Pouzarella alissae, a new species from northwestern California, United States

Abstract

Pouzarella alissae sp. nov. is known only from north coastal California. It is characterized by the evanescent silvery-white layer of fibrils and squamules initially covering the pileus and stipe surface, the dark greyish brown to dark brown innately fibrillose pileus and stipe, a pileipellis of entangled hyphae with cylindro-clavate terminal cells, a suprapellis with cytoplasmic pigments, a subpellis with encrusted pigments, heterogeneous lamellar edges, lageniform to rostrate-ventricose leptocystidia, cheilocystidia and pleurocystidia, abundant aborted basidia, and basidiospores averaging

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ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MYCOTAXON
http://dx.doi.org/10.5248/130.1153
Volume 130, pp. 1153–1164 October–December 2015
Pouzarella alissae, a new species
from northwestern California, United States
David L. Largent1* & Sarah E. Bergemann2
1 Biological Sciences, Humboldt State University, 1 Harpst St, Arcata CA 95521, USA
2 Evolution and Ecology Group, Biology Department, Middle Tennessee State University,
PO Box 60, Murfreesboro TN 37132, USA
* Correspondence to: mrp@humboldt1.com
Abstract —Pouzarella alissae sp. nov. is known only from north coastal California. It is
characterized by the evanescent silvery-white layer of brils and squamules initially covering
the pileus and stipe surface, the dark greyish brown to dark brown innately brillose pileus
and stipe, a pileipellis of entangled hyphae with cylindro-clavate terminal cells, a suprapellis
with cytoplasmic pigments, a subpellis with encrusted pigments, heterogeneous lamellar
edges, lageniform to rostrate-ventricose leptocystidia, cheilocystidia and pleurocystidia,
abundant aborted basidia, and basidiospores averaging <11 µm in length.
Key words —Basidiomycota, Entolomataceae, phylogeny, taxonomy
Introduction
e name Pouzarella is applied to fungi in the Entolomataceae that possess
stipitate–pileate basidiomata with the following features: basidiome typically
mycenoid with a strigose stipe base, pileipellis disc composed of erect to
semi-erect multicellular hyphae, tramal hyphae with encrusted pigments,
nodulose heterodiametric 5–9-angled basidiospores, and the absence of clamp
connections. Phylogenetically, these fungi form a well-supported clade (He
et al. 2013), and Pouzarella Mazzer is applied as a generic concept (Mazzer
1976; Karstedt et al. 2007; Baroni et al. 2008, 2012; Horak 2008) or is treated
as Entoloma subg. Pouzarella (Mazzer) Noordel. (Manimohan et al. 2006,
Noordeloos 2004, He et al. 2013). We treat Pouzarella as a genus because all
species form a monophyletic group with distinctive morphological characters.
Although Pouzarella species are distributed worldwide, they are encountered
most frequently in mesic areas that receive >20 in. annual rainfall (Mazzer
1976). In North America, Pouzarella is common in the eastern United States

