Mycol. Res. 104 (8): 1017–1024 (August 2000) Printed in the United Kingdom. 1017

Hebeloma radicosoides sp. nov., an belonging to the chemoecological group ammonia fungi

N. SAGARA1, T. HONGO2, Y. MURAKAMI3, T. HASHIMOTO4, H. NAGAMASU5, T. FUKIHARU6 and Y. ASAKAWA4 " Life Web, Graduate School of Human and Environmental Studies, Kyoto University, Kyoto 606-8501, Japan. # 3-3-7 Ogaya, Otsu 520-2144, Japan. $ Oita Mushroom Research Institute, Akamine, Mie-machi, Oita 879-7111, Japan. % Faculty of Pharmaceutical Sciences, Tokushima Bunri University, Yamashiro-cho, Tokushima 770-8055, Japan. & The Kyoto University Museum, Kyoto University, Kyoto 606-8501, Japan. ' Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chiba 260-8682, Japan. E-mail: naohikosagara!g53338!sakura.kudpc.kyotoIu.ac.jp

Received 14 April 1998; accepted 4 November 1999.

Hebeloma radicosoides sp. nov. is described, illustrated and characterised. It superficially resembles H. radicosum in having a long- rooting with a well-formed, membranous annulus and in having preference for animal wastes as its resources. Its pileus is yellower than that of H. radicosum, and it fruits after addition of urea to soil, whereas H. radicosum does not. It also fruits on various animal wastes whereas H. radicosum specifically fruits on mole or mouse middens. Unlike H. radicosum, H. radicosoides has no distinct odour. Hebeloma radicosoides also resembles Hebeloma luchuense and H. spoliatum in having a long-rooting stipe and in fruiting on urea-treated soil, but differs from them by having a yellowish scaly pileus and distinct, membranous annulus.

INTRODUCTION In Japan two species have been confused under the name Colour notations in parentheses are taken from Kornerup & . One is true H. radicosum known from Wanscher (1967) and those in quotation marks from Ridgway Europe (Bruchet 1970, Lange 1974) and Japan (Imazeki & (1912). Light-microscopic features were determined from fresh Hongo 1987, Imazeki, Otani & Hongo 1988), and the other material mounted in water or from dried material mounted in has been misidentified as H. radicosum (Imazeki & Hongo 3% KOH aqueous solution (KOH), unless otherwise stated. 1965). This misidentified species, having a long-rooting stipe For SEM, were boiled with 10% KOH solution for similar to H. radicosum, fruits on forest ground after application 5 min, dehydrated through an ethanol series and treated with of urea or any nitrogen compound that releases ammonia on a critical point drying method. decomposition or, under natural conditions, after decompo- sition of an animal body or excrement. It has, therefore, been included in the experimental (‘chemoecological’) group Hebeloma radicosoides Sagara, Hongo & Y. Murak., sp. nov. (Figs 1–5, 9, 11–14) ‘ammonia fungi’ (Sagara 1975, 1992) or in the natural ecological group ‘post-putrefaction fungi’ (Sagara 1995). Pileo 5–7 cm lato, obtuse conico-convexo, dein late convexo vel Here we describe it as a new species, and compare it with H. applanato, frequenter obtuse umbonato, margine primum in curvo, e viscido glutinoso, pallide flavo vel flavido, squamis brunneis appressis radicosum, which does not fruit after experimental nitrogen pconcentrice ornato; carne crassa, alba, sapore levi, odore debili; application, but only on mole or mouse middens (Sagara 1995, lamellis adnexis vel sinuatis, confertissimis, albidis dein sub- Sagara & Senn-Irlet, unpubl. data), and with other rooting cinnamomeis, acie minute fimbriatis; stipite 5–10 cm longo, 0n7–1 cm Hebeloma species. These include H. luchuense (Fukiharu & crasso, aequali vel e basi apicem versus leniter attenuato et subtus in Hongo 1995) and H. spoliatum sensu Imazeki & Hongo (1965, pseudorhizam (6–9 cm) producto, pallide flavido, ad apicem pruinoso- 1987) which, as ammonia fungi, often fruit together with the squamuloso, sub annulo squamis brunneolis revolutis squarroso; new species under the same field conditions. annulo membranaceo, glutinoso, primum albo, demum brunneolo; Ammonia Hebeloma radicosoides sp. nov. 1018

