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Karstenia 47: 29–36, 2007
Additions to the Swedish myxomycete biota
UNO H. ELIASSON and ELISABETH GILERT
ELIASSON, U. H. & GILERT, E. 2007: Additions to the Swedish myxomycete biota.
– Karstenia 47: 29–36. Helsinki. ISSN 0453-3402.
The current inventory of myxomycetes within the frames of the Swedish Taxonomy
Initiative project continues to yield records of species not previously known to Sweden.
A selection of twenty species here formally reported for the fi rst time gives a total of 210
species so far known from Sweden. Among notable records are Echinostelium bisporum
and E. lunatum, the two smallest species of myxomycetes known, fruitbodies of the fi rst
species rarely exceeding 25 m in height. Both species are impossible to detect using
standard methods for myxomycetes and are probably far more common than the few
widely scattered records in the world so far would suggest. Paradiacheopsis longipes
is reported for the third time, previously known only from the Netherlands and China.
Additional records are awaiting publication, but the delimitation and circumscription of
several taxa require further study. Continued fi eld-work supplemented with so-called
moist chamber cultures will further increase the number of species known to occur in
Sweden. Distribution patterns of myxomycetes are discussed briefl y.
Key words: Swedish myxomycetes, Swedish Taxonomy Initiative, Myxomycete dis-
tribution, Arcyria, Echinostelium bisporum, Echinostelium lunatum, Paradiacheopsis
longipes.
Uno H. Eliasson, Department of Plant and Environmental Sciences, Göteborg Univer-
sity, Box 461, SE 405 30 Göteborg, Sweden / uno.eliasson@dpes.gu.se
Elisabeth Gilert, same address / elisabeth.gilert@dpes.gu.se
Introduction
Records of myxomycetes in Sweden date back
to long before the true nature of these organ-
isms was known. Although Linnaeus paid little
attention to lower cryptogams there are three
specifi c Linnaean epithets still in current use.
The epithets are all applied to common and con-
spicuous species, namely Lycogala epidendrum
(L.) Fries, Arcyria denudata (L.) Wettst. and
Diderma radiatum (L.) Morgan. Elias Fries de-
scribed a considerable number of myxomycetes
in several important mycological papers during
the fi rst half of the nineteenth century. About
fi fteen of these species are accepted as distinct
in current myxomycete literature, as are the two
genera Lindbladia and Perichaena. The fi rst
comprehensive summary of Swedish myxomyc-
etes was published by Robert Fries (1912), who
listed 132 species as confi rmed from Sweden.
Santesson (1964) added many new records and
summarized 165 species from Sweden. Dur-
ing the past three or four decades an additional
twenty-fi ve species have been reported in vari-
ous publications and are cited below. Changes
in circumscription and delimitation of taxa have
affected several names in earlier published spe-
cies lists.
Several additional species have been found
during the ongoing inventory within the frames
of the Swedish Taxonomy Initiative project. A
selection of twenty species not previously re-
ported from Sweden is presented in this paper,
making a total number of 210 species herewith
formally reported. Additional records are await-
ing publication but several taxa require extensive
taxonomic discussions. Continued fi eld-work
supplemented by moist chamber cultures will
further increase the number of species known to
occur in Sweden.
30 KARSTENIA 47 (2007)
ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
Summary of species reported from Sweden during the past few decades
The following twenty-fi ve species have been reported from Sweden after the publication of Santes-
son’s (1964) list. Many of these species have been recorded again, some species several times, after
their fi rst published record.
Amaurochaete comata G. Lister & Brândz (Eliasson 1977)
A. trechispora T. Macbr. & G. W. Martin (Eliasson 2000)
Arcyria affi nis Rostaf. (Schinner 1983)
A. fuegiana Aramb. (Nannenga-Bremekamp & Schinner 1986)
A. rubrobrunnea Nann.-Bremek. & Schinner (Nannenga-Bremekamp & Schinner 1986)
A. verrucosituba Nann.-Bremek. & Schinner (Nannenga-Bremekamp & Schinner 1986)
Badhamia apiculospora (Härk.) Eliasson & N. Lundq. (Eliasson & Lundqvist 1979)
Collaria arcyrionema (Rostaf.) Nann.-Bremek. ex Lado (Eliasson 1975 as Lamproderma a.)
Colloderma oculatum (C. Lippert) G. Lister (Eliasson & Sunhede 1972)
Diderma asteroides (Lister & G. Lister) G. Lister (Eliasson 1975)
D. cingulatum Nann.-Bremek. (Eliasson 1975)
D. simplex (J. Schröt.) G. Lister (Schinner 1983)
Didymium anellus Morgan (Eliasson 1975)
Fuligo cinerea (Schwein.) Morgan (Schinner 1983)
Hemitrichia calyculata (Speg.) M. L. Farr (Eliasson 1975)
Licea belmontiana Nann.-Bremek. (Eliasson & Lundqvist 1979)
Perichaena depressa Lib. (Eliasson 1975)
Physarum fl avidum (Peck) Peck (Schinner 1983)
P. nudum T. Macbr. (Schinner 1983)
P. robustum (Lister) Nann.-Bremek. (Schinner 1983)
P. serpula Morgan (Eliasson & Strid 1976)
Stemonaria pilosa Nann.-Bremek. (Nannenga-Bremekamp et al. 1984)
Stemonitis virginiensis Rex (Schinner 1983)
Symphytocarpus amaurochaetoides Nann.-Bremek. (Schinner 1983)
Trichia erecta Rex (Eliasson 1975).
The four Arcyria species listed had all been col-
lected in the province of Lappland north of the
Arctic circle. The records of Arcyria rubrobrun-
nea and A. verrucosituba are the type collections,
so far the sole records in the world. Arcyria fue-
giana represents the second record in the world,
the species was previously known only from the
type collection from Argentina. All four Arcyria
species listed belong to the red-spored group of
species several of which offer taxonomical dif-
fi culties.
Selected species new to Sweden
The following list is a selection of species not
previously reported from Sweden. Records made
from moist chamber cultures have been marked
by “mc”. Dates given for such records are dates
when the substratum was collected. For species
represented by numerous collections only repre-
sentative specimens have been cited.
Arcyria globosa Schwein.
Västergötland: Töllsjö, on bark from living
Quercus, mc, Aug 1989 Eliasson 3664 (GB);
Mölndal, on bark from living Quercus, mc, May
1996 Eliasson 4952 (GB).
31
KARSTENIA 47 (2007) ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
Arcyria stipata (Schwein.) Lister
Närke: Hackvad, on decaying stump of Populus,
Oct 1998 Nilsson (S).
Badhamia affi nis Rostaf.
Västergötland: Göteborg, Kärralund, Aug 1948
Nathorst-Windahl (GB); Istrum, Sept 2003
Lundqvist (S). Uppland: Balingsta, Oct 1987
Lundqvist 17068 (S).
Comatricha ellae Härk.
Västergötland: Alingsås, Östad Säteri, on dead
wood of Quercus, mc, Apr 2007 Eliasson & Sjö-
gren 6675 (GB).
Comatricha longipila Nann.-Bremek.
Halland: Långavaka, on wood and bark of
Quercus, mc, July 2004 Gilert & Eliasson 6273
(GB).
Comatricha tenerrima (M. A. Curtis) G. Lister
Halland: Tjolöholm, on grass, mc, Sept 2005
Eliasson & Gilert 6430 (GB).
Cribraria macrospora Nowotny & H. Neubert
Västergötland: Vårgårda, Klovnasten, on decay-
ing wood, Oct 1970 Eliasson 2744 (GB); Töllsjö,
on decaying wood of Picea, Oct 1992 Eliasson
4669 (GB).
This relatively recently described species
(Nowotny & Neubert 1993) is closely related
to C. macrocarpa and C. argillacea in habit and
ecology and grows preferably on strongly de-
cayed wood and bark of Picea, where fruitings
up to 0.25 m2 in extent have been seen.
Cribraria oregana H. C. Gilbert
Halland: Särö Västerskog, on decaying wood of
Pinus, mc, May 2003 Eliasson 6034 (GB).
Diderma fl oriforme (Bull.) Pers.
Halland: Tjolöholm, at ground level on stump
of Quercus, Sept 2005 Eliasson & Gilert 6399
(GB). Västergötland: Göteborg, on decaying
wood of Quercus, Oct 1943 Nathorst-Windahl
(GB); Alingsås, on branch of Quercus on the
ground, Dec 2006 Sjögren (GB).
Didymium bahiense Gottsb.
Västergötland: Alingsås, Nolhaga, on bark from
living Acer, mc, Aug 2006 Eliasson & Gilert
6610 (GB).
Echinostelium bisporum (L. S. Olive & Stoian.)
K. D. Whitney & L. S. Olive – Figs. 5–7
Västergötland: Mölndal, Stensjön, on dead grass,
mc, Nov 2005 Gilert & Eliasson 6510 (GB);
Töllsjö, on bark from living Acer, mc, Mar 2006
Eliasson & Gilert 6520 (GB).
With fruitbodies rarely exceeding 25 m in
height and producing only two spores, this is the
smallest of all myxomycetes known. The spores
are shed together in characteristic dyads (Fig. 7).
It was originally described as a protostelid of the
genus Cavostelium (Olive & Stoianovitch 1966)
but was later transferred to Echinostelium based
on convincing similarities in ultrastructure and
development (Whitney et al. 1982).
Echinostelium bisporum is too small to be
found using standard techniques for myxomyc-
etes and, therefore, has mainly been recorded in
studies on protostelids. It is too small to be ob-
served in moist chamber cultures even under a
high magnifi cation stereo microscope. Although
probably of worldwide distribution it is believed
to be more common in warmer regions. Original-
ly described on material from Malaysia, Sri Lan-
ka and Florida it has been reported from northern
India (Shadwick & Stephenson 2004) and been
demonstrated to be at least locally abundant in
Central America (Moore & Stephenson 2003). It
also seems to be a common species in Hawaii,
occurring on various kinds of substrata where
protostelids grow (Spiegel, pers. comm.). In Eu-
rope it has previously been verifi ed from Ger-
many (Tesmer et al. 2005).
Echinostelium brooksii K. D. Whitney – Figs.
1, 2
Halland: Särö Västerskog, on bark from living
Quercus, mc, May 2003 Eliasson 6039 (GB).
Västergötland: Göteborg, on bark from living
Robinia, mc, Nov 2003 Eliasson 6071, 6072
(GB); same locality and substratum, mc, Jan
2005 Eliasson 6262 (GB); Alingsås, Mjörn, on
bark from dead Alnus, mc, Apr 2004 Eliasson &
Gilert 6134 (GB); Alingsås, Nolhaga, on bark
from living Acer, mc, Aug 2006 Eliasson & Gil-
ert 6610 (GB).
Echinostelium colliculosum K. D. Whitney &
H. W. Keller
Halland: Särö Västerskog, on bark and bryo-
phytes on bark (together with E. lunatum and
E. minutum), mc, May 2003 Eliasson & Gilert
32 KARSTENIA 47 (2007)
ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
Figs. 1, 2. Echinostelium brooksii (Eliasson 6072). – 1: Fruitbody. Bar: 20 µm. – 2: Upper part of fruitbody further en-
larged, showing spores, strongly pigmented globose columella and peridial collar below columella. Bar: 10 µm. – Figs.
3, 4. Echinostelium lunatum. – 3: Fruitbody with spores and crescent-shaped columella (Eliasson & Gilert 6095b). Bar:
10 µm. – 4: Emptied fruitbody with columella and peridial collar below columella (Eliasson & Gilert 6519). Spores
are still attached to one another by disc-like raised wall thickenings. Bar: 10 µm. – Figs. 5–7. Echinostelium bisporum,
specimens dyed in cotton-blue (Gilert & Eliasson 6510). – 5: Two fruitbodies with spores shed. The columella is fl at-
tened and the stalk has rhizoid-like extensions at base. Bar: 10 µm. – 6: Part of preceding fi gure further enlarged. Bar:
5 µm. – 7: Same picture as in Fig. 6 but focus set at the underlying spore dyad. Bar: 5 µm.
33
KARSTENIA 47 (2007) ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
6095a (GB); Tjolöholm, on bark, mc, Sept 2005
Eliasson & Gilert 6418 (GB).
Echinostelium fragile Nann.-Bremek.
Västergötland: Mölndal, Sisjön, on bark from li-
ving Alnus, mc, Feb 2004 Eliasson & Gilert 6096
(GB); Alingsås, Mjörn, on bark from dead Alnus,
mc, Apr 2004 Eliasson & Gilert 6133 (GB); Gö-
teborg, Botanical Garden, on cone of Larix on the
ground, mc, Mar 2005 Eliasson & Gilert 6310
(GB).
Echinostelium lunatum L. S. Olive & Stoian.
– Figs. 3, 4
Halland: Särö Västerskog, on bark and bryo-
phytes on bark (together with E. colliculosum
and E. minutum), mc, May 2003 Eliasson & Gil-
ert 6095b (GB). Västergötland: Töllsjö, on bark
and bryophytes on bark from living Acer, mc,
Mar 2006 Eliasson & Gilert 6519 (GB).
With fruitbodies 20–50 m in height, this is
the second smallest of all myxomycetes known
and normally not possible to observe even under
a strongly magnifying stereo microscope. Long
known only from Puerto Rico and the United
States this is the fi rst verifi ed record from Europe.
The species is likely to be far more common in
the world than the few records would suggest,
which probably holds for several species within
the same genus.
Licea biforis Morgan
Västergötland: Göteborg, Botanical Garden,
on dead herbs, mc, Mar 2005 Gilert & Elias-
son 6357 (GB); Mölndal, Gunnebo, on dead
grass, mc, Nov 2005 Gilert & Eliasson 6477
(GB); Alingsås, Östad Säteri, on dead stems of
Chamaenerium, mc, Apr 2007 Eliasson & Sjö-
gren 6676 (GB).
Paradiacheopsis longipes Hooff & Nann.-
Bremek. – Fig. 8.
Lappland: Lycksele, on twigs protruding above
snow layer, mc, Apr 2005 Eliasson & Forsberg
6359 (GB).
