Phaeogalera and Galerina in arctic-subarctic Alaska (U.S.A.) and the Yukon Territory (Canada)

Transcription

1 Phaeogalera and Galerina in arctic-subarctic Alaska (U.S.A.) and the Yukon Territory (Canada) E. HORAK Herbarium. ETH, Zollikerstrasse 137, CH-8008 Zurich, Switzerlanrl AND 0. K. MILLER, JR.' Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, I/A , U.S.A. Received April 4, HORAK, E., and MILLER, 0. K., JR Phaeogalera and Galerina in arctic-subarctic Alaska (U.S.A.) and the Yukon Territory (Canada). Can. J. Bot. 70: Eleven taxa of Galerina and Phaeogalera are described. Galerina leptocystis, Galerina subarctica, and Galerina praticola are reported from arctic North America for the first time. Phaeogalra stagnina is only found in very humid, wet meadow tundra associated with Drepatzoclarlus or Calliergon. Galerina arctica is reported for the first time from Alaska and Canada. One species, Galerinapseudocerina, is found only in arctic alpine habitats in Canada and not in the arctic tundra. Two forms of Galerina pseudo~nycenopsis represent the most common taxon observed in Alaskan North Slope wet meadow tundra on peat or associated with Calliergon, Drepanocladus, and Sphagnum. Two species, Galerina clavata and Galerina hypnorun~, are common cosmopolitan taxa, but only G. clavata is frequently encountered on the Alaskan North Slope. The association of the Galerina taxa with mosses is presented and discussed, as well as their occurrence in microhabitats in wet meadow tundra and among polygons in coastal tundra on the Alaskan North Slope. Key words: Galerina, Phaeogalera, Cortinariaceae, Alaska, Yukon Territory, bryophytes. HORAK, E., et MILLER, 0. K., JR Phaeogalera and Galerina in arctic-subarctic Alaska (U.S.A.) and the Yukon Territory (Canada). Can. J. Bot. 70 : Les auteurs dccrivent 11 taxa de Galerina et de Phaeogalera. Le Galerina leptocystis, le Galerina subarctica et le Galerina praticola sont rapportts pour la premibre fois dans I'arctique de I'Amtrique du nord. Le Phaeogalera stagnina n'est retrouvc que dans les prairies de la toundra trbs humide, m&me inondte, associce avec le Drepar~ocladus ou le Calliergon. Le Galerina arctica est rapport6 pour la premibre fois pour 1'Alaska et le Canada. Une espkce, le Galerina pseurlocerina, n'est retrouvce que dans les habitats arctiques alpins du Canada et non dans la toundra arctique. Deux formes du Galerinapseudo~nycenopsis reprcsentent le taxon le plus commun observc sur le versant nord de I'Alaska, dans les prairies humides de la toundra associces avec des Calliergon, Drepanocladus et Sphagnum. Deux espkces, le Galerina clavata et le Galerina kypnorum, sont des taxons cosmopolites communs, mais seulement le G. clavata se retrouve frcquemment sur le versant nord de I'Alaska. L'association des taxons de Galerina avec les mousses est prcsentce et discutce aussi bien que leur prcsence dans les microhabitats des prairies de toundra humide et au sein des toundras c8tikres a polygones du versant nord de I'Alaska. Mots elks : Galerina, Phaeogalera, Cortinariaceae, Alaska, Territoire du Yukon, bryophy tes. [Traduit par la ridaction] Introduction Research was initiated under the U.S. Tundra Biome Program In 1970 (Brown and West 1970). The primary location was on the Arctic Coastal Plane in wet meadow tundra near Barrow, Alaska. The major sltes (sites 1, 2, 3, and 4) plotted in Flg. 3 of Brown and West (1970) were located along a moisture gradient. On sites 1, 2, and 4 a number of 6 x 6 m study plots were selected and marked for study. Plots were established within specific land features such as the polygonal centers and interpolygonal troughs (Miller and Laursen 1974; Rastorfer et al. 1974~). In addition, plots were situated in several areas of disturbance both natural and artificial. The Simpson Oil Seeps are natural phenomena, and the Schultz Fertilizer plots located at Barrow, Alaska are artificial disturbances. A commercial (N-P- K) fertilizer was applied to the Schultz site at a rate equal to 448 kglha (Brown and West 1970). Sites also were established in arctlc tundra at Prudhoe Bay on the North Slope and at Eagle Summit (Anderson 1974) within subarctic alpine tundra. Intensive study of the higher fungi was carried out by Miller and Laursen (1974, 1978) and Laursen (1975) on these sites. Extensive fieldwork also took place elsewhere in arctic tundra including Cape Simpson, 'Author to whom correspondence should be addressed. Meade River, Point Hope, Icy Cape, and Umiat on the Colville River. Studies were also done in subalpine tundra in 1967 and 1969 by 0. K. Miller in cooperation with the Arctic Institute of North America in the vicinity of Kluane Lake in the Yukon Territory of Canada. Study sites included the Skolai Pass at the headwaters of the White River, Slims subalpine tundra above the Slims River, and the Kaskawulsh Nunatak (Miller 1987) located within the Kaskawulsh Glacier in the Donjak Mountains. Colors were recorded when possible by comparison with the plates of Ridgway (1912). Ridgeway colors are given in parentheses, e.g., (vandyke brown). Collections are all deposited at Virginia Polytechnic Institute and State University (VPI) unless otherwise stated. General ecology Ten of the 11 species of Galerina and Phaeogalera were found, at least once, in the vicinity of Barrow in arctic tundra, with the exception of Galerina pseudocerina Smith & Singer. Galerina pseudocerina was recorded only in subalpine tundra at Eagle Summit north of Fairbanks and in the Skolai Pass on the Alaska-Yukon border. Three species Phaeogalera stagnirza (Fries) Pegler and Young, Galerina clavata (Velenovskj) Kiihner, and Galerina praticola (Moeller) Orton, are strongly

2 HORAK AND MILLER - m - West East - Old Beach Ridge Site (-1 - s4- Footprint Cr /- Distance (m) - Plot 450 (-) South Distance (m) - Plot FIG. 1. Typical landform and topography in a wet meadow - tundra community at Barrow, Alaska. (a) East-west profile. (b) North-south profile. I associated with three moss genera (Drepanocladus, Calliergon, and Campyliurn) and are only found in arctic tundra. Campyliutn also only occurs with the three genera of mosses listed above, but the number of reports are limited. Gnleritza pseudomycenopsis Pilit & Nannfeldt form 1 and G. pseudomycenopsis Pilit & Nannfeldt form 2 are associated with 6 and 12 genera of mosses, respectively, and can be characterized as generalists. Galerina subarctica Smith & Singer recorded only in arctic tundra is reported many times with Sphagnum (Gulden 1980) and was also occasionally associated with four other genera of mosses. There are few records of Galeritzn leptocystis Wells & Kempton, two collections of which were associated with Polytrichum. Galerina sp. 1, collected only.. once in the Schultz Fertilization Plots near Barrow, Alaska,. was growing with Meesia triq~ietra (Hook. & Tagl.) Aongstr. Galerina hypnor~im (Schrank:Fries) Kiihner has three records and was recorded with Sphagnum sp. and Dicranum sp. All species of Galerina and Phaeogalera reported from Alaska and Canada have a constant association with mosses. It is also clear that the majority of species of Galeritza have strong host preferences. In several cases the actual attachment of the fungus to the moss suggests that the anamorph is probably decomposing the dead and senescent portions of the moss. In other cases the growth was directly on the peat and rotten debris of unknown plants. Figure 1 illustrates the typical landform and topography within the wet meadow tundra community. The locations of six species of Galerina are typical examples of the microniches that were occupied. The harsh environment of the old beach ridge, polygon ridges (plots 422 and 428), and the wet, cold polygon troughs (plot 424) did not support Galerinas. Favored habitats included the wet meadow (416, 420, 428) and north exposures (421 and 425) that are often partly sheltered by dense grass, sedges, and Salix species. At Barrow, in the wet tundra meadow community, one finds that the bryomass is composed of 76.6% pleurocarpous mosses, 20.7 % acrocarpous mosses, and 2.7 % leafy liverworts (Rastorfer et al. 1974a, 1974b, 1974c, 1978). Thalloid liverworts are absent and sphagnum mosses are very rare. No species of Galerina were found associated with leafy liverworts, although liverworts were present on all of the plots sampled. Acrocarpous mosses are minor associates of Galeritza at Barrow and included three taxa: Dicranum sp., Otzcophorus vvahletzbergii Brid., and Polyrricl~utn cotnmune Hedw. Eleven genera of pleurocarpous mosses (Table 3 in Rastorfer et al. 1974~) were the most frequent in wet meadow tundra. Species of Galerinn were commonly associated with only 4 of the 11 genera. These included Otzcophorus, Camp~lliutn, Calliergotz, and Drepanocladus. The latter three genera of pleurocarpous mosses made up more than "69% of the total bryomass (on the average)" (Rastorfer et al. 1974~). These three moss genera were the dominant or only ones associated with Phneogalera stagnina, Galerina arctica (Singer) Nezdojminogo, G. praticola, and G. clavata. We have no records of association with the other seven moss genera. The abundant generalists, G. pseudotnj~cet~opsis forms 1 and 2, were associated with these three moss genera as well as other genera, including acrocarpous mosses. Galerina leptocystis, Gnlerina sp. 1, and G. subarctica were rarely found, and with one exception, only at the Barrow site, which was by far the most intensively collected area during our arctic field studies. In no case were these species of ~alerina found associated with the dominant genera of pleurocarpous mosses listed above. Galerina pseudocerinn was not found in the arctic tundra on the North Slope. Sphagnum is the only associated moss with G. pseudocerina in the subarctic sites of Eagle Summit and the Skolai Pass. Sphagnum is rarely encountered in the wet meadow tundra on the North Slope of Alaska (Rastorfer et al. 1974~). In conclusion, it appears that many of the species of Galerina from arctic and subarctic habitats have strong or specific associations with specific genera and species of mosses. In addition, these are most often pleurocarpous moss genera.

