1. Some portion of the basidiomata turning green with alkali . . . . 2
Gymnopus section Levipedes
Gymnopus section Levipedes (Fr.) Halling, Brittonia 48: 487. 1996 (1997).
- Subtribus Laevipedes Fries, Epic. Syst. Mycol. 90. 1838.
Collybia section Levipedes Quélet, Mém. Soc. Émul. Montbéliard, sér. II, 5: 96. 1872.
Marasmius subtribus Tergini Fries, Epic. Syst. Mycol. 376. 1838.
Marasmius subsection Tergini (Fr.) Quélet, Mém. Soc. Émul. Montbéliard, sér. II, 5: 220. 1872.
- Basidiomata marcescent to fibrous putrescent. Pileipellis a layer of repent, branched hyphae, often bifurcate, not coralloid or diverticulate, not radially arranged; cells frequently short and broad, more rarely cylindric, narrow and forming a trichodermium.
- Type species: Gymnopus dryophilus (Bull.:Fr.) Murr.
Key to Species of Section Levipedes
2. Hyphae of the lamellar trama without coarse pigment granules . . . . 3
3. Hyphae of the stipe trama without dark pigment encrustions in water mounts or Melzer's; stipe glabrous and dark red brown OR tawny and strigose overall . . . . 4
4. Stipe glabrous and dark red brown . . . . G. erythropus
5. Growing on leaf litter, rotten wood, etc, fruiting season variable . . . . 6
6. Stipe base usually glabrous but occasionally thinly, white pruinose with age; pileus dark reddish brown to honey yellow when fresh; texture somewhat fleshy fibrous . . . . 7
7. Rhizomorphs white; basidiomata dark reddish brown . . . . 8
8. Cheilocystidia filamentous; basidiomata on litter, rotted wood; lamellae crowded . . . . G. dryophilus
Gymnopus alkalivirens (Singer) Halling, Mycotaxon 63: 363. 1997. Micro features
Misapplied name: Collybia plexipes sensu Kauffman, Agar. Michigan 1: 762. 1918, non Fries, Syst. Mycol. 1: 146. 1821.
Spores white in deposit, 5.4-6.5 x 2.2-3.2 µm, lacrymoid, narrowly ellipsoid in profile, ellipsoid to subcylindric in face or back view, slightly wrinkled in water mounts, smooth and pale greenish in alkali, inamyloid, acyanophilous. Basidia 18.4-27 x 5.4-6.5 µm, clavate to subclavate, four sterigmate, greenish in alkali, not siderophilous. Pleurocystidia absent. Cheilocystidia scattered, sometimes inconspicuous, 19.5-72.5 µm long, flexuous cylindric to clavate with a few broad obtuse projections, green in alkali. Lamellar trama subparallel to slightly interwoven, inamyloid; cells 4.2-9.8 µm in diam, occasionally inflated to 16.2 µm, with conspicuous dark brown, granular encrusting pigment in water and Melzer's mounts; pigment soluble in alkali and the tissues discoloring to greenish. Pileus trama loosely interwoven, inamyloid; hyphae 4.2-8.6 µm in diam, with encrusting pigment similar to lamellar trama. Pileipellis a layer of repent, branched hyphae, not diverticulate or coralloid, not radially arranged; hyphae 3.2-5.4 µm in diam, coarsely encrusted with a brown plate-like pigment which is partially soluble in alkali. Stipitipellis a layer of parallel, vertically oriented hyphae; cells 3.2-5.4 µm in diam, encrusted with alkali soluble brown pigment, giving rise to flexuous-cylindric to contorted caulocystidia, becoming greenish in alkali. Stipe tramal hyphae hyaline, encrusted with scattered, brown granular pigment, soluble in alkali. Clamp connections present.
Macrochemical reactions: KOH and NH4OH--all parts green.
