LYOPHYLLUM & Allies in the Pacific Northwest
Including Keys to Staining Species
By Ian Gibson, April 2010
 2010 Pacific Northwest Key Council

TABLE OF CONTENTS
Introduction	1

Microscopic Key to the Staining Species of Lyophyllum	2

Macroscopic Key to the Staining Species of Lyophyllum	4

Description of Species of Lyophyllum, Sagaranella, Sphagnurus, Tephrocybe, Asterophora, Calocybe, Ossicaulis, and Hypsizygus	6

References	13

Index	14

INTRODUCTION

The group of fungi considered here are considered to share a common molecular ancestry and in Dictionary of the Fungi 10th Edition (2008) include 6 of the 8 genera in the family Lyophyllaceae. They include 
1) the familiar Lyophyllum decastes group, 
2) a group of dark-staining Lyophyllum species, 
3) a group of non-blackening species formerly in Lyophyllum, best learned individually (Sagaranella tylicolor, Sphagnurus paluster, Tephrocybe rancida), and 
4) members of Calocybe, Hypsizygus, Asterophora, and Ossicaulis, also best learned individually. 

Lyophyllum and Calocybe species have siderophilous granules in the basidia. The term siderophilous means that the granules darken when heated in acetocarmine, (the same as carminophilous). This is not a practical test for non-specialists but has been used to help organize the taxonomy of related species, and molecular data confirm its usefulness (Hofstetter et al.). This group of five genera also share inamyloid spores without a germ pore, white to pale cream spore deposit, and (for most species) clamp connections.

The genus Lyophyllum has not had a complete treatment in North America, but the staining species were studied by Heinz Clémençon (1982) and by Heinz Clémençon and Alexander H. Smith (1983). The first was a study of type specimens, and the second a key to North American staining species with descriptions of 25 new species. Various reports of individual non-staining species in the Pacific Northwest have appeared, but more undoubtedly occur.

The microscopic key to the staining Lyophyllum species is formed by including from the original key by Clémençon and Smith only the species recorded from the Pacific Northwest. The macroscopic key is more speculative and is based on elements of the descriptions rather than on detailed field work.

All key-based identifications should be checked against the description of the identified species. If staining collections key out to a given species but do not fit the description well, the original Clémençon and Smith key for North America should be consulted. Some of the other species in their key probably occur in the Pacific Northwest as well.

Hofstetter et al.(2002) point out that a subgroup of the species are parasitic. Tephrocybe palustris grows on Sphagnum. Tephrocybe tylicolor grows on mosses and is thought to be an ammonia fungus (growing better in the presence of ammonia). Two other fungi thought to be closely related by ancestry are also parasites: Hypsizygus is a tree pathogen, Asterophora is a fungal parasite, and Calocybe constricta (not recorded from the Pacific Northwest) is often found in fertilized fields. This implied to Hofstetter et al. that these species need nitrates supplied either externally or through parasitism.

Hofstetter et al. found molecular evidence that the species treated here that they studied fell into four groups by ancestry: One included Calocybe carnea, Calocybe fallax, other Calocybe species, and some brightly colored species of Lyophyllum. The second group (which have free gills unlike other groups) includes Tephrocybe rancidum and members of the ant-associated Termitomyces genus (the latter not recorded from the Pacific Northwest). The third group includes all the parasitic species they studied, discussed in the paragraph above. The fourth group includes Lyophyllum decastes and some blackening species including Lyophyllum semitale. The same groups appear in the molecular work of Moncalvo et al.(2002) with the addition of Ossicaulis lignatilis to the parasitic group. The genera included in our key also appear together in the molecular grouping “Lyophyllaceae” in Matheny et al.(2006). Name changes within the grouping will need to be made in future. Hofstetter et al. conclude, “this study suggests that both taxonomy and nomenclature of the traditional Lyophylleae needs to be revised to reflect a more natural classification system”.

Since that time, Tephrocybe tylicolor was moved to Sagaranella tylicolor  (Fr.) V. Hofst., Clemencon, Moncalvo & Redhead, and Tephrocybe palustris was moved to Sphagnurus paluster (Peck) Redhead & V. Hofst., apparently following up on the Hofstetter et al. recommendation.

MICROSCOPIC KEY TO THE STAINING SPECIES OF LYOPHYLLUM

1a	Spores punctate-rough, finely warted, or spiny	L. scabrisporum
1b	Spores smooth	2

2a	Adaxial spore face rhomboidal, supra-apicular plane or depression usually well-developed	L. geminum
2b	Adaxial spore face round to elliptic, with round or pointed base but not rhomboidal	3

3a	Superficial hyphae of stipe frequently agglutinated into massive, erect fascicles; pileipellis 30-60 um thick, gelatinous, with erect, isolated, clustered or intertwisted hairs	4
3b	Superficial hyphae of stipe different; when arranged in erect groups or tufts, then not intertwisted and pileipellis not gelatinous	6

4a	Pileus densely covered with erect, complex knots of braided hyphae	L. furfurellum
4b	Pileus without such knots	5

5a	Pileipellis with a few scattered, cylindrical, short and inconspicuous hairs	L. stenosporum
5b	Pileipellis with separated, dense clusters of small, repent to erect, contorted, clavate, irregular hairs with colorless or brown walls	L. investitum

6a	(3b) Pileus hygrophanous, margin striatulate when wet; spores asymmetrically ovate in side view, thickest towards the apicular end, thinner at the apex, dimensions in the range 6.2-8.7 x 4.3-5.9 um, Q = 1.3-1.8; pileipellis an ixocutis, 30-45 um thick; basidia 25-35 um long	7
6b	Pileus not hygrophanous or margin not striatulate when wet or spores of different shape or pileipellis not gelatinous or thicker than 45 um, or basidia longer than 35 um	8

7a	Odor distinctly of potatoes, pileus dark fuliginous except at margin, fading to grayish brown, surface uneven and pitted	L. conoideospermum
7b	Odor not distinct, pileus very pale watery gray with pallid margin, pallid overall when faded, surface smooth	L. pallidum

8a	(6b) Odor strongly disagreeable rancid-farinaceous; stipe canescent over a pale blue-gray surface by long, cylindrical-wavy hairs, and hollow; basidia 36-40 x 7-8 um; ixocutis 60-80 um thick	L. canescentipes
8b	Odor not strongly disagreeable rancid farinaceous or stipe not canescent or stipe not hollow or basidia of different dimensions, or ixocutis 60-80 um thick not present	9

