov.
spec.
Sloover
=
10
μm.
652
PERSOONIA
Vol.
17,
Part
4,
2001
De
Criteria
tative:
commonly
the
form and
De
bubbles
Bary
media and
al.,
1972).
To
in lactic acid,
are
like Pulvinula
a
author
one
length
(Moravec, 1971 ; Ellis & Ellis,
1998).
ascospores
pm, but
is
Sizes of asci and
Presence
vs.
first order
a
colours
in the
being
a
to
in
paraphyses
of carotenoidbound
mostly
These
are set
the 26
a
linked to
quite
group with
as
out
or
amount
amounts
Coprotus
showing
a
rigidity
or
special
mount-
(Kimbrough
et
blue
cotton
of the spore walls
different
quite
size,
highly
are
variable in
such as
taxa
species
& P.
(H.
13.5-17.5 gm (Aas,
as
(J.
De Sloover
of
Pyroor
1983)
and
J. Mora15-18
as
means are not
99C111)
had smaller
with the type in all other aspects: that
taxon
Coprotus
Crouan)
difficult to handle where
are
slightly deviating
with
apparently
of which the delimitation
devoid of
99C111.
with
some
are
or to
same
in
thick, rarely
small refractive
branched
always
e.g. in
are
(1972)
orange
sign
shades
paraphyses.
on
paraphyses.
to seven
contents,
cell walls.
are
of the
c. median
(textura angularis);
asci with uniseriate
towards basis
golden yellow droplets (t);
a,
(p, q)
b: De
as
well
Obviously
the
species
listed
translucent to white
possess
most
above;
The
among
It should be noted that this
asci. Furthermore, the last nine
cells
the
brought together.
light yellowish
mainly
Here also
m-o.
al.
yellowish
is limited
paraphyses
of fruit-bodies from
optical section;
is
or
refringent droplets,
excipulum
hypothecium
yellow
characteristics
located in the
eight-spored
a, b. Habit
et
bright
be confused. In Table I the 26 known spe-
not to
drying.
to
Coprotus, particularly
aurora,
On the other hand,
pigments
the
of
species
yellow pigments. They usually
of d and e; d.
and
of
of them have ascospores with
any
arduennensis.
paraphyses straight
septate (u)
paraphyses.
sometimes when
in
proposed by Kimbrough
described for instance in C.
as
wall
has
taxa
new
Similar but duller shades have been detected
orange-yellow pigments
species
variable,
so
describing
cells and cell walls. It should be noted that the
excipulum
species sharing
most
(h, l)
data are
greatly. Quantitative
was
hymenium
droplets.
of carotenoids in the
seven
two
pigments
of colourare
of carotenoids
the location
ascospores
in the
origins
bound
apothecia except
u;
of
practice.
oil
hymenium,
that
so
lipid
completely
2.
these criteria
the spores, e.g. in
mounting
the thickness
Such data
agreed
noticed in the
excipulum
species listed;
small
greatest
Fig.
use
criterion to found groups within the genus. Yellow
lipids
different
small group of
are
the
should be considered essential when
common
granules
orange colour of the
cies
to
Pseudombrophila (van Brummelen, 1995)
reported
absence of carotenoid
qualitative
frequently
are
analysis
both in the ascospores and in the
are
by
spores
conditions in
diameteralso vary
apothecial
statistical
simple
but it is far from
as
quali-
yet clear.
not
and
importance given
another. For C. ochraceus
aberrant sample could perhaps indicate
and
quantitative
of their taxonomic features
mentioned. For C. arduennensis one of the collections
(9—) 10.5 (—13)
both
and ascospore form and size,
Krug, 1976).
ascus
to
is
clue
and
&
like spore and
from
Coprotus
one
same
(pers. obs.)
for instance, ascospore
vec
as
Coprotiella (Jeng
or
as
created by
in the
produced
are
ascus
the relative
on
induced in
regarded
Coprotus
extent,
653
Nov.
spec.
different.
