![Phaeocollybia chefensis paratypes. [Lincoln Co.: lln1951109-16 (A-C), rle2007-074 (F), rle2015-21 (H), rle2021-1 (J); Lane Co.: rle2007-129 (D, E); Polk Co.: hc2018-LB4 (I); Benton Co.: lln2091026cc6 (G)]. A-C. Three views of the first specimen from CHEF that fostered the concept for P. tibiikauffmanii in 1995. D. Young robust specimens excavated from Lobster Creek. E. Pristine pseudorhizal tissues display strong magenta reaction in syringaldazine after 15 minutes. F. Second specimen retrieved from CHEF in 2007. G. Solitary specimen retrieved from Conner's Camp on Marys Peak. H. Basidiome at the original CHEF site prior to excavation in 2015. I. The first primordium excavated with its more mature partner in 2018 at Boulder Creek. J. Exeter's 2021 negative syringaldazine reaction from this waterlogged solitaire cautions that adverse environmental conditions do disable this usually reliable diagnostic tool.](https://www.researchgate.net/profile/Lorelei-Norvell/publication/360245442/figure/fig4/AS:1150040110317569@1651202337286/Phaeocollybia-chefensis-paratypes-Lincoln-Co-lln1951109-16-A-C-rle2007-074-F_Q320.jpg)
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MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2022
January–March 2022—Volume 137, pp. 1–30
https://doi.org/10.5248/137.1
Phaeocollybia chefensis sp. nov. and new synonyms for
P. riipes, P. rufotubulina, and P. tibiikaumanii
L L. N*, R L. E, M G,
S-T M, S A. R
1 Pacic Northwest Mycology Service, 6720 NW Skyline, Portland OR 97229 USA
2 2477 SW Maplewood Drive, Dallas OR 97338 USA
3 Molecular Solutions, 4216 N. Castle Ave., Portland OR 97217 USA
4 National Mycological Herbarium (DAOM), Agriculture & Agri-Food Canada,
960 Carling Avenue, Ottawa ON K1A 0C6 Canada
*C : llnorvell@pnw-ms.com
A—Phylogenetic analyses of ITS and RBP2 sequence data from Phaeocollybia
collections made at Cascade Head Experimental Forest in Oregon support recognition
of a new species, P. chefensis. Collections of the new species were previously referred to
P. tibiikaumanii. Sequence analyses also establish that P. tibiikaumanii is a synonym
of P. spadicea, P. riipes is a synonym of P. lilacifolia, and P. rufotubulina is a synonym of
P. californica. A revised general key to Pacic Northwest Phaeocollybia species is provided.
K —Basidiomycota, Hymenogastraceae, nomenclature, Northwest Forest Plan,
taxonomy
Introduction
Phaeocollybia R. Heim (Agaricomycetes, Hymenogastraceae) is a genus of
brown-spored agarics characterized by conic-campanulate pilei, cartilaginous
stipes and pseudorhizae, ornamented ‘beaked’ basidiospores, sarcodimitic
tissue, monovelangiocarpy, and tibiiform diverticula that arise from the
mycelium and primordial pellicular remnants (Norvell 1998a,b; Norvell
& Exeter 2009). e forests of western North America have provided an
exceptionally large number of species in this genus (Murrill 1911; Smith 1937,
1957a,b; Smith & Trappe 1972; Redhead & Norvell 1993; Norvell 1998a,b,
2 ... Norvell & al.
2000, 2002, 2004; Norvell & Redhead 2000, Norvell & Exeter 2007, 2009;
Norvell & al. 2010).
In 1995 the rst author collected a solitary tawny colored Phaeocollybia
from Oregon’s Cascade Head Experimental Forest (CHEF). Although the
specimen’s general morphology resembled that of P. kaumanii (A.H. Sm.)
Singer, it diered by possessing abundant refractive tibiiform cheilocystidia,
prompting application of the provisional name P. tibiikaumanii to the
Oregon coast taxon in Norvell’s 1998 doctoral dissertation.
e author’s research on Phaeocollybia coincided with establishment
of the Northwest Forest Plan (NWFP; USDA-USDI 1994), which listed 14
phaeocollybias among its 234 fungal species of concern (Castellano & al.
1999, 2003; ORBIC 2021). In 1998 transects were established for surveying
epigeous ectomycorrhizal basidiomycetes in Benton County’s Bureau of
Land Management (BLM) Green Peak Density Management Study and
Polk County’s BLM Fungal Chronosequence Study (Norvell & Exeter 2004).
Numerous gregarious to caespitose clusters of P. kaumanii-like specimens
with tibiiform cheilocystidia were collected that matched the solitary 1995
specimen from Lincoln County. Aer morphological comparison with
other specimens from Washington, Oregon, and California, a particularly
well-documented collection from the chronosequence ‘old growth’ study
transect (approximately 64 km from the CHEF site) was selected as type for
P. tibiikaumanii (Norvell 2004).
In 2008, the U.S. Forest Service (FS) & BLM Interagency Special Status/
Sensitive Species Program initiated a project to generate DNA sequence data
for Phaeocollybia specimens (Gordon 2009). Since then, 350 sequences for
the internal transcribed spacer region (ITS1-5.8S rRNA-ITS2) in the nuclear
ribosomal RNA gene cassette and 46 sequences of a portion of the gene
encoding the second largest subunit in the DNA-directed RNA polymerase
(RPB2) were deposited in GenBank. ese data, along with NWFP surveys,
have provided several hundred Phaeocollybia collections for taxonomic and
genetic analysis, and serve as an excellent resource for phylogenetic analyses
and specic primer design.
Initial molecular analyses that clustered P. tibiikaumanii and P. spadicea
sequences within one intermixed clade were received on the same day as
the arrival of the galley proofs for P P N
N A (Norvell & Exeter 2009). Although the vital data arrived
too late for a thorough publication revision, the authors annotated their
keys and commentaries in anticipation of subsequent taxonomic changes.
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 3
Additional ITS sequence analyses conrmed the synonymy of P. spadicea and
P. tibiikaumanii but also supported newly collected material from the original
Lincoln County ‘tibiikaumanii’ site as a distinct species, independent of
P. spadicea.
A revised ITS-based phylogeny of Phaeocollybia was presented in the
Norvell & al. (2010) poster at the International Mycological Congress in
Edinburgh. In this paper we provide additional ITS and RPB2 sequence data
that establish P. chefensis as a new species and formally publish synonymies
for P. californica (= P. rufotubulina), P. lilacifolia (= P. r i ip es ), and P. spadicea
(= P. tibiikaumanii).
Materials & methods
Collections
Specimen collection and examination, ultraviolet inspection, and syringaldazine
spot tests of fresh material followed procedures outlined in Norvell & Exeter (2009),
where morphotaxonomic terms (e.g., tibiiform diverticula, sarcodimitic tissues,
pellicular veil, vertical-monopodial, and rhizomorphic pseudorhizae) are also dened.
Parenthesized color references from Ridgway (1912: e.g., “Pale Pinkish Cinnamon”)
accompany colors of fresh specimens described in general non-standardized color
names in lower case. Microscopical examinations were made of fresh tissues in H
2
O
or (for the type material) dried tissues rehydrated in 6% aqueous KOH. Basidiospores
were taken from the stipe apex. Dimensions of all anatomical cells follow the format
n (number measured): (outlier)low–average–high(outlier) [e.g., basidia n = 19:
25–34.2–41(43) × 6.5–8.1–9.4(9.9) µm].
