Phaeocollybia in western North America 4: Two new species with tibiiform cheilocystidia and Section Versicolores reconsidered

Abstract

Two new phaeocollybias are described from coastal and coastal montane coniferous forests of California, Oregon, and Washington. Phaeocollybia rufotubulina, closely related to the gregarious, hollow-stiped P. californica and P. scatesiae, differs in pileipellis pigments, pseudorhizal branching pattern, and RFLP profile. P. tibiikauffmanii differs from the similarly robust, orange-brown, viscid capped P. kauffmanii in its thick-walled narrow-necked capitulate cheilocystidia. Phenetic and cladistic analyses of restriction data generated from thirty-seven isolates representing P. californica, P. pseudofestiva, P. scatesiae, P. spadicea, P. radicata, and the two new species support the existence of a clade characterized by thick-walled tibiiform cheilocystidia. Characters previously used to diagnose Section Versicolores are re-examined, and the implied phylogenetic separation of the vernal P. pleurocystidiata from the fall-fruiting tibiiform western species is discussed.

Full text

MYCOTAXON
Volume 90(2), pp. 241–260 October-December 2004
Phaeocollybia in western North America 4:
Two new species with tibiiform cheilocystidia and
Section Versicolores reconsidered
1
LORELEI L. NORVELL
llnorvell@pnw-ms.com
Pacific Northwest Mycology Service
6720 NW Skyline Boulevard, Portland OR 97229-1309 USA
Abstract — Two new phaeocollybias are described from coastal and coastal montane
coniferous forests of California, Oregon, and Washington. Phaeocollybia rufotubulina,
closely related to the gregarious, hollow-stiped P. californica
and P. scatesiae, differs in
pileipellis pigments, pseudorhizal branching pattern, and RFLP profile. P. tibiikauffmanii
differs from the similarly robust, orange-brown, viscid capped P. kauffmanii in its
thick-walled narrow-necked capitulate cheilocystidia. Phenetic and cladistic analyses
of restriction data generated from thirty-seven isolates representing P. californica, P.
pseudofestiva, P. scatesiae
, P. spadicea, P. radicata, and the two new species support the
existence of a clade characterized by thick-walled tibiiform cheilocystidia. Characters
previously used to diagnose Section Versicolores are re-examined, and the implied
phylogenetic separation of the vernal P. pleurocystidiata from the fall-fruiting tibiiform
western species is discussed.
Key words — Agaricales, Basidiomycota, Cortinariaceae, ITS, temperate rainforest
Introduction
As part of a continuing study of the genus Phaeocollybia begun in 1991, a large number
of fresh specimens (>1,000 collections) from Pacific coastal temperate rainforests of
western North America have been examined and compared closely with older dried
material (including type specimens). During this research, 160 collections were
molecularly analyzed (Norvell 1998ab, 2000, 2002; Norvell & Redhead 2000), new
insights into the development and biology of the genus gained (Norvell 1998ab), new
morphological characters revealed (e.g., the existence of a universal pellicular veil,
different pseudorhizal morphologies, the presence of sarcodimitism, see Norvell
1998ab), and seven new species named (Norvell 2000, 2002; Norvell & Redhead 2000).
Two additional undescribed species characterized by large ornamented basidiospores,
absence of clamp connections, and narrow-necked thick-walled capitulate tibiiform
cheilocystidia are described below. Similarities among western North American
phaeocollybias characterized by thick-walled tibiiform cheilocystidia are discussed,
1
Based in part on material submitted for a doctoral dissertation at the University of Washington, Seattle WA 98195-5325 USA.

242
and Section Versicolores (Smith 1957b; Singer 1970, 1986, 1987; Bandala and Montoya
1994) is reconsidered.
Materials and Methods
Specimen collection and examination, macrochemical tests, DNA extractions, PCR-
amplification of the ITS1 + 5.8S + ITS2 rDNA region (henceforth referred to as ʻthe
ITS regionʼ), restriction digests and mapping, and phenetic and cladistic analyses using
NTSYS (Rohlf 1993) and Phylip (Felsenstein 1995) were conducted using protocols
outlined by Norvell (1998ab, 2000).
Descriptions are based on data taken from both type and other well-documented
collections. Morphological and developmental terms (
e.g. tibiiform diverticula,
sarcodimitic tissues, pellicular veil, vertical-monopodial and rhizomorphic pseudorhizae)
are explained in Norvell (1998ab). General non-standardized color names in lower case
are accompanied by italicized color references. Ridgway (1912) colors are capitalized
and abbreviated, with a slash “/” or parentheses separating modifiers of the same base
color (e.g. Mikado /Mars /Verona Brown = Mikado Brown, Mars Brown, Verona Brown;
(Pale) Pinkish Cinnamon = Pale Pinkish Cinnamon and Pinkish Cinnamon); bracketed
Munsell (1976) alpha-numeric color ranges (e.g. [2.5Y 6-8/1-4]) follow Ridgway
references. Measurements of anatomical characters were taken from tissues rehydrated
in 6% aqueous KOH unless otherwise noted, and basidiospores from the stipe apex were
measured when spore prints were lacking. Ranges enclosed in parentheses accompany
mean basidiospore dimensions with standard deviations. “L+ll/cm” is the number of
lamellae + lamellulae per cm at the pileus “edge” and at the “midpoint” between edge
and stipe. “Aerial stipe” refers to the portion of the stipe above the ground; “origin”
refers to the pseudorhizal origin at the very base of the basidiome within the substrate.
Herbaria housing cited collections are abbreviated in accordance with Holmgren et al.
(1990), with “PNW” representing the Pacific Northwest Mycology Service herbarium.
Subscripts denote Randon Fragment Length Polymorphism (RLFP) codes; and asterisks
(*) flag Northwest Forest Plan Strategy 1 collections (USDA-USDI 1994, Norvell
1995, Norvell & Exeter 2004) of significance to US governmental forest management
policy. Longitudes and latitudes were either converted from Township-Range-Section
data obtained from USDI-BLM maps using the TRS2LL computer application (Wefald
1995) or obtained electronically from the Yale Peabody Museum of Natural History
GNIS database (YALE). Separate collection dates are not listed for the date-based
numbered Norvell (LLN##), Density Management Study (g##), and Chronosequence
Study (a##) collections
e.g. LLN1921116-02
ruc1
= Norvell November 16, 1992
collection 2, RFLP code ruc1). Collector abbreviations include RLE or Ex (Exeter),
LLN (Norvell), SAR (Redhead), and HDT (Thiers). Vegetation abbreviations include
ABGR (Abies grandis), ABAM (Abies amabilis), BENE (Berberis nervosa), GASH
(Gaultheria shallon), LIDE (Lithocarpus densicarpa), OXOR (Oxalis oregana), PISI
(Picea sitchensis), POMU (Polystichum munitum), PSME (Pseudotsuga menziesii),
SESE (Sequoiodendron sempervirens), TSHE (Tsuga heterophylla).