1154 ... Largent & Bergemann
Table 1. Collections used in the phylogenetic analyses.
New sequences generated for this study are shown in bold.
Square brackets enclose GenBank names diering from those in Fig. 1.
Taxa Collection
identifier
GenBank accession numbers
mtSSU LSU RPB2
Alboleptonia angustospora
[Alboleptonia a. sericella]
MCA1978 GU384583 GU384609 GU384632
Alboleptonia sericella
[Entoloma sericellum]
10 GQ289330 GQ289190 GQ289261
Catathelasma imperiale 11CA001A KR869913 KR869941 KC816816
Claudopus minutoincanus DLL9871 HQ731511 HQ731514 HQ731517
Claudopus parasiticus 10617 TJB KR233902 KR233872 KR233924
Claudopus viscosus DLL9788 HQ31513 HQ31516 HQ31518
Clitocella fallax [Rhodocybe fallax] 25668OKM KR869909 AF261283 KC816937
and Canada, but is rare in the west where it is restricted to a narrow coastal
region extending from southern California to southern British Columbia
(Largent 1994, Mazzer 1976).
Of the y-nine Pouzarella species reported worldwide in Index Fungorum,
four have been described from western North America: P. fernandae (Romagn.)
Largent and P. fulvostrigosa (Berk. & Broome) Mazzer from Washington,
P. araneosa (Quél.) Mazzer from California, and P. versatilis (Gillet) Mazzer
from California, Oregon, and Washington (Largent 1994, Mazzer 1976). In this
report, we describe a third and new species from California, P. alissae.
Materials & methods
Macromorphological and micromorphological features
Techniques and equipment for collecting and describing basidiomata in the eld
and GPS coordinates have been described in Largent et al. (2011a,b), while techniques
for color descriptions using Kornerup & Wanscher (1978) and factors determined from
mathematical analyses in the descriptions are covered in Largent et al. (2013a,b). Dried
specimens were examined microscopically with a research-grade trinocular Nikon
Eclipse Ci-L compound microscope. Microphotographs were made using a Lumenera
Innity 2-5 CCD digital camera, and all microscopic measurements were obtained
using Innity Analyze soware 6.5.0.
DNA sequences and phylogenetic analyses
Sequences of P. alissae were obtained from the mitochondrial small subunit ribosomal
RNA (mtSSU), the nuclear large subunit ribosomal RNA (LSU), and second largest
subunit of RNA polymerase II (RPB2) following protocols described in Largent et al.
(2011b). Sequences were edited using Sequencher 4.2.2 (Gene Codes, USA) and aligned
using MAFFT v. 7 (Katoh & Standley 2013). irty-eight sequences were generated
for this phylogenetic analysis along with sequences obtained from GenBank including
those for the majority of Pouzarella species and from species from other representative
clades as dened in Baroni & Matheny (2011) (Table 1).

Pouzarella alissae sp. nov. (U.S.A.) ... 1155
Clitopilopsis hirneola REH8490 GU384587 GU384611 KC816904
Clitopilus apalus 26394WAT KR869907 KR869936 KC816906
Entocybe haastii [Entocybe sp.] DLL10087 JQ793645 JQ793652 JQ793659
Entocybe trachyospora
[Rhodocybe trachyospora]
5856 TJB GU384605 GU384629 GU384658
Entoloma abortivum 5772 TJB KR233897 KR233869 KR233926
E. albidoquadratum i4 GQ289291 GQ289151 GQ289223
E. bloxamii 219 GQ289294 GQ289154 GQ289226
E. caespitosum GDGM 27564 JQ993070 JQ410327 JQ993078
E. changchunense HMJAU 3886 JQ993061 JQ993095 —
E. crassicystidiatum GDGM 28821 JQ993058 JQ291567 JQ993083
GDGM 27357 JQ993056 JQ291569 JQ993085
E. discoloratum DLL10217 JQ793646 JQ793653 JQ793660
E. fragilum MCA2415 KJ021690 KJ021700 KJ021694
E. furfuraceum GDGM 28818 JQ993062 JQ993094 JQ993084
E. indoviolaceum i13 GQ289312 GQ289172 GQ289243
E. myrmecophilum 231 GQ289314 GQ289174 GQ289245
E. omiense GDGM 27563 JQ993067 JQ410330 JQ993079
E. pallidocarpum GDGM 28828 JQ993074 JQ410331 JQ993080
E. praegracile GDGM 29251 JQ993072 JQ320129 JQ993077
E. procerum 70 GQ289323 GQ289183 GQ289254
E. prunuloides 40 GQ289324 GQ289184 GQ289255
E. readiae 102 GQ289326 GQ289186 GQ289257
E. rhodopolium 8 GQ289327 GQ289187 GQ289258
E. rugosiviscosum DLL9676 JQ793647 JQ793654 JQ793661
E. sericatum 28 GQ289329 GQ289189 GQ289260
E. sinuatum 50 GQ289333 GQ289193 GQ289264
E. subaraneosum KA12-1534 — KJ523137 —
GDGM 28823 — JQ291568 —
E. a. subsinuatum [E. sinuatum] 5349 TJB KR233896 KR233868 AY691891
E. tenuissimum GDGM 28813 JQ993059 JQ993097 JQ993086
GDGM 28814 JQ993060 JQ993096 JQ993087
E. undatum 18 GQ289342 GQ289202 GQ289270
E. valdeumbonatum 189 GQ289343 GQ289203 GQ289271
E. violaceotinctum DLL10088 JQ793643 JQ793650 JQ793657
E. violaceovillosum i2 GQ289345 GQ289205 GQ289273
E. yunnanense GDGM 28815 JQ993057 JQ320128 —
Inocephalus hypipamee DLL10071 JQ624604 JQ624609 JQ624616
I. “lactiuus” 7962 TJB — AF261304 —
I. plicatus DLL9691 JQ624605 JQ624610 JQ624617
I. virescens DLL9772 KR233889 KR233859 KR233918
Lepista nuda 11CA041 KJ021692 KJ021705 KJ136110
Leptonia ambigua DLL9872 JQ756399 JQ756414 JQ756429
L. boardinghousensis DLL10086 JQ756398 JQ756413 JQ756428
L. omphalinoides DLL9673 JQ756400 JQ756416 JQ756431
L. odorifera 6534 TJB KR233899 KR233870 KR233927
L. poliopus DLL10209a JQ756402 JQ756418 JQ756433

