Polish Botanical Journal 53(1): 81–90, 2008
PHLEOGENA FAGINEA (PUCCINIOMYCOTINA, ATRACTIELLALES)
IN POLAND – NOTES ON ECOLOGY AND DISTRIBUTION
ANDRZEJ SZCZEPKOWSKI, ANNA KUJAWA, ANNA BUJAKIEWICZ, JOANNA NITA,
DARIUSZ KARASIŃSKI, MAREK WOŁKOWYCKI & MARCIN S. WILGA
Abstract. The current distribution of Phleogena faginea (Fr.: Fr.) Link in Poland is presented, based on literature data as well as
19 new localities. Six tree species and one fungus as a new substrate for P. faginea in Poland are reported. These new records
confirm the high ecological plasticity of the species, which the authors found in natural stands, managed forests and an urban
park. The ecological preferences of the species are discussed. Wood of Carpinus betulus is the most frequent type of substrate for
P. faginea in Poland. Based on the new distribution data for P. faginea in Poland, a change of its red list category is proposed.
Key words: Phleogena faginea, fungi, Atractiellales, Phleogenaceae, ecology, distribution, Poland
Andrzej Szczepkowski, Department of Mycology and Forest Phytopathology, Warsaw University of Life Sciences – SGGW,
Nowoursynowska 159, PL-02-776 Warsaw, Poland; e-mail: andrzej_szczepkowski@sggw.pl
Anna Kujawa, Field Station of the Research Center for Agricultural and Forest Environment, Polish Academy of Sciences, Turew,
Szkolna 4, PL-64-000 Kościan, Poland; e-mail: annakuja@poczta.onet.pl
Anna Bujakiewicz & Joanna Nita, Department of Plant Ecology and Forest Environmental Protection, Adam Mickiewicz University, Umultowska 89, PL-61-614 Poznań, Poland, e-mail: ascom@amu.edu.pl & malav@wp.pl
Dariusz Karasiński, Department of Mycology, W. Szafer Institute of Botany, Lubicz 46, 31-512 Kraków, Poland; e-mail: darek_
karasinski@op.pl
Marek Wołkowycki, Sacharewo 1, PL-17-200 Hajnówka, Poland; e-mail: wolkm@poczta.onet.pl
Marcin S. Wilga, Automotive and Heavy Machinery Division, Gdańsk University of Technology, Narutowicza 11/12, PL-80-952
Gdańsk-Wrzeszcz, Poland; e-mail: mwilga@mech.pg.gda.pl
INTRODUCTION
Phleogena faginea (Fr.: Fr.) Link is the only member
of the genus Phleogena Link, belonging to the subphylum Pucciniomycotina, class Atractiellomycetes, order Atractiellales and family Phleogenaceae
(Bauer et al. 2006). It is widely distributed in the
temperate and tropical climatic zones. It is known
from Australia and New Zealand, South America,
North America, Asia, and Europe. Its general distribution, taxonomy, morphology and ecology are
described by Wojewoda and Komorowska (1997)
and Wojewoda et al. (1999).
Phleogena faginea is a saprotroph growing
on bark, in bark crevices, or directly on wood of
stumps, trunks and branches of dying or dead (rarer
on living) broadleaved, occasionally coniferous,
trees and shrubs. In Poland it has been noted on
Acer platanoides, Alnus glutinosa, Betula pendula, Carpinus betulus, Corylus avellana, Fagus
sylvatica, Picea abies and Populus tremula, and
on basidiomes of the fungus Fomes fomentarius
(Wojewoda 1977, 2003; Wojewoda & Komorowska
1997; Wojewoda et al. 1999). In other European
countries it is also known from Acer campestre,
Betula pubescens, Carya sp., Fraxinus sp., Ilex
sp., Populus sp., Prunus spinosa, Pyrus communis,
Quercus sp., Salix cinerea and Ulmus sp., and also
from basidiomes of Inonotus obliquus (Kreisel 1987;
Cook 1994; Wojewoda & Komorowska 1997; Holec
2003; Storey 2004). Beyond Europe it was observed
also on Dysoxylon spectabile, Ficus sp., Inga vera,
Podocarpus dacrydioides and Salvadora oleoides
(McNabb 1964; Wojewoda & Komorowska 1997;
López & García 2001). In Brazil it was collected
on construction wood of Cedrela sp. (Möller 1895,
after Oberwinkler & Bauer 1989).
Despite its wide distribution across the world
82
POLISH BOTANICAL JOURNAL 53(1). 2008
and its broad substrate spectrum, P. faginea is a rare
species in many European countries. Only one locality is known from Slovenia (Jurc et al. 2005) and
Norway (Anonymous), 11 localities from the Czech
Republic (Holec 2003), 12 from western Germany
(Krieglsteiner 1991), 13 from Slovakia (Ripková et
al. 2004) and 15 from Denmark (Gøtzsche 1980).
It is uncommon in Great Britain but probably
overlooked (Cook 1994). In some countries such
as Germany (Benkert et al. 1992; Schnittler 1996),
Sweden (Gärdenfors 2000) and Poland (Wojewoda
& Ławrynowicz 1986, 1992, 2006) the fungus is
on the red list of threatened fungi.
MATERIAL AND
METHODS
The current distribution of P. faginea in Poland is described and mapped on the basis of literature data as
well as unpublished information we gathered. In areas
of the frequent occurrence of P. faginea (within the borders of one forest complex), a forest section is considered a single locality. To characterize the substrate of
P. faginea and to determine whether the species prefers
older (thicker) trees, the circumference of standing tree
trunks colonized by the fungus was measured at 1.3 m
height, and that of lying trees at the site of basidiomes.
Specimen descriptions are based on the collected material. Basidiomes were measured fresh, and spores and basidia were taken from dried specimens (WAML 203, 211,
217, 220, 224, 228, 229; MW 1614) and measured in 5%
KOH. Acronyms of herbaria are according to Mirek et al.
