Karstenia 42: 39-48, 2002

Amylolepiota, Clavicybe and , new genera of the

HARRI HARMAJA -

HARMAJA, H. 2002: Amylolepiota, Clavicybe and Cystodermella, new genera of the Agaricales. - Karstenia 42: 39-48. Helsinki. ISSN 0453-3402. lignicola P.Karst., is referred to Amylolepiota Harmaja, n. gen., as Amylolepi­ ota lignicola (P.Karst.) Harmaja, n. comb. Clitocybe clavipes (Pers. : Fr.) P.Kumm. (as the type) and two related are separated from the externally very similar Clitocybe (Fr.) Staude to form a new genus, Clavicybe Harmaja. Clavicybe differs from Clitocybe: (i) the spore surface appears rough ith a high magnification, the anatomy de iates as (ii) the hygrophanity of the fruit body is different and (iii) the gill trama is irregular. A key to the species is given. 3 new nomenclatural combinations in Clavi­ cybe are made: C. avellaneialba (Murrill) Harmaja, C. clavipes (Pers. : Fr.) Harmaja, and C. squamulosoides (P.D.Orton) Harmaja. A third new genus, Cystodermella Har­ maja, is described for a group of species with inamyloid spores segregated from Cysto­ derma Fayod. 12 new nomenclatural combinations in Cystodermella are made: C. adnatifolia (Peck) Harmaja, C. ambrosii (Bres.) Harmaja, C. cinnabarina (Alb. & Schwein. : Fr) Harmaja, C. contusifolia (Pegler) Harmaja, C. cristallifera (Thoen) Harmaja, C. elegans (Beeli) Harmaja, C. granulosa (Barsch : Fr.) Harmaja (type species of Cystodermella), C. japonica (Thoen & Hongo) Harmaja, C. luteohemi­ sphaerica (Dennis) Harmaja, C. myriadocystis (Heinem. & Thoen) Harmaja, C. sipar­ iana (Dennis) Harmaja, and C. subpurpurea (A.H.Sm. & Singer) Harmaja. Special attention was paid to correct nomenclature and author citations. Key words: amyloidity, arthrospores, Clitocybe, , Finland, Floccularia, gill trama, hygrophanity, Lepiota, nuclear DNA content, , Harri Harmaja, Botanical Museum, Finnish Museum of Natural History, PO. Box 47, F/N-00014 University of Helsinki, Finland www.helsinki.fi/people/harri.harmaja/