An easily overlooked species due to its small
size, but very well defi ned through an excellent
combination of characteristics: sporothecae 0.1
– 0.2 mm in diam. on long fi liform stalks up to
ten times the diameter of the sporotheca, stalks
enclosed in a hyaline sheath, spores with widely
spaced but very distinct warts.
Apart from the type collection from the Neth-
erlands (Hooff & Nannenga-Bremekamp 1996)
this species has also been reported from China
(Ukkola et al. 2001).
Paradiacheopsis solitaria (Nann.-Bremek.)
Nann.-Bremek.
Halland: Särö Västerskog, on mosses and lichens
on bark of living Malus, mc, Aug 2006 Eliasson
6583 (GB); Near Falkenberg, on bark, mc, July
2004 Eliasson & Gilert 6424 (GB). Västergöt-
land: Mölndal, Lunnagården, on bark from liv-
ing Alnus, mc, Feb 2004 Eliasson 6089 (GB);
Jonsered, on bark from old Tilia, mc, Sept 2005
Eliasson & Gilert 6422 (GB); Överlida, on bark
of living old Juniperus, Aug 2006 Eliasson 6587
(GB).
Apparently an overlooked but probably com-
mon species so far recorded from south-western
Sweden by numerous specimens (fi eld collec-
Fig. 8. Paradiacheopsis longipes (Eliasson & Forsberg
6359). The diameter of the sporotheca is ca 0.2 mm.
34 KARSTENIA 47 (2007)
ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
tions as well as moist-chamber cultures) on bark
(and lichens and bryophytes on bark) of various
living trees (Juniperus, Alnus, Malus, Populus,
Tilia, Quercus).
Perichaena chrysosperma (Curr.) Lister
Halland: Särö Västerskog, on bark of living
Sambucus nigra, mc, June 2003 Eliasson 6049
(GB). Västergötland: Göteborg, Askim, on bark
of living Malus, mc, Aug 2003 Eliasson 6059
(GB); Göteborg, Botanical Garden, on bark of
living Ulmus, mc, Oct 2003 Eliasson 6070 (GB);
Töllsjö, on bark of living Fraxinus, mc, Eliasson
6561 (GB); Alingsås, Nolhaga, on bark of living
Acer, mc, Oct 2006 Eliasson 6615 (GB).
Apparently a common species at least in
south-western Sweden. It has been obtained nu-
merous times in moist chamber cultures on bark
from a large variety of deciduous trees. As cur-
rently accepted this is a very variable species.
The material cited is rather uniform, consist-
ing of scattered or solitary, globose or subglo-
bose fructifi cations on bark or mosses on bark
of living trees. The capillitium is ornamented
with spinous processes rarely exceeding 2 m in
length. These specimens are quite different from
specimens from tropical and subtropical areas
where fructifi cations are typically plasmodio-
carpous, commonly arcuate or ring-shaped, and
the capillitium has spines up to 6 m long. The
different forms were exhaustively discussed by
Keller (1971) and incorporated within the same
species. The differences are nowadays attributed
to ecotypic variation.
Physarum squamosum L. Flatau & P. Schirmer
Västergötland: Mölndal, vicinity of Lake Sisjön,
on bark of fallen decaying stem of Populus tre-
mula, Oct 2001 Eliasson 5777, 5778b (GB).
This species is characterized by globose spo-
rothecae 0.5 – 0.7 mm in diameter on thick black
stalks and the peridium covered with prominent
whitish calcium tubercles which are brownish in
the centre. It was described from Germany (Fla-
tau & Schirmer 2004) on dead twigs of Sambucus
and bark of living Populus. Two specimens (in
GB) from U.S.A., (Kentucky, on dead branches
of Juniperus virginiana, T. E. Brooks 2806; Iowa,
on bark of living Fraxinus americana, H. W. Kel-
ler 1020), bear an unpublished specifi c epithet,
“tuberculatum”, obviously referring to the prom-
inent ornamentation of the peridium.
Discussion
The publication of the world monograph of
myxomycetes by Martin and Alexopoulos in 1969
was a great stimulus to further studies of these
organisms, which has resulted in an increasing
number of publications and descriptions of new
species during the past few decades. This trend is
likely to continue although better understanding
of the biological system of these organisms and
the infl uence of environmental conditions on the
structure of the fruitbodies will probably warrant
modifi ed circumscriptions of several species and
reduce several published names to synonyms.
The general adoption of the moist-chamber
technique, where a piece of a suitable substratum
is placed under favourable ambient conditions
in a petri dish, has strongly contributed to the
number of new species described during the past
few decades. This technique allows species with
minute and fragile fruitbodies to develop more or
less protected from various arthropods and oth-
er disturbing factors, and has demonstrated the
presence of a number of species too small to be
observed in the fi eld. As a consequence, propor-
tionally more species have been described during
recent decades in genera with minute fruitbodies,
such as Licea and Echinostelium, than in genera
with larger fruitbodies. In Martin and Alexopou-
los (1969) four species of Echinostelium were
accepted (apart from E. roseum which was later
proved not to belong here), whereas Lado (2001)
lists fourteen species. Additional species are like-
ly to be found. This paper reports fi ve species of
Echinostelium new to Sweden, all found in moist
chamber cultures. At least three additional spe-
cies of this genus are likely to be found here.
It is diffi cult to get a true picture of the distri-
bution of organisms that are so small that they
cannot be seen with the naked eye. There is a
general tendency for minute organisms to be
more widespread the smaller they are. Many
species represented by a single or a few records
are probably far commoner than the number of
records would suggest. This certainly holds for
many small myxomycetes where there are sev-
eral examples where a species long known from
only the type collection or from only one part of
the world may unexpectedly be found in a new
locality very distant from previous records, not
rarely on a new continent. There is no reason to
believe that, for example, Paradiacheopsis lon-
35
KARSTENIA 47 (2007) ELIASSON & GILERT: SWEDISH MYXOMYCETE BIOTA
gipes would be as rare as the widely separate
records from Europe and China would hint. It
simply is too small to be easily seen, even with
a hand lens, and is likely to be found, perhaps
worldwide, where conditions are suitable. Echi-
nostelium bisporum and E. lunatum are even bet-
ter examples where size itself excludes detection.
For recording such minute fruitbodies methods
used for protostelids have to be applied (Moore
& Spiegel 1995, Tesmer et al. 2005).
One of the most interesting Swedish records
reported during the past few decades is that of
Arcyria fuegiana from north of the Arctic circle
(Nannenga-Bremekamp & Schinner 1986). As
already mentioned this is the second record in the
world, the species being known previously only
from the type collection from Tierra del Fuego in
southern Argentina (Arambarri 1972). We have
not seen any material of this species. Neverthe-
less, the records give a clear hint not only that
further fi eld-work is needed in the northern al-
pine areas of Sweden, but also that the distribu-
tion patterns and morphological variation of the
red-spored Arcyria species are incompletely un-
derstood. This is further corroborated by the two
likewise red-spored species A. rubrobrunnea and
A. verrucosituba, both known only from their re-
spective type collections from north of the Arctic
circle.
As already mentioned, this paper brings the
total number of myxomycete species known to
Sweden to 210 but several records remain to be
published. Although an inventory will never be
complete the rising curve of number of species in
Sweden is likely to level off perhaps somewhere
around 230. The northern alpine areas are still
insuffi ciently studied. A few so-called nivicolous
myxomycetes, species developing fruitbodies
under or close to melting snow, have been re-
corded, but considering the large number of spe-
cies of this ecological group known from other
parts of Europe, Sweden´s alpine region can be
expected to harbour several nivicolous species
of genera such as Diderma, Lamproderma and
Lepidoderma yet to be recorded.
Acknowledgements: We are grateful to Dr. Grazina Ada-
monyte and Mr. David Mitchell for valuable and construc-
tive comments on an earlier version of the manuscript.
Their points of view signifi cantly improved the paper.
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Karstenia 47: 37–48, 2007
Fleshy fungi of the genera Armillaria, Pleurotus,
and Grifola as habitats of Coleoptera
DMITRY S. SCHIGEL
SCHIGEL, D. S. 2007: Fleshy fungi of the genera Armillaria, Pleurotus, and Grifola as
habitats of Coleoptera. – Karstenia 47: 37–48. Helsinki. ISSN 0453-3402.
The paper deals with adult beetles attracted by wood-rotting agarics of the genera Armil-
laria and Pleurotus, plus those collected on the oak-decaying polypore Grifola frondosa.
The interrelations of 14 species (110 samples) of fungi and 78 species (1691 individu-
als) of beetles are discussed. More close relationships, such as fungivory of larvae of
Coleoptera are also treated. In southern Finland larvae of Cychramus (in Armillaria)
and Triplax (in Pleurotus) are able to utilize fl eshy wood-rotting fungi. The majority of
beetles were recorded as adults, including 52 (67%) species of Staphylinidae.
Key words: Basidiomycota, Coleoptera, decay, ecology, Finland, fungivory.
Dmitry S. Schigel, Botanical Museum, Finnish Museum of Natural History, P.O. Box 7,
FI–00014 University of Helsinki, Finland. E-mail: dmitry.shchigel@helsinki.fi
1. Introduction
Woody plants make most of the forest biomass,
and, alive or dead, become eventually utilized by
various organisms, including insects and fungi.
Polypores are the main group of wood-decom-
posing fungi especially at early stages of wood
decay. Many agarics continue the decay process
at later stages. Polypores usually have sturdy
and long-lasting fruit bodies, and even the soft-
est species last in nature longer than water-rich
agarics. This has consequences on insect fauna:
beetles are diverse on polypores, while diptera
larvae mostly utilize mushrooms.
My fi eld work in the autumn 2006 coincided
with the peak fruiting time of Armillaria and
ample material of beetles could be harvested.
The present paper focuses on beetles linked with
Armillaria borealis Marxm. & Korhonen. My
goal was to explore the similarity of beetle com-
munities on wood-rotting agarics in different
conditions, in order to study the dependence of
beetles on host species, and to review their life
strategies and adaptations in comparison to poly-
pores of southern Finland.
Previously six consistency classes of fungal
fruit bodies could be separated among polypores
(Schigel et al. 2004), and the role of substrate rar-
ity (Schigel & Toresson 2005), and the seasonal
dynamics of the attracted beetles (Schigel et al.
2006) were studied. The treatment would be in-
complete without a report on Coleoptera found
on wood-rotting fl eshy fungi.
2. Materials and methods
Armillaria borealis and associated Coleoptera
were collected during a polypore inventory in the
Kolovesi National Park, southern Finland (Fig.
1). The study was organized by the Finnish For-
est and Park Service (Metsähallitus) in Septem-
ber 2006. Supplementary materials from other
agaric genera derive from the collecting trips to
southern Finland in 2001–2006. These include
the Åland Islands, Helsinki, the Ruissalo oak
forest in Turku, and several locations in Etelä-
Häme (Table 1). All material was collected in the
southern boreal zone (sensu Ahti et al. 1968) of
Finland.
The Kolovesi National Park lies in the for-
ested archipelago of Saimaa, one of the largest
38 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
lakes in Finland. The landscape is strongly infl u-
enced by the Ice Age and is dominated by cliff
walls facing the lake, rugged terrain, and forest-
ed depressions along the lakesides, favoured by
beavers. The woody vegetation is dominated by
Norway spruce (Picea abies), birch species (Be-
tula spp.), aspen (Populus tremula) and Scots
pine (Pinus sylvestris). Rock outcrops are cov-
ered by pine woodlands interrupted with small
spruce swamps, and to a minor extent herb-rich
forests. The high volume of dead wood increases
the conservation value of these fairly old forests,
more than a half of them being over one hundred
years of age.
All fungal specimens were collected for
identifi cation, and dried in mushroom dryers
with ventilated air at +40–45° C. In diffi cult
cases provisional names were later approved
by microscopic study of sections mounted in
Cotton Blue or Melzer’s reagent at ×1250 mag-
nifi cation and phase contrast illumination. Fun-
gal collections are preserved in the Herbarium
of the Botanical Museum, Finnish Museum of
Natural History, University of Helsinki (H);
their nomenclature follows Salo et al. (2005)
and Niemelä (2005).
Clusters of fruit bodies (Fig. 2) were studied
for beetles. Adult beetles were collected and
counted separately from larvae. All adult bee-
tles and samples of larvae were preserved in
70% alcohol for identifi cation. Fungi which con-
tained additional beetle larvae were removed and
placed for rearing in plastic 0.5-litre containers.
Each container was quarter-fi lled with gardening
peat to serve for pupation media. Perforated lids
allowed slow drying of the fruit bodies. Rearing
chambers were not moistened during the rear-
ing. Collections made in summer–autumn were
Fig. 1. Clusters of Armillaria borealis fruit bodies on
birch, Kolovesi National Park, Pohjois-Savo. Photo T.
Niemelä, 2006.
Table 1. Finnish localities mentioned in this paper.
Code Locality Commune Province Coordinates
Å Åland Islands several Ahvenanmaa N 60º2´ E 19º58´
H Helsinki Helsinki Uusimaa N 60º12´ E 24º57´
Kt Kotinen Virgin Forest Lammi Etelä-Häme N 61º14´ E 25º3´
Kv Kolovesi National Park Enonkoski Pohjois-Savo N 62º15´ E 28º49´
L Lammi Biological Station Lammi Etelä-Häme N 61º3´ E 25º2´
R Ruissalo oak forest Turku Varsinais-Suomi N 60º25´ E 22º10´
S Susimäki Virgin Forest Ruovesi Etelä-Häme N 61º51´ E 24º14´
kept for 2–3 months in outdoor temperature,
then exposed to room temperature for further
two months, and after that checked for insects.
Beetles are preserved in the Finnish Museum of
Natural History, University of Helsinki; single
individuals have been given to identifi ers (see
Acknowledgements).
Beetle names accord Silfverberg (2004).
Authors of the Latin names are omitted from the
text, but are shown in Table 2 for fungi and Table
3 for beetles. Terminology follows the previous
papers in the series (Schigel et al. 2004, 2006,
Schigel & Toresson 2005).