(1997). The following abbreviations are used for herbaria
not mentioned by those authors: WAML – Herbarium of
the Department of Mycology and Forest Phytopathology
of the Warsaw University of Life Sciences – (SGGW),
DK – private herbarium of Dariusz Karasiński, MW –
private herbarium of Marek Wołkowycki.
Plant names are given after Mirek et al. (2002),
and the names of geographic regions used in the list of
Polish localities follow Kondracki (2002).
DESCRIPTION
OF SPECIMENS AND LOCALITIES
Phleogena faginea (Fr.: Fr.) Link
Figs 1a–d, f & g; Table 1
Handb. Gewächse 3(1): 396. 1833.
Basidiomes typically grow in rows or form
irregular aggregations of even several hundred
individuals; small, up to 12 mm tall, with head
and stalk resembling the sporangia of some
myxomycetes. Young basidiomes club-shaped
with weakly differentiated head and stalk. Head
1–5 mm in diameter, ± circular, spherical, sometimes slightly flattened, less often phaseoliform,
whitish, ochraceous, greyish, and in mature specimens brownish. Stalk up to 8 mm long and 1.5
mm thick, whitish, becoming brown to blackish
with age. Context elastic, then dry with a distinct
smell resembling lovage or maggi sauce persisting
also in herbarium specimens. Basidia cylindrical
and partitioned by 1–4 transversal septa, 16–40 ×
2.5–5 μm. Spores spherical or ± spherical, thickwalled, smooth, ochraceous to light brown in mass,
4–11 × 4–10 μm.
NEW LOCALITIES. POJEZIERZE POŁUDNIOWOPOMORZielona Góra Reserve near Wyrzysk,
Kaczory forest district, sect. 99 (53°07′N/17°15′E),
Galio sylvatici-Carpinetum, on log of deciduous tree
(Quercus sp. or Carpinus betulus), 07 Oct. 2006, leg.
R. Puciata (WAML 227). POBRZEŻE GDAŃSKIE OUTSKIRTS: Zajęcze Wzgórze Reserve, Gdańsk forest district, Sopot forest range, sect. 47 (54°26′N/18°33′E),
Luzulo pilosae-Fagetum, on barkless fragment of Fagus
sylvatica log, 7 Jan. 2007, leg. W. Półtorak (WAML
228); Gdańsk forest district, Renuszewo forest range,
sect. 118, Zielona Dolina valley (54°23′N/18°33′E),
Stellario holosteae-Carpinetum betuli, on rotten log
of Carpinus betulus, 25 Aug. 2006, leg. M. S. Wilga
(WAML 230), on dead standing trunk of Fagus sylvatica, 16 Feb. 2007, leg. M. S. Wilga; Gdańsk forest
district, Matemblewo forest range, sect. 121c, Samborowo valley (54°23′N/18°33′E), Stellario holosteaeCarpinetum betuli, on barkless dead standing trunk of
Quercus robur (Fig. 1b), Sept. 2006, leg. M. S. Wilga
(WAML 229). WZNIESIENIA POŁUDNIOWOMAZOWIECKIE
HEIGHTS: Rogów forest district, Popień forest part, sect.
171d (51°46′37″N/19°55′35″E), mixed forest, on fallen
trunk of Populus tremula, 18 Sept. 2006, leg. A. Kujawa
(WAML 201), on dead standing trunk of Alnus glutinosa, leg. J. Piętka (WAML 202), on log of Alnus glutinosa, leg. A. Szczepkowski (WAML 203) (http://www.
grzyby.pl/gatunki/Phleogena_faginea.html). BESKIDY
ŚRODKOWE MTS: Magurski National Park, SE slope
of Góra Żydowska Mt. (49°27′45.6″N/021°31′41.2″E,
alt. 573 m), Dentario glandulosae-Fagetum, on wood
and bark of dead standing Ulmus glabra trunk, 28
Oct. 2005, leg. D. Karasiński (DK 051028/0712); SE
slope of Kamień Mt. (49°31′09.6″N/021°32′23.8″E, alt.
SKIE LAKELAND:
A. SZCZEPKOWSKI ET AL.: PHLEOGENA FAGINEA IN POLAND
83
Fig. 1. Phleogena faginea (Fr.: Fr.) Link. a – young club-shaped basidiomes on a trunk of Carpinus betulus in sect. 398 of the
Białowieża National Park, b – well-developed basidiomes on trunk of a monumental dead oak in the Gdańsk forest district
(WAML 229), c – basidiomes growing directly on wood of Ulmus sp. log and on rhizomorphs of Armillaria sp. in Hajnówka
forest district, d – basidiomes growing on bark and directly on wood of a dead Ulmus laevis trunk, and on rhizomorphs of
Armillaria sp. in the Morysin Reserve (WAML 220), e – Acer platanoides colonized by P. faginea in Łazienki Park, f – mature
basidiomes in bark crevices on trunk of Padus avium in the Morysin Reserve (WAML 214), g – basidiomes growing directly
on wood, on lower side of fallen Carpinus betulus log in Las Bielański Reserve (WAML 205). Photo: D. Majgier (a, f, g), M.
Snowarski (c), A. Szczepkowski (d, e) and M. S. Wilga (b).
383 m), Alnetum incanae, on dead standing Alnus incana
trunk, 11 Nov. 2006, leg. D. Karasiński (DK 061111/01)
(http://grzyby.strefa.pl/Phleogena_faginea.html). NIZINA
Ś RODKOWOMAZOWIECKA LOWLAND : Warsaw, Las
Bielański Reserve, sect. 8 (52°17′38.4″N/20°57′57.2″E),
Tilio-Carpinetum, on trunk of dying Carpinus betulus, 29
Dec. 2006, leg. A. Szczepkowski (WAML 223), on remains
of standing trunk of Carpinus betulus, 29 Dec. 2006,
leg. A. Szczepkowski, on trunk of dying Alnus glutinosa,
29 Dec. 2006, leg. A. Szczepkowski, Ficario-Ulmetum
84
POLISH BOTANICAL JOURNAL 53(1). 2008
Table 1. Comparison of some characters of Phleogena faginea (Fr.: Fr.) Link from several descriptions.
