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~- - -~ 3.3 ... ····· ·:· ..... .. ·: ...... . 3.2 . : - -~ ~ : : :o-Ph+ldncfuLT~ ; : : : : : : : : : ··1· ........:........ ··:········ ·:· ....... ~ .......·:~6 . : • - Ph~lodn . . : : -- . : niger .....·~: ....... ·:· .. ~ - ... ·:· .. ..... . /· I e : __ ~.- / : ;_./I / ·· · ·secr Phellodoh fu~;:· 2.9 ~ -· . 2.7+-~ ,o . ··. · ~ .. . - -~·Tr 2.8 1 : 2.9 ' ~oi .. . 3.0 : : - ~ -~c 3.1 :I ).' "9 v : v vv : I · O·~r 0o I .. ; ... / .v:..... ' . -i._,~ : ,..- : ~ . . . : ./ 0 W '-· I : 0 . ,... ···· ········· ········ : / : / . I , I: : - · . V . · • : : : . ~.i_ v: 'V :· · V : : · · . v __ j / -~ : : : 3.2 ,. ~-: .- . ., 4 :/ / / / : . ,/ : : . .. .. . . .. 3.3 3.4 3.5 . .. ··:.... .... . : . . . . ·:· .. .. ... ·1· .... ...·:· ........ . : : : : .. ···· ·· ·7,······· · :····· - ..- ~ _... :II ".II /I ....../:'?; ... .... :..... : : : .... ... ... .. Phellodon cdnnatus: ;-/·······T--- ······: ········.· · . . -, : ... ·j· .. .... .-~ ... .... ·:· ........ j· .......t;Jj·V" ..•. T..... - ~L/ 3·0 2.8 - -~ .... .....:......... : ..... . .. : ...... ... ~ . . . ..... ; ...... .. :...... : : : : ·:7-t~o;/ 3.1 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. 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