AMYLOLEPIOTA I am lucky enough to be among the very few peo­ synonymous with L. lignicola as claimed by ple who have seen the unusual Lepiota lig­ Knudsen (1980). nicola P.Karst. in the nature. This species was described in 1879 from southern Finland. Accord­ Agaricalium generis Lepiotae affinis. Ab ea dif~ ing to my judgement, L. lignicola must be classi­ fert sporis amyloideis habitationeque ligno. fied in a separate, new genus as described below. Epicutis e cellulis elongatis instructa. Raris­ sime occurrit in silvis borealibus vetustis. - Ty ­ pus: Amylolepiota lignicola (P.Karst.) Harmaja Amylolepiota Harmaja, n. gen. (Lepiota lignicola P.Karst.). ?Lepiota sect. Amyloideae Singer, Ann. Mycol. 41: 171. 1943 . - Holotype: Lepiota amyloidea medium-sized. Universal veil appar­ Singer. This synonymy is true if L. amyloidea is ently present; presence of partial veil unclear. Pi- 40 HARMAJA, H.: NEW AGARIC GENERA KARSTENIA 42 (2002) leus convex to plane; surface dry, with appressed having rather broadly adnate lamellae: see the to squarrose brown scales. without ring; original illustration of the type species, Agaricus surface dry, below the ring zone with squarrose decorosus Peck (Peck 1873) [Romagnesi (1989) scales like those of pileus. Lamellae free, whitish. suggested that L. lignicola and A. decorosus Flesh white, not changing. Odour not distinctive. would be identical]. In Cystoderma Fayod s. str., Taste mild. Spores pure white in fresh deposit as restricted by me in this paper (see below), the (pale yellow in age), binucleate, ellipsoid; wall universal veil is essentially composed of sphaer­ without germ-pore, hyaline, smooth, distinctly ocysts and the lamellae are not free but adnate. amyloid, cyanophilic, but carminophobic. Basid­ Most probably the new genus contains a sin­ ia with basal clamp, without carminophilic/si­ gle species only; the necessary new combination derophilic granulation; wall carminophobic, with is made below. However, Cystoderma carpati­ a thin cyanophilic inner layer towards apex. cum M.M.Moser, an enigmatic based on Cheilocystidia present, inconspicuous; pleuro­ a single specimen from Poland (Moser 1979), cystidia lacking. Epicutis, also in scales, of cells should be compared when more material is avail­ which are distinctly longer than wide; pigment able. essentially membranal. Hymenophoral trama reg­ ular. Clamp connections present in hyphae of Amylolepiota lignicola (P. Karst.) Harmaja, n. fruit body. Dried pileus, stipe, lamellae, basal comb. mycelium, flesh and spore deposit do not exhibit Coloured illustrations: Karsten 1883: fig. 1; Kor­ fluorescence but retain their colours under ultra­ honen 1991: Sienilehti 43(1 ), back cover violet light (with wave-lengths 254 nm and 366 nm), excepting the tinges of the three first-named Basionym: Lepiota lignicola P.Karst., Bidrag structures which become slightly deeper. Habitat Kannedom Finlands Natur Folk 32 [Ryssl. Hatts­ on decaying wood (always of Betula ?) in forest; vamp. 1]: 547. 1879. - Cystolepiota lignicola causes white rot. (P.Karst.) Nezdojm., Mikol. Fitopatol. 14: 389. The above description is based on my studies 1980.- Holotype: Finland, Etela-Hame, Tammela, of fresh and dried fruit bodies of the type spe­ Mustiala, in ligno mucido betulae, IX.1879 PA. cies. Included are the first reports of the respons­ Karsten (H: Herb. P.A. Karsten no. 2034). -Other es of its basidiocarp to ultraviolet light and of its specimens examined (also when fresh): Finland, spores and basidia to the reagents methyl (cot­ Etela-Hame, Lammi, Evo, virgin forest 'Kotisten ton) blue and acetocarmine. aarnialue', ll.IX.l973 H. Harmaja (H); exactly Amylolepiota is closely related to the agaric the same place, ll.IX.l973 U. Kurimo (H). genus Lepiota Gray (type species L. clypeolaria ?Lepiota amyloidea Singer, Ann. Mycol. 41: (Bull. : Fr.) P.Kumm.); the fruit bodies remind 171. 1943.-Holotype: Russia, Altay Republic (as those of the species of sect. Echinatae Fayod of 'Oirotia, Altai'), Lake Teletskoe (LE, n.v.). Knud­ the latter genus. The lamellae are likewise free in sen ( 1980) synonymizes this species with L. lig­ the new genus, but their proximal ends reach nicola. closer to the stipe apex than in Lepiota. The deci­ sive differences are the amyloid spore wall and A detailed description of the species is given by the lignicolous habit of Amylolepiota. In Lepiota Knudsen (1980, as Lepiota). the spore wall is dextrinoid and the species are In Finland A. lignicola possesses a specific saprophytes on bare humous soil or in non­ habitat ecology: the species is restricted to mesic woody litter. However, L. zenkeri Henn. has an old-growth forest in the southern boreal zone; amyloid spore wall; it has been included in a sec­ the substrate is decaying wood, a fallen log or tion of its own (Singer 1986). L. zenkeri does not larger branch of Betula lying on the ground (it is appear to belong to Amylolepiota: it occurs in unknown whether B. pendula, B. pubescens or tropical Africa on non-ligneous substrate. Cys­ both are concerned). The species is classified as tolepiota Singer differs from Amylolepiota by critically endangered (CR) in Finland according possessing uninucleate inamyloid spores, a pi­ to the latest Red List of Finland (Rassi & al. 2001). leus cortex of sphaerocysts, and non-woody During the courses of the Universi­ habitat. Leucopholiota (Romagn.) O.K.Mill., ty of Helsinki, arranged at Lammi (S. Finland), the T.J. Yolk & Bessette (Miller & al. 1996) differs by fungus was found in the same locality at Kotinen KARSTENIA 42 (2002) HARMAJA, H.: NEW AGARIC GENERA 41

