Micromphale sect. Perforantia Singer. 1948. Sydowia 2: 32.

1) Pileipellis, pileus and lamellar tramae and stipe medullary hyphae embedded in slime matrix; 2) pileipellis either: a) a layer of repent, encrusted hyphae, conspicuously clamped, usually with thickened walls (through gelatinization) and embedded in a slime matrix; or b) well-developed Rameales-structure; 3) cheilocystidia often absent, when present either clavate to utriform or siccus-type broom cell-like; 4) stipes usually less than 1 mm broad but often up to 40 mm long; ratio of pileus breadth to stipe length often large (see G. pinophilus); 5) stipe usually vestured (but see G. glabrosipes), usually appearing minutely barbed (30×); 6) lower stipe of dark, somber color, from dark sooty brown to virtually black; 6) rhizomorphs almost always present, always black, from <2–40 × 0.2–0.7 mm, branched or unbranched; 7) basidiospores without significant dimension differences; 8) species apparently more or less host-specific, fruiting on dead conifer needles or rotting deciduous leafy debris; 9) pleurocystidia consistently present, fusiform, varying somewhat in distal morphology; 10) clamp connections ubiquitous.

Antonín and Noordeloos (1997) recognized Gymnopus sect. Vestipedes subsect. Impudicae, including G. perforans, based, it would seem on the fetid odor of basidiomata of the included taxa. Later (Antonín and Noordeloos 2010), subsect. Impudicae was accepted at section rank, but basically included the same taxa as before. Wilson and Desjardin (2005) indicated that G. perforans formed its own clade sister to that of G. (Micromphale) foetidum and other mainstream odious Gymnopus taxa. Based on the present study, the original Micromphale sect. Perforantia is found to be more closely related to the Gymnopus (Marasmius) androsaceus complex, not inclusive of G. foetidum and other Gymnopus sect. Impudicae.

Molecular phylogenetic information is available for only few taxa accepted (or proposed) here. For “morpho-taxa,” some outline employed to justify their description in this paper is owed the reader. Unfortunately, almost every morphological character includes a caveat for collections which violate one or more characters. The following diagnoses are offered:

(typus auto.): Agaricus (Gymnopus) perforans Hoffm. ex Fr.

Separating characters as follows: 1) pileipellis a layer of repent, encrusted hyphae, conspicuously clamped, usually with thickened (through gelatinization) walls and usually embedded in a slime matrix; 2) cheilocystidia often absent, when present, clavate to utriform; 3) stipe usually vestured; 4) fruiting on conifer needles or broad-leafed debris.

Gymnopus pinophilus R.H. Petersen (see below)

Separating characters as follows: 1) pileipellis a well-developed Rameales structure; 2) cheilocystidia siccus-type broom cells; 3) stipes glabrous-shining; 4) fruiting on needles of Pinus; 5) discrete clade well-separated from subsect. Perforantia.

United States, Washington, Jefferson Co., Olympic National Park, Tunnel Creek, N47°46'44.19", W123°03'25.83", VIII.1975, coll F. VanDeBogart (as Ma. androsaceus), FVDB 3621 (WTU-F-51955).

Bulliformis = bubble-like, referring to the appearance of cheilocystidia at lamellar edge.

1) Morpho-species (no DNA sequences available); 2) stipe vesture a loose thatch of tortuous, often-branched, thick-walled caulocystidia; 2) cheilocystidia bulbous, smooth, with homogenous contents; 3) basidial sterigmata unusually long and slender; 4) pleurocystidia sharply fusiform, with poorly-defined content partitioning; 5) pileipellis involved in slime matrix.

The description which follows is compiled from examination of dried material. Colors when fresh may vary considerably from those below.

Basidiomata (Fig. 1A, B) diminutive, robust. Pileus 3–14 mm broad, convex, usually somewhat flattened over disc and occasionally with suggestion of small umbo, matt, sulcate near margin but not striate; disc near “Verona brown” 6E5; limb and margin near “sayal brown” 6C5 or “cinnamon buff” 6B4; context thin, fragile. Lamellae adnate to adnexed, not pseudocollariate, thin, becoming ventricose by maturity (–2.5 mm broad) and strongly crisped (dry), subdistant to distant with no anastomoses or interveining, total lamellae = 25–38, through lamellae = 13–18, (fresh) near “ochraceous buff” 5A5 to “chamois” 4B4, (dry) between “ecru drab” 16B2, “olive ochre” 2B8 and “deep olive buff” 3C3, suffusing to brown (“sayal brown” 6C5) from edge in age; lamellulae in 1–2 tiers, often rudimentary. Stipe 16–35 × 0.5–0.8(–1) mm, probably terete when fresh, equal, occasionally abruptly changing directions as though adventitious from being broken, subinsititious, pilose-vestured from apex to base, lightly so upward and there with vesture pallid, downward delicately barbed and more so to felty at base, producing a matted–furry appearance, irregularly elongate–maculate with blond vesture and fuscous pileus surface (dried), but somewhat more prominently vestured upward, upward “clay color” 5C6 or “cinnamon” 6B5, downward “snuff brown” 5E8 to “Verona brown” 6E5 at base, at junction with lamellae dark brown and leaving delicate brown ring in pileus trama. Rhizomorphs undetected. Taste and odor not recorded.

Figure 1. 

Gymnopus bulliformis. A, B Basidiomata C, D Basidiospores. Standard bars: A, B = 20 mm; C, D = 5 µm. A, CWTU-F-9305; B, DWTU-F-51955.

Temperate rainforest; substrate uncertain, certainly dead conifer needles, perhaps Tsuga or Abies; mid-summer.

Pileipellis composed of three elements, all involved with a heterogeneous slime matrix (Fig. 3): 1) pileal hairs (Fig. 2) –120 × 3.5–7 µm, cylindrical with slightly enlarged apex, thin- to firm-walled, occasionally internally clamped; 2) repent hyphae 3–9 µm diam, thin- to firm-walled (wall gelatinizing to 1.5 µm thick; Fig. 3C, D), weakly encrusted (crusting material flake-like, often riding on a thin slime sheath, detersile in slime matrix), conspicuously clamped; and 3) significant hyphal segments with numerous secondary septa (Fig. 3A, B). Pileus trama loosely interwoven; hyphae 3–9 µm diam, gelatinizing and with thin slime sheath, conspicuously clamped. Lamellar trama identical. Pleurocystidia (Fig. 4) 22–34 x 7–9 µm, fusiform, unusually sharply pointed, often somewhat swollen in mi-shaft, conspicuously clamped; contents partitioned, but ill-defined. Basidia (Figs 5, 7A) 24–30 × 7–9 µm, broadly clavate, conspicuously clamped, 4-sterigmate; contents heterogeneous with inclusions appearing oily (PhC); sterigmata –8 × 1 µm, awl-shaped. Basidiospores (Fig. 1C, D) (6.5–)7–7.5(–8) × 3.5–4(–4.5) µm (Q = 1.73–2.14; Q^m = 1.88; L^m = 7.20 µm), ellipsoid, subgymnopoid (slightly tapered proximally), smooth, thin-walled, inamyloid. Cheilocystidia (Figs 6, 7B–G) locally common, 15–26 × 9–15 µm, broadly clavate to subglobose, thin- to firm-walled, obscurely clamped, occasionally with one or two vermiform apical outgrowths (usually without); contents homogeneous, appearing empty. Stipe apex medullary hyphae strictly parallel, 5–9 µm diam, thin- to firm-walled but soon gelatinizing (in KOH) so walls become almost indistinguishable, often encrusted with crust material riding on outer surface of slime sheath, obscurely clamped. Stipe apex cortical hyphae 3–6 µm diam, long-celled, conspicuously clamped, thick-walled (wall often obscuring cell lumen), producing a layer of inflated, thick-walled cells and caulocystidia as side branches; caulocystidia from upper stipe (Fig. 8) 10–75 × 4–8 µm, irregularly cylindrical, usually with basal clamp, thick-walled (wall –2 µm thick, with cell lumen irregularly defined), subhyaline. Mid-stipe medullary hyphae strictly parallel, 3.5–12 µm diam, thick-walled (wall –2 µm thick), obscurely clamped, involved in a slime matrix but not with gelatinizing walls; mid-stipe cortical hyphae as above, producing a thatch of tortuous, thick-walled side branches resembling stipe-apex caulocystidia, from which extended caulocystidia (Fig. 9F) arise; caulocystidia –125 × 4–8 µm diam, thick-walled, cylindrical, refringent (PhC), usually oriented parallel to stipe surface.

Figure 2. 

Gymnopus bulliformis. Pileal hairs. Standard bars = 10 µm. WTU-F-51955.

Figure 3. 

Gymnopus bulliformis. Pileipellis elements. A Heterogeneous slime with secondarily septate hyphae B Secondary septa C, D Hyphae with gelatinized walls. Note clamp connections in D right. Standard bars = 10 µm. WTU-F-51955.

Figure 4. 

Gymnopus bulliformis. Pleurocystidia. Note acute apices and poorly-defined partitioned contents. Standard bars = 10 µm. WTU-F-51955.

Figure 5. 

Gymnopus bulliformis. Basidiole and basidia. Note long, awl-shaped sterigmata. Standard bars = 10 µm. WTU-F-51955.

Figure 6. 

Gymnopus bulliformis. Cheilocystidia. Standard bars = 10 µm. WTU-F-51955.

Figure 7. 

Gymnopus bulliformis. A Basidium B–G Cheilocystidia. Standard bars = 10 µm. WTU-F-9305.

Figure 8. 

Gymnopus bulliformis. Caulocystidia from upper stipe. Standard bars = 10 µm. WTU-F-51955.

Figure 9. 

Gymnopus bulliformis. Caulocystidia from lower stipe. Standard bars = 10 µm. WTU-F-51955.

Proposal of another species of Gymnopus (Micromphale) section Perforantia would seem problematic, especially from a region relatively rich in such taxa, but presence and uniqueness of diagnostic characters make such a proposal relatively safe. The diagnosis (above) lists characters which separate G. (Mi.) bulliformis from other members of sect. Perforantia with diminutive basidiomata.

The temperate rain-forests of the North American Pacific coastal region of United States and Canada apparently support several taxa of the androsaceoid/perforantioid complex. Loan of selected specimens (using Mycoportal as filter) under only Ma. androsaceus and Mi. perforans revealed enough new taxa to make intensive collecting worthwhile for such organisms. This might not be surprising, considering the variety of conifer trees, on the dead needles of which such diminutive basidiomata are to be found.

Separation of taxa in Micromphale sect. Perforantia is difficult. Micromphale perforans occurs in Europe and Scandinavia through northern North America, presumably continent-wide. It may be present in northeastern and western Siberia as well, but this is beyond the scope of this paper. From northern California comes M. sequoiae which closely resembles G. bulliformis but is reported as limited to dead foliage of Sequoia sempervirens. Both taxa are reported as lacking (or rare) cheilocystidia, neither taxon exhibits much differentiation in the pileipellis, and caulocystidia are quite similar, although those of M. perforans are reported as exhibiting some denser, more refringent (PhC) pigment at caulocystidial tips. Ordinarily, M. perforans exhibits a black, vestured stipe when mature, but Antonín and Noordeloos (2010) include dark brown shades as well, and M. sequoiae seems also to form a dark brown stipe. Desjardin (1985) used the following characters to separate M. sequoiae from M. perforans: 1) light brown to flesh-colored pileus versus pale pileus respectively; 2) mild odor versus strong, fetid odor respectively; 3) greyish orange to brown pubescent stipe versus black velutinous stipe respectively; 4) caulocystidia without concentrated apical pigment versus caulocystidia with concentrated apical pigment respectively; 5) habitat on Sequoia debris versus conifer needles, usually Picea and/or Abies respectively.

Attempts were made to obtain ITS sequences from both collections of G. bulliformis, both unsuccessful. WTU-9305 yielded sufficient sequence to identify it as a member of the clade /perforantia and blasted weakly to G. perforans in GenBank. Whether G. bulliformis deserves phylogenetic distinction remains questionable until clean sequences can be obtained from “fresh” material.

Washington, Jefferson Co., Olympic National Park, Tunnel Creek, N47°46'44.19", W123°03'25.83", VIII.1975, coll F. VanDeBogart (as Marasmius androsaceus), FVDB 3621 (WTU-F-51955; holotype); Snowhomish Co., Barlow Pass, Barlow Pass study area, N48.02667, W121.44278 (WGS 84), 8.VII.1992, coll S. Trudell (as Micromphale perforans), s.n. (WTU-F-9305).

United States, Connecticut, Middlesex Co., vic. Salem, Devil’s Hopyard State Park, N41°28.937', W72°20.491', 1.IX.2013, coll RHP, TFB 14332 (TENN-F-68183).

folius (Latin) = leaf; phil- (Greek) = to love; preference for broad-leafed leaves.

Similar to Gymnopus (Mi.) perforans: differing in: 1) fruiting substrate of dead deciduous leaves, most commonly Quercus; 2) unique phylogenetic placement (based on ITS sequences); 3) apparent geographic range in eastern North America.

Basidiomata (Fig. 10) diminutive, scattered to (rarely) gregarious. Pileus 3–22 mm diam, shallowly to strongly convex with downturned margin when young, becoming applanate to slightly depressed centrally by maturity (and then with small central dot of “drab” 6D3 or mouse gray), sometimes with small umbo, subtuberculate, matt to minutely plushy (35×) especially outward, when dried commonly sublaccate (glistening with reflected light rather than matt); disc when fresh “pinkish buff” 6A3, “vinaceous buff” 9B2, near “light pinkish cinnamon” 7A2, “Mikado brown” 7C6, to “drab” 6D3; limb “pale pinkish cinnamon” 6A2, “pale ochraceous buff” 4A2, “pale cinnamon pink” 5A2, “light buff” 3A2, “pinkish buff” 6A3, grayish orange 5B3; margin somewhat thick, entire to uplifted, vaguely sulcate–striate to not striate, usually mellowing to entirely tan to pallid brown. Lamellae adnate–adnexed or occasionally attached to a weak pseudocollarium (especially visible when dried and seceded), subdistant to distant, total lamellae = 30–32; through lamellae = 13–16, arcuate, thickish, narrow (not more than 1 mm broad) with no anastomoses or interveining, with dark brown ring around stipe apex, off-white to “pale pinkish cinnamon” 6A2, near “chamois” 4B4 or “pale ochraceous salmon” 3A3, mellowing to “light buff” 3A2, and perhaps with slight tint of necropigment in storage to “light ochraceous buff” 5A4; lamellar edge entire, never marginate nor fimbriate, occasionally minutely laccate. Stipe 12–30 × 0.8–1.2 mm, terete, equal, insititious, at junction with lamellae “vinaceous russet” 8D4, “bister” 5F8, upward concolorous with gills, “cinnamon buff” 6B4, “avellaneous” 7B3 to “tilleul buff” 7B2, soon downward “wood brown” 5A4, “buffy brown” 6D4, “mummy brown” 6F8, “Verona brown” 6E5, “Rood’s brown” 6D5, “clove brown” 6F5, “chaetura black” 2F3 to “bone brown” 7F8, weakly to densely vestured throughout, variably minutely flocculose, especially upward, to barbed with setoid vesture, hollow, downward becoming stuffed; vesture at stipe apex (30×), hyaline, downward becoming pigmented to dull yellow then dull straw-colored, but never black. Rhizomorphs usually present but widely scattered and inconspicuous, filiform (0.2–0.4 mm thick), resupinate on leaf surface (especially more sclerophyllous leaves), and there branching and anastomosing, sometimes orange-brown and diffuse, otherwise black and meandering, short (never more than 6 mm long), unbranched, straight to somewhat curly apically, tapering to flagelliform terminus. Taste usually reported as negligible, occasionally very weak of garlic; odor usually reported as negligible, occasionally resembling boiled cabbage after drying or mildly fetid.