9a	Odor spermatic, Inocybe-like; pileus 1-2.5 cm broad, very cartilaginous; spores short-ellipsoidal without supra-apicular depression, 5.7-8.0 x 4.7-5.5 um, Q = 1.2-1.6; pileipellis gelatinous	L. chondrocephalum
9b	Odor not spermatic; pileus larger	10

10a	Odor of green corn; spores spherical to short-ellipsoidal: all of the following criteria apply simultaneously to at least 90% of the spores of a preparation: Q=1-1.45, difference between spore-length and width at most 2 um, supra-apicular depression absent	L. leptosarx
10b	Odors various or not distinct; spores elongate-ellipsoidal, frequently asymmetrical: at least one of the following criteria applies to at least half of the spores of a preparation: either Q greater than 1.45, or difference between spore length and width over 2 um, or spores with a supra-apicular depression	11

11a	Stipe naked, even under the gills; odor farinaceous-rancid; cap 2.5-4 cm broad, hygrophanous, dark fuliginous, thin; gills adnate, subdistant; spores 6.5-8.7 x 3.5-5.3 um, with rounded base, without supra-apicular depression; basidia 27-33 um long; marginal cells hyphoid, 3-6 um broad, scattered to rare	12
11b	Not all the characters simultaneously present; stipe covered with some hairs or even a tomentum, at least below the gills	13

12a	Pileipellis not gelatinous, hyphae 2-4 um thick, smooth, colorless, radially arranged, covered with a few interwoven hyphae 5-8 um wide and with brown incrusting pigments and with numerous colorless, smooth, contorted or ramified, erect, 2-3 um broad hairs	L. fistulosum
12b	Pileipellis gelatinous, superficial hyphae 2 to 3 um broad and smooth; hairs rare, scattered, inconspicuous	L. gracile

13a	(11b) Cap margin translucent-striate; stipe hollow, pruinose at apex; spores with a pointed apicular base, 6.7-9.3 x 3.3-5.0 um, Q = 1.6-2.2	L. semitale
13b	Other combination of characters, spore base round	14

14a	Pileipellis not gelatinous	L. chamaeleon
14b	Pileipellis gelatinous	15

15a	Odor and taste not distinct	L. piceum
15b	Odor farinaceous	16

16a	Basidia 37-42 um long; hyphae of pileipellis interwoven	L. lubricum
16b	Basidia 29-36 um long	17

17a	Cap with erect, cylindrical to slightly irregular hairs, 30-40 x 3-4 um	L. acutipes
17b	Cap without erect hairs	L. solidipes

MACROSCOPIC KEY TO THE STAINING SPECIES OF LYOPHYLLUM

1a	Tricholomatoid stature, dry cap with center pallid to dingy gray and margin pallid, cap becoming blackish where bruised, pale ashy gray flesh that becomes darker where bruised, close, pale olive gray gills that stain bluish and finally black, pallid fibrillose stipe that is darker where handled or when old, faint disagreeable odor, somewhat cespitose growth, (spores punctate-rough, finely warted or spiny)	L. scabrisporum
1b	Not as above	2

2a	Fresh cap dark sooty, lubricous, and bald, gills broad gray becoming blackish where bruised or when old, stipe whitish gray darkening where bruised and appressed-fibrillose at first, odor faintly farinaceous, growth cespitose	. L. furfurellum
2b	Not as above	3

3a	Odor of potatoes	L. conoideospermum
3b	Odor farinaceous, spermatic, green corn, or mild	4

4a	Spermatic or green corn odor	5
4b	Farinaceous or mild odor	6

5a	Green corn odor, flesh thin, stains blue and finally black	L. leptosarx
5b	Green corn odor, flesh slowly gray when bruised	L. geminum var. olens
5c	Spermatic odor, cap cartilaginous, all parts stain directly black	L. chondrocephalum

6a	(4b) Farinaceous odor	7
6b	Mild odor	14

7a	Cap hygrophanous, gray brown to dark brown, moist to buttery, margin translucent-striate, gills pallid or grayish, stem pruinose at apex, often in mountains	L. semitale
7b	Not with this combination of characters	8

8a	Cap dark sooty-gray, when broken cap staining immediately bluish gray to black, pale gray gills, stem hoary over a pale blue-gray surface	L. canescentipes
8b	Not with this combination of characters	9

9a	Stem solid when mature	10
9b	Stem hollow or stuffed with pith when mature	11

10a	Cap pale gray to blackish, (with erect, cylindrical to slightly irregular hairs, 30-40 x 3-4 um)	L. acutipes
10b	Cap buffy brown to olive brown, (without erect hairs)	L. solidipes

11a	(9b) Cap flesh not changing in color when cut, gills staining bluish gray when cut or bruised	L. gracile
11b	Cap flesh changing in color when cut, gills staining various colors when cut or bruised	12

12a	Gills staining yellowish then bluish	L. chamaeleon
12b	Gills staining black	13

13a	Cap 4-7 cm broad, stipe 6-11 cm x 1.0-1.6 cm at top, narrowing down to pointed, somewhat rooting base, stuffed becoming hollow, pallid soon darkening and dingy, innately fibrillose often lacerating into scales that become cinnamon buff	L. lubricum
13b	Cap 2.5-4 cm broad, stipe 2-5 cm x 0.4-1.0 cm, hollow, wavy and compressed or grooved, fragile, pale dingy gray (blackening when old), bald	L. fistulosum

14a	(6b) Gills somewhat decurrent, caps medium to dark gray (fuliginous or sooty) when moist	15
14b	Gills adnate, caps pale gray or drab when moist	16

15a	Flesh staining blackish	L. piceum
15b	Flesh staining bluish gray	L. stenosporum

16a	(14b) Cap drab when moist, stem surface pallid from thin hoary coating	L. investitum
16b	Cap pale gray when moist, hoary coating on stem not recorded	17

17a	Cap lubricous to viscid when wet, flesh turning bluish gray, stem surface not discoloring noticeably when bruised, (spores elliptic 6.2-7.4(8.5) x 4.5-5.2(6.5) um)	L. pallidum
17b	Cap moist, flesh turning gray, stem surface whitish becoming gray where handled, (spores fusoid-ventricose 8.8-11.6 x 4.1-6.2 um)	L. geminum var. geminum

