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ISSN (print) 0093-4666 © 2014. Mycotaxon, Ltd. ISSN (online) 2154-8889
MYCOTAXON
http://dx.doi.org/./.
Volume 129(1), pp. 119–148 July–September 2014
The Entolomataceae of the Pakaraima Mountains of Guyana 6:
ten new species and a new combination in Nolanea
T W. H*, M. C A,
D L. L, T J. B
1Department of Biological Sciences, Humboldt State University, Arcata, California, USA. 95521
2Department of Botany & Plant Pathology, Purdue University,
West Lafayette, Indiana, USA. 47907
3Department of Biological Sciences, State University of New York—College at Cortland,
New York, USA. 13045
*C : twh5@humboldt.edu
A — Eleven Nolanea species are described from Guyanese collections. e
West African N. acuta comb. nov. is reported for the rst time from the Neotropics, and
N. alboproxima, N. applanata, N. clavata, N. claviformis, N. concentrica, N. furcata, N. mimiae,
N. rava, N. sinuolata, and N. subsulcata are new to science. All of these species occur in
tropical rainforests, either in Upper Potaro River Basin in Guyana’s Pakaraima Mountains
or in the interior lowlands, and are presumed forest litter saprotrophs. Macromorphological,
micromorphological, and habitat data and comparisons with similar taxa are provided for
each. e genus Nolanea has not been previously reported from Guyana.
K — Agaricales, Agaricomycetes, Guayana Highlands, Guiana Shield, saprotrophic
fungi
Introduction
Species in the cosmopolitan genus Nolanea (Fr.) P. Kumm. are easily
recognized as members of the Entolomataceae (Agaricales) due to their pink-
pigmented basidiospores that are angular in all views. Fries (1821) erected
Agaricus “tribus” Nolanea for pink-spored agarics with a mycenoid stature,
and his concept, with renements based primarily in micromorphological
characters, has been accepted by many later agaricologists at either generic (e.g.
Largent 1994; Largent & Baroni 1988; Dennis 1953, 1970; Pegler 1983, 1997) or
subgeneric (Noordeloos 1980, 1992, 2004) rank.
Here we apply the generic concept of Nolanea sensu Largent (1994)
diagnosed by mycenoid or collybioid (rarely tricholomatoid) basidioma stature;
120 ... Henkel & al.
typically glabrous (rarely innately appressed brillose) pileus; the pileipellis
typically a cutis with 1–4 uninated hyphal layers overlying the more inated
hyphae of the outer pileal trama; long-celled hyphal elements typically present
in the pileal, lamellar, and stipe tramas; lipoid globules, brilliant granules, and
pseudocystidia absent; clamp connections typically absent (occasionally rare
or very scarce in the pileipellis); and a +5 urea concentration. Morphologically
similar Inocephalus species can be most readily distinguished from Nolanea by
a pileipellis that intergrades imperceptibly into the pileal trama, a 0 or 0.5 urea
concentration, and (for most Inocephalus species) the combination of abundant
lipoid bodies in the tramal hyphae, pseudocystidia in the hymenium, clamp
connections in the pileipellis, and a distinctly brillose or squamulose pileus
(Largent 1994).
e few molecular systematics studies published on Entolomataceae have
proved rather equivocal regarding the status of Nolanea at the generic or
subgeneric levels (e.g., Co-David et al. 2009; Baroni & Matheny 2011; He et
al. 2013). Other preliminary molecular data support entolomatoid fungi with
No l a n e a’s morphological diagnostic characters as a genus (Bergemann &
Largent unpubl.). Given the current lack of clear, molecular-based resolution
of genera within the Entolomataceae, we have used morphology to distinguish
and describe the new taxa presented here.
New World tropical and subtropical entolomatoid species meeting the
diagnostic requirements of Nolanea sensu Largent (1994) have been found
in the Lesser and Greater Antilles (Murrill 1911; Pegler 1983), Trinidad and
Venezuela (Dennis 1953, 1970), Brazil (Pegler 1997), and elsewhere in South
America (Horak 1977).
We report here eleven Nolanea species for the rst time from Guyana.
One, Nolanea acuta comb. nov., was previously known from Gabon in West
Africa; the other ten species are described as new to science: N. alboproxima,
N. applanata, N. clavata, N. claviformis, N. concentrica, N. furcata, N. mimiae,
N. rava, N. sinuolata, and N. subsulcata.
Materials & methods
Specimens were collected from the Upper Potaro River Basin during the May–
August rainy seasons of 2000, 2003, 2005, 2007, 2009, 2010, and 2011 and the December
rainy seasons of 2004 and 2009. e collecting area is located within a 15 km radius
of a permanent base camp at 5°18′04.8″N 59°54′40.4″W, elevation 710–750 m, in
an undulating valley approximately 15 km east of Mt. Ayanganna (2200 m) and is
densely forested with a mosaic of primary Dicymbe-dominated and mixed forests of
the Eschweilera–Licania association (Henkel 2003). Additional collections were made
during the May–June 2011 rainy season from Guyana’s Upper Demerara River Basin
at Mabura Ecological Reserve within 2 km of a eld station located at 5°09′19.0″N
58°41′58.9″W, elevation 100 m, in monodominant stands of Dicymbe altsonii.
Nolanea spp. nov. (Guyana) ... 121
Methods for eld descriptions, microscopic analyses, and image capture follow
Largent et al. (2008). e formula for Amman’s solution is cited in Largent et al. (1977).
Fungi were eld-dried with silica gel. Color designations follow Kornerup & Wanscher
(1978) with color plates noted in parentheses (e.g., 4A7). Specimens were deposited
in the following herbaria: BRG, HSU, and PUL (Holmgren et al. 1990). Microscopic
structures were measured as previously described (Largent 1994; Largent et al. 2008).
Statistics determined include means of basidiospore length and width ± standard
deviations; E = quotient of length by width indicated as a range variation in n objects
measured; Q = the mean of E-values; n = number of objects measured.
Taxonomy
Nolanea acuta (Romagn. & Gilles) Largent, comb. nov. P
M MB
≡ Rhodophyllus acutus Romagn. & Gilles, Beih. zur Nova Hedwigia 59: 542 (1979)
P 21–32 mm broad, broadly convex with acute umbo; umbo and part
of surrounding disc dark yellowish brown (5F8) because of appressed radial
brils, background color lighter, maturing overall to light blond (4C3), non-
hygrophanous; margin slightly inturned, somewhat rimose-frayed, not striate
or translucent-striate. L subclose, subthick, very nely adnexed,
becoming pinkish from spores (6B4); lamellulae 3 between lamellae, in 2 tiers.
S 70–85 × 3 mm wide at apex, 6 mm at base, tapered upward, whitish with
appressed brown longitudinal brils. B scant, white, appressed
fe lty. S white, stued. O and none.
B distinctly 5–7-angled in prole, dorsal and ventral views,
heterodiametric in all views, 7.8–11.3 × 5.2–7.5 µm (mean = 9.7 ± 0.83 × 6.7 ± 0.59
µm;
E = 1.3–1.8, Q = 1.43 ± 0.09; n = 30); hilar appendage distinct. B
clavate, slightly to strongly tapered at the base, 27.7–33.1 × 7–10 µm (mean =
30.2 ± 1.8 × 8.6 ± 0.88 µm; E = 3.0–4.4, Q = 3.53 ± 0.30; n = 11), 4-sterigmate.
C absent. L of long-
celled, parallel to subparallel hyphae (not measured). P a cutis of
uninated hyphae that are 3.2–10.7 µm wide and rarely uplied;
cylindric to cylindro-clavate, 33.6–73.8 µm long. P
inated beneath the pileipellis, 5–20 µm wide, elsewhere 5–40 µm
wide. S a cutis; hymenial clusters and caulocystidia absent.