1156 ... Largent & Bergemann
L. serrulata [Entoloma serrulatum] 163 GQ289332 GQ289192 GQ289263
L. “subdecurrentiba” [L. sp.] MCA1486 GU384589 GU384623 GU384635
L. tjallingiorum [Entoloma tjallingiorum] 243 GQ289337 GQ289197 GQ289267
L. trichomata 11CA006 KR233873 KR233873 KR233903
L. umbraphila DLL9793 JQ756409 JQ756424 JQ756440
Mycena a. pura 11CA007 KR869914 KR869942 KC816995
Nolanea cetrata DLL9531 KF738927 KF738942 KF771346
N. cf. conferenda [Entoloma conferendum] 6 GQ289300 GQ289160 GQ289231
N. hebes [Entoloma hebes] 46 GQ289310 GQ289170 GQ289241
N. sericea DLL9527 KR233882 KR233851 KR233913
N. strictior var. isabellina
[E. strictius var. isabellinus]
7710 TJB GU384594 GU384618 GU384641
Panellus stipticus 11CA052 KR869915 KR869943 KC816996
Paraeccilia sericeonitida
[Entoloma sericeonitidum]
7144 TJB EF421098 AF261315 EF421016
Pouzarella albostrigosa DLL9641 HQ876557 HQ876535 HQ876513
DLL9663 HQ876558 HQ876536 HQ876514
P. alissae DLL10490 KT074359 KT074357 KT074361
P. araneosa [Entoloma araneosum] 14 GQ289293 GQ289153 GQ289225
P. debilis DLL9784 HQ876550 HQ876528 HQ876506
P. farinosa DLL9900 HQ876537 HQ876515 HQ876494
DLL9934 HQ876538 HQ876516 HQ876495
P. fusca DLL9623 HQ876549 HQ876527 HQ876505
P. lageniformis DLL9895 HQ876545 HQ876523 HQ876501
P. lasia DLL9662 HQ876551 HQ876529 HQ876507
DLL9670 HQ876552 HQ876530 HQ876508
DLL9729 HQ876553 HQ876531 HQ876509
DLL9807 HQ876555 HQ876533 HQ876511
DLL9811 HQ876556 HQ876534 HQ876512
P. farinosa DLL9900 HQ876537 HQ876515 HQ876494
DLL9934 HQ876538 HQ876516 HQ876495
P. nodospora 5716 TJB KT074358 KT074356 KT074360
P. pamiae DLL9794 HQ876539 HQ876517 HQ876496
DLL9808 HQ876540 HQ876518 HQ876497
DLL9834 HQ876541 HQ876519 HQ876498
P. parvula DLL9901 HQ876546 HQ876524 —
P. pilocystidiata DLL9848 HQ876542 HQ876520 HQ876499
P. setiformis DLL9809 HQ876547 HQ876525 HQ876503
DLL9810 HQ876548 HQ876526 HQ876504
Rhodocybe roseiavellanea 8130 TJB KR869901 KR869930 KC816982
R. spongiosa MCA2129 GU384604 GU384628 GU384657
Rhodophana nitellina 7861 TJB KR869900 KR869929 KC816959
Tricholoma avovirens 11CA038 KJ021691 KJ021704 KC816997
Trichopilus porphyrophaeus
[Entoloma porphyrophaeum]
113 GQ289322 GQ289182 GQ289253

Pouzarella alissae sp. nov. (U.S.A.) ... 1157
Figure 1. Maximum-likelihood topology, highlighting the sub-clade within Pouzarella containing
the new species, Pouzarella alissae. Each sequence is labeled with the GenBank-listed isolate identier.
Branches with ≥70% support from 1000 rapid bootstraps are shown. Catathelasma imperiale, Lepista
nuda, Mycena a. pura, Panellus stipticus, and Tricholoma avovirens were used as the outgroup.