3 CAN. J. BOT. VOL Key to Alaskan and Yukon arctic-subarctic taxa of Phaeogalera and Galerina I. Spores smooth, perispore and plage absent, elliptic to distinctly almond-shaped; plcurocystidia absent Spores minutcly warted, or roughend to distinctly warted. perispore and plage distinctive, bean-shaped or oval to almond-shaped (oftcn with beaked rnucro); plcurocystidia present or absent Sporcs elliptic, x pni (-20 X - 10 pm on 2-spored basidia), thick-walled with distinctivc germ pore: clamp connections present; white veil remnants conspicuous on both pilcus and stipc.... I. Plzaeogalerer sragnitla 2. Sporcs distinctly amygdaliform or pyriform, occasionally minutely warted, X pnl (- 14 X -7.5 pm on 2-spored basidia), thin-wallcd, pale yellow-brown, oftcn collapsed, gcrm pore abscnt; clamp connections abscnt; veil rcmnants none Grileritzcr rrrctice~ 3. Spores phaseoliforrn, coarsely warted, perispore and plage absent, x 5-7 pm (- 16 x -8.5 pni on 2-spored basidia); cheilocystidia and caulocystidia lecythiform: clamp connections absent Grzleritla clri~~ertr~ 3.Spores not as above Spores smooth or occasionally minutely roughened, perispore and plage none, distinctly pyriform or amygdaliform x pm; cheilocystidia and caulocystidia lecythiform; clamp conncctions absent Grrlerir~cr arctic.rr 4. Spores warted, shape and (or) size not as above; clamp connections present Spores minutely, but distinctly warted, perisporc and plage present but usually inconspicuous Spores minutely to coarsely warted or marbled, perisporc and plage clearly visible and distinctive Spores x 5-6 pm Grrleritzn leptoq~.stis 6. Spores x 6-7(-7.5) pm Gnlo-irln sribrrrcticri 7. Spores with a conspicuous perispore forming loose to ear-shaped projections around distinctive plage: plcurocystidia a b: sent Spores without loose to ear-shaped perispore, marbled to subvcrrucose, rust brown, plagc distinct; cystidia oftcn with yellowbrown plasnlatic pigment Cheilocystidia x 4-11 pm, narrowly lagenifornl to fusoid, apex slightly swollen but nevcr capitate, hyaline; spores x 5-6 pm Gcil<!ritzci hyp~~oriirtl 8. Cheilocystidia x 7-15 pni, broadly fusoid, utriform, apex conic to subcapitate, often constrictcd, sometimes with yellow-brown plasmatic pigment; spores x pm Gcilerit~ci sp Cheilocystidia and caulocystidia lecythiforni, 25-45(-60) prn long, pleurocystidia absent; spores I1-14 X 7-9 pm, ovoid (to subamygdaliform), without beaked apex. germ pore obscure: stipe lacking conspicuous persistent veil remnants Gerlerit~rr p.se~reloceritlci 9. Cheilocystidia, plcurocystidia and caulocystidia fusoid to lagcniform with gradually tapering neck (- 100) pm long; gill edges often have balloon-shaped or vesiculose cells; spores 10-12(- 13) x 6-8 pm, ovoid or amydalifor~n (with or without beaked apex), germ pore distinct: stipe with pcrsistcnt, membranous ring (cortina) Basidiomes slender pileus - lo(- 15) mm diameter stipes solitary: spores subelliptic, rnucro indistinctive or abscnt; on or anlong (aquatic) moss (most oftcn on Drepcir~oclndti.~ spp. or Ctrlliergotz spp.) on frcq~lently inundated sites Gnlerbznprctticola 10. Basidiomes robust pileus up to 40 mm diameter; stipes solitary: spores ovoid to pip-shaped or aniydaliform; with many niosses on peator peaty soil I I. Spores predominantly ovoid to pip-shaped...%i. Geilerir~cr p.ser~dor~ryce~~~p.si.s (form I) I I. Sporcs predominantly amydaliforni with niore or less distinctive beaked apex or subelliptic b. Gnlerit~a pserrrlottzycetropsis (form 2) Description of taxa 1. P1zaeogale1.a stagnitza (Fries) Pegler & Young Figs. 2-4, 38 Kew Bull. 30: Agaricus stagnirzus Fries, Syst. Mycol. 1: = Plzaeogalera stagrzirza (Fries) Kiihner, Bull. Soc. Mycol. Fr. 88: (invalid, no basionym) ILLUSTRATIONS: Favre (1955); Gulden et al. (1985). DESCRIPTION (OKMIGAL 10932): Pileus up to 22 mm diameter, hemispheric to convex with incurved margin, disc often obtusely campanulate, in mature specimens becoming plane to depressed with upturned margin, color ranging from fuliginous to dark brown (russet to vandyke brown) or pale brown (Mars brown) depending on age and degree of desiccation, margin at first appendiculate from conspicuous white or pale brown fibrils or patches of veil that gradually disappear with age; surface transparently striate when moist, strongly hygrophanous, dry to subviscid, glabrous. separable pellicle absent. Lamellae broadly adnate to emarginate-subdecurrent, at first close becoming spaced with age, up to 3.5 mm wide; concolorous with pileus or paler (honey yellow, cinnamon brown to Dresden brown), edges white fimbriate or even. Stipe up to 45 x up to 3(-5) mm, cylindric, equal above, slightly tapering towards base or subclavate, occasionally compressed; concolorous with pileus or paler at apex and darker at base, apex subpruinose, with white, persistent fibrillose to submembranous veil remnants forming a distinctive belt, below with irregular zones of white fibrils; dry, solid becoming hollow with age. Context of pileus and stipe concolorous with surface of pileus. Odor and taste not distinctive. Spores x pm (from 4-spored basidia), up to 20 x up to 10 pm (from 2-spored basidia), broadly ovoid to elliptic, dark brown, smooth, thick-walled, with conspicuous apical germ pore, plage and perispore absent. Basidia x 8-11 pm, 4-spored, rarely also 2-spored. Cheilocystidia 30-70(-85) x 6-12(-16) pm, polymorphic, ranging

4 HORAK AND MILLER 4 17 in shape from cylindric to subfusoid or subcapitate, occasionally subglobose apex with or without irregular knob-like projections, neck sometimes with one to several constrictions, hyaline, thin-walled, pigment absent; sometimes mixed with clavate to balloon-shaped cells, x pm. Pleurocystidia absent. Caulocystidia rare, shape and size as in cheilocystidia. Pileipellis a cutis of interwoven, cylindric, nongelatinized hyphae, encrusted with rust brown pigment. Clamp connections present on all septa. MATERIAL EXAMINED: U.S.A.: ALASKA: Mead River: among Drepanoclacliis, Calliergon and lichens, 19 Aug. 1972, OKMI GAL (ZT 4497). Barrow: IBP Tundra Biome Sites: near NE Dewline, among Drepanocl~ldus, Calliergon and liverworts, 4 Aug. 1972, OKMIGAL (ZT 4554). Schultz fertilized plot (2 mi SE of NARL), on Drepanoclad~is revolvens, 6 Aug. 1972, OKMIGAL (ZT 4619). Among Drepanocladus, 17 Aug. 1972, OKMIGAL IBP Tundra Biome: Site 2: Calliergo~z, 19 Aug. 1972, OKMIGAL (ZT 4498); on peaty soil among moss (incl. Polyrrichwn) and grass debris on polygon rim, 27 July 1976, OKM Site 4: among moss, 24 Aug. 1971, OKMIGAL 10609; on Dreparzocladus, 23 Aug. 1972, OKMIGAL (ZT 4486); among Drepanoclaclus and Sphagnum, 23 Aug. 1972, OKMIGAL Site 4: plot 418: among moss, 8 Aug. 1972, OKMIGAL (ZT 4484); among Drepanocladus and Calliergon, 8 Aug. 1972, OKMIGAL ECOLOGY: Occurs solitary to occasionally in caespitose clusters. In the course of the present study carried out in the Barrow region, 11 records of P. sragizina have been examined. This comparatively common species occurs almost exclusively only in the most humid sites of the local wet tundra. Accordingly, it is no surprise that eight records have been found in hygrophilous moss associations dominated by either Drepanoclaclus or Calliergon, the latter being responsible for about 15% of the bryomass in the investigated habitats (Rastorfer et al. 1974). The basidiomes of the remaining three collections have been gathered on naked peat (with or without Polytrichutn) or on decaying plant debris. It should be noted that in arctic-alpine localities in Norway (Gulden 1980) and Switzerland (Senn-Irlet 1988b), P. sragtzina also has been recorded in peat bogs with Drepanoclaclus or Calliergorz, also noted as the most frequent mosses. In the Barrow area, there was only one sample of P. sragnina where remnants of Sphagnum have been part of the substrate. It appears that Sphagnum, which occurs scattered only in the extremely moist, low center polygons around Barrow, is rarely chosen as host plant, an observation already made by several authors working in arctic-alpine localities in the northern hemisphere (Favre 1955; Gulden 1980; Redhead 1981; Gulden et al. 1985; Nezdojminogo 1985) and in subantarctic peat bogs of Tierra del Fuego (Horak 1979). So far all Alaskan records of P. stagnina have been recorded from the vicinity of Barrow, except several collections gathered by Wells and Kempton (1969) in the boreal zone further to the South. REMARKS: In the field, P. stagnina is readily recognized by its rather robust, dark brown basidiomes with contrasting white persistent veil remnants both on stipe and margin of pileus (Fig. 38). As a rule, the microscopical characteristics of Alaskan specimens correspond well with those observed on material of European origin. The identification of the species is verified by the smooth (Gulden 1987), elliptical, thickwalled spores measuring pm. Spores borne on rare bisporous basidia can reach up to 20 pm in length. Another distinctive feature of P. stagrzina is the remarkable polymorphic cheilocystidia. The majority of these abundant cheilocystidia (and caulocystidia) are intermixed, however, with broadly fusoid or even vesiculose cells. Occasionally, cheilocystida with forked tips or multiple constrictions can also be observed. ARCTIC-ALPINE DISTRIBUTION: Switzerland (Favre 1955; Senn-Irlet 1987, 1988b; herb. Horak, ZT 4156, unpublished data); France (Kiihner 1972b); Scotland (Dennis 1955); Norway (Blytt 1905; Gulden 1975, 1980; Gulden et al. 1985); USSR: northern Siberia (Nezdojminogo 1985); Jan Mayen (Larsen 1923); SvAlbard (Gulden et al. 1985; Gulden 1987); U.S.A.: Alaska (Wells and Kempton 1969; Miller et al. 1973, 1974; Laursen and Chmielewsky 1982). SUBANTARCTIC DISTRIBUTION: Argentina (Tierra del Fuego; Horak 1979); Antarctica: Palmer Peninsula (Horak 1982), New Amsterdam Is. (Hennings 1906), South Georgia (Pegler et al. 1980). 2. Galerina arctica (Singer) Nezdojminogo Figs. 5-9, 43 Mikol. Fitopatol. 16: Corrinarius arcticus Singer, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. Nauk SSSR, 4: ILLUSTRATION: Gulden and Jenssen (1988). DESCRIPTION: (OKMIGAL 11066): Pileus up to 20 mrn diameter, at first hemispheric becoming convex to broadly umbonate, expanded in mature specimens; color ochre (ochraceous buff) to gold yellow (antimon yellow), disc pale cinnamon brown (tawny), fading to pale yellow-ochre (warm buff); surface dry, glabrous, conspicuously translucent-striate, strongly hygrophanous, in young specimens margin covered with whitish veil fibrils. Lamellae adnexed to adnate, ventricose (up to 4 mm deep), rather distant; at first pale yellow-ochre changing to deep ochre (yellow ochre), edges concolorous, even. Stipe up to 45 x up to 2 mm, cylindric, equal, concolorous with pileus surface; apex distinctly pruinose, otherwise covered with pale veil fibrils, distinct cortina absent; dry, stuffed. Context of pileus and stipe pale yellow-brown, up to 1 mm thick in pileus. Odor and taste not distinctive. Spores x pm, amygdaliform to sublimoniform or pear-shaped, occasionally reniform; very pale yellowbrown to almost hyaline, thin-walled (and therefore often collapsed), smooth to faintly rough, germ pore, plage and perispore absent. Basidia x 8-10 pm, 4-spored. Cheilocystidia x 2-6 pm, apex up to 5 pm diameter, lecythiform, hyaline, rarely with yellow-brown plasmatic pigment. Pleurocystidia absent. Caulocystidia x 5-20 pm, apex up to 10 pm, lecythiform, occasionally not capitate, hyaline. Pileipellis a cutis of repent cylindric hyphae (2-4 pm diameter), slightly gelatinized, encrusted with yellow-brown pigment, occasionally also with yellow-brown plasmatic pigment, subcutis composed of broadly cylindric to fusoid cells, yellow-brown oleiferous hyphae present. Clamp connections absent on all septa, including basal septum of basidia. MATERIAL EXAMINED: CANADA: YUKON TERRITORY: Skolai Pass: among Calliergon sarmentosum 21 July 1967, OKM 5696 (ZT 4494). U.S.A.: ALASKA: Barrow: IBP Tundra Biome Sites: among Canlpylium and mniaceous moss, 17 July 1973, T. Rau, OKM (ZT 4593). I.B.P. Tundra Biome: Site I: among moss and grass remnants, 25 July 1973, OKMIGAL 11481; among Calliergon and grass debris on peaty soil, 15 Aug. 1973, T. Rau and OKMIGAL (ZT 4594). Site 2: plot 2: among aquatic moss (incl. Drepanocladus), 10 Aug. 1971, OKMI