Basidiome dimensions [mm]
Reference
total height
head
stalk
Basidiospores
[μm]
Basidia
[μm]
Pilát 1956
3–6
1–3
–
6–8 × 5.5–7.5
26–30 × 5
McNabb 1964
4–7
1–3
–
8–10 × 7.5–9
25–37 × 4–5
Tallasch & Jahn 1970
2–5
1–3
2–3 × 0.2–0.3
6–8 × 5.5–7.5
–
Wojewoda 1977, 1981
2–11
1–4–(5)
2–8 × 0.2–1.5
4.5–10 × 4.5–10
16–37 × 4–5
Cook 1994
2–8
1–4
1–4 × 0.2–0.3
(4.5–)6–11
× (4–)5.5–7.5(–9)
(16–)18–27(–37)
× (2.5–)3–4(–5)
Character
Hansen & Knudsen 1997
3–6
–
1–3
8–10
25–30 × 4–5
Holec 2003
up to 7
1–3
1.5–4 × 0.2 –1
5.5–8 × 5–6.5
30–40 × 4–5
Ripková et al. 2004
up to 8
1–4
1–4 × 0.2–1
4–7.5 × 4–8
–
2–12
1–5
1–8 × 0.2–1.5
(5–)6–9(–10)
× (4.5–)5–8(–9)
18–35 × 3–4.5
Present studies
minoris, on stump of Ulmus laevis, 29 Dec. 2006, leg.
A. Szczepkowski (WAML 224), on remains of stump
of Alnus glutinosa, 29 Dec. 2006, leg. A. Szczepkowski
(WAML 225); sect. 9 (52°17′17.7″N/20°58′12.4″E),
Tilio-Carpinetum, on fallen trunk of Carpinus betulus,
21 Sept. 2006, leg. D. Majgier (WAML 205) (Fig. 1g),
on two fallen trunks of Carpinus betulus, 29 Dec.
2006, leg. A. Szczepkowski (WAML 222), on two dead
standing trunks of Carpinus betulus, 29 Dec. 2006, leg.
A. Szczepkowski (WAML 221); sect. 12, Tilio-Carpinetum, on one dead standing Carpinus betulus trunk and
on six fallen Carpinus betulus trunks, 21 Oct. 2006,
leg. M. Kozłowski & A. Szczepkowski (WAML 226);
Warsaw, Łazienki Park (52°12′46.1″N/21°01′59.3″E)
on dead bark of living Carpinus betulus trunk, 17 Nov.
2006, leg. D. Majgier (WAML 206), on dead bark and
wood of four living standing Carpinus betulus trunks,
18 Nov. 2006, leg. A. Szczepkowski (WAML 208), on
dead bark of dying Acer platanoides trunk (Fig. 1e),
17 Nov. 2006, leg. A. Szczepkowski (WAML 207);
Warsaw, Morysin Reserve (52°10′34″N/21°55′35″E),
Ficario-Ulmetum minoris, on decorticated branch of
living Alnus incana, 17 Nov. 2006, leg. A. Szczepkowski
(WAML 211), on trunks of three standing broken Alnus
incana, 29 Nov. 2006, leg. A. Szczepkowski (WAML
216, 218), on four logs of Alnus incana, 17. Nov. 2006,
leg. A. Szczepkowski (WAML 212, 213), 29 Nov. 2006,
leg. A. Szczepkowski (WAML 219), on dead bark of
living standing Ulmus laevis trunk, 17 Nov. 2006, leg.
A. Szczepkowski (WAML 210), on dead standing Ulmus
laevis trunk, 17 Nov. 2006, leg. A. Szczepkowski (WAML
209), on dead bark and wood of standing trunk of Ulmus
laevis and also on rhizomorphs of Armillaria sp., 29
Nov. 2006, leg. A. Szczepkowski (WAML 220), on dead
standing broken Ulmus laevis trunk, 17 Nov. 2006, leg.
A. Szczepkowski (WAML 215), on dead standing Padus
avium trunk, 2 Nov. 2006, leg. D. Majgier (WAML 214)
(Fig. 1f), on bark and wood of dead fallen Padus avium
trunk, 29 Nov. 2006, leg. A. Szczepkowski (WAML
217). NIZINA PÓŁNOCNOPODLASKA LOWLAND: Puszcza
Białowieska primeval forest, Hajnówka forest district,
sect. 272 (52°45′N/23°38′E), Tilio-Carpinetum melittetosum, on log of Ulmus sp., directly on wood and on
rhizomorphs of Armillaria sp. (Fig. 1c), 26 Sept. 2006,
leg. M. Snowarski, D. Karasiński & M. Piątek; sect.
412A (52°42′N/23°37′E), managed tree stand growing
in Ribeso nigri-Alnetum, on dead branch of standing Corylus avellana, 20 Feb. 2002, leg. M. Wołkowycki (MW
2244); sect. 416C (52°42′N/23°41′E), managed tree
stand growing in Tilio-Carpinetum melittetosum, on old
stump of Picea abies, 25 Oct. 2006, leg. M. Wołkowycki
(MW 3400); sect. 464D (52°41′N/23°39′E), managed
tree stand growing in Tilio-Carpinetum circaeaetosum alpinae, on dead lying branch of Tilia cordata
and also on rhizomorphs of Armillaria sp., 2 Sept.
2001, leg. M. Wołkowycki (MW 1614); sect. 488C
(52°40′N/23°41′E), managed tree stand growing in
Tilio-Carpinetum, on dead, standing Alnus glutinosa
trunk, 17 Nov. 2000, leg. M. Wołkowycki (MW 1767);
sect. 516D (52°40′N/23°41′E), managed tree stand
growing in Querco-Piceetum, on log of Betula pendula,
11 Oct. 2001, leg. M. Wołkowycki (MW 2133); Sach-
A. SZCZEPKOWSKI ET AL.: PHLEOGENA FAGINEA IN POLAND
arewo hamlet (52°41′N/23°39′E), on Acer platanoides
wood for fuel, 10 Feb. 1998, leg. M. Wołkowycki (MW
1149); Białowieża National Park, Strict Protection Area,
along educational trail in sect. 398, on fallen trunk of
Populus tremula, 25 Sept. 2006, leg. A. Szczepkowski
(WAML 204), on two standing trunks of Carpinus betulus (Fig. 1a), 25 Sept. 2006, leg. A. Szczepkowski, on
two dead standing trunks of Alnus glutinosa, 25 Sept.
