Phaeocollybia in western North America. I. The Phaeocollybia kauffmanii complex

Abstract

Morphological and molecular investigations during a taxonomic reevaluation of the genus Phaeocollybia revealed four new agaric species from British Columbia, Washington, Oregon, and California that are morphologically similar to Phaeocollybia kauffmanii (Smith) Singer. All five species produce large basidiomes with brown pilei, stipes with cartilaginous rinds surrounding dense pith, vertical-monopodial pseudorhizae, large, verrucose or verruculose, apically beaked basidiospores, and thin-walled, clavate cheilocystidia. The most salient morphological characters distinguishing the new species from P. kauffmanii include abundant clamp connections (Phaeocollybia ammiratii), a pink- or purple-brown pileus lacking encrusting pigments in the pileipellis (Phaeocollybia benzokauffmanii), a nonviscid, fibrillose, ochraceous pileus with a trilaminate pileipellis (Phaeocollybia luteosquamulosa), and unusually large basidiospores and subglobose subcapitate pedicellate cheilocystidia (Phaeocollybia redheadii). An emended description of P. kauffmanii accompanies technical descriptions and a key to the five species and newly revealed complex.

Full text

Phaeocollybia in western North America. I. The
Phaeocollybia kauffmanii complex1
Lorelei L. Norvell
Abstract: Morphological and molecular investigations during a taxonomic reevaluation of the genus Phaeocollybia re-
vealed four new agaric species from British Columbia, Washington, Oregon, and California that are morphologically
similar to Phaeocollybia kauffmanii (Smith) Singer. All five species produce large basidiomes with brown pilei, stipes
with cartilaginous rinds surrounding dense pith, vertical-monopodial pseudorhizae, large, verrucose or verruculose, api-
cally beaked basidiospores, and thin-walled, clavate cheilocystidia. The most salient morphological characters distin-
guishing the new species from P. kauffmanii include abundant clamp connections (Phaeocollybia ammiratii), a pink- or
purple-brown pileus lacking encrusting pigments in the pileipellis (Phaeocollybia benzokauffmanii), a nonviscid,
fibrillose, ochraceous pileus with a trilaminate pileipellis (Phaeocollybia luteosquamulosa), and unusually large
basidiospores and subglobose subcapitate pedicellate cheilocystidia (Phaeocollybia redheadii). An emended description
of P. kauffmanii accompanies technical descriptions and a key to the five species and newly revealed complex.
Key words: Agaricales, Basidiomycota, Cortinariaceae, RFLPs, temperate rainforest.
Résumé : Des études morphologiques et moléculaires, conduites au cours d’une réévaluation taxonomique du genre
Phaeocollybia, révèlent l’existence de quatre nouvelles espèces d’agaric en Colombie-Britannique et dans les états de
Washington, Oregon et Californie; ces espèces sont morphologiquement similaires au Phaeocollybia kauffmanii (Smith)
Singer. Les cinq espèces produisent toutes de gros basidiomes avec des chapeaux bruns, des pieds avec une écorce car-
tilagineuse entourrant une moelle dense, des pseudorhizes verticales et monopodiales, de grosse spores verruqueuses ou
verriculeuses avec apicule crochue, ainsi que des cheilocystides claviformes à paroi mince. Les caractères morphologi-
ques les plus distinctifs des nouvelles espèces par rapport au P. kauffmanii sont d’abondantes anses d’anastomose
(Phaeocollybia ammiratii), un pied brun-rose ou brun-pourpre dépourvu de pigments incrustés dans la pellicule du pied
(Phaeocollybia benzokauffmanii), un pied ochre, non visqueux, fibrilleux, pourvu d’une pellicule du pied trilaminée
(Phaeocollybia luteosquamulosa), et des carpophores inhabituellement gros ainsi que des cheilocystides subglobulaires,
subcapitées et pédicellées (Phaeocollybia redheadii). On présente une description amendée du P. kauffmanii et des des-
criptions techniques ainsi qu’une clé pour les cinq espèces et le nouveau complexe découverts.
Mots clés : Agaricales, Basidiomycota, Cortinariaceae, RFLP, forêt ombrophile tempérée.
[Traduit par la Rédaction] Norvell 1076
Introduction
Discovery of Phaeocollybia kauffmanii linked
cryptospecies
The coniferous coastal forests of Washington, Oregon,
California, and British Columbia support one of the richest
concentrations of Phaeocollybia species in the world (Smith
1957; Horak 1977; Redhead and Norvell 1993; Norvell
1998a, 1998b). An endemic species, Phaeocollybia
kauffmanii (Smith) Singer, with basidiomes up to 40 cm tall
and pilei up to 19.5 cm broad, has been recognized as the
largest of all Phaeocollybias (Smith 1957; Horak 1977;
Bandala 1994; Norvell 1998a). This inhabitant of old-
growth forests is additionally significant because of its Strat-
egy 1 designation in the Forest Ecosystem Management As-
sessment Team (FEMAT) reports mandating federal forest
“survey and manage” practices in northern spotted owl habi-
tats (Forest Ecosystem Management Assessment Team
1993; USDA–USDI 1994; Norvell 1995; Castellano et al.
1999). Unfortunately, the unusually large size that makes
P. kauffmanii so recognizable in the field has obscured other
morphological characters so that the name has been misap-
plied to other similar, undescribed taxa in the Pacific North-
west and elsewhere (Norvell 1998a).
Phaeocollybia kauffmanii was first described as a
Naucoria by A.H. Smith (1937) and named in honor of his
mentor, C.H. Kauffman. In 1925, Kauffman explored Wash-
ington’s Olympic Peninsula and Oregon’s Siskiyou Moun-
tains (Kauffman 1930), encountering a robust unidentifiable
Naucoria with a radicating stipe. While exploring the same
region in 1935, Smith discovered what he considered the
same unnamed fungus (Smith 1937). When Smith named the
new species, he designated as the holotype one of his own
more recent (better described and dried) collections from
Oregon (AHS 3523) and listed the Kauffman collections
Can. J. Bot. 78: 1055–1076 (2000) © 2000 NRC Canada
1055
Received October 28, 1999.
L.L. Norvell.2Department of Botany, Box 355325,
University of Washington, Seattle WA 98195, U.S.A.
1Based in part on material submitted for a doctoral
dissertation at the University of Washington, Seattle, Wash.
2Present address: Pacific Northwest Mycology Service, 6720
NW Skyline Boulevard, Portland, OR 97229-1309, U.S.A.
(e-mail: lorelei@teleport.com).
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(CHKauffman 11.X.1925, Lake Quinault, Wash.;
CHKauffman 7.XII.1925, Takilma, Oreg.) among the six
paratypes.
Smith (1937) compared his new species to Naucoria
lugubris (Fries) Heim from Europe and to what he was also
calling N. lugubris from the western United States (AHS
3458), noting that N. kauffmanii produced consistently larger
spores. Twenty years later, in a preliminary monograph on
North American species of Phaeocollybia, Smith (1957)
proposed the name Phaeocollybia spadicea A.H. Smith for
his North American “N. lugubris.” However, he selected one
P. kauffmanii paratype (AHS3339) as holotype of P.
spadicea, at the same time redisposing another
(CHK7.XII.1925) as a third species, Phaeocollybia
californica A.H. Smith, dispositions with which I concur.
Given the inclusion of three taxa in the protologue and the
subsequent inadvertent confusion caused by placing a strik-
ing illustration and description of Gruber 700 (a different
taxon characterized by clamp connections) under the name
“P. kauffmanii”inMushrooms in Their Natural Habitats
(Smith 1949), it is apparent that initially the defining charac-
ters for P. kauffmanii were its large size and brown color.
Unfortunately, the lack of a precise description of
“P. kauffmanii” based upon a single taxon (that represented
by its type, AHS3523) continues to confuse. Arora’s (1986)
observation that “this giant of the genus can usually be rec-
ognized by its size alone” illustrates the common misappre-
hension that any large brown Phaeocollybia represents
P. kauffmanii.
Over 900 Phaeocollybia collections have been made as
part of the current reevaluation of the genus, which was be-
gun in 1991. This research was prompted in part by a collec-
tion of specimens seemingly closely related to P. kauffmanii
sensu typi but with unusually large basidiospores (Norvell
and Redhead 1992; Redhead and Norvell 1993). Examina-
tion of herbarium collections (including Gruber 700)re
-
vealed numerous other specimens identified as P. kauffmanii,
but which, unlike P. kauffmanii, had clamp connections.
(The existence of a kauffmanii-like species with abundant
clamp connections had been noted by J.F. Ammirati in the
1970s but had not been mentioned in any published litera-
ture until the current study (cf. Norvell and Ammirati 1993,
1995, 1996; Norvell 1998a, 1998b)). Evaluation of
basidiospore morphology (Figs. 1–4) and other characters
suggested there were more than just two kauffmanii-like
species in western North America. Comparisons eventually
led to the hypothesis that there are at least five morphologi-
cally distinct taxa characterized by robust stature, viscid
brown pilei, thick fleshy cartilaginous stipes, vertical-mono-
podial pseudorhizae, large (>8 × 5 µm) verrucose, beaked
basidiospores, and thin-walled clavate cheilocystidia
(Norvell and Ammirati 1993). This hypothesis is supported
by internal transcribed spacer (ITS) length polymorphisms,
restriction fragment length polymorphisms (RFLPs), and re-
striction loci (Norvell 1998a). Integration of morphological,
chemical, and molecular data suggests three close species
(P. kauffmanii, Phaeocollybia benzokauffmanii, Phaeo-
collybia redheadii) and two relatively distant species
(Phaeocollybia ammiratii and Phaeocollybia luteo-
squamulosa). These species and their complex are described
below.
Materials and methods
Specimen preparation and examination
Basidiomes were collected following usual protocols except that
all basidiomes were excavated to trace their pseudorhizal origins
(cf. Norvell 1998b). Fresh material was compared with color stan-
dards (Ridgway 1912; Munsell 1976), photographed, and exposed
to a Raytech BS-8 ultraviolet light (UV µ= 354 nm) to determine
fluorescence. Important ephemeral characters and habitat data were
recorded. Small portions of pileus, lamellae, stipe, and pseudorhiza
were spot-tested for macrochemical reactivity with 6% aqueous
KOH, 10% aqueous NH4OH, α-naphthol, FeSO4, guaiac, guaiacol,
and p-dimethylamino-benzaldehyde (PDAB), L-tyrosine, p-cresol,
and syringaldazine following Lennox (1979) and Marr et al.
(1986); only KOH and syringaldazine proved to be informative
within the complex. Surface features, noted using a 16× hand lens
or Wild/Zeiss dissecting microscope, were photographed with a
macrolens on a SLR camera (Olympus OM-1/Canon Eos-Elan) or
with a Nikon SLR camera mounted on a Wild dissecting micro-
scope. Fresh tissues, mounted in soap–water solution (50 mL
H2O + 3 drops detergent), 3–6% aqueous KOH, and (or) Melzer’s
reagent, were examined through a compound microscope (Zeiss
standard or Leitz DMRB). Specimens were dehydrated using low-
heat forced-air portable food dehydrators (Sigg, American Harvest
FD 50/30, Waring 11DF10).
Tissue preparations were rehydrated in moist chambers,
mounted in Melzer’s, soap-water solution, and (or) KOH, and ex-
amined under a Leitz DMRB light microscope using bright-field,
phase-contrast, and differential interference contrast (Nomarski)
optics. Scanning electron micrographs were made with a JEOL
scanning microscope at 10 kV from rehydrated samples fixed in
OsO4, dehydrated in an ethanol series, critically point dried in
Freon TF, and sputter coated with Au/Pd.
DNA isolation, polymerase chain reaction amplification,
and RFLP digests.
Dried lamellae were ground to a fine powder in liquid nitrogen
and DNA was extracted using CTAB (hexadicetyltrimethyam-
monium bromide) procedures (Rogers 1994; Norvell 1998a). ITS4
(5′-TCC TCC GCT TAG TTA TAT GC-3′) and ITS5 (5′-GGA AGT
AAA AGT CGT AAC AAG G-3′) oligonucleotides specific for the
ITS1 + 5.8S rDNA + ITS2 region (hereafter referred to as the “ITS
region”) were used to prime target segments of nDNA in polymer-
ase chain reaction (PCR) amplifications following Bruns et al.