DESCRIPTION OF SPECIES OF LYOPHYLLUM, TEPHROCYBE, SAGARANELLA, SPHAGNURUS, ASTEROPHORA, CALOCYBE, OSSICAULIS, AND HYPSIZYGUS (arranged alphabetically)

Asterophora lycoperdoides	powder-cap
CAP 0.5-2 cm, hemispheric, margin inrolled for a long time; whitish becoming covered with dense brownish powder; dry; flesh thin, whitish-cream. ODOR and TASTE farinaceous. GILLS attached (not free) but often malformed or rudimentary or completely lacking, distant, thick; pale whitish; spore color white when obtainable. STIPE 1-3 cm x 0.3-0.5 cm, more or less equal, bent, soon hollow; whitish becoming brownish in age; silky or cottony to minutely hairy. FRUITING usually grouped or clustered, on dead mushrooms especially Russula and Lactarius. MICROSTRUCTURES spores 3.2-5.8 x 2.0-4.2 um, elliptic, smooth, colorless, also chlamydospores which form the brown powder on the cap, 13-20 x 10-20 um excluding spines, oval to nearly round, verrucose (warty) to bluntly spiny, light brownish; basidia 4-spored; pleurocystidia and cheilocystidia not seen. REMARKS recognized by habitat on mushrooms, small whitish caps which become brown and powdery, and poorly formed gills; Asterophora parasitica has non-powdery, dirty white silky cap and smooth elliptic chlamydospores from well-formed gills.
 
Asterophora parasitica
CAP 0.5-3 cm, convex, then conico-bellshaped to flat or slightly concave, may be umbonate; whitish becoming dark gray brown; smooth to finely fibrillose or appressed silky-floccose; flesh thin, whitish to brownish. GILLS adnate to decurrent, rather distant, becoming thick and fleshy at the base, minutely floccose but waxy-smooth near margin, blunt, at length distorted and anastomosing; well-formed but often poorly developed; spore color white. STIPE 1-6 cm x 0.08-0.4 cm, variable in length, nearly equal or widening slightly downwards, often curved, fibrous, becoming hollow; whitish; cottony with fine hairs or somewhat tomentose especially towards the base. ODOR strong, unpleasant, sour or farinaceous. TASTE typically farinaceous. FRUITING on living or old rotten fruitbodies of Russula and Lactarius species, occasionally other species. MICROSTRUCTURES spores 5-6 x 3-4 um, elliptic, smooth, generally lacking; gills also produce smooth and elliptic to spindle-shaped to oval chlamydospores 12-17 x 9-10 um, typically thick-walled, colorless, inamyloid; cystidia not seen. REMARKS characterized by small size, growth on Russula and Lactarius mushrooms, and non-powdery, dirty white, silky cap; Asterophora lycoperdoides has powdery cap and poorly formed gills.

Calocybe carnea	pink Calocybe 
CAP 1.5-4 cm, convex to flat or slightly umbonate; pinkish to pinkish brown, or varying to dark reddish, or fading to pale tan; dry, smooth; flesh thin, whitish. ODOR and TASTE mild. GILLS adnexed to adnate to slightly decurrent or notched, crowded, narrow; white; spore deposit white. STIPE 1.5-4 cm x 0.2-0.5 cm, more or less equal; colored more or less as cap; smooth or finely fibrillose to felt-like or with fine hairs, sometimes pruinose at top, not viscid. FRUITING scattered or in groups, in lawns, grassy clearings in woods, on moss, or in other open places. MICROSTRUCTURES spores 4-6 x 2-3 um, elliptic, smooth, with droplets; basidia have siderophilous granules; pleurocystidia and cheilocystidia not seen; clamps present. REMARKS recognized by pinkish cap and stem, crowded white gills, and white spore print.
 
Calocybe fallax
CAP 0.5-2(4) cm, convex, umbonate; hygrophanous, yellow to orange-yellow or orange yellow-brown; bald but becoming mealy under a hand lens, to wrinkled; flesh yellow. ODOR faint. TASTE mild. GILLS adnexed to adnate, crowded, narrow to broad, light yellow to brownish yellow. STIPE 1.7-3.7 cm x 0.1-0.4 cm, equal, solid; yellow to yellow-orange, base orange-brown to red-brown; bald or sometimes coated with white to light yellow fibrils, top pruinose, base covered with white felt-like mycelium. FRUITING in forests and alpine habitats. MICROSTRUCTURES spores 3-5 x 2-3 um, elliptic, smooth, inamyloid; cystidia none, clamps present. REMARK The online Species Fungorum, accessed August 16, 2019 gave the current name as Calocybe naucoria (Mirrull) Singer., but MycoBank, accessed the same day, gave the current name as Calocybe fallax.

Calocybe onychina
CAP 2-9 cm, broadly convex to slightly umbonate, becoming nearly flat, margin inrolled when young and often finely scalloped when old; purple-red to wine-red; bald; flesh firm, white to pale yellow. ODOR farinaceous, or somewhat cucumber-like. TASTE mild. GILLS adnate, close, medium breadth, one tier of subgills; dull deep yellow; spore deposit white. STIPE 2-4.5 cm x 0.4-1.5 cm, widening somewhat toward base; dull white with a pinkish to purplish color especially over upper half; dry, hoary. FRUITING several to gregarious, under conifers, June and early July. MICROSTRUCTURES 3-4 x 2-3.5 um, nearly round to broadly elliptic, inamyloid, thin-walled; cystidia absent; clamps absent. REMARKS characterized by purple-red to wine-red cap, bright yellow gills and purplish to pink stem.

