Karstenia 43: 37--44, 2003
Phellodon secretus (Basidiomycota), a new
hydnaceous fungus .from northern pine woodlands
TUOMO NIEMELA, JUHA KJNNUNEN, PERTII RENVALL and DMITRY SCHIGEL
NIEMELA, T. , KINNUNEN, J. , RENVALL, P. & SCHIGEL, D. 2003: Phellodon
secretus (Basidiomycota), a new hydnaceous fungus from northern pine woodland s.Karstenia 43: 37-44. 2003.
Phellodon secretus Niemela & Kinnunen (Basidiomycota, Thelephorales) resembles
Phellodon connatus (Schultz : Fr.) P. Karst., but differs in havi ng a thinner stipe,
cottony soft pileus, and smaller and more globose spores. Its ecology is peculiar: it is
found in dry, old-growth pine woodlands , growi ng in sheltered places under strongl y
decayed trunks or rootstocks of pine trees , where there is a gap of only a few centimeters between soi l and wood. Basidiocarps emerge from humus as needle-like, ca. I mm
thick, black stipes, and the pileus unfolds only after the stipe tip has contacted the
overhanging wood. In its ecology and distribution the species resembles Hydnellum
gracilipes (P. Karst.) P. Karst. It seems to be extremely rare, found in Northern boreal
and Middle boreal vegetation zones, in areas with fairly continental climate.
Key words: Aphyllophorales, Phellodon, hydnaceou s fungi , taxonomy
Tuomo Niemela, Juha Kinnunen & Dmitry Schigel, Finnish Museum of Natural History, Botanical Museum, P.O. Box 7, FIN-00014 University of Helsinki, Finland
Pertti Renvall, Kuopio Natural History Museum, Myhkyrinkatu 22, FIN-70100 Kuopio, Finland
Introduction
Virgin pine woodlands of northern Europe make a
specific environment for fungi. The barren sandy
soil, spaced stand of trees and scanty lower vegetation result in severe drought during sunny
summer months, in particular because such woodlands are usually situated on exposed hillsides,
river banks, and tops of eskers. Pinus sylvestris
grows in almost pure stands, accompanied by single Picea abies and Betula pubescens here and
there. Slowly growing and straight pine trees give
excellent timber, and nowadays fairly little is left
intact of these handsome forests.
Fungal decomposition proceeds slowly in such
dry habitats, and just a few wood-rotting species
can easily occupy dry coarse woody debri s there .
In natural conditions pine trees die of wildfires or
fall down in storms, but quite a lot of them succeed to reach a high age of300- 500(-800) years,
eventually dying while standing. Such dead pine
trees may keep standing for another 200-500
years, losing their bark and thinner branches: in
this way the so-called kelo trees develop, common and characteristic for northern old-growth
pine woodlands. This process was described in
detail by Niemela et a!. (2002), and in that paper
many wood-inhabiting fungi of the kelo trees were
listed.
A team of mycologists from the University of
Helsinki (Yu-Cheng Dai, Juha Kinnunen , Olli
Manninen, Tuomo Niemela, Dmitry Schigel, Olli
Turunen) has inventoried protected old forests
of North and East Finland during the years 19982002. These studies were initiated and organized
by the local offices of the governmental Natural
Heritage Services; for the background of the inventories in Lapland, see Niemela et al. (2003).
38
NIEMELA ET AL.: PHELLODON SECRETUS
This intensi ve and laborious fieldwork revealed a
number of previously unknown fungal species ,
among them the peculiar hydnaceous fungus
described here.
During the preparation of this paper it turned
out that Pertti Renvall had collected the same
species already in the 1980s in eastern Lapland.
Materials and methods
Specimens of the new species were collected by the authors with CO\ orkers. They were photographed in the
field, and fresh character and ecology notes 1 ere made.
Specimens were dried soon after field trips in mushroom
dryer with venti lated 30-40°C temperature.
In addition to the new species, selected voucher materials of related Phellodon species were studied for comparison. All the specimens listed are deposited in the
Botanical Museum of the University of Helsinki (H),
unless otherwise indicated. Herbarium abbreviations are
according to Holmgren et al. (1990).