Collector abbreviations include (Norvell), (Exeter), and (Redhead).
Latitude/longitude geographic coordinates were converted and rounded to four
decimal places from surveyor input using [for TRS] https://www.earthpoint.us/
TownshipsSearchByDescription.aspx and [for UTM] https://www.ngs.noaa.gov/
NCAT/.
Vegetation abbreviations follow those used during Northwest Forest Plan surveys
and include (in all caps for overstory) ABPR (Abies procera Rehder), PISI (Picea
sitchensis (Bong.) Carr.), PSME (Pseudotsuga menziesii (Mirbel) Franco), and TSHE
(Tsuga heterophylla (Raf.) Sarg.) and (in small caps for understory) (Berberis
nervosa Pursh = Mahonia nervosa (Pursh) Nutt., the preferred synonym),
(Gaultheria shallon Pursh), (Oxalis oregana Nutt.), (Polystichum munitum
(Kaulf.) C. Presl), (Vaccinium ovatum Pursh), and (Vaccinium parviorum
Andrews = Gaylussacia baccata (Wangenh.) K. Koch, the preferred synonym).
Material cited in the S sections is housed (unless otherwise
specied) in Norvell’s Pacic Northwest Mycology Service fungarium (PNW) and
will be distributed among DAOM, NY, OSC, PNW, and WTU aer publication.
Herbarium acronyms follow iers (2021).
4 ... Norvell & al.
T 1: PNW Phaeocollybia ITS & RPB2 sequence data. (Type data in bold).
S C./F.
ITS RGBC, S/P;
P
ammiratii 1941028-10-T JN102495 — Clackamas, OR; Norvell
2051018-01 GQ165629 — Skamania, WA; Norvell
OSC 155802 KJ450913 — Coos, OR; Rodenkirk
2007-103 (GQ165627) KU574760 Benton, OR; Exeter
2007-105 GQ165624 KU574759 Polk, OR; Exeter
2008-068 KX574502 — Benton, OR; Exeter
2009-19 JN102493 KU574787 Benton, OR; Exeter
2009-31 JN102494 KU574790 Benton, OR; Exeter
2009-41 KX574503 — Benton, OR; Exeter
attenuata 2071029-51 JN102501 — Benton, OR; Norvell
2007-026 MZ352106 KU574794 Lincoln, OR; Exeter
2007-090 JN102498 — Benton, OR; Exeter
2007-176 (GQ165632) KU574761 Benton, OR; Exeter
benzokaumanii 1921120-01-T GQ165636 — Mendocino, CA; Norvell
2007-035 GQ165636 — Benton, OR; Exeter
2009-15 (JN102502) KU574785 Benton, OR; Exeter
californica MICH 11607-T JN102503 —Del Norte, CA; Smith
OSC 109290 EU669240 — Linn, OR; Smith
OSC 109332 EU846292 — Josephine, OR; Friend
2007-177 GQ165647 KU574764 Benton, OR; Exeter
2010-05 JN102504 Benton, OR; Exeter
#1921116-1 ruf-T GQ165707 —Mendocino, CA; Norvell
# 7500 GQ165708 — Mendocino, CA; Redhead
chefensis 2009-04-T MZ352102 KU57484 Lincoln, OR; Exeter
2091026cc6a MZ352101 — Benton, OR; Norvell
OSC 155803 MK326851 — Polk, OR; Christensen
2007-074 MZ352097 KU574779 Lincoln, OR; Exeter
2007-129 MZ352096 — L ane, OR; Exeter
dissiliens OSC 114217 EU846271 — Coos, OR; Sperling
2008-138 KF219569 KU574767 Benton, OR; Exeter
fallax 1921007-01 JN102512 — Vancouver Isl. BC; Redhead
2004-01 JN102511 — Benton, OR; Exeter
2007-140 MZ352103 KU574791 Benton, OR; Exeter
2007-159 JN102516 KU574795 Benton, OR; Exeter
2007-175 JN102518 KU574796 Benton, OR; Exeter
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 5
S C./F.
ITS RGBC, S/P;
P
gregaria 2091026cg8 JN102520 — Benton, OR; Norvell
2007-061 GQ165652 — Lincoln, OR; Exeter
2008-026 GQ165654 KU574772 Polk, OR; Exeter
2013-07b KJ450915 — Polk, OR; Exeter
kaumanii 1931015-02 JN102542 — Clallam, WA; Leuthy
2006-10 KF219573 KU574782 Benton, OR; Exeter
2006-13 JN102547 — Benton, OR; Exeter
2007-095 JN102524 — Polk, OR; Exeter
2008-027 KF219572 KU574781 Polk, OR; Exeter
2008-045 JN102526 — Lane, OR; Exeter
lilacifolia 1921111-06 KF219576 — Clackamas, OR; Norvell
2071018-12 GQ165657 KU574763 Lincoln, OR; Exeter
^2006-43 KF219580 — Polk, OR; Exeter
^2013-44 — KU574803 Lane, OR; Exeter
luteosquamulosa 2071029-33 GQ165668 KU574774 Benton, OR; Norvell
2071029-43 GQ165667 — Benton, OR; Norvell
2007-016 GQ165663 — Lincoln, OR; Exeter
WTU-F-003084 JN102528 — Snohomish, WA; Ammirati
ochraceocana 2007-033-T GQ165674 KU574775 Polk, OR; Exeter
OSC 134678 GQ165672 — Benton, OR; Villella
OSC 134679 GQ165671 — Tillamook, OR; Paque
2007-009 GQ165673 — Polk, OR; Exeter
olivacea 1921015-03 JN102533 — Jeerson, WA; Norvell
1921122-04 JN102530 — Mendocino, CA; Norvell
1941128-01 JN102532 — Clackamas, OR; Norvell
OSC 109501 EU846281 — Coos, OR; Sperling
OSC 113875 EU846282 — Douglas, OR; Kersens
2007-122 JN102534 KU574798 Benton, OR; Exeter
2007-133 JN102535 KU574799 Lane, OR; Exeter
2008-050 MZ352094 KU574766 Lane, OR; Exeter
2008-051 JN102536 — Benton, OR; Exeter
2008-129 GQ165678 KU574776 Benton, OR; Exeter
oregonensis 2001105-01 GQ165681 — Multnomah, OR; Norvell
OSC 67425 EU846273 — Douglas, OR; Goldenberg
2006-16 GQ165685 — Lane, OR; Exeter
2013-42 —KU574804 Lane, OR; Exeter
6 ... Norvell & al.
S C./F.