Figure 10. 

Gymnopus foliiphilus. Basidiomata. Standard bars = 10 mm.

On dead broad-leafed leaves, most often Quercus leaves, fruiting especially on midribs and petioles of both red and white oak complexes; other adventitious substrates include Acer (at least A. rubrum), Cornus, Magnolia, Rhododendron, 2-needle Pinus; Appalachian Mountain chain from New England through northern South Carolina and northern Georgia, west to Arkansas and south to Gulf Coast; late Spring through early Autumn.

Pileipellis a thin tissue involved in a slime matrix, composed of the following: 1) pileal hairs (Fig. 11) –120 × 2–3.5 µm, smooth, occasionally clamped internally, with telltale evidence of superficial mucoid deposit; 2) repent hyphae slender [3.5–5.5(–13) µm diam], firm-walled (wall -1 µm thick, often somewhat gelatinized), generally radially oriented, conspicuously clamped, mostly smooth, occasionally vaguely ornamented (Fig. 12A–C) with minute flakes in slime next to hyphal wall, commonly very vaguely striped with some profile flakes (PhC), rarely significantly encrusted, hyaline, embedded in a thin layer of mucoid material (amorphous debris expressed in paradermal squashes, PhC); 3) common secondarily septate hyphal segments (Fig. 12D); small, peg-like, small lobes (Fig. 12F, G) common. Subpellis hyphae broader (5–7 µm diam), thick-walled (wall –1.0 µm thick), conspicuously clamped, hyaline, interwoven, with evidence of a mucoid or gelatinous deposit. Pileus tramal hyphae 5–21 µm diam, thin- to firm-walled, with minimal slime matrix, clamped. Lamellar trama loosely interwoven, of uninflated cells (2–)3.5–10 µm diam, firm- to thick-walled (wall –0.7 µm thick), lightly encrusted or with evidence of insoluble mucoid matrix, conspicuously clamped. Hymenium dense, with basidia becoming diaphanous after spore discharge but not disappearing (“husking”). Pleurocystidia (Fig. 13A–D) 25–33 × 6–9 µm, fusiform, conspicuously clamped, serially produced from subhymenial clamp connections. Basidioles clavate; Basidia (Fig. 13E–H) (21–)27–32 × (5–)6–9 µm, clavate to narrowly clavate, clamped, 4-sterigmate (occasional individuals, always semicollapsed, observed with two prolonged sterigmata), obscurely to conspicuously clamped; contents minutely multigranular, not guttulate. Subbasidial cells appearing catenulate, lobose but hyphal (not as lobose as in G. perforans). Basidiospores (Fig. 14) (4.5–)6.5–7(–8) × 2.5–3.5(–4) µm (Q = 1.33–2.80; Q^m = 1.92; L^m = 6.85 µm), gymnopoid to pip-shaped (ellipsoid, not tapered proximally), smooth, thin-walled, inamyloid. Cheilocystidia observed only in rare specimens, clavate, 28–34 × 9–10 µm, broadly clavate, not longer than basidia but broader, hyaline, clamped. Stipe medullary hyphae 4–9 µm diam, irregularly thick-walled (wall –1.5 µm thick) as though encrusted in bands (but not so), hyaline, conspicuously clamped; outer medullary hyphae 3–6 µm diam, hyaline, thick-walled (wall –2 µm thick), free (not adherent), strictly parallel, conspicuously clamped, commonly anastomosed in “H” connections. Stipe cortical hyphae 3.5–8 µm diam, smooth, thick-walled (wall –1.0 µm thick), pigmented in cytoplasm, producing caulocystidia as side branches. Caulocystidia (Fig. 15) 25–>150 × (2.5–)6–9 µm, arising from a tangled thatch of dry, interwoven, very thick-walled (occluding cell lumen), clamped hyphae, a combination of straight and setoid, mixed with other gnarled or curled individuals, densely scattered, thick-walled, usually arising as hyphal terminus or commonly with short abortive branch below, pigmented in cytoplasm (wall subhyaline), usually somewhat broader near origin than at mid-point (6–10 µm diam). Lower stipe caulocystidia (Fig. 16) setoid, 4–7 µm diam, thick-walled (occluding cell lumen), arising as side branches from stipe surface hyphae, gathered into rough synnematal spines.

Figure 11. 

Gymnopus foliiphilus. Pileal hairs. Standard bars = 10 µm. TFB 2800 (TENN-F-49363).

Figure 12. 

Gymnopus foliiphilus. Pileipellis elements. A–C Encrusted hyphae D Hyphae showing secondary septa E Gelatinized hyphal walls F, G Lobate side branches. Standard bars = 10 µm. TFB 2800 (TENN-F-49363).

Figure 13. 

Gymnopus foliiphilus. Pleurocystidia and basidia. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. TFB 11555 (TENN-F-59441).

Figure 14. 

Gymnopus foliiphilus. Basidiospores. Standard bar = 5 µm. A = TFB 11608 (TENN-F-59641); B = TFB 14322 (TENN-F-68183).

Figure 15. 

Gymnopus foliiphilus. Caulocystidia from upper stipe. Standard bars = 10 µm. TFB 2800 (TENN-F-49363).

Figure 16. 

Gymnopus foliiphilus. Caulocystidia from lower stipe. Standard bars = 10 µm. TFB 2800 (TENN-F-49363).

Gymnopus foliiphilus is the most commonly collected North American taxon in sect. Perforantia. Preliminary field identification attempts to distinguish several taxa with similar basidiomata. Substrate segregates G. androsaceus and G. perforans (conifer needles, usually Picea and/or Abies), from G. foliiphilus. Geographic distribution is less secure. Both G. androsaceus and G. perforans are found in Europe and temperate North America, while G. foliiphilus seems limited to eastern North America. From all these taxa, a mimic, Marasmius pallidocephalus, is separated from G. androsaceus with difficulty in the field, based almost solely by lack of clamp connections of the former and phylogenetic placement.

Often in collections of G. foliiphilus, evidence of some bleaching of substrate can be detected. This is not dramatic – not to pale off-white – but distinct nonetheless. The phenomenon cannot be compared to M. perforans because fallen needles of Picea/Abies naturally bleach over time.

In G. folliiphilus, stipes are almost always bicolor, upward with some avellaneous to pinkish shade, downward to dark brown and finally black toward the base (note that the very junction of stipe and lamellae is always dark brown). The relative stipe length of these colors varies considerably, with the upward avellaneous shades from only the uppermost 10% to as much as the upper 50%. To some extent, the density and quality of the stipe vesture also varies, with upper surfaces producing shorter, less setoid, hyaline caulocystidia, often with long, slender, hyaline hyphae producing a sparsely silky or wispy appearance (40×) rather different from the hispid or barbed appearance of the lower stipe caused by setoid caulocystidia often gathered into synnematous sheaves.

A seductive artifact in microscope mounts of hymenial structures are the subbasidial cells. As is typical, basidioles and basidia are produced in “bouquets” by subbasidial hyphae, which usually are tightly packed but which retain hyphal characteristics. In G. foliiphilus, subbasidial hyphae are catenulate or congestedly lobose. The result are structures which mimic the cheilocystidia of numerous Gymnopus taxa, especially in sect. Vestipedes. In G. foliiphilus (as in G. perforans), cheilocystidia are difficult to interpret, and when present, are consummately basidiiform.

Traditional generic characters are not consistent in sect. Perforantia and sect. Androsacei. For example, the pileipellis of G. androsaceus resembles a rameales structure of repent but diverticulate hyphae. Such a pileipellis is also present in Marasmiellus, taxa of which seem to belong to several relatively distantly related clades. In Micromphale, a gelatinous layer within the pileus trama can usually be demonstrated (i.e. M. foetidum, etc.), but in Micromphale sect. Perforantia the gelatinous layer is absent, but “replaced” by a thin slime matrix over and within the pileipellis.

A paper by Farnet et al. (1999) employed an agar medium reputed to promote production of rhizomorphs. For the present study, this medium (whole wheat flour 20 g/L; agar, Bacto 20 g/L; H₂O 1 L) was used for numerous dikaryon isolates of various Marasmius and Micromphale collections. Ancillary to production of rhizomorphs, aerial mycelium of isolates of M. foliiphilus slowly changed from white to bright yellow (“empire yellow” 3A6), while aerial mycelium of M. perforans remained white.

Desjardin (pers. comm.) indicated the possibility that Marasmius insititius Fr. (1838. Epicrisis: 386), fruiting on Quercus leaves in Sweden, might be similar to M. foliiphilus. Marasmius insititius has seen a checkered history. Recently, Antonín and Noordeloos (2010) excluded the epithet from Marasmius because: 1) no type specimen exists; 2) Fries’s description is less than explicit; 3) in spite of Fries’s physical location in central Sweden in 1838, a habitat on Quercus leaves might indicate his exposure to the organism in southern Sweden; and 4) Orton (1960): 303) had dismissed the epithet as a later heterotypic synonym of Marasmius calopus. Desjardin (1989) did not include M. insititius in his type specimen studies, presumably based on the above and its extralimital status for Marasmius of the southeastern United States. Svengunnar Ryman (UPS; pers. comm.) indicates that M. insititius is an unknown entity. It is not the purpose of this paper to attempt to exhume M. insititius, especially as this name was not taken up in the Scandinavian mycota by Noordeloos in Funga Nordica (Knudsen and Vesterholt 2012).

Once informed of our intention to propose a new species to represent the oak-loving relative of Mi. perforans, Desjardin (pers. comm.) graciously supplied extensive notes on three specimens [DED 4329 (TN), DED 4449 (SC), DED 4477 (NC)] and numerous citations of herbarium specimens chiefly listed under Ma. epiphyllus and Ma. insititius from AL, OH, PA, VA (not represented in “specimens examined” below).

UNITED STATES, Arkansas, Baxter Co., vic. Big Flat, Ozark National Forest, Leatherwood Wilderness, N36°02.4', W92°23.2', 23.X.2013, coll RHP (as M. perforans var. quercophilus), TFB 14422 (ITS, TENN-F-69084). Connecticut, Middlesex Co., vic. Salem, Devil’s Hopyard State Park, 41°28.937'N, 72°20.491'W, 1.ix.2013, coll RHP, TFB 14332 (ITS, TENN-F-68183, holotype). Georgia, Rabun Co., vic. Clayton, Warwoman Dell picnic area, 15.VI.1992, coll SA Gordon, TFB 4902 (ITS, TENN-F-51221). Mississippi, Stone Co., Ramsey Springs, Red Creek Wildlife Management Area, N30°46.572', W88°54.815', coll. RHP, TFB 14291 (ITS, TENN-F-68145). North Carolina, Macon Co., vic. Highlands, Rte 106, Blue Valley Overlook, north side of road, N35°01'45.15", W83°16'57.09", 20.VII.1n989, coll RHP, TFB 2800 (TENN-F-49363); vic. Highlands, Bull Pen Rd. at Slick Rock, 29.VI.1992, coll SA Gordon, TFB 4928 (ITS, TENN-F-51244); vic. Highlands, Nantahala Nat. For., Blue Valley, Trail to Pickelseimer’s Falls, 23.VII.1994, coll RHP, TFB 7243 (TENN-F-56223); same location, Road 79, 14.VIII.1999, coll RHP, TFB 10364 (ITS, TENN-F-57923); same location, Forest Rd. 79, N35°01.103', W83°14.697', 1.VIII.2012, coll. RHP, TFB 10463 (TENN-F-67809); same location, Blue Valley Campground, N35°00'45.23", W83°09'29.33", 11.VIII.2014, coll. RHP, TFB 14508; same location, FR 77 gate area, N35°00.243', W83°14.151', 14.VIII.2014, coll RHP, TFB 14531 (TENN-F-69226); same location, start of FR 79, at picnic area, N35°01.085', W83°14.715', coll RHP (as Mi. perforans var. quercophilus), TFB 13875 (ITS, TENN-F-65571); same location, FR 79, N35°01.103', W83°14.697', 1.VIII.2012, coll RHP (as Mi. perforans var. quercophilus), TFB 14063 (TENN-F-67809); Cliffside Lake Rd., N34°04.749', W83°14.150', 30.VIII.2012, coll RHP, TFB 14048 (ITS, TENN-F-65977). South Carolina, Oconee Co., Oconee State Park, Nature Trail, N34°52'07", W83°06'20", 17.V.1991, coll SA Gordon, TFB 3612 (TENN-F-50731); TFB 3615 (TENN-F-50734); Oconee State Park, N39°52'07", W83°06'19", 18.VIII.1992, coll SA Gordon, TFB 5051 (TENN-F-51454); Rte 107 circ. 12 m south of Cashiers, N34°59'37", W83°03'09", 16.VIII.1992, coll RHP, TFB 5435 (TENN-F-51753); vic. Cashiers (NC), Walhalla Fish Hatchery, N34°59.155', W83°04.374', 7.VIII.1996, coll RHP, TFB 8782 (ITS, TENN-F-55210). Tennessee, Blount Co., GSMNP, Spruce Flats, N35°37'18.4", W83°40'26.3", 24.VI.1991, coll RHP & V. Antonín, TFB 3659 (TENN-F-50778); GSMNP, Metcalf’s Bottoms, picnic area, 9.VI.1997, coll RHP, TFB 9166 (ITS, TENN-F-55764); GSMNP, vic Crib Gap, N35°36'45.8", W83°44'42.1", 22.VII.1991, coll & det DE Desjardin (as Micromphale perforans var. quercophilus), DED 5272 (ITS, TENN-F-50013; SFSU); GSMNP, Greenbrier at trailhead to Ramsay’s Cascades, 20.VI.1991, coll SA Gordon, RHP, V. Antonín, TFB 3642 (ITS, TENN-F-50761); Foothills Parkway, Look Rock Campground, 10.VIII.2003, coll KWH & RHP, TFB 11608 (TENN-F-59641); Cocke Co., vic. Cosby, GSMNP, Gabes Mt. Trail, N35°12'33.0", 5.VII.2006, coll M. Padansee, E Lickey, TFB 13242 (TENN-F-61274).

Costa Rica, Prov. San José, km 68 on PanAmerica Highway, “Finca Jaular,” N9°39.597', W83°52.115'. 29.VI.2000, coll. RHP, TFB 9750 (TENN-F-58651).

Similar to Gymnopus foliiphilus, but differing as follows: 1) pileus darker at all ages, smooth (not pebbled), not convex when young nor everted in age; 2) lamellae more numerous and lamellulae in two ranks; 3) lamellar attachment adnexed to adnate (not pseudocollariate); 4) stipe black or bicolored; 5) stipe insertion subinsititious (not insititious); 6) outer pileipellis hyphae often significantly encrusted (not smooth or with flakes).

Basidiomata (Fig. 17A) diminutive, gymnopoid. Pileus 6–14 mm broad, appearing applanate throughout ontogeny but with downturned margin, outward vaguely sulcate–striate, always very thin-fleshed, smooth (but not glabrous), matt (but not velutinous, etc.), dark brown when young (“amber brown” 6D8, “mummy brown” 6F8, “Natal brown” 8E6, “bone brown” 7F8), fading somewhat by maturity (“tawny” 6C6, “wood brown” 7C4, “tawny olive” 5C5), essentially unicolorous (but occasionally with a suggestion of a paler central disc). Lamellae (Fig. 17B) (dried) appearing adnate to adnexed but not pseudocollariate, shallow (–1 mm deep), total lamellae = 30–37, through lamellae = 12–15, “sulphur yellow” 1A2 to “pale ochraceous buff” 4A2, now (dried) near “ochraceous buff” 5A5 to “light ochraceous buff” 5A4 with no evidence of necropigment. Lamellulae in two ranks. Stipe 13–30(–40) × 0.6–1(–1.5) mm, terete, equal, twisted and compressed upon drying, lightly stuffed to hollow, vestured throughout, from minutely flocculose (40×) to barbed but not consistently located – sometimes flocculose vesture apical, sometimes both apical and mid-section and alternating with barbed texture; color never black, often bicolorous, upward concolorous with lamellae, soon downward darker, “tawny” 6C6, finally dull fuscous (“drab” 6D3, “hair brown” 6E3, “citrine drab” 4D5); insertion insititious to subinsititious (very small basal pad – hardly noticeable). Rhizomorphs (Fig. 17A) sparse, slender, meandering over leaf surface before becoming erect, unbranched, never more than 10 mm long. Odor faint of cabbage, stronger with age; taste not recorded.