39
KARSTENIA 47 (2007) SCHIGEL: FLESHY FUNGI AND COLEOPTERA
3. Results
76 forest compartments (metsäkuviot) were sur-
veyed for polypores, and yielded 98 species of
polypores and 48 species of other fungi (Niemelä
& Kinnunen 2006). In the present paper the in-
terrelations of 14 species (110 samples) of fungi
and 78 species (1691 individuals) of beetles are
discussed. It comes as no surprise that as many
as 52 (67%) beetle species in the report belong
to the Staphylinidae, as they usually outnumber
the other beetles in gilled fungi (Tables 2, 3). All
beetles were sampled as adults, and for Cychra-
mus (Figs. 3, 4) and Triplax also larvae were re-
corded. Four kinds of fl eshy wood-rotting fungi
were sampled for Coleoptera:
(1) Among the three species of long-stiped
agarics studied for associated beetles, the most
diverse community (13 species) was found on
Megacollybia platyphylla (Table 2). The bee-
tle Proteinus brachypterus dominated (76% of
total individuals sampled from the fungus) and
was found on every studied long-stiped fungus.
The second-commonest species (10%) Atheta
boleticola was found along with other, mostly
Staphylinidae species represented by 1–2 indi-
viduals each. These include Atheta species and
larger Staphylinidae, such as Lordithon and
Sepedophilus. In the mushroom Hygrophoropsis
aurantiaca four associated beetle species were
registered, two of which (Proteinus brachypte-
rus and Autalia longicornis) were found also on
Fig. 3. Adult Cychramus luteus to colonize juvenile fruit
body of Armillaria borealis, Kumpula, Helsinki. Photo
D. Schigel, 2007.
Fig. 4. Adult Cychramus variegatus on the cap of Armil-
laria borealis, Susimäki Virgin Forest, Etelä-Häme. Pho-
to D. Schigel, 2005.
Fig. 2. Young fruit bodies of Armillaria borealis on birch
snag, Kolovesi National Park, Pohjois-Savo. Photo D.
Schigel, 2006.
40 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
Megacollybia. Unlike these two fungus species,
Volvariella bombycina was characterized by fi ve
Gyrophaena species, of which G. joyioides dom-
inated (76% of total individuals sampled from
the fungus). Gyrophaena bihamata is the only
species found on both Volvariella and Hygropho-
ropsis (Table 2).
(2) In Kolovesi clusters of Armillaria borea-
lis (Fig. 2) were growing simultaneously on dif-
ferent altitudes above ground. The moister and
lower-located clusters were fi rst to disappear,
but richest in beetles. Clusters over 0.5 m above
ground hosted only single individuals of Cery-
lon, in spite of their slightly extended lifetime,
and clusters above 2 m were in most cases intact
and free from beetles. Larvae of Cychramus spp.
are able to utilize the interior of short-living fruit
bodies of Armillaria. Two Cychramus species
were registered on Armillaria borealis – count-
less swarms of the common C. luteus in Kolovesi
and Helsinki, and the more rare C. varieagtus in
Susimäki (Figs. 3, 4). Adult Cychramus luteus
Table 2. Armillaria borealis plus selected other wood-rotting fungi, and their beetles in Southern Finland. Species of
fungi are given in an alphabetical order; IUCN threat categories (Rassi et al. 2001) are shown. B/F = number of beetle
individuals / number of fungal specimens studied. Number of insect specimens found or reared / number of fungal
specimens is given in parentheses after the beetle species name. Adult insects collected in nature indicated in light ital-
ics; larvae or reared adults (beetle – fungus associations meeting the Lawrence (1973) criterion) in bold italics.
# Fungus species B/F Insect records
1Armillaria borealis
Marxm. & Korhonen
665/38 Agathidium discoideum (1/1), Anopleta corvina (1/1), Atheta aeneipennis (10/5),
Atheta boleticola (28/13), Atheta cinnamoptera (1/1), Atheta nigritula (1/1),
Atheta paracrassicornis (2/2), Atheta pilicornis (15/6), Atheta sodalis (18/8),
Atomaria umbrina (8/1), Autalia impressa (53/10), Autalia longicornis (10/6),
Bolitochara pulchra (7/4), Catops nigricans (1/1), Catops nigrita (1/1), Cy-
chramus luteus (205/4), Cychramus variegatus (3/1), Deliphrum tectum (8/2),
Dryocoetes alni (1/1), Gyrophaena orientalis (1/1), Gyrophaena pulchella (1/1),
Lordithon bimaculatus (5/4), Lordithon lunulatus (2/2), Lordithon speciosus
(1/1), Lordithon thoracicus (5/4), Oxypoda alternans (62/14), Proteinus brachy-
pterus (189/14), Rhizophagus dispar (12/7), Rhizophagus nitidulus (3/1), Sepe-
dophilus littoreus (1/1), Sepedophilus marshami (3/2).
2Crepidotus calolepis
(Fr.) P. Karst.
8/2 Deliphrum tectum (1/1), Proteinus brachypterus (7/1).
3Hygrophoropsis auran-
tiaca (Wulfen : Fr.) J.
Schröt.
6/2 Atheta pilicornis (1/1), Autalia longicornis (1/1), Gyrophaena bihamata (1/1),
Proteinus brachypterus (3/1).
4Hypsizygus ulmarius
(Bull. : Fr.) Redhead
44/4 Atomaria affi nis (35/5), Cerylon ferrugineum (4/1), Rhizophagus dispar (5/2).
5Lentinellus castoreus
(Fr.) Kühner & Maire
9/3 Acrulia infl ata (2/1), Agathidium discoideum (3/1), Atheta nigritula (1/1), Atheta
pilicornis (1/1), Rhizophagus dispar (1/1), Sepedophilus marshami (1/1).
6Lentinellus vulpinus
(Sowerby : Fr.) Kühner
& Maire
4/1 Acrulia infl ata (1/1), Atheta crassicornis (1/1), Atheta pilicornis (1/1), Rhizopha-
gus dispar (1/1).
7Megacollybia platy-
phylla (Pers. : Fr.) Kotl.
& Pouzar
140/7 Atheta aeneipennis (2/1), Atheta boleticola (14/3), Atheta paracrassicornis (1/1),
Atheta pilicornis (3/2), Atheta sodalis (1/1), Autalia impressa (3/1), Autalia
longicornis (2/2), Bolitochara pulchra (1/1), Catops nigrita (1/1), Lordithon tho-
racicus (3/1), Oxypoda alternans (1/1), Proteinus brachypterus (107/4), Sepedo-
philus marshami (1/1).
41
KARSTENIA 47 (2007) SCHIGEL: FLESHY FUNGI AND COLEOPTERA
were dominating on younger Armillaria fruit
bodies in Helsinki, and only a few Staphylinidae
were observed. The reverse was seen on mature
fruit bodies of Armillaria in Kolovesi: Proteinus
brachypterus (41% of total individuals sampled
from the fungus) were the most common visitors,
followed by Oxypoda alternans (13%), Autalia
impressa (12%), and Atheta boleticola (6%).
Less frequent were other Staphylinidae, includ-
ing six more species of Atheta, and two species
# Fungus species B/F Insect records
8Panellus serotinus
(Schrad. : Fr.) J.G.
Kühn
1/1 Acrulia infl ata (1/1).
9Pholiota squarrosoides
(Peck) Sacc.
2/1 Cerylon ferrugineum (1/1), Gnathoncus nannetensis (1/1).
10 Pleurotus dryinus
(Pers. : Fr.) P. Kumm
40/6 Atheta castanoptera (1/1), Atomaria affi nis (1/1), Cryptophagus scanicus (5/2),
Rhizophagus bipustulatus (1/1), Rhizophagus dispar (2/1), Rhizophagus nitidulus
(1/1), Sepedophilus testaceus (1/1).
11 Pleurotus ostreatus
(Jacq. : Fr.) P. Kumm.
28/2 Cerylon ferrugineum (1/1), Triplax aenea (27/2).
12 Pleurotus pulmonarius
(Fr.) Quél.
423/31 Agathidium confusum (1/1), Agathidium pisanum (1/1), Atheta aeneipennis
(14/3), Atheta boleticola (1/1), Atheta boletophila (1/1), Atheta crassicornis
(2/2), Atheta pallidicornis (2/1), Atheta paracrassicornis (2/1), Atheta picipes
(13/5), Atheta pilicornis (1/1), Atheta sodalis (2/2), Atheta subtilis (20/5), Atom-
aria affi nis (1/1), Autalia impressa (8/1), Autalia longicornis (18/2), Cerylon fer-
rugineum (4/2), Cryptophagus confusus (1/1), Deliphrum tectum (9/1), Endomy-
chus coccineus (2/1), Glischrochilus hortensis (7/1), Gyrophaena affi nis (1/1),
Gyrophaena angustata (6/3), Gyrophaena fasciata (62/2), Gyrophaena joyi (7/1),
Gyrophaena orientalis (1/1), Gyrophaena poweri (1/1), Gyrophaena strictula
(4/2), Leptusa pulchella (5/2), Lordithon lunulatus (4/4), Lordithon speciosus
(6/3), Lordithon thoracicus (2/2), Lordithon trimaculatus (10/2), Megarthrus
depressus (2/1), Omalium rivulare (2/2), Oxypoda alternans (32/1), Proteinus
brachypterus (60/3), Quedius plagiatus (1/1), Rhizophagus dispar (42/8), Rhiz-
ophagus nitidulus (1/1), Sciodrepoides watsoni (1/1), Tachinus laticollis (3/2),
Tachinus pallipes (2/1), Triplax aenea (31/6), Triplax rufi pes (23/5)
13 Volvariella bombycina
(Schaeff. : Fr.) Singer
76/4 Cerylon histeroides (2/1), Gyrophaena affi nis (2/1), Gyrophaena bihamata (4/2),
Gyrophaena congrua (1/1), Gyrophaena fasciata (2/1), Gyrophaena joyioides
(58/3), Lordithon thoracicus (4/1), Pocadius ferrugineus (3/1).
14 Grifola frondosa (J.
Dicks.: Fr.) Gray, VU
245/8 Aleochara stichai (2/1), Atheta aeneipennis (1/1), Atheta castanoptera (4/1),
Atheta crassicornis (64/1), Atheta gagatina (1/1), Atheta marcida (3/2), Atheta
nigritula (29/1), Atheta paracrassicornis (65/1), Atheta picipes (2/1), Atheta pili-
cornis (5/1), Atheta sodalist (5/1), Autalia longicornis (1/1), Catops coracinus
(5/2), Deliphrum tectum (12/1), Lordithon bimaculatus (1/1), Lordithon lunulatus
(23/1), Lordithon thoracicus (2/1), Omalium rivulare (4/1), Philhygra malleus
(1/1), Philonthus fi metarius (2/1), Philorhizus sigma (1/1), Proteinus brachypte-
rus (9/1), Sepedophilus testaceus (1/1), Tachinus proximus (2/1)
of Sepedophilus. Agathidium discoideum, Atom-
aria umbrina, two species of Catops and two of
Rhizophagus contributed to the family and eco-
logical diversity of the associated species set, a
total of 31 species (Table 2).
(3) Among the log-favouring pleurotoid fungi,
Crepidotus calolepis, Panellus serotinus, Len-
tinellus castoreus, and L. vulpinus were visited
by generalists such as Proteinus brachypterus
and Acrulia infl ata. Unlike other fungi of the
42 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
kind, Hypsizygus ulmarius was ignored by adult
Staphylinidae, but favoured by Atomaria affi nis.
Rhizophagus dispar, numerous on Pleurotus pul-
monarius (Fig. 5) and several polypore species,
was spotted on the majority of the studied pleu-
rotoid fungi. Similarly to Hypsizygus ulmarius,
Pleurotus dryinus and P. pulmonarius attracted
Cryptophagidae, such as Atomaria affi nis, Cryp-
tophagus scanicus and C. confusus. Fruit bodies
of Pleurotus pulmonarius hosted 44 beetle spe-
cies – the highest number in this study. Of these,
dominating visitors were Gyrophaena fasciata
(15% of total individuals sampled from the fun-
gus), Proteinus brachypterus (14%), Oxypoda
alternans (8%) and Atheta aeneipennis (3%).
The list of visitors includes nine Atheta, six Gy-
rophaena, and four Lordithon species.
(4) Of the total 24 beetle species associated
with the red-listed (Rassi et al. 2001) polypore
Grifola frondosa (Fig. 6), the most typical bee-
tles were Atheta paracrassicornis (27% of to-
tal individuals sampled from the fungus) and
A. crassicornis (26%), with frequent Lordithon
lunulatus (9%). The less numerous species were
mainly Staphylinidae, including eight other spe-
cies of Atheta.
Fig. 5. Pleurotus pulmonarius on birch, Aakenustunturi,
Kittilän Lappi. Photo T. Niemelä, 2000.
Fig. 6. Grifola frondosa at the base of oak stump in Ruis-
salo forest, Turku. Photo D. Schigel, 2004.
4. Discussion
4.1. Consistency classes of polypores and
wood-rotting fl eshy fungi
The concept of consistency classes was created
to describe and name the patterns of fungus–in-
sect interactions found in nature (Schigel et al.
2004). This classifi cation refl ects the repeatedly
found beetle species associations and the respec-
tive properties of host fungi (shape and volume of
the fruit body, its durability, toughness, structure,
water contents, and annuality vs. perenniality).