P uniform, light to medium brown in the cytoplasm of the
pileipellis; incrusted hyphae absent. R abundant in the
pileus trama. absent. M in
Amman’s solution: hymenial elements strongly cyanophilic.
E, , — Scattered on soil in Dicymbe corymbosa
forest in the Upper Potaro River Basin of Guyana; originally described from
Gabon.
122 ... Henkel & al.
P . Nolanea acuta (Aime 1314). A. Basidia. B. Basidiospores. C. Pileipellis. Bars = 10 µm.
S . GUYANA. R : P-S. Pakaraima Mountains.
Upper Potaro River Basin, ~15 km east of Mt. Ayanganna, environs of base camp
located on Potaro River one km upstream from conuence with Whitewater Creek at
5°18′04.8″N 59°54′40.4″W, elevation 710–750 m; vicinity of base camp, on soil, 21 June
2000, Aime 1314 (BRG; PUL).
C — Nolanea acuta is distinguished by its combination of relatively
large basidiomata, a dark yellowish brown acutely umbonate pileus, clavate
stipe with brownish longitudinal brils on a whitish ground, heterodiametric
basidiospores averaging 9.7 × 6.7 µm (Q = 1.4), cytoplasmic pigmentation,
and lack of hymenial cystidia. Rhodophyllus acutus from Gabon (Romagnesi &
Gilles 1979) is morphologically identical to Guyanese N. acuta, justifying the
new combination. e discovery of N. acuta in Guyana constitutes a signicant
intercontinental range extension for the species.
Nolanea spp. nov. (Guyana) ... 123
Nolanea acuta is similar to N. solstitialis (Fr.) P.D. Orton [≡ Entoloma
solstitiale (Fr.) Noordel.] in pileus and stipe size, shape, and colors, basidiospore
shape and size, and lack of cheilocystidia, but N. solstitialis can be distinguished
by its translucent-striate hygrophanous pileus and smooth glabrous stipe
(Noordeloos 1980, 1992, 2004).
Because of its basidioma colors, acutely umbonate pileus, and lack of
cheilocystidia, Nolanea acuta resembles members of a species complex centered
around N. papillata Bres., including N. pseudopapillata Pegler [= N. papillata
sensu Dennis. non Bresadola], N. mammifer (Romagn.) Pegler [≡ Rhodophyllus
mammifer Romagn.], and N. apiculata Petch [≡ Entoloma acuminatum E. Horak].
All these species dier from N. acuta primarily by their externally encrusting
pileipellis pigments (Dennis 1970; Horak 1980; Pegler 1983).
Nolanea alboproxima Largent, Aime & T.W. Henkel, sp. nov. P
M MB
Diers from Nolanea proxima by its whitish overall coloration, heterodiametric
basidiospores and lack of incrusting pigment in the pileus tramal hyphae.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~20 km east of Mt. Ayanganna, Tadang Base Camp 2 km south of Potaro River
at 5°16′14.5″N 59°50′39.1″W, elevation 710–750 m; vicinity of base camp, 21 December
2009, Henkel 9116 (BRG, holotype; HSU, isotype).
E: albus (L. adj.) = white; referring to the whitish pileus color that dierentiates
N. alboproxima from N. proxima.
P 6–16.5 mm broad, 3–4.5 mm high, broadly convex to plano-convex to
plane, somewhat attened across apex and subumbilicate, glabrous, translucent
striate from disc to margin, overall pale orangish white (4A2–4A3) to greyish
orange (4B4–4B5) with age, moist; disc hygrophanous with medium grey cast;
margin irregularly crenulate. L subclose, subthin, adnexed to barely
adnate, 3–5 mm long, 0.5–2.0 mm tall, o-white at rst (4A1–4A2), developing
slight pinkish overtones (~5A3) with age; margins concolorous, nearly smooth
under hand lens; lamellulae consistently 3 between lamellae, 0.5–2 mm long,
in 2 tiers. S 25–53 × 1–2 mm at apex, 2–4 mm at base, tapering upward,
translucent, o-white (4A1–4A2) to pale light yellow (4A4), slightly yellowing
(3A3–3A4) where handled, glabrous, hollow. B scant, white,
and hispid at extreme base. O none; not noted.
B distinctly 5–6(–7)-angled and heterodiametric in prole
view, 7.7–10.8 × 5.3–7.8 µm (mean = 9.3 ± 0.7 × 6.8 ± 0.57 µm; E = 1.2–1.7,
Q = 1.38; n = 58). B clavate and tapering to a long, slender base, 30.0–
42.8
× 7.1–13.1 µm (mean = 35.0 ± 4.0 × 9.3 ± 1.5 µm; E = 2.3–5.2, Q = 3.9 ±
0.86;
n = 24), 2–4-sterigmate, abundant on the lamella edge. C
and absent. L subparallel, relatively narrow;
composed of long-celled hyphae, these 39.5–186.1 × 11.1–16.8 µm, poorly
124 ... Henkel & al.
P . Nolanea alboproxima (holotype; Henkel 9116). A. Basidiomata. B. Pileipellis. C. Basidia.
D. Basidiospores. Bars: A = 10 mm; B–D = 10 µm.
separating. P a shallow cutis of 1–3 uninated hyphal layers; hyphae
4–6 µm wide; cylindric to cylindro-clavate, 45.9–180.9 × 4.2–6.7
µm. P inated beneath the pileipellis, 10–14 µm wide,
elsewhere 96.8–273.4 × 8.6–24.8 µm. S a cutis;
absent. R abundant in the stipe trama, rare in the pileus
trama, absent in the lamellar trama. P minimal, apparently
cytoplasmic, and not incrusting in the pileipellis. C absent.
E, , — Scattered on humus or seed husks
in Dicymbe spp. forests; known only from the Upper Potaro River Basin of
Guyana during the December–January rainy season.
A . GUYANA. R : P-S.
Pakaraima Mountains. Upper Potaro River Basin, ~15 km east of Mt. Ayanganna,
environs of base camp located on Potaro River one km upstream from conuence with
Nolanea spp. nov. (Guyana) ... 125
Whitewater Creek at 5°18′04.8″N 59°54′40.4″W, elevation 710–750 m; on buried seed
husk, 2 January 2004, Aime 2416 (BRG; PUL).
C — Nolanea alboproxima is distinguished by its small, plano-
convex, pale yellowish white, translucent-striate pileus, pale yellow stipe,
heterodiametric spores averaging 9.3 × 6.8 µm, and its lack of hymenial cystidia.
Even though the basidioma colors are reminiscent of those predominant in
Alboleptonia, N. alboproxima ts well in Nolanea because it combines a thin
pileal cutis, inated cells in the outermost pileal trama, and lack of clamp
connections. Its overall aspect resembles an o-white form of N. proxima
Largent or N. cetrata (Fr.) P. Kumm. Isodiametric basidiospores and externally
encrusted pileus tramal hyphae distinguish N. proxima from N. alboproxima.
Nolanea cetrata has similarly shaped and sized basidiospores but can be
separated from N. alboproxima by its yellowish brown pileus, silvery striate
stipe, bisterigmate basidia, and distinctive cytoplasmic or vacuolar pigment
(Largent 1994).
Several other species representing other entolomatoid genera share similar
basidioma color, basidiospore size, lack of odor, and lack of clamp connections
with Nolanea alboproxima. Of these, Entoloma purum E. Horak & Desjardin
from Hawaii is distinguished by its suede-like to pubescent pileus, trichodermial
pileipellis, and much smaller basidiomata (pileus 4–10 mm broad, stipe 3–5 ×
0.5–0.7 mm; Horak & Desjardin 1993). Entoloma proprium E. Horak from New
Caledonia can be separated by its mbriate lamellae, distinctive cheilocystidia,
incrusting pigments, and abundant refractive hyphae (Horak 1980).