Virgin Forest in fi ve successive years (1972-1976), antabuse-like reaction (i .e., that of the disulfiram­ probably on the same decaying Betula log! type) results when fruit bodies heated for food In total, specimens examined by me or litera­ are consumpted with alcohol (e.g. Cochran & ture reports that appear reliable show that A. lig­ Cochran 1978, Bresinsky & Bes11985). nicola has been found (i) in three localities in the In a very recent phylogenetic analysis of a inland of South Finland (in the province of EteHi­ great number of and related species Hame, latest in 1985; according to specimens in (Moncalvo & al. 2002), the position of C. cla­ H), (ii) in the Republic of Karelia, Russia (Koti­ vipes was somewhat peculiar which might sup­ ranta & a!. 1998), and (iii) in several localities port the establishment of a separate genus for throughout the Asian part of Russia (Singer 1943 the species. if the synonymy indicated by Knudsen [1980] is Two additional species are included in Clavi­ approved), Vasil' eva 1973, Nezdojminogo 1982, cybe: Clitocybe avellaneialba Murrill and Clito­ Petrov 1986, Kutafyeva 1989 [recorded as L. hys­ cybe squamulosoides P.D.Orton. The affinity of trix Moll. & J.Lange but most probably referring the former species to Clitocybe clavipes was ear­ to the present species], Astapenko & Kutafyeva lier stated by me (Harmaja 1969); the latter was 1990, Lapitskaya 1993, Gorbunova 2001). To my previously thought by me to represent a form of judgement, the records from the U.S.A. (Knud­ C. clavipes (Harmaja 1970). sen 1980) and Mexico (Cifuentes & al. 1989, as L. amyloidea) require further study, as does the lig­ Clavicybe Harmaja, n. gen. nicolous 'Lepiota hispida Lasch' from Italy Clitocybe subg. Clitocybe sect. Clavipedes Har­ (Bresadola 1927). maja, Karstenia 9: 58. 1969. - Type: Clitocybe clavipes (Pers. : Fr.) P.Kumm. CLAVICYBE Clitocybe subg. Infundibuliformes sect. /nor­ natae subsect. Albiphyllae H.E.Bigelow, Beih. A monograph on the genus Clitocybe (Fr.) Staude Nova Hedwigia 72: 193. 1982.-Type: Clitocybe in Fennoscandia (northwestern Europe) consti­ avellaneialba Murrill. tuted my Ph.D. thesis (Harmaja 1969). In that con­ Clitocybe subg. Hygroclitocybe Bon, Doc. tribution, I reported 43 species from the study Mycol. 13(51): 9. 1983.- Type: Clitocybe cla­ area; they were placed in 3 subgenera and 16 sec­ vipes (Pers.: Fr.) P.Kumm. tions, and I considered the genus clearly hetero­ geneous. C. clavipes (Pers. : Fr) P.Kumrn. was as­ Agaricalium genus. Genus Clitocybem in men­ signed to a section of its own. Later I separated a tern revocat. Ab ea inter alia differt sporis sub­ part of the species and transferred them to Lepista tilissime rugulosis tramaque lamellarum e (Fr.) W.G.Sm. or Singerocybe Harrnaja (Harmaja hyphis intricatis instructa. - Typus: Clavicybe 1974a, 1974b, 1976, 1988). Evenaftertheseproce­ clavipes (Pers. : Fr.) Harmaja (Agaricus clavipes dures, Clitocybe still did not appear natural. Pers. : Fr.). Among the Fennoscandian species, Clito­ cybe clavipes was found by me (Harmaja 1969) Agaricales; habitus of basidiocarp as in Clito­ unique in four respects: (i) the pileus is not hy­ cybe. Veil completely absent. Pileus from convex grophanous though the flesh has this character, to plane or somewhat depressed, not hygropha­ (ii) the wet cap flesh begins to dry and fade from nous, brown or grey-brown; surface dry. Stipe the periphery towards the centre, (iii) the trama usually clavate, concolorous with pileus; surface of the lamellae is constructed of interwoven hy­ dry. Lamellae decurrent, whitish. Odour sweetish phae (vs. parallel or subparallel), and (iv) the or indistinct. Taste mild. Flesh hygrophanous; spores contain a distinct, large, refractive oil cap flesh dries from the periphery towards the drop. As in addition Pegler & Young (1971) centre. Spores pure white in fresh deposit (pale found that, with a high magnification under the yellow in age), not sticking to tetrads or with col­ electron microscope (they used the carbon repli­ lapsed walls in mounts made of