243
Taxonomic Descriptions
Phaeocollybia rufotubulina sp. nov. FIGURES 1-2
Pileus usque ad 60 mm latus, convexo-campanulatus, umbonatus, madidus vel viscidulus,
glaber, e rufo rufo-brunneus; caro subaurantia; odor leviter florali vel leviter favaceus;
lamellae aurantio-cremeae, fulvo-brunnescentes e sporis; stipes usque ad 12 mm versus
apicem, usque ad 180 mm longus cum pseudorhiza tenua racemosa, glaber vel fibrillosus,
cum corticem tenuissimo, cavus (tubulosus). Sporae longae, 8.2-10
× 4.5-6 µm, limoniformes,
rugulosae cum rostro et apiculo glabro et cum plaga obscura; ixocutis cum suprapelle
tenue vel moderato, gelatinosa, cum pigmento aurantio incrustato et subpelle cum pigmento
intraparietali et encrustato; cheilocystidia abundans, heteromorpha, tibiiformea et 3 µm lata
versus septam cum collo crasse tunicato angusto et cum capitulae usque ad 1.5 µm lata, vel
clavata leptoderma, gelatinosa, hyalina. Hyphae desunt in texturis omnibus. Typus ad terram
in silvis mixtis: LLN1921116-01ruc1, L. L. Norvell cum G. L. Barron et S. A. Redhead (Jackson
State Forest, Mendocino County, California, USA). Holotypus WTU; Isotypi DAOM, PNW..
Etymology: from the Latin rufus = “reddish” + tubulus = “little pipe”
Selected descriptions and illustrations: Norvell (1998a)
—color photographs, line drawings.
Norvell (1998b)—black & white photographs, line drawings.
Brief summary — Basidiomes densely gregarious, small to medium, more or less
uniformly reddish orange; pilei convex-campanulate, subviscid; young lamellae pale
orange; stipes strict, tubular with very thin cortices, matte to polished, pale orange
to deep purplish red; pseudorhizae sequential-racemose, cord-like, rhizomorphic.
Basidiospores large (~9 × 5 µm), long-limoniform, coarsely warty-rugulose with smooth
elongated apical beaks, dark orange amber in KOH; cheilocystidia abundant, slightly
heteromorphic, thick-walled, refractive, lageniform/tibiiform, occasionally intermixed
with thin-walled, mucronate elements; pileipellis bilaminate with a moderately thin
yellowish suprapellis composed of narrow hyphae spirally encrusted with orange to
orange-brown pigment overlying a thin brownish-orange subpellis; clamp connections
absent. Tissues either negative or very faintly pink (pseudorhizae only) in syringaldazine.
Dried pileus colored copper or purplish bronze, with a metallic sheen.
Expanded description — Pileus 15-62 mm broad, convex-umbonate with acute umbo
and incurved margin when young, expanding to broadly campanulate with acute to low
broad umbo, incurved to straight margin, and incurved edge, glabrous, lubricous to
subviscid, opaque, nonstriate, generally bright reddish to brownish orange
(Xanthine
/ Ochraceous / Apricot Orange, Mars Yellow, Ochraceous Buff), often with slightly
darker disc and edge (Sanfordʼs / Dark Orange Brown, Tawny), brownish streaks
and damaged areas occasionally present, dried pileus metallic, usually copper or
winy-reddish bronze. Context 3-6 mm thick at the disc, pale orangish cream (Pale
Ochraceous Salmon, Pinkish Buff). Odor faintly floral (reminiscent of cultivated Viola)
or like cooked potatoes (similar to Amanita citrina), occasionally sharply farinaceous
after cold storage, then developing a green corn odor on warming. Taste mild and
not distinctive to slightly bitter (reminiscent of raw potatoes). Lamellae
nearly free,
emarginate with a short tooth, ventricose, multi-tiered with 1-3 (young) to 2-6 (mature)
lamellulae irregularly interspersed; 2-5 mm broad (mean length : width ratio 3.5),
subcrowded (L+ll/cm: edge = 19-23; midpoint = 10-12); color pale orangish cream
(Apricot / Ochraceous Buff) when young, maturing to foxy brown (Clay Color,
(Ochraceous) Tawny) with bright rusty orange streaks.

244
Primordial sheath remnants present as connective fibrils between pileus and stipe
(immature) or as occasional, densely scattered, fine, dark reddish brown fibrils on mature
stipe apex. Stipe central to slightly eccentric; aerial length up to 140 mm, combined
length with pseudorhiza up to 180 mm, apex 4-12 mm diam; strict, terete, uniformly
equal except for slight bulbous swelling at ground level above sharply attenuating
cord-like pseudorhiza, glabrous and smooth or covered with fine short reddish-orange
fibrils, dry to lubricous; apex slightly paler than reddish orange pileus when young
(Xanthine Orange, Light Ochraceous Buff, Cinnamon), gradually darkening upwards
to a dark reddish to purplish brown in age (Hayʼs Russet, Kaiser / Liver Brown); hollow,
cortex 1-1.5 mm thick, brittle, cartilaginous; lumen empty from pileus to rhizomorphic
pseudorhiza. Pseudorhiza sequential-racemose, rhizomorphic, 1.5-2 mm thick cord of
undetermined length (strands up to 55 mm have been retrieved); cortex dark reddish
brown (Liver Brown, Clay Color), cartilaginous; medulla dark orange-brown (Clay
Color), compact.
Spore print brown (slightly darker than 7.5YR 4/4 [Sayal Brown]).
Basidiospores
9 × 5 ± 0.6 × 0.3 µm (overall range 8.2-10 × 4.5-6 µm), limoniform
with a prominent straight to slightly tilted beak in profile, ovate with swollen portion
in face view, exosporium thick-walled, coarsely rugulose roughened except over the
apiculus and beak, with less ornamented plage visible in oil immersion; dark to deep
orange-amber in KOH; inamyloid. Cheilocystidia heteromorphic, with thick-walled
and thin-walled lageniform, tibiiform, and clavate elements intermixed; mucronate/
tibiiform elements occasional to frequent, 17-30 µm long with 3 µm wide ventricose
bases below 1-µm wide refractive necks and with/without 1.5 µm diam capituli,
containing viscous pale to dark amber contents; thin-walled elements less frequent,
moderately inflated or broadly clavate, heavily gelatinized, hyaline, developing long
filamentous apical outgrowths in age or in storage. Pleurocystidia absent. Basidia 4-
spored, 25-30 × 5-6 µm with long (to 8 µm) sterigmata, hyaline to pale straw-colored.
Pileipellis a bilaminate ixocutis; suprapellis 120-200 µm thick, the gelatinous matrix
pale orange-yellow and the embedded hyphae long, branched, 2-4 µm wide, highly
gelatinized, roughened, and with spirally encrusting and intraparietal yellowish-
orange to orange brown pigments; subpellis thick (300-450 µm), hyphae (5) 8-15 µm
diam, thick-walled, highly gelatinized, walls orange-yellow to deep orange in KOH.
Stipitipellis a longitudinal ixocutis, hyphae narrow (2-4 µm diam), cylindrical, thick-
walled, highly gelatinized, incrusted with medium to dark orange-amber to orange-red
pigments. Tramal tissues moderately to highly gelatinized, strongly sarcodimitic in
the pseudorhiza and lower stipe with long (95- 200 µm), wide (≤ 20 µm), thick-walled
(≤ 3 µm), cylindrical, rigid yellowish “vessel” hyphae supported by less conspicuous,
shorter (~20 µm), narrower (4-6 µm), branched, thin-walled (< 0.5 µm), flexuous
hyphae; in the pileus both hyphal types also present, but vessel hyphae with thinner
(1 µm) walls; in the lamellae central hyphae parallel, 4-6 µm diam, highly gelatinized,
slightly thick-walled, with refractive septa and pale amber intraparietal pigments; these
giving rise to a compact, 2-6 µm thick subhymenium. Tibiiform diverticula abundant
on mycelium, primordial surfaces, and pseudorhizal pellis; also frequent on vestiges
of pellicular veil on stipe apex. Clamp connections absent in mature basidiomes but
occasionally present in primordial pellis.