Different polypore fruit bodies were divided
into six consistency classes. Perennial fruit bodies
of fomitoid polypores (e.g. Phellinus, Fomitopsis,
Ganoderma) attract beetles from the families Ano-
biidae, some Ciidae, and Tenebrionidae. Corky
and leathery trametoid fruit bodies (e.g. Len-
zites, Funalia, Cerrena) are favoured by species
of Ciidae; tyromycetoid polypores (e.g. Postia,
Amylocystis, Hapalopilus) are consumed by Hal-
lomenus. Both piptoporoid and xanthocroic poly-
pores produce annual, soft, voluminous and when
living water-rich fruit bodies. In spite this similar-
43
KARSTENIA 47 (2007) SCHIGEL: FLESHY FUNGI AND COLEOPTERA
Table 3. Systematic list of beetles attracted to Armillaria borealis and selected other wood-rotting fungi in southern
Finland. FS/IUCN = Frequency score of beetles (Rassi 1993) / threat categories (Rassi et al. 2001), if relevant; threat
categories and scores of the uncommon and rare beetle species in Finland (30 points and higher) are indicated in
bold face; Loc = locality, for details see Table 1: Å = Åland Islands, H = Helsinki, Kt = Kotinen Virgin Forest, Etelä-
Häme, Kv = Kolovesi National Park, Pohjois-Savo, L = Lammi Biological Station, Etelä-Häme, R = Ruissalo Island,
Varsinais-Suomi, S = Susimäki Virgin Forest, Etelä-Häme. Numbers refer to the species of fungi (Table 2); light face =
records of adult beetles; bold face = larvae or rearings.
Taxon FS/IUCN Loc Host fungi
COLEOPTERA
Carabidae Latreille, 1802
Philorhizus sigma (Rossi, 1790) 4 R 14
Histeridae Gyllenhal, 1808
Gnathoncus nannetensis (Marseul, 1862) 20 Kt 9
Leiodidae Fleming, 1821
Agathidium confusum Brisout de Barneville, 1863 6 Kt 12
Agathidium discoideum Erichson, 1845 40 Kv 1, 5
Agathidium pisanum Brisout de Barneville, 1872 10 Kt 12
Catops coracinus Kellner, 1846 10 R 14
Catops nigricans (Spence, 1815) 10 Kv 1
Catops nigrita Erichson, 1837 2 Kv 1, 7
Sciodrepoides watsoni (Spence, 1815) 1 L 12
Staphylinidae Latreille, 1802
Acrulia infl ata (Gyllenhal, 1813) 6 Kv 5, 6, 8
Aleochara stichai Likovský, 1965 40 R 14
Anopleta corvina (Thomson, 1856) 15 Kv 1
Atheta aeneipennis (Thomson, 1856) 2 L, Kt, Kv, R 1, 7, 12, 14
Atheta boleticola J.Sahlberg, 1876 15 Kt, Kv 1, 7, 12
Atheta boletophila (Thomson, 1856) 30 R 12
Atheta castanoptera (Mannerheim, 1830) 100/VU Å, R 10, 14
Atheta cinnamoptera (Thomson, 1856) 2 Kv 1
Atheta crassicornis (Fabricius, 1793) 4 Kv, L, R 6, 12, 14
Atheta gagatina (Baudi, 1848) 6 R 14
Atheta marcida (Erichson, 1837) 100/NT R 14
Atheta nigritula (Gravenhorst, 1802) 10 Kv, R 1, 5, 14
Atheta pallidicornis (Thomson, 1856) 40 Kt 12
Atheta paracrassicornis Brundin, 1954 3 L, Kv, R 1, 7, 12, 14
Atheta picipes (Thomson, 1856) 15 R 12, 14
Atheta pilicornis (Thomson, 1852) 10 Kt, Kv, R 1, 3, 5, 7, 12, 14
Atheta sodalis (Erichson, 1837) 2 L, Kt, Kv, R 1, 7, 12, 14
Atheta subtilis (Scriba, 1866) 1 L, Kt, R 12
Autalia impressa (Olivier, 1795) 60 Kt, Kv 1, 7, 12
Autalia longicornis Scheerpeltz, 1947 4 L, Kt, Kv, R 1, 3, 7, 12, 14
Bolitochara pulchra (Gravenhorst, 1806) 2 Kv 1, 7
Deliphrum tectum (Paykull, 1789) 1 Kt, Kv 1, 2, 12, 14
Gyrophaena affi nis Mannerheim, 1830 2 H, Kt 12, 13
Gyrophaena angustata (Stephens, 1832) 30 R 12
Gyrophaena bihamata Thomson, 1867 15 H, Kt 3, 13
Gyrophaena congrua Erichson, 1837 30 H 13
Gyrophaena fasciata (Marsham, 1802) 2 H 12, 13
Gyrophaena joyi Wendeler, 1924 40 Kt 12
Gyrophaena joyioides Wüsthoff, 1937 10 H 13
Gyrophaena orientalis Strand, 1938 40 Kt 1, 12
Gyrophaena poweri Crotch, 1866 10 Kt 12
Gyrophaena pulchella Heer, 1839 15 Kv 1
Gyrophaena strictula Erichson, 1839 15 R 12
44 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
Taxon FS/IUCN Loc Host fungi
Leptusa pulchella (Mannerheim, 1830) 1 R 12
Lordithon lunulatus (Linnaeus, 1761) 2 Kv, R 1, 12, 14
Lordithon speciosus (Erichson, 1840) 20 Kv 1, 12
Lordithon thoracicus (Fabricius, 1777) 1 H, Kt, Kv, R 1, 7, 12, 13, 14
Lordithon trimaculatus (Fabricius, 1793) 30 Kt 12
Lordithon trinotatus (Erichson,1839) 100/NT Kv, R 1, 14
Megarthrus depressus (Paykull, 1789) 6 L 12
Omalium rivulare (Paykull, 1789) 1 Kt, R 12, 14
Oxypoda alternans (Gravenhorst, 1802) 2 Kt, Kv 1, 7, 12
Philhygra malleus (Joy, 1913) 6 R 14
Philonthus fi metarius (Gravenhorst, 1802) 6 R 14
Proteinus brachypterus (Fabricius, 1792) 1 L, Kt, Kv, R 1, 2, 3, 7, 12, 14
Quedius plagiatus Mannerheim, 1843 2 Kt 12
Sepedophilus littoreus (Linnaeus, 1758) 4 Kv 1
Sepedophilus marshami (Stephens, 1832) 6 Kv 1, 5, 7
Sepedophilus testaceus (Fabricius, 1793) 2 R 10, 14
Tachinus laticollis Gravenhorst, 1802 2 L, Kt 12
Tachinus pallipes (Gravenhorst, 1806) 1 Kt 12
Tachinus proximus Kraatz, 1855 2 R 14
Nitidulidae Latreille, 1802
Glischrochilus hortensis (Geoffroy, 1785) 4 L 12
Cychramus luteus (Fabricius, 1787) 4 H, Kv, S 1
Cychramus variegatus (Herbst, 1792) 15 S 1
Pocadius ferrugineus (Fabricius, 1775) 6 H 13
Monotomidae Laporte de Castelnau, 1840
Rhizophagus bipustulatus (Fabricius, 1793) 6 Å 10
Rhizophagus dispar (Paykull, 1800) 2 Kv 1, 4, 5, 6, 10, 12
Rhizophagus nitidulus (Fabricius, 1798) 30 Å, Kt, Kv 1, 10, 12
Cryptophagidae Latreille, 1802
Atomaria affi nis (F.Sahlberg, 1834) 30 S 4, 10, 12
Atomaria umbrina (Gyllenhal, 1827) 10 Kv 1
Cryptophagus confusus Bruce, 1934 60 Kt 12
Cryptophagus scanicus (Linnaeus, 1758) 6 Å 10
Erotylidae Latreille, 1802
Triplax aenea (Schaller, 1783) 15 Å, R 11, 12
Triplax rufi pes (Fabricius, 1781) 40 R 12
Cerylonidae Billberg, 1820
Cerylon ferrugineum Stephens, 1830 4 R 4, 9, 11, 12
Cerylon histeroides (Fabricius, 1793) 2 H 13
Endomychidae Leach, 1815
Endomychus coccineus (Linnaeus, 1758) 10 Kt 12
Curculionidae Latreille, 1802
Dryocoetes alni (Georg, 1856) 10 Kv 1
45
KARSTENIA 47 (2007) SCHIGEL: FLESHY FUNGI AND COLEOPTERA
ity, their beetles are different: Tenebrionidae and
Erotylidae mostly use piptoporoid polypores (e.g.
Polyporus, Laetiporus), and Tetratomidae mostly
xanthocroic ones (Inonotus, Onnia). Thin imbri-
cate pilei of Trichaptum provide an environment
for a few specialized beetle genera.
The course of changes within fruit body (ju-
venile – mature – old) differs between wood-
rotting fl eshy fungi and polypores. The fi rst and
main difference is the time scale: from days in
agarics, to weeks or even years in polypores.
Mature fruit bodies of agarics are always fertile
and sporulating, which corresponds to polypore
decomposition stage 2 (out of 4). Differently
from agarics, the sporulation period of polypore
fruit bodies is short compared with their lifetime.
The decomposition processes of agarics and
more robust polypores differ in the similar way:
the fast collapse of the ephemeral fruit bodies of
agarics and softer polypores requires swift larval
development and escape for soil pupation, while
the more durable polypores allow more lengthy
larval development and possibility of pupation
on the spot (decomposition stages 3 and 4).
4.2. Agaricoid, armillarioid and pleurotoid
consistency classes
By analogy with polypores, three new consist-
ency classes are needed to describe wood-rot-
ting fl eshy fungi: agaricoid, armillarioid and
pleurotoid. These accord with the groups (1) to
(3) in the previous section 3 (Results). Fruit bod-
ies of these fungi differ from each other in their
longevity, way of growth, location on tree trunk,
and in the numbers of pilei per cluster. These
Table 4. Basidiocarp consistency classes among the wood-rotting fl eshy fungi, and examples of characteristic fungal
and beetle genera in southern European taiga.
Agaricoid
Annual, prominently stipitate, thin, solitary or gregarious, usually on logs or stumps, extremely ephemeral and
quickly decomposed into black slime.
Examples of fungal genera: Hygrophoropsis, Megacollybia, Volvariella.
Examples of beetles associated: adult Staphylinidae: Atheta pilicornis, Atheta boleticola, Autalia longicornis,
Gyrophaena bihamata, Gyrophaena joyioides, Proteinus brachypterus.
NB: The extremely ephemeral nature of the fruit bodies and their erratic appearance in different years make
agaricoid fungi unsuitable for beetle larvae, but visiting adults are numerous and diverse.
Armillarioid
Annual, prominently stipitate, thick and fi brous, usually with tens of fruit bodies packed in commonly rooted
clusters, in lower parts of tree trunks and snags, ephemeral and decomposed into black slime, but in dry condi-
tions dead fruit bodies persist over winter.
Examples of fungal genera: Armillaria, Pholiota.
Examples of beetles associated: Cychramus (including larvae), adult Staphylinidae.
NB: The voluminous and persistent wood-penetrating mycelium guarantees more regular fructifi cation com-
pared to the agaricoid ones. This predictable availability of the host allows Cychramus spp. to breed specifi cally
in the fungus.
Pleurotoid
Annual, sessile or with rudimentary stipe, solitary or packed in commonly-rooted clusters of a few fruit bodies,
on trunks or lower crown; fungus may continue to grow on downed trunks or fallen branches. Fruit bodies some-
times stay dried on the substrate after sporulation.
Examples of fungal genera: Crepidotus, Hypsizygus, Lentinellus, Panellus, Phyllotopsis, Pleurotus.
Examples of beetles associated: Cyllodes and Triplax (including larvae), adult Staphylinidae, Rhizophagus, and
Cerylon.
NB: Similarly to armillarioid fruit bodies, pleurotoid ones host beetle larvae, which fi nally drop to the soil for
pupation (Triplax). Adults of Cerylon and Rhizophagus are repeatedly collected on gills of fully developed living
basidiocarps.
46 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
consistency classes are described in Table 4 and
below.
Agaricoid fungi are a diverse group of mostly
solitary mushrooms of the forest fl oor and heav-
ily decayed wood. These fungi show a variety of
sizes and shapes, but their ephemeral habit and
unclustered way of growth are the characteristic
features. Beetles associated to agaricoid fungi
are mainly generalist adult Staphylinidae. Even
though only a few agaricoid species were docu-
mented for beetles in this study, it is likely that
Staphylinidae will turn out to dominate in future
collections from many other similar mushroom
species.
Armillarioid consistency class by now com-
prises of two genera of fungi, Armillaria and Pho-
liota. The species-rich genus Pholiota is poorly
documented for associated beetles. It seems,
however, that Pholiota does not belong to the
most commonly visited fungal substrates of gen-
eralist beetles. In Armillaria, like in many other
fungi, the surface grazers (adult Coleoptera),
and the fungivorous dwellers inside the moister
fungal context (mostly larvae of Diptera) do not
compete due to their spatial isolation. However,
fungivorous larvae of dwellers sometimes reach
high densities within the mushroom fruit bod-
ies and then competitive interactions within and
among species may occur. The extremely ephem-
eral nature of mushrooms causes the competition
with time to be more critical for survival than the
competition among individuals.
Co-occurrence of two Cychramus species on
Armillaria was documented in Susimäki, with
the unusual prevalence of C. variegatus. During
the lab experiment adults of Cychramus luteus
collected in Helsinki were eager to mate on the
Armillaria fruit bodies. Probably, larvae of the
two Cychramus species are able to develop in
Armillaria, and as seen from the present Finnish
pilot material, the prevalence of a certain spe-
cies changes geographically. There is little doubt
that larvae of Cychramus luteus in southern Fin-
land dwell mainly in Armillaria fruit bodies,
while more data on the biology of C. variegatus
are needed. In addition to fungi, adults of these
Cychramus species are frequently collected on
fl owers during summertime, especially of the
Apiaceae and Rosaceae. Cychramus luteus is
feeding on pollen, not only on fl owers, but also in
bee hives, intruding to open the brood cells. Cy-
chramus luteus does not otherwise affect brood,
honey or combs, and was never found breeding
there (Neumann & Ritter 2004). The compari-
son of my data to the existing literature data is
complicated by the fact that different Armillaria
species were recognized only lately, and in most
published studies they are collectively treated as
“Armillaria mellea” – a species, that does not
occur in Finland. Most probably, however, adult
beetle visitors fl exibly use the available Armil-
laria species, but the Cychramus species may
be selective according to different geographical
regions.