Microscopic studies were done and spores were measured from sections mounted in Cotton Blue (abbreviated
CB): 0.1 mg aniline blue (Merck 1275) dissol ved in 60 g
pure lactic acid; CB+ means cyanophily, CB( +)weak but
distinct cyanophilous reaction, CB- acyanophily. Amyloid and dextrinoid reactions were tested in Melzer's reagent (IKI): 1.5 g KI (potassium iodide), 0.5 g I (crystalline iodine), 22 g chloral hydrate, aq. dest. 20 ml; IKImeans neither amyloid nor dextrinoid reaction. Occasionally also 5% KOH was used as mountant or reagent.
As a rule 30 spores were measured from each specimen
selected for closer scrutiny ; spore spines were excluded
while measuring the dimensions. Measurements were done
using x 1250 magnification, phase contrast, oil immersion ; eyepiece scale bar showed a 1 -~mgrid,
and dimensions were estimated visually with an accuracy of 0.1 ~m.
In presenting the variation of spore size, 5% of the measurements out of each end of the range are given in parentheses. L= mean length (arithmetical mean of all spores),
W= mean width, Q= extreme al ues of the length/ idth
ratios among the studied specimens, and n= the number of
spores measured from given number of specimens.
SEM photographs were taken by Cambridge Scan S-2
microscope (20 Kv, registration of secondary electrons).
Preliminary picture editing was made in MicroCapture
2.2. and further editing in Corel PhotoPaint 9.
The main reference books used were: Coker & Beers
(1951), Nikolae a (1961 ), Maas Geesteranus (1971,
1975), Domanski (1975), Baird ( 1986a, b) , Breitenbach
& Kranzlin (1986), Stalpers (1993), and Hansen & Knudsen (1997). Special colour terms are from Anonymous
(1969), Rayner (1970) and Petersen (1996).
Phellodon secretus Niemela & Kinnunen,
species nova
- Figs. 1- 4
Fungus stipitatus, aculeatus, terrestris, parvus.
Pileus mollis, tomentosus, griseoalbus, azonatus, stipite tenui, spiculoso, nigra, aculeis al-
KARSTENIA 43 (2003)
bidis vel griseoalbis. Hyphae hyalinae, afibulatae; sporae hyalinae, globosae, spiculatae, 2.93.3 X 2. 7-3 f-lln.
Typus: Finland, Pohjois-Karjala Prov., Ilomantsi, Koivusuo, 26.1X .2002 Niemela 7460, Kinnunen & Schigel (holotype, H).
Etymology: secretus (Latin, adj.) = separate,
hidden , set aside, secret, etc., referring to the way
of growth out of sight under fallen trunks.
Basidiocarp terrestrial, stipitate, small, slender
and fragile, single or confluent with 2 or more,
separate stipes supporting a common pileus. Pileus plane or funnel-shaped or irregularly roundish and lobed, 0.9- 3(-5.5) em in diam., very thin,
0.3-1.5(-3 .5) mm, cottony soft throughout, at first
white with ash-grey tint, later darker grey (mousegrey) or with a hue of sepia, evenly coloured
(young) or paler towards the margin (old), but
not zonate. Lower surface at first white, then light
greyish white (pale mouse-grey), spines sharp,
slender, regular, dense, finally 0 .3-0.9(-1.5) mm
long and 0.06-0.12 mm diam. at base. Stipe black,
glabrous, very thin, 0.3-1.8(-2.3) mm and of even
thickness, 10--18 mm long, brittle when dry. A slice
of context stains olivaceous (pale brownish green)
in KOH. Fresh basidiocarp odourless, dry with
very faint spicy scent.