ITS RGBC, S/P;
P
phaeogaleroides 199-064a-T OM065392 — Benton, OR; Exeter
2009-29a MF737171 KU57489 Benton, OR; Exeter
2010-086 KX574499 — Benton, OR; Exeter
piceae MICH 11629-T MF737169 —Tillamook, OR; Smith
2007-178 (MZ352093) KU574762 Mendocino CA; Bojantchev
2009-12 KF219583 — Benton, OR; Exeter
pleurocystidiata 1940330-02-isoT GQ165688 —Clallam WA; Norvell
1930516-01 GQ165686 —Linn, OR; Bailey
2008-002 GQ165687 KU574765 Benton, OR; Exeter
pseudofestiva 1921104-10 KF219588 —Multnomah, OR; Norvell
2007-069 KF219589 KU574797 Lincoln, OR; Exeter
2007-070 KF219590 — Lincoln, OR; Exeter
radicata OSC 112980 EU846275 —Coos, OR; Rodenkirk
2006-19 GQ165696 — Benton, OR; Exeter
2008-131 GQ165695 KU574771 Benton, OR; Exeter
redheadii 2071018-17 JN102544 — Lincoln, OR; Norvell
2091026cg18 JN102541 — Benton, OR; Norvell
2007-106 JN102546 — Polk, OR; Exeter
2008-038 MZ352098 KU574783 Lincoln, OR; Exeter
scatesiae 1921015-19 GQ165701 — Jeerson, WA; McClenaghan
1931104-09 GQ165703 — Lincoln, OR; Norvell
2071029-02 GQ165699 KU574768 Benton, OR; Norvell
2007-151 GQ165700 KU574769 Benton, OR; Exeter
sipei 1971023-69 EU644706 — B enton, OR; Norvell
OSC 96908 EU644707 — Linn, OR; Bacheller
2007-123 GQ165704 KU574773 Benton, OR; Exeter
spadicea OSC 112482 EU697252 — Benton, OR; Giachini
OSC 113791 EU669364 — Douglas, OR; Wetzel
OSC 134542 MZ352099 — Douglas, OR; Sperling
2009-18 JN102550 KU574786 Benton, OR; Exeter
2009-20 JN102551 KU574788 Benton, OR; Exeter
*2011031ox1 tib-T KF219597 KU574780 Polk, OR; Norvell
* 2061114o2-O KF219596 KU574778 Polk, OR; Norvell
*2007-165 (KF219594) KU574777 B enton, OR; Exeter
P. sp. 1 OSC 155805 JN102507 KU574793 Lane, OR; Loring
2010-05 JN102504 — Benton, OR; Exeter
2014-01 KU574726 KU574801 Benton, OR; Exeter
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 7
S C./F.
ITS RGBC, S/P;
P
P. sp. 2 2009-09 KX574498 — Benton, OR; Exeter
2010-109 KX575400 — Benton, OR; Exeter
P. sp. 3 2071029-53 (KF219598) KU574770 Benton, OR; Norvell
2007-100 KX574493 — Polk, OR; Exeter
2007-152 KF219600 — Benton, OR; Exeter
2015-35 KU574725 KU574800 Benton, OR; Exeter
P. sp. 4 2010-009 MZ352104 KU574792 Benton, OR; Exeter
2015-01 KX574505 — Benton, OR; Exeter
2015-06 KU574727 KU574802 Benton, OR; Exeter
P. sp. 5 OSC 155806 MF957115 — Josephine, OR; Scelza
2013-14 KJ450918 — Benton, OR; Exeter
P. sp. 6 OSC 134575 GQ165677 — Linn, OR; Nakvasil
OSC 151647 MH819350 MH823881 Douglas, OR; Scelza
# = P. rufotubulina, ^ = P. riipes, and * = tibiikaumanii in
Norvell (1998ab, 2002, 2004) and Norvell & Exeter (2004, 2007, 2009).
(T = type); ITS sequences enclosed in parentheses not included in F. 1.
DNA amplication & sequencing
Standard protocols and published primers were followed for DNA extraction,
PCR amplication, and sequencing (White & al 1990, Gardes & Bruns 1993, Liu &
al. 1999, Matheny 2005). Amplications of the ITS region from 350 Phaeocollybia
collections representing 34 putative species utilized fungal primers ITS1 and ITS4b
(Gardes & Bruns 1993). Amplications of the RPB gene region from 46 collections
representing 26 putative species utilized the degenerate basidiomycete specic
primers bRPB2-6F and bRPB2-7.1R (Matheny 2005). Additionally, a new RPB2
forward primer, RPB2Phf (5′--3′, slightly downstream
from the bRPB2-6F primer) was designed based on existing Phaeocollybia RPB2
sequences and used successfully where the original primer pair failed aer multiple
attempts. In total 396 (350 ITS + 46 RPB2) new sequences were generated. Additional
ITS and RPB sequences were added from GenBank to provide a comprehensive
phylogeny for Phaeocollybia (Norvell & al. 2010, this publication). T provides
collection and sequence data for PNW specimens, and T 2 provides sequencing
information for extralimital taxa and the outgroup cited in F 1 and 2.
Separate gene regions were initially aligned with Clustal X (ompson &
al. 1997). Subsequent new sequences were aligned manually in MacClade 4.0
(Maddison & Maddison 2000). Taxa for which gene regions were not sequenced
were coded as missing. Sequences were aligned using MAFFT ver. 7 (Katoh & al.
2019) and manually corrected using MEGA X 10.0.0 (Kumar & al. 2018, Stecher &
al. 2020). All new sequences were deposited in GenBank.
8 ... Norvell & al.
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 9
F. 1. Phylogenetic analysis of the ITS1-5.8S-ITS2 rDNA region in strains of Phaeocollybia spp. Branch lengths were determined using the Bayesian
consensus outle. Nodes with posterior probability values <0.90 are indicated in blue. Type strains are indicated in bold.
10 ... Norvell & al.
Sequence analyses
Sequences were analyzed using programs available in Geneious Prime (version
2021.2.2). Sequence data were obtained from NCBI and compiled into a multiple
sequence alignment. e original ITS dataset contained 296 sequences identied as
Phaeocollybia spp. Duplicate sequences were removed. Representative sequences for
each taxon were retained and the nal dataset, including the outgroup, comprised
129 unique sequences. Sequences were trimmed to remove the and
motifs. For the analysis of partial RPB2 gene sequences, the dataset was composed of
49 unique sequences.
Sequences were aligned automatically using the Geneious alignment tool, and the
alignment was rened manually. Phylogenetic analyses were carried out using the
MrBayes 3.2.6 (Huelsenbeck & Ronquist 2001) plugin developed in Geneious. e
following parameters were used: substitution model GTR; rate variation gamma;
gamma categories 4; 5 million generations; subsampling frequency 1000; burnin
T 2: Sequences from extralimital Phaeocollybia spp. and outgroup
used in phylogenetic analyses
S F GB C
christinae MCVE 3539 JF908573 Italy
cidaris O-F-252891 UDB036543 Norway
elegans PDD 72723 KY827313 New Ze aland
festiva (ITS) WTU-F-053245 DQ494682 Norway
— (RPB2) WTU-F-053245 AY509118 Norway
gracilis PDD 88665 KY827314 New Zealan d
graveolens PERTH 5311586 AF501567 Australia
lugubris O-F-253794 UDB037370 Norway
MCVE 14619 JF908574 Ita ly
megalospora SP 445402 KC662116 Brazil
pakistanica-T SWAT 15-1560 KY007615 Pakistan
SWAT 15-1561 KY007616 Pakistan
ratticauda PDD 72678 KY827316 New Zealand
PDD 72544 KY827315 New Zealand
tenuis PDD 72672 KY827318 New Ze aland
Phaeocollybia sp. TRTC 157723 031-09 Costa Rica
AAM A1064 KF041417 Mexico
PDD 71198 KY827317 New Zealand
Galerina
semilanceata
WTU-F-007080 DQ486706 Washington USA
(RPB2 gene) WTU-F-007080 AY337357 Washington USA
All sequences represent the ITS region unless otherwise indicated.