245
FIGURE 1 Phaeocollybia rufotubulina (LLN1921116-01) 1a. Holotype in situ with differentially
developed basidiomes in Jackson State Forest, Mendocino County, California (photo by Scott
Redhead). 1b. Suprapellis hyphae with refractive septa and dark orange encrustations (phase
contrtast). 1c. Tibiiform diverticula on stipitipellis (bright field). Scales = 10 µm.

246
Macrochemical reactions –– Syringaldazine negative to slightly positive (pseudorhiza
develops black tinge after 15 minutes and magenta tinge after 60 minutes);
KOH dark
brown on pileus ; FeSO
4
slightly greenish on all tissues. Fluorescence –– (fresh material
not checked; dried gills brilliant mustard yellow where mature spores absent. RFLP
profile (Extensive data for 6 isolates including type)
–– ITS 690 bp (± 730 bp second
band); unique in Xho1 (500-190); EcoR1 (365-325) diagnostic; also cut in Cfo1, Hinf1,
Nde2, Pal1, and Pvu2; uncut in Rsa1, Sal1.
Ecology, range and distribution — Densely gregarious in loose bundles in sandy loam
under needle duff in mature mixed forest with an overstory of Lithocarpus densifolius,
Sequoia sempervirens, Pseudotsuga menziesii, Abies grandis and shrubby understory
of Rhododendron, Vaccinium ovatum, and Polystichum munitum. Uncommon, yet
often locally abundant. Known from 13 sites (36 collections) in coastal lowlands
from southern Oregon (Lane County) south to Marin County along the north central
California coast. October through January.
OTHER SPECIMENS EXAMINED UNITED STATES. CALIFORNIA: Marin Co. AUDUBON
C
ANYON RANCH (122.6794°W, 37.19308°N)—05.I.1984 CCalhoun84-3737 (SFSU as P.
californica); BOLINAS RIDGE TR NEAR MT TAMALPAIS (122.7847°W 38.095°N)—clustered in soil
under oaks in dense mixed conifer-hardwood forest 17.I.1992
HDT54067 (SFSU as P. radicata);
Mendocino Co.
JACKSON SF ʻALEURIA GLENʼ ON SF RD 409, 1.5–2 MI FROM 408 (123°46ʼW
39°20ʼ28”N)—mature LIDE ABAM PSME TSHE SESE 24.XI.1992 LLN1921124-02ruc3
,
-03,-04,-03,-
06,-07,-08ruc6,-09,-10,-24,-27 by/w GLBarron REHalling HDT (WTU, -02 -08 split w PNW);
S
IMPSON LANE, FORT BRAGG AREA (123.8114°W, 39.4172°N)—duff mixed forest 22.XI.1982
ASMethven2149 (NY as P. radicata); JACKSON SF (123°38ʼ52”W 39°23ʼ33”N)—gregarious
humus mixed woods
14.XI.1967 HDT21575 (SFSU as P. radicata), 25.X.1986 MTSeidl2114
(WTU), mature LIDE ABAM PSME TSHE SESE 21.XI.1992 LLN1921121-06-ruc9 by REHalling HDT
(NY WTU PNW); JACKSON SF NEAR MENDOCINO (123°47ʼ30”W, 39°18ʼ28”N)—soil mature LIDE
ABAM PSME TSHE SESE 22.XI.1980 n.c. (HSC5940 as P. dissiliens); JACKSONSF JCT RDS 408/409
ʻ
MUSHROOM CORNERʼ (123°44ʼW, 39°19ʼN)—humus dense mixed forest 25.XI.1960 HDT08477
(SFSU as P. attenuata), 11.XII.1965
HDT14607 (SFSU as P. attenuata), 05.XI.1967 HDT21404
(SFSU as P. attenuata), 07.XII.1969 HDT24452 (SFSU as P. californica), 03.XII.1971
JFAmmirati06222 (
MICH as P. californica), 31.X.1972 HDT30427 (SFSU as P. californica),
07.XII.1974 HDT33175 (SFSU as P. californica), 02.XI.1975 HDT35261 (SFSU as P. californica),
soil SESE ABAM 13.XII.1990 DEDesjardin5030 (SFSU as P. californica), mature LIDE ABAM PSME
TSHE SESE LLN1921117-08-ruc8 w SAR (DAOM PNW WTU); RUSSIAN GULCH SP (123°46ʼ26”W
39°19ʼ59”N)—mature LIDE ABAM PSME TSHE SESE 25.XI.1992 SAR7454, 7500ruc7 (DAOM
PNW); WOODLANDS CAMP (123.7050°W, 39.3156°N)—04.XII.1971 DEStuntz17100 (WTU);
San Mateo Co. H
UDDART ROADSIDE PARK (122.2947°W 37.4350°N)—duff mixed forest
23.XII.1964 WJSundberg195 (SFSU as P. attenuata); Santa Clara Co. SANTA CRUZ MTNS
(121
°54ʼW, 37°6ʼN)—gregarious terrestrial RAHanks s.n. (SFSU as P. attenuata); Sonoma Co.
SALT POINT SP (123°18ʼ43”W, 38°34ʼ31”N)—mixed woods 25.XI.1972 HDT30756 (SFSU as P.
californica). OREGON: Lane Co. FLORENCE (124°5ʼ55”W 43°58ʼ58”N)—LLN1971031-01 Mt
Pisgah show (PNW).
Comments Phaeocollybia rufotubulina is easily recognized in the field by its
striking reddish-orange color, dense gregarious habit, and strict tubular stipe that splits
longitudinally and curls back into a floret when sliced crosswise (similar to a dandelion
scape). Long, narrow, heavily rugulose-roughened, dark red-brown spores, moderately
thick pileipellis with concentrically dark-encrusted hyphae, and heteromorphic mix
of abundant thick-walled tibiiform and thin-walled clavate cheilocystidia characterize
the species microscopically. No other
Phaeocollybia species tested possesses an Xho1
restriction site. P. rufotubulina appears coastally restricted to southern Oregon (one