1158 ... Largent & Bergemann
Aer exclusion of the RPB2 introns and mtSSU hypervariable regions and manual
optimization, alignment lengths were 592 bp (mtSSU), 1498 bp (LSU), and 1078 bp
(RPB2). A Maximum Likelihood (ML) analysis based on a concatenated dataset with
1000 replicates and specifying a GTRGAMMA model using RAXML-HPC2 v. 8.1.11
(Stamatakis 2014) was carried out with a partitioned dataset for each RPB2 codon (three
partitions) and separate partitions for mtSSU and LSU. One thousand bootstrap (BS)
replicates were generated for assessment of branch supports.
Phylogeny
Monophyly of Pouzarella is well supported (BS = 100, Fig. 1), and Pouzarella
alissae clusters with other representatives of P. sect. Versatiles (P. araneosa and
Entoloma subaraneosum) with signicant support (BS = 99, Fig. 1). ese
three species share similar features including a brillose pileus and stipe and
thin-walled hymenial cystidia. Additionally, all three species have dark brown
basidiomata, evanescent silvery-white coverings on stipe and pileus, appressed
brillose pileipelli that become subglabrous, intracellular pigments in the pileus
suprapellis, hyphae with encrusted pigments in the pileus subpellis and trama,
and lageniform to rostrate-ventricose leptocystidioid cheilocystidia. ey
cluster within a larger clade with other members of Pouzarella sect. Pouzarella
(P. nodospora (G.F. Atk.) Mazzer and Entoloma violaceovillosum Manim. &
Noordel.), but without support (Fig. 1). Whether this lack of support for the
morphologically based infrageneric classication is due to a lack of adequate
taxon sampling or gene resolution will require further research.
Taxonomy
Pouzarella alissae Largent & Bergemann, sp. nov. Plates 1, 2
MycoBank MB 812300
Diers from Pouzarella araneosa by its dark brown pileus, shorter basidiospores, shorter
cheilocystidia, and the presence of pleurocystidia.
Type — United States, California, Humboldt County, Prairie Creek Redwoods State
Park, Intersection of Big Tree Trail and Prairie Creek Highway, in du beneath Acer
circinatum and Polystichum munitum, within 20 m of 41°22′33.15″N 124°00′51.66″W,
62 m elev., 22 November 2014, DL Largent 10490 (Holotype, HSC; GenBank KT074359
(mtSSU), KT074357 (LSU), KT074361 (RPB2)).
Etymology — named for Alissa Allen, who rst discovered this species.
Basidiomata with the pileus and stipe initially covered with silvery-whitish
evanescent layer of brils that forms squamules over the pileus surface, small
pointed squamules on the pileus margin, and longitudinally arranged appressed
brils on the stipe. When fresh, the stipe brils can be rubbed o revealing a
dark brownish brillose layer on the stipe surface. In older basidiomata, this
layer rst disappears from the pileus before eventually disappearing on the

Pouzarella alissae sp. nov. (U.S.A.) ... 1159
Plate 1. Pouzarella alissae (HSC DLL10490, holotype): A. basidiomata (Photo by Noah Segal);
B. cross section through strigose stipe base; C. longitudinal section through strigose stipe base.
Scale bars: A = 8 mm; B, C = 20 µm.
stipe. In silica gel-dried basidiomata, this layer is absent when sections of the
basidiomata are viewed under the compound microscope.
Pileus 8–30 mm broad, 2–7 mm high, conic to campanulate, becoming
convex to broadly convex, then plane, and nally uplied; initially lacking an
umbo or obscurely umbonate, broadly umbonate when mature; opaque (not