Habit, habitat, and distribution: Solitary, scattered, gregarious, or cespitose on soil, among moss, leaf litter, decaying wood debris, or old fern hummocks. Occurring in hardwood forests during June and July in New England, Quebec, Michigan, New York, and Virginia. Also reported from New Jersey occurring in late May (Lincoff, personal communication). Also reported from Mexico by Guzmán et al. (1992).
Discussion: The above description is derived from one previously published (Halling 1979). Since then, additional material has been examined and observations on related taxa have been made (Baroni and Halling, 1989; Halling 1981, 1990, 1996) that include differentiating G. fuscopurpureus, C. kirchneri (G. erythropus), C. obscura, G. semihirtipes, and G. spongiosus.
Gymnopus semihirtipes (Pk.) Halling, Mycotaxon 63: 365. 1997. Micro features Macro image
Collybia semihirtipes (Pk.) Halling, Mycologia 73: 637. 1981.
Spore deposit cream colored. Spores 7.5-8.6(-9.8) x 3.2-4.2(-5) µm, lacrymoid in profile, obovoid in face or back view, smooth, inamyloid, acyanophilous. Basidia 19.5-32.4 x 5.4-7.5 µm, clavate, four sterigmate, not siderophilous. Hymenial cystidia absent. Lamellar trama interwoven, inamyloid; hyphae 3.2-7.5 µm in diam, smooth, thin walled, not encrusted in water or Melzer's mounts. Pileus trama interwoven, inamyloid; hyphae smooth, thin walled, not encrusted in water or Melzer's mounts. Pileipellis a layer of repent, branched hyphae, not diverticulate or coralloid, not radially arranged; hyphae 4.2-10.8 µm in diam, with scattered, yellow brown encrusting pigment. Stipitipellis a layer of parallel, vertically oriented hyphae; cells 3.2-5.4 µm in diam, with dark brown pigmented walls in water or Melzer's mounts, becoming greenish in alkali; giving rise to a scant vesture of scattered caulocystidia, cylindric contorted, more abundant toward the base of the stipe, walls often thickened up to 2.5 µm. Stipe tramal hyphae 4.2-11.8 µm in diam, usually hyaline with dark brown granular encrustations in water or Melzer's mounts, dissolving in alkali and tissues becoming greenish. Clamp connections present in all tissues.
Habit, habitat, and distribution: Scattered to gregarious on humus, among fallen leaves. Collected in May, June, and October.
Discussion: Gymnopus semihirtipes seems to be intermediate between G. spongiosus and G. alkalivirens. Only the stipe trama of G. semihirtipes has the dark brown, alkali soluble, granular encrusting pigment which is so characteristic of all the tramal hyphae of G. alkalivirens. Gymnopus spongiosus is devoid of granular pigment in the tramal tissues. Gymnopus erythropus is related, but differs in a number of features, viz., the alkaline reaction is restricted to the stipe, and pigment granules are located only on the walls of the hyphae forming the stipitipellis and the vesture at the stipe base. Also the granules do not seem to be alkali soluble. Other distinctive features include long, filamentous cheilocystidia and a stipe vesture that is restricted to the base but rarely reaches halfway up the stipe.
Gymnopus spongiosus (Berk. & Curt.) Halling, Brittonia 48: 489. 1996 (1997). Micro features, Macro image
Collybia spongiosa (Berk. & Curt.) Singer, Lilloa 22: 201. 1949 (1951).
Spore deposit white. Spores 6.2-8.4 x 3.5-4.2 µm, lacrymoid to ellipsoid in profile, obovoid in face or back view, smooth, inamyloid, acyanophilous. Basidia 17.5-24.5 x 6.3-9.8 µm, clavate to subclavate, four sterigmate, not siderophilous. Pleurocystidia absent. Cheilocystidia inconspicuous, often collapsed on lamellar edge, 24.5-50 µm long, cylindric to strangulated or irregularly lobed and knobbed. Lamellar trama parallel to interwoven, inamyloid; hyphae smooth, 4.2-7 µm in diam. Pileus trama interwoven, inamyloid; hyphae smooth. Pileipellis a layer of repent, branched hyphae, often bifurcate, not diverticulate or coralloid, not radially arranged; hyphae 3.5-7(-10.5) µm in diam, coarsely encrusted with a yellow brown pigment which does not dissolve in alkali. Stipitipellis a layer of parallel, vertically orient ed hyphae; cells 3.5-4.9 µm in diam, with walls containing a brown pigment, not encrusted, becoming green in alkali, giving rise to long, cylindric to contorted, branched, septate caulocystidia, with walls up to 1.4 µm thick. Stipe tramal hypahe hyaline,not encrusted. Clamp connections present in all tissues.