Monomitic, h yphae simple-septate, hypha!
walls CB+, IKI-, mostly KOH-; hyphae of context and stipe covered with minute, scattered,
amyloid granules. Context hyphae thin-walled, of
even thickness (not inflated), hyaline except close
to lower surface where slightly grey-brown, (2.8- )
3-4.2(-4.7) ~min
diam. (n=30/1), making a spaced
interwoven network where a few hyphae often
run parallel in bundles ; context hyphae olivaceous
in KOH. Stipe surface with parallel, slightly thickwalled, black-brown hyphae with prominent septa,
(3.2-)3.6-5(-5.7) ~min
diam. (n=30/l); of them
long, single hypha! tips arise upwards , but no
differentiated tomentum present; stipe medulla
with brown, regular, subparallel hyphae, (2.7-)3.44.7(-5) ~min
diam. (n=30/l). Tramal hyphae in
spines very thin-walled, fragile, of even thickness,
subparallel, hyaline (pale brownish in inner parts
close to attachment), (2.1-)2 .6-3.5(-3.7) ~min
diam. (n=30/1 ); subhymenium not differentiated.
Hymenium with basidia and basidioles only, basidia clavate with long, rooting base, 21-34 x 4.95.6 ~m
(n=1011), sterigmata 4 , no basal clamp;
basidioles clavate, (16- )20-29 x (3.8- )4.7-5 ~m
(n=1011 ); hypha! tips regular at spi ne apex.
KARSTENIA 43 (2003)
NIEMELA ET AL.: PHELLODON SECRETUS
39
b
1Dpm
d
10pm
Fig. I. Phellodon secretus Niemela & Kinnunen. a) spores, b) hyphae from context, c) ertical section of aculeus,
d) hymenial cells, e) section from stipe surface. Dra\ n by TN in CB from holotype.
Basidiospores globose, thin-walled, hyaline,
CB-orCB(+), IKI- , KOH-, (2.8-)2.9- 3.3(-3.8) x
(2.4--)2.7-3(- 3.2) fA.Ill, L= 3.10 [-tm, W=2.87 f-lill, Q=
1.06- 1.10 (n= 180/6), with separate spines, distinct oblique apiculus, and medium to small guttule.
Specimens studied: Finland. EteUi-Harne Prov.:
Tammela, Mustiala, Syrjaas, 2l.Vill.l866 Karsten
4677; 6.X.l866 Karsten (F. Fenniae Exs. 10: 907,
H; Herb. E. Fries, UPS). Pohjois-Karjala Prov.: Ilomantsi, Hattuvaara, Koivusuo Strict Nat. Res .,
26.IX.2002 Niemela 7460, Kinnunen & Schigel;
40
NIEMELA ET AL.: PHELLODON SECRETUS
KARSTENIA 43 (2003)
Fig. 2. Phellodon secretus Niemela & Kinnun en, basidiocarps in situ. Holotype, approximately natural size,
photo TN.
29.IX.2002 Kinnunen 1727, Niemela & Schigel.
Kittilan Lappi Prov.: Kolari , Yllasjarvi, Tunturipalo, 9.1X.l999 Niemela 6646, 6648 & Renvall
3873 (KUO). Muonio, Akaskero, 22.VIII. l999
Niemela 6508, 6509 & Dai. Sompion Lappi Prov. :
Savukoski, Urho Kekkonen Nat. Park, Jaurujoki,
Peuraselka, 23 .IX.1987 Renvall628.
Ecology and distribution
The new species is very special in its ecology. All
fruit bodies found by us were growing in virgin,
dry pine woodlands, in narrow spaces under fallen pine trees . The wood was either extensively
rotten trunk, or rootstock of long-ago fallen tree,
preferrably kelo (Niemela et al. 2002). Such sheltered places were dry even in rainy days . The
reason of favouring such places may be that there
is less competition between mycelia there, or that
even though the site is dry, its humidity is very
constant throughout the season, because of little ventilation in the narrow gap.
We got an impression that the new species is
mycorrhizal , because basidiocarps clearly arose
from the humus. According to Urmas Koljalg
(pers. comm.), 'so far it is known that all the tested species of the Thelephorales, including the
resupinate genera Pseudotomentella, Tomentella and Tomentellopsis, are mycorrhizal. Therefore
it is logical to deduce that P secretus is mycorrhizal, too.'