2

3

1

Figs 1–3. Hebeloma radicosoides (Hongo 6731 l duplicate of CBM FB-6118). Fig. 1. Habit and section; bar l 1 cm. Fig. 2. Basidiospores; bar l 10 µm. Fig. 3. Cheilocystidia; bar l 10 µm. sporis 8n7–12n6i4n6–6n4 µm, subamygdaliformibus, punctatis (light Holotypus: Japan: Shiga Prefecture, Otsu, on urea-treated microscopy), irregulariter reticulatis vel rugulosis (SEM), pallide ground in the mixed forest of Pinus densiflora and Quercus ochraceis, indextrinoid; basidiis tetrasporis; cheilocystidiis 20–28i5– serrata, 17 Oct. 1983, Y. Murakami, F-182343 (TNS-National 10 µm, numerosis, clavatis vel subclavatis, hyalinis, tenuitunicatis; Science Museum, Tsukuba; see Table 1). hyphis fibulatis. Pileus (2–)5–7(–15) cm broad, obtusely conico-convex at first Specimens examined: see Table 1. with incurved margin, expanding to broadly convex or plane, often obtusely umbonate; surface viscid to glutinous, pale Distribution: Japan (Honshu: Prefectures Fukushima, Ibaraki, yellow to light yellow (4A3–4A4), greyish yellow (4B5) or Shiga and Kyoto; Ryukyus: Iriomote Is.; see Table 1). ‘Light Buff’, with somewhat concentric, brown (6E6–7E6), appressed scales. Flesh thick (1–1n5(–2) cm at the disc), white, Common name: The false mole-nest finder (in English; yellowish (near 4A4) just beneath the pellicle; taste slightly H. radicosum may be called the mole-nest finder), bitter and acrid, odour faintly fungoid when young and faintly Nagaenosugitake-damashi (in Japanese). of burnt sugar when mature. Lamellae adnexed or sinuate, very crowded, 0n3–0n5(–0n7) cm broad, whitish when young, Iconography: No. 143, Plate 23 in Imazeki & Hongo (1965), becoming light brown (6D4), edges minutely fimbriate. Stipe under the name H. radicosum. (2n3–)5–10 cm long above ground level, (0n4–)0n7–1(–1n8) cm thick at the apex, equal or slightly attenuated upward from Hebeloma radicosoides can be distinguished from H. radicosum the thickened ((0n6–)1–1n5(–2n8) cm) and long-rooting by the yellowish pileus, compared to the whitish one in the (6–9(–25) cm) base, yellowish white (3A2) to pale yellow latter (compare Figs 4, 5 with Figs 6–8; see also Fig. 9), by the (4A3), pruinose-squamulose at the apex, with often recurved more crowded lamellae, and by the lower ornamentation in brownish scales below the annulus, solid to stuffed. Annulus the basidiospore surface (compare Figs 11–14 with Figs membranous, glutinous, white then brownish, attached to the 15–18). upper part or near the middle of the stipe. Basidiospores brown Hebeloma radicosoides is distinct from H. luchuense as the (6E6) or ‘Brussels Brown’ in mass, 8n7–12n6i4n6–6n4 µm latter has a brown pileus without scales, a cortina-like, (10n2p0n8 µm: meanp95% confidence limits) (n l 137), evanescent veil and dark brown scales on the stipe (compare subamygdaliform, finely punctate (tuberculate) under the light Figs 4, 5 with Fig. 10; see also Fig. 9). Hebeloma radicosoides is microscope, irregularly reticulate to rugulose under the SEM, distinct from H. spoliatum as the latter has a brownish pileus pale ochraceous in KOH, non-dextrinoid. Basidia 4-spored. without scales and is devoid of a veil. (The species concept of Cheilocystidia 20–28i5–10 µm, numerous, clavate to sub- H. spoliatum sensu Imazeki & Hongo (1965, 1987) followed clavate, hyaline, thin-walled. trama of parallel hyphae. that of Lange (1935–1940), and hence the Japanese H. Pileipellis an ixocutis (ixotrichodermium; term after Smith, spoliatum is most possibly conspecific with H. danicum as Evenson & Mitchel 1983), consisting of hyphae 2n5–7n5 µm illustrated by Moser, Ju$ lich & Furrer-Ziogas (1985–).) thick. Stipe trama of parallel hyphae. Clamp connections present Other Hebeloma species with rooting stipes may include in all tissues. ‘H. birrus’ (Gillet, 1891), ‘H. fusipes’ (Bresadola, reprint 1982), N. Sagara and others 1019