Rhodophyllus vetulus Romagn., R. tortilis Romagn., and R. subsericellus
Pat. [≡ R. platypus Romagn, nom. illegit] from Madagascar and R. exuosipes
Romagn. & Gilles from Gabon resemble N. alboproxima. Rhodophyllus vetulus,
although with a small pileus, resembles an Entoloma s. str. as it lacks inated
hyphae in the outer pileus trama and the stipe is >4 mm broad at the apex.
Rhodophyllus tortilis is distinctive because of its leathery, somewhat scabrous
pileus lacking striations, and overall lack of yellow tones in the basidiomata.
Rhodophyllus subsericellus has a nely pilose, opaque pileus without striations
(Romagnesi 1941, as R. platypus). Rhodophyllus exuosipes is distinguished
by its ≤70 mm long exuous stipe and ≤35 mm broad campanulate pileus
(Romagnesi & Gilles 1979).
Nolanea applanata Largent & T.W. Henkel, sp. nov. P
M MB
Diers from Entoloma bakeri by its brownish orange, translucent-striate pileus and
hollow stipe.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~15 km east of Mt. Ayanganna, environs of base camp located on Potaro River
126 ... Henkel & al.
one km upstream from conuence with Whitewater Creek at 5°18′04.8″N 59°54′40.4″W,
elevation 710–750 m; 17 August 2007, Henkel 8872 (BRG, holotype; HSU, isotype).
E: applanatus (L. adj.) = at, referring to the pileus shape.
P 45–55 mm broad, planate to upturned, with a low umbo, brownish
orange (5C5) throughout, hygrophanous, becoming lighter rst over disc and
then outwards along radial striations, translucent-striate throughout except
over the disc; surface appearing glabrous and moist throughout, under hand
lens with extremely ne appressed radial brils; margin crenulate, irregularly
splitting 1 or 2 times. L subdistant, subthick, abruptly adnexed,
20.5–24 mm long, 4–4.8 mm tall, pinkish tan (4A4–5A4); margins concolorous,
nely and irregularly eroded under hand lens; lamellulae 1–2 between lamellae,
1–4 mm long. S 62–97 × 4–7 mm (centrally), equal over apical ½, enlarging
evenly over lower ½ to 9–11 mm wide at base, concolorous with pileus, glabrous
to nely appressed brillose under hand lens over upper half, cartilaginous and
twisting, with a hollow core. B scant, white, of matted hyphae.
O none; minimal, slightly mealy.
B distinctly 6–7(–8)-angled with the apex rounded and
consistently heterodiametric in prole and dorsiventral views, 9.9–12.6 ×
6.2–8.2 µm, (mean = 10.9 ± 0.6 µm × 7.1 ± 0.5 µm; E = 1.4–1.8, Q = 1.5 ± 0.1;
n = 26). B clavate, 36.3–47.3 × 6.5–12.2 µm, (mean = 43.8 ± 2.9 µm ×
9.7 ± 1.3 µm, E = 3.5–4.9(–6.4), Q = 4.58 ± 0.7; n = 13), 2- or 4-sterigmate.
L nearly parallel; hyphal cells very long, 149.4–542.8 ×
9.0–18.4 µm; subhymenium narrow. C
absent. P a cutis, 1–4 hyphal layers thick; hyphae 2–8 µm wide. P
inated beneath the pileipellis, 8–18 µm wide, elsewhere very
long-celled, 205–807 × 16.7–34.8 µm. S a cutis; hymenial clusters
and caulocystidia absent. R abundant immediately beneath
the pileipellis, present but indistinct in the subhymenium, rare in the lamellar
and stipe tramas. P cytoplasmic, not incrusting in the pileipellis.
C absent.
E, , — Occurring in a pair on root mat in
Dicymbe corymbosa forest on sandy soils; known only from the type locality in
the Upper Potaro River Basin of Guyana.
C — Nolanea applanata is distinguished by its dull esh-tan to
brownish orange, glabrous, hygrophanous, planate to upturned ≤55 mm
broad pileus that is translucent-striate from disc to margin, concolorous stipe
that is twisted, tapering upward, 60–97 × 4–7 mm (apex) to 9–11 mm (base),
6–8-angled basidiospores that average 10.9 × 7.1 µm, and absence of hymenial
cystidia. Despite the rather large basidiomata, N. applanata ts well in Nolanea
with its thin cuticular pileipellis, distinctly inated outer pileal trama hyphae,
Nolanea spp. nov. (Guyana) ... 127
P . Nolanea applanata (holotype; Henkel 8872). A. Basidiomata. B. Pileipellis. C. Hymenium,
basidia, and lamellar trama. D. Basidiospores. Bars: A = 10 mm; B–D = 10 µm.
very long-celled tramal hyphae in the pileus and stipe, as well as the translucent–
striate pileus and twisted-striate, cartilaginous stipe.
While Entoloma bakeri Dennis from Martinique, Guadeloupe, and
Trinidad is similar to N. applanata in basidioma size and stature, basidiospore
dimensions, and absence of clamp connections, it can be distinguished by its
cream–colored, silky-striate, innately brillose pileus and solid stipe (Pegler
1983).
Nolanea clavata Largent & T.W. Henkel, sp. nov. P
M MB 519975
Diers from Entoloma conicoumbonatum by its lack of incrusting pigments on the
hyphae of the pileipellis and pileus trama and basidiospores with a higher length/width
quotient.
128 ... Henkel & al.
P . Nolanea clavata (holotype; Henkel 8870). A. Basidiomata. B. Pileipellis. C. Basidia.
D. Basidiospores. Bars: B = 10 mm; A, C, D = 10 µm.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~15 km east of Mt. Ayanganna, environs of base camp located on Potaro River
one km upstream from conuence with Whitewater Creek at 5°18′04.8″N 59°54′40.4″W,
elevation 710–750 m; vicinity of base camp, 17 August 2007, Henkel 8870 (BRG,
holotype; HSU, isotype).
E: clavatus (L. adj.) = club-shaped, referring to the stipe shape.
P 26–28 mm broad, 16–21 mm high, narrowly to broadly conic with
rounded to subumbonate apex, dull greyish brown to dark brown (6E4–6F4)
Nolanea spp. nov. (Guyana) ... 129
throughout, slightly darker brown over disc (6F5), appressed radially brillose
throughout, brils more compact over disc but hardly erect, somewhat shiny
macroscopically, opaque, not hygrophanous, silky under hand lens, dry to
submoist; margin broadly undulating forming coarse irregular ridges between
wide radial furrows; extreme edge crenulate and lighter greyish brown (~6C3).
L subclose, subthick, deeply adnexed, 11.5–12.0 mm long, 4–5 mm
tall, dull dirty pink; margins concolorous and smooth; lamellulae 1–2 between
lamellae, 1–3 mm long. S 33–50 mm × 3–4 mm (centrally), equal over
apical ½, enlarging evenly over lower ½ to 6–8 mm, forming a subbulbous,
rounded base, light grey brown (6C2–6D2) throughout but lighter over basal
bulb, appearing silky sericeous, smooth but with ne longitudinal striations
under hand lens. B a ne white bloom at extreme base. O
none; strongly farinaceous.