2006, leg. A. Szczepkowski.
LITERATURE
DATA .
POJEZIERZE ZACHODNIOPOPerłówkowe Buki Reserve
near Stargard Szczeciński, Sept. 1995, leg. H. Komorowska (KRAM-F) (Wojewoda & Komorowska
1997; Wojewoda et al. 1999; Wojewoda 2003). SUDETY ŚRODKOWE MTS: surroundings of Bystrzyca
Kłodzka (Schröter 1889). NIZINA ŚLĄSKA LOWLAND:
Wrocław – Strachocin (Schröter 1889). POJEZIERZE
POŁUDNIOWOPOMORSKIE LAKELAND: Bory Tucholskie forest, Cisy Staropolskie Reserve, Sept. 1995, leg.
H. Komorowska & Z. Heinrich (KRAM-F) (Wojewoda
& Komorowska 1997; Wojewoda et al. 1999; Wojewoda
2003). W ZNIESIENIA P OŁUDNIOWOMAZOWIECKIE
HEIGHTS: Rogów forest district, Popień forest part, sect.
171d (51°46′37″N/19°55′35″E), mixed forest, on dead
standing trunk of Populus tremula, 21 Sept. 2004, leg.
J. Nita (WAML 34) (Szczepkowski 2005). KOTLINA
SANDOMIERSKA BASIN: Puszcza Niepołomnicka forest,
Lipówka Reserve, Sept. 1984, leg. H. Komorowska
(KRAM-F), Oct.1994, leg. Z. Heinrich (KRAM-F), Oct.
1994, leg. W. Wojewoda (KRAM-F) (Komorowska 1991,
1995; Wojewoda & Komorowska 1997; Wojewoda et
al. 1999; Wojewoda 2003). POJEZIERZE MAZURSKIE
LAKELAND: Puszcza Borecka forest, surroundings of
Węgorzewo, Aug.1984, leg. Z. Heinrich (KRAM-F),
Sept. 1984, leg. B. Ginko (KRAM-F) (Wojewoda et al.
1999; Wojewoda 2003). BESKIDY WSCHODNIE MTS:
near Ustrzyki Dolne, Sept. 1980, leg. W. Wojewoda
(KRAM-F) (Gumińska & Wojewoda 1988; Wojewoda
& Komorowska 1997; Wojewoda et al. 1999, Wojewoda
2003). POLESIE ZACHODNIE: Czarny Las Reserve near
Parczew, Sept. 1996, leg. H. Komorowska (KRAM-F),
Sept.1996, leg. Z. Heinrich (KRAM-F) (Wojewoda et
al. 1999; Wojewoda 2003). POJEZIERZE LITEWSKIE
LAKELAND: Puszcza Augustowska forest, Starożyn
Reserve near Augustów, Sept. 1974, leg. Z. Heinrich
(KRAM-F) (Wojewoda 1977, 1979, 2003; Wojewoda
& Komorowska 1997; Wojewoda et al. 1999); Jezioro
Kalejty Reserve near Augustów, Oct. 1995, leg. H. Komorowska (KRAM-F) (Wojewoda & Komorowska
1997; Wojewoda et al. 1999; Wojewoda 2003); Perkuć
Reserve near Augustów, Oct. 1995, leg. H. Komorowska
(KRAM-F) (Wojewoda & Komorowska 1997; Wojewoda
MORSKIE LAKELAND :
85
et al. 1999; Wojewoda 2003). ROZTOCZE: Roztoczański
National Park, Nart near Zwierzyniec Reserve, Sept.
1986, leg. M. Ławrynowicz (KRA) (Sałata 1991; Wojewoda & Komorowska 1997; Wojewoda et al. 1999;
Wojewoda 2003). NIZINA PÓŁNOCNOPODLASKA LOWLAND: Puszcza Białowieska primeval forest, Białowieża
forest district, Pogorzelce Reserve, sect. 252D, on log
of Carpinus betulus, 30 Aug. 2000, leg. A. Bujakiewicz & A. Miśkiewicz (KRAM-F) (Bujakiewicz 2003);
Podolany Reserve, sect. 500A, on log of Carpinus betulus, 29 Aug. 2000, leg. A. Bujakiewicz & A. Miśkiewicz
(KRAM-F) (Bujakiewicz 2003); Białowieża National
Park, 13 Oct. 1950, leg. A. Pilát (PRM) (Pilát 1956;
Wojewoda 1979; Wojewoda 2003); sect. 256, Sept.
1987, leg. A. Bujakiewicz & B. Sadowska, Oct. 1990,
leg. A. Bujakiewicz (POZM) (Faliński et al. 1997;
Wojewoda & Komorowska 1997; Wojewoda et al.
1999; Wojewoda 2003); sect. 283, Sept. 1973, leg.
V. Holubová (KRAM-F) (Wojewoda & Komorowska
1997; Wojewoda et al. 1999; Wojewoda 1979; 2003);
sect. 285, leg. A. Skirgiełło (WA – no specimen in
herbarium) (Wojewoda & Komorowska 1997; Wojewoda et al. 1999; Wojewoda 1979, 2003); sect. 314,
Sept. 1977, leg. W. Wojewoda (KRAM-F) (Wojewoda
& Komorowska 1997; Wojewoda et al. 1999; Wojewoda 2003); sect. 369, Aug. 1973, leg. V. Holubová
(KRAM-F), Aug. 1973, leg. W. Wojewoda (KRAM-F)
(Wojewoda 1977, 2003; Wojewoda & Komorowska
1997; Wojewoda et al. 1999); sect. 370, Aug. 1973,
leg. Z. Pouzar (KRAM-F) (Wojewoda & Komorowska
1997; Wojewoda et al. 1999; Wojewoda 2003); sect.