Figure 17. 

Gymnopus foliiphilus var. costaricensis. A Basidiomata and rhizomorphs B Pileus underside showing lamellar spacing C Pleurocystidia D Basidia E Pileipellis hyphae, showing encrustation and lobose side-branches F Basidiospores. Standard bars: A = 5 mm; B = not to scale; C–E = 20 µm; F = 5 µm. TFB 9750.

Fruiting on sclerophyllous leaves (perhaps Quercus) at mid- to high-elevation; early summer.

Pileipellis an intricately interwoven layer of thin-walled, conspicuously clamped hyphae in a heterogeneous matrix of slime (including copious debris), composed of: 1) hyphae 5–15 µm diam (Fig. 18A, B), firm-walled with thin slime sheath with flake-like crust material riding on outside of sheath; 2) hyphae 6–14 µm diam, firm-walled, producing variable small lobes or outgrowths (Figs 17E, 18C–J), occasionally proliferating into ribbon-like extensions (Fig. 18K, L); and 3) occasional strongly encrusted hyphae present; hyphae 4–7 µm diam, encrustation in significant scabs sometimes suggesting stripes, with minimal profile calluses. Lamellar trama loosely interwoven; hyphae 2.5–3.5 µm diam, firm-walled, long-celled, conspicuously clamped (often medallion). Pleurocystidia (Figs 17C, 19A–D) 17–24 × 6.5–8 µm, fusiform, conspicuously clamped; contents homogeneous. Basidioles clavate; basidia (Figs 17D, 19E–H) 18–22 × 7–8, broadly clavate, clamped, 4-sterigmate. Small clots of subgelatinous material in hymenium (scattered, evidenced by copious “sludge” and/or debris); effete basidia and pleurocystidia evacuating contents but retaining uncollapsed walls (“husking”). Basidiospores (Fig. 17F) (6–)6.5–8 × 3–4 µm (Q = 1.75–2.67; Q^m = 2.11; L^m = 7.20 µm), gymnopoid (not tapered proximally), smooth, thin-walled, inamyloid. Cheilocystidia (Fig. 20) 23–29 × 5–9 µm, clavate, basidioid, usually slightly subcapitulate. Stipe medullary hyphae 3–17 µm diam, strictly parallel, perhaps adherent (shearing in plates and with copious floating debris), hyaline, firm- to thick-walled (wall –1.0 µm thick), obscurely clamped, with occasional constricted lobes. Caulocystidia from upper stipe (Fig. 21A) sub-hyaline (pale champagne color in mass, refringent PhC), gnarled (not straight/setoid), thick-walled (wall –2 µm thick), usually clamped internally, gathered into “plaques” (which appears as floccules at 40×). Caulocystidia from flocculose lower stipe (Fig. 21B) -110 µm long, thick-walled (wall –2 µm thick, yellow-refringent; cytoplasm hyaline), usually lobed or branched at base, 7–12 µm broad.

Figure 18. 

Gymnopus foliiphilus var. costaricensis. Pileipellis elements. A, B Firm-walled hyphae with slime sheath and crust material on exterior of the sheath C–J Diverticulate hyphae with individual lobate outgrowths K, L Extended lobes into ribbon-like proliferations. Standard bars = 10 µm. TFB 9750.

Figure 19. 

Gymnopus foliiphilus var. costaricensis. Hymenial elements. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. TFB 9750.

Figure 20. 

Gymnopus foliiphilus var. costaricensis. Cheilocystidia. A Overview of lamellar edge B–E Individual cheilocystidia. Standard bars = 10 µm. JLM 2221 (USAM).

Figure 21. 

Gymnopus foliiphilus var. costaricensis. Caulocystidia. A Individuals from near stipe apex B Individuals from lower stipe. Standard bar = 20 µm.

Smooth pileus surface and encrusted pileipellis hyphae are suggestive of Gymnopus sect. Vestipedes, but obvious slime matrix points to sect. Perforantia. The vestured stipe also might point toward G. perforans or sect. Vestipedes (not toward Gymnopus sect. Androsaceus s.s.).

Traditionally, G. perforans has been accepted as quite variable in substrate. ITS phylogeny indicates separation into several species, with G. perforans s.s. limited to conifer needles (usually Picea/Abies), with segregants on dead deciduous leaves (G. foliiphilus), or needles of Pinus strobus (G. pinophilus) or P. ponderosa (G. ponderosae). Collection TFB 9750 TENN-F-58651 matches this complex micromorphologically, and especially G. foliiphilus in substrate.

Costa Rica, Prov. Heredia, County Barva, District San José de la Montaña, viz. Sacramento, 25.VIII.2014, JLM 2238 (USAM); Prov. San José, km 68 on PanAmerica Highway, Finca Jaular, N9°39.597', W83°52.115'. 29.VI.2000, coll. RHP, TFB 9750 (TENN-F-58651; holotype); Prov. San José, County Dota, District Jardin, viz. La Chonta, Finca Sta. Maria, 4 km on Hwy 2 from intersection at Empalme, N9°41.806, W83°55.763', 14.VIII.2014, JLM 2219 (USAM); Same location, same date, JLM 2221 (USAM).

United States, Washington, Jefferson Co., Irely Lake Area, North Fork Quinault River, N47°33'47.7", W123°40'12.38", date not recorded, coll M.T. Seidl, ident. S Redhead (as Micromphale perforans), MTS 3715 (WTU-F-9293).

fragillior = Latin, more fragile, referring to more diminutive stature than M. perforans.

1) A morpho-taxon (no DNA sequences available); 2) basidiomata slender (stipe 15–25 × 0.4–0.7 mm), diminutive, slender; 3) stipe base (<1 mm) with hirsute, tawny collar; 4) sterigmata stout, subcornute; 5) basidiospores broadly ellipsoid to pip-shaped (Q^m =1.68); 6) stipe medullary hyphae thick-walled, not involved in slime matrix.

Basidiomata (Fig. 22A) slender, diminutive, attached to individual needles of ?Abies grandis.

Figure 22. 

Gymnopus fragillior. A Basidiomata B Basidiospores. Standard bars: A = 20 mm; B = 5 µm. WTU-F-9293.

Pileus 6–10 mm broad, convex to plano–convex, often somewhat centrally depressed, matt, delicately tuberculate; disc (fresh) pale grey brown, (dry) “army brown” 8D5 to “sayal brown” 6C5; limb and margin (fresh) cream, (dry) “wood brown” 7C4; margin thin; pileus trama dull orange to orange–brown in Melzer’s reagent. Lamellae adnate, distant, thickish, in 2–3 tiers, weakly pseudocollariate, seceding upon drying, (fresh) pallid tan, “tilleul buff” 7B2, “olive buff” 3B3, (dry) slightly necropigmented (near “light ochraceous buff” 5A4). Stipe 15–25 × 0.3–0.5 mm, terete, equal, more or less straight, hollow, non-insititious, lightly vestured to delicately furfuraceous throughout, in contact with lamellae dark brown, apically tan to pale cream (“cinnamon buff” 6B4), downward brown, dark brown, base blackish brown (“clove brown” 6F5, “fuscous black” 6F4); stipe base (<1 mm) a minutely hispid collar, “tawny” 6C6 to “ochraceous tawny;” basal pad restricted, “tawny” 6C6; medulla thin, off-white (not white). Rhizomorphs not observed. Odor alliaceous; taste not recorded.

Known only from the holotype collection. “Scattered on outer bark of fallen tree, mostly fruiting from needles fallen on log” (teste Seidl) (perhaps Abies grandis). Seasonality unknown.

Pileipellis a repent, unoriented layer in a thin slime or gelatinous matrix with detersile encrustation material, composed of the following: 1) pileal hairs (Fig. 23B–E) 15–75 × 5–10 µm diam, digitate to vermiform, equal, hyaline, firm- to thick-walled (wall –0.5 µm thick), obtusely rounded at apex; 2) repent hyphae 4–13 µm diam, thin- to firm-walled, conspicuously clamped (but with occasional secondary septa), at pileus surface strongly to moderately encrusted (Fig. 23A); crust material in scabs, occasionally with suggestion of annular pattern, 0.7–2 µm thick, in subpellis as flakes riding on thin slime sheath. Pileus and lamellar tramae loosely interwoven, dull orange to orange–brown in IKI; hyphae 5–12 µm diam, thin- to firm-walled, with thin slime sheath in KOH, conspicuously clamped. Pleurocystidia (Fig. 24) 28–35 × 5–8 µm, fusiform to narrowly fusiform, conspicuously clamped, hyaline; contents homogeneous. Basidioles (Fig. 25A, B) clavate, clamped; basidia (Fig. 25C, D) 28–36 × 7–10 µm, clavate, usually subcapitulate, conspicuously clamped, 4–sterigmate; sterigmata –6 × 2.5 µm, stout, subcornute. Basidiospores (Fig. 22B) (6–)6.5–7.5(–8) × 3.5–4.5 µm (Q = 1.44–2.00; Q^m = 1.68; L^m = 6.90 µm), plump ellipsoid to pip-shaped, smooth, thin-walled, inamyloid. Cheilocystidia not observed. Stipe medullary hyphae strictly parallel, 4.5–14 µm diam, apparently free (not involved in slime or gelatinous matrix), thick-walled (wall–0.7 µm diam, hyaline), obscurely clamped; occasional slender hyphae (4–5.5 µm diam) meandering among broader hyphae. Stipe cortical hyphae 4.5–7.5 µm diam, thick-walled (wall –2.5 µm thick, pigmented, producing caulocystidia as side branches), obscurely clamped. Mid-stipe caulocystidia (Fig. 26) in scattered clusters, setoid, arising as side branches of surface cortical hyphae or as termini, 15–75 × 7–13 µm diam, not clamped, digitate to subvermiform, thick-walled (wall –5 µm thick, pigmented pale yellowish, refringent; PhC). Caulocystidia from stipe base (Fig. 27) densely gregarious, setoid, 20–95 × 7–13 µm, usually straight, tapering somewhat distally, with basal appendage, thick-walled (wall often occluding cell lumen, yellowish, refringent).

Figure 23. 

Gymnopus fragillior. Pileipellis elements. A Encrusted, repent hypha B–E Pileal hairs. Standard bars = 10 µm. WTU-F-9305.

Figure 24. 

Gymnopus fragillior. Pleurocystidia. Standard bars = 10 µm. WTU-F-9305.

Figure 25. 

Gymnopus fragillior. Basidial development. A, B Basidioles C, D Mature basidia. Note stout sterigmata. Standard bars = 10 µm. WTU-F-9503.

Figure 26. 

Gymnopus fragillior. Mid-stipe caulocystidia. A Cluster of caulocystidia B Small individual C Double caulocystidia D–G Individual caulocystidia. Standard bars = 10 µm. WTU-F-9305.

Figure 27. 

Gymnopus fragillior. Stipe-base caulocystidia. Standard bars = 10 µm. WTU-F-9305.

A pileipellis involved in slime matrix, lacking diverticulate hyphae and/or broom cell-like termini, coupled with vestured stipe, indicate placement in Gymnopus (Micromphale) sect. Perforantia. There it is distinguished from G. perforans, G. pallidocephalus and other taxa with similar characters. Unusual, however, are the scattered clusters of caulocystidia, the stipe base with delicately hispid, brown caulocystidia and pip-shaped spores. From Mycetinis salalis comb. prov, also in the moist forests of the Pacific Northwest, G. fragillior is immediately separated by spore dimensions (15–17 × 3.5–4 µm; Q^m = 4.38 in the former), habitat on conifer needles (rotting wood in the former), different pileipellis structures and absence of cheilocystidia (marasmioid in M. salalis). From both G. perforans and G. pallidocephalus, which share habitat on conifer needles, G. fragillior is distinguished by non-black stipe (black, vestured and more robust in G. perforans, black and glabrous–shining in M. pallidocephalus), and common clamp connections (absent in G. pallidocephalus). Basidiomata of G. bulliformis are somewhat more robust, with pileus somewhat paler and prominent, utriform cheilocystidia (absent in G. fragillior).

United States, Washington, Jefferson Co., Irely Lake Area, North Fork Quinault River, N47°33'47.7", W123°40'12.38", date not recorded, coll M.T. Seidl, ident. S Redhead (as Micromphale perforans), MTS 3715 (WTU-F-9293; holotype).

United States, Washington, Snowhomish Co., Hwy 92, Perry Creek Trail, 24.VI.1993, coll GR Walker (as Micromphale perforans), GRM 410 (WTU-F-9309).

Glabro- = Latin, smooth; pes = Latin, foot, referring to the glabrous–shining stipe.

1) A morpho-species (no DNA sequences available; 2) stipe glabrous–shining; 3) fruiting on dead Tsuga needles; 4) lamellae suffusing brown from edge when bruised or dried; 5) stipe 15–25(–50) × 0.4–1 mm, robust; 6) stipe medullary hyphae involved in slime matrix.

Basidiomata (Fig. 28) diminutive with robust stipe. Pileus 4–9 mm broad, plano–convex to plane, matt, smooth, more or less unicolorous, hardly sulcate at margin, more or less unicolorous, dark brown (“carob brown” 7F7, “chestnut brown” 7E4); context near “olive buff” 3B3, reluctantly bruising to near “buffy brown” 6D4 when cut or bruised. Lamellae adnexed to free, distant, with no anastomosis or interveining, thickish, near ”olive buff” 3B3 (dried), bruising to dull “buffy brown” 6D4 from edge when bruised or dried; lamellulae rare, rudimentary. Stipe 23–30(–50) × 0.7–1.2 mm, robust, terete, equal, insititious, glabrous–shining overall, delicately sulcate–ridged, stuffed, apically “buffy brown” 6D4, soon downward “chaetura drab” 2F2 to “chaetura black” 2F3 overall; stipe medulla white. Rhizomorphs extensive but inconspicuous, black, polished, filiform, –14 × 0.2–0.4 mm, curly, branched with branches spur-like. Taste and odor not recorded.

Figure 28. 

Gymnopus glabrosipes. Basidiomata and rhizomorphs. Standard bar = 20 mm. WTU-F-9309.

Fruiting on needles of Tsuga (associated with Abies); mid-summer.

Pleipellis a layer of unoriented, tightly interwoven, repent, strongly encrusted hyphae involved in a gelatinous or slime matrix; superficial hyphae 5–7.5 µm diam, thin-walled, conspicuously clamped, strongly encrusted (Fig. 29A); incrustation material –2 µm thick, in scabs and vaguely annular patterns; subpellis hyphae 6–11 µm diam, thick-walled (wall 1.5–3 µm thick, hyaline), smooth to weakly encrusted; crust material <1 µm thick, in small scabs and vaguely defined annular patterns, sometimes flake-like on surface of slime sheath (Fig. 29B). Inner subpellis hyphae 7–11 µm diam, smooth, thick-walled with wall slowly (in KOH) swelling or gelatinizing (Fig. 29C), conspicuously clamped. Lamellar trama loosely interwoven; hyphae 2.5–3.5 µm diam, thin-walled, conspicuously clamped, often with thin slime sheath. Hymenium composed of pleurocystidia and basidia, gelatinizing (in KOH) from trama outward, eventually reducing basidial and pleurocystidial bases to gelatinous matrix. Pleurocystidia (Fig. 30B–D) 23–30(–42) × 4–7 µm, narrowly fusiform without partitioned contents, conspicuously clamped. Basidioles narrowly clavate; basidia (Fig. 30A) 29–32(–39) × 9–11 (at widest point), clavate, conspicuously clamped, 4–sterigmate; contents homogeneous. Basidiospores not observed. Cheilocystidia not observed. Stipe medullary hyphae appearing dimitic, without slime matrix; 1) 1.5–7.5 µm diam, thick-walled (wall –1 µm thick, hyaline, non-refringent; PhC) (Fig. 31), frequently clamped, often branched, loosely interwoven, meandering among refringent, wider hyphae; and 2) 5–12 µm diam, tapered at both ends, thick-walled (wall gelatinizing –6 µm thick, strongly refringent, PhC), long-celled, clamped. Stipe cortical hyphae strictly parallel, 2–4.5 µm diam, thick-walled (wall –1 µm thick, pigmented yellow–brown), clamped, minutely roughened outward (perhaps a roughened mucoid film). Caulocystidia absent.

