The North American Species of Pholiota

Subgenus Flammuloides Subg. nov.

Pileus viscidus vel glutinosus, glaber vel floccoso-squamulosus. Stipes glaber vel fibrillosus vel subsquamosus. Pleurocystidia conspicua. Subhymenium gelatinosum vel subgelatinosum, rare non-gelatinosum. Typus: Pholiota subfulva.

Pileus viscid to glutinous from a gelatinous layer in the cuticle, hyphal walls usually gelatinizing, glabrous to merely virgate with fibrils, or rarely at first squamulose from superficial veil remnants; margin sometimes appendiculate; stipe very seldom conspicuously scaly but often decorated with colored zones or patches of veil material; pleurocystidia prominent and conspicuously projecting; subhymenium typically gelatinous.

Key to Sections

1. At least some pleurocystidia with walls thickened to 0.5 µ (or much thicker), the wall thickening should be observed at the base of the neck or in the ventricose portion
Section Flammuloides
1. Pleurocystidia consistently thin-walled
2
2. Always fruiting on burned ground around charcoal
Section Carbonicola
2. Not as above, but habitat typically lignicolous, more rarely on soil or humus
3
3. Spores 7-10 x 5-6 µ but if 7-9 x 4-5 µ try this choice also
Section Spumosae
3. Spores smaller 5-7.5 (8) x 3-4.5 (5) µ
Section Lubricae

Section Flammuloides

The diagnostic features of the section are: The viscid to glutinous pileus, typically gelatinous subhymenium, very prominent pleurocystidia, and the walls of the latter in many of the cystidia in a hymenophore thick enough to measure (about 0.5 µ or more) as revived in KOH or in Melzer's reagent. It remains to be seen how much of this thickening shows on fresh material, but from our experience so far it appears that the feature is a reliable one. In addition to the species formally placed in this section P. baptistii, P. spinulifera, P. haerenosa, P. innocua, P. stratosa and P. pseudopulchella show a tendency toward wall thickenings in the cystidia, and in P. innocua thick-walled cheilocystidia are numerous enough to cause one to wonder if eventually some thick-walled pleurocystidia will be found.

Key to Stirpes of Section Flammuloides

1. Subhymenium cellular and not gelatinous
1. Subhymenium gelatinized by maturity
2
2. Spores 7-9 (10) x 3.7-4.5 (6) µ
Stirps Condensa
2. Spores smaller than above
3
3. Pileus pallid to yellowish on disc at first, but many become olivaceous over marginal area
3. Pileus with different coloration
4
4. Pileus dark vinaceous brown on disc at first or very dull red-brown
Stirps Decorata
4. Not colored as above
5
5. Pileus ferruginous, bay, hazel etc. (see P. sublubrica also)
Stirps Ferruginea
5. Pileus with cinnamon to russet or olive brown colors
6
6. Pileus on disc at first russet, cinnamon, tan or clay color
6. Pileus olive to olive-brown or tawny-olive
Stirps Virgata

 

Stirps Virgata

The characteristic olive tones are evident in both the fresh material and the dried specimens. Aside from the color and streaked appearance of the pileus the stirps is very close to stirps Occidentalis. As yet the stirps contains only the type species and P. pseudograveolens, which has a strong odor.

1. With strong odor
1. Not as above

Stirps Olivaceophylla

This stirps is placed here because a few of the pleurocystidia develop walls up to around 0.5 µ thick, but it is to be regarded as borderline. In all features save the lack of gelatinization of the subhymenium it appears to be a group typical of subg. Flammuloides. Both of the species included here need further critical study based on ample collections.

1. Lamellae white when young; pileus chestnut to bister in color from youth to age
1. Lamellae olivaceous young; pileus yellowish tan to whitish

Stirps Adirondackensis

Here we have three essentially pale yellow species all on wood of conifers; two from Idaho and one from New York. The type species has very thick cystidial walls, in P. flavopallida they are relatively thin, and in P. agglutinata they are intermediate.

1. Odor rather strong and penetrating; young lamellae olive-yellow young
1. Odor not distinctive; lamellae white to pallid when young
2
2. Pleurocystidia 40-56 x 12-17 µ
2. Pleurocystidia 55-80 x 8-15 µ

Stirps Condensa

Like the previous stirps, this one is questionably placed in this section because of the variation in the wall thickness of the pleurocystidia. If the thickening in the walls of some cystidia is disregarded, then the species would fall in stirps Spumosa.