Basidiocarps start to develop as black, needle-thin stipes, sharp at their apex . In some cases
tens of such erect stipes were emerging side-byside on humus or sandy soil. The pileus usually
starts to grow only after the tip touches the overhanging wood, and if the distance is too long, the
growth may terminate and only needle-like stipes
are left (e.g., specimen 6509). The growing pileus
spreads along the wood surface above, and becomes lightly attached to it so that when the wood
is lifted up, the stipe sometimes breaks apart and
pileus follows the wood, or the pileus becomes
detached and whole basidiocarp remains standing on the ground beneath.
Our finds imply that the new species is at least
nowadays a rarity of northern pine forests. However, two collections by P.A. Karsten from the
KARSTENIA 43 (2003)
year 1866 (in H and UPS) derive from southern
Central Finland. It seems that Karsten hesitated
to name his material (evidently recollected from a
single site), because he sent the second collection to Fries; no new name was proposed by either Karsten or Fries. The collecting site at Ilomantsi belongs to slightly continental section of
Middle boreal zone (Ahti eta!. 1968). Most collections were made in Finnish Lapland (Northern
boreal zone): in easternmost parts (Savukoski,
Renvall628) and communes of Kolari and Muonio in the west, in forests belonging to the
planned, extensive Y!His-Pallas National Park
(Koivisto 2003), extending in north-south direction throughout western Finnish Lapland. We
have not seen material from other countries.
gracilipes (see K6ljalg & Renvall2000) is just a
fragile rhizomorph or mycelial cord, the present
species has a true, differentiated stipe. The two
species differ clearly in their colours, H. gracilipes
being bright purple-brown.
The closest relative is Phellodon connatus
(Schultz: Fr.) P. Karst. (=P melaleucus (Fr.: Fr.) P.
Karst.), also having a black, glabrous stipe. That
species is usually more robust: both cap and stipe
are thicker and tough both when fresh and dry.
The spicy odour of dry P secretus is faint, hardly
noticeable, while it is strong and pungent in P
connatus. At least in Finland P connatus usually
grows in spruce-dominated forests amongst thick
moss and fairly moist ground; also when found
in pine forests the moss layer is thick in places
where the fruit bodies of P connatus emerge. In
most cases macroscopy and ecology are enough
to separate it from P secretus, but every now and
then dwarf basidiocarps are found. Then spore
size and shape are the differentiating characters
(Fig. 3, Tab. 1); context hyphae of P connatus are
more densely packed than in the new species,
and almost parallel, thus offering another striking
difference.
Related species
In its size, the very thin stipe and the peculiar
way of growth Phellodon secretus resembles
Hydnellum gracilipes (P. Karst.) P. Karst., and in
fact the two species were sometimes found growing in the same forest, under separate tree trunks
but not far from each other. While the 'stipe' of H.
3.4~-
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41
NIEMELA ET AL.: PHELLODON SECRETUS
········.· ·····
· -+r.~_,
3.6
3.7
3.8
3.9
4.0
~m
Fig . 3. Spore dimensions in fo ur species of Phellodon. Spore spines were excluded from the measurements. Each
symbol indicates a single specimen, mean value of 30 spores measured in CB.
42
NIEMELA ET AL.: PHELLODON SECRETUS
KARSTENIA 43 (2003)
Table 1. Spore dimensions of the specimens studied. 30 spores were measured for each specimen.
Spore spines were excluded from the measurements.