Figure 29. 

Gymnopus glabrosipes. Pileipellis hyphae. A Strongly encrusted hypha with vaguely annular pattern B Hypha with gelatinizing walls and thin slime sheath, with incrusting material on the surface of the slime sheath C Smooth hyphae with gelatinized wall. Standard bars = 10 µm. WTU-F-9309.

Figure 30. 

Gymnopus glabrosipes. Hymenial structures. A Basidium: note 2 sterigmata out of focus B–D Pleurocystidia: note basal clamp connections. Standard bars = 10 µm. WTU-F-9309.

Figure 31. 

Gymnopus glabrosipes. Stipe medullary hyphae. Slender generative hyphae among refringent, thick-walled skeletalized hyphae. Standard bar = 10 µm. WTU-F-9309.

Extraction of usable DNA from the two specimens examined was unsuccessful, so phylogenetic placement of G. glabrosipes remains unknown.

Pileipellis involved in slime matrix, pileus tramal hyphae with gelatinizing walls, basidia and pleurocystidial bases strongly gelatinizing and absence of cheilocystidia all are diagnostic of the Mi. perforans complex. Conversely, the glabrous–shining stipe is not characteristic of that clade.

Two pilei were assessed for diverticulate hyphae with no success. Pileipellis was the same in both.

Macromorphologically, basidiomata are reminiscent of those of Ma. thiersii, with dark chocolate brown pileus and insititious, brown–black stipe. From M. thiersii, WTU 9309 differs in glabrous–shining stipe (minutely velutinous in Ma. thiersii), and absence of diverticulate hyphae (present in Ma. thiersii). WTU 9309 is a Micromphale, Ma. thiersii belongs in Marasmius sect. Androsacei.

Although basidiospores have not been observed (in both specimens examined), basidiospores in this clade are quite uniform, and the organism’s identification is not dependent on these statistics.

Oregon, Clackamas Co., road to Mt. Hood at 2–3 mi post, 25.VI.1995, coll MT Seidl, J Roger, N Weber (as Marasmius androsaceus), MTS 4078 (WTU-F-8919); Washington, Snowhomish Co., Hwy 92, Perry Creek Trail, 24.VI.1993, coll GR Walker (as Micromphale perforans), GRM 410 (WTU-F-9309).

Agaricus perforans Hoffm.: S.F. Gray. 1821. Nat. Arr. Brit. Pl. 1: 622.

≡ Agaricus perforans Hoffm.: Fr. 1821. Syst. Mycol. 1: 138.

≡ Micromphale perforans (Hoffm.: Fr.) S.F. Gray. 1821. Nat. Arrang. Brit. Plants 1: 622.

≡ Marasmius perforans (Hoffm.: Fr.) Fr. 1838. Epicric. 385.

≡ Androsaceus perforans (Hoffm. : Fr.) Patouillard. 1887. Hymen. Eur.: 105.

≡ Marasmiellus perforans (Hoffm.: Fr.) Antonín & Noordeloos. 1997. Mycotaxon 63: 366.

(fide Antonín & Noordeloos, 1997): Sweden, Medelpad, Borgsjö, Granbodsos, 31.VIII.1993, M.E. Noordeloos 93137 (L).

1) Fruiting habit on conifer needles, chiefly Picea; 2) pileipellis and subpellis involved in thin mucoid matrix; 3) stipe barbed–vestured at least in lower half; 4) pileipellis lacking diverticulate hyphae and/or broom cells; 5) clamp connections ubiquitous; 6) cheilocystidia rare, broadly saccate to utriform; 7) lamellae few, reduced.

Basidiomata (Fig. 32A) marasmioid. Pileus 5–13(–20) mm broad, plano–convex or subconcave, usually with small papilla, often with shallow central depression, sometimes faintly sulcate, hygrophanous, glabrous to matt, pallid brown to light brown (“wood brown“ 7C4, “drab” 6D3, “buffy brown” 6C–D3–4) fading to pallid white in age. Lamellae rather distant, free, adnexed to narrowly adnate, with no anastomosis or interveining, subarcuate, total lamellae = 13–19, through lamellae = 1–3, segmentiform, 1–3 mm broad, pale pinkish brown, with concolorous or slightly paler, pastel off-white to pallid brownish orange (“drab” 6C4) in age; edge entire. Stipe 15–40 × 0.5–1 mm, insititious, cylindrical, rarely compressed, stiff, filiform, finely sulcate lengthwise, vestured, at apex concolorous with pileus to “deep olive buff” 5B3, in lower parts dark brown (“Brussels brown” 6E5–6) to black, when mature often entirely black; vesture very sparse upward, setose only downward toward very base, which is usually densely hispid. Rhizomorphs (Fig. 32A) abundant, –20 × 0.4–0.6 mm, tapering gradually to <0.2 mm broad, very curly, frequently branched, obviously binding adjacent needles and bits of debris, black, glabrous–shining. Odor fetid, like rotten cabbage; taste “mildly unpleasant,” often tardily alliaceous.

Figure 32. 

Gymnopus perforans. Basidiomata and basidiospores. A Basidioma and rhizomorph B, C Basidiospores. Standard bars: A = 10 mm; B, C = 5 µm. A, BTFB 7272 (TENN-F-53624). CTFB 7477 (TENN-F-53579).

In large troops on fallen needles of Picea, rarely also Pinus and/or Abies, in humus-rich, coniferous plantations; widespread over Europe and Scandinavia (perhaps also in Asia); July to November.

Pileipellis embedded in hyaline slime matrix, a thin ixocutis, composed of the following elements: 1) pileal hairs (Fig. 33A–D) –120 × 3–4.5 µm, common, erect, thin-walled, conspicuously clamped, equal to slightly tapering distally and then with somewhat inflated apex; 2) repent hyphae 3–8.5 µm diam (Fig. 33E), smooth, turgid on pileus surface, compactly interwoven, thin- to firm-walled (not thick-walled), conspicuously clamped; and 3) similar hyphae, occasionally delicately encrusted in stripes with minimal profile calluses. Pleurocystidia (Fig. 34, 36A) 35–42 × 7–8 µm, fusiform, rounded at apex (seldom acutely so), conspicuously clamped. Basidioles (Fig. 35A) clavate; basidia (Figs 35B–D; 36B) (–25)36–45 × 9–11 µm, clavate, often subcapitulate, (2–)4-sterigmate, conspicuously clamped. Basidiospores (Fig. 32B, C) (5.5–)6.5–8.5(–9.5) × (3.5–)4–5 µm (Q = 1.50–2.57; Q^m = 1.95; L^m = 8.10 µm), ellipsoid to narrowly pip-shaped, slightly tapered proximally, thin-walled, smooth, inamyloid. Lamellar edge heterogeneous. Cheilocystidia (Fig. 33C–M) locally abundant, 30–41 × 10–29 µm, short- to long-stalked (stalk 3–10 × 3–4.5 µm, obscurely clamped), ventricose–rostrate, utriform to broadly, usually irregularly saccate to nearly globose, often with small pustulate outgrowths, significantly larger than pleurocystidia or basidia, thin-walled, hyaline; contents heterogeneous. Stipe medullary hyphae 2–9 µm diam, hyaline, strictly parallel, adherent in a slime matrix, conspicuously clamped, thick-walled (wall –1.0 µm thick, irregular on inner wall). Stipe cortical hyphae 4–7 µm diam, yellow–brown (PhC), strictly parallel, thick-walled (wall –2.0 µm thick), exteriorly smooth, vaguely dextrinoid (reddish brown in IKI, BF). Stipe vesture from near stipe apex composed of two elements: 1) broad, scattered, thick–walled caulocystidia as described below; and 2) long, flexuous, thin–walled “hairs” –140 × 1.3–3 µm diam, apparently thin- to firm-walled, arising from clamp connections (not as side branches), hyaline. Caulocystidia (Fig. 37) from lower stipe –90(–140) × 4–8.5(–13) µm, arising as side branches of stipe cortex surface cells, thick-walled (wall –2 µm thick, hyaline), erect, setoid, densely scattered, often in narrow clumps of 2–4 (appearing hispid at 30×), obscurely vacuolated, often internally secondarily septate, moderately dextrinoid (reddish brown in IKI, BF, especially reddish apically).

Figure 33. 

Gymnopus perforans. Pileipellis structures. A–D Pileal hairs E Repent hypha showing clamp connection. Standard bars = 10 µm. TFB 7477 (TENN-F-53579).

Figure 34. 

Gymnopus perforans. Pleurocystidia. Standard bars: A = 10 mm; B = 5 µm. TFB 7272 (TENN-F-53624).

Figure 35. 

Gymnopus perforans. Basidiole and basidia. A Basidiole B–D Basidia. Standard bars: A = 10 mm; B = 5 µm. TFB 7272 (TENN-F-53624).

Figure 36. 

Gymnopus perforans. Hymenial structures. A Pleurocystidium B Basidium C–M Cheilocystidia. TFB 7272 (TENN-F-53624).

In a paper reporting on North American members of Gymnopus sect. Perforantia, inclusion of European G. perforans could be seen as counterintuitive, but not only is the European entity the name-bringer, but recognition that the North American collections are molecularly separable from the European necessitates a description of the European fungus, as above. As discussed also under G. sequoiae, the G. perforans complex seems composed of subclades with minimal base-pair percent separation. Such is the case with “European G. perforans”, which is separated from sequences from eastern North America by 1.65% bp divergence. Studies by Hughes et al. (2009) and others have shown that, in general, separation at species rank might be expected to be at least 3%. Using this standard, eastern North American collections do not warrant species rank. For this reason, we choose to propose the eastern North American collections at subspecies rank (see below under G. perforans subsp. transatlanticus).

Finland, Etelä-Häme Prov., Lammi, Evo, Kotinen Virgin Forest, northwest part, 13.IX.1994, coll RHP, TFB 7477 (TENN-F-53579). Russia, Leningrad Oblast, Nizhne-Svirskiye Reserve, vic Kovkenitsky, N60°41.691', E33°17.965', 30.VIII.1999, coll N.

Psurtseva, TFB 10826 (TENN-F-58295); same location, N60°36.766', E33°3.482', 29.VIII.1999, coll. RHP, TFB 10643 (TENN-F-58251); Novgorod Reg., Valdai Dist. “Red Hill,” Kasnaya Gorka, N58°06'37", E33°13'07", 21.VIII.2003, coll. RHP, TFB 11629 (TENN-F-59592). SWEDEN, Trollhättan, vic Trollhättan, N58°17'01", E12°17'15", 22.IX.1991, coll SA Gordon, TFB 4721 (TENN-F-50318); same location, N58°17'01", E12°17'15", 22.IX.1991, coll SA Gordon, TFB 4722 (TENN-F-50319); Uppland, vic. Uppsala, Gottsundabergen, N59°48'27", E17°37'24", 7.IX.1994, coll. S-G. Ryman, TFB 7272 (TENN-F-53624).

Canada, Quebec, Quebec-Montmorency Co., vic. Beauport, “Camping Municipal de Beauport,” 46°54.022'N, 71°10.507"W, 29.VII.2006, coll E. Lickey, TFB 13319 (TENN-F-61587).

trans- = Latin, on the other side of; -atlanticus = Atlantic Ocean. Referring to distribution across the Atlantic Ocean from the typical subspecies.

Differing from typical subspecies as follows: 1) basidiomata diminutive, generally shorter and slenderer; 2) spores, pleurocystidia, basidia smaller than typical; 3) cheilocystidia infrequent, smaller and less differentiated; 4) fruiting on fallen Picea, Abies and Tsuga needles;5) separable by ITS sequence from European G. perforans.

Basidiomata (Figs 38, 39A) diminutive, marasmioid, often in troops, usually attached to individual conifer needles. Pileus 3–14 mm broad, strongly convex early, expanding to plano–convex, often with somewhat depressed disc, sometimes with small conical umbo, matt to minutely pruinose (especially near margin), occasionally sulcate–striate but usually not so; disc “sayal brown” 6C5, “tawny” 6C6, “ochraceous tawny” 6C5–6, “tawny olive” 6D5–6, “light ochraceous buff” 5A4, “army brown” 8D5, “tilleul buff” 7B2, paler and nearer to “pale ochraceous buff” 4A2, “light ochraceous buff” 5A4, occasionally yellowish near “antimony yellow” 4B6; limb and margin “pale cinnamon pink” 5A2, “pinkish buff” 6A2, “pale pinkish buff” 6A2 near “tilleul buff” 5B3, “pale olive buff” 3B2 to off-white, fading gradually through maturity, finally often off-white overall; context relatively thick on the disc, elsewhere thin and membranous. Lamellae adnate to subdecurrent, distant, arcuate (not ventricose), occasionally forked but with no anastomosis or interveining, with or without pseudocollarium, thickish, narrow (usually less than 1 mm broad), total lamellae 17–23, through lamellae 8–11, often concolorous with pileus, “pinkish buff” 5A3, “pale pinkish cinnamon” 6A2, “light buff” 3A2, “vinaceous buff” 9B2, “light ochraceous buff” 5A4, “tilleul buff” 7B2, after storage often exhibiting a slight blush of cantaloupe (“warm buff” 3A4 to “orange buff” 3A5) as necropigment; lamellar edge entire, not marginate; lamellulae in one rank, poorly developed. Stipe (10–)12–28 × 0.3–1 mm, in two breadth classes (0.3–0.7 mm vs 0.8–1 mm broad, with the latter not always part of collection) but not in length, terete when fresh, ridged and compressed when dry, subinsititious, apex concolorous with lamellae (darker within pileus), a dull reddish brown (near “russet” 7D6), downward soon “mummy brown” 6F8, “bone brown” 7F8, “soot black” 11F3, “chaetura black” 2F3 to visually black, vestured overall (minutely barbed); apical vesture hyaline (appearing white), wispy, downward gathered into minute synnemata and thus appearing barbed, dark brown; surface sometimes bearing small elliptical scars with fimbriate borders, and rarely with rudimentary, acute-tipped rhizomorphs <1 mm long; base subinsititious (with delicate thatch of spikes). Rhizomorphs (Fig. 39B) of two types: 1) –4 × 0.2–0.7 mm, black, typically curly and branched, often with numerous individuals arising from a single needle, coarse for their length, extensively colonizing the undersides of needles; and 2) –0.5 × 0.05–0.1 mm, black, extremely fine, unbranched, often densely gregarious. Odor negligible to fetid (occasionally weakly alliaceous): taste moderately alliaceous.

Figure 37. 

Gymnopus perforans. Caulocystidia. A–F Full-size setoid caulocystidia G, H Small individual caulocystidia. Standard bars = 10 µm. TFB 7477 (TENN-F-53579).

Figure 38. 

Gymnopus perforans subsp. transatlanticus. Basidiomata. Standard bars = 10 mm.

Figure 39. 