1. Basidiocarps obviously lignicolous
2
1. Basidiocarps appearing terrestrial
3
2. Pileus glabrous; on oak wood; spores 5-6 µ broad
2. Pileus decorated with veil remnants; on wood of conifers; spores 4-5 µ wide
3. Spores 5-6 µ wide
3. Spores 3.5-5 µ
4
4. Pileus with a thick gelatinous pellicle; growing under Sequoia in northern California
4. Not as above
4
5. Pleurocystidia 33-50 x 9-15 µ
5. Pleurocystidia 48-72 x 7-14 µ

Stirps Decorata

This is a rather distinct group of dingy reddish species found in our western states on rotting conifer wood or on conifer duff.

1. Pleurocystidia with walls 1.3-3 µ thick
2
1. Pleurocystidia rarely with walls over 1.5 µ thick
3
2. Odor fragrant; caulocystidia present; pleurocystidia 45-75 x 9-17 µ
2. Odor lacking; caulocystidia lacking (?); pleurocystidia 60-115 x 10-17 µ
3. Veil glutinous and leaving an evanescent glutinous annulus
3. Veil floccose (dry) and fibrillose
4
4. Odor not distinctive; stipe below the veil-line with brownish squamules at first, in age the squamules mostly whitish
4. Odor fragrant; veil remnants pallid to yellow below the veil-line

Stirps Ferruginea

As indicated, the species with bright rusty red, orange, and variations of these colors over the disc of the young pileus are placed here. The colors may fade and then the species may resemble those of stirps Occidentalis. If only old material is available for identification try both stirpes to obtain an identification. This stirps might be further divided on the color of the gills at maturity: one group with dull colored gills and the other with yellow-brown gills. Presumably there would be a difference in the color of the spore print but this remains to be checked.

1. Spores mostly 6-7 x 4-5µ; gills with wood brown to fuscous tones when mature and fresh
2
1. Spores usually narrower in relation to length; gills yellow brown
4
2. Odor strong (of iodiform or Inocybe-like); pileus "burnt sienna" fresh
2. Odor not distinctive; pileus ferruginous at first
3
3. Stipe readily staining yellow
3. Stipe not staining as above
4. Pleurocystidia with walls 1.5-4 µ thick
4. Pleurocystidia with walls thinner (0.5-2 µ)

Stirps Occidentalis

Species with cinnamon, tan, russet to clay-colored pilei are placed here. Some in other stirpes fade out to the above colors hence fresh young or barely mature pilei are needed for correct disposition of a collection.

1. Stipe 10-20 mm thick
2
1. Stipe thinner, usually 3-8 mm diam
3
2. Pileus glabrous and glutinous
2. Pileus soon with spot-like or appressed squamules from the breaking up of the cuticle
3. Pileus at maturity rivulose
3. Pileus not as above
4
4. Pileus evenly dull cinnamon; typically on hardwood—Alnus and Acer
4. Pileus paler on margin; on conifer debris
5
5. Spores 7-9 x 4-5 µ
see stirps Condensa
5. Spores smaller
6
6. Pleurocystidia with walls up to 0.5 µ thick
6. Pleurocystidia with walls about 2 µ thick

Section Carbonicola sect. nov.

Ad terram adustam, carbones etc.; pileus viscidus; pleurocystidia conspicua. Typus P. fulvozonata.

This is an ecological grouping of species very similar morphologically. It is now thought that species with this habitat require a high concentration of minerals to induce fruiting-such as released from burned wood. The characters of the section are the habitat in addition to those defining the subgenus. Singer (1963) placed P. carbonaria Smith, which has a red veil, in his section Subflammentes but for this section described the veil as pale colored "not rusty brown, ochraceous brown nor bright yellow, red etc." We do not know his type species, P. subflammans (Speg.) Singer, but assume it does not have a colored veil. We exclude from section Subflammantes P. carbonaria Smith. Sect. Subflammantes is probably identical with Sect. Adiposae of our work. We have not studied the type.

1. Veil in young specimens red to rusty brown or cinnamon
2
1. Veil bright yellow, pallid or absent
4
2. Veil ferruginous red
2. Veil russet to cinnamon
3
3. Taste mild; veil russet
3. Taste very disagreeable; veil pale cinnamon
4. Lamellae ochre yellow; pileus bay red both when fresh and when dried
4. Not as above
5
5. Stipe 5-10 mm thick
6
5. Stipe 1.5-4 (5) mm thick; veil pallid at first
6. Pileus pale buff young; veil orange brown on aging
6. Pileus dark yellow-brown; veil lemon yellow young

Section Spumosae sect. nov.