Table 1. List of Hebeloma radicosoides specimens used for morphological examination (*) and\or those selected for information on distribution, resource preference and host plant range (in alphabetical order of herbarium names and, for each herbarium, in order of specimen numbers)

Specimen no.a\ Resource Plant cover in the habitat, Date of Place of collection for fruitingb represented by dominant species collection Collector

CBM FB-5370* Ryukyus: Iriomote Is. Urea Castanopsis cuspidata var. sieboldii 3 x 1983 T. Fukiharu CBM FB-6002 Ryukyus: Iriomote Is. Urea C. cuspidata var. sieboldii, Quercus miyagii 3 i 1983 T. Fukiharu CBM FB-6118* Kyoto: Kyoto City Urea Tsuga sieboldii, Abies firma, C. cuspidata 2 x 1986 T. Fukiharu CBM FB-15143 Fukushima: Naraha-machi Urea Q. serrata, A. firma 26 ix 1996 T. Fukiharu CBM FB-19513 Kyoto: Kyoto City Peptone Pinus densiflora, Ch. obtusaf 16 v 1972 N. Sagara CBM FB-19515 Kyoto: Kyoto City Sodium glutamate P. densiflora, Ch. obtusa 7 x 1972 N. Sagara CBM FB-19516 Kyoto: Kyoto City (NH%)#CO$ P. densiflora, Ch. obtusa 7 x 1972 N. Sagara CBM FB-19517 Kyoto: Kyoto City Arginine P. densiflora, Ch. obtusa 7 ix 1973 N. Sagara CBM FB-19518 Kyoto: Kyoto City Milk casein P. densiflora, Ch. obtusa 22 ix 1974 N. Sagara CBM FB-19519 Kyoto: Kyoto City Ammonium oxalate P. densiflora, Ch. obtusa 22 ix 1974 N. Sagara CBM FB-19523 Kyoto: Kyoto City Glutamic acid P. densiflora, Ch. obtusa 6 x 1976 N. Sagara CBM FB-19525 Kyoto: Kyoto City (NH%)#HPO% P. densiflora, Ch. obtusa 6 x 1976 N. Sagara CBM FB-19537 Kyoto: Nagaokakyo City Human excrement Q. serrata 3 xi 1981 S. Takayama CBM FB-19540 Kyoto: Kyoto City Vespine wasp nest C. cuspidata 4 x 1982 N. Sagara CBM FB-19547 Kyoto: Miyama-cho Urea Q. serrata, Q. crispula 11 x 1983 N. Sagara CBM FB-19550* Kyoto: Miyama-cho Urea Carpinus laxiflora, Betula grossa 31 vii 1984 N. Sagara CBM FB-19551* Kyoto: Miyama-cho Urea C. laxiflora, B. grossa 28 ix 1984 N. Sagara CBM FB-19553* Shiga: Otsu City Uric acid P. densiflora, Q. serrata 4 x 1984 N. Sagara CBM FB-19555 Shiga: Ibuki-cho Raccoon dog Q. serrata, Castanea crenata 24 viii 1985 Y. Marunishi excrement CBM FB-19563* Kyoto: Kyoto City Urea C. cuspidata 8 ix 1993 N. Sagara CBM FB-19569 Kyoto: Kyoto City Ureaformc C. cuspidata 20 x 1993 N. Sagara CBM FB-19576 Kyoto: Kyoto City Oxamided Q. serrata, Q. glauca 29 vi 1994 N. Sagara CBM FB-19577 Kyoto: Kyoto City Coated ureae C. cuspidata 29 vi 1994 N. Sagara Hongo 1001* Shiga: Otsu City Not studied P. densiflora, Q. serrata 11 ix 1954 T. Hongo Hongo 5545* Shiga: Otsu City Not studied C. cuspidata var. sieboldii 25 ix 1976 T. Hongo Okabe 5092 Ibaraki: Kita-Ibaraki City Mouse excrement Fagus crenata, Q. crispula 11 x 1989 H. Okabe TNS F-182343 (type)* Shiga: Otsu City Urea P. densiflora, Q. serrata 17 x 1983 Y. Murakami