1160 ... Largent & Bergemann
translucent and thus not translucent-striate), dull, scurfy or tomentulose on disc,
innately brillose elsewhere; margin incurved then decurved and eventually
plane; color overall dark brownish grey (4–5F2–3) fading to dark brown
(5F3–4); esh ≤1.5 mm above the stipe, pale greyish brown. Lamellae 6–12
mm long, 1.25–4 mm deep, initially adnexed and sigmoid and later narrowly to
broadly adnate (at times with a small subdecurrent tooth), subdistant to distant,
moderately broad, pale greyish-tan when young, maturing dark greyish-brown
(5F2) with smoky greyish brown (5–7F5) spots and blotches, when fully mature
with a pinkish cast from spores; edge fairly thick, at rst concolorous and
then developing whitish areas with spore maturation; lamellulae 3–4 between
lamellae in two series [one always long (4–8 mm) and one always short
(1–2.5 mm)]. Stipe 33–88 mm long, apex 1–2.5 mm diam. and base 1.25–4.5
mm diam., cylindrical, equal when young and enlarged at the base when
mature, very nely pruinose at the apex and innately and appressed-brillose
elsewhere, in very scattered places the brils separating and becoming matted
in clumps; hairs or setae absent; basal area decidedly strigose up to 7–8 mm
from substrate, brownish with reddish tones; basal tomentum also present,
whitish (in some basidiomata developing orangish areas). Taste and Odor
not distinctive. Spore deposit pinkish to dingy salmon-tan.
Basidiospores with 5–6 distinct angles in side and prole views,
heterodiametric in side, prole, dorsal, and ventral views, 9–12 × 6–8 µm
(m = 10.2 ± 0.8 × 7.2 ± 0.5 µm; E = 1.2–1.8; Q = 1.4 ± 0.1; n/1 = 40). Basidia
cylindro-clavate,
34–47 × 8–13 µm (m = 41.2 ± 3.5 × 9.8 ± 1.5 µm; E = 3.2–6.0;
Q = 4.3 ± 0.7; n/1 = 10), 4-sterigmate; aborted basidia abundant on the lamellar
edge and face, wall thickening at rst aer which the entire contents become
reddish brown in 10% NH4OH or 3% KOH, cylindro-clavate, 21–39 × 4–11 µm
(m = 29.9 ± 4.2 × 6.5 ± 1.6 µm; E = 3.1–8.3; Q = 4.8 ± 1.3; n/1 = 20). Lamellar
edge heterogeneous. Cheilocystidia (leptocystidia) thin-walled, when
present scattered (but oen absent on some lamellae), colorless or staining
reddish brown in alkali solutions, versiform (cylindro-clavate, clavate,
obclavate), 34–62 × 10.5–16.4 µm or lageniform with a short to medium neck
or rostrate-ventricose, 46–656 × 13–23.5 µm (including the neck or rostrum),
rostrum typically short to medium (rarely longer), 3.1–7.0(–17.5) × 3.0–3.5
(–6.0) µm (n/1 = 4)). Pleurocystidia colorless, uncommon and very
scattered, obclavate, acuminate-ventricose, lageniform, or rostrate-ventricose,
40–42.5 × 12–17 µm (including rostrum, which is 6.4–7.0 µm long and 2.7–7.0
µm diam.). Lamellar tramal hyphae subparallel, 125–564 × 9.2–42 µm
(E = 9.2–19.1; n/1 = 10). Pileipellis composed of entangled hyphae, erect and
resembling a trichodermium on the disc, semi-erect around the disc, becoming

Pouzarella alissae sp. nov. (U.S.A.) ... 1161
Plate 2. Pouzarella alissae (HSC DLL10490, holotype): A. pileipellis radial section from disc;
B. stipitipellis with caulocystidium (longitudinal section from mid-stipe); C. basidium (white
arrow), aborted basidium lled with dark pigment (black solid arrow), and cheilocystidia (dotted
arrows); D. basidium (white arrow) vs. aborted basidium (dotted arrow); E. basidiospores.
Scale bars: A, B = 12 µm; C = 15 µm; D = 10 µm; E = 7 µm.