399, Aug. 1973, leg. Z. Heinrich (KRAM-F) (Wojewoda 1979, 2003; Wojewoda & Komorowska 1997;
Wojewoda et al. 1999).
DISCUSSION
COMPARISON OF SPECIMEN DIMENSIONS
We measured the basidiomes, spores and basidia
recorded on eight tree species and on Armillaria
rhizomorphs to calculate the size of individuals
growing on different substrates and to compare
its variability with literature data. No relationship
between substrate type and the dimensions of any
part was found. Measurement values generally
were within the range of values given by other
authors; only the upper-range values of basidiome
measurements reported from Poland (Wojewoda
1977, 1981; this study) are higher than in the literature (Table 1).
86
POLISH BOTANICAL JOURNAL 53(1). 2008
DISTRIBUTION OF PHLEOGENA FAGINEA
IN POLAND
Phleogena faginea has been considered very rare
in Poland except in the Białowieża National Park,
where it was recorded from many forest sections.
As few as six localities of P. faginea were known
in 1979 (Wojewoda 1979): five in the Puszcza
Białowieska primeval forest and one in the Puszcza
Augustowska forest. Eleven new localities of the
fungus were discovered twenty years later (Wojewoda et al. 1999). Wojewoda (2003) added two
historical localities recorded by Schröter (1889)
from Wrocław (Strachocin) and the surroundings
of Bystrzyca Kłodzka. Bujakiewicz (2003) noted
two more localities in the Puszcza Białowieska
primeval forest, and Szczepkowski (2005) published the first record of the species from central
Poland.
Recently we found 19 new localities of P. faginea in Poland. Three are located near Gdańsk,
the northernmost localities in Poland, and five are
in central Poland, that is, in the area from which
this fungus was recorded only recently (Szczepkowski 2005). Another locality is in the Pojezierze
Południowopomorskie lakeland in northwest Poland, eight in the Puszcza Białowieska primeval
forest in the northeast, and the other two in the
south in the Magurski National Park, where the
fungus was found at its highest station in Poland, at
573 m a.s.l. The number of localities of P. faginea
has doubled over the last three years. The records
of new sites are most likely the result of heightened interest in macrofungi and the intensification
of wide-ranging penetrative field studies which
yield more data, and not from the spread of the
fungus. The 19 new localities presented in this
paper significantly increase the information on the
distribution of the species in Poland, revealing
that while still not common, the fungus occurs
throughout Poland (Fig. 2).
So far, 41 localities of the fungus in Poland
are known, mostly in natural plant communities
protected as national parks or nature reserves.
Of these, 11 are in national parks (Białowieski,
Magurski, Roztoczański), 18 in or very near nature
reserves, 9 in better-preserved managed forests,
Fig. 2. Distribution of Phleogena faginea (Fr.: Fr.) Link in
Poland (● – localities known from the literature, ○ – new
localities); 9 – number of localities known from the literature
in the Puszcza Białowieska primeval forest; 8 – number of new
localities in the Puszcza Białowieska primeval forest.
including one tree which is a nature monument,
and one in a historic urban park (Warsaw). Wojewoda et al. (1999) noted that no locality of this
species in Poland had been found in forest plantations, parks, or wooded synanthropic habitats. This
makes our new locality of P. faginea in Łazienki
Park in the center of Warsaw (Fig. 1e) particularly
noteworthy. It is the first record of the species in
Poland in a habitat highly altered by human activity. Although the Park was established on the
site of an ancient natural forest, systematic gardening has been carried out there for many years.
Damage to some trees, including those colonized
by P. faginea, has been treated with fungicides.
The locality of the species in the center of a large
city should encourage researchers to look actively
for P. faginea in other urban agglomerations to
establish whether the species can adapt to habitats
transformed by human activity.
Phleogena faginea is red-listed as endangered
(E) in Poland (Wojewoda & Ławrynowicz 1986,
1992, 2006). In light of the new data on its distribution presented here, we suggest that its threat
category should be changed to ‘vulnerable’ (V),
and we stress the need for further detailed studies
on the its distribution and ecological range.
A. SZCZEPKOWSKI ET AL.: PHLEOGENA FAGINEA IN POLAND
ECOLOGICAL REQUIREMENTS OF PHLEOGENA
FAGINEA
The new records of P. faginea give information on
additional substrates for it. We recorded the fungus
on the following substrates not reported previously
in the Polish literature: wood of Alnus incana,
Padus avium, Quercus robur, Tilia cordata, Ulmus
glabra and U. laevis, and rhizomorphs of Armillaria sp. The genus Tilia is also a new substrate
record for Europe; rhizomorphs of Armillaria
sp. were not recorded as its substrate in Europe,
but were noted as such in North America (Barr
& Bigelow 1968). We used available literature data
and our own observations to characterize substrates
suitable for P. faginea (Table 2) in Poland. Out of
13 species of deciduous trees, one coniferous tree
and two fungi, constituting 37 records (trees) and
accounting for ca 45% of all records, our analysis
showed that wood of Carpinus betulus provided
the most suitable substrate for P. faginea. Interestingly, that tree was most frequently mentioned as
the substrate of P. faginea also in Slovakia (Ripková et al. 2004) and the Czech Republic (Holec
2003). The fungus was noted relatively often in
Poland on Alnus glutinosa (13 records), Alnus incana (9), and Populus tremula (7); it was found
only accidentally on basidiomes of Fomes fomentarius and on old rhizomorphs of Armillaria sp.
In Poland, P. faginea was rarely found on Fagus
sylvatica (3 records), while in some countries of
Western Europe, such as, Germany, Denmark and
Great Britain, that tree species is reported as the
most frequent substrate colonized by this fungus
(Gøtzsche 1980; Kreisel 1987; Cook 1994).