Gymnopus perforans subsp. transatlanticus. A Basidiomata B Rhizomorphs C Basidiospores. Standard bars: A = 10 mm; B = 2 mm; C = 5 µm.

Fruiting in troops on individual dead conifer needles, chiefly Picea, Tsuga and Abies (Abies amabilis, Abies balsamea, Abies fraseri, Pices glauca, Picea rubens, Tsuga canadensis, Tsuga heterophylla and other conifer needles; from high-altitude spruce/fir “islands” of southern Appalachian Mountains to Canadian Shield. (for forms fruiting on dead deciduous leaves, see under G. foliiphilus); late spring (June, southern habitats) to mid-Autumn (September).

Pileipellis involved in a slime matrix with much minute debris, composed of the following elements; 1) pileal hairs (Fig. 40) –80 × 4–7 µm diam, erect hyphal tips, often arising in juxtaposition to a clamp connection, embedded in slime matrix, often subcapitulate; and 2) repent hyphae (Fig. 41A, B) 3.5–10 µm diam, firm-walled, mostly smooth, occasionally encrusted with small scabs with raised profile calluses, conspicuously clamped but with common secondary septa (Fig. 41C, D), subhyaline individually. Pileus trama hyphae loosely interwoven, involved in a slime matrix, conspicuously clamped; lamellar trama loosely interwoven, involved in slime sheaths; hyphae 3–5 µm diam, firm-walled but wall swelling to 1 µm thick, conspicuously clamped. Pleurocystidia (Figs 42, 43) 22–32 × 5–9 µm, fusiform, narrowly fusiform, to irregularly and asymmetrically so with rounded to acute apex, conspicuously clamped, often easily disarticulated; contents distinctly partitioned (Fig. 43) to not so (Fig. 42). Basidioles clavate; basidia (Figs 44, 45A) 20–29 × 5–10 µm, clavate to subcapitulate, 4-sterigmate, conspicuously clamped; contents heterogeneous; effete basidia and pleurocystidia commonly evacuated but not collapsing (“husking”); subbasidial cells lobate by proliferating clamp connections. Basidiospores (Fig. 39C) (6–)6.5–9 × 3–4.5 µm (Q = 1.56–2.50; Q^m = 1.86; L^m = 7.13 µm). ellipsoid, marasmioid, thin-walled, smooth, inamyloid. Cheilocystidia (Fig. 45B–H) absent or widely scattered, 25–48(–50) × 8–19(–23) µm, broadly clavate, obpyriform, occasionally irregularly lobed, without setulae, firm- to thick-walled (wall –0.7 µm thick, hyaline), sometimes easily disarticulated, subrefringent (PhC), obscurely clamped, only occasionally extending beyond hymenial structures; contents homogenous to heterogeneous (with several globose inclusions; PhC), occasionally with one or more small refractive guttules. Caulocystidia from stipe apex –50 × 6–9 µm, arising as side branches of surface hypha, thick-walled (wall –2 µm thick), refringent, hyaline, often secondarily septate. Caulocystidia from lower stipe (Fig. 46) –180 × 8–11 µm, arising as side branches of stipe surface hypha; basal cell usually swollen somewhat, deep brownish yellow (PhC), producing a thick-walled (wall 1–1.5 µm thick) shaft, gradually tapered distally, often constricted within and occasionally with small side lobe, doubtfully secondarily septate, brittle (although usually not straight); pigmentation equally distributed throughout shaft (not congested apically).

Figure 40. 

Gymnopus perforans subsp. transatlanticus. Pileal hairs. Standard bars = 10 µm. TENN-F-29277.

Figure 41. 

Gymnopus perforans subsp. transatlanticus. Pileipellis elements. A Slender hyphae with thin slime sheath and vague incrustation B Stouter hypha with conspicuous clamp connections C, D Pileipellis hyphae with secondary septa B–D Note multigranular gelatino-mucoid matrix. Standard bars = 10 µm. AV 11.06.11.

Figure 42. 

Gymnopus perforans subsp. transatlanticus. Pleurocystidia. Standard bars = 10 µm. AV 11.06.11.

Figure 43. 

Gymnopus perforans subsp. transatlanticus. Pleurocystidia. Note conspicuously partitioned contents. Standard bars = 10 µm. TENN-F-29277.

Figure 44. 

Gymnopus perforans subsp. transatlanticus. Basidia. Standard bars = 10 µm. AV 11.06.11.

Figure 45. 

Gymnopus perforans subsp. transatlanticus. A Basidium B–H Cheilocystidia. Standard bars = 10 µm. TFB 14395 (TENN-F-69056).

Figure 46. 

Gymnopus perforans subsp. transatlanticus. Caulocystidia. Standard bars = 10 µm. TFB 14350 (TENN-F-69000).

Based on direct comparison between European and northeastern North American specimens, basidiomatal dimensions seem to differ. American basidiomata are, in general, shorter with smaller pilei than European basidiomata, but basidiomata of the American collections seem distributed in two size classes, the larger and more robust of which are commensurate with Europe basidiomata.

A limited phylogenetic tree based on ITS sequences (see Fig. 87 below), sequences from northeastern North American collections clearly form a clade separate from sequences from Europe (including Scandinavia). Base-pair separation between these two clades is 1.65%. Although heterogeneous, base-pair separation among northeastern North American sequences is 0.71%. Altogether, collections from Europe and North America are not considered to represent species-rank separation, but proposal of a subspecies seems warranted. Separation of all G. perforans sequences from a clade containing G. sublaccatus and G. sequoiae is more robust (2.30% see Fig. 87 below) but both G. sequoiae and G. sublaccatus fall within North American G. perforans. For more on this situation, see discussion under G. perforans.

In eastern North America, characters which separate G. foliiphilus fruiting on dead deciduous leaves, from G. perforans subsp. transatlanticus fruiting on conifer needles: 1) G. foliiphilus exhibits stipe vesture between villose and barbed, but G. perforans subsp. transatlanticus is distinctly spiked/barbed; 2) pileal hairs in G. foliiphilus are stouter than those of G. perforans subsp. transatlanticus, and secondarily septate (this may be an artifact); and 3) pileus surface hyphae of G. foliiphilus are often encrusted in small scabs, while those of G. perforans subsp. transatlanticus are generally smooth.

In examining numerous specimens eventually accepted as G. perforans subsp. transatlanticus, it was unforeseen that a large number of specimens originally fruited on dead Tsuga needles, a host reported as rare by Antonín and Noordeloos (2010), although this is the case in Europe. Tolerance of this substrate affords the opportunity to subsist at lower elevations (and therefore at higher temperatures) in the southern Appalachian Mountains where spruce/fir forests exist only at the highest elevations (and therefore the coldest temperatures). In the future, it will be interesting to see if this condition extends across North America to the Pacific Northwest, where the spcies of spruce, fir, and hemlock are all different from eastern North American taxa.

In G. perforans subsp. transatlanticus, cheilocystidia may be confused with inflated basidia and/or pleurocystidia. Some shapes of clearly swollen elements assumed to be cheilocystidia are suggestive of sterigmata (but always two – four were not seen), and occasional objects of cheilocystidial size are shaped like fat pleurocystidia (broadly fusiform). Repeated preparations from a single pileus also showed numerous cheilocystidia.

Rhizomorphs are here described as of two types. The coarser of these are present in virtually every specimen observed, while the extremely fine type were seen in perhaps 25% of the specimens. It can be doubted that the second type actually belong to subsp. transatlanticus, but without some proof, they must be described as present.

Canada, New Brunswick, vic. Alma, Fundy Prov. Park, Maple Grove Back Road, N95°35.368', W64°59.013', 20.IX.2013, coll RHP, TFB 14377 (TENN-F-69042); same location East Branch Trail, N45°38.587', W65°06.937', 25.IX.2013, coll Gary Samuels, TFB 14392 (TENN-F-69056); same location, East Branch Trail, N45°38.587', W65°06.937', 25.IX.2013, coll Gary Samuels, TFB 14400 (TENN-F-69064); same location, East Branch Trail, N45°38.587', W65°06.937', 25.IX.2013, coll RHP, TFB 14395 (TENN-F-69056); Newfoundland & Labrador, Labrador, Benedict Mts., 31.VII.2010, coll A. Voitk (as M. androsaceus), AV10.07.31 AV03; Newfoundland, Codroy,Gillis Cabins, N47°52'55", W59°23'36", 11.VI.2011, coll A. Voitk (as Mi. perforans), AV 11.06.11AV01 (TENN-F-69046); Newfoundland, Gros Morne Nat. Park, Stuckless Pond, N49°25'55", W57°43'38", 18.IX.2010, coll A. Voitk, AV 10.09.18 AV 14; Nova Scotia, Wolfville, Acadia University nature trail, 1.VIII.1992, coll SA Gordon, TFB 5017 (TENN-F-51431); Cape Split area, 45°20'02" N, 64°30'02"W, 3.VIII.1992, coll. S.A. Gordon, TFB 5029 (TENN-F-51440); Quebec, Quebec-Montmorency Co., vic. Beauport, “Camping Municipal de Beauport,” 46°54.022'N, 71°10.507"W, 29.VII.2006, coll E. Lickey, TFB 13319 (TENN-F-61587; holotype). United States, Connecticut, Hartford Co., vic. Colchester, Salmon River State Forest, N41°36'46.77", W72°25'27.43", 31.VIII.2013, coll. Sandy Scheine, TFB 14327 (TENN-F-68179); New York, Cortland Co., vic. Marathon, Hoxie Gorge Preserve, Lower Gorge Trail, N42°32.871', W76°4.707', 3 .IX.2013, coll RHP, TFB 14348 (TENN-F-68198); Ulster Co., Frost Valley, Spring Ridge Trail, N41°50'20.7", W74°30'19.9", 9.IX.1989, coll SA Gordon, TFB 2146 (TENN-F-49764); same location, N41°50'20.7", W74°30'19.9", 9.IX.1989, coll SA Gordon, TFB 2151 (TENN-F-49782); North Carolina, Macon Co., vic. Franklin, Standing Indian Campground, 13.VI.1992, coll SA Gordon, TFB 4913 (TENN-F-51231); Highlands, 9.VII.1966, coll L.R. Hesler (as Marasmius sp.), TENN-F-29279; Tennessee, Blount Co., GSMNP, Indian Gap. Appalachian Trail toward Newfound Gap, near top of ridge, 30.VI.2006, coll RHP & KWH, TFB 13121 (TENN-F-61211); GSMNP, Cades Cove, N35°35', W83°50', 4.VIII.1962, coll & det L.R. Hesler (as Marasmius sp.), TENN-F-24923; Sevier Co., GSMNP, Spruce-fir Nature Trail, Clingman’s Dome Rd., N35°34'40.5", W83°28'47.4", 21.VII.1989, coll SA Gordon, TFB 2114 (TENN-F-49259).

United States, North Carolina, Macon Co., vic. Highlands, Blue Valley Campground, N35°00'45.23", W83°09'29.33", 11.VIII.2014, coll KWH, TFB 14511 (TENN-F-69206).

pino- = referring to the genus Pinus; -phil = to love, referring to habitat on the needles of Pinus.

1) Fruiting habit on needles of Pinus strobus in eastern North America; 2) pileipellis structure including occasional diverticulate hyphae and broom cell-like hyphal termini; 3) cheilocystidia of the siccus-type; 4) basidiomata with long, slender stipes and small pilei; 5) stipe glabrous–shining; 6) rhizomorphs hair-like, black, independent of basidiomata.

Basidiomata (Fig. 47) slender, with long stipe and small pileus. Pileus 3–9(–12) mm broad, convex becoming shallowly convex, sometimes centrally depressed, matt, pebbled, subtly sulcate–striate; disc “fawn color” 7C5, 6E7, “vinaceous fawn” 8B4, “ocher red” 9C6 to “brick red” 8D8, outward “avellaneous” 7B3, 6D4–6, “vinaceous fawn” 8B4, “vinaceous buff” 9B2 “buff pink” 7A4, drying nut brown (“tawny olive” 5C5). Lamellae adnexed to adnate, sometimes pseudocollariate (especially in drying), thickish, not ventricose, with little evidence of anastomosis, off-white to “tilleul buff” 7B2, 6C3–4; lamellulae always very short, hardly ventricose. Stipe 27–45(–60) × 0.4–1 mm, stiff, terete, equal, not vestured, glabrous and often shining, insititious, apically concolorous with lamellae (“tilleul buff” 7B2 to “army brown” 8D5), soon “clove brown” 6F5 to black and remaining so, stuffed; stipe medulla white. Rhizomorphs (Fig. 47) –18 × 0.2–0.3 mm, hair-like, curly, black, usually unbranched, arising separately from basidiomata. Odor negligible; taste negligible.

Figure 47. 

Gymnopus pinophilus. Basidiomata on needles of Pinus strobus. Standard bars = 20 mm. Above: TFB 14511. Below: TFB 14059 (TENN-F-67804).

Gregarious on dead needles of Pinus strobus in eastern North America; summer.

Pileipellis involved in a heterogeneous slime matrix, composed of the following elements: 1) repent hyphae radially oriented, 2.5–6 µm diam, firm- to thick-walled (wall –0.7 µm thick, hyaline), firm-walled with a very thin mucoid sheath, conspicuously clamped, weakly to strongly encrusted, with encrustation appearing as vague, subtle stripes or rings with flake-like profile calluses (Fig. 48A) (lying on the thin mucoid sheath); 2) repent hyphae 4–7.5 µm diam, with gelatinized wall (Fig. 48B) (wall –1.5 µm thick, with weak encrustation); 3) occasional repent hyphae 3.5–5.5 µm diam, firm-walled, clamped, diverticulate; diverticula 2–6 × 1–1.5 µm (Fig. 48C), usually dichotomous, not refringent; and 4) common broom cell-like cells, stalked, arbuscular (stalked without distal inflated portion), branched 1–2 times, surmounted by setulae; setulae digitate, often knobby, usually dichotomous, –6 × 1–1.5 µm, sometimes subrefringent (PhC); broom cell-like structures (Fig. 49) easily gelatinizing from base and often visible only as setulae (Fig. 49A,D). Pileus and lamellar tramae loosely interwoven, with heterogeneous slime matrix. Hymenophore (in KOH) involved in a heterogeneous mucoid matrix with copious debris (collapsed spores, bits of effete basidia, etc.). Pleurocystidia (Fig. 50A–D) 24–32 × 5.5–8 µm, fusiform, conspicuously clamped; contents homogeneous, often with poorly partitioned contents. Basidioles clavate; basidia (Fig. 50E–H) 23–31 × 5.5–7 µm, clavate, clamped, 4-sterigmate; contents multigranular. Basidiospores (Fig. 51) probably dimorphic: 1) (3.5–)4–6(–6.5) × (2.5–)3–4(–4.5) µm (Q = 1.11–2.17; Q^m = 1.57; L^m = 5.15 µm); 2) (5.0–)6–8(–8.5) × (2.5–)3–4.5 µm (Q = 1.38–2.40; Q^m = 1.96; L^m = 7.19 µm), broadly ellipsoid, subcylindric to elongate–lacryiform, hardly flattened adaxially, thin-walled, smooth; contents homogeneous to a few scattered minute, refringent granules. Lamellar edge involved with heterogeneous mucoid matrix; cheilocystidia (Fig. 52) 20–34 × 6–9 µm, scattered to common, similar to pileipellis element, broom cell-like, easily gelatinizing, stalked (stalk 2.5–3 µm diam), arbuscular, terminating in a very complex tuft of setulae; setulae –8 × 1–1.5 µm broad), refringent (PhC), digitate, often dichotomous. Outer stipe medullary hyphae involved in a slime matrix, skeletal–generative, 5–12 µm diam, thick-walled (wall –2.0 µm thick), conspicuously clamped, occasionally producing flagelliform hyphal tips –250 × 1–1.5 µm diam, thick-walled near origin, thin-walled apically; stipe cortex hyphae 4–7 µm diam, thick-walled (wall occluding cell lumen), strongly pigmented, non-dextrinoid; surface hyphae minutely roughened. Caulocystidia not observed.