(1990), White et al. (1990), and Rogers (1994). Target DNA was
also amplified directly from spores without extraction following
protocols developed by Norvell (1998a): spores were suspended in
1× TE (10 mM Tris-HCl (pH 7.4), 1 mM EDTA) buffer, vortexed
with glass microbeads, and plunged into liquid nitrogen to fracture
walls. The exposed DNA was amplified and immediately digested
with restriction enzymes.
All 44 DNA samples were singly digested with nine restriction
enzymes (CfoI, EcoRI, HinfI, NdeII, PalI, PvuII, RsaI, SalI, and
XhoI). RFLPs were revealed via electrophoresis in agarose gels
stained with ethidium bromide and photographed under UV illumi-
nation. Basepair lengths (bp) were determined using a natural log
conversion formula applied to bands on gels (Norvell 1998a).
Species descriptions
Descriptions are based on holotypes and RFLP-tested collec-
tions. General nonstandard colors in lower case are accompanied
by italicized color references. Ridgway (1912) colors are capital-
ized 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) Pink-
ish Cinnamon =Pale Pinkish Cinnamon +Pinkish Cinnamon);
© 2000 NRC Canada
1056 Can. J. Bot. Vol. 78, 2000
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bracketed Munsell (1976) alpha-numeric color ranges (e.g., [2.5Y
6-8/1-4]) follow Ridgway (1912) references. Basidiospores from
the stipe apex were measured when spore prints were lacking.
Mean basidiospore dimensions with standard deviations are ac-
companied by ranges enclosed in parentheses. “L+ll/cm” is the
number of lamellae + lamellulae per cm at the “edge” of the pileus
and at the “midpoint” between the edge and the 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.
Technical terms fully explained in Norvell (1998b) but not defined
in Hawksworth et al. (1995) include (i) “pileostipiticarpic mono-
velangiocarpic,” pre-emergent differentiation of first stipe, then
pileus by a basidiome initial contained within a primordial sheath
(“endocarpic” sensu Clémençon 1997); (ii) “tibiiform diverticula,”
refractive, secretory structures on mycelial and cortical hyphae (di-
agnostic for Phaeocollybia); (iii) “vertical-monopodial,” pseudo-
rhizal model describing solitary, unbranched pseudorhizae
originating from undifferentiated noduli sensu Clémençon (1997).
Examined collections
“Specimena selecta!” list significant collections not cited else-
where in the text (cf. Norvell 1998afor an itemized list of collec-
tions examined before 1998). Collections are housed in the
University of Washington Herbarium (WTU) unless otherwise
noted (PNW, PNW Mycology Service Herbarium; cf. also
Holmgren et al. 1990). Depositor and collector abbreviations in-
clude JFA (J.F. Ammirati), GTN (G.T. Norvell), LLN (L.L.
Norvell), SAR (S.A. Redhead), JR (J. Roger), and AHS (A.H.
Smith). RFLP codes are denoted by subscripts, and asterisks (*)
flag Strategy 1 collections (Norvell 1995) that are of significance
to U.S. forest management policy. Separate collection dates are not
listed for date-based collection numbers (e.g., LLN1941106-
18amm16 = Norvell November 6, 1994 collection 18, RFLP code
amm16).
Taxonomy
Phaeocollybia kauffmanii (A.H. Smith) Singer, Rev. Mycol.
5: 11. 1940. Emended Figs. 1–2, 5–11
⬅Naucoria kauffmani [sic] A.H. Smith, Mycologia
29: 52. 1937.
© 2000 NRC Canada
Norvell 1057
Fig. 1. Basidiospore mean dimensions in the Phaeocollybia
kauffmanii complex. Sampled from approximately 250 collec-
tions; means computed from a minimum of 20 spores per speci-
men per collection. (amm, P. ammiratii; ben, P. benzokauffmanii;
kau s. s., P. kauffmanii; lut, P. luteosquamulosa (lut-x, excluded
small-spored collections); red, P. redheadii).
Figs. 2–4. Scanning electron micrographs of basidiospores. Scale
bar=1µm. Fig. 2. Phaeocollybia kauffmanii (holotype). Fig. 3.
Phaeocollybia ammiratii. Fig. 4. Phaeocollybia redheadii
(holotype).
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Pileus: (30) 45–150 (195) mm diam., conic-convex and
usually sharply minutely papillate when young, expanding to
broadly campanulate with broad low umbo and slightly
decurved or occasionally upraised margin, edge inrolled at all
times; surface glabrous, smooth, viscid to heavily glutinous,
opaque, nonstriate, pristine basidiomes most often zonate with
deep orange-brown or bright brownish orange disc (Tawny,
Russet, Amber/Mikado/Mars/Verona Brown [5YR 3-5/4-8];
Kaiser/Sanford’s Brown, Ochraceous Tawny, Hazel [2.5YR 3-
5/6-8]), pale to bright brownish orange margin (Ochraceous
Buff/Orange, (Ochraceous) Tawny, Zinc Orange, Mars Yellow
[6.25YR 5-7/7-10]; Amber Brown, Chestnut, Cinnamon Buff,
Orange Rufous), and deep yellow-brown or orange-brown
edge (Buckthorn/Dresden Brown, Antimony Yellow [10YR 4-
5/5-7]; Russet, (Ochraceous) Tawny, Amber/Argus/Cinnamon
Brown [5YR 3-5/4-8]), occasionally ± uniformly deep orange-
brown or bright brownish orange overall, blackish or deep
brown where damaged; dried pileus generally reddish copper,
frequently with darker vinaceous or blackish pointed disc.
Context: Firm, 3–10 mm thick at the disc, when young pale
yellow cream or smoky pink (Pale Ochraceous Salmon, Pink-
ish/Cartridge Buff, [2.5Y 8-9/1-4]; Pale Pinkish Cinnamon,
Pale Ochraceous Buff [7.5YR 8-9/2-4]), when wet often de-
veloping a 1–3 mm wide brown zone adjacent to the lamellae
or between otherwise confluent pilear and stipe contexts.
Odor: Mildly to strongly farinaceous, occasionally not dis-
tinctive. Taste: Slightly to strongly farinaceous, occasionally
slightly bitter. Lamellae: Free or sometimes attached by a
small tooth, ventricose with end nearest stipe almost truncate,
edges uneven to slightly serrulate, eroded in age, thin, occa-
sionally forked, lamellulae in 5–7 tiers with 2–6 interspersed,
narrow when young, becoming very broad when mature (to
20 mm, mean length:width ratio 4 (young to mature range
1.3–9)), crowded when young, becoming subcrowded in age
(L+ll/cm: 13–38 at edge, 8–20 at midpoint), pale buff when
very young ((Pale) Pinkish/Cartridge Buff [2.5Y 6-8/1-4])
darker as maturing ((Light/Pale) Ochraceous Buff, Pinkish
Cinnamon [6.25YR 6-8/5-8]; Pale Salmon Color), and dull
smoky golden brown in age (Clay Color, Cinnamon Buff,
Avellaneous, Buckthorn/Dresden Brown, Cinnamon [10YR 5-
7/5-7]).Veil: When present evident as scattered fibrils or
scant fibrillose patches on stipe apex. Stipe: Central to slightly
eccentric, terete, aerial length 40–60 (110) mm, combined
length with pseudorhiza up to 400 mm, apex 9–25 mm diam.,
usually widening to 10–37 mm above ground level before ta-
pering downwards to the pseudorhiza; surface matte-smooth,
appressed fibrillose (hand lens) with fine longitudinal
striations, moist, apex pale yellow-brown or buff when young
((Light/Pale) Pinkish Cinnamon [7.5YR 6/6-8/2];
Vinaceous/Light Ochraceous Salmon [5YR 7-8/2-6]), develop-
ing darker tones when mature (Pinkish/Cinnamon Buff [10YR
7-8/4-8]; Cinnamon [7.5YR 4-6/5-7]), below darkening to
deep orange brown (Army/Natal Brown [5YR 3-5/2-5]), when
cut staining orange then brown; cartilaginous rind 1–3 mm
thick, context firm, pale light pink or dull ivory, turning deep
orange-brown when crushed or cut, long, narrow, yellow- or
orange-brown lines occasionally visible when water-soaked.
Pseudorhiza: Generally 2/3 to 5/6 of overall stipe length,
gradually tapering to a blunt origin, overall deep red or orange
brown (Liver Brown [7.5R 2-3/4-5]; Kaiser/Mars/Mikado
Brown, Tawny [5YR 3-5/4-8]) or paler (Light Ochraceous/Cin-
namon Buff [6.25YR 5-7/6-8, 10YR 5-6/5-6]) at the origin.
Spore print: Cinnamon brown (Cinnamon [7. 5YR 5/6],
Buckthorn Brown [10YR 5-6/4-6]).
Basidiospores: 8.5×5±0.4×0.2µm (overall range 7.5–
10 × 4–6, cf. Fig. 1), terete to slightly compressed, inequilat-
erally amygdaliform or limoniform in profile, fusoid-elliptical
or almost naviculate in face view, verruculose to verrucose ex-
cept on the 0.5- to 1.5-µm-long mammiform beaked apex and
eccentric prominent apiculus, suprahilar plage an area of
slightly lower ornamentation visible in scanning electron mi-
crographs but not readily discernible under oil immersion, me-
dium pale to dark amber in KOH, pale yellow-amber in H2O,
and dextrinoid in Melzer’s. Cheilocystidia: Abundant, ema-
© 2000 NRC Canada
1058 Can. J. Bot. Vol. 78, 2000
Fig. 5. Phaeocollybia kauffmanii. Excavated basidiomes from
Jackson State Forest, California, are shown in situ. The vertical-
monopodial pseudorhizae of mature and immature basidiomes
have been exposed down to the nodulus (hidden from view). De-
velopment of the primordium, still covered by its pellicular
sheath, is advanced, with both pseudorhiza and stipe formed and
pilear differentiation begun. Scale bar = 10 mm.
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nating from the lamellar trama to form a dense highly gelati-
nous layer; irregularly cylindrical or narrowly clavate, isolated
cheilocystidia sometimes catenate, terminal lengths variable
(up to 50 µm), septa 2–4 µm diam., in mature basidiomes
these elements intermixed with (occasionally) broadly clavate
or (very rarely) subcapitate (to 6 µm diam.) pedicellate ele-
ments; all elements thin-walled, highly gelatinized, hyaline or
(in older basidiomes) with oily pale amber or brownish con-
tents, often developing long filamentous apical outgrowths in
very old or stored fresh specimens. Pleurocystidia: Absent.
Basidia: (2-) 4-spored, 30–38 × 6–7 [10] µm, clavate, hyaline.
Pileipellis: A bilaminate ixocutis with very thick (200–
700 µm) suprapellis of long, branching, cylindrical, narrow
(2–4 µm), thin-walled, highly gelatinized, hyaline hyphae with
refractive septa, this layer embedded in a thick colorless gelat-
inous matrix and overlying a much thinner (60–180 µm) dull
orange brown to yellow orange subpellis with inflated hyphae
(up to 12 µm from 3–5 µm wide septa) and encrusting,
© 2000 NRC Canada
Norvell 1059
Figs. 6–9. Phaeocollybia kauffmanii. Fig. 6. Habit: longitudinal section showing inrolled pileus edge and cartilaginous stipe cortex.
Scale bar = 5 cm. Fig. 7. Apically beaked basidiospores and basidium. Scale bar = 10 µm. Fig. 8. Thin-walled narrowly clavate
cheilocystidia. (Figs. 6–8 from Oregon’s Mt. Hood National Forest, LLN1941022-04,20). Scale bar = 10 µm. Fig. 9. Basidiospores,
cheilocystidium, basidia (both 4- and 2-sterigmate forms), and basidiole from holotype collected in 1935 near Lake Tahkenitch, Oregon
(AHSmith 3523, MICH). Scale bar = 10 µm.