247
FIGURE 2. Phaeocollybia rufotubulina (LLN1921116-01 Holotype WTU; isotypes DAOM, PNW)
2a. Habit showing sequentially branching cord-like rhizomorphic pseudorhiza. Scale = 5 cm.
2b. Cheilocystidia, basidium, and basidiospores. Scale = 10 µm.
2b
2a

248
collection) and Humboldt, Mendocino, and Marin Counties in California. (For ontogeny
and pseudorhizal anatomy, see Norvell 1998b).
A cryptic species previously misidentified as the morphologically similar
Phaeocollybia
californica A.H. Sm., P. rufotubulina produces more richly pigmented and strongly
ornamented basidiospores, dark orange (not hyaline) concentric encrustations over
subtly wider suprapellicular hyphae, a thinner, darker stipitipellis, and more abundant
tibiiform cheilocystidia. P.californica produces slightly larger, less highly pigmented
basidiomes and stipes with thicker cartilaginous cortices lined with long white fibrils
Phaeocollybia rufotubulina
and P. californica form a complex with another western
species, P. scatesiae A. H. Sm. & Trappe, previously synonymized with P. californica
by Horak (1977). All three species share anatomical similarities (basidiospore form and
size, cheilocystidia) and possess racemose rhizomorphic pseudorhizae. Molecularly, P.
rufotubulina is separated by a 10bp length mutation in the ITS region that results in
a unique RFLP. P. californica
and P. scatesiae, which have identical RFLP profiles,
are differentiated by striking macroscopical and subtle microscopical differences: the
densely clustered, highly glutinous P. scatesiae basidiomes erupt in fasciculate mounds
(usually with scores of tightly clustered basidiomes emanating from single points
on subtending rhizomorphic pseudorhizae) rather than in the ʻgregarious arcsʼ of P.
californica. Additionally, the broadly conic and more sharply umbonate pilei are heavily
glutinous and yellowish brown to dark blackish-brown (not subviscid and brownish-
orange to orange brown), and the spore prints are a paler brown. Microscopically, P.
californica is distinguished by its larger, more rounded, darker basidiospores, less
abundant cheilocystidia, and ʻfurredʼ suprapellicular hyphae (spirally hyaline to pale
amber encrusted) with clearly visible, frequently refractive septa. In P. scatesiae, the
smooth-walled suprapellis hyphae are usually so submerged within a thick gelatinous
matrix that individual septa can be seen only with difficulty. P. scatesiae is the most
widely distributed of the three species, extending from Californiaʼs Mendocino County
the furthest north (to Washingtonʼs Olympic peninsula) and farthest inland (to Oregonʼs
Mt. Hood in the Cascades).
Phaeocollybia tibiikauffmanii sp. nov. FIGURES 3-4
Pileus usque ad 70 mm latus, conico-campanulatus, umbonatus, viscidus, glaber, fulvus
vel brunneo-aurantius; caro crassa, pallida; odor leviter florali vel farinaceus; lamellae
subaurantiae, subroseo-brunnescentes e sporis; stipes usque ad 12 mm latus versus apicem,
usque ad 230 mm longus cum pseudorhiza attenuata, glaber, farctus, versus apicem lamellis
subconcolorus, versus subapicem vinascens. Sporae 7.5–9
× 4.5–5.2 µm, limoniformes,
verrculosae vel leviter punctatae cum rostro longo stricto glabro usque ad 1.5 µm et cum
apiculo excentrico glabro, cum plaga obscura; ixocutis cum suprapelle subcrassa, gelatinosa,
hyalina et subpelle crassa cum pigmento intraparietali, infrequens incrustato; cheilocystidia
tibiiformeae cum collo crasse tunicato angusto et capitulae 1.5–3 µm crassae vel infrequens
clavata leptosperma hyalina. Hyphae desunt in texturis omnibus. Typus ad terram in silvis
mixtis Tsugarum et Pseudotsugarum: L. L. Norvell a2011031ox-01 cum R. L. Exeter (Pedee
BLM Reserve Forest Chronosequence transect, Polk County, Oregon, USA). Holotypus OSC;
Isotypi WTU, PNW.
Etymology: from ʻtibiʼ (< tibia = L. for thigh-bone) relating to its tibiiform (ventricose below
and capitate above) cheilocystidia and ʻkauffmaniiʼ for its similarity to P. kauffmanii.
Selected descriptions and illustrations: Norvell 1998a—color photographs, line drawings;
1998b—black & white micrograP.

249
Brief summary – Basidiomes solitary to fasciculate, moderately large, slightly
farinaceous, deeply rooting; pilei orange tawny to brown, broadly campanulate to
reflexed, viscid; young lamellae pale orange yellow to pale buff; stipes slender, pale
pinkish cinnamon, stuffed with longitudinally splitting cortex surrounding firm to
cottony pith; pseudorhizae vertical-monopodial, frequently fasciculate. Basidiospores
medium-sized (8 × 4.5 µm), limoniform, roughened, with prominent, smooth, tapered
apical beaks; tibiiform cheilocystidia with refractive, thick-walled necks intermixed
with thin-walled lageniform and clavate elements; brownish-orange with diffuse
pigments; clamp connections absent. Tissues immediately magenta in syringaldazine.
Dried pileus copper-colored, sometimes with deep purple red umbo.
Expanded description –– Pileus up to 120 mm wide, broadly conic-campanulate with
conic to acute low umbo, upturned margin and down-turned straight edge; glabrous,
viscid, opaque to slightly hygrophanous, nonstriate; overall brownish orange or foxy
brown (5YR 5/8 – Tawny, Orange Cinnamon) or with tawny disc and orange cinnamon
edge zone, darkening in age (7.5YR4/2, 10YR 8/4 – Verona, Snuff Brown). Context
4-6 mm at the disc and confluent with stipe pith; color when young pale orangish
white (10YR 9/3), becoming drab grey in age (10YR 7/2-6). Odor faintly floral with
farinaceous overtones. Taste mild, not distinctive. Lamellae nearly free, ventricose,
thin with ± even edges, polydymous with 3-7 irregularly interspersed tiers, narrow (4-5
mm broad; average length : width ratio = 5.5), close (L+ll/cm = 19 edge, 10 midpoint);
color pale orangish buff (10YR 6-8/6 = Warm Buff) when young, dull pinkish brown
(5YR 5/6 = Fawn Color) when mature. Remnants of primordial sheath evident as
occasional darker fibrillose patches on stipe apex.
Stipe slightly eccentric, terete, ±
equal above, gradually narrowing below toward pseudorhiza; apex 5-12 mm diam, aerial
length to 100 mm, combined length with pseudorhiza ≤ 230 mm; glabrous except for
darker fibrillose patches, dry, innately longitudinally lined; apex pale to deep pinkish-
orange (7.5 YR 6/5-8; 10YR 7-8/4 = Pinkish Cinnamon, Orange Cinnamon; Warm
Buff), below grading to dull pinkish brown (5YR 5/6 = Fawn Color); cartilaginous
rind 2 mm thick, splitting longitudinally in age; stipitipith compact fibrillose, orangish
white; Pseudorhizal form vertical-monopodial, slightly >2/3 overall stipe length,
gradually tapering to pointed origin, origin ferruginous to dark brown (2.5 YR 4/6-
8 = Ferruginous, Sanfordʼs Brown); interior pith firm, brown where water-soaked,
otherwise concolorous with stipitipith.
Spore print
dull pinkish brown (Fawn Color, 5YR 4-5/4-6).
Basidiospores
8 × 4.5 ± 0.3 µm (range 7.5–10 × 4–5.5 µm), limoniform with a
protruding beaked apex in profile, fusoid-elliptical in face view, verruculose to punctate
roughened except on 0.5-1 µm long beaked apex and eccentric apiculus, suprahilar
plage an indistinctly bordered area of lowered ornamentation discernible under oil
immersion, orange-amber in KOH, ochraceous in H2O, inamyloid (very young spores
still attached to basidia faintly dextrinoid in Melzerʼs). Cheilocystidia abundant,
lageniform/tibiiform elements abundant, occasionally diverticulate, necks refractive,
thick-walled; capituli usually present; occasional clavate elements intermixed, thin-
walled; tibiiform elements 17-32 (50) µm long, branched at or above ultimate and
penultimate 2-3 µm diam septa, often swelling above septa to 3-5 µm before narrowing