Figure 48. 

Gymnopus pinophilus. Pileipellis elements. A Encrusted hyphae showing crust material in slime sheath B Hyphae with gelatinized walls C Diverticulate hypha showing dichotomous diverticula. Standard bars = 10 µm. DED 4491 (TENN-F-54665).

Figure 49. 

Gymnopus pinophilus. Broom cell-like hyphal termini of pileipellis. A Diverticula in slime matrix B, C Individual elements D Gelatinizing broom cell-like termini. Standard bars = 10 µm. DED 4491 (TENN-F-54655).

Figure 50. 

Gymnopus pinophilus. Hymenial elements. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. DED 4491 (TENN-F-54665).

Figure 51. 

Gymnopus pinophilus. Basidiospores. Standard bars = 5 µm. A = TFB 14097. B = TFB 10459 (TENN-F-67804).

Figure 52. 

Gymnopus pinophilus. Cheilocystidia. A Cluster of two cheilocystidia B–H Individual cheilocystidia. Standard bars = 10 µm. TFB 7234 (TENN-F-53553).

Desjardin (1989) and Gordon (1994) treated G. pinophilus as an unnamed subset of Ma. androsaceus. The latter was perceived as fruiting on three substrata, hardwood leaves, needles of Picea/Abies, and needles of Pinus. Gordon and Petersen (1997) understood that M. androsaceus s.l. in eastern North America could be divided into three sexually interINcompatible groups, one of which (mating group III) was represented by only a single collection (TFB 5627 TENN-F-53488 from Idaho). With subsequent collecting and DNA sequence production, this collection was determined as G. pinophilus. Mata et al. (2007) showed that Marasmius sect. Androsacei (represented by M. androsaceus) was actually embedded within Gymnopus, and Noordeloos and Antonín (2008) formally transferred the section as Gymnopus sect. Androsacei. Furthermore, Desjardin (1990) and Desjardin and Horak (1997) reported considerable morphological and phenological variation within the North Temperate Androsacei, in part by circumstantial evidence, based on the data above. Present phylogenetic analyses now show that the small alliance of G. pinophilus and G. ponderosae (Gordon’s mating group III) is more closely related to the Mi. perforans complex than to the Ma. androsacei complex.

Observation on TFB 14097 (TENN-F-67846) revealed two sets of dimensions of spores, and TFB 10459 showed that these spores are formed by basidia in the same hymenium – thus spores must be judged as dimorphic. One cause for this might be 2- versus 1-nucleate condition of individual spores, but all seem to be produced by 4-sterigmate basidia.

Observations of pileipellis of TFB 10459 showed that broom cell-like hyphal termini are ephemeral – apparently they wash or gelatinize away, so when they were absent from some pilei of various collections, the strongly ornamented repent hyphae remained.

Recently, TFB 14097 was established in dikaryron and monokaryon cultures and a self-cross was performed (see Gordon and Petersen 1997 for methods). From an assay of 20 putative single-basidiospore isolates (SBIs), 12 clampless putative monokaryons were selected and paired in all combinations. As expected, a tetrapolar mating system was revealed. A¹B¹ = 5*, 6; A²B² = 1*, 3, 4, 8; A²B¹ = 2*, 7, 9, 11, 12; A¹B² not represented in the sample (* indicates a tester strain). Most growth of donors was submerged (except for structures noted below). Most pairings exhibited very subtle “barrage” or “flat” contact zones, most easily envisioned with the naked eye against back-lighting. Barrage = somewhat congested growth in “mustaches” (not within contact zone). Flat = slightly congested hyphae on both sides of a lightly overgrown crevasse.

On malt extract (Difco, 15 g/L) agar (Difco, Bacto, 20g/L) most SBIs produced: 1) pure white spherical hyphal masses (<1– 2 mm diam; appearing as minute “snowballs”). These structures are narrowly attached to the agar-surface mycelium and are composed of loosely interwoven hyphae with a mucoid medulla, pure white inside and out, easily crushed (like a cotton ball); and 2) minute ganglia of hyphae scattered over agar surface in mucoid matrix. SBIs include two hyphal types: 1) 4–6.5 µm diam, thin-walled, frequently septate, hardly inflated; and 2) roughly arbuscular, 1.5–2.5 µm diam, thin-walled, as though a ramealis structure but with branches longer, apparently opposite or sub-opposite.

Canada, Nova Scotia, Kejimkujik National Park, Grafton Lake Loop Trail, 5.VIII.1992, coll S.A. Gordon, TFB 5034 (TENN-F-53487). United States, New York, Franklin Co., Malone, Franklin Academy High School nature trail, 14.VII.1992, coll Scott Gordon, TFB 4975 (TENN-F-53478). North Carolina, Haywood Co., GSMNP, Cataloochee Cove, Schoolhouse area, N35°37'43.93", W83°06'44.18", 5.IX.1992, coll RHP, TFB 5547 (TENN-F-52480); same location, 9.IX.1987, coll DE Desjardin, DED 4491 (TENN-F-54665); Jackson Co., vic. Cashiers, Panthertown Valley, N35°09'34.63", W83°00'35.75", 24.VI.1992, coll. JE Johnson, TFB 6311 (TENN-F-53128); Macon Co., vic. Highlands, Cliffside Lake, end of road, N34°04.749', W83°14.150', 4.VIII.2012, coll. KWH, TFB 14097 (TENN-F-67846); Cliffside Lake Campground, N35°04'44.92", W83°14'12.90", 3.IX.1986, coll D.E. Desjardin. DED 4107 (TENN-F-54643; SFSU); Highlands, Highlands Biological Station, N35°03'09.72", W83°11'19.98", 9.VIII.1966, coll LR Hesler, LRH 29279 (TENN-F-29279); vic. Highlands, Bull Pen Rd., Ellicott Rock Wilderness trailhead, N35°01.010', W83°08.190', 20.VII.2011, coll RHP, TFB 13913 (TENN-F-65808); Horse Cove, Walking Stick Rd., “Double Bridges,” N35°00.983', W83°09.619", 31.VII.2012, coll RHP, TFB 14059 (TENN-F-67804); Bull Pen Rd., Ellicott Rock Wilderness trailhead, N35°01.010', W83°08.190', 20.VII.2011, coll RHP, TFB 13913 (TENN-F-65808); Nantahala Nat. For., Blue Valley Campground, N35°00'45.23", W83°09'29.33", 11.VIII.2014, coll KWH, TFB 14511 (TENN-F-69206; holotype); vic. Franklin, Standing Indian Campground, 19.VII.1994, coll DBG Nichol, TFB 7629 (TENN-F-53659). South Carolina, Oconee Co., Burrell’s Ford campground, N34°58.408', W83°06.726, 12.VIII.2014, coll Highlands Biol. Station class, TFB 14517 (TENN-F-69212).

United States, California, Humboldt Co., Rte 299, Grey’s Falls Campground, N40°54.422', W123°42.420', 16.XI.1996, coll RHP, TFB 9020 (TENN-F-55669).

ponderosae = Latinized, referring to Pinus ponderosa, fruiting habitat of the species.

1) fruiting on dead needles of Pinus ponderosa; 2) pileipellis with diverticulate hyphae and broom cell-like hyphal termini; 3) pileipellis broom cell-like hyphal termini and cheilocystidia arbuscular, with conspicuous stalk but hardly swollen distal portion; 4) rhizomorphs conspicuous, hairlike, curly, unbranched, –35 × 0.1–0.4 mm; 4) spores perhaps dimorphic.

Basidiomata (Figs 53, 54A) scattered on needle duff, usually solitary on individual needles, occasionally two per needle. Pileus 3–13 mm broad, applanate but with wide inflexed margin, occasionally slightly depressed, delicately tuberculate, not umbonate, subtly sulcate–striate, matt; disc brown (“Mars brown” 8F7, “bister” 5F8), grayish avellaneous to dull deep avellaneous; limb light brown (“sayal brown” 6C5, “tawny olive” 5C5) to near “tilleul buff” 7B2. Lamellae adnexed to adnate, subdistant, not anastomosing, not pseudocollariate but seceding and appearing pseudocollariate after drying, subventricose, total lamellae (Fig. 54B) = 22–28, through lamellae = 13–15, (fresh) light brownish orange (6C4–3) to avellaneous, (dry) gray with hint of olive (“deep olive buff” 3C3, “light grayish olive” 30B2); lamellulae in one rank, rudimentary. Stipe 24–40(–60) × 0.5–0.8 mm broad, glabrous–shining over all, tapering gradually downward from mid-section to base, insititious, upward “avellaneous” (concolorous with lamellae, 7B3), socket at lamellar junction dark brown, downward black (“fuscous black” 6F4, “chaetura black” 2F3). Rhizomorphs (Fig. 54A) common, –35 × 0.1–0.4 mm, hair-like, black, glabrous–shining, kinked but not branched, attached to substrate with basal pad, sparse, not attached to basidiomata but closely associated. Odor negligible; taste negligible.

Figure 53. 

Gymnopus ponderosae. Basidiomata on dead needles of Pinus ponderosa. Standard bar = 10 mm. Mushroom Observer CA391914.

Figure 54. 

Gymnopus ponderosae. A Basidioma and rhizomorphs B Lamellar schematic C Basidiospores. Standard bars: A = 10 mm; C = 5 µm. B not to scale. TFB 9020 (TENN-F-55669).

Fruiting on dead needles, twigs and debris of Pinus ponderosa (three-needle pine); distribution probably following distribution of Pinus ponderosa (at least California, Idaho, Washington); late autumn.

Pileipellis constructed of three elements involved in a minimal slime matrix over central areas: 1) hyphae 3–7 µm diam, firm- to thick-walled [wall 0.7–1 µm thick, subhyaline singly, pigmented dull yellow–olive brown in mass (PhC)], conspicuously clamped, strongly encrusted (Fig. 55A, B); crust material in thick scabs with profile calluses –1.5 µm thick or as annular ornamentation (Fig. 52B) or with profile calluses –1 µm thick or minute “flakes” adherent to a slime sheath; 2) free-form hyphae 4–6 µm diam, tibiiform to meandering, thin- to firm-walled (wall –1 µm thick, hyaline), often diverticulate with a combination of lobes and diverticula (Figs 55C,D, 56A, B); diverticula –8 × 1–1.2 µm, digitate, often dichotomous, subrefringent; lobes –8 × 2–3 µm at base, fin-shaped (triangular), non-refringent; and 3) broom cell-like termini (Fig. 56C–F), arbuscular, stalked (stalk 4–13 × 3–5 µm), hardly or not inflated distally, branched in 1–2 ranks, beset with setulae 3–12 × 1–1.5 µm, digitate, often somewhat catenulate, often dichotomous, subrefringent, apparently free (not in slime). Pileus trama loosely interwoven; hyphae 2.5–6.5 µm diam, firm-walled, conspicuously clamped, weakly encrusted in “flakes.” Pleurocystidia (Fig. 57A–D) 24–37 × 7–9.5 µm, fusiform, conspicuously clamped; contents homogeneous, often with partitioned contents at apex. Basidioles clavate to ampulliform; basidia (Fig. 57E–H) (25–)32–38 × (7–)8–10 µm, clavate, often with slight constriction to produce capitulate morphology, clamped, 4-sterigmate; contents minutely heterogeneous. Basidiospores (Fig. 54C) possibly dimorphic: 1) (5–)5.5–7(–8.5) × (3–)3.5–5 µm (Q = 1.11–2.43; Q^m = 1.67; L^m = 6.2 µm); 2) (6.5–)7–8.5 × 3.5–4.5 µm (Q = 1.75–2.43; Q^m = 2.01; L^m = 7.7 µm), broadly ellipsoid to rotund–ellipsoid, pip-shaped, smooth, thin-walled, inamyloid; contents minutely granular. Cheilocystidia (Fig. 58) 18–35 × 10–17 µm, broom cell-like, stalked [stalk (9–)15–27 × 4–5 µm, sometimes minutely roughened], without dilated distal portion, branched in 1–2 ranks, obscurely clamped; setulae –8 × 1–1.5 µm, often gnarled and/or branched, subrefringent (PhC). Stipe medullary hyphae 3–10 µm diam, hyaline, thick-walled (wall –2 µm thick), obscurely clamped, with minimal slime matrix, strictly parallel. Stipe cortical hyphae 4–10 µm diam, thick-walled [wall –2 µm thick, pigmented yellow brown (PhC)], obscurely clamped; surface hyphae encrusted in minute scabs, making surface cells appear rough. Caulocystidia not observed.

Figure 55. 

Gymnopus ponderosae. Pileipellis structures. A Strongly encrusted hypha B Hypha with annular encrustation C Thick-walled diverticulate hypha D Thick-walled, lobate diverticulate hypha. Standard bars = 10 µm. TFB 9020 (TENN-F-55669).

Figure 56. 

Gymnopus ponderosae. Pileipellis structures. A, B Diverticulate hyphae C–F Broom cell-like hyphal termini. Standard bars = 10 µm. TFB 9020 (TENN-F-55669).

Figure 57. 

Gymnopus ponderosae. Hymenial structures. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. TFB 9020 (TENN-F-55669).

Figure 58. 

Gymnopus ponderosae. Cheilocystidia. A Cluster of cheilocystidia B–I Individual cheilocystidia. Standard bars = 10 µm. TFB 9020. (TENN-F-55669).

Collections accepted as four separate species (G. “scoticus” nom. prov., G. “adventitius” nom. prov., G. pinophilus, G. ponderosae) fruit on dead needles of Pinus spp. Of these, the former two are found in Gymnopus sect. Androsacei, while the latter two belong in sect. Perforantia. Both G. ponderosae and G. adventitius fruit on needles of Pinus ponderosae and superficially resemble one another. Both exhibit black, glabrous–shining stipes and dark brown pilei, but the pileipellis of G. ponderosae comprises repent, encrusted hyphae, diverticulate hyphae and broom cell-like hyphal termini, while G. adventitius pileipellis does not show diverticulate hyphae and broom cell-like hyphal termini. Moreover, stipes of G. adventitius often produces adventitious rhizomorphic structures from wounds, not seen in G. ponderosae.

The small clade comprising G. pinophilus and G. ponderosae is found sister to that of core sect. Perforantia. Pileipellis construction, which includes diverticulate hyphae and pileipellis “broom cells” is characteristic of a “Rameales structure,” indicative of sect. Androsacei, but hardly that of traditional Perforantia, although involved in minimal slime. Marasmioid cheilocystidia also separate this pine-loving alliance from sect. Perforantia. Rhizomorphs, while not typically as long as basidiome stipe, are common, obvious and arise from the same needles as basidiomata.

TENN53488-TFB5627 (Idaho, G. ponderosae) was the only representative of mating group III of “Marasmius androsaceus” by Gordon (1994) and Gordon and Petersen (1997). The taxonomic concept of the latter, however, was broad ([see Desjardin and Petersen (1989); Gilliam (1976)], including two separate taxa now known to be members of sect. Androsacei (as well as G. ponderosae in sect. Perforantia), and perhaps other organisms for which haploid isolates were unavailable for pairing experiments.

The possibility of dimorphic spores is reported here also for G. pinophilus. Although reported for other mushroom groups (notably hygrophoroids) the cause of this phenomenon is not known in the group treated here. Basidia are uniformly 4-spored and basidia are conspicuously clamped, indicative of normal nuclear number and behavior.