5 418 CAN. J. BOT. VOL. 70, 1992 FIGS Phaeogalern stagnina (VPI 10932). Fig. 2. Spores. Fig. 3. Basidia. Fig. 4. Cheilocystidia. FIGS Galeritza arctica (VPI 11066). Fig. 5. Spores. Fig. 6. Basidia. Fig. 7. Cheilocystidia. Fig. 8. Caulocystidia. Fig. 9. Pileipellis. Scale bar = 10 pm for spores, 20 pm for basidia and cystidia, and 40 pm for pileipellis. GAL 10418; plot 10: on moss, 10 Aug. 1971, OKMIGAL 10 Sept. 1969, Kuhner (holotype of G. griseipes (ZT 4483). Site 4: among Drepanocladus, 17 Aug. 1972, Kuhner); Vanoise, Pralognon, Lac des Vaches, 2300 m, 6 Sept. OKMIGAL (ZT 4485); among moss, 23 Aug. 1972, 1973, det. Kuhner (herb. Lamoure). OKMIGAL FRANCE: Vanoise, Lac des Evettes, 2500 m, ECOLOGY: Occurs solitary, never caespitose, but it may be

6 HORAK AND MILLER 419 gregarious. The single Yukon and the seven Alaskan collections of G. arctica have been observed in close association with the typical wet tundra mosses Campyliutn, Calliergotz, and Drepatzoclaclus, i.e., under similar ecological conditions as reported by Gulden (1987) from localities in Svilbard. REMARKS: The examined material represents the first records of G. arctica in Alaska and Canada. Previously the area of distribution of this common high arctic Galeritza has been restricted to the northern Soviet Union (including its offshore Siberian Islands; cf. Nezdojminogo 1982 and Svilbard (Gulden 1987; Jalink and Nauto 1988). Despite several discrepancies (which will be discussed elsewhere) with the original description (Singer 1938), the taxonomy of this species closely follows the interpretation proposed by Gulden (1987). 'The brilliant yellow to ochre color of the basidiomes makes G. arctica a macroscopically distinctive species that can be mistaken for the similar, and in arctic regions sympatric, G. clavata (see below). The identity of the former species is readily confirmed by its pale and distinctly amygdaliform spores whose thin walls (often collapsed in mounts) are either roughened or covered only by minute scattered warts. Furthermore, G. arctica is characterized by the lack of clamp connections, large lecythiform cheilocystidia, and caulocystidia that occasionally can be filled or encrusted with yellow-brown pigment in KOH. The present Alaskan material has been compared with specimens collected by Gulden (herb 0, ) and found to be identical in all relevant details. Gulden and Jenssen (1988) considered G. arctica to be conspecific with G. griseipes Kuhner (19726), originally described from the Savoyan Alps in France. The reexamination of both the type material and an additional collection from the same general region (herb. Lamoure, 6 Sept. 1973) revealed that the French Galerincl is actually a closely related but clearly separate species. The two taxa (G. griseipes and G. arctica) are distinguished by the size and the ornamentation of the spores. For that reason, the occurrence of G. arctica in the Alps has to be deleted from the records. ARCTIC-SUBARCTIC DISTRIBUTION: Russia: Novaja Semlya (Singer 1938), Franz Josef Land, Severnaja Zemlya, Dickson Island, New Siberian Islands, E-coast of Taimyr Peninsula (Nezdojminogo 1982); Svllbard (Gulden 1987; Gulden and Jenssen 1988; Jalink and Nauta 1989). 3. Galerina clavata (Velenovsky) Kuhner Figs , 39 Encycl. Mycol. 7: 171, 1935 Galerina fragilis var. clavata Velenovsky, Ceske Houby, 548, 1921 MISAPPLIED NAME: Galerina heterocystis (Atk.) Smith & Singer, ILLUSTRATION: Gulden and Jenssen (1988). DESCRIPTION: (OKMIGAL 10733): Pileus up to 16 mm diameter, convex with faint umbo becoming broadly convex with shallow depression at center; color orange (orange rufous), brown (hazel) at disc and over striations to pale yellow-brown over the margin; moist, translucently striate, hygrophanous, whitish veil remnants at margin fugaceous. Lamellae adnate to emarginate, subdistant, pale ochre (apricot buff), edges subfimbriate and concolorous. Stipe up to 43 X up to 2 mm, cylindric, equal or slightly enlarged at base; pale yellow-brown (like margin of pileus), densely covered with minute, white, appressed fibrils of veil, disappearing with maturity. Context concolorous with the pileus, thin in pileus and stipe. Odor and taste not distinctive. Spores x 6-7 pm, elliptic to subphaseoliform, supra- hilar depression always distinctive, yellow-brown, marbled or minutely warted to coarsely warted, germ pore and perispore absent. Basidia x 8-12 pm, 4-spored, clamp connection absent. Cheilocystidia x 5-15 pm, fusoid-capitate to lecythiform, apex up to 11 pm diameter, thin-walled, hyaline. Pleurocystidia none. Caulocystidia up to 100 x up to 13 pm, slender fusoid-capitate, apex up to 13 pm diameter, hyaline, thin-walled. Pileipellis a cutis of short cylindric hyphae (2-6 pm diameter), encrusted with yellow-brown pigment, oleiferous hyphae rare, differentiated terminal cells (pileocystidia) on epicuticular hyphae absent. Clamp connections absent on all septa. MATERIAL EXAMINED: U.S.A.: ALASKA: Pt. Hope: among moss along stream, 17 Aug. 1974, OKMIGAL Barrow: IBP Tundra Biome Sites: among Campylium and? Tomenthypnum, 29 July 1972, OKMIGAL (ZT 4620); among Mnium Hedwig and plant debris, 17 Aug. 1980, G. Gulden (herb. 0, ); among moss, 19 Aug. 1980, G. Gulden (herb. 0, ). Willey Post, 16 mi SW of Barrow: among Campylium cf. arctici~tn and Drepannocladus, 5 Aug. 1972, OKMIGALIN. Wieland 10900; among D. revolvens (C. M.) Warnst., Calliergon and Bryum, 5 Aug. 1972, OKMIGALI N. Wieland Schultz fertilized plots, 2 mi SE of Barrow: among D. revolvetzs and Bryum, 6 Aug. 1972, OKMIGAL (ZT 4618); among Messia triquetra (Hook. & Tayl.) Alongstr., 6 Aug. 1972, OKMIGAL 10916B. Gas Pipe Ridge: among Calliergon and Carnpylium cf. arcticum, 28 Aug. 1974, OKMIGAL (ZT 4592); OKM (ZT 455 1). IBP Tundra Biome: Site 1: among Campyliutn sellatum var. arcticum (Williams) Sav.-Ljub. and grass debris on peaty soil, 10 Aug. 1972, OKMIGAL Site 2: plot 1 : among moss, 10 Aug. 1971, OKMIGAL (ZT 4488). Site 4: among Tomenthypnum nitens (Hedw.) Loeske in drained pond bottom, 6 Aug. 1971, OKM (cf. G. tnniophila in Miller et al. 1973) (ZT 4500); among Drepanocladus, 22 July 1972, OKMIGAL (cf. G. heterocystis in Miller et al. 1973) (ZT 4491); among moss on peaty soil, 31 July 1972, OKMI GAL Site 4: plot 421: among moss, 1 Aug. 1972, OKMIGAL (cf. G. subatztzulata in Miller et al. 1973). Site 7: among Calliergon and Camnpylium arcticum, 13 Aug. 1973, OKMIGALIT. Rau Site 12: among Calliergotz and Cinclidiuttz, 16 Aug. 1973, OKMIGALIT. Rau ECOLOGY: The long list of published records proves that in arctic-alpine habitats G. clavata is a widely distributed species. The 18 collections from arctic Alaska provide evidence that this taxon is one of the most frequently occurring Galerina species. Regarding its ecology, the majority of records have been registered in the wet tundra near Barrow among mosses belonging to Campylium, Calliergon, and Drepannocladus. In the investigated sites several representatives of these three moss genera are most commonly encountered both in interpolygonal troughs and in polygon centers (Rastorfer 19746). Their combined bryomass accounts for about 70% whereas decaying Cattlpylium alone contributes roughly half of the bryogenous substrate. To a much lesser degree members of Bryum, Cinclidiutn, and Mrzium can be considered as further host plants, and finally Tomenthyphum and Meesia Hewdig are the least recorded mosses that serve as substrates for G. clavata. Our data recording substrate relationships and host preferences compares well with those reported from several alpine localities in the Swiss Alps (Kiihner 19726; Senn-Irlet 1987) and in arctic environs in northern Europe (Bresinsky 19666; Gulden 1980). Moss associates recorded in these regions

7 420 CAN. J. BOT. VOL. 70, 1992 FIGS Galerina clavnta (VPI 10733). Fig. 10. Spores. Fig. 11. Basidia. Fig. 12. Cheilocystidia. FIGS Galerina leptocystis (VPI 10587). Fig. 13. Spores. Fig. 14. Basidia. Fig. 15. Cheilocystidia. Fig. 16. Caulocystidia. FIGS Galerinn subnrcticn (VPI 10339). Fig. 17. Spores. Fig. 18. Basidia. Fig. 19. Cheilocystidia. Scale bar = 10 Frn for spores, 20 Krn for basidia and cystidia. include species in the following genera: Acrocladium, Crato- subarctic bogs where it was seen on several mosses including neuron, Hyloconziurn, Paludella, and Philonotis. Sphagnum. Based upon the available literature, however, The first Alaskan record of G. clavata was published by G. clavuta is rarely associated with Sphagnurn in the arctic- Wells and Kempton (1969) who described this agaric from alpine environment viz. in Norway (Gulden and Lange 1971;