Hypsizygus tessellatus
(= Hypsizygus marmoreus (Peck) H.E. Bigelow; Hypsizygus elongatipes (Peck) H.E. Bigelow)
CAP 2-14 cm broad, convex becoming nearly flat; pinkish cream with distinctive darker, round water spots over the center; moist, smooth without hairs; flesh firm but not tough; white to pinkish buff. GILLS “adnexed (notched)”, with a thin line on the upper stem, subdistant, broad, veined; buff to pinkish buff; spore deposit buff. STIPE 4-22 cm x 0.4-2.0 cm, nearly equal or gradually narrowing toward base, often curved or bent; white; smooth except near the top where there is white down, and the base, which has white stiff hairs; veil absent. FRUITING on poplar or occasionally other hardwoods, usually several stems from a common base. MICROSTRUCTURES spores 4-5(6) x 4-4.5 um, round to nearly round, smooth. REMARKS Much confusion has surrounded this species. A number of field guides give Pleurotus ulmarius as a synonym of this species, but the synonymy is incorrect. Redhead(1986) says Hypsizygus tessellatus is a species that often grows cespitosely, in North America on poplar and sugar maple, occasionally on Betula, Ulmus, Abies, and Fagus, has a guttate-marmorate cap when fresh, (Bulliard's 'tessellatus' emphasizes the spots, Peck's 'marmoreus' the lines between, but cap not cracked in a tessellated manner), and has small round to nearly round spores 4-5(6) x 4-5 um. Hypsizygus ulmarius (Bull.: Fr.) Redhead in contrast, he continues, usually grows solitarily, in North America frequently on elm and box elder (Acer negundo), occasionally on Populus and other Acer species, has a smooth not guttate cap when fresh which often becomes areolate-cracked with age and has larger nearly round to broadly elliptic spores (5)5.5-6(7) x 5-5.5(6) um. Both species are found in Eurasia; H. ulmarius according to Redhead(1984) is found in North America in areas east of the continental divide and generally central latitudes while H. tessellatus (Bull. ex Fr.) Singer is found is found from coast to coast and north to the Yukon. The description above is derived from the Miller(1979) description of Pleurotus elongatipes, except the habitat and microstructures which are from Redhead. The Miller(2006) description of Hypsizygus tessulatus is very close to this one and appears to be based on it. Notice that the name is more often spelled Hypsizygus tessulatus. 

Lyophyllum acutipes
CAP 3-8 cm, hygrophanous, pale gray-brown becoming blackish, fading to sordid pale gray where not blackened; bald, lubricous; flesh thin, pliant, cartilaginous, darkening where bruised. GILLS adnate becoming adnexed, close, 3 tiers of subgills, narrow but broad in large caps, dingy gray when young, black-spotted when old. STIPE 5-8 cm x 0.8-1.6 cm, narrowed downward to pointed base, solid; whitish varying to grayish white, darkening where handled or bruised; dry, unpolished. ODOR farinaceous. TASTE slightly farinaceous. FRUITING type gregarious in moss under pine. MICROSTRUCTURES spores 7.2-8.5 x 4.2-4.8 um.

Lyophyllum canescentipes
CAP 4-10 cm, dark sooty-gray drying dirty gray, staining immediately bluish gray to black when broken; bald; flesh stains bluish gray to black immediately when broken. GILLS adnate, close to subdistant, moderately broad; pale gray. STIPE 4-8 cm x 0.5-1.1 cm, hollow, canescent over a pale blue-grey surface, scarcely darkening below. ODOR and TASTE strongly disagreeable farinaceous. FRUITING clustered to gregarious on soil. MICROSTRUCTURES spores 6.9-9.1 x 4.7-6.4 um. REMARKS Other listed staining Lyophyllum species with hoary stems lack the unpleasant farinaceous odor.

Lyophyllum chamaeleon 
CAP 3-4.5 cm, hygrophanous, near blackish brown to clove brown when moist, slowly fading to olive brown or grayer; moist, bald, margin slightly pubescent (downy) at first, rigid-cartilaginous because of thick cuticle; flesh pallid grayish, slowly staining gray or darker when cut. GILLS pale when young, becoming drab, broadly adnate-subdecurrent, close, narrow, staining yellowish and then bluish. STIPE 6-10 cm x 1.0-1.5 cm, narrowing downward sometimes, stuffed; whitish-gray, soon dingy gray where handled; silky-fibrillose to unpolished, more or less longitudinally striate. ODOR and TASTE farinaceous. FRUITING type cespitose under pine. MICROSTRUCTURES spores 7.1-8.8 x 3.5-4.4 um.

Lyophyllum chondrocephalum 
CAP 1-2.5 cm, hygrophanous, dark fuliginous (sooty) to blackish when moist fading to dingy yellowish gray, blackening where cut or bruised; moist, bald; flesh thin and pliant. GILLS somewhat adnexed, subdistant, broad; drab or nearly so when young, glaucous, blackening when bruised. STIPE 2-3 cm x 0.4-0.8 cm at top, widened downward, often compressed, hollow; colored as cap or paler, blackening with injury; thin coating of grayish, silky fibrils giving a canescent somewhat pruinose appearance at first, bald when old. ODOR spermatic. TASTE disagreeable. FRUITING type was cespitose along an old road on soil. MICROSTRUCTURES spores 5.7-8.0 x 4.7-5.5 um. 

Lyophyllum conoideospermum
CAP 9 cm across, hygrophanous, dark sooty except margin watery gray, fading to sordid grayish brown; moist, bald; flesh thin, cartilaginous, gray, changing to bluish gray when cut or bruised. GILLS broadly adnate to subdecurrent, moderately close, 3 tiers of subgills, gills broad, broadest in middle; pale drab staining blackish, edges staining blackish. STIPE 13 cm x 1.5 cm, stuffed; pale gray; bald or long-striate. ODOR distinctly of potatoes. TASTE slightly disagreeable to mild. FRUITING type single to gregarious on humus. MICROSTRUCTURES spores 7.0-8.7 x 4.3-5.9 um. 

Lyophyllum decastes (= Lyophyllum aggregatum)	fried chicken mushroom
CAP 3-15 cm across, gray-brown to tan or yellowish brown with silky or silvery streaks (color varies from this if forms are included which some regard as separate species); smooth, soapy; flesh firm; white, not staining. GILLS adnate to slightly decurrent, close, moderately narrow to moderately broad; white to grayish. STIPE 3-10(20) cm x 0.8-2.5 cm, cylindric to clavate, sometimes narrowed toward base, solid, often twisted or off-center, consistency fibrous, tough; white at top becoming brownish toward base, longitudinally fibrillose, white-pruinose at top. ODOR and TASTE mild. FRUITING in clusters on ground, sometimes in fairy rings, more rarely single, in waste places, grassy areas, wood edges, paths, or in forests. MICROSTRUCTURES spores 5-7 x 5-6.5 um, nearly round. REMARKS Arora includes the following in a Lyophyllum decastes group: Lyophyllum connatum with white or whitish cap (easily confused with Clitocybe dilatata); Lyophyllum loricatum with blackish-brown to dark brown cap when young, often with hoary sheen or metallic luster, and thick cartilaginous cap cuticle, fading with age to paler brown or tan and then indistinguishable from L. decastes, (Arora). Lyophyllum fumosum is another color form that is sometimes considered a separate species, (Trudell & Ammirati). Lyophyllum semitale is similar in color to the common brown forms of L. decastes, but "is somewhat smaller, grows singly, in groups, or small clusters, and turns black in age or when bruised, although often the change is slow", and spores are larger (6.5-9 x 3-4.5 um), (Trudell & Ammirati).