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 11
F. 2. Phylogenetic analysis of the partial RPB2 gene region in strains of Phaeocollybia spp.
Branch lengths were determined using the Bayesian consensus outle. Nodes with posterior
probability values <0.90 are indicated in blue. Type strains are indicated in bold.
frequency 50%; heated chains 4; heated chain temperature 0.2; unconstrained branch
lengths GammaDir (1, 0.1, 1, 1); and shape parameter exponential (10). e outgroup
was Galerina semilanceata PBM 1398 (WTU); DQ486706 for the ITS analysis and
AY337357 for the RPB2 analysis, respectively). Phylogenetic trees were drawn using
the Bayesian consensus outle and annotations were added using Inkscape 0.92.4.
Phylogenetic results F –
Norvell & al. (2010) presented the rst comprehensive ITS phylogeny for the
genus based on 300 Phaeocollybia sequences (including two from Australia—
P. graveolens B.J. Rees & K. Syme, P. ratticauda E. Horak—and P. festiva (Fr.)
12 ... Norvell & al.
R. Heim from Norway) and the outgroup—Hebeloma radicosum (Bull.)
Ricken, Psilocybe semilanceata (Fr.) P. Kumm., Galerina marginata (Batsch)
Kühner—revealing a cohesive clade of 69 putative taxa with the Galerina-
like P. phaeogaleroides clade in the basal-most position within Phaeocollybia.
e current ITS tree (pruned to eliminate duplicates) conrms the existence
of three synonymies, several dicult species complexes, and six unnamed
species (F. 1). e three synonymies supported are P. riipes = P. lilacifolia
(>99.7% similarity), P. rufotubulina = P. californica (>99.5% similarity), and
P. tibiikaumanii = P. spadicea (>99.8% similarity (F 1, 2).
e comprehensive ITS phylogeny (not shown) generated from 296
sequences (plus the outgroup Inocybe pallidicremea Grund & D.E. Stuntz [as
I. lilacina (Peck) Kauman)] and Galerina semilanceata (Peck) A.H. Sm. &
Singer) and the trimmed 129-sequence ITS tree (F. 1) both support 29 PNW
species and 14 extralimital putative taxa. A familial relationship between
Phaeocollybia and Galerina is also supported, with both genera now classied
in Hymenogastraceae (Matheny & al. 2006, Kalichman et al. 2020).
e successful amplication of far fewer (46) RPB2 sequences prohibits
generating a robust multigene consensus tree. Nevertheless, both ITS (F. 1)
and RPB2 (F. 2) trees support the same clades. With insucient coverage
across the genus for 28S sequence data and with the two-gene phylogeny
well supported by convincing morphological characters, we do not include a
28S-based phylogeny at this time.
Taxonomy
Phaeocollybia chefensis Norvell & Exeter, sp. nov. F –
IF
Diers from Phaeocollybia kaumanii by its tibiiform cheilocystidia and from P. spadicea
by its uniformly carrot orange to orange-tawny coloration, larger basidiospores, and
solitary to scattered (never gregarious or cespitose) habit.
T: Oregon, Lincoln County, Cascade Head Experimental Forest, W of Hwy 12, 30 m S
Tillamook Co. line, 45.0447°N 123.9153°W, 245 m asl, 27.x.2009, Tsuga heterophylla (old
and young), Picea sitchensis, Vaccinium parviorum, Polystichum munitum, 2009-04
(holotype OSC 155804, isotype PNW; GenBank MZ352102, KU574784).
E: derived from the acronym (CHEF) for the type locality, Cascade Head
Experimental Forest.
P 70–80 mm diam, broadly conic-campanulate with acute papillate
umbo, upturned inner margin, down-turned outer margin, and straight edge;
glabrous, glutinous in rain with the gluten at times somewhat milky, non-striate,
overall brownish orange or foxy brown (Tawny, Orange Cinnamon); dried
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 13
F. 3. Phaeocollybia chefensis (Holotype, 2009-04). T : excavated specimens with intact
vertical monopodial pseudorhizae. B: Two views of the type collection in situ in Cascade
Head Experimental Forest, Lincoln County, Oregon. Five collections of 1–2 specimens were
collected from this same site during 1995–2021.
pileus uniformly copper metallic. C ~6 mm at the disc and conuent
with stipitipith, pale orangish white. O faintly oral with farinaceous
overtones; mild, not distinctive. L nearly free, ventricose,
thin with ± even edges, polydymous with 3–7 irregularly interspersed tiers of
lamellulae, narrow (4–5 mm, with average length/width ratio 5.5), close, pale
orangish bu (Warm Bu) when young, developing darker spots in maturity.
V sparse (when present, evident only as occasional darker brillose
Ron Exeter
14 ... Norvell & al.
remnants on stipe apex). S slightly eccentric, terete, 80–90 mm above
ground level, overall length including pseudorhiza ≤200 mm, apex 10–12
mm diam, ± equal above, gradually narrowing below toward pseudorhiza;
glabrous except for occasional brils, moist, nely longitudinally lined, apex
pale to deep pinkish orange (Pinkish Cinnamon, Orange Cinnamon) below
grading to dull pinkish brown (Fawn Color), 2 mm thick cartilaginous rind
surrounding compact brillose orangish white stipitipith, rind sometimes
with small perpendicular separations producing ~5 mm long recurved rind
patches. P vertical-monopodial, ≤2/3 overall stipe length,
gradually tapering to pale salmon-colored pointed to blunt origin, rm pith
brown where water-soaked, otherwise concolorous with stipitipith. S
dull pinkish brown (Fawn Color).
B (n = ): 8–8.9–10 × (5)5–5.6–5.8 µm, Q = (1.45)1.52–
1.65–1.81(1.84), limoniform with a protruding beaked apex in prole,
fusoid-elliptical to amygdaliform in face view, apical callus 0.5–1 µm
long (occasionally abrupt or more oen tapering to end), ornamentation
verruculose to verrucose except on smooth apical callus and eccentric
apiculus, suprahilar plage an indistinctly bordered area of lowered
ornamentation (oil immersion); orangish amber in KOH (ochraceous in
H2O, based on examinations of paratypes). B (n = 19): 4-spored,
curved; broadly clavate above narrower base, 25–34.2–41(43) × 6.5–8.1–
9.4(9.9) µm, sterigmata 2.0–3.0–3.7 µm long, curved; hyaline to dull orangish
brown, guttulate, granular, or uniformly oily. C 24.2–28.7–34
µm long, diameters (basal septum) 2–2.6–3 × (stomach) 4.9–5.7–6.9 × (neck)
1–1.3–1.6 × (capitulum) 1.5–2.0–2.6 µm (n = 13), abundant, intermixed with
basidia, secretory; capitulate tibiiform (primarily) and lageniform, (only
occasionally intermixed with thin-walled clavate elements), bases hyaline,
necks and capitula refractive, thick-walled, pale amber. P
absent except for isolated cheilocystidia scattered occasionally on the gill face.
L hyphae parallel, 65–80 × 3–6 µm, thin-walled, hyaline
inated, subgelatinized, narrowing toward gill edges to 2–3 µm diam and
giving rise to the subhymenial layer. S c. 25 µm thick, hyphae
2–4 µm diam, tightly packed, parallel, cylindrical, hyaline, gelatinized.