Pleurotoid fungi are found from the forest
fl oor high up to branches, and they are more
long-lasting than the other gilled wood-rotting
mushrooms. Pleurotus may produce fruit bodies
high in the tree crown, while others, like Crepi-
dotus and Panellus mostly occupy logs and fallen
branches. The most long-lasting, beetle-rich and
best-studied fungi in this consistency class are
the Pleurotus species, and probably the most in-
teresting beetle genus in relation to Pleurotus is
Triplax. In Nordic countries larvae of three Tri-
plax species feed on Pleurotus fruit bodies. Still
another, T. russica (Linnaeus, 1758) develops in
Inonotus obliquus. Triplax russica and other bee-
tle associates avoid the sterile conks of Inonotus
obliquus, and come to colonize the sporulating
fruit bodies as soon as the covering birch bark
cracks. In the present material from southern
Finland, two out of three Pleurotus-associated
Triplax species were discovered. Triplax aenea
and T. r ufi pes were reared in equal numbers from
P. pulmonarius. Triplax aenea larvae also utilize
P. ostreatus (Table 2).
Triplax scutellaris Charpentier, 1825, missing
from this material, was earlier registered by me
in Finnish Lapland (Pisavaara and Salla) coloniz-
ing Pleurotus pulmonarius and P. dryinus. This
data overview shows the specifi city of the three
Triplax species to Pleurotus, but also their abil-
ity to fl exibly use alternative host species. Five
adults of a rare, near-threatened species (Rassi
et al. 1993, 2001), Agathidium pallidum (Gyl-
lenhal, 1827), were collected from two samples
of Pleurotus dryinus in eastern Finnish Lapland
(Salla: Julmoiva) in 2005. Another Pleurotus-
dependent species, Cyllodes ater has now been
rediscovered in eastern Finland (Yakovlev &
Hokkanen 1995). Beetles whose larvae develop
rapidly in ephemeral substrates may have several
generations a year depending on substrate avail-
47
KARSTENIA 47 (2007) SCHIGEL: FLESHY FUNGI AND COLEOPTERA
ability. It is unclear if two or more generations of
Triplax can complete their cycle within a single
growth season in North Europe.
In Russian Far East Zaitsev and Kompantsev
(1987) list Cyllodes, Triplax, and Eutriplax as
obligate beetle dwellers whose larvae primarily
feed on Pleurotus spp. These associations were
detected also in other parts of Eurasia with such
species as Triplax rubrica, T. collaris, T. lepida,
and T. ru fi pes. Monophagous larvae of Triplax
gracilenta are associated with East Asian Pleu-
rotus citrinopileatus. Obligate inhabitants of
Pleurotus in Russian Far East (Cyllodes, Triplax
and Eutriplax) demonstrate how the life cycles
are correlated with the longevity of fungal hosts.
Larvae abandon decaying fruit bodies to pupate
in soil. The pattern of colonisation and destruc-
tion is governed by host moisture: dry carpo-
phores are mostly utilized by Coleoptera (Tri-
plax, Eutriplax, Cyllodes), but wet ones are used
by Diptera. Dry pilei of Pleurotus are similar in
their consistency with piptoporoid polypores:
Dacne larvae occupy Pleurotus, but also live
in Piptoporus betulinus, Laetiporus sulphureus,
and Spongipellis litschaueri. Very dry and hard
fruit bodies of Pleurotus are durable enough to
support larvae of Platydema dejeani and Ennear-
thron sp. (Zaitsev & Kompantsev 1987).
Grifola frondosa – This oak-decaying poly-
pore makes a special case as its way of growth
and associated beetles resemble clustered wood-
rotting agarics, rather than polypores. This fun-
gus was observed in the Ruissalo oak forest. Late
time of the fruit body growth combined with rar-
ity may explain the absence of beetle larvae in
Grifola frondosa. Voluminous clustered struc-
ture, fi brous fruit body, limited longevity, and a
habit of growing at the base of oak trees make
the fungus a meeting point for late-season adults
of mycophilous Staphylinidae. Such ecological
role of Grifola frondosa may be compared with
Armillaria in Kolovesi.
4.3. Beetles of wood-rotting agarics,
a summary
Certain parallelism in seasonality can be seen be-
tween wood-rotting mushrooms and polypores.
Beetle larvae were absent from annual ephemer-
al polypores, and from agaricoid fungi. In annual
sturdy polypores and pleurotoid agarics the peak
of beetle activity fell on the living fruit bodies.
Characteristics of armillarioid fungi are some-
what intermediate, and so is the set of associated
beetles.
Differently from beetle-rich annual ephemeral
and annual sturdy polypores, wood-rotting fl eshy
fungi hosted only the rapidly growing larvae of
Cychramus and Triplax. These beetles are spe-
cifi c to only one host-fungus genus, Armillaria
and Pleurotus, respectively. The structure and
seasonality of fruit bodies of fl eshy wood-rotting
fungi make Staphylinidae the main group of adult
beetle visitors. Typically one to three species of
rove beetles outnumber the other species.
Annual fruit bodies of wood-rotting fungi pro-
vide beetles with an ecological fast-food service:
these nutritional resources are often available for
the visitors after they have hatched from pupae,
but before their breeding hosts appeared. Only
a few beetle species are adapted to use these
ephemeral fungi as their breeding media.
5. Conclusions
Agaricoid, armillarioid, and pleurotoid fungi
make three new consistency classes among the
wood-rotting fungi. Together with polypores,
discussed in previous papers of the series, fruit
bodies of fl eshy wood-rotting agarics provide
a wide range of habitats for fungivorous Cole-
optera, both adults and larvae. Our knowledge
of fungivorous beetles and their host fungi has
grown considerably, but some aspects of their
ecology remain neglected. Among such topics is
the impact of frost and drought on the seasonal
dynamics of fungus–beetle systems. Ecologically
frost and drought have a lot in common (periods
of hibernation), but they require different physi-
ological adaptations from fungi and beetles. Fruit
bodies of ephemeral fungi can not resist extreme
conditions; mycelium of some species react to
a sudden change in temperature by starting to
fruit in time unsuitable for beetle colonization,
while perennial fruit bodies withstand repeated
seasonal changes and attacks of beetles. I plan to
expand my studies in fungus–beetle interactions
into different climatic zones, and to other biogeo-
graphical aspects of the fungivory of Coleoptera,
their life strategies and functional morphology.
The present pilot study covered fruit bodies
of just a few wood-rotting fl eshy fungi. Larvae
of many beetle species depend on decayed wood
penetrated by fungal mycelium. Other fungal
48 KARSTENIA 47 (2007)
SCHIGEL: FLESHY FUNGI AND COLEOPTERA
substrates still require closer look: stereoid fungi
may have similar substrate characteristics as the
thin Trichaptum basidiocarps; boletes are prob-
ably visited by adult Staphylinidae generalist,
like those on agarics. Ecology of beetles of both
soft and sturdy Ascomycetes, the specialized in-
habitants of Gasteromycetes, consumers of an-
amorphic fungi and fungi-like habitats such as
Myxomycetes are to a large extent unknown.
Acknowledgements: Tuomo Niemelä kindly supervised
the writing, and collaborated in the fi eld together with
Juha Kinnunen and Petri Oinonen, all from the Univer-
sity of Helsinki. Viktor B. Semenov (E. I. Marcinovsky
Institute of Medical Parasitology and Tropical Medicine,
Moscow), and Nikolay B. Nikitsky (Moscow State Uni-
versity) identifi ed selected diffi cult beetle specimens. The
PUTTE research programme for defi ciently known and
threatened forest species (Finnish Ministry of Environ-
ment) is thanked for supporting the study.
References
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parative Zool. 145: 163–212.
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association of Cychramus luteus (Coleoptera: Nitidul-
idae) with honeybee (Apis mellifera) colonies. – Api-
dologie 35: 665–666.
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puiston kääpäinventointi. – Metsähallitus, Etelä-
Suomen luontopalvelut. 19 pp.
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tera) frekvenssipisteet 1.1.1960 – 1.1.1990. – Maail-
man Luonnon Säätiön WWF Suomen Rahaston Ra-
portteja 6: 1–136.
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(eds.) 2001: The 2000 Red List of Finnish species. –
Ministry of the Environment & Finnish Environment
Institute. 432 pp.
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(eds.) 2005: Suomen helttasienten ja tattien ekologia,
levinneisyys ja uhanalaisuus. – The Finnish Environ-
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pores of western Finnish Lapland and seasonal dy-
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Manninen O. 2004: Polypores and associated beetles
of the North Karelian Biosphere Reserve, eastern Fin-
land. – Karstenia 44: 35–56.
Schigel, D.S. & Toresson, H.G. 2005: New records of
Polyporus pseudobetulinus, a rare polypore fungus
(Basidiomycota, Aphyllophorales) in Scandinavia,
and notes on associated beetles. – Memoranda Soc.
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Silfverberg, H. 2004: Enumeratio nova Coleopterorum
Fennoscandiae, Daniae et Baltiae. – Sahlbergia 9:
1–111.
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leoptera, Nitidulidae) found again in Finland. – Ento-
mol. Fennica 5: 203–204.
Zaitsev, A.I. & Kompantsev, A.V. 1987: Complexes
of Coleoptera and Diptera, associated with carpo-
phores of wood-rotting fungi of genus Pleurotus (Fr.)
Quél. in Eastern Siberia and Far East. [
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Karstenia 47: 49–54, 2007
The genera Typhula and Pistillaria (Typhulaceae,
Aphyllophorales) in Finland. A check-list of the
species
ANTON SHIRYAEV and HEIKKI KOTIRANTA
SHIRYAEV, A. & KOTIRANTA, H. 2007: The genera Typhula and Pistillaria (Aphyllo-
phorales) in Finland. A check-list of the species. – Karstenia 47: 49–54. 2007. Helsinki.
ISSN 0453-3402.
Twenty two Typhula (Pers.) Fr. species and one Pistillaria Fr. species are reported from
Finland. Five species are new for the country, viz. Typhula capitata (Pat.) Berthier, T.
culmigena (Mont. & Fr.) J. Schröt., T. hyalina (Quél.) Berthier, T. spathulata (Peck)
Berthier, and T. struthiopteridis Corner. Most of the species are common. The localities,
habitats, substrates and short ecological notes for the new species are given.
Key words: Distribution, Finland, Pistillaria, Typhula.
Anton Shiryaev, Institute of Plant and Animal Ecology RAS, 8 March str. 202, 620144,
Ekaterinburg, Russia. E-mail: 26081978@mail.ru
Heikki Kotiranta, Finnish Environment Institute, Research Department, Research Pro-
gramme for Biodiversity, P.O. Box 140, FI-00251 Helsinki, Finland. E-mail: heikki.ko-
tiranta@ymparisto.fi
Introduction
Most of the 23 species of Typhula and Pistillaria
found in Finland are saprotrophic, more seldom
they are parasites of grasses and herbs. P. A.
Karsten published several papers (e.g. Karsten
1879, 1881a, 1881b, 1881c, 1882, 1899) on new
species and notes on species which he mostly
collected in the vicinity of Mustiala (Tammela).
He did not give detailed information on the dis-
tribution of the species in Finland, because no
other material was available for him. Some more
recent papers also list Typhula species. Ulvinen
et al. (1981) listed species from the province of
Koillismaa (Ks), and Ohenoja (1996) from Lap-
land, and according to Knudsen (1997) 17 spe-
cies were known in Finland. However, the dis-
tribution of most of the species was not given by
Knudsen (1997).
Material and methods
This is the fi rst study where the whole Finnish material,
altogether 390 specimens, of Finnish and some other her-
baria (H, OULU, TUR, SVER, TAA) was examined.
The biological provinces and collecting sites are ac-
cording to Heikinheimo and Raatikainen (1981). The
provinces are illustrated also in Hansen and Knudsen
(1997: 28). The nomenclature of Typhulaceae follows
Berthier (1976) and that of vascular plants Hämet-Ahti et
al. (1998). The following abbreviations for substrates are
used: Epilobium refers to E. angustifolium, Pinus to Pi-
nus sylvestris, Picea to Picea abies, and Populus to Popu-
lus tremula. Species new to Finland are marked with an
asterisk (*). Notes of the distribution, hosts and biology
of the new species are partly based on Russian material,
made by the senior author, and have not been published
before. We believe that the biology of the species in Fin-
land is very similar to that in adjacent areas in Russia. The
identifi cation of the species is made by the senior author.
50 KARSTENIA 47 (2007)
SHIRYAEV & KOTIRANTA: TYPHULA AND PISTILLARIA IN FINLAND
List of species
Pistillaria paradoxa (P. Karst.) Corner
EH: Tammela, Mustiala, decayed twigs of Pi-
nus, 8.XIII.1865 Karsten 1297, 1298, 1300
(lectotype, H), decayed twigs of Pinus, 5.I.1866
Karsten 1299 (topotype, H).
Typhula athyrii Remsberg
Syn. Typhula todei Fr. : Fr.
EH: Tammela, Mustiala, Haarankorpi, dead pet-
ioles of Athyrium sp., 1.IX.1869 Karsten 3164
(H).
*Typhula capitata (Pat.) Berthier
EH: Luhanka, Lempää, dead Calamagrostis
arundinacea, 25.IV.2007 Shiryaev 7000 (SVER).
InL: Utsjoki, Kevo, dead grasses, 30.VIII.1973
Ulvinen (OULU F23311).
Typhula caricina P. Karst.
EH: Tammela, Haarankorpi, Carex vesicaria,
16.IX.1869 Karsten 1291 (lectotype, H), Tam-
mela, Mustiala, C. vesicaria, 8.IX.1870 Karsten
1292 (syntype, H). PH: Virrat, Hauhuu, Siko-
saari, Salmela, decaying leaves of Carex sp.,
27.IX.1995 Kytövuori 95-1599 (H).
Typhula crassipes Fuckel
Syn. T. corallina Quél. & Pat.
V: Turku, Impivaara, decaying leaves of Populus,
20.IX.1981 Huhtinen 81/226 (TUR 68946). EH:
Nokia, Kankaantaka, decaying leaves of Betula
pendula and Populus, 18.IX.1977 E. Ohen-
oja (OULU F49269), Tammela, Haarankorpi,
decaying leaves of Populus, 2.IX.1869 Karsten
(H), Tammela, Mustiala, decaying leaves of Al-
nus sp., 20.X.1891 Karsten 1286 (holotype of T.
cylindracea P. Karst., H). OP: Muhos, Muhos-
perä, Isterinkoski, dead grasses, 4.VII.1970 E.