B distinctly 5–6-angled, apex rounded and heterodiametric in
prole and dorsiventral views, 8.7–11.9 × 5.7–8.2 (mean = 10.2 ± 0.7 × 6.9 ± 0.5
µm; E = 1.4–1.7, Q = 1.5 ± 0.1; n = 30). B (2–)4-sterigmate, short clavate,
tapering basally, 31.1–41.0 × 8.5–12.3 µm (mean = 34.9 ± 2.9 µm × 10.6 ± 1.1
µm; E = 2.9–3.9, Q = 3.3 ± 0.2; n = 14). C and
absent. L subparallel and broad; cells 72.2–166.6 ×
11.2–19.9 µm, E = 4.7–11.2. P a cutis, 1–4 hyphal layers thick; hyphae
1.5–6 µm wide; upturned hyphal ends rare to absent. P
slightly interwoven, inated beneath the pileipellis, 15–20 µm wide, elsewhere
53.6–132.7 × 15.4–32.4 µm (E = 1.9–5.0). S a cutis; hymenial
clusters and caulocystidia absent. S long and broad,
104.8–412.2 × 13.5–24.6 µm, (E = 4.3–30.5). P not observed in
water mounts, not obvious in 3% KOH, likely cytoplasmic and uniform in the
suprapellis of the pileus; not obvious in the stipitipellis. R
in radial section abundant in the pileal trama near the subhymenium; rare to
absent elsewhere. C absent.
E, , — Solitary on humic layer of forest oor
under Dicymbe corymbosa; known only from the type locality in the Upper
Potaro River Basin of Guyana.
C — Nolanea clavata is distinguished by its relatively large basidioma
with conical, subumbonate, grey brown to dark brown, radially appressed-
brillose pileus, clavate, light grey brown stipe, strongly farinaceous taste,
absence of hymenial cystidia, and heterodiametric, 5–6-angled basidiospores
averaging 10.2 × 6.9 µm. e radially appressed–brillose, tall, conic pileus and
clavate stipe of N. clavata are reminiscent of species of the entolomatoid genus
Inocephalus. However, N. clavata lacks lipoid globules and clamp connections,
features typical of Inocephalus species, and has a thin pileipellis of repent hyphae
overlying distinctly inated tramal hyphae, features diagnostic for Nolanea.
130 ... Henkel & al.
Because of the similar pileus shape and color, basidiospore size, and lack
of cheilocystidia, N. clavata resembles Entoloma substrictior (Singer) E. Horak
from Argentina and Chile and Entoloma cystopus (Berk.) Sacc. from India and
Malaysia. However, E. substrictior has much larger basidiomata with a ≤50 mm
broad strongly striate pileus with a rugose surface (Horak 1977). Entoloma
cystopus is also much larger, with a pileus up to 45 mm broad and stipe up to
140 mm long, and becomes pale livid yellow when dry (Horak 1980). Entoloma
conicoumbonatum Hesler resembles N. clavata in basidioma size and colors and
has similarly shaped basidiospores, but diers in having incrusting pigments
on the hyphae of the pileipellis and pileus trama and basidiospores with an
E = 1.1–1.3 and Q = 1.15 (Hesler 1967; Noordeloos 1988).
Nolanea claviformis Largent & Aime, sp. nov. P
M MB
Diers from all described species of Nolanea by its light brown, concentrically ridged
pileus, orange-white lamellae, dark brown stipe, clavate cheilocystidia, and two types of
pileus pigment.
T: Guyana. Region 10: Upper Demerara-Berbice. Mabura Ecological Reserve: ~100
m northwest of Mabura eld station at 5°09′19.0″N 58°41′58.9″W, elevation 100 m; in
Dicymbe altsonii monodominant stand #1, 23 May 2011, Aime 4279 (BRG, holotype;
PUL, isotype).
E: claviformis (L. adj.) = club-shaped, referring to the clavate cheilocystidia.
P 7–25 mm diam, 7–17 mm high, campanulate-conical, concentrically
ridged with a broad umbo, satiny light brown (5D4), felty, with appressed
longitudinal silky brils under hand lens; margin suggestively striate, crimped,
and felty. L 2 mm tall, narrow, adnate, close to crowded, orange white
(5A2); margins broadly scalloped under hand lens; lamellulae numerous,
dicult to count. S 25–69 × 1.5–2.5 mm, equal, concolorous with pileus,
silky; pruina scant at apex under hand lens. B appressed, white,
extending upward. C thin, hollow in stipe. B none.
O none; none, mealy in texture.
B distinctly 4–5(–6)-angled in all views, isodiametric
in polar view, isodiametric to heterodiametric in prole view, 7.3–10.9 ×
6.8–8.6 µm (mean = 9.0 ± 0.8 × 7.5 ± 0.48 µm, E = 1.0–1.4, Q = 1.20 ± 0.10;
n = 31). B clavate to cylindro-clavate, 28.2–42.5 × 9.5–13.0 µm (mean
= 35.2 ± 4.8 × 10.7 ± 1.2 µm, E = 2.9–3.9, Q = 3.28 ± 0.4; n = 7), 4-sterigmate.
C abundant, forming a sterile layer, clavate, colorless, 41.4–57.6
× 8.7–15.7 µm (E = 4.0–4.8; n = 6). P absent. L
subparallel, cells relatively short and broad, 50.8–130.0 × 9–14.0
µm (n = 4). P a cutis, 4–5 hyphal layers thick; hyphae 2.0–7.5 µm
wide; cylindric to cylindro–clavate, narrow. P
slightly inated beneath the pileipellis; cells 65.9–128.2 × 8.7–14.6 µm,
Nolanea spp. nov. (Guyana) ... 131
P . Nolanea claviformis (holotype, Aime 4279). A. Basidiomata. B. Basidiospores. C. Cheilo-
cystidia. D. Pileipellis. E. Caulocystidia. Bars: A = 10 mm; B–D = 10 µm.
elsewhere up to 20 µm wide. S a cutis between scattered hymenial
clusters at the apex; in hymenial clusters clavate, colorless,
132 ... Henkel & al.
similar to basidia but more narrow, 30.9–43.4 × 4.7–12.2 µm (E = 3.6–6.6,
Q = 5.0; n = 5). S long; cells 72.3–768.4 × 13.9–30.1 µm
(n = 4). R absent. P minutely to distinctly
incrusting the walls of the more slender hyphae in the pileal trama, faintly
cytoplasmic in the pileipellis hyphae. C absent.
E, , — Scattered on litter of forest oor in
Dicymbe altsonii forests on sandy soils; known only from the type locality at
Mabura Ecological Reserve, Guyana.
C — Nolanea claviformis is characterized by its concentrically ridged
pileus, narrow crowded lamellae, and two types of pigment in the pileipellis.
Refer to the comments sections for N. sinuolata and N. concentrica for
discussions of species similar to N. claviformis.
Nolanea concentrica Largent & T.W. Henkel, sp. nov. P
M MB
Diers from Nolanea bicoloripes by its abundant cheilocystidia and much shorter stipe.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, 15–20 km east of Mt. Ayanganna, Tadang Base Camp 2 km south of Potaro River
at 5°16′14.5″N 59°50′39.1″W, elevation 710–750 m; vicinity of base camp, 19 December
2009, Henkel 9106 (BRG, holotype; HSU, isotype).
E: concentricus (L. adj.) = ringed, referring to the concentrically ridged,
umbonate pileus umbo.
P 18 mm broad, 7 mm tall, broadly conic with concentrically ridged umbo,
apex submammilate but rounded, brown (6D5–6D6), slightly darker (6E6) over
disc, shiny sericeous and moist throughout, nely translucent-striate over outer
2/3; margin nely roughened. L subthin, crowded, adnate, 8.3 mm
long, 1.5–2.0 mm tall, pink; margins concolorous, roughened-cystidiate under
hand lens. S 42 mm × 1.5 mm, equal, tan brown (5C4–5C5) throughout,
smooth with ne suberect brils under hand lens. B lacking.
