Nova Hedwigia 000 0 0 0000 0000 Wartchow 0916 Proof-1

C Nova Hedwigia, Vol.
000 (2024), Issue 0-0, 000–000

Published online January 2024
Early Access-Article
Studies on types and additional materials of Amanita

from Brazil
Felipe Wartchow
Universidade Federal da Paraíba, Departamento de Sistemática e Ecologia/CCEN, CEP:
58051-970, João Pessoa, PB, Brazil; fwartchow@yahoo.com.br
With 7 figures
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Abstract: Amanita is a well established genus in both morphologically and molecularly, with three
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recognized subgenera: Amanita, Amanitina and Lepidella.. In Brazil, the genus was infrequently
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studies during the Twentieth Century, but recent efforts reported more taxa from this country. Thus,
this work aims to perform type studies in Amanita and to determine the current name of some ma ma-
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terials deposited in Brazilian herbaria with more accuracy. The descriptions of the type materials
were updated using modern methods. As result the isotypes of Amanita sulcatissima (sect. Amani-
ta), A. coacta, A. craseodearma,and A. crebresulcata (sect. Vaginatae) are designed here as lecto-
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type since their holotypes were lost or destroyed. Other specimens identified under Amanita were
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studied and when possible, their generic status updated because some of them do not correspond to
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Amanita,, but to genera belonging Agaricaceae. Descriptions, discussions and drawings are provid
provid-
ed.
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Keywords: Agaricomycetes; Agaricomycetidae; Amanitaceae; the Neotropics; taxonomy

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Introduction
Amanita Pers. is a well established genus in both morphologically and molecularly. Bas
(1969) defined this genus as having bilateral lamella trama, presence of acrophysalides,
and the schizohymenial development of the basidioma (Yang & Oberwinkler 1999). Re-
garding molecular phylogeny, although changes in its infrageneric systematic, Tulloss et
al. (2016) and Cui et al. (2018) clearly showed that the genus is monophyletic, recogniz-
ing many sections distributed among the subgenera Amanita, Amanitina (E.-J. Gilbert)
E.-J. Gilbert and Lepidella Beauseign.
Studies on this genusin Brazil were infrequently performed during the Twentieth Century.
The most relevant studies were carried out by the Dutch mycologist Cornelius Bas, who
© 2024 J. Cramer in Gebrüder Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de

Germany DOI: 10.1127/nova_hedwigia/2024/0916 0029-5035/2024/0916 $ 0.00
eschweizerbart_xxx
2 Felipe Wartchow
described seven new species and a provisory name from the states of Amazonas in North-
ern (Bas 1978), and later Paraná in Southern (Bas & de Meijer 1993) regions of this
country. However, more species were reported for this country in the 21st century (see
summary by Wartchow 2017). This study aims to analyze the (1) type specimens of names
described by Bas (1978) and Bas & de Meijer (1993) for investigate the typification status
and redescribe them using modern methodology (Tulloss & Lindgren 2005); and (2) to
study additional exsiccates identified as Amanita deposited in some Brazilian herbaria to
perform an accuracy determination of these specimens.
Material and methods
Type specimens and additional materials deposited at CEPEC, INPA, IPA, JPB, L, MBM,
PACA, SP and URM (Thiers, continuously updated) were requested and studied. Mor Mor-
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phological studies, and biometric measurements and biometric variables were performed
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basing in the emended methodology of Tulloss & Lindgren (2005). Some terms used for
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Amanita are summarized by Wartchow & Gamboa-Trujillo (2012) and Wartchow et al.
(2013). For Amanita sp. 1 and Amanita sp. 2 the macroscopic description of the exsiccates
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are based in Singer’s field notes (in bold) and my direct observation on dry basidiome
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following Yang (2000), except for Amanita crebresulcata that is a compilation of all ma-
terials examined. For ‘habits’ and ‘habitats’ sections of the descriptions, I re-wrote them
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directly from Bas (1978), Bas & de Meijer (1993), and Simmons et al. (2002). Color
codes and names follow Maerz & Paul (1950) – M&P and Online Auction Color (2004)
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– OAC. Nomenclature and generic and infrageneric names and concepts follow Tulloss et
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al. (2016) and Cui et al. (2018).