a Herbarium abbreviations: CBM, Natural History Museum and Institute, Chiba; Okabe, personal collection by Hiroaki Okabe, Forestry and Forest Products Research Institute, Tsukuba, Ibaraki 305-0903, Japan; Hongo, personal collection by Tsuguo Hongo, the second author of this paper; TNS, National Science Museum, Tsukuba. b Besides the substances listed here, ammonia (aqueous), calcium cyanamide and ammonium formate were effective for producing this fungus (Sagara 1975), although the specimens from these substances have not been preserved. c–e Slow-acting fertilizers, produced as follows: c ureajformaldehyde (Sumitomo Chemical Co., Ltd); d ammoniajprussic acid (Ube Industries, Ltd); e urea coated with synthetic resin (Sumitomo Chemical Co.). f Chamaecyparis obtusa is not ectomycorrhizal, so could not be the host of H. radicosoides; others could be (see text). Ammonia fungus Hebeloma radicosoides sp. nov. 1020

4 5

6 7 8

9 10

Figs 4–10. Basidiomata of Hebeloma radicosoides (Figs 4, 5, 9), H. radicosum (Figs 6–8) and H. luchuense (Figs 9, 10). Fig. 4. Young # basidiomata of H. radicosoides growing on a urea-treated plot (part of the type TNS F-182343; urea (1 kg m− ) applied on 14 Jan. 1983; photograph on 17 Oct. 1983; bar l 5cm;H. spoliatum in the background). Fig. 5. H. radicosoides collected from a urea plot (CBM # FB-19550; 1 kg m− applied on 24 Dec. 1982; photograph on 31 July 1984; bar l 10 cm). Fig. 6. Young basidiomata of H. radicosum growing on deserted middens of a mole (bar l 5 cm). Figs 7, 8. H. radicosum collected from mole nesting sites (bars l 10 cm). Fig. 9. H. radicosoides with yellowish scaly pilei (at left, CBM FB-5370), and H. luchuense with brown, smooth pilei (at right), fruiting together # on a urea plot (342 g m− applied on 26 Jan. 1983; photograph on 2 Oct. 1983; bar l 5 cm). Fig. 10. H. luchuense collected from a # urea plot (686 g m− applied on 16 Aug. 1982; photograph on 1 Jan. 1983; bar l 5 cm).

‘H. radicatum’ (Bresadola, reprint 1982), ‘H. cylindrosporum radicosoides. In addition Smith et al. (1983) do not include any var. pseudoradicatum’ (Bon 1979) and ‘H. syrjense’ (Lewis & comparable species with membranous annulus and rooting McGraw 1984). All these species have brownish pilei without stipe. scales and thus can again easily be distinguished from H. According to the classification of subgenera of Hebeloma by N. Sagara and others 1021

11 12 13 14

15 16 17 18

Figs 11–18. Basidiospores of Hebeloma radicosoides (Figs 11–14) and H. radicosum (Figs 15–18). Figs 11–12. H. radicosoides (Fig. 11 l CBM FB-19551, Fig. 12 l CBM FB-19563; in lactophenol; bars l 10 µm). Figs 13–14. H. radicosoides (SEM) (Fig. 13 l CBM FB-19551, Fig. 14 l CBM FB-19563; bar in Fig. 13 l 5 µm; bar in Fig. 14 l 2 µm). Figs 15–16. H. radicosum (in lactophenol; bars l 10 µm). Figs 17–18. H. radicosum (SEM) (bar in Fig. 17 l 5 µm; bar in Fig. 18 l 2 µm).