dry gills, fairly ca method), the spores of this species have a large, a proportion of them always broadly fusi­ roughened surface, I planned to create for C. cla­ form; uninucleate (Kuhner 1945); wall without vipes a genus of its own in the 1970's. Moreover, germ-pore, hyaline, inamyloid, cyanophobic C. clavipes has been found "poisonous": the (Singer 1972, Harrnaja 1974a, 1976, 1979b), rugu- 42 HARMAJA, H.: NEW AGARIC GENERA KARSTENIA 42 (2002) lose under the electron microscope at least in the Discussion type species (Pegler & Young 1971, Bigelow Macroscopically, the species of this new genus 1981); contents with one distinct oil drop; hilar are very similar to the larger non-hygrophanous appendix large, ca. 0.9-1.1 x 0.7-0.9 !J.m. Cystidia species of Clitocybe (sensu Harmaja 1976). As of any kind absent. Pileus cortex made of epicutis mentioned above, Clavicybe differs from Clito­ and subcutis of subparallel hyphae; pigment es­ cybe in two fundamental points: (i) the spore wall sentially situated within cells of epicutis. Hymen­ is rough when observed with the electron micro­ ophora1 trama irregular, i. e., composed of inter­ scope (at least in the type species) vs. smooth woven hyphae (textura intricata). Clamp con­ and (ii) the hyphae of the gill trama are interwo­ nection abundant everywhere in the basidiocarp. ven vs. running in parallel-subparallel direction Dried pileus, stipe, lamellae, basal mycelium and as in Clitocybe. Also Bigelow (1982) describes flesh and of the type species do not exhibit fluo­ the gill trama of C. clavipes as interwoven, like­ rescence but retain their colours under ultravio­ wise that of C. avellaneialba (as does Smith let light (with wave-lengths 254 nm and 366 nm), 1949). Moreover, there are two differences in the excepting the tinges of the three first-named hygrophanity of the fruit body as described in structures which become slightly deeper. Sapro­ the introductory section above. These unusual phytes which decay plant litter, especially nee­ properties of hygrophany are apparently related dles and leaves, sometimes herbaceous litter or to each other and they may ultimately result from woody substrates. some anatomical peculiarity of the whole fruit body. The interwoven gill trama may also be in­ Clavicybe avellaneialba (Murrill) Harmaja, n. volved. comb. The direction in which the hyphae are running Basionym: Clitocybe avellaneialba Murrill, in the trama of the lamellae is diagnostic in the Mycologia 5: 207. 1913. - Material from the Agaricales, especially at the generic level, e.g. U.S.A. studied (Harmaja 1969). characterizing the genus Camarophyllus (Fr.) P.Kumm. (Singer 1986). Clavicybe clavipes (Pers. :Fr.) Harmaja, n. comb. Of the three species included in Clavicybe, C. clavipes is distributed over a wide area in the Basionym: Agaricus clavipes Pers., Syn. meth. northern hemisphere, C. avellaneialba is re­ fung. : 353.1801: Fr., Syst. mycol. 1: 86. 1821.­ stricted to western North America, and the poor­ Clitocybe clavipes (Pers. : Fr.) P.Kumm., Fiihr. ly known C. squamulosoides has been found in Pilzk.: 124. 1871.- Omphalia clavipes (Pers.: Fr.) temperate western Europe. In addition, a fungus Que!., Enchir. fung. 20. 1886. which is sometimes called as "the pale-brown form of Clitocybe clavipes" occurs in the tem­ Clavicybe squamulosoides (P.D.Orton) Harmaja, perate deciduous woods of Central Europe. A n. comb. good coloured photograph of the last-named Basionym: Clitocybe squamulosoides P.D .Orton, fungus was published by Phillips (1981, p. 48, as Trans. Brit. My col. Soc. 43: 187. 1960. - Isotype Clitocybe clavipes). This taxon may be a fourth examined (Harmaja 1970): U.K., England, Surrey, species of Clavicybe, or it falls within the varia­ Witley Common, 13.XII.1956 P.D. Orton (H). bility of C. squamulosoides. Also the typical dark grey-brown C. clavipes decays, besides needles, leaf litter, often leaves of Betula; I my­ self have found it in the leaves of the beech (Fa­ gus sylvatica) near Femsjo, Sweden. KARSTENIA 42 (2002) HARMAJA, H.: NEW AGARIC GENERA 43