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Results
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Exsiccates of the following taxa described by Bas (1978) and Bas & de Meijer (1993)
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were analyzed: Amanita lanivolva

lanivolva, A. sulcatissima, A. xerocybe (subg. Amanita sect.
Amanita), A. coacta, A. craseoderma, A. crebresulcata (subg. Amanita sect. Vaginatae),
A. gralipes (subg. Lepidella sect. Lepidella), A. campinaranae, and A. phaea (subg.
Amanitina sect. Validae). Except by A. grallipes, all other revised specimens correspond
to type material or belong to original collection. Additional exsiccates deposited in some
Brazilian herbaria were also analyzed with their identification corrected at least at genus
level.
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Studies on types and additional materials of Amanita 3
Subgenus Amanita section Amanita
Amanita lanivolva Bas, Persoonia 10: 13. 1978.

MycoBank: MB#308564
Fig. 1A–D
From protologue: ‘Basidiomata solitary, medium-sized, fairly slender. Pileus 50–75 mm,
plano-convex with broad central depression, subumbonate or not, moderately dark
slightly more reddish brown [M&P 15C2 “Clove”] with centre slightly more reddish
brown [M&P 8H11 “Congo”], and white interstriation at margin, glabrous, sub-viscid; in
dried specimens matt to somewhat shiny particularly at centre, outside centre slightly in-
nately radially fibrillose under strong lens; margin broadly sulcate-striate (40–60% of
ratio); context thin white, unchanging; universal veil lacking. Lamellae free, crowded,
narrow (probably less than 3 mm) white to pale grayish, probably with concolorous edge
but in dried state edges pale buff to pale ochraceous brown; lamellulae truncate to
obliquely truncate unevenly distributed. Stipe 75–100 × 5–6 mm, tapering upward, with
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slightly bulbous base, hollow, white, fragile, smooth and glabrous; context white, un un-
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changing; partial veil absent; universal veil saccate, membranous, loosely sheathing basal
30–35 mm of stipe, grey to fuscous grey [M&P 15A7 “Soaptone” to 15C8 “Chucker
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brown”], but lower half paler to whitish, tomentose lanose-tomentose; in dried specimen
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cylindrical to somewhat ventricose, adnate to lower 8–10 mm of stipe, with internal
volval limb a fimbriate 1–1.5 mm wide rim situated in angle between external volval limb
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and stipe (“unitangent volva”).’ Odorless; taste not checked by Bas.

Basidiospores [50/1/1] (6.5–)7–9 × (5–)5.5–7(–7.5) µm, ((L = 8 µm; W = 6 µm; Q =
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(1.21–)1.29–1.46(–1.49), Q = 1.34), inamyloid, hyaline, colorless, broadly ellipsoid to

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ellipsoid, smooth, thin walled, adaxially sub concavo rare flattened, with more or less at
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tenuate to some rounded apex; apiculus obtuse, small, sub lateral to sub apical; contents
including one guttule (not well observed in the material). Basidia 28–35 × 8–11 µm,
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clavate, four sterigmata 2–3.5 µm high, clamped at base. Subhymenium well rehydrated,
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cellular up to 35 µm thick with 2–3 layers of inflated cells 12–25 × 11–20 µm; wst-near
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= up to 25 µm; wst-far = up to 36 µm. Lamella trama difficult to rehydrating in the