O mildly fungoid; not obtained.
B distinctly 4–5-angled in all views, isodiametric in polar
view, in prole and dorsiventral views isodiametric to heterodiametric, 6.5–9.0
× 5.1–6.9 µm (mean = 7.7 ± 0.64 × 6.0 ± 0.52 µm; E = 1.13–1.61, Q = 1.3 ± 0.13;
n = 28). B cylindro-clavate, 24.2–30.6 × 6.7–9.5 µm (mean = 27.4 ± 2.0 ×
8.1 ± 0.8 µm; E = 3.4 ± 0.47, Q = 2.8–4.2; n = 13), tapering abruptly near base to
1.5–4.6 µm wide, 4-sterigmate; sterigmata 1.9–4.3 µm long. C
abundant, forming a sterile layer, hyaline, broadly aciculate to obclavate at rst,
ultimately long ventricose–rostrate, 34.4–121.9 × 5.8–16.1 µm (mean = 79.3 ×
10.3 µm; E = 7.75, Q = 5.9–9.6; n = 10). P absent. L
subparallel, cells relatively short and broad, 69.8–121.7 ×
10.1–13.7 µm. P a cutis, 4–5 hyphal layers thick; hyphae 2.0–6.2 µm
Nolanea spp. nov. (Guyana) ... 133
P . Nolanea concentrica (holotype; Henkel 9106). A. Basidioma and pilei. B. Basidiospores.
C. Basidium. D. Pileipellis and pileal trama. E. Cheilocystidia. Bars: A = 10 mm; B–D = 10 µm.
134 ... Henkel & al.
wide; cylindric to cylindro-clavate. P
beneath the pileipellis slightly inated, 8.7–14.6 µm wide; cells elsewhere
50.9–128.2 × 6.7–13.8 µm. S not studied. R
absent. P minutely incrusting the more slender hyphae in
the pileal trama, faintly cytoplasmic within the pileipellis hyphae. C
absent.
E, , — Solitary on litter of forest oor in mixed
Dicymbe spp. forest; known only from the type locality in the Upper Potaro
River Basin of Guyana.
C — Nolanea concentrica is distinguished by its dark brown, broadly
conic pileus that is shiny sericeous, nely translucent-striate, and submammilate
with a concentrically ridged umbo, crowded narrow lamellae, smooth equal
stipe, abundant broadly aciculate to ventricose-rostrate cheilocystidia, and
4–5-angled basidiospores averaging 7.7 × 6.0 µm. Nolanea bicoloripes Largent
& iers from western North America is similar to N. concentrica in the
distinctive, concentrically ridged, umbonate, silky-sericeous pileus, narrow
equal stipe, and some micromorphological features, but diers in its longer
(≤140 mm) stipe and lack of cheilocystidia (Largent 1994).
Both N. claviformis and the sympatric N. concentrica have a concentrically
ridged or beveled pileus, white lamellae, 4–5-angled basidiospores, and
incrusting and cytoplasmic pigments in the pileipellis. Nolanea claviformis is
distinguished by its clavate cheilocystidia, larger (~9.0 × 7.5 µm) basidiospores,
and larger (28.3–42 5 × 9.5–12.0 µm) basidia. See the comments under
N. sinuolata for additional information.
Nolanea subsolstitialis Largent from Washington, N. infula (Fr.) Gillet from
the Netherlands and Sweden, and N. solstitialis from Europe also have small
basidiospores, a beveled or concentrically ridged pileus, and a subpellis that
is only slightly inated and lack clamp connections. However, all three species
lack cheilocystidia and have dierent stipe dimensions than N. concentrica. In
addition, N. solstitialis has a distinct, abundant external incrusting pigment,
N. solstitialis has intracellular pigment, and N. infula possesses both pigment
types (Noordeloos 1980; Largent 1994).
Nolanea furcata Largent & T.W. Henkel, sp. nov. P
M MB
Diers from Entoloma infula var. chlorinosum by its non-papillate pileus, faint chlorine
odor, and abundant cheilocystidia.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, 15–20 km east of Mt. Ayanganna, Tadang Base Camp 2 km south of Potaro River
at 5°16′14.5″N 59°50′39.1″W, elevation 710–750 m; 0.5 km SW of Tadang Base Camp,
24 December 2009, Henkel 9139 (BRG, holotype; HSU, isotype).
Nolanea spp. nov. (Guyana) ... 135
P . Nolanea furcata (holotype; Henkel 9139). A. Basidiomata. B. Hymenophore.
C. Basidiospores. D. Basidia. E. Pileipellis and pileus tramal hyphae with external encrustations.
F. Cheilocystidia. Bars: A, B = 10 mm; C–F = 10 µm.
136 ... Henkel & al.
E: furcatus (L. adj.) = forked, referring to the irregularly forked lamellae.
P broadly conic with subacute to rounded apex, 13–23 mm broad, 7–16
mm tall, greyish brown (6E3–E4), hygrophanous to lighter greyish brown
(5D2–D3) over disc, appearing smooth macroscopically; surface under hand
lens a dense, very ne brillose mat, this uniform throughout with no erect
elements evident, not translucent-striate; margin slightly inrolled, otherwise
entire and light grey; moist. L thin, crowded, adnate with a decurrent
tooth, 6.5–15 long, 1–2 mm tall, forking irregularly, white to light pink (6A1–A2);
margins concolorous, highly roughened-cystidiate under hand lens; lamellulae
numerous, too crowded to count. S subequal, tapering very slightly toward
apex, 74–87 mm × 3–4 mm (centrally), base 4–6 mm, concolorous with the
pileus over upper ¾, smooth, nely longitudinally brillose under hand lens.
B over basal ¼ of stipe a dense, white brillose mat. P
light grey. S grey, hollow. B none.
O faintly of detergent or chlorine; pleasantly fungoid.
B distinctly (4–)5-angled and subisodiametric to
heterodiametric in prole view, 6.6–8.4 × 5.1–7.2 µm (mean = 7.5 ± 0.4 ×
6.0 ± 0.5 µm; Q = (1.09–)1.14–1.46, E = 1.26 ± 0.1; n = 25). B clavate,
strongly tapered basally, 27.6–35.6 × 5.8–11.1 µm (mean = 30.8 ± 2.3 × 8.5 ± 1.6
µm; Q = 2.95–5.1, E = 3.8 ± 0.8; n = 12); base 2.7–5.2 µm wide; 4–sterigmate;
sterigmata 1.4–3.2 µm long. C abundant, forming a sterile layer
on the gill edge, uniformly cylindro-clavate, colorless, 47.9–74.3 × 3.7–5.9 µm.
P absent. L subparallel, relatively
slender and uniform in width; cells 54.8–112.2 × 3.7–6.1 µm. P
a cuticular layer of largely repent, entangled, relatively narrow hyphae;
cylindro-clavate, 20.1–49.3 × 4.1–9.3 µm (n = 11). P
beneath the pileipellis slightly to distinctly inated; cells
38.0–134.4 × 6.2–13.8 µm (n = 8), not measured elsewhere. S with
an entangled hyphal layer similar to the pileipellis, with hymenial clusters at the
stipe apex; abundant, somewhat opaque, similarly-shaped as,
but longer than the cheilocystidia, 20–137.8 × 3.2–6.3 µm (n = 7). R
abundant in the pileal trama, absent in the lamellar trama and the
subhymenium. P of two types in the pileipellis: uniformly
cytoplasmic and faintly brownish and with external encrustations; pileus trama
hyphae strongly externally encrusted. C absent.
E, , — Scattered on litter of forest oor in
mixed Dicymbe spp. forest; known only from the type locality in the Upper
Potaro River Basin of Guyana.