A key to the species of Clavicybe

1 Spores ca. 6-9 x 3.5- 5.3 flm, variable in shape; on litter of needles or leaves ...... 2 1' Spores ca. 8-10 x 4.0-5.5 flm, broadly fusiform; mostly on much decayed woody substrates (western North America) ...... C. avellaneialba

2 Cap and stipe dark in colour, grey-brown or sepia brown; stipe distinctly clavate; in litter of needles or leaves; widely distributed ...... C. clavipes 2' Cap and stipe pale brown or avellaneous; stipe hardly enlarged downwards; mostly in leaf litter; temperate woods of western Europe ...... C. squamulosoides

CYSTODERMELLA Fayod ( 1889) included partly unrelated species in dermella Harmaja (type C. granulosum) is de­ his new genus Cystoderma Fayod (see Smith & scribed below for the latter species group. Singer 1945). Heinemann & Thoen (1973) delimit the genus in a most appropriate way, and their Cystodermella Harmaja, n. gen. delimitation is essentially the current one. An universal veil possessing a thick outer layer of Agaricus subg. Lepiota [sect.?] Granulosi Fr., spherocysts is the conspicuous main diagnostic Hymenomyc. eur. : 35 . 1874. - Lepiota [sect.?] character of the genus. However, since the 1930's Granulosae (Fr.) Quel., Enchir. fung .: 7. 1886. ­ it has been known that the species of the genus Cystoderma sect. Granulosa (Fr.) Locq., Bull. can be assorted to two groups: those with amy­ Soc. Linn. Lyon 14: 88. 1945.- Cystoderma subg. loid spores and those with an inamyloid spore Granulosa (Fr.) Wasser, Fl. grib. Ukrainy: 209. wall. The lectotype of the genus is C. amianthi­ 1980. -Type: Agaricus granulosus Batsch: Fr. num (Scop.) Konrad & Maubl. with amyloid Cystoderma sect. Cinnabarina Heinem. & spores (selected by Smith & Singer 1945). Gener­ Thoen, Bull. Soc. Mycol. France 89: 23. 1973. - ally, the genus has been divided into two sec­ Type: Agaricus cinnabarinus Alb. & Schwein. :Fr. tions: sect. Cystoderma and sect. Granulosa (Fr.) Locq. (type C. granulosum (Batsch : Fr.) Agaricalium generis Cystodermatis proxima. Kuhner) (see e.g. Singer 1986). Ab eo inter alia differt tunica sporarum totaliter For decades, I have personally been interest­ inamyloidea. - Typus: Cystodermella granulo­ ed in Cystoderma and made research on the ge­ sa (B atsch : Fr.) Harmaja (Agaricus granulosus nus. Now I have (i) evaluated the taxonomic Batsch :Fr.). weight of the spore difference within Cystoder­ ma mentioned above, (ii) considered some previ­ Agaricales; habitus of basidiocarp as in Cysto­ ously unused, less exact differences or tenden­ derma. Universal veil present. Pileus convex or cies present in the genus (see Table 1), (iii) ap­ plane; surface finely to coarsely granulose. Stipe plied the results of a recent contribution on the covered with veil remnants except at apex; ring nuclear DNA content of some species (Saar & mostly lacking, sometimes present. Lamellae nar­ Kullman 2000) as well as (iv) taken into consider­ rowly to broadly adnate. Flesh of stipe and pileus ation the results of the phylogenetic analysis of of continuous elasticity. Conidia (arthrospores) Moncalvo & al. (2002). My judgement, based on absent from context of pileus. Odour indistinct. the above, is that Cystoderma, as currently un­ Taste mild. Spores white in deposit, binucleate derstood (Heinemann & Thoen 1973, Singer (Saar & Kullman 2000: three species examined, 1986), has to be split into two genera: Cystoder­ the type included), ellipsoid; wall without germ­ ma s. str. will comprise species that possess amy­ pore, smooth, inamyloid, cyanophilic (at least in loid spores while a new genus has to be estab­ type species: Singer 1972). Cystidia of lamellae lished for species with inamyloid spores. Cysto- mostly absent; when present they are 'harpoon- 44 HARMAJA, H.: NEW AGARIC GENERA KARSTENIA 42 (2002) like' due to encrusted crystals. Cortex of pileus violet light (with wave-lengths 254 nm and 366 and stipe (apex excluded) formed by dry velar nm; no difference to C. amianthinum; these are layer essentially composed of sphaerocysts. Hy­ apparently the first observations on the respons­ menophoral trama of more or less parallel hy­ es of Cystoderma s. lato fruitbodies to ultraviolet phae. Clamp connections present in hyphae of light). Saprophytes of non-ligneous, rarely ligne­ fruit body. Dried pileus, stipe, lamellae, flesh and ous, plant litter; sometimes among mosses at the basal mycelium of the type species do not exhibit same time but probably not truly muscicolous. fluorescence but retain their colours under ultra-