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intraparietal and intercellular pigments. Stipitipellis: A narrow
(100–150 µm) compact layer of parallel, short-branched,
thick-walled, narrow (2–4 µm), highly gelatinized, hyaline
hyphae slightly inflated at the septa and encrusted with dark
brown gelatinous pigments. Pseudorhizal pellis: Approxi-
mately 300 µm thick, of melanized or dark brown hyphae with
moderately coarse encrusting and brownish (often coagulated)
intraparietal pigments; outermost hyphae bearing densely
abundant tibiiform diverticula. All tramal tissues:
Sarcodimitic to some degree, highly gelatinized, hyaline or
pale amber, and with oleiferous hyphae present throughout; in
the pseudorhiza strongly sarcodimitic, with parallel, cylindri-
cal, long (>200 µm), wide (30 µm), thick-walled (3 µm), rigid
vessel hyphae intermixed with and supported by less conspic-
uous highly branched, thin-walled (up to 0.5 µm), narrow (2–
3µm) flexuous hyphae; in stipe apex flexuous hyphae widen-
ing to 3–4 µm with vessel hyphae becoming narrower (7–
15 µm) and thinner walled (1 µm); in the pileus both hyphal
types also present; in the lamellae parallel, the central region
of long, 3–4 µm diam., thick-walled hyphae intermixed with
occasional highly branched, narrow, flexuous hyphae flanked
by a narrow zone of 2- to 3-µm-wide elements giving rise to a
compact subhymenium. Tibiiform diverticula: Abundant on
mycelia and primordial and pseudorhizal pelli and frequent on
pellicular sheath remnants on stipe apex, up to 15 (20) ×
1µm, hyaline to slightly straw-colored, highly refractive with
no septum between base and hypha, with or without
(sub)globose head and (or) apical droplet. Clamp connec-
tions: (True and false), rare to occasional in stipitipellis and
pseudorhizal pellis, absent in all other tissues.
MACROCHEMICAL REACTIONS: Syringaldazine: immediately
deep magenta overall (restricted to pseudorhiza or patchy
when very young); KOH: orange pileus cuticle becoming
brown. Fluorescence: young basidiomes with dull white to
pale yellow lamellae; mature basidiomes with brilliant (pale)
yellow lamellae, orange pileal context, purplish stipe context
(apex) and dull white pseudorhizal cortex.
RFLP PROFILE (FIGS. 10–11): ITS region of eight isolates =
740 bp (DNA from holotype too degraded for RFLP analy-
sis). PalI(235-230-105-90-80), the most informative en-
zyme. Also cut by CfoI, EcoRI, HinfI, NdeII; uncut by
PvuII, RsaI, SalI, and XhoI.
© 2000 NRC Canada
1060 Can. J. Bot. Vol. 78, 2000
Fig. 10. Uncut ITS, HinfI, and PalI RFLP profiles in the Phaeocollybia kauffmanii complex (restriction bands depicted on a linear
scale). Abbreviations as in Fig. 1.
Fig. 11. PvuII, RsaI, and Sal RFLP profiles in the Phaeocollybia kauffmanii complex (restriction bands depicted on a linear scale). Ab-
breviations as in Fig. 1.
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ECOLOGY AND DISTRIBUTION: Mycorrhizal. Solitary, scattered
or gregarious; early to late autumn; often in loamy soils with
high humus content in coastal or inland mesic closed-canopy
mixed coniferous rainforests; less frequent in poor soils in
open mixed coniferous–deciduous (Pinus, Pseudotsuga,
Lithocarpus) forests. Widely distributed and relatively com-
mon (149 collections verified to date), from southern British
Columbia to northern California (Mendocino County) and
east to Idaho.
HOLOTYPE: United States, Oregon, Douglas County, Lake
Tahkenitch (124°8′48′′W 43°47′50′′N). November 18, 1935,
AHSmith3523kau8 (MICH 11572).
Specimena selecta!
CANADA: British Columbia: Vancouver Island, Upper
Carmanah Valley, LLN1931030-26 (DAOM). UNITED
STATES: California: Del Norte County, Crescent City,
23.XI.1956 AHS55616* BIsaacs (MICH); Mendocino County,
Jackson State Forest Road 409 approximately 2.5 km from
Road 408, LLN1921124-12kau1 SAR GLBarron (DAOM,
WTU); Shasta County, Castella, LLN1961129-04a JFA. Idaho,
Boundary County. RHPetersen s.n.(TENN 23245). Oregon:
Clackamas County, Wildcat Mountain, Mt. Hood National
Forest, Road 2609, LLN1941028-02 w JR; Multnomah
County, Mt. Hood National Forest, Larch Mountain summit,
LLN1931024-09kal5 wGTN; Polk County, PeDee Bureau of
Land Management Reserve Chrono Study, LLN1981104.O2-
01 w RLExeter (PNW); Tillamook County, Cascade Head Ex-
perimental Forest, FR 1861, LLN1931114-01kau2 wGTN.
Washington: Clallam County, Olympic National Park, Rugged
Ridge Trail, LLN1921023-01kab2, -09kau6; Grays Harbor
County, Olympic National Park, Lake Quinault, Colonel Bob’s
Trail, LLN1941002-01kaa13 WSteglich; King County,
Snoqualmie Pass, Asahiel Curtis campground, 4.X.1978
WJSundberg3641 (SIU); Pierce County, Mt. RainierNational
Park, Carbon River, 06.X.1952 AHS40667* (MICH);
Snohomish County, Mt. Baker-Snoqualmie National Forest,
Barlow Pass, 07.IX.1992 JFA10696kab12, LLN1920921-01kab1;
Whatcom County, Silver Fir Camp, 08.XI.1987 JFA9662* (as
P. piceae).
Selected illustrations
Smith (1937): black and white photograph (presumably of
type); Bandala (1994): line drawings, scanning electron mi-
crographs of type basidiospores; Norvell (1998b): develop-
mental drawing. (N.B. I have not examined the specimens
identified as P. kauffmanii in Smith (1973), Smith and Weber
(1980), Bessette and Sundberg (1987), and Phillips (1991).)
Comments
In the field, P. kauffmanii sensu stricto can be differenti-
ated from other species in the complex by its much brighter
(almost orange when young), viscid pileus that never devel-
ops ochraceous (q.v. P. luteosquamulosa) or drab (q.v.
P. benzokauffmanii) tones. The vinaceous red-brown acute
umbo and metallic copper margin of the dried pileus is also
diagnostic for P. kauffmanii (see Table 1). Phaeocollybia
kauffmanii produces the smallest basidiospores in the com-
plex (with a mean of 8.5 × 5 µm for 122 collections, cf.
Fig. 1). The P. kauffmanii holotype has a preponderance of
narrowly clavate cheilocystidia (Fig. 9); the more bulbous
cheilocystidia depicted by Smith (1957) are much rarer than
implied by his illustration. Other key characters include a
strong magenta reaction in syringaldazine, pileal supra-
pellicular hyphae embedded in a thick gelatinous colorless
matrix, presence of encrusting pigments in the subpellis, and
an absence of clamp connections in pileal and lamellar tis-
sues.
Although P. kauffmanii sensu stricto is relatively common
within the complex, only five (including type and one
paratype) of the original 16 collections cited in Smith’s 1957
monograph remain identified as P. kauffmanii. On a draft of
that monograph (on file at MICH), Smith noted after
P. kauffmanii to “check all collections for clamps,” suggest-
ing that he had detected clamp connections in some of his
collections. He apparently made no redispositions of those
collections into different taxa. Six of the sixteen collections
cited in 1957 have clamped hyphae in the basidiomes. These
collections are redisposed below under P. ammiratii.Five
others (including three paratypes) contain large-spored
basidiomes redetermined as P. redheadii (q.v.). As noted
above, Smith (1957) reassigned the two remaining paratypes
to species well outside the kauffmanii complex.
Phaeocollybia kauffmanii has also been reported from
eastern North America (Hesler 1949; Bigelow and Barr
1966). Redhead and Norvell (1993) noted that Hesler17217
(TENN) represented an unidentified taxon and not
P. kauffmanii. Re-examinations of Bigelow13570 (NY) from
Vermont reveal that the clamp connections previously re-
ported by Redhead and Norvell (1993) were in fact misinter-
pretations of flexuous hyphal branch apices in sarcodimitic
tissues (Norvell 1998a; Norvell and Redhead 2000). Until
collections of mature basidiomes are made, however, spe-
cific identification of the immature Vermont material still re-
mains tentative. Phaeocollybia kauffmanii is also reported as
the most common Phaeocollybia in the Pinus–Abies forests
in Mexico (Guzmán et al. 1987; Bandala-Muñoz et al. 1989;
Bandala 1994).
Phaeocollybia ammiratii sp. nov. Figs. 1, 3, 10–16
Pileus 40–80 mm latus, campanulatus, umbonatus vel
papillatus, viscidus, glaber, fulvus; caro pallida; odor
cucumeris vel farinaceus; sapor amarus; lamellae luteolae vel
luteofuscae, fulvo-brunnescentes e sporis; stipes 25–60 × 6–
15 mm supra terram, usque ad 300 mm longus cum
pseudorhiza attenuata, glaber, farctus, versus apicem lamellis
subconcolorus, versus subapicem vinascens. Sporae 8.4–9.2 ×
5–6 µm, amygdaliformes, verruculosae vel verrucosae cum
rostro mammiformi, glabro usque ad 1–1.5 µm et cum apiculo
excentrico glabro; ixocutis cum suprapelle crassa, gelatinosa,
hyalina et subpelle cum pigmento diffuso et/vel aliquando
intraparietali, infrequens encrustato; cheilocystidia cylindracea
vel angustoclaviformia, 4 µm lata, gelatinosa, hyalina.
Hyphae fibulatae in texturis omnibus. Ad terram in silvis
coniferarum mixtarum.
HOLOTYPUS: United States, Oregon, Clackamas County, Mt.
Hood National Forest, Wildcat Mountain, October 28,
1994, L.L. Norvell 1941028-10amm15 cum J. Roger (WTU,
isotypus PNW).
ETYMOLOGY: In honor of Dr. Joe Ammirati, who first noted
the presence of a P. kauffmanii-like species with clamp con-
nections in the Pacific Northwest.
© 2000 NRC Canada
Norvell 1061
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Pileus: 40–80 (115) mm, convex with tightly inrolled outer
margin, expanding to broadly campanulate with low, fre-
quently papillate umbo and incurved to straight outer margin,
surface glabrous, viscid to glutinous when wet, opaque,
nonstriate, when young dull ochre to pale butterscotch overall
(Antimony Yellow, Isabella Color, Ochraceous Tawny [10YR 4-
5/6-8]) or distinctly zonate with red-brown disc (Russet, Ve-
rona Brown [5YR 3-4/4-8]), tawny outer margin (Tawny,
Sayal/Amber Brown [5YR 4-5/7-9]), and pale yellow-brown
edge (Buckthorn Brown [10YR 4-5/6-7]), becoming tawny to
butterscotch overall in maturity; dried pileus distinctly metal-
lic, pale copper. Context: Confluent in pileus and stipe, 3–8
(12) mm thick at disc, very pale pink- or yellow-white (Pale
Pinkish Cinnamon [10YR 7-8/2]), when wet often developing
a 1–3 mm wide dark brown band adjacent to lamellae. Odor:
Distinctly cucumber-farinaceous when cut or crushed. Taste:
Somewhat unpleasant and often bitter, reminiscent of bitter
cucumbers. Lamellae: Free to narrowly attached, ventricose,
thin, even young edges soon becoming serrulate; lamellulae in
3–7 irregularly intermixed tiers, narrow and crowded when
young, broader (to 10 mm, (mean length:width ratio 3.3–6.3)
and close in mature specimens (L+ll/cm: 11–30 at edge, 9–20
at midpoint), orange-cream (Light Ochraceous/Cream/Cinna-
mon/Pale Pinkish Buff [2.5Y 6-8/26]) when young, orange
tones later obscured by cinnamon spores. Veil: Occasionally
evident as fibrillose patches on aerial stipe or young pileus
margin. Stipe: Central to eccentric, aerial length 25–60
(105) mm, combined length with pseudorhiza up to 300+ mm,
© 2000 NRC Canada
1062 Can. J. Bot. Vol. 78, 2000
P. kauffmanii P. ammiratii P. benzokauffmanii P. luteosquamulosa P. redheadii
Young pileus color Orange to orange
brown Tawny yellow to
butterscotch Drab to purple-
brown Ochre Pale tawny to red-
brown
Pileus moisture Viscid/glutinous Viscid/glutinous Viscid/glutinous Moist to dry Viscid/glutinous
Pileus surface Smooth Smooth Smooth Appressed-scaly Smooth
Dried pileus color Metallic copper with
dark vinaceous
acute umbo
Metallic coppery
gold Matte black Matte ochre to
dirty gold Shiny mahogany or
dark metallic copper
Syringaldazine
(magenta) All tissues strong Pseudorhiza
weak/moderate All tissues strong Negative All tissues strong
Clamp connections Very rare
(stipitipellis) Abundant Absent Absent Very rare (stipitipellis)
Pileipellis Bilaminate Bilaminate Bilaminate Trilaminate Bilaminate
Suprapellis Thick clear gelati-
nous matrix Thick clear gelati-
nous matrix Moderate clear
gelatinous
matrix
Thin yellow layer
over gelatinized
clear mediopellis
Thick clear gelatinous
matrix
Mean spore
dimensions (µm) 8.5×5 9×5.5 9×5.5 9.8×6 10×6
Dominant cheilo-
cystdial form Mix of narrow and
medium clavate Cylindrical to nar-
rowly clavate Mix of narrow and
medium clavate Clavate with
inflated apices Subglobose subcapitate
pedicellate
Restriction enzymes Pal, Pvu, Rsa, Sal Pal Rsa Hinf, Pal, Sal Pal, PvuII
Note: Boldface indicates characters that are unique in the complex; italics indicates a character that is secondarily diagnostic.