250
to 8–15 × 0.5–2 µm necks, necks either thick-walled and refractive or thin-walled, with
or without 1.5–3 µm diam capituli, hyaline. Pleurocystidia absent.
Basidia 4-spored,
27–32 × 7–8 µm, hyaline. Pileipellis a bilaminate ixocutis; suprapellis 100–120 µm
thick, hyphae radially aligned, 2–6 µm diam, highly gelatinized, hyaline, hyphae within
the top 50 µm of suprapellis cylindrical, 2 µm diam, closely compacted and collapsed,
in lower portion 2–6 µm wide, thin-walled, often inflated, smooth; subpellis 170–200
µm thick, brownish-orange, hyphae 2–8 µm diam, thin-walled, inflated above refractive
septa, with diffuse and often encrusting orange pigments, occasional orange oleifers
intermixed. Stipitipellis hyphae subgelatinized, thin-walled, long, 2–6 µm diam, pale
brownish green (diffuse pigments soluble in KOH). Pseudorhizal pellis hyphae 2–4
µm diam, gelatinized, dark orange brown. Tramal tissues gelatinous, oleifers present
throughout; in the pseudorhiza slightly sarcodimitic, “vessel” hyphae infrequent, only
slightly thick walled (≤0.5 µm thick), long, 10–20 µm diam; flexuous hyphae abundant,
thin-walled, branched, 2–6 µm wide, irregularly inflating; in the stipe and pileus
monomitic; in the lamellae hyphae parallel, 65–80 × 3–6 µm, thin-walled, inflated,
subgelatinized, hyaline, toward the hymenium narrowing to 2–3 µm diam and giving
rise to a rudimentary subhymenium. Tibiiform diverticula
abundant on mycelium and
pseudorhizal pellis, less frequent on fibrillose pellicular remnants on stipe apex, 4–20 ×
0.5–1 µm, hyaline. Clamp connections absent in all tissues.
Macrochemical reactions
—Syringaldazine positive (immediately magenta) on all
tissues; KOH positive (immediately dark brown); FeSO
4
greenish. Fluorescence––
fresh material not tested; dried lamellae dull yellow orange (one small area in one a
brilliant orange yellow)
RFLP profile (Fragmentary data for 1 isolate amplified from outlier Washington
collection)—ITS 715 bp; Hinf1 (395-320) diagnostic, also cut in Cfo1, EcoR1, Nde2,
Pvu2; uncut in Rsa1, Xho1; data missing for Pal1, Sal1.
Ecology, range and distribution: Solitary to closely gregarious in humus under mature
second- and old-growth Tsuga heterophylla, Pseudotsuga menziesii,
and/or Picea
sitchensis among Polystichum munitum, Oxalis oregana, Eurhynchium oreganum.
Known from 13 sites (43 collections) in low-lying to montane areas from Oregonʼs
mid-coast to Washingtonʼs Olympic peninsula. October to December.
OTHER SPECIMENS EXAMINED UNITED STATES. OREGON: Benton Co. Ex200-
283 (PNW); GREEN PEAK BLM DENSITY MANAGEMENT STUDY (123.4551°W, 44.366°N) — PRE-
TREATMENT ʻCLEARCUTʼ 2000ʼ 65yo PSME TSHE POMU g1981124c1-04 LLN RLE TFennell (OSC
PNW); ʻHIGH RETENTIONʼ 1900ʼ 65yo PSME TSHE POMU g1991208hx.03 LLN RLE SAR (PNW
DAOM), g2001127hx3 LLN RLE (PNW); KLICKITAT BLM UNIT 3 (TRS: 3S7W21NWSW) — 60yo
PSME TSHE GASH 30.X.2000 Ex200-065 (PNW), 16.XI.2000 Ex200-252 (PNW); KLICKITAT
BLM UNIT 6 (TRS: 3S7W9SWSE) — 50yo PSME TSHE POMU 31X.2000 Ex200-082,-090 (PNW),
14.XI.2000 Ex200-187b (PNW); KLICKITAT BLM UNIT 7 (TRS: 3S7W21NWNE) –– 60yo PSME
TSHE GASH 30.X.2000 Ex200-070 (PNW); RUNNING BEAR BLM UNIT (TRS: 13S7W5, 123.5631°W,
44.4686
°N) — 200YO PSME TSHE POMU LLN1981109.108E RLE (PNW OSC), 6.XI.2000
Ex200-103 (PNW), 28.X.2002 Ex2002-16 (PNW). Lincoln Co. CASCADE HEAD EF, 100M S
TILLAMOOK CO. LINE (123°54ʼ55”W, 45°2ʼ41”N) — late-succ 2nd growth PSME TSHE POMU OXOR
LLN1951109- 19 (WTU PNW); FOGARTY CREEK SP (124.0444°W, 44.8425°N) — mid-succ PISI
TSHE LLN1951109-22 (PNW). Linn Co. BLM CASCADE RA KEEL FLATS (TRS: 12s1E23) –– PSME
BENE POMU 6.XII.1999 Ex199-BK by KScott 991206-KF-5-KS1 (PNW OSC). Polk Co. MARYʼS
PEAK RA BLM UNIT (TRS: 9S7W1NENW) — 2000ʼ 200yo PSME TSHE GASH 26.XI.2001 Ex2001-
112,115
(PNW); PEDEE BLM RESERVE CHRONOSEQUENCE STUDY (123.4885°W, 44.7913°N)

251
FIGURE 3 Phaeocollybia tibiikauffmanii (Holotype, a2011031ox-01). Cluster of primarily mature
basidiomes excavated in lab to expose origins of vertical pseudorhizae. Scale = 1 cm.
Inset: Detail
showing new primordium (arrow) arising from fungal mass at excavated pseudorhizal origin.