Comparison of the genera Cystoderma and Cystodermella

Table 1. A comparison of Cystoderma Fayod and Cystodermella Harmaja. The exact differences are given in bold. See the text immediately below for details and references.

Cystoderma Cystodermella

Spore wall in Melzer's amyloid inamyloid 'Harpoon' cystidia absent present in some species Arthrospores in fruit body and/or Present in some species (e.g. type) unknown mycelium culture Liability to Squamanita type species susceptible resistant Bryophily often strong often not apparent Ploidy level 2x 3x or 6x (both type species studied) Phylogeny type species clusters near type species clusters Floccularia Pouz. near Ripartitella Sing.

The spore wall of C. superbum Huijsman is com­ amyloid spores, contained higher amounts of monly reported to be amyloid in a small area only, DNA: C. adnatifolium (Peck) Harmaja and C. above the hilar appendix (e.g. Heinemann & granulosum were on 3x level and C. cinnabari­ Thoen 1973). However, according to the obser­ num (Alb. & Schw. : Fr) Konrad & Maubl. (as C. vations of Pegler & Young (1971) the spore wall terreii (Berk. & Broome) Harmaja) was on 6x level. is weakly amyloid throughout, but the area men­ The species of Squamanita Imbach (Agari­ tioned (the plage) is strongly amyloid. cales) parasitize other agarics. They develop chi­ Kuhner (1969) reported the occurrence of co­ meric which are mixtures of the nidia (arthrospores) in the flesh of the fruit body structures and tissues of the fruit bodies of the and/or in the mycelium culture of some Cystoder­ host and the parasite (Harmaja 1988, Redhead & ma taxa with amyloid spores. Heinemann & Thoen a!. 1995). Observations on these chimera fruit­ (1973) and Harmaja (1979) likewise noted the pres­ bodies and the agaric species growing in their ence of the arthrospores in the basidiocarps of immediate neighbourhood suggest that always some amyloid-spored species, including the type. when a Cystoderma s. lato is contaminated with Saar & Kullman (2000) analyzed the nuclear Squamanita, it is C. amianthinum (Smith & DNA content (genome size, ploidy level, C value) Singer 1948, Watling 1974, Reid 1983, Harmaja of six species of current Cystoderma from the 1988, Lange & Liess¢e 1989, Stridvall & Stridvall spore nuclei. They found that all three species 1994, Redhead & al. 1995) or C. carcharias (Erik­ with amyloid spores (C. carcharias (Pers.) Kon­ sen 1997). rad & Maubl., C.jasonis (Cooke & Massee) Har­ In the woods or on rock outcrops, species of maja and C. amianthinum, the type of the genus) Cystoderma s. str., in particular, very often grow were on 2x ploidy level while the nuclei of three in a moss carpet with their basal mycelium tightly other species, belonging to the group with in- attached to living mosses. My field experience KARSTENIA 42 (2002) HARMAJA, H.: NEW AGARIC GENERA 45 even suggests that two species, at least (C. lilaci­ Smith & Singer (1945), Heinemann & Thoen pes Harmaja and C. saarenoksae Harmaja) are (1973; some species are treated in sect. Cinnaba­ obligately associated with species of Polytri­ rina Heinem. & Thoen) and Singer ( 1986). chum s. str. (Harmaja 1979a, 1985). At least 12 species of Cystodermella are well In the comprehensive phylogenetic work of defined, the area of the genus extending from the Moncalvo & al. (2002) Cystoderma amianthi­ tropical to the arctic zone. Cystoderma s. str. num and C. chocoanum Franco-Molano (like­ comprises roughly 17 species at present; the dis­ wise with amyloid spores) clustered with Floccu­ tribution of the genus is subcosmopolitan