Table 1. Key characters of the Phaeocollybia kauffmanii complex.
Fig. 12. Phaeocollybia ammiratii. Freshly excavated basidiomes of holotype on Wildcat Mountain, Mt. Hood National Forest, Oregon.
Arrowheads indicate location of diagnostic vinaceous “flush” on lower stipe and upper pseudorhiza. Scale bar=5cm.
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apex 6–15 (18) mm diam., swollen and ventricose when im-
mature, ± equal to ground level when mature; surface
appressed fibrillose (hand lens) with occasional detached fi-
brils, longitudinally lined, and dry; apex generally concolorous
with gills, pale cream or pinkish white (Vinaceous/Light Pink-
ish Cinnamon, Ochraceous Buff [7.5YR 6-8/4-8]) when young,
often flushed with orange in age, lower stipe grading to tawny
with a characteristic dingy vinaceous tinge above and below
ground level (Mikado/Kaiser Brown, Fawn Color, Cinnamon
Drab [5YR 4-6/1-6]); stuffed, 2–4 mm thick cartilaginous rind,
context firm, pale cream to pale pinkish white (Pale Pinkish
Cinnamon, Chamois [7.5YR 8/2]); dried stipes distinctly bur-
gundy. Pseudorhiza: Vertical-monopodial, up to 5/6 of over-
all length, gradually tapering to a fleshy narrow straight or
loosely coiled blunt origin, vinaceous to tawny above a pale
buff to salmon origin.
© 2000 NRC Canada
Norvell 1063
Figs. 13–16. Phaeocollybia ammiratii (holotype). Fig 13. Habit: longitudinal sections (two left) and exterior view (right). Note that the
mature pileus edge in this less robust species may be downturned, but not inrolled as elsewhere within the complex. Scale bar=5cm.
Fig. 14. Clamp connections (arrowheads) on cylindrical to narrowly clavate cheilocystidia. Scale bar = 10 µm. Fig. 15. Basidia with
clamp connections (arrows). Scale bar = 10 µm. Fig. 16. Verrucose-ornamented limoniform basidiospores. All Phaeocollybia
basidiospores have a smooth apical callus, but not all are “beaked” as those produced within the kauffmanii complex. Scale
bar=10µm.
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Spore print: Pale cinnamon brown (Pinkish Cinnamon
[7.5YR 6/4-6]).
Basidiospores: 9 × 5.5 ± 0.3 × 0.4 µm (overall range 8.4–
9. 2 (10) × 5–6 (6.5) µm), terete or slightly compressed,
amygdaliform to limoniform with inconspicuous to pro-
nounced apical beak in profile, fusoid-elliptical in face view,
verruculose to verrucose except on the 1- to 1.5-µm-long
mammiform beaked apex and eccentric apiculus, suprahilar
plage an area of decreased ornamentation barely visible with
oil immersion, amber or orange-amber in KOH, paler in H2O,
weakly dextrinoid in Melzer’s. Cheilocystidia: Cylindrical to
narrowly clavate, thin-walled, abundant, emanating from the
lamellar trama to form a distinct gelatinous layer, 2–5 µm
wide with apex usually only slightly swollen, lengths variable,
from very short to >45 µm long, often with medallion clamp
connections, walls heavily gelatinized, usually hyaline, occa-
sionally with pale yellowish oily contents, often developing
long filamentous apical outgrowths in extreme age or after
long storage. Pleurocystidia: Absent. Basidia: (2-) 4-spored,
projecting up to 10 µm above developing hymenium, 29–35
(40) × 7–8 µm, hyaline. Pileipellis: A bilaminate ixocutis
with a 150–400 µm thick suprapellis of radially aligned, cy-
lindrical, narrow (3–5 µm), hyaline hyphae with medallion
clamp connections, this layer loosely embedded in a clear ge-
latinous matrix and overlying a moderately thick (120–
300 µm) orange subpellis of inflated hyphae (5–12 µm wide),
intercellular and intraparietal pigments present. Stipitipellis:
A 200-µm-thick layer of parallel, long (>200 µm), branched,
cylindrical, highly gelatinized, thick-walled, brownish hyphae.
Pseudorhizal pellis: Similar to stipitipellis, outermost hyphae
bearing abundant tibiiform diverticula. All tramal tissues:
Gelatinized and hyaline; in the pseudorhiza strongly
sarcodimitic, with wide, thick-walled, cylindrical, rigid vessel
hyphae intermixed with and supported by less conspicuous,
thin-walled, branched, flexuous hyphae; from stipe to pileus
flexuous hyphae increasing in number as vessel hyphae be-
come more narrow, thin-walled, and distorted; in the lamellae
parallel, the central region composed of 2- to 10-µm diam.,
thick-walled hyphae intermixed with occasional thin-walled,
narrow, branched flexuous hyphae, flanked by a narrow zone
of 2-µm-wide hyphae giving rise to a compact subhymenium.
Tibiiform diverticula: Abundant on mycelia and primordial
and pseudorhizal pelli, frequent on pellicular sheath remnants
on stipe apex, 4–20 × 0.5–1 µm, hyaline, highly refractive,
aseptate, with or without (sub)globose head and (or) apical
droplet. Clamp connections: Frequent to abundant, medal-
lion-like, readily visible on suprapellicular hyphae and cheilo-
cystidia, less visible but present at basidial bases; also present
in tramal tissues and stipitipellis.
MACROCHEMICAL REACTIONS: Syringaldazine: pseudorhizal
cuticle alone weakly reactive (pale magenta after 30 min);
KOH: no change or slightly brown. Fluorescence: intense
pale yellow in young lamellae, becoming less intense as
spores obscure lamellar faces during maturation.
RFLP PROFILE (FIGS. 10–11): ITS region for 13 isolates includ-
ing type = 715 bp. PalI (400-225-90) most informative en-
zyme. Also cut by CfoI, EcoRI, HinfI, and NdeII. Uncut by
PvuII, RsaI, SalI, and XhoI.
ECOLOGY AND DISTRIBUTION: Presumably mycorrhizal. Soli-
tary to clustered in the early autumn, under mixed conifers
(Picea, Tsuga, Abies) or in mixed deciduous–coniferous for-
ests (Pinus, Lithocarpus, Pseudotsuga). Not uncommon (89
collections verified to date) in humid coastal and inland re-
gions in southern British Columbia, Washington, Oregon,
and northern California.
SPECIMENA SELECTA!: CANADA: British Columbia: Mt.
Seymour, Vancouver Watershed, LLN1931006-01amb3
SGamiet. UNITED STATES: California: Humboldt County,
Big Lagoon County Park, LLN1921119-01amb2 w
DLLargent, SAR (DAOM, WTU); Mendocino County Jack-
son State Forest near Mendocino, 19.XI.1960 HPeters192
(SFSU as P. radicata); Shasta County, Castella,
LLN1961129-01 JFA; Sonoma County, Camp Meeker WD,
Camp Heera, 13.I.1968 DLLargent3584 (HSC 2682* as
P. kauffmanii). Oregon: Benton County, Bureau of Land
Management Green Peak Density Study, LLN1981124C2-02
w RLExeter; Clackamas County, Mt. Hood National Forest
Road 2609, Wildcat Mountain, LLN1941028-14amm14 w JR;
Multnomah County, Mt. Hood National Forest, Larch Moun-
tain summit, LLN1921104-08amb4,w GTN SAR PWTrimm;
Tillamook County, Cascade Head Experimental Forest Road
1861, LLN1941106-18amm16 w GTN. Washington: Clallam
County, Olympic National Forest Sol Duc Road,
LLN1931011-02amb7 STrudell; Olympic National Park Sol
Duc Up Valley, LLN1921031-01amb8 GLBarron; King
County, Asahiel Curtis State Park, LLN1931008-01amb6
SClarke. Pierce County, Mt. Rainier National Park, Lower
Tahoma Creek, 23.IX.1948 AHS31664* (MICH, WTU
DEStuntz04719, both as P. kauffmanii); Snohomish County,
Mt. Baker-Snoqualmie NF Barlow Pass, 07.IX.1992
JFA10697amb11, LLN1920913-06amb9,-08amb10 GRWalker,
LLN1931020-02amb5 w GLBarron, SAR (DAOM, WTU);
Thurston County, Capitol Fall Creek Campground,
LLN1961203-02 RWGaines.
Selected illustrations
Smith (1949, as P. kauffmanii): stereoscopic color trans-
parency. (Possibly also McKenny and Stuntz (1987, as
P. kauffmanii).)
Comments
Phaeocollybia ammiratii can with difficulty be differenti-
ated in the field from P. kauffmanii by its slightly more frag-
ile stature, straight to only slightly inrolled mature pileus
margin, and characteristically vinaceous lower aerial stipe.
The dried pileus cuticle usually has a gold metallic sheen
unique within the complex. The pseudorhizal pellis is the
only portion of the basidiome mildly reactive to
syringaldazine, which distinguishes it from P. redheadii,
P. benzokauffmanii, and P. kauffmanii (where all tissues turn
rapidly deep magenta). Young ochre-colored forms of
P. ammiratii may be confused in the field with
P. luteosquamulosa, otherwise distinguished by a nonviscid
appressed fibrillose pileus and a negative syringaldazine re-
sponse.
Phaeocollybia ammiratii is clearly separated from the rest
of the complex microscopically by the presence of frequent
clamp connections that are particularly noticeable on the
cheilocystidia and narrow suprapellicular hyphae. The dis-
tinctive PalI RFLP pattern (Fig. 10) clearly separates
P. ammiratii from the other 25 tested Phaeocollybia species.
© 2000 NRC Canada
1064 Can. J. Bot. Vol. 78, 2000
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Worldwide, of 24 species characterized by clamp connec-
tions (Norvell 1998a; Rees and Syme 1999), 14 produce
smaller or differently shaped basidiospores, one thick-
walled, tibiiform cheilocystidia, and four very small fruiting
bodies. Phaeocollybia deceptiva Smith and Trappe (1972,
based on a single 1968 Smith collection from Idaho) also
produces clamp connections and large basidiospores, but ex-
amination of the holotype (AHS77000, MICH) reveals that it
lacks the tibiiform diverticula, gelatinized pileipellis, carti-
laginous stipe cortex, and true cheilocystidia characteristic
of Phaeocollybia and more properly belongs in Cortinarius
subgenus Telamonia (J. Ammirati, personal communication;
Norvell 1998a).
Three remaining species—Phaeocollybia amazonica
Singer (1961) from Brazil, Phaeocollybia spoliata Horak
(1974) from India, Phaeocollybia oligoporpa Singer (1987)
from Costa Rica and Colombia—share certain characters
with P. ammiratii. However P. amazonica has distinct
striations over 1/4 of its pileus, a narrower hollow stipe with
an abruptly attenuating pseudorhiza, a nongelatinized cutis,
smaller and less ornamented basidiospores, and much
shorter basidia (Singer 1961). Bandala et al. (1998) exam-
ined the holotype of P. spoliata and noted clamp connections
and its macroscopic similarity to P. kauffmanii. The Indian
species, however, produces subviscid pilei, hollow stipes,
fusoid or subcapitate cheilocystidia, larger basidiospores,
and encrusting pigments in the subpellis.
While P. ammiratii shares overall stature, bitter taste, wa-
termelon-like odor, pseudorhizal morphology, and cap color
with P. oligoporpa (Singer 1987; Horak and Halling 1991;
Bandala 1994; Bandala and Montoya 1994), significant differ-
ences, including the latter’s association with Quercus spp.,
support separation of the two species. Examination of the
Costa Rican holotype (F1052230) reveals relatively rare
clamp connections in the suprapellis, encrusting pigments in
the pileipellis, larger (10 × 6 µm) and less highly ornamented
spores, more variable cheilocystidia, blackish to dark purplish
red dried pilei, and a much darker brown (Sayal Brown) spore
print. (The “highly heteromorphic cheilocystidia,” not as vari-
able as implied in Singer’s (1987) type description, appear to
result from apical regenerations, possibly caused by extended
storage of the basidiomes before drying, cf. Norvell 1998b).