& M. Ohenoja (OULU F49293). Ks: Kuusamo,
decaying stems of Actaea spicata, 29.VIII.1988
Ulvinen (OULU F22652).
*Typhula culmigena (Mont. & Fr.) J. Schröt.
V: Korppoo, Bonäs, dead stem of Verbascum
thapsus, 17.VII.1948 Kari (TUR 30810). U:
Nurmijärvi, church village, dead stems of Fes-
tuca pratensis, 29.IX.1986 Askola 1902 (TUR
53340). EK: Sippola, Ruotila, Epilobium,
7.IX.1947 Kujala (TUR 062431). EH: Padas-
joki, Vesijako Strict Nat. Res., fallen stems of
Epilobium, 13.IX.2001 Parmasto (TAA(M)
180732). Tammela, Mustiala, decaying grasses,
2.VII.1867 Karsten (H). OP: Oulu, Nuottasaari,
Equisetum arvense, 7.X.1965 Ulvinen (OULU
F49288). Pudasjärvi, Jaurakkajärvi, dead stems
of Epilobium, 18.VII.1988 E. Ohenoja (OULU
F45805); Ks: Kuusamo, Liikasenvaara, Kor-
vasvaara, spruce dominated herb-rich forest,
Epilobium, 23.VIII.1978 E. Ohenoja (OULU
F49299).
Typhula erythropus (Pers. : Fr.) Fr.
V: Lohja, Orsnäs, decaying leaves of Salix cap-
rea, 1.X.1937 Laurila (H). Parainen, Degerholm,
decaying leaves, 8.X.1967 A. & K. Pohjola
(TUR 30812). Turku, Hirvensalo, leaves of
Populus, 20.IX.1983 Huhtinen 83/317 (TUR
73707). St: Noormarkku, decaying leaves of
Alnus incana, 20.IX.1937 Laurila (H, TUR
30644). EH: Lammi, decaying petioles of de-
ciduous trees, 18.IX.1990 E. Ohenoja (OULU
F34126). Luhanka, Lempää, decaying leaves
of Populus, 24.IV.2007 Shiryaev 7001 (SVER).
Tammela, decaying leaves of Acer platanoides,
15.VIII.1868 Karsten (H). Tammela, Haaranko-
rpi, deciduous leaves, 24.IX.1878 Karsten (H).
Tampere, Peltolammi, decaying leaves of Al-
nus sp., 9.IX. 1988 Söderholm (H). Tampere,
Rusko, Polunmäki, dead petioles of Alnus sp.,
18.IX.2005 Söderholm 3669 (TUR 173242). EP:
Isojoki, Lauhanvuori, Huhtakorpi, mesic conif-
erous forest, leaves of A. incana, 21.X.2005 E.
Ohenoja (H). Kn: Paltamo, Oikarila, deciduo-
us litter, 22.IX.1974 E. & M. Ohenoja (OULU
F38685). OP: Kiiminki, Kalkkialue, Vasikka-
suo, decaying leaves of Betula sp., 23.IX.1969
Ulvinen (OULU F49317).
Typhula graminum P. Karst.
EH: Luhanka, Lempää, decaying Calamagrostis
arundinacea, 24.IV.2007 Shiryaev 7002, 7003
(SVER). Tammela, Mustiala, X.1867 Karsten
(H, holotype);
*Typhula hyalina (Quél.) Berthier
EH: Tammela, Mustiala, dead stems of Dactylis
glomerata, 18.VIII.1869 Karsten (H), meadow,
dead stems of Carex sp., 21.IX.1869 Karsten
(H).
Typhula incarnata Lasch ex Fr.
Syn. T. elegantula P. Karst.
51
KARSTENIA 47 (2007) SHIRYAEV & KOTIRANTA: TYPHULA AND PISTILLARIA IN FINLAND
EH: Luhanka, Lempää, decaying leaves of Ca-
lamagrostis arundinacea, 23.IV.2007 Shiryaev
7004 (SVER). Tammela, Mustiala, Elytrigia re-
pens, VIII.1868 Karsten 1294, 1295 (isotype of
T. elegantula, H), Calamagrostis sp., 6.X.1867
Karsten 1477, 1478 (H). PS: Kiuruvesi, Mäen-
pää, Secale cereale, 9.V.1948 Huttunen (H).
Maaninka, Festuca pratensis, 15.V.1948 Jama-
lainen (H). OP: Oulu, Hintta, decaying grasses,
2.X.1975 E .Ohenoja (OULU F49351). Ylis-
taro, Phleum pratense, 5.VI.1946 Jamalainen
(H).
According to Jamalainen (1957) and Mäkelä
and Mäki (1980) T. incarnata is frequent in some
years on cultivated winter cereals. However, the
herbarium material is scanty.
Typhula ishikariensis Imai var. ishikariensis
ES: Mikkeli, Secale cereale, 3.V.1948 Yli-
mäki (H), Mikkeli Agr. Stat., Secale cereale,
29.IV.1948 Koskinen (H). Savonlinna, Nojan-
maa, grasses, 1947 Ylimäki (H). PH: Karstula,
Vastinki, Phleum pratense, 1.V.1948 Marttinen
(H). KP: Ruukki, Secale cereale, 31.V.1946,
3.VI.1946, 31.VI.1946 Jamalainen (H). PeP:
Rovaniemi, Apukka, decaying grasses, 11.V.2000
Wäli (TUR 159943). KiL: Kolari, Phleum pra-
tense, 7.VI.1946 Jamalainen (H).
Typhula lutescens Boud.
PH: Keuruu, Pihlajavesi, decaying Equisetum
sp., 28.IX.1995 Kytövuori 95-1664 (H). Kn:
Puolanka, Paljakka, Jatkonsuo, decaying herbs
and mosses, 7.IX.1979 Eurola (OULU F49274).
OP: Kiiminki, Isohalmeenmaa, decaying
Equisetum sp., 5.IX.1968 M. Ohenoja (OULU
F49307). Kiiminki, limestone area, Equisetum,
23.IX.1969 Ulvinen (OULU F49308), Raivio
spring, calcareous soil, E. palustre, 5.IX.1968
Ulvinen (OULU F49306). PeP: Tervola, Peura,
Equisetum sp., 3.IX.1970 M. Ohenoja (OULU
F49310). InL: Utsjoki, Kevo, E. fl uviatile,
30.VIII.1973 Ulvinen (OULU F23320). Uts-
joki, Njallajärvi, mosses and decaying herbs,
13.IX.1967 Kankainen (TUR 30815).
Typhula micans (Pers. : Fr.) Berthier
Syn. T. anceps P. Karsten
V: Kakskerta, Harjattula ön, Trifolium mon-
tanum, 17.VIII.1936 Kari (TUR 72321); St:
Tyrvää (Tyrvis), decaying Cirsium arvense,
14.IX.1859 Karsten 1289 (H); EH: Kangasala,
Vihtinen, decaying leaves of Angelica sylvestris,
07.VIII.1994 Söderholm 2229 (TUR 113330),
Luhanka, Lempää, dead stems of Tussilago far-
fara, 25.IV.2007 Shiryaev 7005, 7006 (SVER),
Tammela, Mustiala, decaying leaves of Salix
sp.,14.IX.1870 Karsten 1285 (holotype of T. an-
ceps, H), decaying Cirsium arvense, 30.VI.1871
Karsten 1290 (H), Sedum acre, 23.IX.1872
Karsten 1306 (H).
Unlike Berthier (1976) and Knudsen (1997),
we believe that T. anceps is a synonym of T. mi-
cans, because we could not found reliable distin-
guishing characteristics between these species.
Typhula olivascens Berthier
EH: Nokia, Tottijärvi, Tolppa, decaying petioles
of Pteridium aquilinum, 27.IX.1995 Kosonen
(herb. Berthier; specimens not seen).
Typhula phacorrhiza (Reichard : Fr.) Fr.
V: Koski, Hongisto, litter in mixed forest,
30.IX.1990 Heinonen 99-90 (TUR 101595).
Kustavi, Kiparluoto, Riihivainio, decaying
leaves and other plant remnants, 25.IX.1994
Mäkinen 94-1124 (TUR 114336). Turku, dead
leaves of Betula sp., 17.X.1993 Vauras 9021
(TUR 109955), compost heap,17.X.2006 Vauras
25584 F (TUR 176466). U: Nurmijärvi, church
village, litter, 15.IX.1986 Askola 2022 (H), deci-
duous litter, 22.IX.1990 Askola 2735 (H). Sipoo,
Hindsby, decaying leaves of Populus, 23.09.1978
Saarenoksa 32578 (H). St: Noormarkku, decay-
ing leaves of Betula and Alnus sp., 18.IX.1937
Laurila (TUR 30785). EH: Kalvola, Taljala-
Niemi, Kalliomaa Nat. Res., decaying leaves,
23.IX.1980 Alho (TUR 97507). Lammi, Hauhia-
la, litter, 3.IX.1984 Harmaja (H), Pappilankylä,
decaying leaves of Populus, 7.IX.1983 Harmaja
(H). Tammela, decaying leaves of B. pendula,
1.V.1972 E. Ohenoja (OULU F49272). Tam-
pere, Peltolammi, deciduous litter, 31.VIII.1988
Söderholm 1513 (OULU F49315). OP: Kiimin-
ki, Murtoinsaaret, deciduous litter, 5.IX.1968 Ul-
vinen (OULU F49345), 8.IX.1968 M. Ohenoja
(OULU F49352). Muhos, Muhosperä, deciduous
litter, 17.IX.1976 E. Ohenoja (OULU F49349).
Typhula quisquiliaris (Fr. : Fr.) Henn.
U: Kirkkonummi, mixed forest, with Corylus,
Picea, Populus, Sorbus, decaying petioles of Pte-
ridium aquilinum, 25.08.1982 Saarenoksa 46282
(H). EH: Kangasala, Vatiala, Kisapirtti, dead
52 KARSTENIA 47 (2007)
SHIRYAEV & KOTIRANTA: TYPHULA AND PISTILLARIA IN FINLAND
petioles and leaves of P. aquilinum, 29.VIII.1995
Söderholm 2439 (TUR 115285). Tammela, Mus-
tiala, P. aquilinum, VIII.1866 Karsten (H). KP:
Siikajoki, Paavola, decaying petioles of P. aqui-
linum 29.VIII.1965 Kankainen (TUR 97944).
Typhula sclerotioides (Pers.) Fr.
V: Merimasku, decaying herbs, VIII.1860
Karsten 1803 (H). U: Espoo, decaying grasses,
IX.1937 Frey (H). EH: Tammela, Mustiala,
herbs, VIII.1865 Karsten (H). Kn: Paltamo,
Oikarila, deciduous litter, 22.IX.1974 E. & M.
Ohenoja (OULU F49344). Sotkamo, Kontin-
joki, Lauttolampi, decaying leaves of Popu-
lus, 10.IX.1986 Ulvinen (OULU F49275). OP:
Oulu, Toppila, Eteläsatama, deciduous litter,
1.X.1965 Ulvinen (OULU F49346). Ks: Po-
sio, Livojärvi, herb-rich forest, decaying leaves
of Prunus padus, 28.IX.1982 Ulvinen (OULU
F49350). InL: Utsjoki, Tsharsjoki, delta island,
decaying leaves of Betula sp. and Salix lanata,
15.VIII.1962 Kankainen (TUR 28665).
Typhula setipes (Grev.) Berthier
Syn. T. gyrans Fr., T. ovata P. Karst., T. pusilla
Fr.
V: Turku, Yliopistonmäki, decaying herbs,
11.X.1999 Huhtinen 88/110 (TUR 127939). U:
Helsinki, leaves of Salix fragilis, autumn 18??
Karsten 1304 (H). St: Noormarkku, Poosjoki,
decaying leaves of Populus, 25.IX.1937 Laurila
(H). EH: Tammela, Mustiala, decaying leaves of
Betula sp., 22.X.1868 Karsten 1479 (H), leaves
of Salix viminalis, 20.X.1891 Karsten 1480 (H).
Tampere, Koivistonkylä, decaying leaves of Be-
tula sp., 6.X.1987 Söderholm 1476 (H). KP: Pie-
tarsaari (Jakobstad), decaying leaves of Populus,
21.X.1862 Karsten 1301 (H). OP: Muhos, Lep-
piniemi, dead leaves of Prunus padus, 29.X.1974
E. Ohenoja (OULU F49291). Oulu, Korkeasaari,
decaying leaves of Alnus sp., 19.X.1969 Ulvin-
en (OULU F49287), Orikari, beach, decaying
leaves of Alnus sp., 24.IX.1965 Ulvinen (OULU
F49289, 49290).
*Typhula spathulata (Peck) Berthier
Syn. T. fruticum P. Karst.
OP: Oulu, decaying petioles and stems of Rubus
idaeus, 5.X.1965 Ulvinen (OULU F49285).
The type material of T. fruticum is missing
from H.
*Typhula struthiopteridis Corner
St: Kankaapää, Venesjärvi, Sinahmi, on decaying
petioles of Matteuccia struthiopteris, 1.IX.1936
Laurila (TUR 30807, 30811). EH: Tammela,
Haarankorpi, dead petioles of M. struthiopteris,
1.IX.1869 Karsten (H).
Typhula cf. subvariabilis Berthier
OP: Muhos, Muhosperä, Pekkala, dry herb-rich
forest among mosses, 14.VIII.1968 M. Ohenoja
(OULU F49296). KiL: Kittilä, Aakenus, Home-
vuotso Forest Res., old herb-rich spruce forest,
decaying leaves of Sorbus aucuparia, 12.IX.1983
Ulvinen (OULU F49303).
Typhula uncialis (Grev.) Berthier
Syn. Clavaria typhuloides Peck, Pistillaria peta-
sitites Imai
If otherwise not stated, the substrate is decaying
stem of Epilobium angustifolium.