P : a bilaminate ixocutis with a ≥300 µm thick
with elements tightly compact in freshly dried material (but
readily separating in gel matrix aer 11 years in fungarium), hyphae >40 ×
1.5–5.0 µm, radially aligned, sinuous, long-branching, gelatinized, frequently
(spirally) gel encrusted, hyaline; c. 500 µm thick, gelatinized, vessel
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 15
F. 4. Phaeocollybia chefensis paratypes. [Lincoln Co.: 1951109-16 (A–C), 2007-074 (F),
2015-21 (H), 2021-1 (J); Lane Co.: 2007-129 (D, E); Polk Co.: 2018-LB4 (I); Benton
Co.: 2091026cc6 (G)]. A–C. ree views of the rst specimen from CHEF that fostered the
concept for P. tibiikaumanii in 1995. D. Young robust specimens excavated from Lobster Creek.
E. Pristine pseudorhizal tissues display strong magenta reaction in syringaldazine aer 15 minutes.
F. Second specimen retrieved from CHEF in 2007. G. Solitary specimen retrieved from Conner’s
Camp on Marys Peak. H. Basidiome at the original CHEF site prior to excavation in 2015. I. e
rst primordium excavated with its more mature partner in 2018 at Boulder Creek. J. Exeter’s
2021 negative syringaldazine reaction from this waterlogged solitaire cautions that adverse
environmental conditions do disable this usually reliable diagnostic tool.
a–c, e–g: Lorelei Norvell. d, h ,j: Ron Exeter i: Heidi Christensen.
16 ... Norvell & al.
F. 5. Phaeocollybia chefensis (Holotype, 2009-04): A. Colorless suprapellis in gelatinous
matrix overlying a pigmented subpellis [400×]. B. Pileipellis under oil [1000×]. C, D. Suprapellis
hyphae (gel encrustations detailed in C). E. Lightly sarcodimitic pileus tramal tissues with
relatively thin-walled vessel hyphae. F. Oleiferous hyphae in stipititrama (just below pellis
hyphae). G. Strongly sarcodimitic pseudorhizal trama with thick-walled vessel hyphae and thin-
walled exuous (arrow) hyphae.
hyphae with thinner walls; hyphae long, 4–12 µm diam at septa and inating
to ≤25 µm; pigments orange to dark brownish orange, diuse to occasionally
encrusting, soluble in KOH (forming droplets in mountant); small hyaline
crystals scattered infrequently throughout. S hyphae 75–100 ×
1.5–3(6) µm, parallel aligned, gelatinized, pigments diuse, pale amber to
reddish in places. R cells long × 4–10 µm, pale to dark amber,
heavily gel- and/or red-brown pigment-encrusted. T : lightly
(pileus) to noticeably (stipe, pseudorhiza) sarcodimitic, gelatinized, ±hyaline
except darkening in pseudorhiza; fusoid (sometimes tapering
to obtuse end), 25–100 × 3–13 (at septa), rigid walls 1–2 µm thick;
generally winding or curving around vessels, more or less cylindrical
but sometimes appearing attened, thin-walled, ≤50 µm long, diameters
narrowest (1–2 µm) in pileus and broadest (3–5 µm) in stipe;
occasional to frequent in tramal tissues throughout, sinuous,
aseptate, 3–10 µm diam, thin-walled, contents dull brown, oily. T
infrequent on aerial stipe apex, lageniform to (more oen)
tibiiform, frequent to abundant on rhizopellis, 8–20 µm long, 1–2 at base,
narrowing in refractive neck area before expanding when capitula (1–2
µm diam) present; arising directly from hypha and lacking basal septum,
secretory, hyaline, refractive to very pale amber. C absent
in all tissues.
S : strongly positive (pseudorhizal origin dark
burgundy in ve minutes; stipe (cross-section) and pileus and lamellae
leaching magenta aer ten minutes). F fresh material (of
holotype) not tested; dried lamellae dull yellow orange (one small area of one
specimen a brilliant orange yellow).
A —UNITED STATES, O, Benton Co.
Conner’s Camp, 44.5060°N 123.5565°W, 762 m asl, 200yo PSME TSHE 26.x.2009
2091026cc6 (MZ352101). Lane Co. Lobster Valley, 44.2268°N, 123.6148°W, 427
m asl, 200yo PSME young TSHE , 6.xi.2007 2007-129 (MZ352096). Lincoln
Co. Cascade Head EF, 30 m S Tillamook Co. Line, 45.0447°N 123.9153°W, 245 m asl,
~120yo PISI TSHE : 9.xi.1995 1951109-19; 18.x.2007 2007-074
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 17
Lorelei Norvell
18 ... Norvell & al.
(MZ352097, KU574779); 2.xi.2015 2015-21 (KX574495); 14.x.2021 2021-01.
Polk Co. Boulder Creek 44.8983°N 123.4995°W, 823 m asl, ~80yo ABPR PSME TSHE
10.x.2018 HChristensen 2018-LB4 (OSC 155803; MK326851).
E : autumn (October–November); basidiomes solitary
or in pairs in needle du under mature Abies procera, Tsuga heterophylla,
Picea sitchensis and/or Pseudotsuga menziesii with understory of Gaultheria
shallon, Oxalis oregana, Polystichum munitum, Vaccinium ovatum, and/or
V. parviorum. Known thus far from only ve sites along the Oregon coast and
in the Oregon coast range.
Discussion
Phaeocollybia chefensis
Diagnostic characters include a bright orange to brownish orange viscid
conic-campanulate pileus, seemingly robust stature that in age becomes limp
or accid, stued stipe, vertical monopodial pseudorhiza, medium-sized
limoniform verrucose basidiospores, thick-walled tibiiform cheilocystidia, and
lack of clamp connections. ITS and RPB2 sequence analyses strongly support
P. chefensis as an independent species (F 1, 2).
Morphologically, all paratypes match the type, except for the smaller (7.5–
8–9 × 4–4.5–5.2 µm) basidiospores measured in the immature 1995 specimen
(1951109-19), which in part explains why Norvell (2004) did not detect
the selection of a P. spadicea collection as type until aer sequence analyses.
e 1995 collection has not been sequenced, but sequences from the four
subsequent collections from the identical site strongly support the earlier
collection as P. chefensis. e molecularly conrmed collections imply Tsuga
heterophylla as the primary ectomycorrhizal associate; possible secondary
associates include Picea sitchensis at the type locality (245 m asl), Pseudotsuga
menziesii at the Lobster Creek (427 m asl) and Conner’s Camp (762 m asl)
sites, and Abies procera and Pseudotsuga menziesii at Boulder Creek (823 m
asl). Lamellae of fresh specimens examined under UV exhibited the strong
yellow uorescence characteristic of all PNW phaeocollybias except for the
uniquely non-uorescing P. gregaria A.H. Sm. & Trappe.
As noted above, Phaeocollybia chefensis closely resembles P. kaumanii
macroscopically, while microscopically it is quite similar to P. spadicea and
P. pseudofestiva A.H. Sm., Phaeocollybia chefensis shares with P. kaumanii a
brownish orange viscid pileus, stued stipe, vertical-monopodial pseudorhiza,
magenta syringaldazine reactivity, and morphologically similar basidiospores,
but the presence of refractive thick-walled tibiiform cheilocystidia and
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 19
Fig. 6 Phaeocollybia chefensis (Holotype, 2009-04 (,,); 2007-074 (,); 2007-129
(,): A, B. Cheilocystidia. C. Granular basidia, sterigmata, and orangish brown basidiospores
with smooth apical beaks (arrow). D. Basidiospores with older basidia lacking guttules or granules.