The two Colombian P. oligoporpa collections were found to
differ slightly from the holotype: in Halling 6165 (NY) clamp
connections, found only at basidial bases, were absent from
more uniformly shaped cheilocystidia and suprapellicular
hyphae, while the variable cheilocystidia in Halling 6137
(NY) recall those of the type (occasionally seeming to break
into arthrospores), clamp connections are rare and
suprapellicular hyphae more cohesive.
Four MICH collections cited by Smith (1957) as belong-
ing to P. kauffmanii—Gruber700 (Larch Mountain, Oreg.,
21.X.1945), AHS17545 (Mt. Angeles, Wash., 04.X.1941,
mistakenly cited as AHS17547 by Smith), AHS19467
(Salmon River, Oreg., 06.X.1944), AHS20173 (Rhododen-
dron, Oreg., 24.X.1944)—are now assigned to P. ammiratii.
Phaeocollybia benzokauffmanii sp. nov. Figs. 1, 10–11, 17–21
Pileus 40–145 mm latus, convexo-campanulatus,
umbonatus, viscidus, glaber, avellaneus; carno pallida; odor
farinaceus vel leviter florali, sapor farinaceus vel amarus;
lamellae fumeo-cretaceae, fulvo-brunnescentes e sporis;
stipes 35–100 × 7–23 mm supra terram, usque ad 250 mm
longus cum pseudorhiza gradatim attenuata, ventricosa,
farcta, glaber, versus apicem avellaneus, versus subapicem
rufobrunneosens. Sporae 8.2–10 × 5.6 µm, amygdaliformes,
verruculosae cum rostro glabro usque ad 0.5 µm projecto et
cum apiculo glabro; ixocutis cum suprapelle gelatinosa,
hyalina, et subpelle cum pigmento diffuso; cheilocystidia
anguste vel late claviformia, 3–7 µm lata, gelatinosa,
hyalina. Fibulae desunt in texturis omnibus. Ad terram in
silvis mixtis.
HOLOTYPUS: United States, California, Mendocino County,
Van Damme State Park, November 20, 1992, L.L. Norvell
1921120-01ben1 cum S.A. Redhead et G.L. Barron (WTU;
DAOM, PNW isotypi).
ETYMOLOGY: From benzo- (Ridgway color “Benzo Brown”)
+kauffmanii.
Pileus: 40–145 mm wide, convex to plano-umbonate with
broad umbo, incurved margins, edge strongly inrolled at all
stages of development, glabrous, smooth, opaque, nonstriate,
heavily glutinous, either a mixture of various shades of pink-
ish to purplish brown overall (Benzo/Wood/Army/Natal
Brown, Cinnamon Drab, Avellaneous, Fawn Color [5YR 3-
5/2-5]) or trizonate with dark disc, paler margin, and even
paler drab edge (Dark Vinaceous/Pale Brownish/Light Drab
[5YR 4-5/2-3.5]), older specimens with brown-black spots
and patches; dried pileus matte black overall, occasionally
with dull orange-brown outer margin. Context: firm, conflu-
ent through pileus and stipe and pileus, at disc up to 11-mm
thick, purple-white or pink-white (Light (Cinnamon) Drab,
(Pale) Pinkish, or Cartridge Buff [5YR 6/4, 1.25Y 7-8/1-2]),
when water-soaked darker overall (Drab, Light Vinaceous
Cinnamon) and often with a 1- to 3-mm-wide dark drab
band adjacent to the lamellae. Odor: Slightly floral or
sweetly farinaceous. Taste: Farinaceous or reminiscent of
bitter cucumbers. Lamellae: Free or attached by a small nar-
row tooth, occasionally forking, ventricose with uneven
edges, with lamellulae irregularly interspersed in 5–7 tiers,
broad (up to 15 mm, mean length:width ratio 3.1 [range 1.9–
6]), crowded (L+ll/cm: 17–28 at edge, 7–14 at midpoint),
white to pale pink or ash gray when very young, developing
smoky drab tones as maturing (Vinaceous/Pinkish Buff,
Avellaneous, (Light) Drab [10YR 5-7/2-4]), and dark yellow-
ish gray in age (Tawny Olive, Wood/Buckthorn Brown,
Isabella Color, Saccardo’s Umber [10YR 4-5/4-7]).Veil:Of
-
ten evident as fibrillose patches on aerial stipe. Stipe: Cen-
tral to slightly eccentric, terete or slightly compressed, aerial
length 35–100 mm, combined length with pseudorhiza up to
250 mm, apex 7–23 (30) mm wide, equal or slightly tapering
or ventricose and swelling to 28 (42) mm at ground level,
aerial surface appressed fibrillose (hand lens) and occasion-
ally bearing scattered purplish-brown fibrillose patches, lon-
gitudinally lined, dry; apex pale pink-gray or purple-gray
(Buff Pink, (Light) Cinnamon Drab, Fawn Color, (Light)
Vinaceous Fawn, Vinaceous Buff/Cinnamon [5YR 5-6/3-5])
darkening below to deep purple-brown or red-brown
(Army/Benzo/Bone/Hay’s/Hessian/Liver Brown, Cinnamon
Drab, Fawn [5YR 3-5/2-4, 10R 2. 5-3.5/3-5]); stuffed, 2–
3 mm thick cartilaginous rind occasionally splitting length-
wise to reveal proliferating, firm, pallid context with dark
cartilaginous intrusions; context staining dark brown where
© 2000 NRC Canada
Norvell 1065
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bruised. Pseudorhiza: Vertical-monopodial, generally1/2 to
2/3 total length of basidiome, tapering gradually to a fleshy
blunt or loosely curled origin, deep red-brown overall (Ha-
zel, Hay’s/Liver Brown [10R 2-4/3-5]) or paler (Cinna-
mon/Warm Buff [10YR 6-8/5-6]) at the origin.
Spore print: Pale purplish to dull yellowish brown (Drab
to Buckthorn Brown [10YR 5/6]).
Basidiospores:9×5.5±0.3µm (overall range 8.2–10 ×
5–6 µm), terete or slightly compressed in end view, fusoid-
elliptical in face view, sub-limoniform in profile, verruculose
to verrucose except on 0.5-µm-long beaked apex and distinct,
eccentric apiculus, medium amber in KOH, paler in H2O,
dextrinoid in Melzer’s. Cheilocystidia: Abundant, emanating
from the lamellar trama to form a dense highly gelatinous
layer (unpigmented gill edge visible to the naked eye), thin-
walled, hyaline, lengths variable, narrowly and broadly
clavate elements intermixed, apices generally 3 µm (young) to
5–7 µm (mature) wide above relatively long, 2–3 µm wide
pedicels, often with long filamentous apical outgrowths in ex-
treme age or after long storage. Pleurocystidia: absent.
Basidia: 4-spored, 27–38 × 6–7.5 µm, clavate above long, 2-
µm-wide pedicel, projecting up to 5 µm above developing
hymenium, granular to vacuolate, hyaline. Pileipellis:A
bilaminate ixocutis with a thick (100–300 µm) suprapellis of
radially aligned, sparingly branched, cylindrical, narrow (3–5
(6) µm), highly gelatinized, hyaline hyphae, this layer tightly
compacted in a thick gelatinous matrix and overlying a nar-
rower (50–100 µm) dull orange brown subpellis of slightly
thick-walled, 5–8 µm wide, heavily gelatinized hyphae with
intracellular pigments. Stipitipellis: A 70- to 150-µm-thick
layer of parallel, highly gelatinized, long, sparingly branched,
thick-walled, cylindrical, narrow (2–3 µm), hyaline to dark
brown hyphae with refractive septa. Pseudorhizal pellis: Simi-
lar to stipitipellis, outermost hyphae bearing abundant
tibiiform diverticula. All tramal tissues: Highly gelatinized
and hyaline; in pseudorhiza and stipe strongly sarcodimitic,
with thick-walled (up to 2 µm), wide (up to 30 µm), cylindri-
cal to slightly fusoid, rigid vessel hyphae intermixed with and
supported by less conspicuous, highly branched, thin-walled
(0.5 µm), narrow (2–4 µm), flexuous hyphae; in stipe apex
the flexuous hyphae widening (to 6 µm) and predominating,
walls of the vessel hyphae increasingly thinning; in the pileus
vessel hyphae absent; in the lamellae, parallel, the central re-
gion of 2–3 (5) µm diam., thin-walled hyphae flanked by a
narrow zone of 2- to 30-µm-wide elements giving rise to a
compact subhymenium. Tibiiform diverticula: Abundant on
mycelia and primordial and pseudorhizal pelli, also frequent
on pellicular sheath remnants on stipe apex. Clamp connec-
tions: Absent in all tissues.
MACROCHEMICAL REACTIONS: Syringaldazine: immediately
deep magenta overall; KOH: dull brown. Fluorescence:in-
tense pale yellow young lamellae, purple pileus edge.
RFLP PROFILE (FIGS. 10–11): ITS region of five isolates (in-
cluding type) = 715 bp. Unique in RsaI (630-85); PalI (450-
105-90-70) and PvuII (625-90) also informative. Also cut by
CfoI, EcoRI, HinfI, NdeII; uncut by SalI and XhoI.
ECOLOGY AND DISTRIBUTION: Mycorrhizal. Gregarious, origi-
nating at the albic horizon in podzolic sandy or humic soils
© 2000 NRC Canada
1066 Can. J. Bot. Vol. 78, 2000
Fig. 17. Phaeocollybia benzokauffmanii. Fresh robust, pristine basidiomes (holotype), in situ within a mixed Pinus–Lithocarpus forest
in Van Damme State Park, California, are shown. Their characteristically drab pilei and whitish lamellae are unique within the com-
plex. Scale=5cm.
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in ancient or mature second-growth mesic mixed closed can-
opy coniferous forests in early to mid-autumn. Uncommon
to rare (20 collections verified to date) in coastal and inland
forests of Washington, Oregon, and north-central California.
Specimena selecta!
UNITED STATES: California: Humboldt County, Lord
Ellis Summit, 01.XII.1986 DeShazer370 Holgren (HSC
10443 as P. spadicea); Mendocino County, Jackson State
Forest near Mendocino LLN1921122-03bek3 THorton. Ore-
gon: Clackamas County, Mt. Hood National Forest Road
2609, Wildcat Mountain, LLN1941022-05 SSpencer;
Tillamook County, Cascade Head Experimental Forest Road
1861, LLN1941106-02 w GTN. Washington: Clallam
County, Olympic National Park, La Push, LLN1931015-
10ben5 PKroeger BMcAdoo; Olympic National Park Rugged
© 2000 NRC Canada
Norvell 1067
Figs. 18–21. Phaeocollybia benzokauffmanii. Fig. 18. Habit: Exterior views contrasting robust and slender forms within the same col-
lection (holotype). The appressed fibrillose remnants of the primordial sheath on the stipe (arrow) are typical. Scale bar=5cm.
Fig. 19. Beaked amygdaliform-limoniform basidiospores. The plage (arrow)—an area of lowered ornamentation above the apiculus on
the spore face—can be found to some degree in all species of the complex. Scale bar = 10 µm. Fig. 20. Thin-walled clavate
cheilocystidia. Scale bar = 10 µm. Fig. 21. Basidia. Scale bar = 10 µm.
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Ridge Trail, LLN1921023-05ben4 w SAR GLBarron EFox;
Pierce County, Mt. Rainier National Park Ipsut Creek Trail,
LLN1930924-01ben3 w SAR (DAOM, WTU).
Selected illustrations
Norvell (1998bas P. benzokauffmanii nom. prov.): scan-
ning electron micrographs, black and white photographs of
pseudorhizae with primordium and tibiiform diverticula.
Comments
Mistaken for P. kauffmanii by earlier collectors, the often
more massive basidiomes of P. benzokauffmanii are found in
similar habitats and also have narrow to moderately clavate
cheilocystidia, large limoniform spores, and no clamp con-
nections. The characteristic drab coloration that distin-
guishes P. benzokauffmanii in the field is unique within the
complex and found elsewhere in the region only in
Phaeocollybia gregaria A.H. Smith & Trappe and the small-
spored Phaeocollybia oregonensis A.H. Smith & Trappe. Al-
though microscopically close to P. kauffmanii (also strongly
syringaldazine positive), P. benzokauffmanii produces
slightly longer, broader basidiospores, has slightly broader,
more cohesive hyphae within a thinner suprapellis, and lacks
encrusting pigments in the subpellis. Its ITS region, unlike
that of P. kauffmani, is cut by RsaI.