Lyophyllum fistulosum
CAP 2.5-4 cm; hygrophanous, dark sooty color, fading gradually to dingy brown; moist, bald, margin opaque; flesh thin, cartilaginous, watery sooty and slowly staining black. GILLS adnate, subdistant, broad; dingy gray, paler than cap, edges soon staining black where bruised. STIPE 2-5 cm x 0.4-1.0 cm, hollow, wavy and compressed or grooved, fragile; colored as gills, blackening when old; bald, not pruinose at top. ODOR and TASTE disagreeable, somewhat farinaceous. FRUITING type cespitose on soil. MICROSTRUCTURES spores 6.5-8.7 x 4.1-5.3 um, elliptic to slightly oval in face view.

Lyophyllum furfurellum
CAP 3-9cm, hygrophanous, dark sooty color, fading to dark dingy gray with cinnamon buff tinge at least over disc; lubricous when moist, bald; flesh thin, cartilaginous (from rigid cuticle), sooty then pallid, darkening at least slightly where bruised. GILLS broadly adnate to subdecurrent becoming adnexed or seceding, moderately close, 3 tiers of subgills, gills broad; pallid dingy ash gray, becoming blackish where bruised or when old. STIPE 3-8cm x (0.6)0.8-1.5cm, more or less equal, hollow; whitish gray at first, darkening when bruised; appressed-fibrillose to lacerate-fibrillose, becoming more or less bald. ODOR faintly farinaceous. TASTE mild. FRUITING type was cespitose under pine on soil. MICROSTRUCTURES spores 7.8-9.8 x 3.8-4.6 um, oval to elliptic.
 
Lyophyllum geminum
CAP 3-6 cm across, hygrophanous, pale smoky gray, fading to pale alutaceous; moist, bald; flesh thin, pliant, pallid, slowly staining gray when bruised. GILLS broadly adnate, close to subdistant, broad; whitish, edges staining grayish. STIPE 3-4 cm x 0.8-1.2 cm at top, equal to narrowly club-shaped; whitish becoming gray where handled; bald. ODOR not distinct, although var. olens from Michigan has a green corn odor. TASTE not distinct. FRUITING type of var. geminum was single to scattered on sandy soil. MICROSTRUCTURES spores 8.8-11.6 x 4.1-6.2 um, broadly fusoid-ventricose. REMARKS Lyophyllum pallidum is another staining Lyophyllum that is hygrophanous pale gray, but it has elliptic spores. 

Lyophyllum gracile
CAP 2.5-3 cm across, hygrophanous, disc dark sooty, margin buffy brown, cap fading to dingy grayish brown; moist, bald, faintly striate on extreme margin; flesh thin, colored as surface, no color change on bruising or cutting. GILLS bluntly adnate to depressed-adnate, subdistant, 0.5-0.8 cm broad, ventricose; pallid becoming only slightly darker but staining bluish gray when cut or bruised. STIPE 3-5 cm x 0.35-0.5 cm, equal down to tapered, somewhat rooting base, solid or with slight tubule; pallid watery gray in upper part, darker in lower part; at first bald with a few appressed silvery fibrils near base, when old appearing fibrous-striate. ODOR and TASTE rancid-farinaceous. FRUITING type closely gregarious on soil under conifers in mixed woods. MICROSTRUCTURES spores 6.8-8.6 x 3.5-5.0 um, elliptic to ovoid.

Lyophyllum investitum
CAP 3-4.5 cm across, evenly drab when moist, fading to ash-gray; lubricous, bald, margin faintly striate; flesh 0.2-0.3 cm thick, equal, watery, becoming dingy gray when bruised. GILLS horizontal, subdistant, 2 tiers of subgills, gills about 0.5 cm broad; ash-gray, edges darkening to dingy gray when bruised. STIPE 5-8 cm x 0.4-0.7 cm at top, hollow, fragile, bases arising in large numbers from a common origin, not arising from one point and not attached to a fleshy mass; pallid from a thin hoary coating, pallid grayish at maturity. ODOR and TASTE mild. FRUITING type cespitose under larch and pine. MICROSTRUCTURES spores 7.7-9.5 x 4.0-5.6 um, oval with narrower base or elliptic.

Lyophyllum leptosarx
CAP 1-4 cm across, hygrophanous, pale sooty to sooty brown, streaked with grayish pallid in fading; moist, obscurely translucent-striate before fading; flesh thin, staining blue and finally black when injured. GILLS adnate, close, broad; pallid grayish brown, becoming near olive buff, spotted blackish when old. STIPE 2-3.5 cm x 0.3-1.0 cm, hollow, fragile; colored as cap margin, darkening to olive brown from base up; bald. ODOR strong of green corn. TASTE mild. FRUITING cespitose to scattered on soil. MICROSTRUCTURES spores 5.4-7.2 x 4.5-5.9 um, short-elliptic. REMARKS Lyophyllum geminum var. olens from Michigan and L. luteogriseascens Clemencon & A.H. Sm. from Michigan (with creamy white cap and gills that stain yellow to gray when bruised) both have green corn odor; L. chondrocephalum has spermatic odor.

Lyophyllum loricatum	frosty Lyophyllum
CAP 3-13 cm across, pliant, cartilaginous; hygrophanous, dark olive-brown to dark gray-brown or chestnut-brown; satiny or with a hoary sheen, often veined, ribbed, bumpy, or wrinkled, as if armored with a thick cartilaginous skin, the cuticle breaking up into tiny granules as the surface expands, revealing the white background and giving the surface a spotted or speckled look; flesh elastic, tough to cartilaginous (especially cuticle - when broken the cap gives a distinct snapping sound), whitish. GILLS adnate to somewhat notched or sometimes short-decurrent as a tooth, close, broad, 3-7(11) subgills between each pair of gills; whitish to gray-whitish. STIPE 3.5-10 cm x 0.7-1.5 cm, cylindric, solid; cream to pale brownish, gray-brown when old; almost smooth, longitudinally fibrillose, white-powdered at top; often fused and not only at the base. ODOR herbaceous or mild. TASTE mild, at times somewhat peppery or burnt. FRUITING usually cespitose, along edges of forests and paths, in forests, gardens, and parks, primarily under hardwoods, on bare soil or among grass or herbs. MICROSTRUCTURES spores 5-7 x 4.5-7 um, nearly round to round, smooth. REMARKS Lyophyllum loricatum is like a dark version of L. decastes and when old resembles that species.