8 HORAK AND MILLER 42 1 Gulden 1980) and in Greenland (Lange 1957; Kobayasi et al. 1971). REMARKS: The 18 collections of G. clavata examined demonstrate that in arctic Alaska this species is an inherent member of moss communities in the moist humid sites observed in the local wet tundra (Fig. 1) where it regularly occurs together with P. stagnina and G. arctica. Galerirza arctica may be readily mistaken for G. clavata (Fig. 39) whose macroscopical characters are similar regarding size, shape, color, and fugaceous veil remnants. The two taxa, however, are readily separated by their microscopic features, since the spores of the latter species are phaseoliform and rather coarsely warted. As a rule, the basidia are 4-spored and occasionally clampless. One also encounters 2-spored basidia that produce large spores measuring between pm in length. See G. arctica above for comparison. ARCTIC-ALPINE DISTRIBUTION: Italy (Bon 1987); France (Remy 1964; Kiihner 1972b; Lamoure 1982; Kiihner and Lamoure 1986); Switzerland (Favre 1955; Senn-Irlet 1987, 1988b); Sweden ('?Bresinsky 19660; Kiihner 1972b; Jacobsson 1984); Norway (Gulden 1975, 1980; Gulden and Lange 1971;.. Kiihner 19720); USSR (Tomilin 1971; Vasilkov 1972; Stepanova and Tomilin 1972, 1973; Nezdojminogo 1982, 1984, 1985); Hebrides (Watling and Richardson 1971 ; Dennis : 1986); Faeroes (Moeller 1945); Iceland (Hallgrimsson and I Kristinsson 1965); Greenland (Lange 1957; Kobayasi et al. / 1971; Watling 1977; Lamoure et al. 1982); SvPlvard (Gulden ; Gulden and Jenssen 1988; Fellner and Landa 1989); i I U.S.A.: Alaska (Laursen et al. 1972, 1977; Miller et al. 1973, (cf. G. stagilina, OKM 10864; G. tnniophila, OKM I I I 10359); Bunnell et al. 1980; Laursen and Ammirati 1982; Laursen and Chmielewsky 1982). 4. Galerina leptocystis Wells and Kempton Figs Lloydia, 32: ILLUSTRATION: Wells and Kempton (1969). DESCRIPTION: (OKM/GAL and OKM/GAL 10587): Pileus up to 8 mm diameter, conic (also in mature specimens); color brown to dark brown; surface hygrophanous, translucently striate, dry, smooth, margin without veil remnants. Lamellae adnexed, sinuate; pale brown to yellow-brown. Stipe up to 27 x nim, cylindric, equal; dark brown, covered with minute, scattered whit~sh fibrils forming an evanescent superior annulus. Context of pileus and stipe brown. Odor and taste not distinctive. Spores x 5-6 pni, pip-shaped to slender amygdaliform, pale yellow, roughened to minutely punctate, plage indistinct, perispore absent or thin, clearly seen only as bulging fringe bordering plage, germ pore absent. Basidia x 8-10 pm, 4-spored. Cheilocy stidia x 5-9 pm, slender fusoid with elongated neck, rarely subcapitate, thinwalled, hyaline. Pleurocystidia absent. Caulocystidia up to 70 x up to 8 pm, polymorphic, fusoid, subcapitate or tapering towards apex. Pileipellis a cutis of short cylindric hyphae (up to 10 pm diameter), encrusted with pale yellow pigment, epicutis composed of hyaline cylindric nongelatinized hyphae, differentiated terminal cells absent. Clamp connections present on all septa. MATERIAL EXAMINED: U.S.A.: ALASKA: Barrow: IBP Tundra Biome Sites: among moss (incl. Polytrichurn commune) on peaty soil, 23 Aug. 1971, OKMIGAL (ZT 4581); aniong Polytrichurn, lichens and grass debris on peaty soil, 4 Aug. 1973, OKM (ZT 4595). Anchorage, Kenai Peninsula, on Sphagrzum, 9 July 1966, Wells-Kempton 2966 (HOLOTYPE of G. leptocystis, MICH). ECOLOGY: The two Barrow collections have not been found on Sphagnum (cf. type material from southern Alaska) but on peaty soil among Polytrichurn, lichens, and plant debris, which indicates rather dry site conditions. REMARKS: This is the first record for arctic North America. Taste and odor are not fully documented for both collections and they lack, therefore, important macroscopical data for identification. The most distinctive microscopic features are the pale yellow, amygdaliform spores that are devoid of a germ pore. The plage on the flattened supraapicular zone is surrounded by inconspicuous perisporial remnants that are best recognized by the bulged margin of the plage and by the scattered small blisters near the apiculus. The wall is either punctate-roughened or merely covered with minute warts. Except for the ecological conditions of the Barrow habitat, the two arctic Alaskan records key out with G. leptocystis that was originally described by Wells and Kempton (1969) from a Sphagnurn bog at Kenai Peninsula, south of Anchorage. In every way the microscopic characters fit those observed on the type specimen of G. leptocystis observed in material of OKMIGAL and OKM (see also the discussion under G. subarctica (5)). ARCTIC DISTRIBUTION: U.S.A.: Alaska (for records see above). 5. Galerirza subarctica Smith & Singer Figs , 40 Monogr. Galerirza, DESCRIPTION: (OKM 11207): Pileus up to 18 mm diameter, at first convex with pointed umbo, becoming convex to expanded with age; dark orange brown (argus brown) to argillaceous brown (antique brown), paler towards margin; moist to subviscid, appressed fibrillose, translucently striate, hygrophanous. ~amellae adnexed to emarginate-submarginate, subdistant; pale ochre (honey yellow) to pale ferruginous brown (cinnamon brown); edges uneven, concolorous. Stipe up to 30 x up to 2.5 mm, cylindric, equal; concolorous with pileus surface or paler (chamois), at base sometimes pale greenbrown (light yellowish olive), whitish fibrillose veil remnants forming an indistinct evanescent cortina; dry, becoming hollow with age. Context concolorous with pileus, up to 2 mm thick at disc. Odor and taste not distinctive. Spores x pm, amygdaliform, yellow-brown or rust brown, faintly roughened or minutely punctate, plage and gerni pore distinctive, perispore poorly developed but often forming a bordered plage area. Basidia x 7-8 pm, 4-spored. Cheilocystidia x 6-14 pm, lageniform to fusoid with subcapitate neck, hyaline, thin-walled. Pleurocystidia absent. Caulocystidia x 5-15 pm, polymorphic, mostly subclavate or like cheilocystidia. Pileipellis a cutis of cylindric hyphae (up to 15 pm diameter), nongelatinized and encrusted with yellow-brown pigment, terminal cells of velar hyphae not differentiated. Clamp connections present on all septa. ECOLOGY: At Barrow 6 of 10 collections of G, subarctica were found in association with Sphagnum. In several samples Dicrarzum, Oncophorus, and Polytrichum have also been observed. In addition, among remnants of lichens and the liverworts, Anastrophyllurn and Blepharostorna have been part of the substrate colonized by this Galerina. These ecological data indicate that in the wet tundra (at least around Barrow) G. subarctica is one of the most frequent species in a close relationship with Sphagnum.

9 422 CAN. J. BOT. VOL REMARKS: This is the first record in arctic North America. In 1980, G. Gulden collected material at Barrow that was published (Laursen and Ammirati 1982) as G. hypophaea Kuhner and G. pumila var. subalpina Smith. In the course of our studies, these three collections have been reexamined and found to belong to one taxon. It also turned out that eight more collections of the same species are among the Galerina material gathered in the Barrow region. Without exception, the spore walls in all the above-mentioned samples are definitely rough to minutely warted. In the original diagnoses, however, the spores of both G. lzypophaea and G. purnila var. subalpina are described as smooth. Consequently, Gulden's specimens (Gulden and are considered to be misidentified and these two Galeritla collections have to be taken from the list of arctic Alaskan records. In this connection, it must be pointed out that the spores in the Norwegian material (mentioned by Gulden 1980) are actually described as "punctate to marbled." Under these circumstances, it remains open to question whether or not the Alaskan fungi are conspecific with the Scandinavian ones. In summary, the 11 specimens cited above from Barrow are tentatively considered to represent G. subarctica. The microcharacters of this species, originally described from the area around Anchorage, fit those observed on the basidiomes from northern Alaska in all respects. The only notable differences of the two extant collections (Wells and Kempton 1969), however, are the larger size of the type specimen and the habitat that is said to be in spruce-hemlock forests in the vicinity of Anchorage. MATERIAL EXAMINED: U.S.A.: ALASKA: Icy Cape: among Oncophorus wahlenbergii, Blepharostoma trichoplzyll~lm, lichens, and plant debris on peaty soil, 17 Aug. 1974, OKMI GAL Barrow: IBP Tundra Biome Sites: on Sphngnutn sp., 22 Aug. 1980, G. Gulden (herb ). E-shore of Lake Evrulivik, 6.5 mi S. of NARL, on Splzagnutn sp., 3 Aug. 1971, OKM (ZT 4482). Bird Census Transect: on Sphagnum sp., 19 Aug. 1972, OKMIGAL Gas Pipe Ridge: among Sphagnum,? Otzcoplzorus, and? Dicrarzutn, 28 Aug. 1974, OKMIGAL 11967; among Splzagnum, 28 Aug. 1974, OKMIGAL (ZT 4590). IBP Tundra Biome: Site 1 : plot 4: among Dicratzuni (not Sphagnum) in low center polygon, 2 Aug. 1971, OKMIGAL (ZT 4584). Site 4: among Sphagnurn, 23 Aug. 1972, OKMIGAL (ZT 4487). Site 4: plot 422: among Dicranunz cf. elongatipes and Anastrophyllum minnrum), 19 Aug. 1974, OKMIGAL Anchorage, 9 Oct. 1959, Wells (MICH, 59-3; HOLOTYPE of G. subarctica). ARCTIC DISTRIBUTION: Known only from Alaska. 6. Galer-ina hypnorurn (Schrank: Fries) Kiihner Figs , 41 Encycl. Mycol. 7: Agaricus hypnorum Schrank, Bayr. F1. 2: Galerina calyptrata Orton, Trans. Br. Mycol. Soc. 43: ILLUSTRATIONS: Kuhner (1935); Gulden (1980, 1987). DESCRIPTION: (OKM 10870): Pileus up to 15 mm diameter, hemispheric with strongly incurved margin, becoming convex with obtuse umbo; color brown with orange tinge to argillaceous brown (antique brown to buckthorn brown) or (isabel color), turning pale ochre-yellow (Naples yellow) upon drying; dry, translucently striate, hygrophanous, smooth, towards margin with appressed whitish evanescent fibrils of veil. Lamellae adnate, distant, up to 4 mm wide; concolorous with pileus, edge uneven, whitish. Stipe up to 40 X up to 2(-5) mm, cylindric, equal or tapering to swollen base; pale ochre (warm buff) to pale orange (apricot buff), at first with scattered whitish fibrils of veil that disappear with age; dry. Context of pileus and stipe concolorous. Odor and taste not distinctive. Spores x 5-6 pm, pip-shaped to slender amygdaliform, pale rust brown, smooth or minutely roughened or faintly punctate, plage and germ pore distinctive, perispore conspicuous and often ear-shaped around plage, often also with blisters near apex. Basidia x 7-9 pm, 4-spored, occasionally 2-spored. Cheilocystidia x 4-11 pm, fusoid to lageniform, apex not or only slightly swollen, hyaline thin-walled. Pleurocystidia absent. Caulocystidia in shape and size as cheilocystidia. Pileipellis a cutis of cylindrical hyphae up to 15 ym diameter, nongelatinized with encrusted pale yellow-brown pigment surface covered by scattered hyaline, cylindrical universal veil hyphae 3-6 pm diameter, without differentiated terminal cells. Clamp connections present on all septa. MATERIAL EXAMINED: U.S.A. : ALASKA: Icy Cape: among Dicranum on peaty soil in polygon trough, 17 Aug. 1974, OKMIGAL (ZT 4591). Barrow: IBP Tundra Biome: Site 5: among Sphagnum, 3 Aug. 1972, OKM (ZT4553). ENGLAND: DEVONSHIRE: Bickleigh, Plym Valley, among Sphagnum, 29 Aug. 1956, Orton 883 (E; HOLOTYPE of G. calyptrata). ECOLOGY: In the tundra surrounding Barrow only two collections of Galerina lzyprzorum have been made. The substrate in one sample was Sphagrzurn whereas peat with Dicranurn were the predominant matrix of the other. Both hosts are repeatedly mentioned as being associated with the species present (Gulden 1987). REMARKS: Solitary but often gregarious. Microscopically, G. hypnorurn is readily distinguished by its smooth spores with a well-defined plage and conspicuous perispore that covers the wall with large blisters or loose lumps (cf. illustrations in Kuhner 1935). The spores of G. Ieptocysris, also reported from arctic Alaska, are similar in shape and size but differ from the former taxon by the absent or occasionally only poorly developed perispore. In arctic-alpine and subantarctic habitats, G. h~pnorunz appears to be the most widely distributed species of the genus. The available data indicate that this taxon is not restricted to a specific host substrate but has been observed on many different mosses, including Sphagnurn and liveworts. Reexamining the type material of G. calyptratn Orton (1960), no taxonoinically reliable differences have been discovered to separate convincingly this taxon from G. hypnorurn s. Kuhner (1935). Accordingly G. calyptrata is considered here as a later synonym (Arnolds 1982; Gulden 1987). In addition, Gulden (1987) reports that Rostrup's (1906) material listed as G. hyprzorurn is actually a taxon belonging to the G. pseudornycenopsis complex. With regards to the putatively worldwide distribution of G. hypnorunz, it can be expected that after critical evaluation of the relevant voucher specimens several further synonyms will be established. ARCTIC-ALPINE DISTRIBUTION: France (Remy 1964: "f. bispora"; Kuhner 1972a); Switzerland (Boudier and Fischer 1895; Favre 1955; Senn-Irlet 19886); Scotland (Dennis 1955); Shetland Is. (Dennis and Gray 1954); Hebrides (Dennis 1986); Sweden (Lange 1946; Bresinsky 1966b; Kuhner 1972a), Norway (Gulden 1980); Finland (Karsten 1882;