Of the other nine Phaeocollybia species with similar col-
oration, only P. gregaria is morphologically and anatomi-
cally close. Both are found in Oregon’s Cascade Head
Experimental Forest and produce tightly gregarious
basidiomes with similarly colored glutinous pilei, ash gray
lamellae, similar basidiospores, and no clamp connections.
Phaeocollybia gregaria, however, produces smaller
basidiomes with conic pilei, “thin and cartilaginous pliant”
contexts (Smith and Trappe 1972), a taste described as never
bitter, and very distinctive, long, irregularly narrow
cheilocystidia. The dried P. gregaria type specimens are also
more fragile than P. benzokauffmanii exsiccati, with acute,
brassy metallic pilei and longitudinally splitting stipes.
RFLP profiles differ completely.
Phaeocollybia luteosquamulosa sp. nov. Figs. 1, 10–11, 22–26
Pileus 25–95 mm latus, e conico-convexo campanulatus,
humidulus vel siccus, squamulosus, glaber, fulvus; caro
pallida; odor lenis; sapor frequenter amarus; lamellae
luteolae vel ochraceae, fulvo-brunnescentes e sporis; stipes
30–80 × 8–15mm supra terram, usque ad 400 mm longa
cum pseudorhiza attenuata, ventricosus, subsquamulosus,
farctus, versus apicem lamellis subconcolorus, versus sub-
apicem fulvobrunnescens. Sporae 9–10 × 5–6.2 µm, amyg-
daliformes, verruculosae cum rostro glabro usque ad 1.5 µm
et cum apiculo excentrico glabro; ixocutis cum suprapelle
tenui cum pigmento intraparietali et encrustato, mediopelle
gelatinosa hyalina et subpelle cum pigmento diffuso, intra-
parietali et encrustato; cheilocystidia cylindraceae ad 3 µm
late vel claviformia, subcapitata ad 9 µm lata versus apicem,
gelatinosa, hyalina. Fibulae desunt in texturis omnibus. Ad
terram in silvis mixtis.
HOLOTYPUS: United States, California, Mendocino County,
Jackson State Forest, November 24, 1992, S.A. Redhead
7481lut3 cum G.L. Barron (WTU, isotypus in DAOM).
ETYMOLOGY: From luteo- (yellow) + squamulosa (little
scales).
Pileus: 25–95 mm wide; convex when young, expanding to
conic-umbonate or broadly campanulate with incurved margin
and tightly inrolled edge; surface moist to dry, hygrophanous,
± nonstriate, sparsely silky with short appressed scales on
outer margin and edge, some shade of ochre yellow overall
(Ochraceous Tawny/Buff, Antimony/Light Orange Yellow, Raw
Sienna, [8.75YR 5-8/7-9]), often with darker disc (Russet,
Dresden/Antique Brown [8.75YR 4-5/5-7]), brighter/paler
outer margin (Antimony Yellow, Amber/Buckthorn Brown
[10YR 5-7/6-8]), darker/brighter damaged areas (Russet, Su-
dan Brown [5YR 4-5/6-7]), and (or) slightly darker/brighter
scales (Tawny, Ochraceous Orange [5YR 4.5-6/8-10]); dried
pileus matte, dark orange-brown, often with dull orange mar-
gin. Context: Firm, confluent in pileus and stipe, up to
11 mm thick at the disc, creamy. Odor: Not distinctive to
faintly raphanoid when cut or crushed. Taste: Not distinctive,
or (more often) bitter. Lamellae: Free to almost free, ±
ventricose with slightly uneven edges, lamellulae in many ir-
regular tiers, narrow when young, becoming broad (up to
11 mm, with mean length:width ratio of 3.0 (range 2.3–8.7),
crowded (L+ll/cm: 25–27 at edge, 10–18 at midpoint), yel-
low-cream or orange-cream when young (Cartridge Buff, Pale
Ochraceous Salmon, Light (Ochraceous) Buff [10YR 8-9/2-3,
5YR 7-8/46]), darkening to yellow-brown in age (Cinna-
mon/Ochraceous Buff [1.25Y 6-7/5-6, 7.25YR 6-7/7-8]).Veil:
Evident (i) on unexpanded specimens as isolated fibrils con-
necting pileal margin to stipe and (ii) in mature basidiomes as
scattered, small appressed pileal scales (more conspicuous
when dry) and scattered fibrillose patches on the aerial stipe,
yellow-orange brown (Tawny, Ochraceous Orange [5YR 4.5-
6/8-10]).Stipe: ± central, terete or slightly compressed, aerial
length 30–80 (100) mm, combined length with pseudorhiza
up to 400 mm, apex 8–15 mm, equal or widening up to
20 mm just above ground level; surface matte, finely longitu-
dinally lined, dry, appressed fibrillose (hand lens), apex occa-
sionally bearing scattered orange brown fibrillose patches,
buff apex (Orange/(Light) Ochraceous/Warm/Pinkish Buff
[6.25YR 7-8/6-8, 10YR 7-8/4-6]) grading to pale orange
brown (Orange Cinnamon, Cinnamon Rufous [5YR 4-6/7-9])
at ground level, staining orange (Ochraceous/Zinc Orange
[5YR 5-6/10-12]) where bruised; stuffed, cartilaginous rind
(2–3 mm thick) surrounding firm, cream-colored context.
Pseudorhiza: Vertical-monopodial, generally 2/3 to 5/6 of
overall length, gradually tapering to a loosely coiled or
straight blunt origin, dark red-brown (Kaiser/Carob Brown,
Orange Cinnamon, Xanthine Orange, Russet, Cinnamon
Rufous [5YR 4-5/6-9.5, 2.5YR 3-5/5-8]) above paler
(Ochraceous Buff, Light Ochraceous Salmon [5YR 6-7/7-8])
or darker (Chestnut Brown, Ferruginous [2.5YR 3-4/5-8]) ori-
gin.
Spore print: Pale cinnamon brown ((Light) Pinkish Cin-
namon [7.5YR 6-7/4-6]).
Basidiospores: 10×6±0.5×0.4µm (9–11 × 5–6.8),
terete in end view, broadly amygdaliform to broadly limoni-
form in profile, verruculose, with exosporium irregularly
projecting beyond outline except on 1–1.5 µm long mammi-
form apex and distinct eccentric apiculus, medium amber in
KOH, paler in H2O, inamyloid and nondextrinoid in
Melzer’s. Cheilocystidia: Abundant, emanating from the
© 2000 NRC Canada
1068 Can. J. Bot. Vol. 78, 2000
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lamellar trama to form a distinct gelatinous layer; up to
38 µm long, 3 µm diam. at septa, thin-walled, gelatinized,
hyaline, apices either cylindrical and not appreciably in-
flated or broadly clavate or slightly subcapitate up to 9 µm
diam.; long filamentous apical outgrowths frequently present
in very old or stored fresh specimens. Pleurocystidia:Ab
-
sent or present only as occasional 1- to 2-µm-wide hyphidia
in mature basidiomes. Basidia: 4-spored, 28–43 × 7–9 µm,
projecting up to 10 µm above developing hymenium,
hyaline. Pileipellis: A trilaminate ixocutis with brilliant or-
ange yellow oleiferous hyphae running throughout, remnants
of the pellicular sheath forming a yellow suprapellis of 1–4
radially aligned, 5 µm wide, slightly gelatinized, rarely
diverticulate hyphae encrusted with KOH-soluble, brilliant
yellow-orange pigments and with yellow intraparietal pig-
ments noticeably concentrated at the refractive septa; this
layer overlying a 75–110 µm thick, colorless mediopellis of
unbranched, cylindrical, 2–5 µm wide, highly gelatinized
hyphae and a 100-µm-thick, orange-yellow subpellis of thin-
walled, ventricose and inflated (to 15 µm diam.), gelatinized
hyphae with intraparietal, diffuse, and encrusting pigments.
Stipitipellis: A compact parallel layer of long, cylindrical,
thick-walled, narrow (4 µm wide), highly gelatinized hyphae
with orange-yellow pigments that form tiny crystals in KOH
mountants. Pseudorhizal pellis: Similar to but darker than
stipitipellis, bearing abundant tibiiform diverticula on outer-
most hyphae. All tramal tissues: moderately gelatinized; in
the pseudorhiza strongly sarcodimitic, long (>200 µm), wide
(up to 15 µm), thick-walled (up to 2 µm), rigid, hyaline ves-
sel hyphae intermixed with and held in place by highly
branched, narrow (2–3 µm), thin-walled (approx. 0.25 µm),
flexuous hyphae; in stipe apex both types widening to 30 µm
(vessel) and 6 µm (flexuous), the vessel hyphae becoming
thinner walled (to 1 µm); in the pileus not noticeably
sarcodimitic, hyphae inflated, thin-walled and orange; in the
lamellae parallel, the central region of cylindrical, 3- to 6-
µm-diam., thin-walled, hyaline hyphae flanked by a narrow
zone of 2–3 µm diam., hyaline elements giving rise to a
compact subhymenium. Tibiiform diverticula: Rare to oc-
casional on pileal scales, frequent on stipe fibrillose patches,
and prolific on the pseudorhizal pellis, up to 30 × 2 µm,
aseptate, highly refractive, hyaline, with or without a
(sub)globose head and (or) apical droplet. Clamp connec-
tions: Absent in all tissues.
MACROCHEMICAL REACTIONS: Syringaldazine: no reaction;
KOH: no macroscopical reaction, but frequently producing
0.5-µm-wide crystals in mounts of pellicular tissues. Fluo-
rescence: young lamellae intense whitish yellow, context
dull yellow-orange pileus, stipe context dull purple.
RFLP PROFILE (FIGS. 10–11): ITS region of nine isolates includ-
ing type = 730 bp. Unique in HinfI (400-190-140) and SalI
(535–195). Also cut by CfoI, EcoRI, NdeII, PalI, and PvuII;
uncut by RsaI and XhoI.
ECOLOGY AND DISTRIBUTION: Presumably mycorrhizal. Soli-
tary to clustered in humic podzolic soils at the onset of the
autumnal mushroom fruiting season in inland mature second
© 2000 NRC Canada
Norvell 1069
Fig. 22. Phaeocollybia luteosquamulosa. Freshly excavated basidiomes from Jackson State Forest, California (holotype), are shown.
The nonviscid pileus with appressed fibrillose patches (not seen), bright ochraceous ground colors, and contrasting russet “bruises”
(also on aerial stipe) help identify this species. Scale bar=5cm.
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growth or ancient (virgin) mesic closed-canopy coniferous
rainforests (Tsuga heterophylla (Raf.) Sarg., and Abies spp.)
in Washington and Oregon or mixed coniferous–fagaceous
forests (Tsuga, Abies, Lithocarpus, and Sequoia) in coastal
northern California. Rare (23 collections verified to date).
SPECIMENA SELECTA!: UNITED STATES: California: Del Norte
County, Stout Grove, Jedediah Smith State Park, 05.XII.1937
AHS9426* p.p. (MICH as P. kauffmanii); Mendocino County,
Jackson State Forest Road 409 approx. 2.5 km from 408,
“Aleuria Glen,” LLN1921124-01lut1 REHalling, HDThiers. Or-
egon: Multnomah County, Mt. Hood National Forest, Larch
Mountain summit, LLN1921104-09luu7 wGTN SAR PWTrimm
(PNW), LLN1931024-06lut4,-10alut2 wGTN, LLN1931105-
09lut6 wSAR, LLN1941006-11lut7, LLN1941011-18lut9. Wash-
ington: Lewis County, Rainbow Falls State Park near PeEll,
04.XI.1990 NLaycock s.n. Snohomish County, Mt. Baker –
Snoqualmie National Forest, Barlow Pass, 07.IX.1992
JFA10700lut8, SATrudell 250-92-04lut10.
© 2000 NRC Canada
1070 Can. J. Bot. Vol. 78, 2000
Figs. 23–26. Phaeocollybia luteosquamulosa. Fig. 23. Habit: exterior view of mature basidiomes and longitudinal section of immature
specimen. The appressed fibrillose patches on the pileus and stipe (arrows) are remnants of the ruptured primordial gelatinous pellicle.
Scale bar = 5 cm. Fig. 24. Beaked amygdaliform basidiospores. Scale bar = 10 µm. Fig. 25. Basidium. Scale bar = 10 µm. Fig. 26.
Broadly clavate cheilocystidia. The apical extension (arrow) on one is an artifact of extended storage. Scale bar = 10 µm.