Lyophyllum lubricum
CAP 4-7 cm across, hygrophanous, dark to light watery gray soon becoming dark sooty, fading to grayish; lubricous but not viscid; flesh gray-brown to dingy gray, watery gray when moist, staining black when old or where bruised. GILLS rounded adnate to narrowly adnexed, seceding, close to subdistant, about 1 cm broad near stem; pale gray, staining black. STIPE 6-11 cm x 1.0-1.6 cm at top, narrowing down to pointed somewhat rooting base, stuffed becoming hollow; pallid soon darkening and dingy; innately fibrillose often lacerating into scales that become cinnamon buff. ODOR and TASTE farinaceous. FRUITING type gregarious on soil under conifers. MICROSTRUCTURES spores 6.8-9.2 x 3.5-4.7 um, elliptic to slightly oval.

Lyophyllum pallidum
CAP 3-6 cm across, hygrophanous, pale watery gray on disc, pallid on margin, fading to pallid overall; lubricous to viscid when wet, bald and shining, margin striate; flesh thin, pliant, colored as surface, changing to bluish gray when cut or bruised. GILLS bluntly adnate, moderately close, 2 tiers of subgills, gills moderately broad, broadest at stem; white to pallid and staining pale bluish gray when cut or bruised.. STIPE 5-8 cm x 0.8-1.0 cm, equal or narrowing at base, hollow, pliant; whitish gray, paler than cap and with drier appearance, not discoloring noticeably where bruised; obscurely striate. ODOR mild. FRUITING type gregarious on soil under conifers. MICROSTRUCTURES spores 6.2-7.4(8.5) x 4.5-5.2(6.5) um, elliptic, smooth. REMARKS Lyophyllum geminum is another staining species that is hygrophanous pale gray but it has spindle-shaped spores.

Lyophyllum piceum
CAP 4-8 cm, hygrophanous, dark sooty fading to dingy grayish brown then pale ash gray; bald, margin pruinose, becoming striate before fading; flesh thin, pliant, cartilaginous, watery gray becoming pallid, changing to blackish when bruised or cut. GILLS subdecurrent, close to subdistant, 2-3 tiers subgills, gills narrow becoming moderately broad, broadest near stem; pale drab and blackening where bruised. STIPE 4-8 cm x 1-2 cm, pallid to dark gray, paler in upper part; unpolished, more or less longitudinally striate, sometimes scurfy in upper part. ODOR and TASTE mild. FRUITING type single to scattered under conifers. MICROSTRUCTURES spores 9.0-11.1 x 4.5 -5.7 um, ovate to long-elliptic. REMARKS Lyophyllum rugulosum Clemencon & A.H. Sm. from California has lubricous, often finely wrinkled, watery gray, blackening cap, pale olive buff blackening gills, mild odor, and spores 6-7.7 x 3.4-4.7 um.

Lyophyllum scabrisporum
CAP 4-9.5 cm across, center pallid to dingy gray and darker when old, margin pallid, cap becoming blackish where bruised; dry, disc streaked without visible fibrils, margin minutely pubescent at first, opaque at all stages; flesh pale ashy gray but darker where bruised. GILLS sinuate-adnate, close, moderately broad, broader in middle; pale buffy brown, pale olive gray, staining bluish and finally black. STIPE 3-5 cm x 1.0-1.3 cm, equal above narrow base, solid, somewhat cartilaginous, pallid, darker where handled or when old; appressed-fibrillose, fibrillose-scurfy toward top. ODOR faint and disagreeable. TASTE mild. FRUITING type was somewhat cespitose under conifers and alder. MICROSTRUCTURES spores 6.2-7.9 x 2.7-3.5 um, cylindrical with rounded ends, finely punctate, roughened but not warty. REMARKS The other known staining Lyophyllum species with rough spores are L. leucophaeatum (P. Karst.) P. Karst., with basidia 17-20 um long, and even rather than eroded gill edges, and L. oldae (Svrček) Clémençon with mycenoid rather than tricholomatoid stature, cap less than 2.5 cm across, and elliptic spores wider than 4.5 um.

Lyophyllum semitale
CAP 2-10 cm, hygrophanous, beige-brown or gray brown to dark bister brown or sooty, staining dark gray to black eventually where injured; bald, smooth, dull, bald, moist, somewhat buttery, opaque to translucent-striate; flesh thin, whitish to watery gray-brown, blackening. GILLS adnate to somewhat decurrent, close, broad, 3 subgills between neighboring gills; cream to gray or pale brown, staining yellowish then bluish and finally black after several hours. STIPE 4-10 cm x 0.3-1.5 cm, equal or at times somewhat widened toward base or also slightly fusiform, solid, pithy, becoming hollow; similar in color to cap, white to pale gray, becoming dingy ocherish or grayish, blackening when old; longitudinally striate and/or fibrillose. ODOR rancid-farinaceous. TASTE mild to rancid-farinaceous. FRUITING gregarious or in small clusters under conifers or hardwoods. MICROSTRUCTURES spores 6.5-9.5 x 3.2-5 um, elliptic, smooth. REMARKS Lyophyllum semitale is fairly common in mountains but also found elsewhere in the Pacific Northwest. A slow blackening reaction may make it difficult to recognize. Lyophyllum chamaeleon also has gills that stain yellowish then bluish, but L. chamaeleon tends not to stain as darkly finally, and cap is not translucent-striate.