252
— 50yo 1900ʼ PSME TSHE POMU 1.XI.2000 Ex200-091 w LLN (PNW); 150yo 1770ʼ PSME TSHE
POMU a1981104o1-01 & o2-02–04 LLN RLE CHibbler (PNW), 5.X.2000 Ex200-045 (PNW),
a2001018o1-15 & o2-25, a2001101o2-30, a2011031o2-03,04,05,06,08 & ox-02, a2011114o1-
01,02,03 & ox-01,
a2021015o2-01, a2021113o2-01,02,03 LLN RLE (PNW). WASHINGTON:
Clallam Co.
KLAHANIE CMPGRD, OLYMPIC NF (124.3000°W, 47.9613°N) — >250yo PISI TSHE
LLN1921018-04
usps3
STrudell (WTU PNW).
Comments A tawny, viscid, conic-campanulate pileus, moderately large size, stuffed
tapering stipe, medium-sized punctate-roughened basidiospores, heteromorphic
cheilocystidia with thick-walled tibiiform intermixed with thin-walled elements, and lack
of clamp connections diagnose P. tibiikauffmanii. Its size, pileus color, tapering stuffed
stipe, vertical-monopodial pseudorhiza, syringaldazine reactivity, and basidiospore size
resemble those of Phaeocollybia kauffmanii (A.H. Sm.) Singer, but the presence of
thick-walled tibiiform cheilocystidia, differences in pigment topography, and absence
of heavily gelatinized, strongly sarcodimitic tissues in the pileal trama clearly separates
P. tibiikauffmanii from the earlier named species. In the field P. tibiikauffmanii
is
differentiated by its more slender aspect and lack of strongly inrolled pileus edge.
Anatomically, P. tibiikauffmanii shares affinities with Phaeocollybia spadicea A.H. Sm.
and P. pseudofestiva A.H. Sm. Young specimens of P. spadicea are readily differentiated
by their dark to blackish brown pilei lacking orange hues, drab stipes usually covered with
recurved, agglutinated, cinnamon-colored fibrillose patches, negative syringaldazine
reactivity of the pileus and lamellar tissues, and smaller basidiospores. Phaeocollybia
pseudofestiva — with similar sized basidiospores, tibiiform cheilocystidia, positive
syringaldazine reactivity, pileus shape, and odor — is distinguished by smaller olive-
green basidiomes with slightly more prominently beaked basidiospores and lacking
thin-walled cheilocystidia.
Molecular data for this species were generated from the single Washington collection,
initially thought to represent an orange form of P. spadicea. The 715 bp ITS region is
much shorter than that estimated for P. kauffmanii, and the Cfo1, EcoR1, Hinf1 and
Nde2 fingerprints support separation of P. tibiikauffmanii from both P. spadicea
and P.
pseudofestiva.
Cystidial morphology and Section Versicolores
There are two distinct cystidial forms represented in Phaeocollybia: thin-walled, ≥2
µm diam, cylindrical / clavate elements versus thick-walled, refractive, narrow-necked
(≤ 1 µm diam) lageniform / tibiiform elements. Clémençon does not directly address
phaeocollybian cystidia in his treatment of hymenomycete anatomy (Clémençon 1997),
but the two cystidial types are similar to those he classifies as lamprocystidia and
leptocystidia within the non-secretory alethocystidia. Such cystidial classification is,
however, complicated by the fact that most western phaeocollybian cystidia both gelify
and appear to serve in a secretory capacity.
A. H. Smith, the first to classify
Phaeocollybia at the subgeneric level (Smith 1957),
divided the genus into two sections — Phaeocollybia, for species characterized by
thin-walled, clavate cheilocystidia, and Versicolores, for species characterized by
thick-walled, narrow-necked, lageniform to tibiiform cheilocystidia. After finding

253
FIGURE 4 Phaeocollybia tibiikauffmanii (LLN1951109-19). 4a. Habit of young basidiome
showing lamellar attachment, uplifted pileus, and pseudorhizal form. Scale = 1 cm
. 4b. Tibiiform
cheilocystidia and basidiospores. Scale = 10 µm.
4a
4b

254
new species from the neotropic western hemisphere, Singer (1970, 1986) revised
Smithʼs sections and added three new ones. Singer separated his sections based on
cheilocystidial morphology + basidiospore length + clamp connections:
TABLE 1. Phaeocollybia sections sensu Singer 1970, 1986
SECTION CHEILOCYSTIDIA
BASIDIOSPORE
LENGTH
CLAMP
CONNECTIONS
Phaeocollybia
clavate to cylindrical > 6.5 µm
—
Subattenuatae
variable, non-capitate > 6.5µm
+
Versicolores
acute or apically capitate > 6.5 µm
—
Radicatae
(sub)capitate < 6.5 µm
+
Microsporae
variable; oft clavate / cylindrical < 6.5 µm
—
Most recently, Bandala and Montoya (1994) proposed two subgenera — Subgenus
Phaeocollybia (with Singerʼs ʻunclampedʼ sections, Phaeocollybia, Versicolores, &
Microsporae) and Subgenus
Fibulophaeocollybia (with Singerʼs ʻclampedʼ sections,
Subattenuatae & Radicatae).
Molecular support for section Versicolores
Amplification and enzymatic digestion of the ITS region from 160 isolates representing
27 Phaeocollybia species and five out-taxa (Norvell 1998a, 2000, 2002; Norvell &
Redhead 2000) revealed nine different ITS lengths within Phaeocollybia (Norvell
1998a). Most ITS lengths and restriction fragment length polymorphisms (RFLPs)
support morphologically based species hypotheses. Table 2 summarizes ITS lengths,
number of restriction sites, and RFLP profiles for eight tibiiform species. Although
preliminary cladistic analyses of the data obtained from western North American
specimens indicate that taxonomic classification at sectional and subgeneric level is
still premature, they do imply at least one monophyletic clade diagnosed by cystidial
morphology in Phaeocollybia. Seven out of eight species characterized by thick-walled
cystidia have isolates with relatively short (<715 bps) ITS regions (T
ABLE 2). The eighth
species (P. pleurocystidiata) is, however, characterized by the longer (740 bp) ITS
region more commonly found in thin-walled cystidiate species.
While RFLP ʻfingerprintʼ patterns have proved extremely useful for species diagnoses,
DNA sequencing should provide more phylogenetically reliable information (
cf.
Norvell 1998a for a discussion of problems associated with inferring phylogenies from
RFLP generated data). Cfo1 and EcoR1 RFLP fingerprints for all 160
Phaeocollybia
isolates show exceedingly subtle differences that are clearly correlated to ITS length
variation. EcoR1 halves the ITS regions in P. californica, P. scatesiae, and the unallied
P. piceae A.H. Sm. & Trappe, making the enzyme an effective diagnostic tool, if not
a phylogenetically informative one. Hinf1 and Pal1 produce far more informative
fingerprints with relatively easily mapped restriction sites. P. spadicea isolates have