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Selected descriptions and illustrations
Norvell (1998bas P. luteosquamulosa nom. prov.): black
and white photographs of pileal scales, pellicular veil,
cheilocystidia, and basidiospores.
Comments
Phaeocollybia luteosquamulosa is recognized in the field
in dry conditions by its squamulose, dry to merely lubricous,
yellow-ochre pileus, and nondistinctive odor. Other distin-
guishing characters are its negative syringaldazine reaction
and trilaminate pileipellis with the patchy, narrow, brilliant
yellow suprapellis. The P. luteosquamulosa ITS region pro-
duces unique HinfI, PalI, and SalI banding patterns within
the complex.
No other known Phaeocollybia shares the robustness, pig-
mentation, cuticular structure, and spore morphology of
P. luteosquamulosa. Of the 14 species described as having
pileal scales, only two—Phaeocollybia coniuncta Horak
(1980) and Phaeocollybia caudata Horak and Halling
(1991)—have dry to moist pilei similar in color. The Indian
P. coniuncta is described as a small species with dirty yel-
lowish-brown pileus, slender, hollow stipe, cylindrical
cheilocystidia, and normally cespitose habit. The Columbian
and Costa Rican P. caudata is smaller and has a hollow
stipe, thread-like pseudorhiza, small ellipsoid spores, and
thick-walled tibiiform cheilocystidia.
Two collections—Ammirati 6018 (MICH) from Del Norte
County, Calif., Norvell A1991202O1-02 (PNW) from Benton
County, Oreg.—are very closely affiliated with P. luteosqua-
mulosa. They share with it a similar stature, moist to dry
fibrillose pileus, trilaminate pileipellis with a very thin bril-
liant yellow pigmented suprapellis, and similarly shaped
clavate cheilocystidia. However the Chestnut Brown pilei and
considerably smaller basidiospores (8 × 4.5, Fig. 1) are atypi-
cal. These collections are excluded from P. luteosquamulosa
pending further investigation.
Phaeocollybia redheadii sp. nov. Figs. 1, 4, 10–11, 27–31
Pileus 45–140 mm latus, campanulatus, umbonatus,
viscidus, glaber, fulvus; caro pallida; odor farinaceus; sapor
farinaceus et amarus; lamellae pallide roseocremeae, fulvo-
brunnescentes e sporis; stipes 40–100 × 10–23 mm supra
terram, usque ad 300 mm longa cum pseudorhiza attenuata,
ventricosus, glaber, farctus, versus apicem lamellis subcon-
colorus, versus subapicem fulvobrunnescens. Sporae 9.8–11
(12) × 5.5–6.5 µm, amygdaliformes, verrucosae cum rostro
mammiformi glabris usque ad 1 µm et cum apiculo
excentrico glabro; ixocutis cum suprapelle crassa gelatinosa
hyalina et subpelle cum pigmento diffuso et intraparietali et
encrustato; cheilocystidia anguste vel late claviformia,
subcapitata et pedicellata, 6–10 (15) µm lati versus apicem,
gelatinosa, hyalina. Hyphae fibulatae in stipitipelle, raro. Ad
terram in silvis Pinacearum mixtarum.
HOLOTYPUS: Canada, British Columbia, Vancouver Island,
Upper Carmanah Valley, October 24, 1991, L.L. Norvell
1911024-02rec1 cum S.A. Redhead (WTU, isotypus DAOM).
ETYMOLOGY: In honor of Dr. Scott Redhead, who first
noted a large-spored “variant” of P. kauffmanii.
Pileus: (23) 45–140 mm wide, conic-convex when young
with decurved margin, expanding to broadly campanulate
with broad low umbo, decurved margin, extreme edge
inrolled at all times; surface glabrous, smooth, viscid to
heavily glutinous, nonstriate, pristine basidiomes either
warm orange-brown overall (Verona/Sayal Brown, Warm
Sepia [5YR 3-5/3-5, 7.5YR 4-5/5]) or zonate with deep or-
ange brown to dark blackish brown disc (Ve-
rona/Bone/Snuff/Clove Brown, Warm Sepia [5YR 3-5/3-5,
7.5YR 4/2-4]), dark yellowish brown or brownish orange
margin (Sayal/Snuff/Cinnamon/Verona/Mikado Brown, Rus-
set [7.5YR 4-5/46, 5YR 3-5/4-8]), and dark brown edge
(Warm Sepia, Verona/Mikado/Snuff Brown [5YR 3-5/3-6]),
older basidiomes becoming blackish or purplish brown over-
all, moldy or frost-exposed basidiomes frequently develop-
ing dull purplish or greenish brown tones (Benzo/Army
Brown, Saccardo’s Umbo, Isabella Color [5YR 4/2-4, 2.5YR
3-5/3-5]), damaged areas generally blackish, dried pileus
with blackened umbo and metallic, dark bronze margin, or
shiny, deep mahogany brown overall. Context: Firm, con-
fluent in pileus and stipe, up to 12 (15) mm thick at the disc,
when young, pale yellow cream or pinkish white (Car-
tridge/(Pale) Pinkish Buff [2.5Y 7-8/1-4]), when old and (or)
watersoaked dark yellow brown (Snuff/Sayal Brown [7.5YR
4-5/4-5]), staining orange or deep orange-brown when cut
(Xanthine Orange, Verona Brown [5YR 4-5/4-5(9)]), when
wet often developing a 1–3 mm wide, darkened zone adja-
cent to the lamellae. Odor: Strongly farinaceous, often
slightly floral when cut or crushed. Taste: Strongly farina-
ceous (cucumber-like), glutinous pileus cuticle slightly to
extremely bitter. Lamellae: Free or attached by a small
tooth, ventricose with broad central portion, occasionally
forked, lamellulae in (3) 5–7 irregular tiers and distributed in
wedge-shaped patterns, broad in mature specimens (up to
15 mm, with mean length : width ratio 4.6 (range 3.2–8.3,
14 when very young)), crowded (L+ll/cm: 15–24 for edge,
8–14 for midpoint), pinkish white or yellow cream when
very young (Pale Pinkish/Cartridge/Ochraceous/Tilleul Buff
[2.5Y 8/2, 7.5YR 8/1-2, 10YR 9/2]), maturing to brownish
gray-yellow or yellow-brown (Pinkish/Cinnamon Buff, Cin-
namon [10YR 6-7/4-5, 7.5YR 4-7/4-7]), and dark yellow-
brown to deep brown-yellow in age (Sayal/Snuff Brown,
Tawny Olive, Tawny Olive [7.5YR 3-5/4-5, 1.25Y 4-5/4-5]).
Veil: Occasionally evident as fibrillose patches on aerial
stipe. Stipe: Central to slightly eccentric, terete to slightly
compressed, aerial length 40–100 mm, combined length in-
cluding pseudorhiza up to 340 mm, apex wide (10–23 mm
diam.), occasionally equal or tapering downward, usually
ventricose and widening to 12–30 mm just above ground
level, smooth or appressed fibrillose (hand lens), longitudi-
nally lined, moist, apex pale to medium pinkish buff in
young specimens ((Light) Pinkish/Light Vinaceous/Orange
Cinnamon [7.5YR 6-8/4-6, 5YR 5-8/4-6]), grayish orange- or
yellow-brown in old or damaged specimens ((Vinaceous)
Cinnamon, Cinnamon Drab, Avellaneous [5YR 5-6/3-6,
7.5YR 5-6/4-7]), darkening below to deep orange-brown
(Ferruginous, Kaiser/Mikado/Verona Brown, Pale Orange
Cinnamon, Warm Sepia [2.5YR 4/4-8, 5YR 3-6/3-5]), stain-
ing deep red-brown; stuffed, 2–4 mm thick cartilaginous
cortex surrounding firm context, context frequently prolifer-
ating so abundantly as to split cortex lengthwise, cream or
pinkish white in young specimens (Cartridge/Pale Pinkish
Buff [2.5Y 8/0-2]), light to dark yellowish brown in age
((Pale Pinkish) Cinnamon [7.5YR 5/7, 8-9/2-4]).
© 2000 NRC Canada
Norvell 1071
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Pseudorhiza: Vertical-monopodial, generally 2/3 to 5/6 of
overall stipe length, gradually tapering to a blunt fleshy ori-
gin, overall deep red- to rusty-brown overall (Liver Brown,
Bay [7 5YR 2-3/4-5]; Kaiser/Pecan Brown [2.5YR 3-4/5-6])
or with pink-orange (Salmon Color) origin, origin arising
from a nodulus surrounding buried rootlets.
Spore print: Cinnamon brown (Cinnamon [7. 5YR 5/6]).
Basidiospores: 10.5×6±0.4×0.3µm (8.5–12 × 5–
7µm), terete to almost angular in end view, elongated inequi-
laterally amygdaliform in profile view, elongate-elliptical in
face view, verrucose everwhere except on the 1-µm-long
mammiform beaked apex and eccentric prominent apiculus,
suprahilar plage an area barely visible in oil immersion but
easily visible in scanning electron micrographs (Fig. 4); or-
ange to medium amber in KOH, paler in H2O, dextrinoid in
Melzer’s. Cheilocystidia: Abundant, emanating from the
lamellar trama to form a dense, highly gelatinous layer,
lengths up to 35 µm, broadly clavate elements intermixed
with subcapitate elements with 6–10 (15) µm wide globose
heads atop (2) 3–4 µm wide pedicels of variable length,
sometimes catenulate with swelling also noted near apices of
the penultimate elements, thin-walled, highly gelatinized,
hyaline, often developing long filamentous apical outgrowths
in very old or stored fresh specimens. Pleurocystidia:Ab
-
sent. Basidia: (2-) 4-spored, 29–50 × (6) 7–8 (10) µm, mod-
erately to narrowly clavate with pedicel tapering to the 1.5–3
µm-wide basal septum, hyaline. Pileipellis: a bilaminate
ixocutis with a 200–600 µm thick suprapellis of branching,
cylindrical, narrow (2–6 µm wide), thin-walled, hyaline
hyphae, this layer loosely embedded in a clear gelatinous ma-
trix and overlying a 75–200 µm thick, yellow orange to dull
orange-brown subpellis of unbranched, thick-walled, 5–8 µm
wide, inflated (to 14 µm) hyphae with refractive septa and en-
crusting, intraparietal and diffuse pigments; oleiferous hyphae
present throughout pileipellis. Stipitipellis: A compact layer
of parallel hyphae gradually intergrading with the stipititrama,
of 4–9 µm wide hyphae with thick, hyaline to orange-brown
walls, rarely with true or pseudo-clamp connections.
Pseudorhizal pellis: Up to 200 µm thick, hyphae 20 µm wide,
with coarse, dark-brown encrustations, often with coagulated
brownish contents; outermost hyphae bearing tibiiform
diverticula, clamp and pseudo-clamp connections rare. All
tramal tissues: Highly gelatinized, colorless to pale amber,
and containing occasional dark amber oleifers; in the
pseudorhiza strongly sarcodimitic, with parallel, long
(>200 µm), wide (up to 40 µm), thick-walled (up to 4 µm),
rigid vessel hyphae supported by inconspicuous, branched,
thin-walled (0.5 µm), narrow (2–3 µm), flexuous hyphae; in
the stipe both hyphal types with thick (1–2 µm) walls, vessel
hyphae narrowing to 20 µm; in the pileus all hyphae in a
loose radial arrangement, 15–20 µm wide, with 0.5–1 µm
thick walls; in the lamellae parallel, with the 200–400 µm
central region of relatively thick-walled (approx. 1.0 µm), 3–
10 µm wide hyphae occasionally intermixed with highly
branched, narrow, thin-walled flexuous hyphae flanked by a
compact zone of 2–3 µm wide elements giving rise to a com-
pact subhymenium. Tibiiform diverticula: Abundant on my-
celia and primordial/pseudorhizal pelli and frequent on
pellicular sheath remnants on stipe apex, up to 25 × 1
(1.5) µm, hyaline to pale straw, highly refractive, aseptate,
with or without a (sub)globose head and (or) apical droplet.
Clamp connections: Sporadic and rare in stipitipellis, absent
in all other tissues.
MACROCHEMICAL REACTIONS: Syringaldazine: lamellae and
pseudorhizal pellis immediately deep magenta; KOH: posi-
tive (immediately black- or green-brown). Fluorescence:
lamellae: Violet-white (young), intense pale or mottled yel-
low (maturing), brilliant intense yellow under spores (fully
mature). Context: Dull orange (primordia), purple-orange
(pileus), blue (stipe apex).