E. Comparison of verrucose spore ornamentation and less ornamented plage regions (arrows).
F. Abundant tibiiform diverticula on pseudorhizal pellis.
Lorelei Norvell
20 ... Norvell & al.
the absence of heavily gelatinized, strongly sarcodimitic elements in the
stipe and less extensive gelatinous matrix in the suprapellis clearly separate
P. chefensis from P. kaumanii. In the eld the much longer-lived P. kaumanii
can be distinguished by its larger (at times massive) and more robust stature,
sharp farinaceous odor and taste, strongly inrolled mature pileus edge, and
frequently gregarious habit.
Phaeocollybia spadicea, sharing many morphological features that led to
the selection of an orange form as type for P. tibiikaumanii, diers in its
darker tawny (“date-colored”) to frequently blackish brown pileus, abundant
rough brillose patches (invariably covered with tibiiform diverticula) on
the stipe apex, closely gregarious habit, and (oen) negative reactivity of the
pileus and lamellar tissues to syringaldazine. Specimen age and condition
may explain the variable syringaldazine reactivity in both P. chefensis and
P. spadicea.
Phaeocollybia pseudofestiva, which shares general basidiospore and
cheilocystidial morphology, syringaldazine reactivity, and odor and taste
is easily separated from P. chefensis by its dark green pileus, smaller size,
more prominently beaked basidiospores, cord-like pseudorhiza, and closely
gregarious habit.
Phaeocollybia tibiikaumanii and P. spadicea
As noted above, comparisons of ITS and RPB2 sequences from collections
designated as either P. spadicea or P. tibiikaumanii based on the presence/
absence of orange coloration (Norvell 1998a,b,c, 2004; Norvell & Exeter 2009)
revealed that all (including the holotype) except one (F. 3) P. tibiikaumanii
collection represented P. spadicea. Subsequent morphological comparisons
have shown that P. spadicea pileus colors range along a foxy orange to tawny to
deep brown continuum. Additionally, all seven P. chefensis collections occurred
singly or in pairs in contrast to P. spadicea, which typically occurs in closely
gregarious to cespitose clusters; Alexander Smith’s notes archived in MICH
describe an “orange” spadicea that he provisionally named P. “caespitosa”
(a name he also provisionally applied to what was to become P. scatesiae A.H.
Sm. & Trappe; Norvell 1995). us, we propose the synonymy:
Phaeocollybia spadicea A.H. Sm., Brittonia 9: 215, 1957. F.
= Phaeocollybia tibiikaumanii Norvell, Mycotaxon 90: 248, 2004.
S Phaeocollybia spadicea (listing those previously cited as
P. tibiikaumanii in Norvell (2004) and/or Norvell & Exeter (2004, 2009)—UNITED
STATES. O: Benton Co. Conner’s Camp 44.5060°N 123.5565°W, 762 m asl,
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 21
F. 7. Phaeocollybia spadicea [208-55 (); 2011031o2-1 (); 1921015-10 ()].
A. Gregarious cluster on Oregon’s Marys Peak (as P. tibiikaumanii in Norvell & Exeter
2009). B. Excavated P. tibiikaumanii holotype from the Polk County Chronosequence Study.
C. “Typical” brown-capped P. spadicea specimens in Hoh Valley, Olympic National Park.
200yo PSME TSHE, 5.xi.2005 2008-55 (KF219592); Ernest Creek 44.4185°N
123.5165°W, 610m asl, 200yo PSME TSHE 15.xi.2007 2007-165 (KU574777,
KF219594); Running Bear 44.4774°N 123.5769°W, 696 m asl, 60yo PSME TSHE,
6.xi.2000 DCalver 200-103 (KF219595). Polk Co. Pedee Chronosequence Study
44.8113°N 123.5212°W, 460 m asl, 150yo PSME TSHE 4.xi.1998 (all by
) 1981104o1-01, 1981104o2-02–04 (with CHibbler); 5.x.2000 200-
045; 18.x.2000 2001018o1-15,o2-25; 1.xi.2000 200-091, 2001101o2-30;
31.xi.2001 2011031ox1 (P. tibiikaumanii holotype; KF219597, KU574780),
2011031o2-03,04,05,06,08, 2011031ox-02; 14.xi.2011 2011114o1-01,02,03,
2011114ox-01; 15.x.2002 2021015o2-01; 13.xi.2002 2021113o2-01,02,03;
14.xi.2006 2061114o2-O (KF219596, KU574778), 2006-47, 16.x.2007 rle2007-
060 (KF219593). W: Jeerson Co. Hoh Valley @ Twin Creek 47.8307°N
124.0012°W, 180 m asl, ~400yo PISI TSHE 15.x.1992 1921015-10
spds1
(w ) (WTU-F-003698; previously cited and RFLP-conrmed as P. spadicea).
A Phaeocollybia P. —UNITED
STATES. O: Benton Co. Green Peak BLM Density Management Study 44.366°N
123.455°W, (all collections by & ): Clear-cut transect (pre-treatment)—
610m asl, 65yo PSME TSHE 24.xi.1998 1981124c1-04 (w TFennell); High
Retention transect 579m asl, 65yo PSME TSHE — 8.xii.1999 1991208hx.03
(w ) (PNW, DAOM), 27.xi.2000 2001127hx3; Klickitat BLM Unit-3 44.4406°N
123.5673°W, 60yo PSME TSHE 30.x.2000 200-065; 16.xi.2000 RLE200-252;
Klickitat Unit-6 T.S. 44.4548°N 123.5473°W, 50yo PSME TSHE : 30.x.2000
200-082,83,90, 14.xi.2000 200-187b; 44.4260°N 123.5472°W, 60yo PSME
TSHE : 30.x.2000 200-070; R B T.S. 44.4774°N 123.5770°W,
200yo PSME TSHE 9.xi.1998 1981109.108e w , 6.xi.2000 200-103;
28.x.2002 2002-16. Linn Co. Keel Flats T.S. 44.5133°N 122.6518°W, PSME
6.xii.1999 KScott 199-BK. Polk Co. Pedee Crk head waters near Cold Springs
44.8233°N 123.4862°W, 610m asl, 200yo PSME TSHE 26.xi.2001 RLE 2001-
112,115. W: Clallam Co. Olympic NF Klahanie Campground 47.9657°N
124.3057°W, 250yo PISI TSHE 18.x.1992 STrudell 1921018-4
usps3
(WTU-F-003646).
c: Elixabeth Fox.
a: Ron Exeter. b: Lorelei Norvell.
22 ... Norvell & al.
Phaeocollybia rufotubulina and P. californica
On November 16, 1992, numerous densely gregarious orange
phaeocollybias in Jackson State Forest (Mendocino County, California) were
collected and referred to P. californica. is rst highly informative collection
contained numerous pin-head primordia that helped establish the presence
of a universal veil for all Phaeocollybia species and introduced the ‘sequential-
racemose’ pseudorhiza arising from a horizontal ‘mother rhizomorph’ within
the soil (Norvell 1998b, 2004).
Restriction Fragment Length Polymorphism (RFLP) analyses of the ITS
region from type materials supported separation of P. rufotubulina from
P. californica and P. scatesiae (Norvell 1998a), indicating a new species and
supporting the synonymy of P. scatesiae under P. californica proposed by Horak
(1977). e 2009 ITS sequence analyses, however, clustered P. californica and
P. rufotubulina within one intermixed clade well-separated from P. scatesiae,
a brownish species macroscopically easily distinguishable in the eld from
the two orange phaeocollybias. Phaeocollybia scatesiae is now supported as
independent from P. californica (F ,).