Macrochemical reactions: KOH and NH4OH green on stipe surface.
Habit, habitat, and distribution: Solitary, scattered, gregarious or cespitose on leaf and needle litter in deciduous, mixed or pine forests; rarely on sand. Occuring from July to October in eastern North America, and also during February and December in Louisiana and Florida.
Discussion: The most striking feature of this species is the rusty brown, tomentose stipe with an incrassate spongy base. The color contrasts markedly with that of the pileus which is soon pale. Also, the reaction of the stipe surface to alkali is an important characteristic. Other species with similar stipes that might be confused with Gymnopus spongiosus are G. biformis, G. semihirtipes, and G. pinastris. Gymnopus biformis, however, does not have alkali-soluble pigments, the pileus is much darker, and, in addition to diverticulate branchlets in the pileipellis, it has cheilocystidia which are quite conspicuous. Also, G. biformis usually grows on soil. Gymnopus pinastris, on the other hand, appears to be confined to needle litter of spruce, fir, and pine, has an alliaceous odor and lacks a green reaction to alkali. It also has extremely thick-walled caulocystidia. Gymnopus semihirtipes is easily differentiated from G. spongiosus because of the dark brown, granular pigment that encrusts the hyphal walls in the stipe trama.
Gymnopus erythropus (Pers.:Fr.) Antonín, et al., Mycotaxon 63: 364. 1997.
Marasmius erythropus (Pers.:Fr.) Quél., Mém. Soc. Émul. Montbéliard, sér. 2, 5: 221. 1872.
Collybia erythropus (Pers.:Fr.) Kummer, Führ. Pilzk. 115. 1871.
Collybia kuehneriana Singer, Persoonia 2: 24. 1961.
Collybia marasmioides (Britzelm.) Bresinsky & Stangl in Stangl & Bresinsky, Z. Pilzk. 35: 67. 1969.
Basidiospores white in deposit, 6-8.5 x 3 5-4 µm, lacrymoid in profile, ovoid to ellipsoid in face and back views, inamyloid, smooth, acyanophilous, slightly greenish in alkali. Pasidia 21-30 x 4.5-7 µm, clavate to subclavate, 4-sterigmate, not siderophilous. Pleurocystidia not differentiated. Cheilocystidia subclavate, strangulated, furcate to subclavate with a short mucro when young, becoming rather inconspicuous with age and then collapsing or sometimes with a filamentous prolongation at the apex, this latter sometimes branched, 25-40 ILm long. Lamellar trama of parallel to interwoven, inamyloid, mostly cylindric hyphae, 3.5-8.5 Lm diam., with age becoming branched and inflated and some with walls thickened to 3.5 m, lacking encrusting pigments and not green in alkali. Pileus context loosely interwoven, inamyloid, with elements 6.8-12.8 (-21) µm broad, sometimes slightly thickwalled like the lamellar trama elements. Pileipellis a layer of repent, branched hyphae, often bifurcate, not diverticulate or coralloid, not radially arranged; hyphae (3.4-)6.8-15.5(-21) µm diam., some elements finely encrusted with pigment. Stipitipellis composed of vertically oriented, thin-walled, cylindric hyphae, 2.5-5.1 µm diam., with some rusty brown, granular, intercellular pigment that becomes green, but does not dissolve, in alkali. Caulocystidia abundant, long, and tangled at the stipe base, sometimes with walls encrusted with rusty brown pigment. Stipe trama composed of hyaline, thick-walled hyphae, usually cylindric but sometimes inflated to 17 Lm in diam., not encrusted with pigment and not changing in alkali. Clamp connections present in all tissues.