255
Species ITS
Cfo
1
Eco
R1
Hinf
1
Nde
2
Pal
1
Pvu
2
Rsa
1
Sal
1
Xho
1
P. californica
(2)
700
400
300
350
350
355
345
390
465
140
95
610
90
0 0 0
P. pleurocystidiata
(4)
740
420
320
390
350
390
350
405
450
190
100
630
80
30
0 0 0
P. pseudofestiva
(4)
700
400
300
360
340
365
335
405
440
165
95
605
95
0 0 0
P. radicata
(2)
710
400
310
380
330
390
320
–––
480
120
100
––– ––– 0 –––
P. rufotubulina
(6)
690
400
290
370
320
365
325
400
435
160
95
600
90
0 0 1
P. scatesiae
(9)
700
400
300
350
350
355
345
390
465
140
95
610
90
0 0 0
P. spadicea
(13)
710
410
300
375
335
395
125
110 80
275
480
120
110
615
95
0 1 0
P. tibiikauffmanii
(1)
715
405
310
380
335
390
325
410 –––
625
90
0 ––– 0
Numbers in each species cell = number of isolates tested. Numbers elsewhere = fragment lengths (in base
pairs) estimated per RE (ITS, Cfo1, EcoR1, Hinf!, Nde2, Pal1, PVU2) or number of restriction sites (Rsa1, Sal1,
Xho1). RFLP profiles in gray boxes are unique for the 25 Phaeocollybia species tested (Norvell1998a); only the
top band is given for Nde2. Isolates not cited elsewhere include Ph. californica
—HOLOTYPE AHSmith55610cal2
(CA) (
MICH); Ph. pleurocystidiata—HOLOTYPE LLN1940330-02ple1 (WA), LLN1930516-01ple2 (OR), MTSeidl
914ple3 (CA), DAOM128270ple4 (CA) (DAOM); Ph. pseudofestiva —LLN1921116-02pse1* (CA), LLN1921121-
01pse6 (CA), LLN1911123-02pse8* (OR), LLN1921123-8pse11(CA); Ph. radicata—AHSmith3763rad2
(CA) (
MICH); Ph. scatesiae—LLN1931104-09sca1 (OR), ISOTYPE KScates1136sca2 (OR), LLN1921104-14sca3
(OR), LLN1921123-01sca4 (CA), LLN1921015-19*sca5 (WA), LLN1921015-16*scc5 (WA),
HOLOTYPE
AHSmith 79286*sca6 (OR) (
MICH11630), LLN1931104-02sca7 (OR), CGoetz33sca9* (OR); Ph. spadicea—
LLN192.11.02-03spd1 (OR), LLN1931023-06spd2 (OR), LLN1921015-26spd3 (WA), LLN1941106-24spg4
(OR), LLN1931104-04sps4 (OR), LLN1921110-22spd5 (OR), LLN1931104-06sps5 (OR), LLN1931114-07spd6
(OR), LLN1931023-07spd7 (OR), LLN1921120-02sps6 (CA), LLN1931114-13spd8 (OR); LLN1921031-
03spd4 (WA), [
DAOM] LLN1921015-10sps1 (WA). Four additional Ph. pseudofestiva isolates from Washington
and British Columbia with subtly different banding patterns were not included in cladistic analyses. Collections
at WTU unless otherwise noted.
instantly recognizable RFLP profiles that are unique within the genus (Hinf1) or within
the thick-walled clade (Pal1). All isolates within this clade have Pvu2 loci
and lack
Rsa1 loci. Only P. spadicea isolates have Sal1 and only P. rufotubulina isolates have
Xho1 sites. None of the nine tested enzymes produce data that support separation
between the P. californica and P. scatesiae (both with estimated 700 bp ITS regions),
two species that are otherwise clearly anatomically and morphologically distinct.
TABLE 2. RFLP-generated data isolated from tibiiform cystidiate western North
American phaeocollybias
Morphological characters have been mapped onto an unrooted Fitch-Margoliash +
Neighbor-joining consensus tree generated from ITS length and restriction site data
from 24
Phaeocollybia species and Stagnicola perplexa (Orton) Redhead & A. H. Sm.
(Figure 5). Six out of eight ʻtibiiformʼ Pacific Northwest phaeocollybias––P. californica,
P. pseudofestiva, P. radicata, P. rufotubulina, P. scatesiae, and P. spadicea––occur
in one clade. Molecular data for P. tibiikauffmanii were too fragmentary to include

256
in the analyses. The eighth species, P. pleurocystidiata, is unique among western
phaeocollybias in its possession of abundant thick-walled tibiiform pleurocystidia. Its
additionally distinct vernal phenology (exhibited by only one other western species,
P. phaeogaleroides Norvell) suggests a possible source for genetic isolation from the
seven other thick-walled species, all autumnal fruiters.
The four species at the tips of the autumnal ʻtibiformʼ clade in Figure 5 have
subtending, branching, cord-like rhizomorphic pseudorhizae and stipes that soon
become hollow. Among other western Phaeocollybias, only Phaeocollybia dissiliens
A. H. Sm. & Trappe
and P. sipei A. H. Sm. (also hollow stiped) have been confirmed
with cord-like pseudorhizae. Both P. spadicea and P. tibiikauffmanii (as well as the
out-lying P. pleurocystidiata) produce basidiomes characterized by vertical monopodial
pseudorhizae and stipes always stuffed with firm to spongy pith. Somewhat more
problematical is placement of P. radicata, which is characterized by clamp connections
and extremely small ellipsoid spores. Here molecular data, obtained from only one
isolate, were fragmentary and conclusions regarding the place of P. radicata within the
clade must be considered premature.
Restriction site data do support a clade diagnosed by thick-walled morphology (Figure
5). Exclusion of the vernal Phaeocollybia pleurocystidiata from that clade, however,
challenges Section Versicolores as circumscribed by both Smith (1957) and Singer
(1970, 1986). Inclusion of the clamped P. radicata in the clade likewise challenges
Section Versicolores as revised by Singer (1970, 1986, 1987) and accepted by Bandala
and Montoya (1994). Recent phylotrees containing
Phaeocollybia sequence data (from
the ITS region in Rees et al., 2002, and large ribosomal subunit in Moncalvo et al., 2002,
and Rees et al., 2003) offer little insight into sectional relationships within the genus.
The phylogeny generated by Moncalvo et al. (2002) from 877 homobasidiomycete
taxa (including four north temperate phaeocollybias) recognizes the /phaeocollybia
clade in ʻeuagarics incertae sedisʼ. The phylotree also supports separation of subgenera
Phaeocollybia and Fibulophaeocollybia as defined by Bandala and Montoya (1994),
placing the ʻclamplessʼ P. attenuata & P. redheadii (Section
Phaeocollybia) and P.
jennyae A. H. Sm. (Section Microsporae) apart from the ʻclampedʼ P. dissiliens (Section
Radicatae). Rees et al. (2002) included two southern hemisphere representatives—P.
ratticauda E. Horak (Section Microsporae) and P. graveolens B. J. Rees & Syme (Section
Radicatae)—in their preliminary phylogeny of Gymnopilus species and related genera,
while Rees et al. (2003) included sequences from the south temperate P. ratticauda and
north temperate P. jennyae in their phylogenetic analyses of 78 species from brown-
spored genera. While all three phylogenies imply monophyly for
Phaeocollybia, the
omission of taxa representing sections
Subattenuatae and Versicolores leaves the Smith
(1957) and Singer (1970, 1986) sectional hypotheses untested. Given the preliminary
nature of current molecular analyses, revision of phaeocollybian subgeneric taxa
awaits analysis of sequence data representing all five ʻtraditionalʼ sections from both
hemispheres.