and laria albolanaripes (G.F.Atk.) Redhead (with extends from the arctic through the tropics to amyloid spores) while C. granulosum clustered subantarctic regions. In both genera, north tem­ in another clade with Ripartitella brasiliensis perate and boreal zones with a not too continen­ (Speg.) Singer (with inamyloid spores and often tal climate display the greatest diversity known harpoon cystidia). at present; my observations suggest that several undescribed species exist. For treatments of Discussion world species, consult Smith & Singer (1945), Heinemann & Thoen (1973) and Singer (1986). The presence of a partial veil in Cystodermella Those species are transfetTed to the genus remains to be ascertained. In C. adnatifolia, in Cystodermella which are familiar to me or which particular, a whorl of white hyphae remains at the appear distinct enough and well described in the 'ring zone' of the stipe. It is unclear whether this literature. tissue represents a separate partial veil or is formed from the inner layer of the universal veil. New combinations in Cystodermella Besides Cystoderma s. str., also Ripartitella Singer is closely related to Cystodermella. Ri­ Cystodermella adnatifolia (Peck) Harmaja, n. partitella differs from Cystodermella: (i) a per­ comb. fect universal veil is lacking, (ii) the cortical lay­ Basionym: Lepiota adnatifolia Peck, Bull. New ers of the pileus and stipe are composed of elon­ York State Mus. Nat. Hist. 54: 947. 1902. -Armil­ gated cells instead of sphaerocysts, and (iii) the laria adnatifolia (Peck) Kauffman, Pap. Michi­ spores are verruculose (Singer 1986). gan Acad. Sci. 2: 60. 1923.- Cystoderma granu­ The placement of current Cystoderma within losum var. adnatifolium (Peck) A.H.Sm. & Sing­ the Agaricales is somewhat problematic: it has er, Pap. Michigan Acad. Sci. 30: 90. 1945.- Cysto­ been included in the families , Lepio­ derma adnatifolium (Peck) Harmaja, Karstenia taceae and (e.g., Heinemann & 14: 122. 1974. Thoen 1973, Kuhner 1980, Singer 1986, Harmaja 1979a). Pegler & Young (1969, 1971) published two Cystodermella ambrosii (Bres.) Harmaja, n. interesting observations: in C. amianthinum the comb. structure of hilar appendix of the spore is of the nodulose type, and the spore wall appeared thin Basionym: Armillaria ambrosii Bres., Fungi Tri­ and got easily collapsed under vacuum when car­ dentini 1: 27. 1881. - Cystoderma ambrosii bon replicas were produced. As they pointed out, (Bres.) Singer, Ann. Mycol. 41: 170. 1943. both these characters of C. amianthinum show affinities to the fungi of Tricholomataceae with Cystodermella cinnabarina (Alb. & Schwein. their hyaline thin-walled spores rather than to Fr.) Harmaja, n. comb. Agaricaceae/Lepiotaceae. Unfortunately they did Basionym: Agaricus A. [unranked Lepiotae] not examine the spore ultrastructure of C. granu­ granulosus var. ee cinnabarinus Alb. & Sch­ losum or any other species with inamyloid spores. wein., Consp. Fung. Lusat.: 147. 1805. -Agaricus In the phylogenetic study of Moncalvo & al. cinnabarinus (Alb. & Schwein. : Fr.) Fr., Syst. (2002) the proper placements of the type species mycol. 3: Index: 12. 1832.- Lepiota cinnabarina of Cystoderma and Cystodermella among the (Alb. & Schwein. : Fr.) P.Karst. , Bidrag Kanne­ euagarics remained unresolved. dom Finlands Natur Folk 32 (Ryssl. Hattsvamp. For additional information of Cystodermella: 1]: 14. 1979. - Armillaria cinnabarina (Alb. & see the descriptions of the section Granulosa in Schwein. : Fr.) Kauffman, Pap. Michigan Acad. 46 HARMAJA, H.: NEW AGARIC GENERA KARSTENIA 42 (2002)