Pileus viscidus; pleurocystidia fusoide ventricosa; sporae 7-11 x 4-6 . Typus: Pholiota spumosa.

Key to Stirpes*

*If any pleurocystidia show some wall thickening up to about 0.5 µ try section Flammuloides also.

1. Growing on Sphagnum or clumps of moss in bogs
1. Habitat various but not as above
2
2. Odor pungent to fragrant; lamellae pale yellow young; many pleurocystidia with orange red content revived in KOH
Stirps Graveolens
2. Not as above
3
3. Stipe 1-3.5 mm thick; pileus 1-3.5 cm wide
Stirps Scamba
3. Pileus wider and stipe thicker
4
4. Spore deposit testaceous; taste bitterish
Stirps Subamara
4. Not as above
5
5. Pileus appressed-squamulose at least around the disc (see P. paludosella also)
Stirps Stratosa
5. Pileus glabrous or merely virgate
Stirps Spumosa

Stirps Sphagnicola

1. Spores 8-10 x 4.5-6 µ
2
1. Spores 7-9 x 4-4.5 µ
3
2. Gills whitish when young; pileus with reddish or reddish-brown spotted center; caulocystidia absent
2. Gills very soon yellowish; pileus disc tawny to dark yellow-brown; caulocystidia present
3. Pileus tinged flesh color when young and moist, pallid when faded
3. Pileus orange-fulvous over disc, greenish yellow over marginal area

 

Stirps Graveolens

1. Only one species

Stirps Scamba

The slender stipe and relatively wide spores along with the short pleurocystidia of many species characterize the group. See P. subtestacea also.

1. Spore wall 0.5 µ (or slightly more) thick
1. Spore wall 0.25 µ, thick approximately
2
2. Veil tawny as judged by flecks on pileus and zones on the stipe
2. Veil not as above
3
3. Pileus chrome yellow
3. Pileus not colored as above
4
4. Pileus dark cinnamon to vinaceous brown or at least reddish tawny on the disc
5
4. Pileus essentially pale yellow to pale buff or pallid young
6
5. Disc of pileus dark vinaceous brown to vinaceous drab
5. Disc of pileus reddish tawny (see P. condensa also)
6. Taste bitterish; spores 7-9 x 4.5-5.5 µ
6. Taste mild; spores mostly slightly larger
7
7. Pileus slimy viscid; veil rudimentary
7. Pileus merely viscid; veil fairly well developed
8
8. Spores 9-11 x 5-6 µ; subhymenium not gelatinous
8. Spores 7-9 (10) x 4.5-5.5 µ; subhymenium gelatinous

Stirps Subamara

1. Only one species

Stirps Stratosa

Pholiota bakerensis will key out here if no thick-walled cystidia are observed.

1. Only one species

Stirps Spumosa

1. Pileus pellicle gelatinous but thin and easily obliterated. Veil very thin; spores with apical pore distinct (some spores obscurely truncate)
1. Not as above
2
2. Pileus dingy yellow to olive-yellow (finally) on margin and tawny to dingy yellow-brown on disc
2. Pileus bright yellow on margin and disc bay to orange or red
3
3. Stipe 2-4 mm thick; veil fibrillose and copious, yellowish becoming pallid
3. Stipe 4-15 mm; veil thin
4
4. Caulocystidia absent
4. Caulocystidia present at stipe apex (see P. subflavida also)

Section Lubricae (Fr.) Singer, emended

Lilloa 22: 516. 1951.

Pleurocystidia prominent but not of the type generally known as chrysocystidia; spores 5-8 (9) x 3-4.5 (5) µ; pileus glutinous to viscid. Type: P. lubrica.

Key to Stirpes

1. Taste bitter
1. Taste mild to farinaceous
2
2. Pileus color in bright ferruginous-red, orange-red or brilliant orange-yellow range
2. Pileus color darker or duller at least over disc
3
3. Pileus cinnamon to dark cinnamon brown, subtestaceous or bay-brown; marginal area not yellow (may be pallid to tan, etc.)
Stirps Lubrica
3. Pileus variously colored but at least the pileus margin some shade of yellow varying to pallid (old or faded specimens of stirps Fibrillosipes will key here also)
4
4. Pileus muticolored-mixed green, violaceous, purple-drab and yellow at different stages
Stirps Polychroa
4. Not as above
5
5. Typically on decaying wood; spores 3.5-4.5 µ (see stirps Lenta also)
5. Typically near decaying wood or on very decayed wood
6
6. Spores 2.8-3.5 (4) µ broad
Stirps Innocua
6. Spores 4-4.5 (5) µ broad
Stirps Lenta

Stirps Astragalina

The bitter taste is the central feature. We recognize one species in the group at present. Flammula amara Fr. sensu Kauffman probably belongs here but we finally decided to exclude it pending further study.