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specimen, but obviously bilateral; wcs = ± 20 µm; abundant filamentous hyphae 4–8 µm
diverging from the central stratum gradually to rather abrupt, lacking more inflating hy-
phae, unbranched and rarely branched, frequently with clamps at the common septa; vas-
cular hyphae absent. Marginal tissue of lamellae sterile; up to 68 µm thick, two layered,
yellowish brown, with the outer layer covered by short chain of hyphae with terminal
clavate bodies up to 10 µm wide and the internal layer consisting of globose broadly
clavate or ovoid brownish cells (although more difficult to localize) moderately thick
walled and granular contents. Pileus context not satisfactorily rehydrated; acrophysalidi-
oid bodies up to 22 × 14 µm, broadly clavate, inconspicuous; filamentous hyphae 2.5–
10 µm, plentiful, very interwoven, commonly branched, septade, clamped; vascular hy-
phae not observed or lacking. Stipe context very well rehydrated, distinctly acrophysa-
lidic; acrophysalides 190–300 × 40–55 µm, plentiful, very conspicuous, clavate; filamen-
tous hyphae 4–8 µm, longitudinally oriented, rarely branched, septate, clamped; vascular
eschweizerbart_xxx
4 Felipe Wartchow
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Fig. 1. A–D. A. lanivolva. A. Universal veil from sac (after crushed). B. Basidiospores. C. Basidia and
subhymenium. D. Dried basidioma (paratype). E–H. A. sulcatissima. E. Basidiospores. F. Basidium,
basidioles and subhymenium. G. Universal veil from stipe base (after crushed). H. Universal veil from
pileus showing the adjacent. Scale bar is 10 µm for microstructures and 10 mm for basidioma.
eschweizerbart_xxx
hyphae 5 µm wide, unbranched, rare in the middle of the context turning more abundant
near the surface. Pileipellis a cutis differentiated into two layers; suprapellis up to 90 µm
thick at centre with radially to gradually erect filamentous hyphae 3–5 µm wide, distant,
somewhat with inflation knobs, slightly gelatinized, colorless, with clamps in all septa;
subpellis up to 112 µm thick at centre, with filamentous hyphae 2.5–9 µm wide, radially
oriented, most dark colored; vascular hyphae absent. Universal veil on pileus absent; on
stipe base volval limb made up of a 200 µm thick exclusively with filamentous hyphae
2.5–5 µm, very interwoven, hyaline and vascular hyphae rare up to 4 µm wide; the outer
surface covered by filamentous hyphae 3–15 µm wide, mostly grayish brown pigment
and slightly thick walled and terminal globose, piriform to broadly ovoid cells 27–115 ×
18–95 µm mixed with some clavate or broadly clavate (e.g. 95–105 × 32–45 µm) and
some elongate (e.g. 137 × 20 µm) or fusiform mucronate (e.g. 95 × 35 µm), yellowish
brown to brown and some pale colored, thick walled 1.7–2.5 µm; the internal surface is
immersed in a strongly gelatinous matrix. Partial veil absent.
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Habitat: “Fairly common on the ground in secondary tropical rain forest near Neea
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(Nyctagianaceae; the collection R. Singer B 9897 definitively show to be connected with
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short roots of Neea sp.), with nearby Rubiaceae, Sapindaceae, Euphorbiaceae, Legumi-
Legumi
nosae, Flacourtiaceae and Palmae in Amazonia” (Bas 1978). This species also occurs in
Bas 1978). p
“Riverine swamp forest on flat ground dominated by Dicymbe altsonii and D. corymbosa
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with scattered associate hardwoods” (Simmons
Simmons et al. 2002).
2002).
Material examined: BRAZIL. Amazonas, sede do INPA, 29 May 1977, R. Singer B
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9897 (INPA 66716, paratype!).

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Observation on exsiccate: The exsiccate consists in a well preserved specimen with