United States, California, Humboldt Co., Rte 299, Grey’s Falls Campground, N40°54.422', W123°42.420', 16.XI.1996, coll RHP (as Marasmius sp.), TFB 9020 (TENN-F-55669; holotype). Idaho, Bonner Co., vic. Priest River, Priest River Experimental Forest,25.IX.1992, coll RHP (as Marasmius sp.), TFB 5627 (TENN-F-53488); same locale, same date, coll RHP (as Marasmius sp.), TFB 5633 (TENN-F-52411); Valley Co., vic. McCall, N44.911 W116.097, 16.IX.1977, coll. SD Libonati-Barnes (as Marasmius androsaceus), SDLB 1461 (WTU-F-9083). Washington, Pend Oreille Co., Roosevelt Grove, N48°45'15.89", W117°03'30.33", 29.VI.1993, coll L. Norvell (as Marasmius androsaceus), LN93.06.29-16 (WTU-F-8918).

Costa Rica, Prov. San José, San Gerardo de Dota, Albergue de Montaña, Savegre, 9°33'2" N, 83°48'27"W, 21.VI.1995, coll RHP, TFB 7879 (TENN-F-53736).

Latinized; resembling Pyracantha, referring to spike-like setulae on pileipellis broom cell-like hyphal termini.

1) A morpho-species (no DNA sequences available); 2) broom cell-like hyphal termini of pileipellis coarsely thorny; 3) subhymenial hyphae with slime sheath and flake-like encrusting material; 4) rhizomorphs well-developed, curly, sparingly branched; 5) stipe minutely pruinose overall; 6) fruiting on sclerophyllous deciduous leaves.

Basidiomata (Fig. 59A) diminutive, arising independent from but gregarious with rhizomorphs. Pileus 3–6 mm broad, strongly convex and in one case slightly umbonate, matt, minutely tuberculate, subtly sulcate or not so; disc “wood brown” 7C4; limb and margin “vinaceous buff” 9B2 to ”tilleul buff” 7B2. Lamellae adnexed (Fig. 59A), usually weakly pseudocollariate, thickish, subdistant, narrow (<1 mm broad), slightly ventricose, without anastomosis or interveining, without necropigment, total lamellae = 31–33, through lamellae = 10–11, “vinaceous buff” 9B2; lamellar edge entire, not marginate. Stipe 24–38 × 0.3–0.5 mm, appearing glabrous–shining but under magnification (35×) very finely pruinose over total length (pruinosity hyaline), stuffed (medulla off-white), insititious, apically concolorous with lamellae, downward quickly “army brown” 8D5, downward “Natal brown” 8E6. Rhizomorphs (Figs 59A, 60) common, mostly resupinate on leaf surface (both upper and lower side), 0.1 mm broad (extremely slender), only occasionally branched when resupinate; aerial rhizomorphs – 17 × 0.2–0.3 mm, gyrose to curly in aerial portions and frequently branched, glabrous–shining, now strap-shaped where free (aerial portions compressed in drying, not resupinate portions), functionally black in resupinate portions, brass-brown (reminding of M. “flavipes” nom. prov. from Tasmania) to “bone brown” 7F8 in aerial portions. Taste and odor not recorded.

Figure 59. 

Gymnopus pyracanthoides. A Basidiomata and rhizomorphs B Basidiospores. Standard bars: A = 20 mm; B = 5 µm. TFB 7879 (TENN-F-53736).

Figure 60. 

Gymnopus pyracanthoides. Broad-leafed habitat. Lines indicate resupinate rhizomorphs. Scale = ×1. TFB 7879 (TENN-F-53736).

Known from only the type specimen; fruiting on dead sclerophyllous leaves (probably Quercus) at high elevation; summer.

Pileipellis (Fig. 61) a dense thatch of diverticulate hyphae and hyphal termini; parent hyphae 2–2.5 µm diam, thin- to firm-walled, hyaline, not involved in slime matrix, producing elongate, jointed, digitate side branches 2–7 × 0.8–1.5 µm, rarely but conspicuously clamped. Pileus and lamellar tramal hyphae 2–5 µm diam, firm- to thick-walled (wall –0.7 µm thick, hyaline), producing thin slime sheaths, ornamented with external “flakes” of encrusting material, infrequently but conspicuously clamped. Pleurocystidia (Fig. 62A–D) 21–28 × 5–7 µm, fusiform, with narrowly rounded to acute apex, abundant, thin-walled, conspicuously clamped. Basidioles clavate; basidia (Figs 62E–H, 63F) 19–26 × 6–7 µm, clavate, sometimes very slightly subcapitate, 4-sterigmate, obscurely clamped. Subhymenium often gelatinizing, including bases of basidia and pleurocystidia. Basidiospores (Fig. 59B) 6–7.5 × (3–)3.5–4 µm (Q = 1.63–2.33; Q^m = 1.94; L^m = 6.85 µm), ellipsoid to elongate pip-shaped, smooth, thin-walled, inamyloid. Lamellar edge fertile; cheilocystidia(Fig. 63A–E) scattered, obscure, 22–32 × 5–8 µm, subcapitate to subampulliform, obscurely clamped, hardly projecting beyond basidia, hyaline. Stipe medullary hyphae 3.5–7.5 µm diam, firm-walled, conspicuously clamped, strictly parallel, with minimal slime matrix, hyaline. Stipe cortical hyphae 4–7 µm diam, thick-walled (wall –1.5 µm thick, pigmented), at surface producing side branches as caulocystidia. Caulocystidia (Fig. 64) setoid, –85 × 5–7.5 µm diam, arising as side branches (not clamped), tapering slightly distally, refringent (PhC), in IKI moderately dextrinoid (PhC), yellow-brown (BF).

Figure 61. 

Gymnopus pyracanthoides. Pileipellis elements. A Cluster of broom cell-like hyphal termini B–G Individual broom cell-like hyphal termini. Standard bars = 10 µm. TFB 7879 (TENN-F-53736).

Figure 62. 

Gymnopus pyracanthoides. Hymenial elements. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. TFB 7879 (TENN-F-53736).

Figure 63. 

Gymnopus pyracanthoides. Hymenial elements. A–E Individual cheilocystidia F Basidium. Standard bars = µm. TFB 7879 (TENN-F-53736).

Figure 64. 

Gymnopus pyracanthoides. Caulocystidia. A–D Individual caulocystidia E Caulocystidial turf. Standard bars = 10 µm. TFB 7879 (TENN-F-53736).

Presence of slime sheaths surrounding pileus and lamellar tramae as well as the dimensions and shape of cheilocystidia are characteristic of taxa in sect. Perforantia. Conversely, pileipellis presents a very distinctive thatch of broom cell-like hyphal termini rather than repent, encrusted hyphae in a slime matrix as is seen throughout the section. Such a differentiated pileipellis might also qualify for some infrageneric groups of Marasmiellus but would be expected to be without slime deposition. Unfortunately, DNA sequences could not be produced from the only known collection, so molecular placement remains unknown.

Cheilocystidia in G. pyracanthoides resemble those of G. bulliformis (q.v.). They also resemble those described for G. trabzonensis. They may represent some stage of basidial development because basidioles also are clavate, but basidioles are present throughout the lamellae face, while the structures here described as cheilocystidia are found along the lamellar edge. No other differentiated cheilocystidial structures were observed, but some taxa in sect. Perforantia are known to lack differentiated cheilocystidia.

Slime is produced as a thin sheath surrounding individual hyphae, with ornamentation appearing as flakes riding on the exterior of the gelatinized wall. It also appears to obliterate subhymenium. Likewise, stipe medullary hyphae exist in minimal (but present) slime. There is little evidence of a slimy (or gelatinized) matrix in the pileipellis.

Costa Rica, Prov. San José, San Gerardo de Dota, Albergue de Montaña, Savegre, 9°33'2"N, 83°48'27, 21.VI.1995, coll RHP, TFB 7879 (TENN-F-53736; holotype).

United States, Washington, Grey’s Harbor Co., vic Amanda Park, Forest service Rd. 2140, “Higley’s Swamp,” 16.X.1992, coll RHP & KWH, TFB 5886 (TENN-F-51994).

Lake Quinault and the Quinault River, Washington, the vicinity of the holotype collection.

1) Fruiting on Thuja debris (less on associated Abies and Tsuga needles); 2) rhizomorphs short, stout, arising and terminating in small black pads; 3) stipe fuscous brown-black, vestured, especially downward; 4) pileal hairs often ornamented with annular rings; 5) pileipellis constructed of incrusted hyphae, and hyphae with gelatinizing walls; 6) cheilocystidia not observed.

Description. Basidiomata (Fig. 65A) marasmioid, diminutive. Pileus 3–15 mm broad, convex to plano-convex, matt, very short-striate on downturned margin; disc now near “sayal brown” 6C5, outward now dull cinnamon buff, near “avellaneous” 7B3 to “vinaceous buff” 9B2. Lamellae (Fig. 65A) adnate to adnexed, close to subdistant, seceding somewhat upon drying and then appearing pseudocollariate, subventricose (–1.2 mm broad), thickish, with no anastomoses; total lamellae = 30–32; through lamellae = 11–14, now concolorous with pileus, bleeding slightly from dark brown stipe apex; lamellulae short, in 1–2 ranks; lamellar trama under magnification (40×) appearing glassy (probably gelatinized). Stipe 15–30 × 0.7–1.2 mm, subinsititious, terete, equal, hollow, minutely vestured upward (35×), downward becoming pruinose to minutely hispid, upward “cinnamon” 6B5 to “orange cinnamon” 7B6, soon brown and downward “fuscous black” 6F4 to “chaetura black” 2F3 (not totally black anywhere), remaining darker than lamellae through attachment to pileus flesh; medulla off-white. Rhizomorphs (Fig. 65A) apparently common on Thuja branchlets (incidental on Abies needles), –10 mm × 0.3–0.8 mm (relatively stout for length), arising from a small black pad (and terminating similarly on adjacent needles), commonly branched (usually short, spur branches), colonizing adjacent needles; resupinate black hyphae individual, meandering over substrate surface. Taste and odor not recorded.

Figure 65. 

Gymnopus quinaultii. A Basidioma and rhizomorphs B Basidiospores C Caulocystidia. Standard bars: A = 10 mm; B = 5 µm; C = 20 µm. TFB 5886 (TENN-F-51994).

Fruiting on dead needles of Thuja plicata and Abies grandis; Autumn.

Pileipellis composed of the following elements: 1) pileal hairs (Fig. 66) –100 × 3–4.5 µm, erect, usually more or less straight, firm-walled, often with basal clamp connection, often weakly ornamented, especially in annular rings (Fig. 66D); 2) repent, unoriented, interwoven hyphae 3.5–8 µm diam, thin-walled but wall gelatinizing to 2–3 µm thick (Fig. 67C, D), with vague outer wall, obscurely clamped; and 3) weakly to strongly encrusted hyphae (Fig. 67A, B) 4.5–8 µm diam, thin-walled but semi-gelatinizing; incrustation in scabs and/or vague stripes, with strong but rough profile calluses; narrower hyphae encrusted as though with many minute crystals (not coarse scabs). Lamellar trama loosely interwoven; hyphae 3–5.5 µm diam, firm-walled, clamped. Pleurocystidia (Fig. 68) common, 28–34 × 6–8 µm, fusiform, conspicuously clamped; contents homogeneous or partitioned. Basidioles (Fig. 69A) clavate; basidia (Fig. 69B–D) (25–)30–33(–37) × 6.5–9 µm, clavate, subtly subcapitulate, 4-sterigmate, clamped. Basidiospores (Fig. 65B) (6.5–)7.5–9(–9.5) × (3.5–)4–4.5 µm (Q = 1.63–2.50; Q^m = 1.99; L^m = 7.82 µm), plump–ellipsoid, marasmioid (somewhat tapered proximally), flattened adaxially, smooth, thin-walled, inamyloid; contents vaguely univacuolate (PhC). Cheilocystidia not observed. Stipe medullary hyphae strictly parallel, perhaps coherent, 2–9 µm diam, firm- to thick-walled (wall – 1.0 µm thick, hyaline), conspicuously clamped. Stipe cortical hyphae 4–7 µm diam, strictly parallel, minutely roughened, thick-walled (wall –2 µm thick), obscurely clamped; stipe vesture setoid, hardly gathered into synnemata but usually densely scattered. Caulocystidia (Fig. 65C, 70) –180 × 3.5–7 µm, occurring as side branches of stipe surface hyphae, irregularly rooted, often gnarled at base, tapering distally to rounded apex, thick-walled (wall 1.5–3 µm thick, often occluding cell lumen, dull yellow-ochre, highly refringent), yellowish (PhC), occasionally internally clamped and/or secondarily septate.

Figure 66. 

Gymnopus quinaultii. Pileal hairs. Standard bars = 10 µm. TFB 5886 (TENN-F-51994).

Figure 67. 

Gymnopus quinaultii. Pileipellis elements. A, B Encrusted hyphae C, D Gelatinizing hyphae. Standard bars = 10 µm. TFB 5886 (TENN-F-51994).

Figure 68. 

Gymnopus quinaultii. Pleurocystidia. Note apical partition of contents. Standard bars = 10 µm. TFB 5886 (TENN-F-51994).

Figure 69. 

Gymnopus quinaultii. Hymenial elements. A Basidiole B–D Basidia. Standard bars = 10 µm. TFB 5886 (TENN-F-51994).

Figure 70. 

Gymnopus quinaultii. Caulocystidia. Note origin as side branches and internal secondary septa. Standard bars = 10 µm. TFB 5886 (TENN-F-51994).

G. quinaultii clearly belongs in sect. Perforantia. The vestured stipe is not black but fuscous brown. Spores are somewhat large for sect. Perforantia, and host association (Thuja/Abies) is different (not spruce/fir needles).

Based on habitat on Thuja debris, an ancillary study was made of Collybia thujina Kauffman (MICH!), which immediately led to examination of material of Marasmius filopes Peck (NYS!) and Collybia piceina Kauffman (MICH!). Prior treatments had been published by Hesler (1959), Gilliam (1976), Redhead (1980) and Desjardin (1989). Basidiomata of all correctly identified collections were significantly smaller and more slender than those of G. quinaultii, and pileipellis organization was of a well-developed Rameales-structure, not similar to that in G. quinaultii. Desjardin (1989) agreed with Redhead that M. filopes, C. piceina and C. kauffmanii were taxonomic synonyms, all correctly placed in Marasmiellus by Redhead. I agree with this assessment, leaving G. quinaultii as a separate taxon.

Canada, British Columbia, vic. Whistler, Paradise Valley between Whistler and Squamish, 49°50'35" N, 123°09'25"W, 6.X.1990, coll RHP & LWH, TFB 3400/4 (TENN-F-49650). United States, Washington, Grey’s Harbor Co., vic Amanda Park, Forest service Rd. 2140, “Higley’s swamp,” 16.X.1992, coll RHP & KWH, TFB 5886 (TENN-F-51994; holotype).

Micromphale sequoiae Desjardin. 1985. Mycologia 77: 894–895.

United States, California, Mendocino Co., Jackson State Forest, junction of state roads 408 & 409, 13.XI.1982, coll & det D.E. Desjardin, DED 1740 (SFSU-F-000711).

(fide Desjardin 1985). 1) fruiting on needle debris of Sequoia sempervirens; 2) light brown to flesh-colored, rugulose pileus; 3) lamellae concolorous with pileus; 4) odor mild; 5) grayish-orange to brown pubescent stipe; 6) pileipellis of filamentous hyphae in a slime matrix (no Rameales-structure); 7) poorly developed rhizomorphs.