Habita, habitat, & distribution: Scattered to cespitose on decaying wood. Known from both eastern and western USA.
Discussion: First reported as Collybia kirchneri by Baroni and Halling (1989) for North America, where it may be rare but widespread (known from California and New York). However, because it is related to that complex of species around Gymnopus dryophilus, and shares some macroscopic resemblance to, and habitat preference (rotten wood) with Gymnopus acervatus some earlier misidentifications are possible.
Gymnopus erythropus is easily distinguished from G. dryophilus and G. acervatus by the distinctive green reaction of the stipe surface when tissues are mounted in alkali solutions and observed under the compound microscope. At least four other species in section Levipedes develop green color reactions in some part of the basidioma when subjected to alkali: G. alkalivirens, G. fuscopurpureus, G. spongiosus, and G. semihirtipes (Halling, 1979, 1981, 1983, 1990). However, in G. alkalivirens, G. fuscopurpureus, and G. semihirtipes, the dark brown intercellular pigment granules are not confined to the stipe surface (also located in at least the stipe tramal tissues), but are soluble and turn greenish in alkaline solutions; pigment granules are lacking in G. spongiosus. In G. erythropus, the dark brown intercellular pigment granules are generally few, restricted to the caulocystidia and stipe surface, and do not dissolve in the presence of alkali.
Gymnopus earleae Murrill, N. Amer. Flora 9: 364. 1916. Micro features #1, Micro features #2, Macro image
Collybia earleae (Murr.) Coker & Beardslee, J. Elisha Mitchell Sci. Soc. 37: 89. 1921.
Habit, habitat, and distribution: Scattered to gregarious or cespitose on wet soil. Occurring during May, June, and July in the northeast, April in Alabama. Reported by Coker and Beardslee (1921) as appearing several times in May. Also reported from California (Vilgalys and Miller, 1983).
Discussion: Gymnopus earleae is distinguished macroscopically from small forms of G. dryophilus by the smaller size, darker colors (especially the lamellae), a habitat on soil, and its tawny, strigose stipe base. Microscopically, inflated cheilocystidia (as opposed to filamentous) are distinctive. Murrill's (1911) illustration of Collybidium dryophilum is G. earleae.
Gymnopus kauffmanii (Halling) Halling, Mycotaxon 63: 364. 1997. Micro features, Macro image
Collybia kauffmanii Halling, Mycol. Mem. 8: 1983.
Spores (5.6-)6.4-7.7 x 2.8-3.5 µm, lacrymoid to subellipsoid in profile, obovoid to ellipsoid in face or back view, smooth, inamyloid, acyanophilous. Basidia 17.5-28 x 4.8-6.2µm, clavate, four sterigmate, not siderophilous. Pleurocystidia absent. Cheilocystidia scattered, infrequent, somewhat inconspicuous, 31.5-42 µm long, clavate to irregularly cylindric with filamentous, apical prolongations. Lamellar trama parallel, inamyloid; hyphae 3.5-12 µm in diam, smooth, thin walled. Pileus trama interwoven, inamyloid; hyphae 4.2-12 µm in diam, smooth, thin walled. Pileipellis a layer of repent, branched hyphae, often bifurcate, not radially arranged; hyphae (4.2-)5-10.5(-12) µm in diam, thin walled, with brown pigment localized in the walls, breaking up to form scattered encrustations. Stipitipellis a layer of parallel, vertically oriented hyphae; cells 2.8-5(-8) µm in diam, smooth, moderately thick walled, giving rise to a dense vesture of caulocystidia which are 2.8-5.6 µm in diam, cylindric, occasionally somewhat flexuous, smooth and thin walled. Clamp connections present in all tissues.