Given the diculty encountered in isolating DNA from the P. californica
holotype (obtained in 2011), it is probable that the 58-year-old P. californica
DNA isolated in 1994 was contaminated by DNA isolated in the same run
from the much younger P. scatesiae isotype. We hereby propose the synonymy:
Phaeocollybia californica A.H. Sm., Brittonia 9: 216, 1957.
= Phaeocollybia rufotubulina Norvell, Mycotaxon 90: 243, 2004.
Norvell & Exeter (2009) presented P. californica (pp. 63–70) and P. rufotubulina
(pp. 175–180) as separate species; all material related to P. rufotubulina
throughout that monograph as well as those in Norvell (1998a,b, 2004) should
now be referred to P. californica.
Phaeocollybia riipes and P. lilacifolia
During a two-month long collecting expedition through western North
American coastal rainforests 1992, several small lilac-gilled tawny to dark
brown-capped phaeocollybias were collected and provisionally determined
as P. lilacifolia. Except for accid small basidiomes and unusually small
basidiospores, the specimens morphologically matched Smith’s P. lilacifolia
type from Washington’s Mt. Rainier Park (Smith 1957a). However, 1994
restriction digests produced a RFLP prole unique to these collections.
Unable to extract DNA from the Washington P. lilacifolia holotype or Smith’s
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 23
Cascade Head Experimental Forest paratype, Norvell (1998a, 2002) proposed
the name P. riipes (honoring its unique RFLP ngerprint) for the specimens
that were morphologically separated by unusually small basidiospores and
stature.
During 2001–2013, numerous small P. riipes specimens were collected
from the Polk County old growth chronosequence transect; not until 2007,
however, was the rst recognizable P. lilacifolia collected from Lincoln
County’s Saddle Bag Mountain, c. 25 km southeast of Cascade Head
Experimental Forest. e Saddle Bag collections comprised robust healthy
specimens with normal-sized basidiospores. In 2009, ITS phylogenetic
analyses clustered P. lilacifolia sequences and all but one P. riipes sequences
within one clade, a clade supported by our current ITS and RPB2 phylogenies.
Disposition of 4592sl (MZ352100), a Loring collection from Josephine
County (initially identied as P. r i ip e s), remains unresolved at this time.
Although we have not yet successfully isolated DNA from Smith’s
P. lilacifolia holotype or paratypes, the small “limp” stature, poor condition,
and age of the 1992–1994 collections suggest that their peculiar RFLP
proles resulted from an unknown contaminant. e following synonymy
is proposed:
Phaeocollybia lilacifolia A.H. Sm., Sydowia Beih. 1: 59, 1957.
= Phaeocollybia riipes Norvell, Mycotaxon 81: 102, 2002.
All keys, descriptions, and photos of P. riipes in Norvell (1998a, 2002) and
Norvell & Exeter (2009) should be referred to P. lilacifolia.
Revised general key to Pacic Northwest Phaeocollybia species
Basidiospore size ranges of measurements taken from holotype specimens are
cited for species; means of measurements taken from molecularly conrmed
specimens are cited for complexes.
1. Basidiomes small: pileus ≤50 mm diam, and never green or drab;
stipe apex usually ≤4 mm diam and never drab ........................... 2
1. Basidiomes medium to large: pileus usually ≥50 mm diam;
stipe apex usually >5 mm diam ......................................... 5
2. Basidiospores ellipsoid, ornamentation minutely punctate and
small apical callus detectable in 1000× oil immersion;
clamp connections present ............................................. 3
2. Basidiospores limoniform, ornamentation verrucose and
protruding apical beak visible without oil immersion lens;
clamp connections absent .............................................. 4
24 ... Norvell & al.
3. Spores small (5–6 × 3–3.5 µm); cheilocystidia lageni-/tibiiform,
necks narrow and thick-walled, abundant but inconspicuous,
dicult to see in dense gelatinous matrix;
basidiome collybioid, stipe pliable .............................. P. radicata
3. Spores larger (~10 × 6 µm); cheilocystidia thin walled,
narrowly clavate, extending well beyond hymenium;
basidiome mycenoid, stipe fragile and easily broken . . .
P. phaeogaleroides complex
4. Stipe and pseudorhiza eshy, not shiny or brittle;
cheilocystidia tibiiform, necks and capitula refractive and thick-walled;
pleurocystidia present and, frequent on gill faces;
spores 8–11 × 5–7 µm, tapering to straight beaks;
phenology vernal ...................................... P. pleurocystidiata
4. Stipe polished, corneous, soon hollow;
pseudorhiza lateral monopodial, brittle and wire-like (criniform);
cheilocystidia clavate, thin-walled; pleurocystidia absent;
spores 7–8.5 × 5–5.5 µm, big-bellied with abrupt tilted beaks;
phenology autumnal .........................................P. attenuata
5. Young pileus green, rapidly aging to brown or brownish olive .................. 6
5. Young pileus ochre, orange, tawny, brownish, or drab (never green) ............. 8
6. Young lamellae violet; spores ~9 × 5.3 µm, moderately beaked,
in face view fusoid or naviculate, verrucose to marbled;
cheilocystidia clavate, thin-walled, apices swollen to subcapitate,
only rarely forming liform apical outgrowths . . . . . . . . . . . . . . P. fallax complex
6. Young lamellae creamy to yellowish bu; spores limoniform with
pronounced beak, +/- ovate in face view, rugulose to warty roughened;
cheilocystidia clavate or tibiiform ....................................... 7
7. Spores 7.5–8 × 4.5–5 µm, with long, projecting beaks;
cheilocystidia tibiiform with narrow refractive necks ......... P. pseudofestiva
7. Spores ~10 × 6 µm, ‘turtle-backed’ with abruptly protruding
eccentric refractive beaks; cheilocystidia thin-walled, clavate,
liform apical outgrowths frequent in age . . . . . . . . . . . . . . . . P. olivacea complex
8. Spores ellipsoid, short (<7.5 µm), punctate-roughened;
cheilocystidia clavate, thin-walled ....................................... 9
8.
Spores limoniform, verruculose to warty;
cheilocystidia tibiiform or clavate
....................................... 11
9. Basidiome drab to gray, robust; stipe ≤20 mm diam, stout, rmly stued;
pseudorhiza eshy; taste and odor cucumber-farinaceous;
spores 6–7.5 × 4–4.5 µm; all tissues soon deep magenta in
syringaldazine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. oregonensis
9. Basidiome orange to auburn colored, fragile; stipe ≤13 mm diam,
slender, hollow; pseudorhiza cord-like; taste and odor mild;
pileus & lamellae syringaldazine negative . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 25
10. Clamp connections abundant; pileus orange with yellowish margin,
viscid, conic-campanulate; cheilocystidia lamentous to narrowly clavate;
spores 5.5–6.5 × 3.5–4.3 µm ................................... P. dissiliens
10. Clamp connections absent; pileus uniformly auburn colored,
subviscid, campanulate with a raised blunt umbo;
cheilocystidia narrowly clavate with long pedicels and subcapitate apices;
spores 5.5–6.5 × 3–4 µm .......................................... P. sipei
11. Pileipellis 3-layered with colorless gelatinized middle layer
between yellow to yellow-orange top and bottom layers;
pileus minutely scaly (appressed), dry to subviscid (never glutinous) ........ 12
11.