Singer (1986), the subgenus Myxocybe is characterised by the 1995 and unpublished data). H. radicosum has not, therefore, membranous annulus, well formed and persistent, and by the been included in the ammonia fungi (Sagara 1995). stipe that has a pseudorhiza. Morphologically, H. radicosoides Under natural conditions, H. radicosoides fruits on forest belongs to this subgenus, which has so far comprised solely ground that has happened to be disturbed by deposition of H. radicosum. nitrogenous animal matter (Table 1), such as a dead cat (Kuroyanagi et al. 1982), vespine wasp nest (Sagara et al. 1985), human excrement (Sagara & Takayama 1982), raccoon ECOLOGY dog excrement (Sagara 1989) and wood mouse excrement Hebeloma radicosoides fruits on forest ground after experimental (Sagara, Okabe & Kikuchi 1993). On the other hand, H. addition of aqueous ammonia or its substitutes, i.e. urea (Figs radicosum fruits specifically on mole or mouse middens (Figs 4, 5, 9), calcium cyanamide, ammonium carbonate, ammonium 20, 21; Sagara 1992, 1995; ‘midden’ in this paper means a site phosphate (dibasic), ammonium formate, ammonium oxalate, of repeated urination and\or defecation). sodium glutamate, glutamic acid, arginine, peptone, casein Hebeloma spoliatum shows, in field experiments as well as (from milk), and uric acid (Table 1; Sagara 1975, 1992). On under natural conditions, a similar resource preference to H. the contrary, H. radicosum does not fruit in response to such radicosoides, but can easily be distinguished by its morphology chemical treatment. This was true even when, in simulation of as stated previously. Hebeloma luchuense has been known only location by mole or mouse middens (see below), or a continual from urea-treated plots, and can also be distinguished by its supply of nitrogen from a mole or mouse, urea or some morphology. Other rooting Hebeloma species are not known fertilizers that release ammonia more slowly than urea, e.g. to have such ecological characteristics. Consequently, these ureaform, oxamide and coated urea, had been applied to species are excluded from the following discussion. deeper soils near the fruiting sites of H. radicosum; the Both H. radicosoides and H. radicosum seem to form treatments induced the fruiting of H. radicosoides and\or H. ectomycorrhizas under those special conditions brought about spoliatum, but not of H. radicosum (Fig. 19, Table 1; Sagara by the addition of nitrogenous material (Sagara et al. 1993, Ammonia fungus Hebeloma radicosoides sp. nov. 1022

Sagara 1995). Their growth may be confined to ectomy- crustulinol whereas H. radicosum does not (Hashimoto, corrhizal forests (Table 1; Sagara 1995). Asakawa & Sagara unpublished). This compound resembles the toxic principles known from (Bernardi et al. 1983) and H. spoliatum (Fujimoto, Katano CHEMISTRY & Yamazaki 1992). Hebeloma radicosoides has minimal odour (fungoid or of burnt sugar) whereas H. radicosum has a strong odour ‘like prussic acid or sweetish’ (Smith 1908) or ‘like cherry laurel or bitter CULTURE ’ (Rea 1922). Fresh basidiomata of H. radicosoides In tube cultures on a yeast glucose medium (glucose 20 g, dry collected from urea plots in Kyoto City, Kyoto Pref., on 16 yeast 5 g, agar 15 g, tap water 1000 ml) at room temperature Sep. 1993 and those of H. radicosum collected from a mole (10–20 m), H. radicosoides does not develop aerial hyphae nesting site in Miyama-cho, Kyoto Pref., on 9 Oct. 1992 were while H. radicosum does. As the cultures age, mycelia of H. extracted with ethyl ether, and their extracts were analysed radicosoides remain white while those of most H. radicosum using a Hewlett–Packard capillary GC-quadrupole MS system strains become brown. Both fungi form basidiomata in pure (Model HP-5890) fitted with a DB-17 column, where the oven culture (Ohta 1998). temperature was programmed from 80 mC to 250 m at −" 10 m min . This study of odourous substances showed that H. radicosoides does not contain volatile substances at REMARKS appreciable levels whereas H. radicosum contains 1-octen-3-ol, phenylacetaldehyde and N-formylaniline at high levels (Fig. As a result of the recognition of H. radicosoides as a distinct 22), and that the well-known characteristic odour of H. species the name ‘H. radicosum’ used until 1985 in the series radicosum is yielded when 1-octen-3-ol and phenylacet- of papers on the ammonia fungi (Sagara 1973, 1975, 1976, aldehyde are mixed. 1-Octen-3-ol is known to be quite 1978 (Fig. 5A), 1984, Sagara & Takayama 1982, Kuroyanagi widespread among fungi (Kaminski, Stawicki & Wasowicz et al. 1982, Sagara et al. 1985) should be corrected to 1974), and phenylacetaldehyde to be mainly responsible for H. radicosoides. The name ‘H. radicosoides’ has already been the smell of Phallus impudicus gleba (Kikuchi, Kadota & Nishi tentatively used for this species in some publications (Sagara 1984). Further analyses revelaed that H. radicosoides contains 1989, 1992, Sagara et al. 1993), and ‘Hebeloma sp.’ was also 3β-acetyl-12,21-anhydro-2α-(3h-hydroxy-3h-methyl)glutaryl- used (Sagara 1995).