257
FIGURE 5 Phylogenetic relationships of 8 tibiiform cystidiate species (bold starred) among
themselves and 15 other western
Phaeocollybia species and the outgroup, Stagnicola perplexa,
are shown in an unrooted consensus tree generated from Fitch-Margoliash + Neighbor trees
derived from ITS lengths and estimated Cfo1, Hinf1, Pal1, Pvu2, Rsa1, Sal1 and Xho1 restriction
sites in 150 isolates. Important missing Pal1 and Sal1 data from the only representative tested
prevented inclusion of P. tibiikauffmanii (not shown), while P. radicata (shown) was mapped from
suspect molecular data. Sectional diagnostic characters noted above include CYSTIDIAL FORM (heavy
bar = thick-walled tibiiform cheilocystidia), CLAMP CONNECTIONS (heavy dashed bar), and SPORE
MORPHOLOGY (medium solid bar = small ellipsoid punctate). Lines map differing PSEUDORHIZAL
MORPHOLOGIES (solid = vertical monopodial; narrow-dashed = racemose; wide-dashed = criniform
lateral-monopodial).
oregonensis
ammirati
sipei
rifflipes
piceae
Stagnicola
(outgroup)
fallax
olivacea
luteosquamulosa
‘similis’
attenuata
redheadii
kauffmanii
benzokauffmanii
olivaceaB
gregaria
pseudofestivaB*
rufotubulina*
pleurocystidiata*
(vernal)
spadicea*
pseudofestiva*A
californica*
scatesiae*
radicata*

258
Removal of Phaeocollybia olivacea
from Section Versicolores
One final observation should be made with respect to tibiiform cystidia. Smith
(1957), noting their rarity in other genera, felt that refractive, thick-walled, narrow-
necked cystidia represented a recently derived form in Phaeocollybia. Given their
morphological similarity to the generically significant tibiiform diverticula that are an
integral part of the primordial pellicular sheath (Norvell 1998ab), however, it seems
equally possible that such elements are plesiomorphic and thin-walled cheilocystidia
are derived. If so, then thin-walled more or less clavate cheilocystidia could be regarded
as modified extensions of the central hyphae in the lamellar trama that have evolved to
assume the same secretory functions as their refractive, thick-walled antecedents.
While many western phaeocollybias appear to possess only one cystidial type,
others (e.g. P. rufotubulina, P. spadicea
, P. tibiikauffmanii) possess both thick-walled
lageniform/tibiiform narrow-necked elements and thin-walled clavate elements. Smith
(1957) noted that Phaeocollybia olivacea A.H. Sm.––which he described as having
thin-walled cystidia––also tends to develop thick-walled ʻforerunnersʼ. Bandala and
Montoya (1994), who also observed thick-walled elements when they examined the
holotype, used this to support their transfer of P. olivacea into Section Versicolores. My
examination of all P. olivacea type specimens (Norvell 1998ab), however, uncovered
only one specimen bearing heteromorphic cheilocystidia. In that specimen, the far
more abundant thin-walled clavate elements were sporadically topped with long
thin extensions that were even more rarely capitulate. Thickening of cystidial walls
in this specimen was even more rare. I regard the extensions atop the thin-walled
elements in this specimen as apical regenerations of innately thin-walled elements––a
condition frequently found in old or improperly stored phaeocollybias with thin-walled
cheilocystidia (Norvell 1998ab).
Here the infrequent occurrence of thickened walls—again found very rarely in only
one type specimen of P. olivacea
—is more significant than the slightly more numerous
presence of narrow-necked (but thin-walled) extensions. In any case, present RFLP
data (Norvell 1998a; Figure 5) support exclusion of P. olivacea from the thick-walled
clade and thus from Section Versicolores s. a.
Acknowledgements
Thanks to Joe Ammirati for guidance, financial support, and laboratory facilities at the University
of Washington, Scott Redhead for general support, field assistance, and insightful discussions,
Ron Exeter for good-natured field assistance and innumerable P. tibiikauffmanii collections, and
Ron Exeter, Dick Korf, and Scott Redhead for reviewing this manuscript. David Arora, George
Barron, Roy Halling, Steve Trudell, and the late Harry Thiers generously donated collections of
P. rufotubulina or P. tibiikauffmanii. Joe Ammirati (WTU), Bill Denisen & Joey Spatafora (OSC),
Dennis Desjardin & Harry Thiers (SFSU), Bob Fogel & Robert Shaffer (MICH), Christel Goetz,
Roy Halling (NY), Dave Largent (HSC), Janet Lindgren, Maggie Peeters, Ron Petersen, Scott
Redhead (DAOM), Judy Roger, Maggie Rogers, Kit Scates-Barnhart (deceased), Michelle Seidl,
and Tom Volk are thanked for loaning or donating additional specimens cited in this paper. Luca
Comai, Amy Denton, David Hibbett, Yajuan Liu, François Lutzoni, George Mueller, John Stiller,

259
and Gene Takimoto provided guidance or materials for the molecular work and/or phenetic and
cladistic analyses. Field support provided by the Olympic Natural Resources Center (Forks, WA)
and the Cascade Head Experimental Forest Research Station (Otis, OR) and financial support
from the Daniel E. Stuntz Memorial Foundation, Oregon-Washington Reciprocity Award, and the
Mycological Society of America Graduate Fellowship are gratefully acknowledged.
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