Specimen
Spore dimensions
Spore L x W
Q
Phellodon secretus
Karsten -1677
1866 Karsten (UPS)
Kinnunen 1727
Niemela 6508
Niemelii 6509
Niemelii 6646
Niemela 6648
Niemela 7460
Renvall 628
(2.9- )3.0-3.3(-3.5)
3.0-3.3
(2.9-)3.0-3.4(- 3.6)
(2.8-)2.9- 3.3(-3.4)
3.0-3.3
(2.9- )3.0-3.2(-3.3)
(2.8-)2.9- 3.2(- 3.3)
2.8-3.1(- 3.5)
3.0-3.3(-3.7)
X
X
X
X
X
X
X
X
X
(2.5-)2.6--3.0
(2.8- )2.9- 3.0
(2.5- )2.7- 3.0(-3.1)
(2.6--)2.8-3.1
2.8-3.0(- 3.1)
2.7-3.0
(2.4-)2.6--3.0(-3.1)
2.6--3.0(- 3.1)
(2.5-)2.7- 3.9(-3.1)
3.!5
3.15
3.17
3.11
3.17
3.04
3.03
3.01
3.13
2.86
2.94
2.90
2.92
2.94
2.87
2.78
2.82
2.87
1.10
1.07
1.09
1.06
1.08
1.06
1.09
1.07
1.09
(2.6--)2.8-3.1(-3.3)
(2.7-)2.8- 3.1
(2.6--)2.9- 3.2(-3.5)
2.8-3.2(- 3.7)
(2.5- )2.6--3.0(- 3.1)
(2.8-)2.9-3.2
(2.6--)2.9-3.2(- 3.3)
(2.6--)2.8-3.0(- 3.1)
(2.8-)2.9-3.1(- 3.2)
(2.7- )2.8-3.0(- 3.1)
(2.8-)2.9-3.1(- 3.2)
(2.7-)2.9- 3.3(-3.6)
(2.6--)2.8- 3.0(- 3.3)
(2.9- )3.0-3.3(-3.4)
(2.6--)2.8-3.1(-3.2)
3.32
3.35
3.62
3.70
3.37
3.38
3.39
3.38
3.40
3.41
3.27
3.65
3.36
3.51
3.55
X
2.97
2.92
3.00
3.01
2.87
2.98
3.00
2.94
2.98
2.92
2.97
3.04
2.97
3.08
2.98
1.12
1.15
1.21
1.23
1.18
1.13
1.13
1.15
1.14
1.17
1.10
1.20
1.13
1.14
1.19
3.0-3.6(- 3.7)
(2.9-)3.0-3.7(- 3.9)
(2.8-)3.0-3.3(-3.4)
(2.7- )2.9-3.1(-3.5)
(2.8- )2.9-3.2(- 3.5)
(3.0-)3.1-3.7(- 3.9)
(3.0-)3.1-3.7(-3.8)
(2.9- )3.0-3.3(- 3.6)
3.66
3.80
3.61
3.58
3.47
3.79
3.81
3.57
X
3.21
3.25
3.13
3.02
3.08
3.28
3.35
3.09
1.14
1.17
1.15
1.19
1.12
1.16
1.14
1.15
3.86
4.13
3.97
X
X
3.38
3.74
3.26
1.14
1.10
1.22
4.45
X
3.84
1.16
X
X
X
X
X
X
X
X
X
Phellodon connatus
Haikonen 20367
1934 Hayren
Karsten 2038
Kinnunen 1518
1985 Koski-Kotiranta
Kytovuori 871177
Kytovuori 921484
Kytovuori 922480
Kytovuori 922648
Kytovuori 981743
Kytovuori 982290
1944 Malmstrom
Saarenoksa 29993
1960 Saltin
1977 Ulvinen
(2.9-)3.0-3.8(-4.0)
(3.0-)3.1-3.7(-4.0)
(3.2-)3.3-4.0(-4.1)
(3.1-)3.3-4.0(-4.1)
(3.0-)3.1-3.8(-4.0)
(3.1- )3.2-3.7(-3.9)
(3.1- )3.2- 3.6(-4.0)
(3.1-)3 .2- 3.6(-3.8)
3.2- 3.6(-3.8)
(3.0-)3.1-4.0(-4.1)
(3.0-)3.1-3.5(-3.7)
(3.2-)3.4-4.0
(3.0-)3.2-3.6(- 3.8)
(3.1- )3.2-3.8(-4.0)
(3.1-)3.2-4.0(-4.1)
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Phellodon niger
Askola 407
Askola 1811
Askola 2492
1943 Hayren
Niemelii 5682b
Niemela 1712
Niemelii 2310
1948 v. Schulmann
(3.2-)3 .3-4.0(-4.1)
(3.1- )3.4-4.0(-4.1)
(3.1- )3.2-4.0
(3.0-)3.2-4.0(-4.2)
3.1-3.8(-4.0)
(3.1- )3.4-4.1(-4.3)
(3 .3-)3 .4-4.0
(3.0-)3.2-4.0
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Phellodon tomentosus
KytOvuori 901839
KytOvuori 902033
Korhonen 11230
(3.5-)3.7-4.0(-4.2)
3.9-4.3(-4.7)
(3.6--)3.8-4.2
X
X
(3.0-)3.2-3.6(-3.7)
(3.4-)3.5-3.9(-4.0)
(2.8-)3.0-3.6(-3.7)
(4.0-)4.2-4.8(- 5.2)
X
(3.2- )3.6-4.0(-4.3)
X
X
Phellodon alboniger
1961 Hintikka
KARSTENIA 43 (2003)