Hornbeam trees occur practically throughout
Poland and are absent only from lower and upper
montane forest, growing above 600 m only exceptionally, while beeches hardly ever occur in
northeast Poland (Boratyńska & Boratyński 1990;
Boratyńska 1993), from where more than half of
the localities of P. faginea are known; this explains
its more frequent records on Carpinus betulus than
on Fagus sylvatica in Poland. Both tree species
have similar ecological requirements but their life
strategies differ. Hornbeam is a typical admixture species, encountered in different forest types
87
or forming lower layers in mixed stands where
it dies earlier as a shorter-lived species than, for
instance, beech (Dzwonko 1990; Faliński & Pawlaczyk 1993; Jaworski 1995), becoming available
earlier as substrate for P. faginea.
On the other hand, the area covered by stands
with beech dominant is ten times greater that of
hornbeam, and covers ca 366,000 ha in Polish
forests, that is, 4.2% of the forest area in Poland,
while hornbeam is reported from only 37,000 ha
(0.4%) (Grzesiak et al. 2007). A more in-depth and
broader search for this fungus in beech forests and
in communities where both beech and hornbeam
are present, might produce more records of the
fungus on beech wood.
Analysis of the literature data on the vertical
distribution of P. faginea in Europe shows that
it occurs mainly in lowlands and foothills. Its
locality from the Magurski National Park in Poland at 573 m is the highest site reported in print.
Similar data are given from Slovakia (Ripková
et al. 2004), where the highest site is located at
550 m. The highest locality in the Czech Republic
is at only 350 m (Holec 2003) and in Austria at
340 m (Scheuer & Poelt 1995).
Phleogena faginea is reported most often from
protected areas and is therefore considered to be
a relict of old-growth forests (Wojewoda 1999).
We found the fungus growing on thick old trunks
as well as thinner (younger) trees and shrubs. The
circumference of the measured trunks and stumps
ranged from 0.2 to 2.94 m (Table 2) and we could
not identify any clear-cut tendency to colonize the
oldest trees. It also seems that P. faginea has no
preferences for a particular height on the trunk.
On standing trees we observed the fungus growing
from the root swelling up to as high as 3.5 m.
It was even found growing on roots of Prunus
spinosa in the Czech Republic (Holec 2003), and
at 4–5 m above the ground on a broken trunk of
Fagus sylvatica in Germany (Tallasch & Jahn
1970). On the other hand, we noticed that basidiomes of P. faginea on fallen trees, logs, and
branches were on the lower side in most cases.
The fungus was less often found growing laterally, and only accidentally on the upper side of
the substrate.
88
POLISH BOTANICAL JOURNAL 53(1). 2008
Table 2. Characteristics of substrate of Phleogena faginea (Fr.: Fr.) Link from Polish localities.
Species
Number
of trees
Circumference [m]
Form of substrate
References*
Carpinus betulus
37
0.95–2.80
dead standing trunk, trunk of dying tree, remains of standing trunk,
fallen trunk, dead bark and wood of living standing tree, rotten log;
dead bark of living standing tree, fissures of bark of living tree, dead 1, 2, 3
fallen trunk, decayed fallen trunk, dead, fallen branch; dead bark of
living standing tree, butt of trunk
Alnus glutinosa
13
0.40–1.95
dead standing trunk, log, remains of the stump, trunk of dying tree;
dead bark of living standing tree, dead standing trunk, dead fallen trunk,
decayed fallen trunk
1, 2
Alnus incana
9
0.20–1.15
dying tree; decorticated branch of living tree, trunk of standing broken
tree, log
1
Populus tremula
7
0.45–1.20
dead standing tree, lying tree; dead fallen trunk, bark of decayed
trunk
1, 2
Ulmus laevis
5
0.25–2.90
stump, dead bark of trunk of living tree, dead standing trunk, dead
standing broken trunk
1
Acer platanoides
3
1.20–2.30
dead bark of trunk of dying tree, wood for fuel; standing trunk
1, 2
Fagus sylvatica
3
1.10
barkless log, dead standing trunk; decayed trunk
1, 2
Corylus avellana
2
0.40
dead standing branch; bark of dead branch
1, 3
Padus avium
2
0.25–0.35
Picea abies
2
2.80
old stump; dead standing trunk
1, 2
Betula pendula
2
1.90
log; dead standing trunk
1, 2
Quercus robur
1
2.49
dead, without bark, standing trunk
1
Tilia cordata
1
1.30
dead, fallen branch
1
Ulmus glabra
1
1.50
wood and bark of dead standing trunk
1
Unidentified trees
2
0.80
log; dead, fallen deciduous trunk
Fomes fomentarius
–
–
basidiocarp growing on Carpinus betulus
2
Armillaria sp.
–
–
rhizomorphs growing on dead standing trunk and log of Ulmus laevis
and on lying branch of Tilia cordata
1
dead standing, trunk; dead bark and wood of lying tree
1
1, 2
* 1 – present data; 2 – Wojewoda & Komorowska 1997; 3 – Wojewoda et al. 1999
According to Oberwinkler and Bauer (1989),
P. faginea prefers angiosperm substrates of hard
wood and bark on standing trees or only partly
decayed lying logs. Tallasch and Jahn (1970)
describe the occurrence of P. faginea on a trunk
of Fagus sylvatica with Bjerkandera adusta and
on a lying trunk of the same species colonized
by Pleurotus ostreatus, Trametes gibbosa and
T. hirsuta. We also observed that basidiomes of
P. faginea could often be found on trees and logs
colonized by other fungi causing white wood rot
(e.g., Armillaria sp., Chondrostereum purpureum,
Fomes fomentarius, Inonotus radiatus, Panellus
serotinus, Phellinus sp., Pleurotus ostreatus,
Schizophyllum commune), as well as brown wood
rot (Fomitopsis pinicola). The occurrence of P.
faginea on various tree species of different ages
and on other fungi, as well as its tolerance of
wood decay of different degrees and types, shows
high ecological plasticity and a high survival po-
A. SZCZEPKOWSKI ET AL.: PHLEOGENA FAGINEA IN POLAND
tential in managed forests and other substitute
habitats.
Phleogena faginea was observed in Poland
from August to October (Wojewoda & Komorowska 1997; Wojewoda et al. 1999). We
noted the occurrence of its fresh fruitbodies in
November, December, January and February
during an exceptionally mild autumn and winter.