Sci. 2: 60. 1923. - Cystoderma cinnabarinum 4 [Nov. mycol. noviss.]: 57. 1947; = ?]- Cystoder­ (Alb. & Schwein. : Fr.) Konrad & Maubl., Icon. ma granulosum (Batsch : Fr.) KUhner, Botaniste sel. fung. 3(3): pl. 238. 1927. 17: 125. 1926. Agaricus terryi ('Terreii') Berk. & Broome, Ann. Mag. Nat. Hist. 4(6): 462. 1870.- Cystoder­ Cystodermella japonica (Thoen & Hongo) Har­ ma terryi (Berk. & Broome) Harmaja, Karstenia maja, n. comb. 18: 30. 1978. Basionym: Cystoderma japonicum Thoen & Bon ( 1999) correctly points out that, in the in­ Hongo, Trans. Mycol. Soc. Japan 26: 23 . 1985. dex of Systema mycologicum, published at the end of its third part, Fries used and even sanc­ Cystodermella luteohemisphaerica (Dennis) tioned the epithet cinnabarinus at the specific Harmaja, n. comb. level, so that epithet shall be used for the present species rather than terryi as I earlier (Harmaja Basionym: Cystoderma luteohemisphaericum 1979a) suggested. Dennis, Kew Bull. 15 : 109. 1961.

Cystodermella contusifolia (Pegler) Harmaja, n. Cystodermella myriadocystis (Heinem. & comb. Thoen.) Harmaja, n. comb. Basionym: Cystoderma contusifolium Pegler, Basionym: Cystoderma myriadocystis Heinem. Kew Bull., Addit. Ser. 9:410. 1983. & Thoen, Bull. Soc. Mycol. France 89 : 14. 1973.

Cystodermella cristallifera (Thoen) Harmaja, n. Cystodermella sipariana (Dennis) Harmaja, n. comb. comb. Basionym: Cystoderma cristalliferum Thoen, Basionym: Lepiota sipariana Dennis, Kew Bull. Bull.Jard. Bot. Belg. 39: 185. 1969. 7: 488. 1953.- Cystoderma siparianum (Dennis) Thoen, Bull. Jard. Bot. Belg. 39: 190. 1969.- Ri­ Cystodermella elegans (Beeli) Harmaja, n. comb. partitella sipariana (Dennis) Dennis, Kew Bull., Addit. Ser. 3: 58. 1970. Basionym: Armillaria elegans Beeli, Bull. Soc. Roy. Bot. Belgique 59: 111. 1927.- Cystoderma C. subpurpurea (A.H.Sm. & Singer) Harmaja, n. elegans (Beeli) Thoen, Bull. Jard. Bot. Belg. 39: comb. 188. 1969. Basionym: Cystoderma subpurpureum A.H.Sm. Cystodermella granulosa (Batsch : Fr.) Harmaja, & Singer, Mycologia 40: 457. 1948. n. comb.

Basionym: Agaricus granulosus Batsch, Elench. Acknowledgements: The Lammi Biological Station of fun g. l : 170. 1783: Fr., Syst. my col. 1: 24. 1821. ­ the University of Helsinki has provided me excellent Lepiota granulosa (Batsch : Fr.) Gray, Nat. ar­ research facilities during many years. Thanks are due to rang. Brit. pl. 1: 602. 1821. - Mastocephalus gran­ the curators of those herbaria which have lent material ulosus (Batsch : Fr.) Kuntze, Rev. gen. pl. 2: 860. for my studies during years. Professor Teuvo Ahti made useful improvements to the manuscript. Mr. Heino Van­ 1891. - Armillaria granulosa (Batsch : Fr.) ska, Lie. Phil., kindly checked the Latin of the generic Kauffman, Pap. Michigan Acad. Sci. 2:60. 1923. descriptions. [non Armillaria granulosa Vel., Opera Bot. Cech. KARSTENIA 42 (2002) HARMAJA, H .: NEW AGARIC GENERA 47

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