B s

N

19 20 21

Figs 19–21. Ecological features of Hebeloma radicosoides and H. radicosum. Fig. 19. H. radicosoides fruiting on a buried mixture of urea and litter, with long rooting stipes (3 cm thickness of L and L–F horizons, collected from the surrounding area, and 62n5 g of urea were buried together at the 25i25 cm bottom (arrow) of a 30 cm deep hole in a Pinus densiflora-Quercus serrata mixed forest, Otsu City, Shiga Pref., on 14 Jan. 1983; photograph on 17 Oct. 1983; s l H. spoliatum). Figs 20, 21. H. radicosum fruiting on a mole’s deserted midden: Fig. 20. Basidioma (B) beside the mole’s nest (N); Fig. 21. The same basidioma, having grown on the midden around a tunnel (arrow) in the vicinity of the nest. Thick lines on the scale at 10 cm intervals. N. Sagara and others 1023

TIC: ST0914–1.D 33·86 Hebeloma radicosoides 100000

80000

60000

25·59 Abundance 40000

20000

0 5 101520253035 Time

TIC: 92100904.D 26·95 Hebeloma radicosum 14·59 NHCHO 700000 26·32 26·05 600000 OH 5·88 500000 CH2CHO 23·37 34·62 27·93 400000

9·88 Abundance 300000

200000 14·25 28·54 32·06 7·76 100000 4·28 12·42 15·97 28·12

0 5 10 15 20 25 30 35 Time Fig. 22. Gas chromatograms of ethyl ether extracts from Hebeloma radicosoides (top) and H. radicosum (bottom), showing little production of volatile substances with a low-retention time in the former and much production in the latter. The substances identified for the latter are 1-octen-3-ol (Rt l 5n88), phenylacetaldehyde (Rt l 9n88) and N-formylaniline (Rt l 14n59).

ACKNOWLEDGEMENTS Bresadola, J. (reprint 1982) Iconographia Mycologica, Vol. IV. Massimo Candusso, Saronno. Thanks are due to Mr Sakae Takayama, Ms Yasue Marunishi, Dr H. Okabe, Bruchet, G. (1970) Contribution a l’e! tude du genre Hebeloma (Fr.) Kummer; Mr Kazumasa Yokoyama and Dr Y. Suzuki for supplying specimens, to partie spe! ciale. Bulletin Mensuel de la SocieTteT LinneTenne de Lyon 39 (6) Mr E. Nagasawa for discussion on basidiospore morphology of H. radicosoides. supplement: 1–132. and to the Editor Dr I. C. Tommerup and anonymous referees for improving Fujimoto, H., Katano, Y. & Yamazaki, M. (1992) Isolation, identification and the manuscript. The senior author (N.S.) is grateful to Drs D. A. Reid (Kew), pharmacological studies on three toxic metabolites from a mushroom, R. Watling (Edinburgh), M. Moser (Innsbruck) and R. L. Shaffer (Ann Arbor) Hebeloma spoliatum. Chemical and Pharmaceutical Bulletin 40: 869–872. for allowing the survey of their herbarium or personal specimens, and again Fukiharu, T. & Hongo, T. (1995) Ammonia fungi of Iriomote Island in the to Drs Watling and Moser for personal discussions on the taxonomy of some southern Ryukyus, Japan and a new ammonia fungus, Hebeloma luchuense. Hebeloma species. N.S. wishes to dedicate this paper to the late Prof. Emeritus Mycoscience 36: 425–430. Dr E. J. H. Corner (Cambridge, UK), who died on 14 Sep. 1996, for constant Gillet, C.-C. (1891) Champignons de France. Les HymeTnomyceZ tes, 3rd Part. A. encouragement throughout his last 30 years, and to Profs. Emeritus Drs Herpin, Alançon. R. P. Korf (Ithaca) and K. Tubaki (Tsukuba) for stimulation. Imazeki, R. & Hongo, T. (1965) [Coloured Illustrations of Fungi of Japan, Vol. II.] Hoikusha, Osaka. [In Japanese.] Imazeki, R. & Hongo, T. (1987) [Coloured Illustrations of Mushrooms of Japan, REFERENCES Vol. I.] Hoikusha, Osaka. [In Japanese.] Imazeki, R., Otani, Y. & Hongo, T. (1988) [Fungi of Japan.] Yama-Kei, Tokyo. Bernardi, M. de, Fronza, G., Gianotti, M. P., Mellerio, G., Vidari, G. & Vita- [In Japanese.] Finzi, P. (1983) Fungal metabolites XIII: New cytotoxic triterpene from Kaminski, E., Stawicki, S. & Wasowicz, E. (1974) Volatile flavor compounds Hebeloma species (Basidiomycetes). Tetrahedron Letters 24: 1635–1638. produced by molds of Aspergillus, Penicillium, and Fungi Imperfecti. Applied Bon, M. (1979) Fungorum Rariorum Icones Coloratae, Part II. J. Cramer, Vaduz. Microbiology 27: 1001–1004. Ammonia fungus Hebeloma radicosoides sp. nov. 1024