Similar phenological observations were reported
from Slovakia, where the fungus was found from
July to December (Ripková et al. 2004). Oberwinkler and Bauer (1989) suggest that its time
of occurrence may be the reason for the rarity
of collections. We observed that basidiomes of
P. faginea could persist for a long period. In the
localities in Warsaw, some basidiomes produced
in 2006 were still observed in July 2007.
ACKNOWLEDGEMENTS. We thank Marek Kozłowski, Dariusz Majgier, Dr. Marcin Piątek, Dr. Jacek Piętka, Waldemar Półtorak, Robert Puciata, and Marek Snowarski
for information about localities of Phleogena faginea
and for permission to publish their photographs in this
paper, and Dr. Anna Ronikier and Dr. Marcin Piątek for
information on specimens in KRAM-F, and the Directors of the Białowieża National Park for permission to
collect basidiomes from the Strict Protection Area. We
also thank the anonymous reviewer for useful remarks
on the manuscript.
REFERENCES
ANONYMOUS 2007. The Norwegian Mycological Database.
[March 2007]. http://www.nhm.uio.no/botanisk/sopp/
index.html.
BARR M. E. & BIGELOW H. E. 1968. Martindalia unmasked.
Mycologia 60(2): 456–457.
BAUER R., BAGEROW D., SAMPAIO J. P., WEISS M. & OBERWINKLER F. 2006. The simple-septate basidiomycetes:
a synopsis. Mycological Progress 5: 41–66.
BENKERT D., DÖRFELT H., HARDTKE H.-J., HORSCH G.,
K REISEL H., K RIEGLSTEINER G. J., L ÜDERITZ M.,
RUNGE A., SCHMID H., SCHMITT J. A., WINTERHOFF W.,
WÖLDECKE K, ZEHFUSS H.-D., EINHELLINGER A., GROSS
G., GROSSE-BRAUCKMANN H., NUSS I. & WÖLFEL G.
1992. Rote Liste der gefährdeten Großpilze in Deutschland.
Deutsche Gesellchaft für Mykologie e.V. Naturschutzbund
Deuschland e.V. (NABU), Eching.
BORATYŃSKA K. 1993. Systematics and geographical distribution. In: W. BUGAŁA (ed.), Grab zwyczajny Carpinus
89
betulus L., pp. 17–50. Polska Akademia Nauk Instytut
Dendrologii w Kórniku & Sorus, Poznań – Kórnik (in
Polish with English summary).
BORATYŃSKA K. & BORATYŃSKI A. 1990. Systematics and
geographical distribution. In: S. BIAŁOBOK (ed.), Buk
zwyczajny Fagus sylvatica L., pp. 27–73. Polska Akademia
Nauk Instytut Dendrologii w Kórniku & Państwowe
Wydawnictwo Naukowe, Warszawa – Poznań (in Polish
with English summary).
BUJAKIEWICZ A. 2003. The Białowieża Forest – refuge for endangered macrofungi. Parki Narodowe i Rezerwaty Przyrody 22(3): 323–346 (in Polish with English summary).
COOK P. 1994. Profiles of Fungi 62: Phleogena faginea (Fr.:
Fr.) Link. Mycologist 8(3): 107.
DZWONKO Z. 1990. Ecology. In: S. BIAŁOBOK (ed.), Buk
zwyczajny Fagus sylvatica L., pp. 237–328. Państwowe
Wydawnictwo Naukowe, Warszawa – Poznań (in Polish
with English summary).
FALIŃSKI J. B. & PAWLACZYK P. 1993. Outline of ecology. In:
W. BUGAŁA (ed.), Grab zwyczajny Carpinus betulus L., pp.
157–263. Sorus, Poznań – Kórnik (in Polish with English
summary).
FALIŃSKI J. B., MUŁENKO W., BUJAKIEWICZ A. & MAJEWSKI T.
(eds) 1997. Crytptogamous plants in the forest communities of Białowieża National Park. Ecological atlas. (Project
CRYPTO 4). Phytocoenosis 9 (N.S.), Supplementum Cartographiae Geobotanicae 7: 427.
GÄRDENFORS U. (ed.) 2000. Rödlistade arter i Sverige – The
2000 Red List of Swedish Species. ArtDatabanken, SLU
Uppsala.
GØTZSCHE H. F. 1980. Phleogena faginea in Lyngby Ĺmose.
Svampe 1: 6–7 (in Danish with English summary).
GRZESIAK M., BUDNA E., GRZYBOWSKA L., KARCZEWICZ A.
2007. Forestry 2007. Information and Statistical Papers.
Główny Urząd Statystyczny, Warszawa (in Polish with
English summary).
GUMIŃSKA B. & WOJEWODA W. 1988. Grzyby i ich oznaczanie. 4 ed. Państwowe Wydawnictwo Rolnicze i Leśne,
Warszawa.
HANSEN L. & KNUDSEN H. (eds) 1997. Nordic macromycetes.
Vol. 3. Heterobasidioid, Aphyllophoroid and Gastromycetoid Basidiomycetes. Nordsvamp, Copenhagen.
HOLEC J. 2003. Phleogena faginea – abundant occurrence in
the Podyjí National Park and notes on its distribution in
the Czech Republic. Mykologické Listy 84–85: 33–37 (in
Czech with English summary).
JAWORSKI A. 1995. Charakterystyka hodowlana drzew leśnych.
Gutenberg, Kraków.
JURC D., PILTAVER A. & OGRIS N. 2005. Fungi of Slovenia:
species and distribution. Studia Forestalia Slovenica 124:
1–497.
KOMOROWSKA H. 1991. The Tricholomataceae (Agaricales)
90
POLISH BOTANICAL JOURNAL 53(1). 2008
of the Niepołomice Forest. Folia Societatis Scientiarum
Lublinensis, Biologia 30(1–2): 55–62 (in Polish with English summary).