10 HORAK AND MILLER FIGS Galerina hyptzor~rm (VPI 10870). Fig. 20. Spores. Fig. 21. Basidia. Fig. 22. Cheilocystidia. FIGS Galeritza sp. 1 (10916 A). Fig. 23. Spores. Fig. 24. Basidia. Fig. 25. Cheilocystidia. FIGS Galerina pselldoceritza (VPI 5722). Fig. 26. Spores. Fig. 27. Basidia. Fig. 28. Cheilocystidia. Fig. 29. Caulocystidia. Scale bar = 10 ym for spores, 20 ym for basidia and cystidia. Kallio and Kankainen 1966); Russia (northern Siberia: Lebedeva Iceland (Rostrup 1903; Larsen 1932; Lange 1949); Greenland 1927; Vasilkov 1970; Mikhailovski 197%; Nezdojminogo (Rostrup 1888, 1891, 1894; Allescher and Hennings 1897; 1984, 1985); Caucasus (Singer 1931); Faeroes (Boergesen and Ferdinandsen 1910); Jan Mayen (Reichardt 1886; Hariot Ostenfeld-Hansen ; Rostrup 1901 ; Moeller 1945); 1893; Rostrup 1897; Larsen 1924); Svilbard (Lindblom 1841 ;

11 424 CAN. J. BOT. V Karsten 1872; Michelmore 1934; Dobbs 1942; Gulden 1987); U.S.A. : Alaska (Miller et al. 1974); Canada (Dearness 1923); Subantarctic distribution: Kerguelen Is. (Berkeley 1877h; Pegler et al. 1980), Tierra del Fuego (Horak 1979, 1982). 7. Galerina sp. 1 Figs , 42 DESCRIP~ION (OKMIGAL 10916A): Pileus up to 18 mm diameter, hemispheric to umbonate, surface translucently striate, hygrophanous, dry, smooth, with minute whitish veil remnants at margin. Lamellae adnexed to broadly adnateemarginate, edges white, fimbriate. Stipe cylindric, dry, apex pruinose, with scattered appressed whitish fibrils from veil, not forming a cortina. Spores x pm, amygdaliform, rust brown, minutely warted to marbled, plage and germ pore distinctive, perispore conspicuous, often ear-shaped around plage, occasionally with blisters towards apex. Basidia x 7-9 pm, 4-spored. Cheilocystidia x 7-15 pm, polymorphic but usually fusoid-capitate to broadly utriform, sometimes fusoid with several constrictions, wall hyaline, often filled with yellow plasmatic pigment. Caulocystidia not observed. Pileipellis a cutis of cylindric hyphae (up to 16 pm diameter), nongelatinized hyphae encrusted with yellow pigment, terminal cells of velar hyphae not differentiated. Clamp connections present on all septa. MATERIAL EXAMINED: U.S.A.: ALASKA: Barrow, Schultz fertilized plots, 2 mi SE of NARL, among Meesia triquetra on peaty soil, 6 Aug. 1972, OKMIGAL 10916A (ZT 4583). ECOLOGY: In the wet tundra at Barrow, this unnamed Galerina has been observed only once. It occurred in a fertilized plot (Brown and West 1970) on peaty soil among M. triquetra. REMARKS: Figure 42 illustrates this unidentified Galerina for which there are insufficient descriptive notes (e.g., taste and odor) available. Nevertheless, this record is remarkable because of its very distinctive microscopic characters. The rather large, minutely warted spores with obvious perisporal blisters are most noteworthy. This morphological feature points to a possible taxonomic relationship with G. hypnorurn. The shape and size of the conspicuously large and polymorphic cheilocystidia actually do not fully support this concept because they are considerably larger and broader than the locally gathered material of G. hypnorum. This Galerina does not appear to be in the keys published both by Smith and Singer (1964) and Wells and Kempton (1969). ARCTIC-ALPINE DISTRIBUTION: Only known from Barrow, arctic Alaska, U.S.A. 8. Galerina pseudocerina Smith & Singer Figs Mycologia, 50: ILLUSTRATIONS: Gulden (1980); Senn-Irlet (1987). DESCRIPTION (OKM 5722): Pileus up to 10 mm diameter, at first conico-convex becoming convex with low umbo; color dark cinnamon brown (Mikado brown) becoming pale orangebrown towards the margin; moist, hygrophanous, translucently striate veil remnants absent. Lamellae adnate to broadly emarginate, distant to subdistant; concolorous with pileus surface or pale ochre (cinnamon buff), edges minutely fimbriate, concolorous. Stipe up to 45 x up to 2.5 mm, cylindric, equal or gradually attenuated upwards, base not bulbous; concolorous with pileus surface, covered with minute fibrils of veil especially towards base, persistent veil remnants absent; dry, solid, very brittle. Context thin, concolorous with pileus surface. Odor and taste not distinctive. Spores x 7-9 pm, broadly ovoid to subamygdaliform, dark rust brown, coarsely marbled or verrucose, plage and perispore distinctive, thick-walled, occasionally with apical pore. Basidia X 9-11 pm, 4-spored. Cheilocystidia x 3-10 pm, lecythiform, apex up to 5 pm diameter, sometimes forked or constricted, thin-walled, sometimes with yellow-brown plasmatic pigment. Pleurocystidia absent. Caulocystidia up to 60 x up to 12 pm (apex up to 7 pm diameter), shape same as cheilocystidia. Pileipellis a cutis of cylindric subgelatinized hyphae (1-4 pm diameter), differentiated terminal cells absent, walls of subcuticular hyphae encrusted with rust brown or yellow-brown pigment, oleiferous hyphae absent. Clamp connections present on all septa. MATERIAL EXAMINED: CANADA: YUKON TERRITORY: Skolai Pass: among moss on soil, 5100 ft, July 1967, D. Scott, OKM 5814 (ZT 4599); among moss, lichens, and grass remnants on peaty soil, 22 July 1967, OKM 5694, OKM 5695, OKM 5697, OKM 5698, OKM 5719, OKM 5720, OKM 5721, OKM 5722 (ZT 4493). U.S.A. : COLORADO: San Juan Mountains, Trout Lake, Smith (MICH; HOLOTYPE of G. pseudocerina). ECOLOGY: Occurs solitary to subcaespitose. The examined material consists of eight collections all gathered on the same day and in the same remote locality. No data about the plant association at the actual site are available. Information in the literature reveals that this species prefers base-rich habitats characterized by plant associations belonging to the Dryadion and Salicetum retuso-reticulatae. REMARKS: At present there is some confusion about the taxonomic concept of G. pseudocerina and accordingly the distributional data mentioned in the literature have to be reconsidered. The presented interpretation of this taxon is strictly based on the following microscopic characters observed on the type material: (i) spores distinctly amygdaliform, often with mucronate or beaked apex, coarsely marbled to verrucose, supraapicular depression with well-defined plage; (ii) cheilocystidia (and caulocystidia) lecythiform, fusoid with elongate neck and conspicuous capitate apex, up to 40 pm long; and (iii) clamp connections present. After careful evaluation of the arctic Alaskan material and all published records, several times also supported by reexamination of voucher specimens, it can be concluded that typical G. pseudocerina so far has been recorded only from arctic-alpine localities in Switzerland (Kuhner 1972~; Senn-Irlet 1987, 1988a), Svilbard (Gulden 1980, 1987 p.p.), Canada, and U.S.A. Gulden (1987, Fig. 4) when discussing her ample collections of G. pseudocerina, considered to be the most common species of Galerina in Siberia (Nezdojminogo 1982), France (Kuhner and Lamour 1986), and Svilbard, recognized populations not only with amagdaliform but also ovoid spores. Arctic collections studied from the Skolai Pass on the Yukon- Alaskan border and Eagle Summit in central Alaska have been observed to have spores that are mostly ovoid but amygdaliform in some samples. Gulden and Jenssen (1988) published a detailed description (including an instructive plate and drawings) of specimens found in Svilbard. The illustrated material (GG 107/86), however, is characterized by having ovoid to almost subglobose spores with coarse warts. A reexamination of the pertinent material confirms Gulden's (1987) observations in all details. Finally in 1990, one of us (E.H.) collected material in the alpine zone of the Dolomites (Italy, ZT 4148) whose micros-