C — Nolanea furcata is distinguished by its conic, dark brownish
grey, hygrophanous, opaque pileus, narrow, crowded, irregularly forked
lamellae that are white initially, faintly detergent-like or chlorine-like odor,
Nolanea spp. nov. (Guyana) ... 137
abundant cylindro-clavate cheilocystidia and caulocystidia, cytoplasmic and
externally incrusted pigment in the pileipellis, and relatively small (~7.5 ×
6.0 µm) 4–5-angled basidiospores. Entoloma infula var. chlorinosum (Arnolds
& Noordel.) Noordel. from Europe shares a dark brown pileus and stipe, the
two types of pileipellis pigments, somewhat similarly shaped basidiospores,
and chorine-like odor but fundamentally diers from N. furcata in its
papillate pileus, persistent and much stronger chlorine-like odor, and lack
of cheilocystidia (Noordeloos 1992). For additional information on Nolanea
species similar to N. furcata, refer to the comments for N. sinuolata.
Nolanea mimiae Largent & Aime, sp. nov. P
M MB
Diers from Nolanea mammosa var. venezuelana by its non-papillate pileus and presence
of pleurocystidia.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~15 km east of Mt. Ayanganna, environs of base camp located on Potaro River
one km upstream from conuence with Whitewater Creek at 5°18′04.8″N 59°54′40.4″W,
elevation 710–750 m; up Potaro River from base camp on line to Lance’s plot near
Blackwater Creek, 25 July 2010, Aime 4024 (BRG, holotype; PUL, isotype).
E: In honor of Mimi Chin, long-time Guyana eld assistant and nder of the
type collection.
P 19 mm broad, 11 mm tall, conical, lacking an umbo, glabrous
macroscopically but faintly appressed-brillose under hand lens, opaque, not
hygrophanous, not translucent striate, dark brown (6E7), slightly darker over
the disc; margin crenulate and lighter. L subthin, close, adnate, 13
mm long, 3 mm tall, pale yellow (4A3) initially, later with pink spots; margins
concolorous with face, roughened-cystidiate under hand lens. S 60 mm ×
1.5 mm, equal, dark yellowish brown (5F8), glabrous, inserted; trama hollow.
B lacking. B none. O indistinct;
none at rst, then somewhat soapy.
B distinctly 5-angled, isodiametric to heterodiametric
in prole view, 8.4–10.9 × 6.7–8.8 µm (mean = 9.8 ± 0.6 × 7.8 ± 0.5 µm;
E = 1.1–1.5, Q = 1.26 ± 0.10; n = 34). B cylindro-clavate, 27.6–32.0 ×
10.5–13.5 µm (mean = 30.0 ± 1.5 × 11.7 ± 1.0 µm, E = 2.2–2.9, Q = 2.6 ± 0.3;
n = 6), 4-sterigmate. C scattered to abundant, colorless,
lageniform, 27.4–68.8 × 6.4–18.4 µm (E = 3.6–5.8; n = 9). P
infrequent to scattered, similar in shape to cheilocystidia, 26.4–64.1 × 5.0–14.1
µm
(E = 2.4–10.5; n = 14).
L subparallel; cells 187.4–
480.6 × 8.6–29.5 µm (n = 4). P a cutis, 5–7 hyphal layers thick; hyphae
3.7–8.1 µm wide; cylindro–clavate, 26.9–37.1 × 2.5–7.6 µm
(E = 4.4–10.8; n = 5). P beneath the pileipellis slightly
inated, 5.7–14.3 µm wide; cells elsewhere inated, relatively long and tapered
138 ... Henkel & al.
P . Nolanea mimiae (holotype; Aime 4024). A. Basidioma. B. Basidiospores. C. Cheilocystidia.
D. Pileipellis. E. Cytoplasmic pigments in the pileipellis hyphae. Bars: A = 10 mm; B–E = 10 µm.
at both ends, 53.7–394.1 × 15.3–30.2 µm (n = 7). S at the apex
with rare clusters of basidia, and rare terminal cells, these slightly out-turned,
cylindro-clavate, 30.8–42.6 × 3.4–6.2 µm (n = 4), elsewhere a cutis. S
56.9–377.8 × 10.9–23.9 µm (n = 7). R
glistening in 3% KOH, abundant in the pileal trama, absent in the lamellar
trama; not studied in the stipe trama. P cytoplasmic in the hyphae
Nolanea spp. nov. (Guyana) ... 139
of the pileipellis, in the form of abundant dark brown, plaque-like agglutinated
clots, absent in all other tissues. C absent.
E, , — Solitary on litter of forest oor in mixed
Dicymbe spp. forest; known only from the type locality in the Upper Potaro
River Basin of Guyana.
C — Nolanea mimiae is distinguished by its conical, non-umbonate,
opaque, non-hygrophanous, dark brown pileus, pale yellow lamellae, dark
yellowish brown stipe, scattered lageniform cheilocystidia and pleurocystidia,
cytoplasmic pigment in the pileipellis, and nearly heterodiametric 5-angled
basidiospores averaging 9.8 × 7.8 µm. ese characters, along with the
cytoplasmic pigment in the form of plaques or clots, and the mild then soapy
taste, make N. mimiae unique among previously described Nolanea species.
Nolanea mammosa var. venezuelana Dennis from Venezuela [≡ Rhodophyllus
venezuelanus (Dennis) Singer from Chile] and Entoloma taedium E. Horak
from Chile are somewhat similar to N. mimiae because of their lageniform
cheilocystidia and basidioma colors. Nolanea mammosa var. venezuelana
diers from N. mimiae by its papillate pileus and lack of pleurocystidia (Singer
1969; Dennis 1970). Entoloma taedium diers by its distant pallid lamellae, lack
of pleurocystidia, and cytoplasmic pigment in the cheilocystidia (Horak 1977).
Nolanea rava Largent & Aime, sp. nov. P 9
M MB
Diers from Entoloma acutopallidum by its greyish yellow pileus, 5–6-angled
basidiospores, obclavate to rostrate-ventricose cheilocystidia, and abundant
pleurocystidia.
T: Guyana. Region 10: Upper Demerara-Berbice. Mabura Ecological Reserve: ~100
m northwest of Mabura eld station at 5°9′19.0″N 58°4′58.9″W, elevation 100 m; in
Dicymbe altsonii monodominant stand #1, 27 May 2011, Aime 4312 (BRG, holotype;
PUL, isotype).
E: ravus (L. adj.) = greyish-yellow, referring to the color of the pileus and
stipe.
P 13 mm broad, 10 mm tall, conical with a tiny, nipple-like umbo that is 1
mm long, dry, silky, at rst greyish yellow (4B4), remaining so on the disc and
umbo, elsewhere becoming pale greyish yellow (4B3), hygrophanous, shallowly
sulcate and translucent-striate to disc, glabrous; margin decurved, broadly
undulating in outline. L thin, close, adnexed, 11.5 mm long, 3–5 mm
tall, esh-colored (6B3) with masses of concolorous basidiospores visible under
hand lens; margins entire, concolorous and undulating; lamellulae 3–5 between
lamellae. S 37 × 2 mm at apex, 3 mm base, somewhat attened and tapered
from base to apex, glabrous but obscurely longitudinally striate under hand
lens, greyish yellow (4B4) like the mature pileus disc. B absent.
140 ... Henkel & al.
P . Nolanea rava (holotype; Aime 4312). A. Basidioma. B. Basidiospores. C. Hymenium with
basidia. D. Pileipellis and pileal trama. E. Cheilocystidia. Bars: A = 10 mm; B–D = 10 µm.
C not examined. B yellowish near the pileus
margin. O none; not obtained.