V: Koski, Halikkola, 5.VII.2000 M-L. & P. Hei-
nonen 567-2000 (TUR 136891). Koski, Hongis-
to, 8.VII.1990 Heinonen 27-90 (TUR 101269),
31.VII.1993 Heinonen 61-93 F (TUR 112433).
Paimio, Kruuvainen, 22.VII.2002 M-L. & P.
Heinonen 16-2002 (TUR 160278). Parainen,
Kirjalaön, Trollgärda, 2.IX.2005 M-L. & P. Hei-
nonen 108-2005 (TUR 174325). Särkisalo, För-
by, 23.IX.1994 Huhtinen (TUR 111165). EK:
Sippola, Ruotila, 19.VII.1947 Kujala (OULU
F49268). EH: Kangasala, Lehtola, 27.VII.2006
Söderholm 3721 (TUR 178523). Korpilahti,
Oittila, 20.VIII.1986 Ulvinen (OULU F49327).
Luhanka, Lempää, 25.IV.2007 Shiryaev 7008
(SVER). Pälkäne, Mallasvesi, 18.VIII.1993
Kytövuori 93-292 (H). Sysmä, Onkiniemi,
Uotila, Epilobium sp., 11.VIII.1987 Haikonen
8730 (H). Tammela, Mustiala, 4.IX.1867 Kar-
sten 1288 (H), 10.X.1881 Karsten 1476 (H).
Tampere, Kaukajärvi, Selkämäki, 31.VII.1976
Söderholm 147 (H), Rusko, 28.VII.1985 Sö-
derholm 1202 (TUR 80222). ES: Kangasniemi,
18.IX.2004 Jakobson 4310 (H). Rantasalmi,
Kolkonpää, 8.VIII.1998 Huhtinen 98/60 (TUR
124413); EP: Vimpeli, Moskovankallio, Epi-
lobium sp., 18.VIII.1993 Jakobson (H); PH:
Äänekoski, Parantala. Aittoniemi, 17.VII.1981
Storbacka (OULU F49273). PS: Heinävesi,
Vihtari, Lehtomäki, Epilobium sp., 12.VIII.2003
Haikonen 22697 (H). Lapinlahti, Haminanhar-
ju, 17.VIII.1971 Pohjola 71-528 (TUR 76489).
PK: Ilomantsi, Mekrijärvi, 26.VIII.1996 M-L.
53
KARSTENIA 47 (2007) SHIRYAEV & KOTIRANTA: TYPHULA AND PISTILLARIA IN FINLAND
& P. Heinonen 68-96 (TUR 116902). KP: Haa-
pavesi, Mieluskylä, VIII.1970 Koskela (OULU
F49333). OP: Kiiminki, 19.VIII.1967 Ulvinen
(OULU F49337), limestone area, 4.X.1966
Ulvinen (OULU F49332). PeP: Kemi, Ajos,
19.VIII.1986 Kamula (OULU F49294). Rova-
niemi, Jaatila, 17.VIII.1999 E. Ohenoja (OULU
F46574). Rovaniemi, Tennilä, Vieno Viiri,
22.VII.1981 E. Ohenoja (OULU F49301). InL:
Utsjoki, 25.VIII.1964 Mäkinen (TUR 71527).
Russia. Murmansk Region. Ks: Salla, Kisajo-
ki, 22.VII.1937 Laurila (H).
Typhula variabilis Riess
U: Helsinki, Käpylä, decaying leaves of Acer
platanoides, 15.XII.2002 Shiryaev 7009 (SVER).
EH: Vilppula, Keuruu, Ukonselkä lake, decaying
Equisetum fl uviatile, 30.VII.1999 Järvinen 1641
(TUR 131531). OP: Kiiminki, Pöksälänkangas,
deciduous litter, 22.IX.1968 M. Ohenoja (OULU
F49347). Oulu, Hupisaaret, decaying leaves
of Prunus padus, 29.IX.1965 Ulvinen (OULU
F49343, TUR 30746). Ks: Kuusamo, Oulanka,
Liikasenvaara, rich brook-side forest, litter of
Betula and Salix, 24.VIII.1978 Ulvinen (OULU
F49321). InL: Utsjoki, Kevo, deciduous litter,
28.VIII.1973 Ulvinen (OULU F23322).
Notes on some species
Knudsen (1997) reports two species, viz. Typhu-
la muelleri (Sauter) Corner and Typhula trifolii
Rostrup to occur in Finland, but because no her-
barium material of them was available, we could
not confi rm their existence in Finland. Accord-
ing to Knudsen (1997), T. sclerotioides sensu
Karsten (1882) is T. muelleri. Karsten (1882)
reports T. sclerotioides to occur in Norway on
Gentiana purpurea, which does not belong to the
fl ora of Finland.
All the species new for Finland are known
also in adjacent areas in Russia (Archangel Re-
gion, Murmansk Region, Leningrad Region and
Karelian Republic) and the notes of their ecology
are largely based on that data. It must be noted
that many of the species range also much further
to the south (even semi-deserts), but here only
the biological requirements in boreal zone and
subarctic or subalpine areas are described.
Typhula capitata
A common boreal species with a holarctic dis-
tribution. It grows from hemiboreal zone in the
south to the subarctic/subalpine areas in the north
up to the Barents sea but is rare in the north. Its
habitat requirements are not especially strict, but
mostly it has been collected from southwestern
slopes (on Alopecurus, Calamagrostis) or an-
thropogenic sites (on Deschampsia). It prefers
wet, open habitats close to rivers or lake shores,
and has not been found inside coniferous forests.
Fruiting time in hemiboreal zone is October, and
in more northern areas from mid-August to the
fi rst frosts.
Typhula culmigena
A common boreal species with a holarctic dis-
tribution. It grows from hemiboreal zone in the
south to subarctic/subalpine areas in the north,
where it is not as common as in the boreal zones.
It prefers open, sunny sites, like coasts, mead-
ows, road sides etc., and has a very wide range of
substrates, if compared with the closely related
T. uncialis, which almost exclusively grows on
Epilobium. Typhula culmigena has been found
from herbs and grasses like Actaea, Angelica,
Cicerbita, Dactylis, Epilobium, Heracleum, Fes-
tuca and Secale and on dead leaves of Alnus,
Betula and Populus. Fruitbodies can be found
from spring to the fi rst snow cover.
Typhula hyalina
A common boreal species, known only in Eura-
sia. It is widely distributed from hemiboreal
zone to subarctic areas, where it has, however,
been found only twice from Carex species. In
subalpine area in Murmansk region on Khibiny
Mountains (Gory Hibiny) it grows on Carex spp.
and Juncus spp. but is not common. In boreal
Pinus forests it prefers Melica nutans, and in
Picea forests Carex spp., Juncus spp. and Dac-
tylis glomerata. The fruiting period in south bo-
real–north boreal zones is from late August to the
fi rst snow cover. In hemiboreal forests it infests
Allium, Bromus, Dactylis, Carex, Melica and
Juncus in wet localities. The fruiting starts in the
beginning of October.
Typhula spathulata
A common species from temperate zone to sub-
arctic areas, but is more common in the south.
In hemiboreal zone T. spathulata grows on twigs
54 KARSTENIA 47 (2007)
SHIRYAEV & KOTIRANTA: TYPHULA AND PISTILLARIA IN FINLAND
of several deciduous trees and bushes, like Acer
platanoides, Alnus glutinosa, A. incana, Cytisus,
Salix spp., Syringa spp., Quercus robur, Ulmus
spp. and Rubus idaeus. It favors western and
north-western, somewhat moist slopes and the
substrate is not covered by decaying leaves. The
fruiting period is from the mid-September to the
mid-October.
In southern boreal to northern boreal zones T.
spathulata prefers twigs of Alnus spp., Prunus
padus, Salix spp. and Rubus idaeus, but is never
collected from Populus, Betula spp. or Tilia cor-
data. The habitats are moist slopes, hill tops and
openings in coniferous forests. The fruiting pe-
riod begins at the end of August and continues
till the middle of September.
In subarctic areas T. spathulata is rare, but
grows on Alnus incana, Salix spp. and Rubus
idaeus. It prefers shady northern or north-west-
ern slopes and grows sometimes under leaf litter.
It has never been collected from bogs or perma-
nently wet habitats. The fruiting period is short,
from the middle to the end of August.
Typhula struthiopteridis
A common boreal species, and known at least
from Estonia, European part of Russia, Ural
Mountains, the Caucasus, and Siberia. In east-
ern Fennoscandia collected from hemiboreal and
southern boreal zones exclusively from dead
petioles of Matteuccia struthiopteris.
In hemiboreal zone it prefers humid hill tops,
and is never found close to water courses. In this
zone it is also more frequent than in the north and
the fruitbodies are larger. From southern boreal
zone northwards the habitats are normally conif-
erous forests, less often deciduous forests, and
it is very rare along brook-sides or lake shores.
Fruiting period begins at the end of August and
continues till the fi rst frosts.
Acknowledgements: The curators of the following her-
baria, H, TUR, TAA, SVER, OULU, and Esteri Ohenoja
(Oulu), are warmly thanked for the loans, as well as the
staff especially in the Botanical Museum, University of
Helsinki, for the kind help for the senior author during his
visit. The author Kotiranta thanks the Ministry of Environ-
ment for a research grant (PUTTE, YM131/5512/2002).
References
Berthier, J. 1976: Monographie des Typhula Fr., Pistil-
laria Fr. et genres voisins. – Bull. Mens. Soc. Linn.
Lyon 45: 1–233.
Hämet-Ahti, L., Suominen, J., Ulvinen, T. & Uotila, P.
(eds.) 1998: Retkeilykasvio. – Finnish Mus. Nat.
Hist., Bot. Mus., Helsinki. 656 pp. (In Finnish).
Hansen, L. & Knudsen, H. (eds.) 1997: Nordic acromyce-
tes 3. Heterobasidioid, aphyllophoroid and gastromy-
cetoid Basidiomycetes. – Nordsvamp, Copenhagen.
444 pp.
Heikinheimo, O. & Raatikainen, M. 1981: Ruutukoor-
dinaattien ja paikannimien käyttö Suomessa. (Grid
reference and names of localities in the recording of
biological fi nds in Finland.) – Notulae Entomol. 61:
133–154.
Jamalainen, E. A. 1957: Overvintering of Gramineae-
plants and parasitic fungi II. On the Typhula sp. fungi
in Finland. – J. Sci. Agric. Soc. Finland 29: 75–81.
Karsten, P. A. 1879: Symbolae ad Mycologiam Fennicam
6. Basidiomycetes De B. – Medd. Soc. Fauna Flora
Fennica 5: 15–46.
Karsten, P. A. 1881a: Fungi novi, lecti atqve descripti a P.
A. Karsten. – Hedwigia 20: 177–179.
Karsten, P. A. 1881b: Hymenomycetes Fennici. – Acta
Soc. Fauna Flora Fennica 2(1): 1–40.
Karsten, P. A. 1881c: Enumeratio Boletinearum et Po-
lyporearum Fennicarum, systemate novo disposita-
rum. – Rev. Mycol. 3(9): 16–19.
Karsten, P. A. 1882: Rysslands, Finlands och den Skan-
dinaviska Halföns hattsvampar 2. Pip-, tagg-, hud-,
klubb- och gelésvampar. – Bidrag Känned. Finlands
Natur Folk 37(9): 1–257.
Karsten, P. A. 1899: Finlands basidsvampar. Floristiska
handböcker för nybegynnare. – Soc. Fauna Flora Fen-
nica, Helsingfors Centraltryckeri. Helsingfors. 186
pp., 9 pls.
Knudsen, H. 1997: Typhulaceae Jülich. – In: Hansen, L.
& Knudsen, H. (eds.), Nordic macromycetes 3. Hete-
robasidioid, aphyllophoroid and gastromycetoid Basi-
diomycetes, 256–260. Nordsvamp, Copenhagen.
Mäkelä, K. & Mäki, L. 1980: The occurrence of micro-
mycoglora in the stem base and roots of cereals in
southern Finland. – Ann. Agric. Fenniae 19: 87–222.
Ohenoja, E. 1996: A check-list of the larger fungi in Inari
Lapland (NE Finland) and in Finnmark (NE Norway).
– Kevo Notes. 11: 1–26.
Ulvinen, T., Ohenoja, E., Ahti, T. & Alanko, P. 1981: A
check-list of the fungi (incl. lichens) of the Koillis-
maa (Kuusamo) biological province, N.E. Finland.
– Oulanka Reports 2: 1–64.
Karstenia 47: 55–59, 2007
Notes on some rare polypores, found in Russia 3.
Genera Anomoloma, Hyphodontia, Lindtneria, and
Sistotrema
WJACHESLAV A. SPIRIN and IVAN V. ZMITROVICH
SPIRIN, W. A. & ZMITROVICH, I. 2007: Notes on some rare polypores, found in Rus-
sia 3. Genera Anomoloma, Hyphodontia, Lindtneria, and Sistotrema. – Karstenia 47:
55–59. Helsinki. ISSN 0453-3402.
Anomoloma albolutescens (Romell) Niemelä & K.H. Larss. is found for the fi rst time
in European Russia. Ecology and distribution of this species are briefl y reviewed. Poria
buxi Bondartsev is a later synonym of Hyphodontia fl avipora (Cooke) Sheng H. Wu.
Hyphodontia radula (Schrad. : Fr.) E. Langer & Vesterholt was found to be widely dis-
tributed in hemiboreal European Russia. Lindtneria trachyspora (Bourdot & Galzin)
Pilát and Sistotrema dennisii Malençon are reported as new to Russia. Sistotrema brun-
neolum Spirin & Zmitr. is described as a new species closely related to S. alboluteum
(Bourdot & Galzin) Bondartsev & Singer. The new species is characterized by relatively
fi rm, brownish, porioid basidiocarps, two- or four-sterigmate basidia, and short-cylindri-
cal to ellipsoid, thick-walled spores. The isolated position of both S. brunneolum and S.
alboluteum within the genus Sistotrema is discussed.
Key words: corticioid fungi, polypores, virgin forest.