Lyophyllum solidipes
CAP (3.5)5-11 cm, hygrophanous, margin pale watery gray, disc darker buffy brown to olive brown and soon with blackish areas when old, sometimes blackish brown overall; lubricous to subviscid when wet, bald, moist, margin more or less striate; flesh thin, pliant, colored as surface, gradually staining dark gray to black when cut or bruised. GILLS abruptly adnexed, moderately close, 2-3 tiers of subgills, gills broad near stem, tapering toward margin; pale olive buff and gradually darker gray, staining dark gray where cut or bruised. STIPE 7-12 cm x 0.8-1.5 cm at top which is sometimes slightly enlarged, equal down to narrow base, solid; pallid becoming grayish and blackening when bruised; bald or slightly fibrillose-punctate in upper part, longitudinally striate. ODOR and TASTE farinaceous. FRUITING type cespitose along trail among Douglas-fir. MICROSTRUCTURES spores 6.5-8.1 x 3.7-4.7 um, elliptic.

Lyophyllum stenosporum
CAP 3-5 cm, hygrophanous, sooty color fading to ashy brown and drying dingy gray-brown; lubricous and shiny, bald; flesh staining bluish gray. GILLS short decurrent, distant to subdistant, broad; pale gray to dingy brown, staining where bruised. STIPE 2-3 cm x 0.5-0.8 cm at top, club-shaped; whitish; pruinose at top, bald below. ODOR and TASTE mild. FRUITING type gregarious under conifers. MICROSTRUCTURES spores 8.3-10.3 x 4.0-4.7 um, narrowly elliptic to narrowly oval, smooth.

Sagaranella tylicolor, (=Tephrocybe tylicolor,= Lyophyllum tylicolor)
CAP 0.5-2 cm across, always with an distinct umbo, margin sharp, projecting somewhat beyond gills; hygrophanous, red-brown to yellow-brown or dark gray-brown to almost black, toward margin paler when moist; dull to shiny, silky-floccose when young but later slightly fibrillose, not striate to heavily grooved almost to center; flesh thin, gray-brown and watery when moist. GILLS finely adnexed to almost free, 22-25 reaching stem, broad, 3 subgills between neighboring gills, edge may be scalloped; whitish, light gray, yellowish, brownish, or gray-black. STIPE 3-6 cm x 0.1-0.2 cm, equal, narrowing downward, or somewhat rooting; hollow, fragile, somewhat twisted; silvery white, sometimes with a faint brownish tint (especially toward base), sometimes gray; smooth, white silky fibrillose, satiny, longitudinally striate, top somewhat white-floccose. ODOR slightly radish-like. TASTE mild, slightly farinaceous. FRUITING single to gregarious or in rings, on rotting flesh, rotting mushrooms, human feces, on leaves or needle beds, on mosses, on humus. MICROSTRUCTURES spores 5.8-8 x 4.8-6 um, broadly elliptic to nearly round, spiny.

Sphagnurus paluster (=Tephrocybe palustris,= Lyophyllum palustre)	marsh Lyophyllum
CAP 1-3 cm, hygrophanous, olive brown as a rule but often more umber brown or ocher brown and sometimes pale watery gray; grooved, smooth, not lubricous, at first covered by a faint white bloom but soon bald and watery; flesh thin, soft, watery, without distinct color change when crushed or bruised. GILLS adnate or finely adnexed or with a slight tooth, close to subdistant, narrow to broad, rather thick at times and interveined, often forked, 1-7 subgills between neighboring gills; whitish to pale smoky gray. STIPE 4-10 cm x 0.1-0.5(0.7) cm, equal, fragile, hollow, base sometimes slightly widened; gray to gray-brown, paler than cap; white-pruinose when young becoming bald and watery, or may be gray-whitish-fibrillose. ODOR farinaceous. TASTE mild, farinaceous. FRUITING single to gregarious among Sphagnum in bogs, sometimes growing on moss, sometimes in ditches. MICROSTRUCTURES spores 6-8 x 4-4.5 um, elliptic, smooth, inamyloid. REMARKS Sphagnurus paluster is characterized by parasitism on Sphagnum. It kills Sphagnum in distinct areas around it where moss becomes gray: fruitbodies often occur close to the living moss.

Tephrocybe rancida  (=Lyophyllum rancidum)
CAP 2-5 cm, always with distinct, obtuse umbo; gray-brown, black-gray with a steel-blue tint, not hygrophanous; smooth, satiny, innately radially fibrillose, sometimes whitish-pruinose, sometimes finely wrinkled; flesh thin, whitish when young becoming gray. GILLS strongly notched to almost free, 36-42 reaching stem, broad, 3-5(7) subgills between neighboring gills; brown-gray to ocher-gray. STIPE 4-10 cm x 0.2-0.5 cm, equal, with a fusiform root-like extension up to 5 cm long, stem hollow, sometimes off-center; gray to brown-gray, top paler; smooth to slightly longitudinally silky-fibrillose, top white-pruinose. ODOR strongly farinaceous-rancid. TASTE mild, farinaceous. FRUITING single to gregarious in hardwood and coniferous forests. MICROSTRUCTURES spores 6.1-8.4 x 3-4.5 um, elliptic. REMARKS fairly common.

Ossicaulis lignatilis (= Pleurotus lignatilis)
CAP 1.5-12(24) cm across, convex to pulvinate (cushion-shaped) initially, becoming flat to shallowly depressed, with inrolled to incurved edges; grayish to dirty whitish to grayish or gray-brownish, often chalky-white, with or without faint rose to pale vinaceous tints on disc; pruinose to mealy, or rarely somewhat tomentose, dry, opaque; flesh fleshy and tough. GILLS adnate to adnexed or decurrent by short tooth, crowded to subcrowded, up to 3 tiers of subgills, moderately narrow (up to 0.6 cm broad); white but drying yellowish white; spore deposit white. STIPE 1-6(18) cm x 0.15-1(6) cm, variable, often short and strongly curved, usually swollen in lower part, usually off-center, often with clustered aborted small caps at base; usually chalky-white, occasionally with faint vinaceous-rose tint; when old browning somewhat; surface similar to cap, often slightly more matted-roughened at top, otherwise like suede. ODOR fungal, may be farinaceous when first cut. TASTE mild to strongly fungal or slightly farinaceous. FRUITING mainly on stumps or dead or living trunks of old growth hardwoods and occasionally conifers. MICROSTRUCTURES spores 4-6 x 3.0-3.5 um, elliptic to broadly elliptic, smooth, inamyloid; basidia 1-spored, 2-spored, 3-spored, 4-spored.