1162 ... Largent & Bergemann
prostrate towards the margin; pileocystidia (terminal cells) on or around the
disc cylindro-clavate with a rounded apex 93–184 × 9–18 µm (m = 131.9 ± 31.1
× 12.6 ± 2.4 µm; E = 8.6–14.3; Q = 10.6 ± 2.1; n/1 = 11), towards and at the
margin/edge longer and tapered, 166–218 × 14–17 µm. Pileal tramal
hyphae subparallel and entangled, 49–310 × 3–30.5 µm (m = 152.2 ± 71.6
× 13.7 ± 9.4 µm; E = 4.9–46.3; Q = 15.9 ± 12.3; n/1 = 13). Stipitipellis at
the apex composed of hymenial elements (= pruinose), below the apex hyphae
primarily prostrate, 2–4 cells deep but occasionally out-turned in some
areas; caulocystidia (terminal cells) cylindro-clavate and similar in shape
to the pileocystidia, 67–136 × 9–14 µm (m = 89.4 ± 24.4 × 11.2 ± 1.9 µm;
E = 5.8–12.2; Q = 8.2 ± 2.5; n/1 = 6). Stipe tramal hyphae nearly parallel,
somewhat entangled, 9–67 µm diam. (n/1 = 7). Oleiferous hyphae rare in
the pileal trama. Brilliant granules scattered to somewhat abundant in
the basidia. Lipoid globules absent. Pigments brownish, cytoplasmic and
parietal on the apical 1–4 cells in the pileus and stipe suprapelli, encrusted as
rings in the pileus and stipe subpelli, and decidedly encrusted on all tramal
hyphae. Clamp connections absent.
Ecology & distribution — Solitary, in clumps or loose clusters in wet
du under Acer circinatum Pursh and Polystichum munitum (Kaulf.) C. Presl in
Prairie Creek Redwoods State Park.
Comments — Morphologically, P. alissae, P. araneosa, P. celata Mazzer,
P. versatilis, and Entoloma subaraneosum Xiao L. He & T.H. Li are placed in
Pouzarella sect. Versatiles Mazzer because of their innately brillose pileus
and stipe, thin-walled hymenial cystidia, and basidiospores that average <13
µm in length. Pouzarella araneosa and E. subaraneosum can be dierentiated
from P. alissae by their longer spores (9.5–15 µm for P. araneosa; 9.5–13 µm for
E. subaraneosum), longer cheilocystidia (45–100 µm) with much longer necks,
and lack of pleurocystidia (He et al. 2013; Mazzer 1976; Noordeloos 1992,
2004).
In North America, P. celata, described from Michigan, and P. versatilis,
described from Europe (Noordeloos 1979, 1992, 2004; as Entoloma versatile
(Gillet) M.M. Moser) and reported from western North America including
California (Mazzer 1976) and Washington (Largent 1994), are similar in
size and also produce dark brown colored basidiomata. Both species can be
distinguished from P. a l i s s a e by their abundant and longer cheilocystidia and
pleurocystidia and absence of silvery white evanescent brils and squamules on
the pileus. Pouzarella celata also diers by its longer (11–14 µm) basidiospores
and glabrous pileus surface that is streaked by innate silky grey brils while
P. versatilis is distinguished by its abundant pleurocystidia, olivaceous green

Pouzarella alissae sp. nov. (U.S.A.) ... 1163
tints or olivaceous brown pileus color (Noordeloos 2004, Largent 1994), and
the metallic luster of the dried pileus (Mazzer 1976).
Pouzarella araneosa, reported from California under Acer circinatum,
consisted of one basidiome with a missing stipe base (Largent 1994). e
collection diers from P. alissae and P. araneosa by the absence of a silvery
brillose layer, its medium dark brown (5–6F4) pileus, its translucent-striate
pileal margin, and its broader and longer cheilocystidia; until more material
is obtained, the diagnosis of the collection formerly reported as P. araneosa
remains doubtful.
Acknowledgments
Materials required to complete this manuscript were supported by the Largent
family trust, and we are particularly grateful for the support of Pamela Largent. e
DNA sequences generated in this study are based upon work supported by the National
Science Foundation under Grant No. DRI 0922922 awarded to Dr. Sarah Bergemann.
Comments by the two reviewers, Dr. Timothy J. Baroni and Dr. Joseph F. Ammirati,
and by the Nomenclature Editor, Dr. Shaun Pennycook, were also helpful. We wish to
thank especially Noah Siegel for allowing us to use his illustration of P. alissae in the eld
and to see his description of the species in his forthcoming book.
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