Basidiomata (Figs 71, 72A) pliant, marcescent, reviving. Pileus 6–12 mm broad, when young convex to campanulate, often with a short, acute umbo, in age becoming broadly convex to plano-convex with or without a central papilla, occasionally plane with a shallow central depression; margin when young decurved or slightly incurved, even, entire, in age becoming straight, wavy, crenate, rugulose-striate to rugulose-sulcate ¼ of the distance to center; surface dry to moist, dull glabrous, hygrophanous, at first light brown 7D4–5 overall, rarely with disc reddish brown 8E5–7, in age disc remaining light brown or fading to brownish orange 6C3–4; margin in age fading to brownish orange, greyish orange 6B2–3 or orange white 5A2, in age rarely colored buff overall with a slightly darker disc; pileus trama light brown to brownish orange, soft, up to 1 mm thick. Lamellae adnate, free in age or rarely attached to a pseudocollar, close to subdistant, narrow to medium broad (up to 1 mm), rarely anastomosing or intervenose, total lamellae = 25–27, through lamellae = 14–16; at first pale greyish orange 6B2, fading in age to pale orange white 5–6A2, typically concolorous with the pileus margin at maturity; edge even, entire, concolorous; lamellulae in 1–2 series. Stipe 20–43 × 0.7–1.5 mm broad, terete or rarely apically compressed and cleft, equal or tapered downward, hollow, cartilaginous, insititious; context concolorous with stipe surface; apex pruinose, off-white, downward pubescent and often with furfuraceous base, when young, apical portion pale greyish orange 6B2, central portion light brown 7D4–6, base dark brown 7F5–7 to rusty brown, in age apex becoming pale brownish orange 7C3, central portion becoming brown 7E4–5, base becoming dark brown 7–8F4–8 or occasionally dark brown overall in age. Rhizomorphs (Fig. 69A) short, slender, black, poorly developed, scattered; sterile stipes rare. Taste strongly alliaceous after 1–2 minutes; odor mild or rarely slightly fetid when old and wet.

Figure 71. 

Gymnopus sequoiae. Basidiomata in natural habitat. A Courtesy Michael Wood B Ryan Snow, Mushroom Observer. Standard bars = 20 mm.

Figure 72. 

Gymnopus sequoiae. A Basidiomata and rhizomorphs B Basidiospores. Standard bars: A = 10 mm; B = 5 µm. TFB 14620 (TENN-F-69325).

Scattered to gregarious on branchlets and leaves of Sequoia sempervirens; presumably throughout the range of S. sempervirens (at least northern California); October–February.

Pileipellis of inner limb up to 60 µm thick, involved in a slime matrix; slime material heterogeneous with copious crystal suspension, transparent, not totally soluble in KOH, hyaline; hyphae (Fig. 73A) 3.6–11 µm diam, repent, unoriented, interwoven, smooth, thick-walled (wall –1.5 µm thick), conspicuously clamped (Fig. 73B) but with frequent secondary septa (Fig. 73C). Pileus trama loosely interwoven; hyphae smooth, non-gelatinized, 4.2–7.2 µm broad, with hyaline to pale yellowish, inamyloid walls up to 1.5 µm thick. Lamellar trama interwoven, of two hyphal types: 1) filamentous hyphae 3.5–7 µm diam, firm-walled, clamped, not incrusted or gelatinizing; and 2) free-form hyphae, inflated to 13 µm diam, often articulating with neighboring hyphal segments, firm- to thick-walled (wall –0.7 µm thick, hyaline); contents with scattered inclusions (PhC). Pleurocystidia (Fig. 74A–D) 21–31 × 7–8 µm, fusiform, clamped; contents homogeneous, dense (PhC). Basidioles broadly clavate, often becoming ampulliform, clamped; contents multigranular; basidia (Fig. 74E–H) (21–)25–30 × (6–)9–11 µm, clavate, hyaline, (2–) 4-sterigmate; contents multigranular; sterigmata up to 4.8 µm long. Basidiospores (Fig. 72B) (6–)6.5–7.5(–8) × 3–4 µm (Q = 1.63–2.00; Q^m = 1.81; L^m = 6.50 µm) ellipsoid to lacrymoid, hyaline, smooth, inamyloid, white in deposit. Lamellar edge basically fertile; cheilocystidia (Fig. 75) common but scattered, 27–33 × 4.8–6.6 µm, clavate or ventricose-rostrate, occasionally submammilate, hyaline and thin-walled, projecting up to 11 µm beyond basidia. Stipe medullary hyphae (Fig. 76A) free (walls not gelatinized), 3–7.5(–11.5) µm diam, thick-walled (wall –1.0 µm thick, hyaline), conspicuously clamped. Stipe cortical hyphae (Fig. 76C) 4–8.5 µm diam, thick-walled (wall –1.0 µm thick), strongly incrusted in thick scabs and annuli, pigmented (yellow-brown, PhC), easily shattering in squash mounts. Caulocystidia at stipe apex (Fig. 76B, D, E) 10–140 × 9–13 µm, arising from incrusted surface hyphae with somewhat constricted attachment, subventricose and usually tapering slightly distally, thick-walled (wall –2.5 µm thick, hyaline), often strangulate, often secondarily septate and/or clamped. Caulocystidia from stipe base, (Fig. 77B–F) 25–170 × 9–13 µm, versiform, irregular in outline with obtuse apices; walls brown, evenly pigmented, –1.2 µm thick.

Figure 73. 

Gymnopus sequoiae. Pileipellis structures. A Hyphae with subgelatinized walls in slime matrix B Clamp connection C Secondary septa. Standard bars = 10 µm. TFB 14620 (TENN-F-69325).

Figure 74. 

Gymnopus sequoiae. Hymenial structures. A–D Pleurocystidia E–H Basidia. Standard bars = 10 µm. TFB 14620 (TENN-F-69325).

Figure 75. 

Gymnopus sequoiae. Cheilocystidia. A, B Clusters of cheilocystidia C–F Individual cheilocystidia. Standard bars = 10 µm. TFB 14620 (TENN-F-69325).

Figure 76. 

Gymnopus sequoiae. Stipe apex structures. A Stipe medullary hyphae C Strongly incrusted cortical hyphae C Young caulocystidium with shagreened surface B, E Individual caulocystidia. Standard bars = 10 µm. TFB 14620 (TENN-F-69325).

Figure 77. 

Gymnopus sequoiae. Lower stipe structures. A Stipe surface free-form cells with one caulocystidium B–F Individual caulocystidia showing broad-based origin and secondary septa. Standard bars = 10 µm. TFB 14620 (TENN-F-69325).

Care was taken to demonstrate all stages of basidiole maturation to demonstrate the difference between young basidioles and cheilocystidial structures. Basidioles are abundant over all of the lamellar surface and are subspherical at the earliest stage, soon becoming broadly clavate and developing a subcapitulate upper portion. Basidia remain broadly clavate throughout spore development. Contents of such structures are consistently multigranular. Although cheilocystidia are similar in dimensions and appearance, contents are homogeneous (PhC), and such structures are found only at the lamellar edge. Conversely, pleurocystidial structures are fusiform from their earliest state, merely elongating to mature size and shape. Contents are homogeneous except for a vacuolated area in midsection (perhaps nucleus; PhC).

Caulocystidia arise as side branches of stipe surface, incrusted hyphae. Early stages of caulocystidial development often bear a shagreened surface but soon becoming smooth. A unique character is the frequent internal secondary septation, as well as occurrence of a clamp connection near caulocystidial origin.

California, Humboldt Co., Redwood National Park, Davidson Rd., N41°12'51", W124°00'12", 24.X.1992, coll & det DE Desjardin, DED 5546 (SFSU-F-025665); Redwood National Park, Davidson Rd., N41°12'51", W124°00'12", 24.X.1992, coll & det DE Desjardin, DED 5546 (SFSU-F-025665); Mendocino Co., vic. Fort Bragg, Simpson Lane, 28.IX.1986, coll & det H.D.Thiers, HDT 50541 (SFSU-F-025669); Jackson State Forest, Hwy 408 at junction with road to Mendocino Woodlands, 21.XI.2015, coll & det DE Desjardin, DED 8802/TFB 14620 (TENN-F-69325); Jackson State Forest, along Hwy 409 car 1 mi from junction of Hwy 408, 18.XI.1995, coll & det DE Desjardin, DED 6316 (SFSU-F-025662); same location, 13. Nov. 1982, DED 1740 (holotype); same location, Hwy 409, 13.XII.1990, coll & det D.E. Desjardin, DED 5023 (SFSU-F-025663); same location, “Aleuria Glen,” 29.X.1990, coll H.D. Thiers, det D.E. Desjardin, DED 5012 (SFSU-F-025668).

Canada, British Columbia, Victoria, Saanich Peninsula, Observatory Hill, N48.5262°, W123.422°, 4.XI.2011, coll & det O. Ceska (as Marasmius androsaceus), UBC-F-25212.

sub- = Latin: less than; laccatus = Latin: appearing polished or varnished, referring to varnished appearance of dried pileus.

1) Fruiting on needles of Thuja (and Pseudotsuga); 2) cheilocystidia rare, small, clavate, smooth, without setulae; 3) stipe near ochraceous buff above, downward rusty brown; 4) pileal hairs usually smooth, rarely roughened; 5) dried pileus sublaccate; 6) pileipellis of smooth, unencrusted, repent hyphae in slime matrix; 7) rhizomorphs inconspicuous, with minute black basal pad.

Basidiomata (Fig. 78A) diminutive. Pileus 2–8 mm broad, convex to plano-convex, minutely laccate, vaguely tuberculate, not striate or sulcate; disc about “sayal brown” 6C5 to “ochraceous buff” 5A5; limb and margin about “light ochraceous buff” 5A4 to “tilleul buff” 7B2. Lamellae (Fig. 78A) pseudocollariate (dried), adnate, distant, thickish, total lamellae = 20–23, through lamellae = 9–11, now (dried) “tilleul buff” 7B2 to “olive buff” 3B3; lamellulae in a single rank. Stipe 18–25 × 0.6–0.8 mm, terete, equal, hollow to lightly stuffed, subinsititious, minutely shaggy to silky above, downward becoming minutely barbed (35×), above about “ochraceous buff” 5A5, downward through “army brown” 8D5, increasingly dark to rusty brown (“Prout’s brown” 5F6 to “Vandyke brown” 7E6), sometimes slightly expanded at base; stipe medulla white. Rhizomorphs (Fig. 78C) (if produced) inconspicuous, often represented only by minute, black basal pads. Taste and odor not recorded.

Figure 78. 

Gymnopus sublaccatus. A Basidiomata showing adventitious rhizomorphs and stipe insertion to pileus B Basidiospores C Substrate scales with back spots and rhizomorphs. Standard bars: A = 20 mm; B = 5 µm; C = not to scale. UBC 25212.

Fruiting on needles of Thuja plicata, less often on Pseudotsuga menziesii; British Columbia; Autumn to early Winter.

Pileipellis involved in a slime matrix which includes detersile encrusting material, of the following elements: 1) pileal hairs (Fig. 79) –150 × 3–6 µm, erect, firm-walled, smooth or rarely weakly roughened, slightly subcapitulate; 2) repent hyphae 4.5–7.5 µm diam, firm-walled, conspicuously clamped but often secondarily septate, without discernable slime sheath. Pileus and lamellar tramae loosely interwoven; hyphae 3–5.5 µm diam, smooth, firm-walled, without slime sheath, conspicuously clamped. Pleurocystidia (Fig. 80A–D) 25–35(–41) × 7–8 µm, fusiform to clavo-fusiform, without partitioned contents but with vague vacuolated area in midsection (PhC), conspicuously clamped. Basidioles (Fig. 80E) clavate; basidia (Fig. 80F–H) 24–27 × 8–9 µm, clavate, 4-sterigmate, clamped; contents multigranular at maturity; effete pleurocystidia and basidia emptying but not collapsing (“husking” Fig. 81B); subbasidial hyphae (Fig. 81A) rupturing at clamp connections, appearing beaded. Basidiospores (Fig. 78B) (6.5–)7–7.5(–8) × (3.5–)4–5 µm (Q = 1.40–1.88; Q^m = 1.67; L^m = 7.20 µm), rotund-ellipsoid, smooth, thin-walled, inamyloid. Cheilocystidia (Fig. 82) very locally common, usually rare to absent, 14–22 × 8–12 µm, clavate to utriform, thin-walled, obscurely clamped; contents homogeneous. Stipe medullary hyphae (Fig. 81C) 4–12 µm diam, strictly parallel, apparently free (no discernable gelatinized matrix, hyaline, thick-walled (wall –0.7 µm thick, hyaline, non-gelatinized), obscurely clamped, often with small side-branches ranging from lobate to rudimentarily branched. Stipe cortical hyphae 4–8 µm diam, strictly parallel, apparently free, thick-walled (wall –1.2 µm thick, especially at stipe surface), pigmented (yellow-brown, KOH + PhC), appearing moderately dextrinoid (IKI + PhC), or not so (IKI + BF), often producing broad-based side branches. Caulocystidia (Figs 83, 84) –45 µm at stipe apex, –140 µm at stipe base, × 5.5–10 µm, ranging from lobate to elongate-digitate, without basal clamp, often internally secondarily septate, sometimes branched in one rank, thick-walled [wall –4 µm thick, often occluding cell lumen, pigmented (yellow-brown PhC)].

Figure 79. 

Gymnopus sublaccatus. Pileipellis elements. A–E Pileal hairs F Secondary septa of pileipellis hypha. Standard bars = 10 µm. UBC 25212.

Figure 80. 

Gymnopus sublaccatus. Hymenial elements. A–D Pleurocystidia E Basidiole F–H Basidia. Standard bars = 10 µm. UBC 15356.

Figure 81. 

Gymnopus sublaccatus. A Subbasidial hyphae appearing beaded B Effete hymenial structures without collapse (“husking”) C Stipe medullary hyphae. Standard bars = 10 µm. UBC 25212.

Figure 82. 

Gymnopus sublaccatus. Cheilocystidia. Standard bars = 10 µm. UBS 15356.

Figure 83. 

Gymnopus sublaccatus. Caulocystidia from stipe apex. Standard bars = 10 µm. UBC 25212.

Figure 84. 

Gymnopus sublaccatus. Caulocystidia from stipe base. Standard bars = 10 µm. UBC 25212.

With slime matrix covering the pileus surface and involving hymenial structures, with characteristic clavate cheilocystidia and with vestured stipe, UBC 25212 seems certain to belong in sect. Perforantia. There it joins G. perforans, G. foliiphilus and G. sequoiae with the same general characters. From Ma. androsaceus, G. sublaccatus differs in pigmented (not black), vestured stipe. If its substrate preference is limited to Pseudotsuga, this constitutes another difference. Finally, pileipellis of G. androsaceus is characterized by diverticulate hyphal segments and broom cell-like hyphal termini, unlike that of G. sublaccatus which lacks these structures.

Basidiomata of G. sequoiae resemble those of M. sublaccatus in stature and size, but seems limited to fruiting on needles of Sequoia sempervirens in northern California. Cheilocystidia are shaped like an incandescent bulb, sometimes slightly askew, but otherwise quite typical of cheilocystidia in this taxonomic complex (compare, for example, those of G. bulliformis and of G. sequoiae). Difficult to distinguish from immature basidia, such cheilocystidia are less obscure after several sightings.

Canada, British Columbia, Queen Charlotte Islands, Burnaby Island, Section Cove, N52°24'35", W131°19'55", 8.IX.2006, coll PK Kroeger (as Marasmius androsaceus), PK 489 (UBC F18168); Queen Charlotte Islands, N52°21'23", W131°24'24", 11.IX.2008, coll P. Kroeger (as Marasmius androsaceus), PK 5904 (UBC F16670); Queen Charlotte Islands, Ellen Island, N52°09', W131°06', 10.IX.2004, coll & P. Kroeger, B. & C. Kendrick, J. Brown, (as Marasmius androsaceus), PK 2932 (UBC F15356); Queen Charlotte Islands, Ross Island, N52°10', W131°07', 2.IX.2006, coll P. Kroeger (as Marasmius androsaceus), PK 4347 (UBC F17675); Victoria, Saanich Peninsula, Observatory Hill, N48.5262°, W123.422°, 4.XI.2011, coll & det O. Ceska (as Marasmius androsaceus), UBC-F-25212 (holotype).