quite
be viewed
only
must
sample
be
mind those artifacts
our
Quantitative criteria,
one
Depending
may
have been
Thelebolaceae like
from
excipulum
consistently
are
they
(Baral, 1992): they
nemataceae
delimit species of
to
content.
classification
resulting
ing
used
Coprotus arduennensis,
colour and size,
apothecia
paraphyses
Sloover:
to
e.
multispored species.
section of an
ectal
irregularly
apothecium
excipular cells;
biseriate
or clavate at apex
Sloover 00C114;
f-l.
spores; p-
(r), exceptionally
c-u:
De
Sloover
654
PERSOONIA
Concerning pigmentation Coprotus
six
C.
with
species
In
the
Coprotus
like C. disculus
1977)
C.
or
and ectal
ness
(Thind
in C.
et
layers
up
to
a
cells
in C. ochraceus where
the
excipulum along
marginatus.
margins
Table I. Location
of carotenoids in
in
++
=
al.,
large
is absent
though
they
1972).
may
Ectal
clearly elongated
are
cells
or
in
walls of
Kimbr., Luck-Allen
Phillips) Kimbr.,
Phillips)
glaucellus; (Rehm) Kimbr.
glaucellus
C.
C.
marginatusKimbr.,
most
C.
granuliformis (P.
&
& H.
Crouan) Kimbr.
Luck-Allen
dhofarensis Gené,
C.
Luck-Allen
leucopocillum Kimbr., Luck-Allen
C.
C.
duplus Kimbr.,
C.
sexdecimsporus; (P.
sexdecimsporus
C. breviascus
C.
C. vicinus:
C.
C.
C.
& H.
H.
1 (H.
(H.
C. ochraceus
C.
(H.
(H.
Kimbr.,
Crouan)
Luck-Allen
C.
C.
sphaerosporus
only
species
of
(E. Marchai)
(E.
(Boud.)
C.
C.
sarangpurensis K.S.
C. uncinatus Y.Z.
C.
C.
C.
1) Aas,
Aas, 1983;
1983;2)
2)
1)
=
in small
top-
amount,
oil
oil
with
gut-
Ascospores
Excipulum
slightly yel-
and/or cells
cells
walls
walls
yellow
&
& Cain
5
5
(+)
(+)
(+)
(+)
(+)
(+)
(+)
+
(+)
(+)
+
(+)
(+)
++
++
(+)
(+)
(+)
5
5
& Cain
&
& Cain
5
5
5
5
5
& Cain
Cain
++
+
5
++
3
& Cain
5
5
Kimbr. 5
5
5
5
S.C. Kaushal
& Kimbr.
7
2
9
9
J.M. Moravec
Bell &
Bell
Coprotus: (+)
100 pm
most
++
Thind
Wang
subcylindrosporus
subcylindrosporus
to
++
++
& Cain
Kimbr.
Kimbr.
Kimbr.
C. trichosurussA.E.
A.E. Bell
up
of its kind, where
+
Luck-Allen
Luck-Allen
(Heimerl)
rhyparobioides (Heimerl)
C. albidus
C.
pm,
+
'• 5
Moravec '•
Gibson &
& Kimbr.
(Fuckel) Kimbr.,
C. winteri
C.
ochraceus.
cells of the ectal
species reaching
one
5
4
4
& Krug
baeosporus Jeng
baeosporus
Jeng&Krug
C.
C.
goes from 50-
in C.
1978)],
8
Luck-Allen
Luck-Allen
C.
C.
C. niveus
C.
al.,
two
layers
(+)
C. arduennenesis
C.
arduennenesis ; J.R. De Sloover
C. luteus
C.
six
or
lowish
lowish
5
5
& Cain
Crouan)
Crouan) Kimbr.,
& P.
five
to
slightly elongated [8-30
et
e.g.
thick-
(+)
Kimbr.
Crouan) Kimbr.
(Velen.) Kimbr.,
Kimbr.,
& P.
reported,
of 25, from
out
thickness
5
& Cain
& Cain
Luck-Allen
(Boud.) Kimbr.,
aurora
Krug,
medullary
excipulum
reported
5
5
& Cain
ElShafie &
& Guarro
Luck-Allen
Kimbr.,
species
&
(+)
& Cain
Cain
C.