12 HORAK AND MILLER 425 copic and macroscopic features were identical with those given in Gulden and Jenssen (1988). The Italian specimens have been found on limestone among Dt-yrls and calciphilous Salix, i.e., under ecological conditions similar to the records of Svilbard. Based on these facts we came to the conclusion that in the same habitat G. pseudoceritza (s. Smith & Singer) can occur together with another similar-looking, obviously closely related and so far undescribed species that can be separated on the basis of its peculiar spore characters. To support this hypothesis, all published records of "G. pseudocerina" have been critically screened. As a result the ovoid-spored double of G. pseurlocerina definitely also occurs in northern Siberia (Nezdojminogo 1982), Svilbard (Reid 1979; Gulden 1987 p.p.; Gulden and Jenssen 1988), Italy (Horak ZT 4148, unpublished), Canada (Yukon, Skolai Pass, OKM 58 14), and Alaska (Eagle Summit, OKM 10015). Below all those, doubtful records have been listed that either lack information about the shape of the spores or that probably relate to data drawn from mixed collections: France (Kuhner 1965, 1972a; Bon 1985; Kiihner and Lamoure 1986), Scotland (Watling 1983), Norway (Kuhner 1965, 1972a), Sweden (Kuhner 1972a), USSR (Nezdojminogo 1984), Greenland (Watling 1983), SvPlbard (Gulden 1987 p.p.), and Canada (Alex Heiberg Is: Parmelee 1963). ARCTIC-ALPINE DISTRIBUTION: Switzerland (Kuhner 1972a; Senn-Irlet 1987, 1988a); Norway (Gulden 1980); Svilbard (Gulden 1987 p.p.); U.S.A. : Colorado, Rocky Mts. (Smith and Singer 1958). TIze Galerina pseudomycenopsis (= G. moelleri) complex In this paper, we do not discuss our view regarding the taxonomical and nomenclatural problems concerning G. pseudomycenopsis. The pertinent literature has many divergent concepts about this taxon (and its numerous synonyms) and at the present moment we consider it still too premature to make proposals on how to delimit this polymorphic species against a swarm of closely related taxa. To resolve many open questions, much more research has to be carried out not only in the field but also in the laboratory. Nevertheless, there is the possibility of splitting G. pseudomycerzosis s.1. into two microscopically well-defined subspecific groups (form 1 and form 2). At the same time, the recommendation to keep these two forms apart is regarded as an invitation to fellow mycologists to check for the microscopic characters in D. pse~ldornycenopsis material. Unfortunately, many published records of arctic-alpine and antarctic G. pseudomycenopsis lack reliable data about the actual shape of the spores. For that reason, the specimens cited in the following publications cannot be related to either of the proposed forms but nevertheless broaden the list of discoveries for this species as conceived in its wide sense: France (Kuhner and Lamoure 1986), Norway (Gulden and Lange 1971; Gulden 1975), Sweden (Bresinsky 1966b), Russia: northern Siberia (Nezdojminogo 1984), Greenland (Lange 1957; Bas 1960; Petersen 1977; Watling 1977; Lamoure et al. 1982), SvPlbard (Fellner and Landa 1989; Jalink and Nauta 1989), U.S.A.: Alaska (Laursen and Ammirati 1982: cf. G. molleri; Laursen and Chmielewsky 1982), Canada: Ellesmere Is. (Rostrup 1906, as G. hyptzorum, fide Gulden 1987), Antarctica: S Georgia (Dennis 1986), S Orkney Is. (Pegler et al. 1980). As indicated in the key, form 1 and form 2 of G. pseudomycenopsis cannot be separated by macroscopic features. Regarding the examined arctic Alaskan material, however, these two forms can be readily distinguished by the shape of the spores. Several authors (Kuhner 1972a; Kuhner and Lamoure 1965; Gulden 1987) examined numerous populations of "G. pseudotnycetzopsis" and finally came to the conclusion that the spore shape can range from ovoid (or pip-shaped) to amygdaliform. Concerning our material from Alaska, the data obtained are in full accordance with the findings by the above workers. It must be emphasized, however, that within a given collection (i.e., one basidiome), only one spore type is present. Based upon our observations, no apparent intergrading between the extreme shapes of the spores has been noticed. Accordingly, one can speculate that a particular spore morphology is correlated with certain ecotypes of G. pseudotnycet~opsis. At Barrow, the ovoid to pip-shaped spores of form 1 have been detected in about one-third (11 = 38) of the 106 randomly collected samples of G. pseudomycet~opsis. The remaining 68 collections are distinguished by amygdaliform spores. Relying upon personal observations extracted from type material, the specimens allotted to form 2 represent typical material of G. pseudornycenopsis, which has been repeatedly described under different names viz. Pholiota pumila ss. Moeller (1945), Galer-ina pumila var. oreitla Favre (1955), G. pseudopumila Orton (1960), G. moelleri Bas (1960), and probably several more. The biologically interesting observation that at Barrow separate populations of G. pseudorrzycenopsis forms 1 and 2 do actually coexist side by side in the same habitat is not unique, as the following example demonstrates. In 1955, Favre described G. pumila var. oreirzn (=syn. G. pseuclomycenopsis) from Val Sesvenna in the Swiss National Park. His drawings in the original description obviously relate to a taxon having amygdaliform to sublimoniform spores of form 2. Subsequently, Bas (1960, Fig. 16 right) and Horak (unpublished) reexamined the type material and confirmed the observations of Favre (1955). In 1980, however, the senior author (E.H.) collected G. pseudomycetlopsis near the type locality of var. oreinn in the Swiss National Park. The spores of these specimens have no mucronate apex and hence belong to form 1 circumscribed above. These findings prove once more that in nature sympatric strains representing different ecotypes (?) can actually occur simultaneously at the same site. 9a. Galerina pseudotnycenopsis Pilat & Nannfeldt Figs , 44 Friesia, 5: (Form 1: spores pip-shaped, mucro absent) ILLUSTRATIONS: Bas (1960); Kuhner and Lamoure (1965); Horak (1979, as "G. riparia"); Reid (1979); Nezdojminogo (1982); Gulden et al. (1985 p.p.). DEscRrPTIoN (OKM 10549): Pileus up to 40 mm, hemispheric or obtusely conic becoming convex or umbonate; color yellow-brown turning deep brown in older specimens, occasionally with distinctive olive tinge, slightly translucently striate when moist, hygrophanous; subviscid to greasy, waxy, occasionally with pale fibrillose veil remnants along margin. Lamellae broadly adnate to subdecurrent, more or less triangular in shape in young specimens; pale yellow-brown, edges even, concolorous. Stipe up to 40 x up to 4 mm, cylindric, equal, concolorous with or paler than the pileus; membranous ring persistent, white to pale brown, below annulus with white to pale brown fibrils and patches from veil; dry, solid becoming hollow with age. Context up to 3 mm thick in pileus. Odor

13 426 CAN. J. BOT. VOL FIGS Galerirza pseuclomyce~lopsis Form 1 (VPI A). Fig. 30. Spores. Fig. 31. Basidia. Fig. 32. Cheilocystidia. FIG. 33. Gnlerinn pser~clotnycerlo~~sis form 2 (VPI 11051). Spores. FIGS Gnlerirza prniicoln (VPI 10729). Fig. 34. Spores. Fig. 35. Basidia. Fig. 36. Cheilocystidia. Fig. 37. Pileipellis. Scale bar = 10 pnl for spores, 20 pm for basidia and cystidia, and 40 pnm for pileipellis. and taste not distinctive or slightly farinaceous. Spores 10-12(- 13) x 6-8 pm, broadly ovoid but without mucro (occasionally also subamygdaliform), rust brown, coarsely marbled or verrucose, plage and perispore distinc- tive, thick-walled with an apical germ pore. Basidia x 8-10 pm, 4-spored, occasionally 1- to 2-spored. Cheilocystidia 30-70(- 110) x 7-16 pin, fusoid with elongated neck, apex rounded, rarely slightly swollen or conic, hyaline

14 ' HORAK AND MILLER or with yellow-brown plasmatic pigment; often intermixed with clavate to vesiculose cells. Pleurocystidia and caulocystidia scattered, size and shape as cheilocystidia. Pileipellis a cutis of cylindric, subgelatinised, hyaline hyphae (1-4 pm diameter), differentiated terminal cells absent, walls of subcuticular hyphae with yellow-brown to rust brown incrusted pigment, oleiferous hyphae absent. Clamp connections present on all septa. MATERIAL EXAMINED: CANADA: Northwestern Territory: Devon Island, 75 N, 85 W, 30 July 1973, J. Ryan, OKM 11700A. U.S.A. : ALASKA: Prudhoe Bay: among Tomerzfhypnuin niterzs and Bryum, 20 July 1973, M. Williams, OKM Barrow: Imikpak Lake, E shore, 314 mi E of NARL: on peat, 3 Aug. 1971, P. Penhole, B. Breedlove & W. Campbell, OKM IBP Tundra Biome Sites: on peat, 22 Aug. 1980, EH, ZT 028; on peat in swamp, 22 Aug. 1980, EH, ZT 029; among moss on peat, 23 Aug. 1971, OKM (ZT 4555); among moss and Sphagnum on peat, 23 Aug. 1971, OKM (ZT 4582); among moss on peat, 23 Aug. 1972, OKMIGAL 11211; among moss on peat, 4 Aug. 1973, OKMIGAL & T. Rau 11570; among Calliergon and?pseurlobryum, 27 Aug. 1974, OKMI GAL (ZT 4563); among moss on exposed peat scarp, 27 Aug. 1974, OKMIGAL (ZT 4564). Pintail Polygon Plot: among moss and grass debris on peat, 30 July 1971, OKMI GAL (ZT 4569); on mossy-grassy peat, 3 1 July 1971, OKMIGAL (ZT 4566). Schultz fertilized plot: among moss on peat, OKMIGAL 10817, OKMIGAL IBP tundra Biome: Site 1: on peat among moss and grass debris in lower center polygon, 3 1 July 1971, OKM (ZT 4489); among moss on peaty sandy soil, 9 Aug. 1971, J. Reed, OKM 10407; among moss on peat, 20 July 1972, OKMIGAL 10703; among grass debris on peaty soil, 21 July 1972, OKMI GAL 10724; among Polyfriclzum, Dicranurn, and lichens on peat, 28 June 1973, S. Ashkenasi, OKM Site 1: plot 1 I: among moss and grass debris, 12 Aug. 1971, OKMIGAL Site 2: among Sphagnum, 22 Aug. 1972, OKMIGAL Site 2: plot 10: on humus, 10 Aug. 1971, OKMIGAL 10431; plot 30: among moss, 10 Aug. 1971, OKMIGAL 10450; plot 36: among moss on peaty soil, 10 Aug. 1971, OKMIGAL Site 4: among moss on peat, 6 Aug. 1971, OKMIGAL 10362; among Ca1- liergon and Drepanoclarius on peat, 24 Aug. 1971, OKMI GAL (ZT 4577), OKM 10604; among moss, 19 July 1972, OKMIGAL 10686; among moss on peat, 24 July 1972, OKMIGAL 10761; among moss and grass debris on peat, 13 July 1973, T. RauIOKM 11422, OKM 11423; among moss and grass debris on peat, 17 July 1973, OKMIGAL 11444; among moss and grass debris on peat, 1 Sept. 1973, T. Raul OKM Site 4: plot 10: among moss on peat, 9 Aug. 1971, OKMIGAL 10401; plot 416: on peat among moss, lichens, and Salix rotundifolia, 28 July 1972, OKMIGAL 10806; plot 417: on peat, 8 Aug. 1972, OKMIGAL 10945; plot 421: among moss on peat, 8 Aug. 1972, OKMIGAL Site 5: among?pohlia on peat, 3 Aug. 1972, OKMI GAL Site 7: among Calliergon and Drepanocladus in lower center polygon, 5 Aug. 1971, OKMIGAL ZT 4600); on peat, 13 Aug. 1973, OKMIGAL Site 12: among moss on peat, 23 July 1972, OKMIGAL In Miller et al. (1973) the above OKM: 10279, 10345, 10450, 10458, 10549, 10563, 10603, are recorded as "G. subannulafa". SWITZERLAND: National Park, Val Sesvenna, in swamp among moss and Salix herbacea, 2360 m, 5 Sept. 1980, Horak (ZT 192). ECOLOGY: Occur in clusters or solitary. Twenty-eight of 38 collections of form 1 have been recorded on peat as the principal substrate; for eight samples the association with mosses is as follows: Calliergon, Dreparzoc/adus, and Sphagrzum occurred most often in these habitats that have to be regarded as the most humid sites of the local wet tundra. In Svilbard (Ohenoja 1971) the two mosses dominant in arctic Alaskan records also are reported to be very common in connection with G. pseudornycerlopsis (form 1). ARCTIC-ALPINE DISTRIBUTION: Faeroes (Moeller 1945 p.p. ; Bas 1960 p.p.); Sweden (Kuhner and Lamoure 1965 p.p.); Norway (Kuhner and Lamoure 1965 p.p.; Gulden 1980 p.p.; Gulden et al p.p.); Finland (Kuhner and Lamoure 1965 p.p.); Sweden (Jacobsson 1984); Greenland (Lange 1957 p.p.); USSR (Nezdojminogo 1982); SvPlbard (Ohenoja 1971; Reid 1979). ANTARCTIC DISTRIBUTION: Antarctica: Palmer Peninsula (Singer and Corte 1962); Tierra del Fuego (Horak 1979, as G. riparia, 1982). 9b. Galerina pseudomycerzopsis Pilat & Nannfeldt Figs. 33, 45 Friesia, 5: (Form 2: spores amygdaliform, apex mucronate, morphology typical for taxon) ILLUSTRATIONS: Moeller (1945 p.p.); Bas (1980 p.p.); Favre (1955); Singer and Corte (1962); Kuhner and Lamoure (1965 p.p.); Pilit and Nannfeldt (1954); Gulden (1980 p.p., 1987 p.p.); Gulden et al. (1985). DESCRIPTION: (OKMIGAL 11926): Pileus up to 40 mm diameter, hemispheric to convex with obtuse umbo becoming broadly campanulate, margin sometimes upturned in aged specimens; color argillaceous to pale ochre-brown (ochraceous buff) becoming cinnamon brown (buckthorn brown), turning paler upon drying, surface hygrophanous, subviscid, waxy, smooth, margin occasionally with whitish veil remnants. Lamellae moderately crowded, broadly adnate to emarginate-subdecurrent, up to 8 mm wide, cinnamon brown with rust brown tinge, edges whitish, serrulate-fimbriate. Stipe up to 60(-80) x up to 5 mm, cylindric, equal or subclavate towards base, often compressed; concolorous with pileus surface, often becoming darker at base; below white to pale brown with several whitish appressed fibrillose zones from universal veil below the white to pale brown persistent membranous annulus; dry, solid becoming hollow. Context pale ochre-brown, thick in pileus (up to 3 mm diameter). Odor and taste nutty to subfarinaceous. Spores 10-12(- 13) x 6-8 pm, distinctly amygdaliform (to slender pip-shaped), rust brown, coarsely marbled to verrucose, plage and perispore distinctive, thick-walled with apical germ pore. Basidia x 6-9 prn, 4-spored, occasionally also 5- to 7-spored. Cheilocystidia x 7-16 pm, polymorphic, slender fusoid, tapering towards rounded or subcapitate apex, thin-walled, hyaline or sometimes with yellow-brown plasmatic pigment, often intermixed with clavate to vesiculose cells. Pleurocystidia and caulocystidia scattered, shape and size as cheilocystidia. Pileipellis a cutis of cylindric, subgelatinized hyaline hyphae (1-4 pm diameter), differentiated terminal cells absent, walls of subcuticular hyphae encrusted with rust brown or yellow-brown pigment, oleiferous hyphae absent. Clamp connections present on all septa. MATERIAL EXAMINED: U.S.A.: ALASKA: Demarcation Point,