SCHRÖTER J. 1889. Die Pilze Schlesiens. Erste Hälfte. In:
F. COHN (ed.), Kryptogamen-Flora von Schlesien. 3(1). J.
U. Kern’s Verlag, Breslau.
KOMOROWSKA H. 1995. Grzyby wielkoowocnikowe. In:
Z. MIREK & J. J. WÓJCICKI (eds), Szata roślinna Parków
Narodowych i Rezerwatów Polski Południowej. Polish
Bot. Stud. Guidebook Series 12: 44–48.
STOREY M. 2004. BioImages: The Virtual Field-Guide (UK).
www.bioimages.org.uk/html/P5/P52109.php.
KONDRACKI J. 2002. Geografia regionalna Polski. Wydawnictwo
Naukowe PWN, Warszawa.
TALLASCH H. & JAHN H. 1970. Phleogena faginea (Fr.) Link
im Naturscchutzgebiet ‘Hasbruch’ bei Bremen. Westfälische Pilzbriefe 8(2): 31–35.
KREISEL H. (ed.) 1987. Pilzflora der Deutschen Demokratischen
Republik. Basidiomycetes (Gallert-, Hut-, und Bauchpilze).
VEB Gustav Fischer Verlag, Jena.
KRIEGLSTEINER G. J. 1991. Verbreitungsatlas der Großpilze
Deutschlands (West). Band 1: Ständerpilze. Teil A: Nichtblätterpilze. Verlag E. Eugen, Stuttgart.
LÓPEZ A. & GARCÍA J. 2001. Phleogena faginea (Fries) Link.
Funga Veracruzana 42. http://www.scribd.com/doc/40664/
FUNGA-VERACRUZANA-Num-42-Phleogena-faginea.
MCNABB R. F. R. 1964. New Zealand Tremellales. 1. New
Zealand J. Bot. 2(4): 403–414.
MIREK Z., MUSIAŁ L. & WÓJCICKI J. J. 1997. Polish herbaria.
Polish Bot. Stud. Guidebook Series 18: 1–116.
MIREK Z., PIĘKOŚ-MIRKOWA H., ZAJĄC A. & ZAJĄC M. 2002.
Flowering plants and pteridophytes of Poland. A checklist.
W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków.
MÖLLER A. 1895. Protobasidiomyceten. Bot. Mitt. Tropen 8:
1–152.
OBERWINKLER F. & BAUER R. 1989. The systematics of gasteroid, auricularioid Heterobasidiomycetes. Sydowia 41:
224–256.
PILÁT A. 1956. Phleogena faginea (Fr.) Link – prachovečník
bukový v Karpatach. Česká Mykol. 10(2): 91–94 (in Czech
with Latin summary).
RIPKOVÁ S., KUČERA V., ADAMČÍK S. & MEREĎA P. 2004.
The occurrence of Phleogena faginea in Slovakia. Bulletin
Slovenskej Botanickej Spoločnosti 26: 19–22 (in Slovakia
with English summary).
SAŁATA B. 1991. Polish Mycological Symposium on ‘Flora,
ecology and geographical distribution of fungi.’ Folia Societatis Scientiarum Lublinensis, Biologia 30(1–2): 3–9.
SCHEUER C. & POELT J. 1995. Phleogena faginea (Fr.: Fr) Link.
Mycotheca Graecensis 2. http://www.uni-graz.at/~scheuer/
Exsiccata/MYCOGRAEC.html.
SCHNITTLER M. 1996. Zu den Roten Listen der Pilze Deutschlands. Schriftenreihe Vegetationsk. 28: 369–376.
SZCZEPKOWSKI A. 2005. Rogów 2004. Agricola 60: 40–41.
WOJEWODA W. 1977. Podstawczaki (Basidiomycetes). Trzęsakowe (Tremellales). Uszakowe (Auriculariales). Czerwcogrzybowe (Septobasidiales). Państwowe Wydawnictwo
Naukowe, Warszawa – Kraków.
WOJEWODA W. 1979. The geographical distribution of the
Polish tremellaceous fungi in Poland. Acta Mycol. 15(1):
75–144 (in Polish with English summary).
WOJEWODA W. 1981. Basidiomycetes (Podstawczaki). Tremellales (Trzęsakowe). Auriculariales (Uszakowe). Septobasidiales (Czerwcogrzybowe). Państwowe Wydawnictwo
Naukowe, Warszawa – Kraków.
WOJEWODA W. 2003. Checklist of Polish Larger Basidiomycetes. W. Szafer Institute of Botany, Polish Academy of
Sciences, Kraków.
WOJEWODA W. & KOMOROWSKA H. 1997. Notes on Phleogena
faginea (Fungi, Atractiellales). Fragm. Florist. Geobot.
42(1): 153–160.
WOJEWODA W., HEINRICH Z. & KOMOROWSKA H. 1999. New
localities and new host for Phleogena faginea (Fungi,
Atractiellales) in Poland. Fragm. Florist. Geobot. Ser.
Polonica 6: 199–202 (in Polish with English summary).
WOJEWODA W. & ŁAWRYNOWICZ M. 1986. Red list of
threatened macrofungi in Poland. In: K. ZARZYCKI,
W. WOJEWODA & Z. HEINRICH (eds), List of threatened
plants in Poland. 1 ed., pp. 45–82. W. Szafer Institute of
Botany, Polish Academy of Sciences, Kraków.
WOJEWODA W. & ŁAWRYNOWICZ M. 1992. Red list of
threatened macrofungi in Poland. In: K. ZARZYCKI,
W. WOJEWODA & Z. HEINRICH (eds), List of threatened
plants in Poland. 2 ed., pp. 27–56. W. Szafer Institute of
Botany, Polish Academy of Sciences, Kraków.
WOJEWODA W. & ŁAWRYNOWICZ M. 2006. Red list of the
macrofungi in Poland. In: Z. MIREK, K. ZARZYCKI,
W. WOJEWODA & Z. SZELĄG (eds), Red list of plants and
fungi in Poland, pp. 53–70. W. Szafer Institute of Botany,
Polish Academy of Sciences, Kraków.
Received 13 April 2007