RFLP PROFILE (FIGS 10–11): ITS region of five isolates includ-
ing type = 730 bp. PalI (220-220-110-90-90) and PvuII
(640-90) were the most informative enzymes. Also cut by
CfoI, EcoRI, HinfI, NdeII; uncut by RsaI, SalI, and XhoI.
© 2000 NRC Canada
1072 Can. J. Bot. Vol. 78, 2000
Fig. 27. Phaeocollybia redheadii. Freshly excavated young
basidiomes from Snohomish County, Washington, are shown.
The red-brown tones of the pileus help differentiate this species
from the characteristically brighter, orange to orange-brown
P. kauffmanii. Scale bar=5cm.
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ECOLOGY AND DISTRIBUTION: Mycorrhizal. Solitary to scat-
tered or gregarious during early to late autumn, originating
at the albic horizon in humic podzolic soils in coastal and
inland ancient or mature second-growth mesic mixed closed
canopy Tsuga-Abies forests. Fairly common (109 collections
verified to date) from southern British Columbia to northern
California.
Specimena selecta!
CANADA: British Columbia: Vancouver Island, Lower
Carmanah, LLN1921009-20 wSAR EFox (DAOM, WTU).
UNITED STATES: California: Del Norte County, Jedediah
Smith State Park, 05.XII.1937 AHS9426 p.p. (MICH as
P. kauffmanii). Oregon: Benton County, Mary’s Peak,
29.IX.1966 JMTrappe934 (BPI, NY, OSC 20072, SFSU all as
P. kauffmanii); Clackamas County, Mt. Hood National Forest
Road 2609, Wildcat Mountain, LLN1941028-07rec6 wJR
(PNW); Lincoln County, Cascade Head Experimental Forest,
LLN1941106-04rec5 wGTN; Multnomah County, Mt. Hood
National Park Larch Mountain, LLN1931024–03rek5 wGTN,
LLN1931105-01rec7 wSAR. Washington: Clallam County,
Olympic National Park Rugged Ridge, LLN1921023-34; Jef-
© 2000 NRC Canada
Norvell 1073
Figs. 28–31. Phaeocollybia redheadii. Fig. 28. Habit: exterior view and longitudinal sections. Scale bar=5cm.Fig. 29. Large, rotund
limoniform beaked basidiospores (note plage areas). Scale bar = 10 µm. Fig. 30. Pedicellate and catenulate cheilocystidia with charac-
teristically swollen subglobose apices. Scale bar = 10 µm. Fig. 31. Basidium (sterigmata not fully extended). Scale bar = 10 µm.
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© 2000 NRC Canada
1074 Can. J. Bot. Vol. 78, 2000
ferson County, Olympic National Park Twin Creek Research
National Area, LLN1921017-21rek4 wGTN OLNorvell
PWTrimm; Snohomish County, Mt. Baker – Snoqualmie Na-
tional Forest, Barlow Pass, LLN1920921-04rec2,-07reo2.
Selected descriptions and illustrations
Norvell (1992 as P. kauffmanii): Black and white photo-
graph. Redhead and Norvell (1993, holotype as
P. kauffmanii): Black and white photographs and line draw-
ings; Norvell (1998bas P. redheadii nom. prov.): Black and
white photographs of excavation and mycorrhizal evidence.
Comments
The authors who first reported P. redheadii from Canada
as P. kauffmanii (Norvell 1992; Norvell and Redhead 1992;
Redhead and Norvell 1993) suggested that this large-spored
variant might be a new species. Subsequent comparisons of
microscopic features and molecular data support dividing the
orange-brown members of the P. kauffmanii complex into
three taxa—the obviously clamped P. ammiratii and the es-
sentially unclamped P. kauffmanii sensu stricto and
P. redheadii. These species are easily confused. The gener-
ally less robust stature and vinaceous stipes of P. ammiratii
help to separate this species in the field. The difference be-
tween the distinctly orange young pileus of P. kauffmanii
and the darker apricot brown young P. redheadii is a subtle
one, however, making the two difficult to separate macro-
scopically.
Microscopically, P. redheadii is easily separated from
P. kauffmanii by its larger basidiospores and distinctive pedi-
cellate (sub)capitate cheilocystidia (cylindrical to narrowly
clavate in P. kauffmanii, cf. Figs. 9 and 30). Phaeocollybia
redheadii basidiospores, the largest in the complex, do occa-
sionally intergrade with those of P. luteosquamulosa and
P. benzokauffmanii (Fig. 1). This overlap may complicate dif-
ferentiating P. redheadii from P. benzokauffmanii (which oc-
casionally also produces subcapitate inflated cheilocystidia)
when P. redheadii pilei have developed a purplish-brown cast
from cold or fungal pathogens; here encrusting pigments in
the pileipellis identify P. redheadii. Within the complex,
P. redheadii has the only ITS region cut by PvuII and uncut
by RsaI, SalI, and XhoI.
Of the six extralimital Phaeocollybia species with basidio-
spores averaging approx. 10 × 6 µm, three have clamp con-
nections and the fourth has tibiiform thick-walled
cheilocystidia. Of the two remaining taxa, P. muscicolor
Horak (1977), from New Guinea, produces small olive
basidiomes with hollow stipes and even larger basidiospores.
Phaeocollybia quercetorum Singer (1987) from Costa Rica
is similar in color to P. redheadii, but is generally smaller in
stature, has a hollow stipe, smells of phenol, and is associ-
ated with Quercus-dominated forests.
Three of the four paratype collections in Smith’s (1937)
original description of Naucoria kauffmani [sic]—Kauffman
X.11.1925 (Lake Quinault, Wash.), AHS2495 (Cape Flattery,
Wash., 18.IX.1935), AHS3888 (Eureka, Calif., 9.XI.1935)—
and three collections cited under P. kauffmanii in Smith’s
1957 monograph—AHS3727 p.p. (Orick, Calif.,
03.XII.1935), AHS49479 (Larch Mountain, Oreg.,
25.X.1954), AHS56489 (Prairie Creek, Redwood State Park,
Calif. 09.XII.1954)—are now reassigned to P. redheadii.
The Phaeocollybia kauffmanii complex
Technical description (Table 1, Figs. 1–4 and 10–11)
Basidiomes are generally large and robust (mature pilei >
40 mm diam. and stipe apices > 10 mm diam.); stipes have a
conspicuous dense pith and are never hollow; pseudorhizae
are vertical-monopodial; development is pileostipiticarpic
monovelangiocarpic; basidiospores are large (>8 × 5 µm),
amygdaliform or limoniform with moderately coarse orna-
mentation; cheilocystidia are indeterminate, thin-walled,
clavate, producing long, narrow, apical extensions during se-
nescence or storage.
Key to species
1a. Pilei dry to moist (never glutinous), appressed-scaly, ochraceous yellow when mature; pileipellis trilaminate with
narrow bright orange-yellow suprapellis (retained pellicular sheath), relatively narrow (75–100 µm thick) colorless
gelatinized mediopellis, and duller orange yellow subpellis; all tissues negative in syringaldazine...............
................................................................P. luteosquamulosa
1b. Pilei glabrous (never scaly), viscid to glutinous, not ochraceous yellow when mature; pileipellis bilaminate with
highly gelatinized hyaline suprapellis and pigmented subpellis; at least some tissues immediately or tardily magenta
in syringaldazine................................................................ 2
2a. Clamp connections present throughout, notably conspicuous on the suprapellicular hyphae and cheilocystidia and
inconspicuous (obscured) at the basidial bases; pseudorhizal pellis weakly magenta after 30 min in syringaldazine . .
.................................................................... P. ammiratii
2b. Clamp connections absent in suprapellicular hyphae, cheilocystidia, and basidial bases but sometimes present in
stipitipellis; all tissues immediately magenta in syringaldazine..................................3
3a. Pilei and stipe apices dull pinkish to purplish-brown (young pilei glabrous, viscid, drab; stipe apex pink to drab);
clamp connections absent; subpellis with dull brown diffuse pigments.....................P. benzokauffmanii
3b. Pilei orange, pale tawny, brownish orange to orangish brown, lacking drab to pinkish-brown colors; clamp con-
nections occasionally present in stipitipellis; subpellis with brownish-orange encrusting pigments..............4
4a. Basidiospores generally approx. 10.5 × 6 µm (overall range 8.5–12 × 5–7 µm); cheilocystidia generally
pedicellate and capitate to subcapitate (subglobose heads frequent); young pilei pale tawny, becoming red-brown to
medium dark brown in age..................................................P. redheadii
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4b. Basidiospores generally approx. 8.5 × 5 µm (overall range 7.5–10 × 4–6 µm); cheilocystidia irregularly cylindri-
cal to narrowly clavate, rarely pedicellate or subcapitate; young pilei bright to dull orange (apricot brown), becoming
tawny or orange-brown in age...............................................P. kauffmanii
Comments
The complex is generally associated with ectomycorrhizal
conifers (Tsuga heterophylla,Abies spp., Picea sitchensis
(Gon.) Carr., Pseudotsuga menziesiii (Mirbel) Franco, and
Pinus contorta Dougl.), although habitats become more
complex in California with the introduction of Lithocarpus
densiflorus (Hook. and Arn.) Rehder and the nonecto-
mycorrhizal Sequoia sempervirens (D. Don) Endl. Represen-
tatives are frequently found in the heart of a closed-canopy
mesic rainforest, either in ancient preserves (e.g., British Co-
lumbia’s Carmanah Valley, Washington’s Barlow Pass and
Olympic National Park, California’s Jedediah Smith State
Park) or in fully mature second growth (e.g., Oregon’s Mt.
Hood National Forest and Cascade Head Experimental For-
est, California’s Big Lagoon State Park and Jackson State
Forest). There, ericaceous (Vaccinium or Rhododendron)
species and ferns (primarily Polystichum munitum (Kaulf.)
Presl.) frequently inhabit the understory, and competing her-
baceous ground cover is generally sparse, with sandy or
loamy substrates topped by heavy needle litter, or tightly
packed or scattered mosses. The dominant soil is an acidic
spodozol, generally with large amounts of woody debris and
roots intermixed with rock and sand. Both P. ammiratii and
P. kauffmanii have been collected from more open Pseudo-
tsuga menziesi – Lithocarpus densiflorus forested areas in
more inland montane regions of northern California.
There are only subtle distributional differences. The two
most common and widely distributed species—P. kauffmanii
and P. redheadii—and the less common P. ammiratii have
been collected in British Columbia, Washington, Oregon,
and northern California. The rare P. benzokauffmanii and
P. luteosquamulosa have not yet been collected from British
Columbia. Only P. kauffmanii has been collected in Idaho.
Acknowledgments
I thank J.F. Ammirati for guidance, financial support, and
laboratory facilities, S.A. Redhead for field assistance and
numerous illuminating discussions, and both for critical
manuscript reviews. I also thank J.E. Trappe for assistance
with the Latin diagnoses, T.J. Baroni, R. Currah, and R.E.
Halling for helpful reviews, and Agriculture & Agri-Food
Canada for permission to publish Dr. Redhead’s photographs
of P. kauffmanii, P. benzokauffmanii, and P. luteosquam-
ulosa. Specimens and (or) photographs were donated or
loaned by J.F. Ammirati (WTU), C. Ardrey, G.L. Barron, W.
Denisen (OSC), D.E. Desjardin (SFSU), R.L. Exeter, R.
Fogel and R. Shaffer (MICH), R.W. Gaines, S. Gamiet, C.
Goetz, R.E. Halling (NY), P. Kroeger, D.L. Largent (HSC),
B.E. Lippert, C. Leuthy, O.K. Miller (VPI), G. Mueller (F),
M. Peeters, R.H. Petersen (TENN), S.A. Redhead (DAOM),
J. Roger, M.M. Rogers, A.E. Rossman (BPI), C. Scates-
Barnhart, S. Spencer, W. Steglich, W.J. Sundberg (SIU), I.
Tavares (UC), and H.D. Thiers (SFSU). Guidance during
scanning electron microscopy or molecular work was given
by L. Comai, A. Denton, Y. Liu, G. Mueller, K. Nakasone,
B. Reine, S.O. Rogers, M.T. Seidl, J. Stiller, and G.
Takimoto. Field support was provided by the Lake
Cowichan Field Station (Lake Cowichan, B.C.), Olympic
National Resource Center (Forks, Wash.), and Cascade Head
Experimental Forest Research Station (Otis, Oreg.). Finan-
cial assistance was provided by the D.E. Stuntz Memorial
Foundation, the Mycological Society of America, and G.T.
Norvell.
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