NIEMELAET AL.: PHELLODON SECRETUS
Phellodon tomentosus (L. : Fr.) Baker is browncoloured (fawn, clay-buff, hazel, sienna etc.), with
zonate upper surface, larger than the new species, and grows on open ground in gregarious or
ring-shaped groups. The other related European
species, Phellodon niger (Fr. : Fr.) P. Karst. and P
confluens (Pers.) Pouzar have a well-developed
tomentum in their stipe, and their spores are larger than in P secretus (Fig. 3, Tab. 1).
The species name of Hydnum occultum Britz.
somehow points to the present species, but it
was described to be 'fibrous, corky, woody ', and
having tints of yellow and brown; Maas Geesteranus (1960) suspected it to be an uncommon col our variety of Phellodon connatus. Karsten identified his collection of the new species (Karsten
4677, listed above) as Hydnum cyathiforme
Schaeff., \ hich is considered to be a synonym of
Phellodon tomentosus (Maas Geesteranus 1975):
' stipe rusty, aculei rosy ; common' (Fries 1821).
Original collection of Hydnum hepaticum Kalchbr. (UPS) was studied; the specimen is badly preserved, but it is much too robust to be our species, and Maas Geesteranus (in herb.) has measured the spores to be 4.5-4.9 x 4 .3 ~-tm.
There are very many other, old, poorly understood names for stipitate hydnums, often missing authentic collections and identifiable by brief
macroscopic descriptions and hand-made habit
illustrations only. They were sur eyed in particu-
Fig. 4. Phellodon secretus Niemela & Kinnunen , spore
ornamentation . SEM from holotype, x 15 000, prepared by OS.
43
lar from Fries (1821) and Maas Geesteranus (1958,
1960, 1964,1971, 1975), butnoneofthemseemed
to match our new species. If this turns out to be a
northern species, as it seems now, the probability
of an older name is small.
Phellodon sinclairii (Berk.) G.H. Cunn., found
in New Zealand, has larger spores than our species, 3.6-4.5 x 3.1-3.8 ~-tm
(Maas Geesteranus
1971).
Hydnum pygmaeum Yasuda resembles our species in being small and having a dark stipe; the
English translation of the description was published by Maas Geesteranus ( 1971: 31 ), who suspected this to be another synonym of P cannatum. The stipe was said to be 'brown to black'
and the cap surface 'rather dark brown, with soft
dense hairs ', which does not fit well with P secretus. We tried to get the type material on loan from
TNS (Tokyo) , but in vain .
Related species studied: Phellodon connatus:
Finland. Varsinais-Suomi Prov.: Pohja, Dalkarby,
3.IX.1960 Saltin. Uusimaa Prov.: Espoo, Luukki,
20. VIII.1985 Koski-Kotiranta. Helsinki, Laajasalo,
28.IX.1934 Ha_yren ; Vuosaari, Kallvikudde,
2.IX.l993 Saarenoksa 29993. Tuusula, Klemetskog, Miitlikivenmiiki, 2l.IX.1944 Malmstrom.
Satakunta Prov.: Tyrviiii, 4.IX.1859 Karsten 2038.