Nolanea spp. nov. (Guyana) ... 141
B distinctly 5–6-angled in all views, isodiametric to
heterodiametric in prole and dorsiventral views, angles at times somewhat
rounded, 9.4–11.9 × 7.6–9.8 µm (mean = 10.5 ± 0.6 × 8.5 ± 0.49 µm; E = 1.0–1.3,
Q = 1.24 ± 0.08; n = 34). B cylindro-clavate, hardly tapered, 29.8–43.0 ×
10.5–13.5 µm (mean = 36.8 ± 3.7 × 11.6 ± 0.9 µm; E = 2.4–3.6, Q = 3.2 ± 0.4; n
= 12); 4–sterigmate; sterigmata up to 5.0 µm long. C abundant,
colorless, obclavate to rostrate-ventricose, base rounded, 26.7–62.0 × 9.0–26.8
µm (mean = 42.7 ± 11.9 × 15.3 ± 6.1 µm; E = 1.9–4.9, Q = 3.0 ± 1.1; n = 8);
rostrum 8.2–35.9 µm long × 4.3–11.6 µm wide at the base, 2.0–5.2 µm wide
at the apex. P abundant, similar in shape but longer than the
cheilocystidia, colorless, 26.5–88.8 × 12.5–25.2 µm (mean = 49.8 ± 19.1 × 18.5
± 4.2 µm, E = 1.6–3.4, Q = 2.6 ± 0.6; n = 11). L long
and tapered at both ends; cells 53.4–498.7 × 8.3–25.4 µm (n = 9). P
a cutis, 2–3 hyphal layers thick; hyphae 4.0–7.5 µm wide;
cylindric and at times somewhat swollen at the apex. P
slightly inated beneath the pileipellis, 10.3–14.3 µm wide; cells overall 44.2–
270.5 × 3.4–14.3 µm (n = 6). S a cutis; hymenial elements and
distinct terminal cells absent. S not studied. R
absent. P minimal, cytoplasmic in the pileipellis and
stipitipellis. C absent.
E, , — Solitary on litter of forest oor in
Dicymbe altsonii forests on sandy soils; known only from the type locality at
Mabura Ecological Reserve, Guyana.
C — Nolanea rava is distinguished by its conical, acutely umbonate,
silky, hygrophanous, translucent-striate greyish yellow pileus and concolorous
stipe, nearly obclavate to rostrate-ventricose cheilocystidia and pleurocystidia,
nearly heterodiametric 5–6-angled basidiospores averaging 10.5 × 8.5 µm,
and general lack of microscopic pigmentation. is combination of characters
makes N. rava unique among previously described Nolanea species.
e overall basidioma aspect of Nolanea acuta resembles that of Entoloma
acutopallidum E. Horak & Cheype from French Guiana (Horak & Cheype
2008). Entoloma acutopallidum diers from N. rava by its orangish clay-colored
pileus, white stipe, cuboid basidiospores, clavate to vesiculose cheilocystidia,
and lack of pleurocystidia.
Nolanea sinuolata Largent, Aime & T.W. Henkel, sp. nov. P
M MB
Diers from all described species of Nolanea by its nearly concolorous dark brown
pileus, lamellae, and stipe, cylindro-clavate cheilocystidia and caulocystidia that are
slightly wavy to sinuous in outline, and externally encrusted pigment on the hyphal
walls of the pileipellis and lamellar trama.
142 ... Henkel & al.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~15 km east of Mt. Ayanganna, environs of base camp located on Potaro River
one km upstream from conuence with Whitewater Creek at 5°18′04.8″N 59°54′40.4″W,
elevation 710–750 m; 1 km NE of base camp, sandy zone E of Benny’s Ridge, 29 August
2009, Henkel 8900 (BRG, holotype; HSU, isotype).
E: sinuolatus (L. adj.) = faintly waved, referring to the cheilocystidia that are
faintly wavy or sinuous in outline.
P 11–24 mm broad, 5–12 mm tall, broadly conic with broadly rounded
and attened umbo, initially dark brown (6F4–6F5) throughout, then lighter
concolorous (6E6 or 5E6) on the umbo and paling to light brown (5D6) toward
the margin, glabrous macroscopically, under hand lens a contiguous mat with
minute suberect elements over disc, moist, outer 1–2 mm of margin with
faint translucent striations; marginal edge decurved, entire in areas, elsewhere
very nely crenulate. L subthin, subcrowded, adnate, brown at rst
(concolorous with the pileus), then with pinkish overtones (5C4), nally dirty
dull pink (6A2–6B2) with basidiospore maturation; margins whitish, densely
cystidiate under hand lens; lamellulae 1–2 between lamellae, 0.5–3 mm long.
S 50–71 × 1.5–1.75 mm, equal, glabrous macroscopically, very nely
twisting longitudinally striate under hand lens with a whitish basal bloom,
otherwise dark yellowish brown (5F8) becoming dark brown (6F5–6F6)
throughout, inserted, cartilaginous, hollow. B whitish and
scant. O none or faintly spermatic; not obtained.
B distinctly 5–6-angled in all views, in prole view
isodiametric to heterodiametric, 6.7–9.5 × 5.3–7.9 µm (mean = 8.1 ± 0.6 ×
6.8 ± 0.5 µm; E = 1.0–1.4, Q = 1.21 ± 0.1 subisodiametric; n = 47). B
cylindro-clavate and hardly tapered, rather narrow, 24.2–40.3 × 6.2–9.9 µm
(mean = 34.0 ± 4.3 × 8.0 ± 1.12 µm; E = 3.2–5.8, Q = 4.33 ± 0.81; n = 17),
4–sterigmate. C abundant, forming a sterile layer, colorless,
narrowly cylindro-clavate and wavy to sinuous in outline, occasionally with a
capitulate apex, 26.6–73.2 × 3.7–8.2 µm (mean = 49.7 ± 12.6 × 6.1 ± 1.18 µm; E
= 3.9–12.8, Q = 7.23 ± 2.2; n = 22). P absent. L
subparallel to parallel, composed of broad hyphae, these 56–86 × 10–19 µm,
along with narrow hyphae, these 83–85 × 7.2–7.4 µm (n = 9). P a
cutis but with a few loosely entangled clusters of cylindro-clavate terminal cells,
semi–erect on the disc, repent elsewhere; near disc 26.5–70.2
× 4.0–9.8 µm (mean = 43.8 ± 12.40 × 6.4 ± 2.22 µm; E = 4.61–14.32, Q = 7.41
± 2.89; n = 12), near margin 51.2–88.2 × 4.0–7.0 µm (n/1 = 3). P
beneath the pileipellis rather short-celled and somewhat inated; cells
38.9–138.8 × 7.0–17.9 µm (n = 9, not studied elsewhere). S with
abundant clusters of caulocystidia, these similar in shape to the cheilocystidia;
28.8–77.9 × 3.4– 6.2 µm (mean = 45.5 ± 13.37 × 4.7 ± 0.99 µm;
E = 5.0–13.5, Q = 9.9 ± 2.7; n = 12). S broad in the center;
Nolanea spp. nov. (Guyana) ... 143
P . Nolanea sinuolata (holotype; Henkel 8900). A. Basidioma. B. Basidiospores. C. Pileus
tramal hyphae with external encrustations. D. Cheilocystidia. E. Caulocystidia. Bars: A = 10 mm;
B–E = 10 µm.
cells 81.0–228.0 × 13.5–20.3 µm (n = 5) and narrow near the stipitipellis; cells
168.3–223.2 × 3.0–3.5 µm (n = 5). R absent. P
obvious and dark brown in 3% KOH in spot plates, dark brown in microscopic
144 ... Henkel & al.
sections, distinct and encrusting on the outer portions of the hyphae in the
pileipellis, pileus trama, and lamellar trama. C absent.
