The North American Species of Pholiota

154. Pholiota highlandensis (Pk.) comb. nov.

Agaricus carbonarius Fries, Obs. Myc. 2: 33. 1818.
Flammula carbonaria (Fr.) Kummer, Der Führer in die Pilzkunde, p. 82. 1871.
Flammula highlandensis (Pk.) Peck, New York State Mus. Ann. Rept. 50: 138. 1897.
Gymnopilus carbonarius (Fr.) Murrill, Mycologia 4: 256. 1912.
Pholiota carbonaria (Fr.) Singer, Agaricales, p. 517, 1951.
Dryophila carbonaria (Fr.) Quélet, Enchir. Fung., p. 70. 1886.

Illustrations: Text figs. 330, 332-335; pls. 67a, 70b, 72.

Pileus 2-4 (6) cm broad, viscid, convex, becoming somewhat depressed, at times with a low umbo, fulvous or reddish-cinnamon-brown to reddish brown, the margin usually paler, colors varying "russet," "tawny," "pecan brown," "verona brown," or in age almost "mummy brown," glabrous except for deciduous veil-remnants along the margin, hygrophanous and fading to various shades around "ochraceous-buff," margin even. Context rather thin, yellow to subconcolorous; odor not distinctive, taste slightly disagreeable or none.

Lamellae adnate or rounded-adnate, pallid to pale-yellowish when young, becoming "snuff-brown" or "cinnamon-brown," broad, close, edges even or eroded.

Stipe (1) 2-4 cm long, (2) 3-6 mm thick, apex whitish to yellowish at first, becoming dingy brownish, lower portion pallid then dark brown (darker than apex), with zones or patches of the veil which is pallid yellow to "cinnamon-buff" (pale buff) and forms an evanescent, fibrillose annular zone, more or less glabrescent.

Spores "cinnamon-brown" in deposit (American material); 6-8 x 4-4.5 µ, smooth, apical pore distinct, mostly elliptic but varying to ovate in face view, in profile subelliptic to obscurely inequilateral, wall about 0.3 µ thick, dark rusty brown (near cinnamon brown) in KOH, paler and more ochraceous in Melzer's reagent.

Basidia 25-32 x 7-8 µ, subcylindric to clavate, 4-spored, yellowish to hyaline in KOH, yellowish in Melzer's sol. (in American material often 18-24 x 5-6 µ). Pleurocystidia 38-65 (70) x 7-15 µ, fusoid-ventricose, apex obtuse, smooth, thin-walled, content often ochraceous in KOH. Cheilocystidia like pleurocystidia but smaller (30-50 x 7-12 µ) but varying to subfusoid to nearly clavate, content yellow to hyaline. Caulocystidia 45-88 x 7-12 (16) µ cylindric, clavate or fusoid-ventricose.

Gill trama of a central area of subparallel floccose hyphae, cells short to long, walls thin smooth and yellowish to hyaline in KOH; subhymenium a well defined layer of hyaline branched narrow (± 2µ) smooth hyphae. Pileus cutis a thick gelatinous pellicle of yellow narrow (2-3 µ) hyphae many appearing to have frequent refractive transverse septa; hypodermium of floccose distinctly encrusted tawny hyphae 4-12 µ in diam. Context hyphae compactly interwoven, having inflated cells and thin hyaline to yellow walls. All hyphae inamyloid. Clamp connections present.

Habit, Habitat, and Distribution: On burned-over soil, or on charred wood, New York, Maryland, Tennessee, Florida, Texas, Michigan, Colorado, Wyoming, Idaho, Washington, Oregon and California: reported from Jamaica and from Europe, spring-fall, winter (south).

Observations: This species grows on burned areas and varies somewhat in its characteristics. The pileus is brown, usually some shade of reddish brown, with the margin paler. One collection (Smith 65054, from Idaho) is obviously strongly faded, doubtless from exposure to sun and wind. The pileus flesh is pallid or yellowish. Peck (1897) described Agaricus (Flammula) highlandensis as differing from Agaricus (Flammula) carbonarius in its whitish flesh; but, in his description, he admits that the flesh is sometimes tinged with yellow under the cuticle. In P. highlandensis the lamellae are broad, in contrast to the narrow gills of another carbon-loving species, P. carbonaria Smith. The veil in P. highlandensis is pallid to yellowish or buff, becoming cinnamon brownish when sprinkled with spores; in P. carbonaria Smith the veil is cinnabar red to ferruginous, fading to ochraceous. In P. highlandensis the veil may be rudimentary or if better developed leaves on the stipe remnants which appear as scattered fibrillose remains or at times as thin yellowish patches (Smith 19894, from Oregon). The microscopic characters, although rather constant, show occasional variations. The spores usually are up to 7.5 µ long, but in Smith 51513, from Colorado, the upper limit is 8 µ, as it was in a collection of Bresadola's. In nearly all collections studied the pleurocystidia have a slender simple neck; but in Smith 17717, from Washington, some were forked and branched. The structure of the subhymenium at once distinguishes P. harenosa from this species.

A study of the "Highland" (near Highland Falls, N. Y.) collection which must be accepted as the type gave the following microscopic data.

Spores 6.5-8 x 4-4.5 µ, ellipsoid to ovoid in face view, inequilateral in profile, smooth, germ pore not evident. Basidia 23-27 x 5-6 µ, 4-spored. Pleurocystidia 47-72 x 7-12 µ, ventricose, at times slightly constricted, sometimes brown; cheilocystidia 33-44 x 6-10 µ, ventricose or subfusoid. Gill trama a mediostrate, hyphae slightly interwoven, hyphae 5-8 µ broad. Subhymenium a gelatinous zone. Pileus trama radial. Cuticle gelatinous, of loosely interwoven, pale brown hyphae, sometimes lightly incrusted. Clamp connections present. Pileus and gill trama yellowish brown or pale rusty in KOH; rusty brown in Melzer's reagent. Caulocystidia none.

Kauffman however had studied a specimen of P. prolixa under this name and Peck himself had sent Kauffman such a specimen. We designate those specimens having pleurocystidia 47-72 x 7-12 µ as the type since they unquestionably came from burnt ground.

Material Examined: ALABAMA: Burke 680; Hesler 22978; CALIFORNIA: Cooke 18160; Copeland 455; Rea H-887 (MICH); COLORADO: Smith 51513; FLORIDA: Hesler 16152, 21600; IDAHO: Slipp UIFPH 1545, 3105 (both MICH); Smith 44665, 44803, 54354, 54446, 58605, 59449, 64933, 65019, 65054; LOUISIANA: Hesler 3958; MICHIGAN: Bartelli 2418; Harding 350, 390, Smith 32-10, 32-221, 32-221a, 32-513, 33-99, 1197, 4558, 6120, 33289, 33695, 71441; MARYLAND: Kauffman 6-16-19; NEW MEXICO: Barrows July 1955; NEW YORK: The type; Smith 3617, 7929, 19891, 19894, 55353; TENNESSEE: Hesler 4259, 4260, 10128, 18845, 21329, Smith 21647 (MICH); TEXAS: Thiers 1488; WASHINGTON: Smith 3063, 17717, (MICH) 14145; WYOMING: Smith 34357, 34434; ENGLAND: Austwick 403; GERMANY: Syd. Mycoth. Ger. 1411, 1812; SWEDEN: Fung. Suec. 20-X-1945; 5-V-1946.