REFERENCES

Arora, David. 1986 Mushrooms Demystified Second Edition. Ten Speed Press, Berkeley.
Clémençon, Heinz. 1982. “Type studies and typifications in Lyophyllum (Agaricales). I. Staining Species.” Mycotaxon 15: 67-94.
Clémençon, Heinz, Alexander H. Smith. 1983. “New Species of Lyophyllum (Agaricales) from North America and a key to the known staining species.” Mycotaxon 18(2): 379-437.
Hofstetter, Valérie, Heinz Clémençon, Rytas Vilgalys, and Jean-Marc Moncalvo. 2002. “Phylogenetic analyses of the Lyophylleae (Agaricales, Basidiomycota) based on nuclear and mitochondrial rDNA sequences.” Mycol. Res. 106(9) 1043-1059.
Matheny, P. Brandon, Judd M. Curtis, Valérie Hofstetter, M. Catherine Aime, Jean-Marc Moncalvo, Zai-Wei Ge, Zhu-Liang Yang, Jason C. Slot, Joseph F. Ammirati, Timothy J. Baroni, Neale L. Bougher, Karen W. Hughes, D. Jean Lodge, Richard W. Kerrigan, Michelle T. Seidl, Duur K. Aanen, Matthew DeNitis, Graciela M. Daniele, Dennis E. Desjardin, Bradley R. Kropp, Lorelei L. Norvell, Andrew Parker, Else C. Vellinga, Rytas Vilgalys, and David S. Hibbett.  2006.  “Major clades of Agaricales: a multilocus phylogenetic overview.”  Mycologia 98(6): 982-995.
Miller Jr., Orson K. 1979. Mushrooms of North America. E.P. Dutton. New York Third Printing.
Miller Jr., Orson K., Hope H. Miller. 2006. North American Mushrooms. A field guide to edible and inedible fungi. Falcon Guide.
Moncalvo, Jean-Marc, Rytas Vilgalys, Scott A. Redhead, James E. Johnson, Timothy Y. James, M. Catherine Aime, Valérie Hofstetter, Sebastiaan J.W. Verduin, Ellen Larsson, Timothy J. Baroni, R. Greg Thorn, Stig Jacobsson, Heinz Clémençon, and Orson K. Miller Jr. 2002. “One hundred and seventeen clades of eugarics.” Molecular Phylogenetics and Evolution 23 357-400.
Redhead, S.A. 1984. “Mycological observations 13-14: on Hypsizygus and Tricholoma.”  Trans. Mycol. Soc. Japan 25: 1-9.
Redhead, S.A. 1986. “Mycological Observations 15-16: On Omphalia and Pleurotus.” Mycologia 78(4): 522-528.
Singer, Rolf.  1986.  The Agaricales in Modern Taxonomy  4th Edition.  Koeltz Scientific Books, Koenigstein, Germany.  981 pp.  
Trudell, Steve, Joe Ammirati. 2009. Mushrooms of the Pacific Northwest. Timber Press.

INDEX
		Key Lead	Page
ASTEROPHORA  Ditmar
	A. lycoperdoides  (Bull.) Ditmar	6
	A. parasitica  (Bull. ex Pers.) Singer	6
CALOCYBE  Kühner ex Donk
	C. carnea  (Bull.) Donk	1, 6
	C. fallax  (Sacc.) Redhead & Singer	1, 6
	C. onychina  (Fr.) Donk	7
HYPSIZYGUS  Singer
	H. tessellatus  (Bull. ex Fr.) Singer	7
LYOPHYLLUM  P. Karst.
	L. acutipes  Clémençon & A.H. Sm.	Mic17a, Mac10a	4, 5, 7
	L. canescentipes  Clémençon & A.H. Sm.	Mic8a, Mac8a	4, 5, 7
	L. chamaeleon  Clémençon & A.H. Sm.	Mic14a, Mac12a	4, 5, 8, 11
	L. chondrocephalum  Clémençon & A.H. Sm.	Mic9a, Mac5c	3, 4, 8, 9
	L. conoideospermum  Clémençon & A.H. Sm.	Mic7a, Mac 3a	3, 4, 8
	L. decastes  (Fr.) Singer	1, 2, 8, 10
	L. fistulosum  Clémençon & A.H. Sm.	Mic12a, Mac13b	3, 5, 8
	L. furfurellum  Clémençon & A.H. Sm.	Mic4a, Mac2a	2, 4, 9
	L. geminum  Clémençon & A.H. Sm.	Mic2a, Mac5b, Mac17b	2, 4, 5, 9, 10
	L. gracile  Clémençon & A.H. Sm.	Mic12b, Mac11a	3, 5, 9
	L. investitum  Clémençon & A.H. Sm.	Mic5b, Mac16a	2, 5, 9
	L. leptosarx   Clémençon & A.H. Sm.	Mic10a, Mac5a	3, 4, 9
	L. loricatum  (Fr.) Kühner ex Kalamees		8, 10
	L. lubricum  Clémençon & A.H. Sm.	Mic16a, Mac13a	4, 5, 10
	L. pallidum  Clémençon & A.H. Sm.	Mic7b, Mac17a	3, 5, 9, 10
	L. palustre  (Peck) Singer - see Tephrocybe palustris
	L. piceum  Clémençon & A.H. Sm.	Mic15a, Mac15a	4, 5, 10
	L. rancidum  (Fr.) Singer - see Tephrocybe rancida
	L. scabrisporum  Clémençon & A.H. Sm.	Mic1a, Mac1a	2, 4, 11
	L. semitale  (Fr.) Kühner ex Kalamees	Mic13a, Mac7a	2, 3, 4, 8, 11
	L. solidipes  Clémençon & A.H. Sm.	Mic17b, Mac10b	4, 5, 11
	L. stenosporum  Clémençon & A.H. Sm.	Mic5a, Mac15b	2, 5, 11
	L. tylicolor  (Fr.) M. Lange & Sivertsen - see Tephrocybe tylicolor
TEPHROCYBE  Donk
	T. palustris  (Peck) Donk	1, 12
	   = Lyophyllum palustre  (Peck) Singer
	T. rancida  (Fr.) Donk	1, 12
	   = Lyophyllum rancidum  (Fr.) Singer
	T. tylicolor  (Fr.) M.M. Moser	1, 12
	   = Lyophyllum tylicolor  (Fr.) M. Lange & Sivertsen
OSSICAULIS  Redhead & Ginns
	O. lignatilis  (Pers.) Redhead & Ginns	2, 12
	   = Pleurotus lignatilis  (Pers.) P. Kumm.