15 CAN. J. BOT. VOL FIG. 38. Phaeogalera stagnina (VPI 10932). FIG. 39. Galerina clavata (VPI 10812). FIG. 40. Gnleritz~ subnrcticr~ (VPI 11207). FIG. 41. Galeritla hypnorutn (VPI 10870). FIG. 42. Galerina sp. 1 (VPI A). FIG. 43. Galerina nrcticn (VPI 11066). Scale bar = 10 rnm. Beaufort Lagoon, Nuvagapak Pt.: among moss on peat, 30 July 1976, H. Miller & OKM, OKM 15299; among T. nitens, 1971, B. M. Murray, OKM F Cape Simpson: among Polytrichum strictum, Oncophorus wahlenbergii, Sphagnum, 28 July 1976, H. Miller & OKM, OKM 15307; among Sphagnum, 28 July 1976, H. Miller & OKM, OKM and grass debris on peat, 20 July 1973, T. Rau, OKM 11465; (ZT 4568); among moss and Salix on peat, 28 July 1976, among moss and dead grass on peaty soil, 24 July 1973, H. Miller & OKM, OKM 15327; among Sphagnum sp., E. Schofield, OKM 11479; Cape Simpson near oil seep 28 July 1976, H. Miller & OKM, OKM Mead River, No. 2: among moss, lichens and dead grass on peat, 28 July about 60 mi S of Barrow: among grass debris on soil, 19 Aug.

16 HORAK AND MILLER , OKMIGAL 11089; among moss on soil, 30 Aug. 1974, OKMIGAL 12019; among Sphagrzum on peat, 30 Aug. 1974, OKMIGAL 12031; on peaty soil, 30 Aug. 1974, OKMIGAL Lake Umiat: among dead grass on peaty soil, 16 Aug. 1971, OKMIGAL (ZT 4598). Barrow: IBP Tundra Biome Sites: Among Polytrichum, B. triclzopilyllurn,?oncoplzorus and grass debris, 23 Aug. 1971, OKMIGAL 10550; among lichens and grass on soil, 4 Aug. 1973, T. Rau & OKMIGAL 11567; among moss on peat, 4 Aug. 1973, T. Rau & OKMIGAL 11568; among moss and grass on peat, 4 Aug. 1973, T. Rau & OKMIGAL (ZT 4596); among moss on peat, 4 Aug. 1973, T. Rau & OKMIGAL 11571; among moss and grass debris, 4 Aug. 1973, T. Rau & OKMIGAL 11573; among dead grass on disturbed peat, 27 Aug. 1974, OKMIGAL (ZT 4565). Pintail Polygon Plot: among moss on peaty soil, 31 July 1971, OKMIGAL 10255; among moss (incl. Polytrichurn) on peaty soil in low center polygon, 3 1 July 1971, OKMIGAL 10293; among moss on peaty soil in polygon trough, 31 July 1971, OKMIGAL (ZT 4570). Bird Census Transect: among Sphagnum on peat, 19 Aug. 1972, OKMIGAL (ZT 4607); among moss on peat, 19 Aug. 1972, OKMIGAL (ZT 4606). Schultz fertilized plot: among grass debris on peat, 30 July 1972, OKMI GAL 10820; among grass debris on peat, 30 July 1972, OKMIGAL IBP Tundra Biome: Sites 1-4: among moss and dead grass on peat, 22 Aug. 1971, OKMIGAL (ZT 4556); among Canzpyliutn cf. arctic~ltn on sandy soil, 22 Aug. 1971, OKMIGAL (ZT 4557); among moss and grass debris on humus, 23 Aug. 1971, OKMJGAL 10547; on moist grassy humus, 23 Aug. 1971, OKMIGAL 10554; among grass debris on peaty soil, 23 Aug. 1971, OKMIGAL Site 1: among moss and dead grass on peat, 9 Aug. 1971, J. Reed, OKM 10412, OKM 10413; among moss and grass debris, 29 Aug. 1971, T. Rau, OKM Site 1: plot 6: on mossy humus, 12 Aug. 1971, OKMI GAL 10475, OKM Site 4: among Drepanoclad~u and Sphagnum, 20 July 1972, OKMIGAL 10708; among Ochrolechin, Lopadiutn and grass debris on peat, 20 July 1972, OKMIGAL 10711; among moss on peat, 21 July 1972, OKMIGAL 10732; among Cnlliergotz on peat in polygon trough, 23 July 1972, OKMIGAL 11249; among moss on peat, 24 July 1972, OKMIGAL 10772; among moss and grass debris on peat, 24 July 1972, OKMIGAL 10773; among Drepanoclad~u uncitintus, mniaceous moss and grass debris on peat, 26 July 1972, OKMIGAL 10784; on peat, 17 Aug. 1972, OKMIGAL 11051, OKM 11069; among Splzngnurn, 13 Aug. 1973, OKMIGAL 11613; among moss on peat, 1 Sept. 1973, T. Rau, OKM 11705; among moss and grass debris on peat, 23 Aug. 1974, OKMIGAL ; among Polzlin and grass debris on polygon top, 23 Aug. 1974, OKMI GAL (ZT 4606); among Polytriclzunz and grass in wet meadow, 23 Aug. 1974, OKMIGAL 11876; on exposed peat on bank, 27 Aug. 1974, OKMIGAL 11937; among Pohlia and Polytrichum, lichens, and dead grass, 5 Sept. 1974, OKMIGAL Site 4: plot 2: among moss, 9 Aug. 1971, OKMIGAL 10382; among Drepanoclad~u, Calliergorz and grass debris on peat, 9 Aug. 1971, OKMIGAL 10383; plot 12: on Drepanocladus, 9 Aug. 1971, OKMIGAL 10404; plot 416: among moss (Pogonatlrm) and Dupontia, 24 July 1972, OKMIGAL 10770; among moss on peat, 1 Aug. 1972, OKMIGAL 10852; among grass debris on peat, 8 Aug. 1972, OKMIGAL 10947; plot 418: among moss and Sphagnum on peat, 1 Aug. 1972, OKMIGAL 10834; plot 419: among D. ~incinatus, and grass debris on peat, 24 July 1972, OKMIGAL 10771; plot 420: among moss on peat, 24 July 1972, OKMIGAL 10765; plot 421: on peat, 8 Aug. 1972, OKMIGAL 10963; plot 425: among Pohlin, minaceous moss, dead grass and Salix on peat, 8 Aug. 1972, OKMIGAL 10933, plot 428: on peat, 24 July 1972, OKMIGAL Site 5: on peat, 3 Aug. 1972, OKMIGAL 10859; among Polytrichurn comnzune on peaty soil, 3 Aug. 1972, OKMJGAL Site 7: among moss and dead grass on peat, 10 Aug. 1972, D. Smith & M. Williams, OKM FAEROES: OSTERO: Slattaratinde, 17 July 1938, Moeller (C; HOLOTYPE of Plz. putnila s. Moeller). In Miller et al. (1973) the above OKM: 10382, 10383, 10404, 10412, 10475, 10518, 10532, 10550, 10554, (no specimens) are recorded as G. subnnnulnta. SWITZERLAND: National Park, Haut Va Sesvenna, 2400 m, Favre 1720 (G; HOLOTYPE of G. purniln f. or-einn Favre); same locality, 2550 m, 20 Aug. 1943, Favre 172n (G). SWEDEN: Abisko, ad castram militum loco turfoso in Hyprzis udis, 21 June 1948, Pilat-Nannfeldt (PR ; HOLOTYPE of G. pse~hrnycet~opsis PilAt & Nannfeldt); Norrbotten, Laktatjakka, 28 Aug. 1985, Moser (IB). ECOL.OGY: Occurs solitary or in clusters. Compared with G. pseudonzycenopsis (form l), the ecological data for the Barrow specimens of form 2, which fits the type material most closely, is better known from the statistical point of view. In total 68 collections with amygdaliform spores have been examined. In 43 cases the basidiomes were in close contact with peat or peaty soil, and in 24 samples the associated mosses have been identified. It is noteworthy that in about half of these records (n = 11) Sphagrl~lm and Drej~anoclndus have been distinctly dominant. Other typical mosses (several species of Calliergorz and Cnrnpyliunz) regionally found in wet localities of the tundra (Rastorfer et al. 1974b) occurred only in a few sites. However, the high preference of form 2 for Sphngtzunz and Drepanocladus is counterbalanced by nine records of Polytriclz~~ttz and Poizlia that as a rule characterize rather dry edaphic conditions. ARCTIC-ALPINE DISTRIBUTION: France (Kuhner and Lamoure 1965 p.p. ; Kiihner 1972a, p.p.); Switzerland (Favre 1955; Kuhner p.p.); Norway (Kuhner p.p.; Kuhner and Lamoure 1965 p.p.; Gulden 1980 p.p.; Gulden et al p.p.); Sweden (Kuhner 1972a p.p.; Kuhner and Lamoure 1965 p.p.); Faeroes (Moeller 1945 p.p.; Bas 1960 p.p.); Svilbard (Gulden 1987 p.p.); U.S.A.: Alaska (Laursen et al. 1972; Miller et al. 1973, 1974; Laursen et al. 1972; Laursen et al. 1977); Antarctic distribution: Palmer Peninsula (Singer 1968; Singer and Corte 1962). 10. Galeritza praticola (Moeller) Orton Figs , 46 Trans. Br. Mycol. Soc. 43: = Plzoliota prnticoln Moeller, Fungi Faeroes, 1 : ILLUSTRATIONS: Moeller (1945); Bas (1960). DESCRIPTION (OKMIGAL 10729): Pileus up to 20 mm diameter, at first hemispheric to obtusely conic becoming broadly umbonate-convex; color rust brown-orange (ferruginous) to dark hazel brown (kaiser brown), paler upon drying, surface hygrophanous, subviscid, waxy, translucently striate when moist, smooth, towards margin with whitish, fibrillose, veil remnants. Lamellae adnate to broadly adnate, moderately distant: color pale ochre (ochraceous buff) to cinnamon brown with rust brown tinge, edges white, subserrulate. Stipe up to 30(-40) x up to 3 mm, cylindric, equal or gradually enlarged towards base; concolorous with or paler than the pileus; whit-