E, , — Solitary on litter of forest oor in mixed
Dicymbe spp. forest; known only from the Upper Potaro River Basin of Guyana.
A . GUYANA. R : P-S.
Pakaraima Mountains. Upper Potaro River Basin, ~15 km east of Mt. Ayanganna,
environs of base camp located on Potaro River one km upstream from conuence with
Whitewater Creek at 5°18′04.8″N 59°54′40.4″W, elevation 710–750 m; vicinity of base
camp, 1 July 2003, Aime 2280 (BRG; PUL).
C — Nolanea sinuolata is diagnosed by its conic, broadly umbonate,
dark brown pileus, brown, thin crowded lamellae, dark brown stipe, encrusted
pigment on the pileus tramal hyphae, cylindro-clavate cheilocystidia with
faintly wavy to sinuate sides, and 5–6-angled basidiospores averaging 8.1 ×
6.8 µm. Nolanea sinuolata, N. furcata, N. claviformis, and N. concentrica (all
described here as new) share the following features: a dark brown, conic or
conic-campanulate, macroscopically glabrous but sericeous to faintly brillose
pileus with a rounded, submammilate umbo; thin, close to crowded lamellae;
a tough, cartilaginous stipe concolorous with the pileus; basidiospore shape;
presence of cheilocystidia; an only slightly inated subpellis; encrusted pigment
on the outer hyphal walls of the pileipellis; and lack of clamp connections.
Among these species, N. sinuolata can be diagnosed by the brown lamellae
when young, the encrusted pigment on lamellar tramal hyphae, cylindro-
clavate cheilocystidia with slightly wavy walls and occasionally capitulate apex,
the absence of cytoplasmic pigment in the pileipellis, and the lack of concentric
ridges in the pileus. Nolanea furcata is distinguished by the white lamellae
that fork toward the margin, the abundant tomentum covering the basal ¼
of the clavate stipe, the chlorine odor, and a cytoplasmic pileipellis pigment.
Nolanea claviformis and N. concentrica have a concentrically ridged or beveled
pileus, white lamellae, and also have a cytoplasmic pigment in the hyphae of
the pileipellis. Nolanea claviformis is distinguished from N. concentrica by
the clavate cheilocystidia, larger (~9.0 × 7.5 µm) basidiospores, and larger
(28.3–42 5 × 9.5–12.0 µm) basidia. Nolanea concentrica has aciculate to
rostrate-ventricose cheilocystidia, smaller (~7.7 × 6.0 µm) basidiospores, and
smaller (24.2–30.6 × 6.7–9.5 µm) basidia.
Nolanea subsulcata Largent & T.W. Henkel, sp. nov. P
M MB
Diers from Entoloma guatopoanum by its glabrous pileus and stipe.
T: Guyana. Region 8: Potaro-Siparuni. Pakaraima Mountains. Upper Potaro River
Basin, ~15 km east of Mt. Ayanganna, environs of base camp located on Potaro River
one km upstream from conuence with Whitewater Creek at 5°18′04.8″N 59°54′40.4″W,
elevation 710–750 m; vicinity of base camp in Dicymbe forest, 1 June 2005, Henkel 8825
(BRG, holotype; HSU, isotype).
Nolanea spp. nov. (Guyana) ... 145
P . Nolanea subsulcata (holotype; Henkel 8825). A. Basidiomata. B. Pileipellis. C. Basidia.
D. Basidiospores. Bars: A = 10 mm; B–D = 10 µm
E: sub– and sulcatus (L. adj.) = somewhat grooved, referring to the pileus
surface texture.
P 16–22 mm broad, 5–10 mm tall, broadly conic with low, rounded umbo,
brownish orange (6C4 “red-haired”–7C4) throughout, hygrophanous over disc
with age, striate to subsulcate over outer 4/5, more smooth over immediate disc;
surface glabrous and nearly translucent when fresh, under hand lens appearing
nely longitudinally appressed-brillose, disc with no distinct vestiture, moist;
margin nely and irregularly crenulate at sulcations; extremely
thin, less than 1 mm throughout. L subthin, subdistant, adnexed,
brownish orange (7C4); margins concolorous, smooth; lamellulae 1 between
lamellae, 1–2 mm long. S 56–80 × 2–2.5 mm, equal, light greyish orange
(6B3 “esh”–6B4), appearing smooth but with ne appressed longitudinal
brils under hand lens, cartilaginous. B absent. O and
not noted.
B distinctly 6–8-angled in prole, dorsal, and ventral views,
sides slightly concave, heterodiametric, 8.2–12.3 × 5.1–7.2 µm, (mean = 9.7 ±
146 ... Henkel & al.
0.9 × 6.4 ± 0.51 µm; E = 1.3–1.8, Q = 1.5 ± 0.14; n = 31). B subcylindric
to subclavate, barely tapering toward base, 26.9–37.3 × 5.7–10.6 µm, (mean =
30.0 ± 3.0 × 8.3 ± 1.2 µm; E = 2.7–4.8, Q = 3.7 ± 0.63; n = 13), 2 or 4-sterigmate.
C absent. L subparallel,
composed of broad, long-celled hyphae (not measured). P a cutis
of 1–3 uninated hyphal layers; hyphae 3.2–10.0 µm wide;
cylindric to cylindro–clavate, 31.0–74.2 µm long. P
beneath the pileipellis moderately inated, 5–18 µm wide, elsewhere 5-40
µm wide. S a cutis, lacking hymenial clusters or caulocystidia.
R abundant in the pileus trama, scattered in the lamellar
trama. P cytoplasmic, faintly evident in 3% KOH. C
absent.
E, , — Solitary on humic litter layer of forest
oor in Dicymbe corymbosa forest; known only from type locality in the Upper
Potaro River Basin in Guyana.
C — Nolanea subsulcata is distinguished by its broadly conic,
umbonate, subsulcate, translucent-striate, hygrophanous pileus with a
paler crenulate margin; equal stipe; overall brownish orange to esh colors
throughout; lack of hymenial cystidia and clamp connections; and 6–8-angled,
heterodiametric basidiospores averaging 9.7 × 6.4 µm. Entoloma guatopoanum
(Dennis) E. Horak [≡ Leptonia guatopoana Dennis] from Venezuela, which also
has a sulcate-striate similarly colored pileus and heterodiametric basidiospores
measuring 9–11 × 6–8 µm and lacks hymenial cystidia and clamp connections,
can be distinguished from N. subsulcata by its pruinose pileus and its white,
minutely pruinose stipe with hyaline downy hairs (Horak 1977). Rhodophyllus
lutensis Romagn. & Gilles from Gabon has similarly sized, shaped, and colored
basidiomata but diers from N. subsulcata in its non-hygrophanous, non-
striate pileus, and much smaller (7–8.2 × 5–6 µm) basidiospores (Romagnesi
& Gilles 1979).
Acknowledgments
is research was made possible by grants from the National Geographic Society’s
Committee for Research and Exploration and NSF DEB-0918591 to TWH, and an
Explorer’s Club Exploration and Field Research Grant to MCA. Dillon Husbands
functioned as Guyanese local counterpart and assisted with eld collecting, descriptions,
and specimen processing. Mimi Chin, C. Andrew, L. Williams, V. Joseph, F. Edmund,
and L. Edmund provided eld assistance in Guyana. Research permits were granted
by the Guyana Environmental Protection Agency. Our special thanks go to Drs. Ron
Petersen and Clark Ovrebo for their critical comments and revisions, and Dr. Shaun
Pennycook for nomenclatural edits. is paper is number 206 in the Smithsonian
Institution’s Biological Diversity of the Guiana Shield Program publication series.
Nolanea spp. nov. (Guyana) ... 147
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