C. disculus
C.
as
5
Luck-Allen
Kimbr.,
or
(Thind
is the
dhofarensis
selected
in
true
amount.
C.
C. dextrinoideus
C.
species
excipulum
tules
& W.
particularly
least indistinct as
seven
isodiametric
yellow
(Cooke
(Cooke
at
or
for
reach 55 |om
vicinus,
baeosporus (Jeng
well differentiated, the
Paraphyses
C. lacleus
lacteus
this is
could reach 425 pm
usually
are
C.
it is rather well differentiatedwith
only reported
et
In this respect C.
+
present,
Even
the group of the other
C. ochraceus.
few others like C.
1978)
medullary excipulum
medullary
a
to
i.e. C. breviascus,
maximum of four in C. vicinus and up
sexdecimsporus (Kimbrough
except
in
2001
related
seems
poorly developed:
al.,
are
4,
paraphyses,
particularly
hypothecium
al., 1978).
et
baeosporus,
While basal and
=
a rule
as
the
layers. Moreover,
95 pm, while the
in C.
is
al., 1978), yet
et
and the number of its
layers
in C.
excipulum
(Thind
and
baeosporus,
sarangpurensis (Thind
in C. ochraceus
in the
Part
17,
arduennensis
golden yellow droplets
C. luteus, C.
aurora,
Vol.
6
Kimbrough, 1973; 3)
Gibson &
&
et al., 1972; 6) Moravec, 1971; 7)
5) Kimbrough et
4) Jeng
Jeng & Krug, 1976, 1977;
Kimbrough, 1980; 4)
Thind et
Gene
al., 1978;
1978; 8)
8) Gené
et
al., 1993; 9)
9) Wang,
Wang,
1994
De
cells
on
cells
exceeding
oped
than the
and the
size
Coprotus arduennensis,
the upper flank of the ectal
the surface of the
one
of those in
pulum
with
as
excipulum elongate upwards forming
Here the
hymenium.
described for C.
marginatus,
largest
that
are
reported
thus far,
enlarged
in C. arduennensis. It appears that C.
a
more
devel-
nearby paraphyses
have
dhofarensis paraphyses
where the
species
200
|im)
of about the
are
marginatus,
marginal
cells
margin
C.
larger
are
same
than the
also
order
at
largest
considerably
and C. arduen-
dhofarensis
particularly
are
a
well-differentiatedexci-
a
are even
The cells of the raised
marginatus.
group of three
of long
fringe
known in the genus, in C. ochraceus. The sizes of cells
in C. arduennensis (exceeding
in C.
a
is far
structure
but its cells look like the
arduennensis shows
Coprotus. Coprotus
ones
excipular margin
nensis form
marginal
isodiametric basal cells up to 45 pm. which
globose
the
655
Nov.
spec.
described here for C. arduennensis, while C.
ones
typical
of size
the
Sloover:
well
developed
in different ways.
Thus C.
Table I
arduennensis shares the
having paraphyses
as
C. ochraceus,
to
C.
aurora.
the smaller ascospores and
sufficient
to
and
especially
baeosporus. Moreover,
The wide
species
most
the
indicated in
richly pigmented
pronounced yellow
particularly
the
straight cylindrical paraphyses,
large
swollen
excipular
C. arduennensis from these other
distinguish
the
is another feature that C. arduennensis has in
apothecium
with C. ochraceus and C.
characteristics with the six
same
yellow guttules
C. luteus and C.
aurora,
orange colour of the
with
cells
are
the
common
larger asci,
consistent, and
species.
ACKNOWLEDGEMENTS
thanks
My
branch)
A.
due to
are
for its invaluable
Fraiture
literature
Dr. J.
Brummelen
van
advice
and for
(Nationaal
checking
(Jardin Botanique National, Meise) helped
and Paul
Pirot corrected the Latin
Herbarium
and
quickly
so
in
Nederland,
accurately
many
ways
Universiteit Leiden
the submitted material.
when
searching
Mr.
for the relevant
diagnosis.
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