17 430 CAN. J. BOT. VOL FIG. 44. Galerinapseudornycetzopsis Form 1 (VPI 10703). FIG. 45. Galerina pseudomjcenopsis Form 2 (VPI 11051). Frc. 46. Galerina praticoln (VPI 10729). Scale bar = 10 mm. ish fibrillose to submembranaceous annulus subpersistent and often eroded with age, below with several appressed whitish fugaceous zones of veil; dry, solid becoming hollow. Context of pileus thin, pale ochre-brown. Odor and taste not distinctive. Spores 10-12(- 13) X 6-8 pm, subelliptic to amygdaliform or slender, apex occasionally mucronate, rust brown, coarsely marbled to verrucose, plage and perispore distinctive, thick-walled with apical germ pore. Basidia (30-45 x 7-9 pm, 4-spored, occasionally 2-spored. Cheilocystidia X 7-15 pm, polymorphic, slender fusoid, elongated neck gradually tapering with rounded or subcapitate apex, occasionally constricted, thin-walled, hyaline or with yellowbrown plasmatic pigment, occasionally intermixed with clavate to vesiculose cells. Pleurocystidia and caulocystidia scattered, shape and size as cheilocystidia. Pileipellis a cutis of cylindric, subgelatinized, hyaline hyphae (1-4 pm diameter), differentiated terminal cells absent, walls of subcuticular cells encrusted with yellow-brown to orange-rust brown pigment, oleiferous hyphae absent. Clamp connections present on all septa. MATERIAL EXAMINED: CANADA: YUKON TERRITORY: Skolai Pass: among moss, 22 July 1967, OKM 5711 (ZT 4496). U.S.A.: ALASKA: Mead River, ca. 60 mi S. of Barrow: among moss and grass on peaty soil, 30 Aug. 1974, OKMIGAL (ZT 4609). Barrow: Cape Simpson: near oil seep No. 2, among moss (incl. ~icrat~utn sp.) on peaty soil, 28 ~ul; 1976, H. Miller & OKM, OKM (ZT 4627). IBP Tundra Biome Sites: Schultz fertilized plots, about 2 mi SE of NARL: among moss on peaty soil, 30 July 1972, M. Allessio, OKM (ZT 4579); among Drepanocladus, 30 July 1972, M. Allessio, OKM 10827; among Drepanocladus, B. trichophyllutn (L) Dumort. on peaty soil, 30 July 1972, M. Allessio, OKM 10828; among grass debris and Drepanocladus and grass on peaty soil, 3 Aug. 1972; OKMIGAL 10913; among moss, 6 Aug. 1972, OKMIGAL Pintail Polygon Plots: among moss and grass on peaty soil in low center polygon, 31 July 1971, OKMIGAL (ZT 4490). IBP Tundra Biome: Site 1: among moss on peaty soil, 8 Aug. 1971, J. Reed, OKM (ZT 4585). Site 1: plot 11: among moss and grass remnants, 12 Aug. 1971, OKMIGAL (ZT 4587). Site 2: among moss and grass remnants, 20 July 1972, OKMIGAL (ZT 46 15). Site 2: plol 4: among Carnpylium arcticutn and mniaceous moss, 10 Aug. 1971, OKM~GAL 10420, plot 30: on moss detritus, 10 AU~. 1971, OKMIGAL Site 4: among Drepanocladus, 24 Aug. 1971, OKMIGAL (ZT 4572); among moss on peaty soil, 25 Aug. 1971, OKMIGAL (ZT 4614); on D. revolvens among Dupontia-Carex, 22 July 1972, OKMI GAL (ZT 4580); among moss and lichens, 17 Aug. 1972, OKMIGAL Site 4: plot 416: among D. uncinatus and Polytrichum alpinum) and lichens on peaty soil, 8 Aug. 1972, OKMIGAL 10969; plot 417: on moss among Carex- Dupontia in polygon trough, 23 July 1972, OKMIGAL 11250; plot 418: among Calliergon in polygon trough, 24 July 1972, OKMIGAL 10769; among Drepanocladus and Calliergon in polygon trough, 26 July 1972, OKMIGAL Site 5: on D. revolvens, 3 Aug. 1972, OKMIGAL (ZT 4589). Site 7: among D. revolvens, Sphagnum and grass debris on peat, 27 July 1973, OKMIGAL Site 12: among Riccardia pinquis (L) S. F. Gray and Drepanocladus, 23 July 1972, OKMIGAL (ZT 4610); among Drepanocladus, 23 July 1972, OKMIGAL (ZT 4605); among Drepanocladus, 23 July 1972, OKMIGAL 10751; among Calliergon, Cinclidium, and Drepanocladus and grass debris on peat, 13 Aug. 1973, OKMIGAL In Miller et al. (1973) OKM 10376, 10448, 10482, 10601, are recorded as G. subannulata. FAEROES: Nolso 25 Aug. 1938, Moeller (C; HOLOTYPE of Pholiota praticola Moeller). COMPARATIVE MATERIAL: GERMANY: BAVARIA: To~z, Unterbuchen, among Sphagnum, May 1963, Bresinsky (M; HOLO- TYPE of G. beinrothii Bresinsky). ECOLOGY: Galerina praticola occurs solitary or occasionally

18 HORAK AND MIL1.ER 43 1 in clusters. The plots in the tundra near Barrow yielded 28 ) samples of G. praticola, with comparatively few of its basidiomes inhabiting peat or peaty soil. Contrary to the very similar sy mpatric G. pseudonzycenoides, the majority of G. prnticola have been in direct contact with mosses. In about half of the 28 collections, the fungus is associated with either D. uizcinatus or D. revolvens as substrate. On the list of the most preferred mosses, Drepanocladus is followed by Calliergon, Car?zpylium, and Sphagnum. In the Alaskan wet tundra, all four bryophytes are restricted in their occurrence to the most humid and often inundated sites both in low center polygons and interpolygonal troughs (Rastorfer et al ). Further field work is needed to determine if these assessed host relationships of G. praticola are of ecological significance. REMARKS: This is the first record for arctic North America. After examining the type material of this distinctive Galerinn, the same conclusions as previously published by Bas (1960) have been reached. This species is characterized by its slender habit, distinctly umbonate pileus, and its ovoid to ellipsoid spores that lack a mucro at the apex (cf. original illustrations in Moeller 1945). Based upon taxonomically relevant features,. the present Alaskan material essentially agrees with Moeller's.. G. praticola. In regard to gross morphology and ecology, G. praticola appears like G. unicolor (Fries) Kiihner (19726) that, however, is distinctly separated by its much smaller spores. For this reason we (in agreement with Orton 1960) do not support the opinion of Gulden (1980) who synonymizes G. pruticola with G. ~inicolor and also G. beinrothii Bresinsky (1966a). Taxonomically regarded as closely related to the G. marginatacomplex, the spores of the former taxon ought to be comparatively small (see above). Furthermore, the reexamination of the type specimens of G. bienrothii revealed that the spores of this species differ markedly from those of G. prnticola (at least according to our presented interpretation). ARTIC-ALPINE DISTRIBUTION: Faeroes (Moeller 1945), Hebrides (Watling and Richardson 1971; Dennis 1986); Scotland (Watling 19810); Greenland (Lange 1957). For lack of specimens the identity of the following Alaskan records of Galerinn are now considered doubtful: G. macro-... spora (cf. Miller et al. 1974; Laursen and Chmielewsky 1982) and G. stordcllii (cf. Miller et al. 1974; Laursen and Chmielewsky 1982). All cited specimens are kept in the herbarium of 0. K. Miller at VPI and (or) a few are in the Herbarium of E. Horak et ZT. Acknowledgements The authors are very grateful to Barbara Murray (Herbarium, Fairbanks, Alaska) for the identification of numerous moss samples. We also thank C. Bas (Leiden), G. Gulden (Oslo), D. Lamoure (Lyon), R. Kiihner (Lyon), G. Laursen (Fairbanks), and M. Moser (Innsbruck) for providing valuable herbarium material for comparison. Furthermore, for the loan of type specimens we are greatly indebted to the curators of the herbaria C, FH, K, M, MICH, and PR. We thank the Arctic Institute of North America through its Icefield Ranges Research Project and the National Science Foundation, IBP Tundra Biome Program (GV X1) for their support. We wish to thank the Department of Biology, Virginia Polytechnic Institute and State University for sponsoring Dr. E. Horak as a Visiting Professor in Allescher, A,, and Hennings, P Pilze aus den1 Umanakdistrikt In: C. Vanhoffen's Botanischen Ergebnissen der von der Gesellschaft fiir Erdkunde zu Berlin unter Vanhoffen's Sammlungen bearbeitet. A. Kryptogan~en. Bibl. Bot. 42: Anderson, J. H Plants, soils, phytocoenology and primary production of the Eagle Summit tundra bion~e site. U.S. Tundra Biome Data Rep. 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