Etelii-Savo Prov.: Joutseno, Konnunsuo, Leppiilii,
4.IX.1987 Kytovuori 871177. Kerimiiki, Ruokojiirvi, Louhi, 8.IX.1998 Kytovuori 981743. Pohjois-Savo Prov.: Savonranta , Muhamiiki ,
19.IX.1990 Haikonen 20367. Pohjois-Karjala
Prov.: Eno , Uimaharju, 19.IX.1992 Kytovuori
922648. Liperi, Viinijiirvi, Ahonkylii, 18.IX.l992
Kytavuori 922480 . Oulun Pohjanmaa Prov.:
Ylikiiminki, Karahka, 15.IX.1977 Ulvinen. Koillismaa Prov.: Kuusamo, Oulanka Nat. Park,
21.VIII.1992 Kytavuori 921484. Kittiliin Lappi
Prov.: Kolari, Akaslompolo, Kesiinki, 2l.IX.2001
Kinnunen 1518. Sweden. Smaland Pro .: Vastra
Ed, Viistervik, 27 .IX.J998 Kytavuori & Kytavuori
982290. Germany. 'Hydnum melaleucum Fr., Germania, in silvis montosis abiegnis Thuringiae ad
Saalfeldiam, Klotzsch Herb. Myc. n. 122, Septbr.
Klotzsch et Opatowski' (Herb. E. Fries, UPS).
Austria. 'Hydnum melaleucum, bsterrike, Gabelberg prope Grein, 24.VIII.1866 Heufler' (Herb. E.
Fries, UPS).
Phellodon niger: Finland. Uusimaa Prov.:
Elimaki, Mustila, 1948 Schulmann. Espoo, Luukki, 3.IX.1993 Niemela . Nurmijiirvi , Parkkimiiki,
25.IX.1988 As kola 2492 ; Pitkiimiiki, 10.IX.1977
44
NIEMELA ET AL.: PHELLODON SECRETUS
Askola 407; Kiljava, 22.IX .1985 Askola 1811.
Tuusula, NummenkyHi, 9.IX.1943 Hiiyren. EtelaHame Prov.: Lammi, Evo, Kotinen Virgin Forest,
14.IX.1979 Niemela 1712; Alinen Rautjarvi,
4.IX.1981 Niemelii 2310.
Phellodon tomentosus: Finland. VarsinaisSuomi Prov. : Karjaa, NE ofK1even, U:ivkullaudden, 25.IX.1990 Kytovuori 902033. Kemio , Pedersa, 2l.IX.1990 Kytovuori 901839. Tammisaari,
Bromarv, 17 .IX.1992 Korhonen 11230.
Phellodon confluens: Sweden. Ostergotland
Prov. : Vadstena, 23.IX.l985 Kytovuori 851344.
Gryt, 16.IX.1950Nannfeldt 11181. U.S.A. Michigan: Washtenaw County, Waterloo, 10.X.l961
Hintikka.
Phellodon alboniger: U.S.A. Michigan: Wilderness State Park, 17 .IX.l961 Hintikka.
Hydnum hepaticum: Hungary. 'Hydnum hepaticum nov. sp. Ungern : Scepusii, Oct. 1860 C.
Kalchbrenner 173' (Herb. E. Fries, UPS).
Acknowledgements: Staff of the Natural Heritage Services, in particular Paivi Paalamo (Rovaniemi) and Maarit
Simila (Lieksa) are thanked for inviting us to carry out
mycological inventories in the Y!His-Aakenus area and
Koitajoki Nature Reserve, respectively, and for arranging us excellent working conditions. Yu-Cheng Dai, Olli
Manninen, Kari Steffen and Olli Turunen accompanied
us in fieldwork. Teuvo Ahti revised the Latin. Pertti
Salo helped in finding the Karsten collections. Notes
and improvements to the manuscript by Urmas K6ljalg
are gratefully acknowledged. The author DS wishes to
thank Drs. A. Davydovich, A. Bogdanov and N. Zvonkova (Laboratory of Electron Microscopy, Biology Faculty, Moscow State University) for help. Important specimens from the Fries herbarium (in UPS) were obtained
on loan . Research grant from the Ministry of En ironment (YM131155 12/2002) enabled us to carry out thi s
study.
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