Pileipellis 2-layered, with colorless top layer and variously pigmented lower layer;
pileus bald, smooth (never appressed scaly), subviscid to glutinous ......... 13
12. Spores large, 9–11 × 5–6.8 µm;
pileus dry to greasy, yellowish ochre to ochraceous gold;
frequently associated with Abies . . . . . . . . . . . . . . . . . . . . . . . . P. luteosquamulosa
12. Spores smaller, 7.2–8.7 × 4.4–5.5 µm;
pileus greasy to subviscid, tawny ochraceous or tawny;
associated with Pseudotsuga or Tsuga ....................... P. ochraceocana
13. Cheilocystidia tibiiform, broad bases thin-walled,
capitula and narrow necks refractive and thick-walled;
stipe hollow or stued ................................................ 14
13. Cheilocystidia variably cylindrical to clavate,
lamentous apical secondary growth occasional in older specimens
but always lacking refractive thick-walled necks;
stipes stued with rm pith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
14. Mature stipe tubular and hollow; basidiomes closely gregarious to fasciculate,
arising from a branching rhizomorphic cord . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14. Mature stipe stued with rm compact pith; solitary to closely gregarious,
arising from unbranched eshy (not cord-like) pseudorhiza ................. 16
15. Pileus heavily glutinous, acutely conic-campanulate, yellowish to dark brown;
habit in dense fasciculate mounds; suprapellis thick, colorless,
hyphae kinked, not spirally pigment-encrusted and with septa obscured
by thick gel matrix; spores 8.5–9.5 × 4.5–5.5 µm .................... P. scatesiae
15. Pileus moist to viscid, obtusely convex-campanulate, red orange to tawny;
habit in troops, arcs, or (rarely) fasciculate mounds; suprapellis compact,
amber to orange, hyphae spirally pigment-encrusted with refractive septa;
spores 8.5–10 × 5–6 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. californica
16. Pileus foxy orange to orangish brown; stipe apex orange cinnamon colored;
habit solitary or paired, never gregarious; all tissues soon magenta in
syringaldazine; spores 8–10 × 5–5.8 µm ......................... P. chefensis
16. Pileus tawny to dark brown; stipe apex pallid to drab;
habit gregarious to fasciculate; only pseudorhiza magenta in
syringaldazine; spores 7.5–9 × 4.5–5 µm . . . . . . . . . . . . . . . . . . . . . . . . P. spadicea
26 ... Norvell & al.
17. Clamp connections frequent throughout, most easily seen
in pileus suprapellis
and on cheilocystidia;
pileus campanulate with oen papillate umbo,
tawny ochraceous to tawny;
cheilocystidia cylindrical to narrowly clavate;
spores 8.4–9.2 × 5–6 µm ..................................... P. ammiratii
17. Clamp connections lacking (rarely in stipitipellis);
pileus shape & color, cheilocystidial shape, and spore size varied . . . . . . . . . . . 18
18. Young lamellae whitish, smoky gray, or deep violet;
young pileus colors tawny, brownish-pink or drab;
pileus and lamellae magenta in syringaldazine ........................... 19
18. Young lamellae pinkish, orangish, or yellowish;
young pileus colors rarely pink or drab;
syringaldazine reactivity varied . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19. Spores 7.5–9.4 × 4.5–5.5 µm; young lamellae intensely bluish lilac to violet;
pileus tawny to deep brown; taste not distinctive;
pseudorhiza syringaldazine negative ............................ P. lilacifolia
19. Spores 9 × 5.5 µm;
young lamellae white (pinkish) or ash gray, never lilac or violet;
pileus pinkish, drab, or purplish brown; taste ‘bitter-cucumber’ farinaceous;
all tissues soon deep magenta in syringaldazine ........... P. benzokaumanii
20. Young lamellae never uorescing under UV;
pileus yellow tan, soon darker (cocoa brown); young stipe ivory tan,
lower stipe staining orange or orange banded at ground level;
habit densely gregarious;
syringaldazine negative;
spores 9–10 × 4.5–6 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gregaria
20. Young lamellae uorescing under UV;
pileus orange or tawny; young stipe apex bu, orange, or pale cinnamon,
lower stipe colors similar to apex, habit scattered to gregarious,
syringaldazein negative or positive; spores 8–12 µm long ................. 21
21. Basidiome small to moderately large; stipe apex ≤12 mm diam (slender),
stipe stued but larval infested, leaving a hollow rind at ground level;
pileus conic umbonate, subviscid, uniformly bright apricot-/peach-colored;
all tissues negative in syringaldazine; pileipellis hyphae colorless,
spirally gel-incrusted, subpellis pigments KOH soluble, diuse;
spores 8–9.5 × 5–6 µm, beaks short (≤5 µm), straight ............... P. piceae
21. Basidiome large to massive; stipe apex 10–25 mm diam (robust),
stipe long-lived, stued with rm, insect-free pith;
pileus campanulate with obtuse umbo, viscid to glutinous, orange to tawny;
all tissues strongly magenta in syringaldazine;
pileus subpellis pigment-encrusted, pigments not KOH soluble;
spores with slightly tilted long (≤1.5 µm) beaks . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Phaeocollybia chefensis sp. nov. (U.S.A.) ... 27
22. Spores 7.5–10 × 4–6 µm, limoniform, beak forward-tilted;
pileus +/- uniformly orange, tawny, or orangish brown,
typically drying with a burgundy-colored peaked umbo;
lower stipe and pith staining rst orange, then orangish brown;
cheilocystidia cylindrical or narrowly to broadly clavate,
rarely pedicellate and subcapitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. kaumanii
22. Spores 8.5–12 × 5–7 µm, amygdaliform with long ‘belly’, beak back-tilted;
young pileus with reddish tawny umbo, amber margin, and tan edge,
eventually uniformly mahogany or dark reddish brown;
lower stipe and pith staining reddish brown;
cheilocystidia clavate to
subcapitate with subglobose apices,
pedicellate, catenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. redheadii
Research on Pacic Northwest phaeocollybias is ongoing; a fully revised second
edition of P P N N A will be
released aer several additional taxa have been published.
Acknowledgments
e Pacic Northwest Phaeocollybia consortium wishes to acknowledge earlier
assistance given by individual collectors, government surveyors and agencies,
fungarium curators (BPI, DAOM, F, HSC, MICH, O, OSC, NY, SFSU, TENN, UBC,
UC, WTU), technicians, and others cited in full on p. 205 of Norvell & Exeter (2009).
We also acknowledge the excellent molecular work by M.N. Louise Lefebvre in 2010
at Agriculture & Agri-Food Canada in Ottawa. Kelli Van Norman (Interagency
Special Status/Sensitive Species Program of the Forest Service (Region 6) PNW
Region and Oregon/Washington Bureau of Land Management) is thanked for
facilitating molecular sampling of government Phaeocollybia collections; particular
thanks are due Darci Rivers-Pankratz (also of ISSSP) for providing data on specimens
inventoried for the Northwest Forest Plan, facilitating microscopical reevaluation,
and forwarding Heidi Christensen’s Boulder Creek collection for examination.
We also thank Dr. Joe Ammirati (University of Washington, Seattle, USA),
Dr. Egon Horak (Innsbruck, Austria), and Dr. P. Brandon Matheny (University
of Tennessee, Knoxville, USA) for their greatly appreciated expert presubmission
reviews. Egon Horak is further thanked for sharing his SEM-based basidiospore
measurements for selected type specimens.
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