1. Only one species

Stirps Fibrillosipes

This group contains the non-bitter bright red, orange-red to orange or brilliant yellow species.

1. Stipe 6-12(15) mm thick (see P. spinulifera also)
2
1. Stipe 3-6(8) mm thick
4
2. Veil with an outer gelatinous layer
2. Veil dry and fibrillose
3
3. Veil remnants on pileus pallid tan to yellowish; spores ochraceous tawny in KOH
3. Veil remnants whitish; spores in KOH merely pale yellowish
4. Growing on soil; lamellae narrow
4. Not with above combination of features
5
5. Some fibrils on the stipe brilliant orange
5. Not as above
6
6. Spores in KOH pale rusty cinnamon
6. Spores merely yellowish in KOH

Stirps Lubrica

We group here the species without distinct yellow tones over the margin of the pileus. This is in accord with the Friesian descriptions but not with the illustration of Fries in the Icones.

1. Stipe 1-2.5 mm thick
2
1. Stipe (3) 4-12 mm or more thick
3
2. Pileus testaceous
2. Pileus dull cinnamon color
3. Caespitose under larch; spores 7-9 x 3.5-4 µ
3. Not as above
4
4. Young lamellae avellaneous; veil yellow, copious
4. Not as above
5
5. Pileus virgate with dark brown streaks; stipe rooting and with much sand adhering (as in Laccaria trullisata)
5. Not as above
6
6. Lamellae almost blackish brown when mature; pileus reddish cinnamon; on the ground under oak
6. Not as above
7
7. Lamellae yellow when young
8
7. Lamellae not yellow at first
9
8. Veil gelatinous; stipe viscid when young
8. Veil dry
9. Lamellae pallid when young (see P. fulvozonata and P. bakerensis also )
9. Lamellae dull brown when young

Stirps Polychroa

1. Only one species

Stirps Crassipedes

The distinctly lignicolous species not included in any previous stirps are placed here.

1. Pileus clay-color to pinkish-buff
2
1. Pileus distinctly yellow at least on the margin
3
2. Pileus context slowly staining cinnamon-buff when cut; pleurocystidia 60-104 µ long; subhymenium thin and cellular (not gelatinous)
2. Pileus context white and unchanging; pleurocystidia 40-80 µ long
3. Pileus spotting green when handled
3. Not as above
4
4. Pileus dark yellow brown but drying bay-red; lamellae sulphur yellow w hen young
4. Not as above
5
5. Pleurocystidia 40-56 x 12-17 µ; spores 5-6 x 3-3.5 µ
5. Pleurocystidia 38-73 x 7-21 µ; spores 5.5-7 x 3.5-4.5 µ

 

Stirps Innocua

Terrestrial or appearing terrestrial and the spore width 2.8-3.5 (4) µ are the two features in combination which delimit this group. See P. perniciosa also; it has long pleurocystidia but actually they are chrysocystidia.

1. Lamellae pallid but soon becoming pale greenish yellow
1. Not as above
2
2. Some spores with a dark violet-brown coagulated content as revived in Melzer's; pleurocystidia 38-52 x 8-15 µ
2. Not as above
3
3. Stipe 3-5 mm thick; pileus broadly conic
3. Stipe (4) 5-10 mm thick
4
4. Pileus margin slightly greenish when bruised; pileus context white
4. Not as above
5
5. Pleurocystidia 30-46 x 10-15 µ, utriform caulocystidia not distinctive      
5. Pleurocystidia 50-70 x 11-15 µ; fusoid-ventricose; caulocystidia conspicuous and versiform (fig. 459)

Stirps Lenta

Terrestrial species or basidiocarps appearing to be terrestrial (not located directly on wood such as logs, stumps, or branch material above ground): spores 4-5 µ wide; colors various but not as in stirpes Lubrica, Fibrillosipes, or Polychroa.

1. Pileus conic, in age with a conic umbo 
2
1. Pileus convex to plane, rarely with a low umbo
3
2. Lamellae ferruginous at maturity
2. Lamellae clay-color at maturity
3. Pileus pallid to grayish and developing yellow tones in age
3. Pileus yellow (at least on margin when young)
4
4. Lamellae pallid when young (see P. lubrica also)
4. Lamellae yellow at first