Wjacheslav Spirin, St. Petersburg University of Humanities, Fuchika 15, 192238
St.Petersburg, Russia
Ivan Zmitrovich, Komarov Botanical Insitute RAS, Popova 2, 197376 St. Petersburg,
Russia
Introduction
This paper is devoted to studies on some poly-
pores closely related or belonging to the genera
where most of the species have smooth, tubercu-
late or odontioid (i.e. non-poroid) hymenophore,
or the so-called corticioid fungi. Most of the spe-
cies treated here are extremely rare, ephemeral,
and therefore data on their ecology and substrate
preferences are very scanty. A new porioid Sis-
totrema species, S. brunneolum, is described.
Material and methods
The microscopic routine used in this study was described
in Spirin and Zmitrovich (2003). The material is depos-
ited in the mycological herbarium of the Botanical Muse-
um, University of Helsinki, Finland (H) and in Komarov
Botanical Institute, St. Petersburg, Russia (LE). The plant
communities of the research area were described by Spi-
rin (2005), and the polypore names are given according to
Niemelä (2005) and Niemelä et al. (2007).
Species descriptions
Anomoloma albolutescens (Romell) Niemelä &
K.H. Larss.
Syn. Anomoporia albolutescens (Romell)
Pouzar
First Russian fi nds of this very rare polypore
were reported from Altai Region (Bondartseva
1972), and for more than 30 years A. albolute-
scens remained known in the country only from
Siberia. Our record treated below is the fi rst one
for European Russia. The species was collected
in very old (over 200 years old) southern-taiga
56 KARSTENIA 47 (2007)
SPIRIN & ZMITROVICH: RARE RUSSIAN POLYPORES
forest, dominated by Picea abies, Abies sibirica,
and Tilia cordata. The substrate was very rotten
spruce trunk with a large cavity inside; intensive-
ly developing fruitbodies were seen there.
The present record agrees well with current
species concept (Niemelä 1994). Basidiospores
are (3.9–) 4.1–5.2 (–5.4) × 3.1–3.5 (–3.6) m,
ellipsoid, with fl at ventral side, amyloid, acy-
anophilous.
Anomoloma albolutescens is very rare ev-
erywhere, and found in Sweden (type locality,
Romell 1911), Norway, Finland, Russian Far
East, Kazakhstan, China, and North America
(Niemelä 1994).
Specimens examined: Russia. Nizhny
Novgorod Reg.: Sharanga Dist., Kilemarsky
Nat. Res., Picea abies, 17.VIII.2004 Spirin 2137
(H, LE, W.S.). Finland. Etelä-Häme: Vesijako
Strict Nat. Res., P. abies, 9.VIII.1981 Kotiranta
2886 (H).
Hyphodontia fl avipora (Cooke) Sheng H. Wu
Syn. Schizopora fl avipora (Cooke) Ryvarden
Poria buxi was described as a new species
by Bondartsev (1940), and was later combined
by him into the genus Aporpium Bondartsev &
Singer (Bondartsev 1953). For a long time, this
name remained forgotten: both Donk (1974) and
Ryvarden (1991) gave no data on a contempo-
rary position of this taxon. Bondartseva (1998)
was of opinion that Poria buxi is identical to Oli-
goporus sericeomollis (Romell) Bondartseva.
Type specimen (LE 25311) consists of two
pieces of porioid fungus with corky, buff-co-
loured tubes; pores are angular, ca. 4–6 per mm.
Microscopically, this specimen is characterized
by clamped narrow hyphae bearing swollen api-
ces at the dissepiment edges; basidiospores are
ellipsoid, (4.0–) 4.1–5.2 (–5.3) × (2.6–) 2.7–3.4
(–3.6) m, inamyloid. Our conclusion is that it
belongs to Hyphodontia fl avipora (Cooke) Sheng
H. Wu, which is widely distributed in southern
Europe.
Selected specimens: Russia. Krasnodar Reg.:
Hosta, Buxus sempervirens, 20.IX.1938 Va s i -
lyeva (LE 25311, holotype of Poria buxi). Ni-
zhny Novgorod Reg.: Arzamas Dist., Pustynsky
Nat Res., Quercus robur, 2.VII.2000 Spirin (LE
213649); Lukoyanov Dist., Razino, Betula pube-
scens and Q. robur, 9 – 11.VII.2005 Spirin 2362,
2385 (H), Sanki, Quercus robur, 18.VIII.2006
Spirin 2613 (H); Sharanga Dist., Kilemarsky
Nat. Res., Tilia cordata, 22.VIII.2004 Spirin
2273 (H).
Hyphodontia radula (Schrad. : Fr.) E. Langer &
Vesterholt
Syn. Schizopora radula (Schrad. : Fr.) Hallen-
berg
This species was not known in Russia for a
long time (Bondartseva 1998). First records were
published by Niemelä et al. (2001) and Spirin
(2002). Hyphodontia radula has a remarkably
wide distribution in hemiboreal zone in Russia,
preferring broad-leaved forests; its main sub-
strate is Quercus robur.
Modern descriptions of this species were pub-
lished by Niemelä (1987) and Vampola (1990).
These authors questioned the miticity of the
basidiocarps as a distinguishing character be-
tween H. radula (‘monomitic’) and H. paradoxa
(‘dimitic’), which agrees well with our results.
Evidently, the dominance of thick-walled skel-
etal-like hyphae depends strongly on environ-
mental conditions. The specimens collected in
exposed habitats have clearly ‘dimitic’ construc-
tion, with dominating sclerifi ed hyphae without
clamps. The best differentiating character of H.
radula is the spore length, not exceeding 5.5
m (the measurements must be made in Cotton
Blue).
Selected specimens: Hyphodontia radula.
Russia. Nizhny Novgorod Reg., Lukoyanov
Dist., Sanki, Quercus robur, 7.VIII.2005 Spirin
2310 (H); Kurley, Q. robur, 13.VIII.2005 Spi-
rin 2421 (H); Nizhny Novgorod City, Q. robur,
29.X.2000 (LE 213128). – Hyphodontia para-
doxa. Russia. Nizhny Novgorod Reg., Lukoya-
nov Dist., Kurley, Q. robur, 13.VIII.2005 Spirin
2425 (H).
Lindtneria trachyspora (Bourdot & Galzin)
Pilát
This rare fungus is known from North Amer-
ica and some European countries (Gilbertson &
Ryvarden 1986, Ryvarden & Gilbertson 1993);
here it is published as new to Russia. It is eas-
ily identifi ed in the microscope due to coarsely
ornamented brownish spores 6.8–8.2 (–8.5) ×
6.4–7.6 (–8.0) m, strongly cyanophilous in ju-
venile state. Basidia of L. trachyspora are very
large, suburniform or bladder-shaped, and com-
prise numerous strongly cyanophilous granules.
This feature links Lindtneria with members of
57
KARSTENIA 47 (2007) SPIRIN & ZMITROVICH: RARE RUSSIAN POLYPORES
the corticioid genus Cristinia Parmasto (Jülich
1982).
Specimen examined: Russia. Primorye Reg.:
Khasansky Dist., Kedrovaya Pad’ Nat. Res.,
Pinus koraiensis, 21.VIII.2005 Psurtseva (LE
214902).
Sistotrema alboluteum (Bourdot & Galzin)
Bondartsev & Singer
This rare saprotrophic fungus was found
in Russia quite recently (Kotiranta & Mukhin
1998). To the fi rst locality from Kamchatka (Rus-
sian Far East), we add here another record from
Nizhny Novgorod Region (European Russia).
S. alboluteum produces bright-yellow, small,
porioid basidiocarps on strongly decomposed
wood of both conifers and angiosperms, or even
on debris. In the microscope it is easily identifi ed
due to the 2–4-sterigmate urniform basidia and
large, thick-walled, subglobose spores, 4.6–6.3
m in diam. See notes to S. brunneolum.
Specimens examined: Russia. Nizhny
Novgorod Reg.: Bor Dist., Kerzhensky Nat.
Res., very rotten Betula, 24.IX.1998 Spirin (LE
212644, H).
Sistotrema brunneolum Spirin & Zmitr., spec.
nova. – Fig. 1.
Carpophorum resupinatum, porioideum, fra-
gile, colore pallido-ochraceum vel brunnescens.
Systema hypharum monomiticum; hyphae fi bula-
tae. Basidia urniformia, cum 2 – 4 sterigmatibus.
Sporae crassitunicatae, brevicylindriceae vel el-
lipsoideae, 4.6 – 6.7 × 2.5 – 2.9 μm.
Holotype: Russia. Nizhny Novgorod Reg.:
Sharanga Dist., Kilemarsky Nat. Res., Populus
tremula, 17.VIII.2004 Spirin 2127 (LE 215000,
isotype in H).
Etymology: brunneolum (Lat., adj.) – with
brown hues in its colour.
Basidiocarps short-living annual, resupinate,
widely effused (up to 15 cm in longest dimen-
sion), soft-waxy and fragile in fresh condition,
soft-corky and relatively hard when dry. Margin
distinct, fi rm (of cardboard-like consistence),
cream to pale-ochraceous or even brownish, 2–4
mm wide; rhizomorphs absent. Pore surface at
fi rst cream to pale ochraceous, later resinous
brown (especially marginal areas), even; pores
angular, sometimes merging together, lacerate
and strongly elongated on sloping substrate,
(3–) 4–6 per mm, with thin, entire or minutely
denticulate orifi ces. Section: subiculum distinct,
white or pale-cream, soft-corky, 1–2 mm thick;
tubes concolorous with pore surface, soft-waxy
when fresh, hardening in dry state, 1–5 mm
thick. Odour faint, fungoid; taste mild.
Hyphal structure monomitic; all hyphae thin-
walled, with abundant clamps, moderately cy-
anophilous.
Subiculum: Hyphae hyaline, richly branched
and abundantly septate with clamp connections,
irregularly arranged, 3.5–7 m wide, some hy-
phal cells with numerous oily droplets.
Tubes. Hyphae hyaline, branched at sharp
angles, with abundant septa and large clamps,
sometimes with greasy drops, 2.5–4 m wide;
trama irregular to subparallel. No cystidia. Ba-
sidia urniform, large, 15–22 × 4.5–6 (–7) m,
with 2 or 4 sterigmata (up to 6 m long), content
homogeneous. Basidioles of the same size as ba-
sidia, bladder-shaped.
Fig. 1. Sistotrema brunneolum: hymenium and spores.
Scale bar = 10 µm.
58 KARSTENIA 47 (2007)
SPIRIN & ZMITROVICH: RARE RUSSIAN POLYPORES
Spores: Basidiospores short-cylindrical to nar-
rowly ellipsoid, with fl at or slightly rounded ven-
tral side, (4.3–) 4.6–6.7 (–7.3) × (2.3–) 2.5–2.9
(–3.4) m, thick-walled, inamyloid, moderately
cyanophilous.
Distribution and ecology: – Nowadays S.
brunneolum is known from the type locality only.
The site represents old undisturbed southern taiga
forest where Anomoloma albolutescens was col-
lected, too (see above). The species was found on
still standing dead trunk of Populus tremula. Evi-
dently, this species is ephemeral and appears only
in rainy seasons. Visually, the rot produced by S.
brunneolum is white; however, this fact must be
confi rmed with cultural studies.
Identifi cation and taxonomy: – Undoubtedly,
our new species is very close to S. alboluteum.
The reasons to describe a new species are the
peculiar macromorphology and disagreeing
spore dimensions. Superfi cially, S. brunneolum
is strongly reminiscent of Ceriporiopsis species,
especially C. resinascens (Romell) Domaski.
Common features are ochraceous-brownish co-
lours of pore surface contrasting with paler mar-
gin, and the number of pores per mm. Unlike Sis-
totrema alboluteum, our new species is relatively
fi rm, and has no rhizomorphs. However, in the
microscope these two species are very similar,
but differ clearly by spore dimensions: globose
to subglobose in S. alboluteum (Q = 1.0–1.13)
versus short-cylindrical to ellipsoid in S. brun-
neolum (Q = 1.87–2.1). Two other porioid Sis-
totrema species known in Europe, S. muscicola
(Pers.) S. Lundell and S. dennisii Malençon, dif-
fer in having 6 – 8-sterigmate basidia and much
smaller spores.
Sistotrema alboluteum and S. brunneoum
share an isolated position within the genus due
to constantly porioid basidiocarps, cyanophilous
hyphae, large basidia with two or four sterigma-
ta, and thick-walled cyanophilous spores. A few
Sistotema species have cyanophilous hyphae and
spores; this character links them with the genus
Botryobasidium Donk (Jülich & Stalpers 1980,
Kotiranta & Saarenoksa 1990). Probably, the cur-
rent generic concept of Sistotrema is very wide,
and splitting into more homogeneous unities
might be welcome. Unfortunately, a very small
number of the Sistotrema species are included in
molecular studies (Larsson et al. 2004, Binder
et al. 2005), and therefore we cannot make any
clear conclusions.
Sistotrema dennisii Malençon
New to Russia. This very rare species is found in
some European countries and North Africa (Ry-
varden & Gilbertson 1994, Niemelä et al. 2001).
Our specimen was collected on fallen, decorti-
cated Picea abies, and is characterized by 4–6–8-
spored, urniform basidia, and short-cylindrical,
slightly curved spores, 3.1–4.3 × 2.1–2.6 m. Hy-
phae are clamped and almost acyanophilous.
Specimens examined: Russia. Nizhny
Novgorod Reg.: Sharanga Dist., Kilemarsky Nat.
Res., Picea abies, 17.VIII.2004 Spirin 2123 (LE,
H). Finland. Etelä-Häme: Janakkala, P. abies,
8.IX.2000 Salonen 6 (H).
Acknowledgments: We are very grateful to Prof. Tuomo
Niemelä (Helsinki) for reviewing of the manuscript and
valuable remarks on it. Prof. Teuvo Ahti (Helsinki) kindly
revised the Latin. Also we would like to thank Dr. Na-
dezhda Psurtseva (Komarov Botanical Institute, St. Pe-
tersburg, Russia), who furnished us with a specimen of
Lindtneria trachyspora.
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