Kikuchi, T., Kadota, S. & Nishi, A. (1984) Odorous metabolites of an acellular Sagara, N. (1978) The occurrence of fungi in association with wood mouse slime mold, Physarum polycephalum Schw., and a basidiomycete, Phallus nests. Transactions of the Mycological Society of Japan 19: 201–214. impudicus Pers. Chemical and Pharmaceutical Bulletin 32: 797–800. Sagara, N. (1984) On ‘Corpse Finder’. McIlvainea 6(2): 7–9. Kornerup, A. & Wanscher, J. H. (1967) Methuen Handbook of Colour, 2nd edn. Sagara, N. (1989) [Mushrooms and Animals.] Tsukiji-shokan, Tokyo. [In Eyre Methuen, London. Japanese.] Kuroyanagi, E., Honda, S., Yoshimi, S. & Sagara, N. (1982) [The appearance Sagara, N. (1992) Experimental disturbances and epigeous fungi. In The Fungal of Hebeloma radicosum from a buried cat carcass.] Transactions of the Community, 2nd edn. (G. C. Carroll & D. T. Wicklow, eds): 427–454. Mycological Society of Japan 23: 485–488. [In Japanese with English Marcel Dekker, New York. summary.] Sagara, N. (1995) Association of ectomycorrhizal fungi with decomposed Lange, J. E. (1935–1940) Flora Agaricina Danica. 5 vols. Copenhagen. animal wastes in forest habitats: a cleaning symbiosis? Canadian Journal of Lange, L. (1974) The distribuiton of macromycetes in Europe. Dansk Botanisk Botany 73 (Supplement 1): S1423–S1433. Arkiv 30(1): 1–105. Sagara, N., Kitamoto, Y., Nishio, R. & Yoshimi, S. (1985) Association of two Lewis, D. P. & McGraw, L., Jr. (1984) Studies on Big Thicket . The Hebeloma species with decomposed nests of vespine wasps. Transactions of Southwestern Naturalist 29: 257–264. the British Mycological Society 84: 349–352. Moser, M., Ju$ lich, W. & Furrer-Ziogas, C. (1985–) Colour Atlas of Basidiomycetes. Sagara, N., Okabe, H. & Kikuchi, J. (1993) Occurrence of an agaric fungus Gustav Fischer, Stuttgart. Hebeloma on the underground nest of wood mouse. Transactions of the Ohta, A. (1998) Fruit-body production of two ectomycorrhizal fungi in the Mycological Society of Japan 34: 315–322. Hebeloma Mycoscience genus in pure culture. 39: 15–19. Sagara, N. & Takayama, S. (1982) [Hebeloma radicosum appearing on the old Rea, C. (1922) British Basidiomycetae. Cambridge University Press, Cambridge, latrine of a boy-scouts’(?) camping site]. Nature Study 28: 99–100. [In UK. [Reprinted in 1968 by J. Cramer.] Japanese.] Ridgway, R. (1912) Color Standards and Color Nomenclature. Published Singer, R. (1986) The Agaricales in Modern Taxonomy, 4th edn. Koeltz Scientific privately: Washington, D.C. Books, Koenigstein. Sagara, N. (1973) Proteophilous fungi and fireplace fungi (A preliminary Smith, A., Evenson, V. S. & Mitchel, D. H. (1983) The Veiled Species of report). Transactions of the Mycological Society of Japan 14: 41–46. Hebeloma in the Western United States. The University of Michigan Press, Sagara, N. (1975) Ammonia fungi – a chemoecological grouping of terrestrial Ann Arbor. fungi. Contributions from the Biological Laboratory Kyoto University 24: Smith, W. G. (1908) Synopsis of the British Basidiomycetes. British Museum, 205–276. London. Sagara, N. (1976) [Supplement to the studies of ammonia fungi (1).] Transactions of the Mycological Society of Japan 17: 418–428. [In Japanese with English summary.] Corresponding Editor: I. Tommerup