Habit, habitat, and distribution: Scattered to gregarious on leaf litter of hardwoods. Appearing during late August and September.
Discussion: The sordid-tan pileus, a pallid, dense stipe vesture, the marcescent-pliant texture, lack of a reaction to alkali, a leaf litter subtrate, and the inconspicuous cheilocystidia differentiate Gymnopus kauffmanii from other taxa in section Levipedes.
Gymnopus dryophilus (Bull.:Fr.) Murrill, N. Amer. Flora 9: 362. 1916. Micro features #1, #2 Macro image
Marasmius dryophilus (Bull.:Fr.) Karsten, Krit. Ofvers. Finlands Basidsvamp. 103. 1889.
Collybia dryophila (Bull.:Fr.) Kummer, Führ. Pilzk. 115. 1871.
Collybidium dryophilum (Bull.) Murrill, Mycologia 3: 101. 1911.
Spore deposit white to pale yellowish white (2A2) when fresh, pale yellowish when dry. Spores (4.8-)5.6-6.4(-7) x 2.8-3.5 µm, lacrymoid to ellipsoid in profile, ellipsoid to obovoid in face or back view, smooth, inamyloid, acyanophilous. Basidia 14-18 x 5.6-7 µm, clavate to subclavate, four sterigmate, not siderophilous. Pleurocystidia absent. Cheilocystidia 15.4-49 µm long, scattered to abundant, often collapsing along lamellar edge in older basidiomata, clavate contorted to diverticulate or irregularly lobed, sometimes furcate. Lamellar trama parallel, inamyloid; hyphae 4.2-10.5 µm in diam, smooth, thin walled, at times slightly inflated. Pileus trama interwoven, inamyloid; hyphae 4.9-11.2 µm in diam, frequently branched, cylindric, smooth, thin walled. Pileipellis a layer of repent, branched hyphae, often bifurcate, not diverticulate or coralloid, not radially arranged; hyphae 4.2-10.5(-14) µm in diam, with scattered, brown encrusting pigment, thin walled. Stipitipellis a layer of parallel, vertically oriented hyphae; cells 2.2-5 µm in diam, smooth, thin walled, rarely giving rise to cylindric caulocystidia, up to 7 µm in diam. Clamp connections present in all tissues.
All parts of the basidiomata occasionally parasitized by species of Syzygospora
Macrochemical reactions: Guaiac--pileus context slowly green.
Habit, habitat, and distribution: Scattered, gregarious or sometimes cespitose on humus or well decayed wood in conifer-hardwood forests. Appearing from June through September. Widely distributed.
Discussion: Gymnopus dryophilus is perhaps the most commonly collected and most widely distributed species in the genus. Because of its widespread occurrence, G. dryophilus is often illustrated in mushroom field guides (Smith 1949, 1963, Miller 1972, Groves 1975) and more often than not is described as edible (Kauffman 1918, Smith 1949, 1963, Kühner and Romagnesi 1953, Groves 1975). In these works, where a technical description is provided, a comparison of G. dryophilus to R. butyracea is invariably made and often there is a suggestion that these two taxa intergrade with one another. This confusion may be due, in part, to the fact that these species often are growing at the same time and in similar habitats. Moreover, a large number of apparently closely related species, varieties, and forms of both species has been described which seem to be, for the most part, imperceptibly distinct. However, Vilgalys and Miller (1983, 1987) have presented convincing evidence that there are several taxa reminiscent of G. dryophilus that deserve recognition. Originally, these included C. brunneola Vilgalys and Miller and C. subsulphurea Peck in North America and C. aquosa (Bull.:Fr.) Kummer, C. alpina Vilgalys and Milller, and C. ocior (Pers.) Vilgalys and Miller in Europe. Since then, Vilgalys (1991) has suggested subspecific status for C. brunneola (under C. ocior) and C. subsulphurea (under C. alpina) based on population genetics and DNA hybridization. Excellent descriptions can be found in Vilgalys and Miller (1983).