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detailed field notes given by Singer. Dr. Dirce L. Komura (pers. corresp.) kindly sent me
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the images of the holotype (R. Singer B 9843) and apparently it is also well preserved.
However, the exsiccate analyzed here is neither the holotype nor isotype, but a paratype.
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Remarks: Amanita lanivolva characterized among sect. Amanita by the unusual presence
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of a membranous-saccate universal veil recovering the entire basal bulb ((Tulloss & Hal-
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ling 1997).
). This structure is also observable in the analyzed exsiccate (pers. observ.). As
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additional characteristics of this species, the reddish brown pileus, broadly ellipsoid to
ellipsoid basidiospores, and proliferating clamp connections in basidia and universal veil
are also characteristic (Bas 1978). Recent collection of A. lanivolva from Guyana was
described with small cluster of gray, conical or even patches of velar remnants at centre
of pileus, and slightly longer basidiospores [40/3] 7.4–9.8 × 5–6.6(–7.2) µm, Q = 1.30–
1.65, Qm = 1.40–1.55 (Simmons et al. 2002).
Another South American species with similar volval behavior is A. pseudospreta Raithelh.
from Nothofagus forest in Argentina. However, it differs in the gray pileus, presence of
partial veil and the larger globose to subglobose to broadly ellipsoid basidiospores
[40/1/1] (8.8–)9–11.2(–12.2) × (7.8–)8.5–10.2(–11.2) µm, L = 10.2 µm, W = 9.3 µm,
Q = (1.00–)1.02–1.18(–1.56), Q = 1.10 (Tulloss & Halling 1997, Raithelhuber 2004).
eschweizerbart_xxx
6 Felipe Wartchow
Tulloss & Yang (2023) reported at least two other taxa of sect. Amanita with saccate volva
covering the entire basal bulb: A. rhodophylla Beeli and A. aurea (Beeli) E.-J. Gilbert.
The first taxon differs at least in the yellowish white pileus and pink lamellae. Amanita
aurea differs in the golden yellow pileus, yellow lamellae and basidiospores (5–)5.5
(–5.8) × (4.5–)5.3 µm. Pegler & Shah-Smith (1997) reported A. aurea taxon from Zambia
with basidiospores measuring 4–5 × 3.5–4.5 µm, L = 4.7 µm, W = 3.8 µm, Q = 1.12.
Regarding infrageneric placement of A. lanivolva, preliminary evidence rising this ques-
tion after suspicious of a small roundish bulb in the exsiccate (Bas 1978). Later Tulloss &
Halling (1997) observed similar structure in the type of A. pseudospreta. This question
was only solved after Simmons et al. (2002) discovered small, but distinct bulb inside the
saccate volva of fresh Guyanese specimens of this species. These conclusions were cor-
roborated by recently phylogenies performed by Mighell et al. (2021).
Amanita sulcatissima Bas, Persoonia 10: 9. 1978.
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MycoBank: MB#308595
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Fig. 1E–H
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From protologue: ‘Basidiomata rather small and short-stemmed, solitary. Pileus 40 mm
wide, plane-convex with slightly flattened centre, at first sordid whitish, later becoming
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ochraceous, at centre fairly dark, slightly olivaceous tinged ochraceous brown (M&P
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14J8 ‘Maple sugar’) to slightly paler and slightly more reddish ochraceous brown (M&P
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interstria-
14J9 ‘Mummy’), at margin pale buff gray (M&P 11C3 ‘sheepskin’) with pallid interstria
tion, in dried state matt to somewhat shiny and (even under strong lens) without visible
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fibrillose structure; margin densely sulcate-striate (14–18 grooves per 10 mm of circle

half way sulcate zone); context fleshy, white to whitish, unchanging; universal veil as
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large patch (in dried state minutely tomentose and tending to disrupt in vague small
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patches with very pale arachnoids edges), pale brownish gray (M&P 13P6 ‘almond’).
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Lamellae free, narrow, pale grayish buff (M&P 11E4 ‘maple’; slightly darker than Muns.
2.5 Y 8/4), in dried state with slightly darker (brownish ochraceous) edge; lamellulae
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scarce, truncate. Stipe 45 × 4.5 mm, tapering upward, gray (M&P 13C4 ‘handman’;
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Muns. 10 YR 6/4-5/4) with paler apex, minutely fibrillose, lower part rather glabrous,
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whitish with small ochraceous-brown spots; bulb ca. 7 mm high and 10 mm wide, rather
abrupt, slightly flattened subglobose; context solid (hollow in dried state), white to whit-
ish, unchanging; partial veil absent; universal veil arising from upper part of bulb, ap-
pressed, pale brownish gray, with sub tomentose to sub lanose surface, at one side of stipe
forming a thin sub membranous limb up to 10 mm high above bulb, but for the rest form-
ing thin vaguely delimited patches not reaching so high as limb. Odor absent and taste
not recorded’.
Basidiospores [60/1/1] (6–)7–9.5(–10) × (4.5–)6–7(–7.5) µm, (L = 9.1 µm; W = 6 µm;
Q = (1.20–)1.28–1.69(–1.81), Q = 1.42), inamyloid, hyaline, colorless, broadly ellipsoid
to ellipsoid infrequently to elongate, usually adaxially flattened, smooth, thin walled;
apiculus obtuse, sub lateral to sub apical; contents including one large guttule. Basidia
32–43 × 12–13.5 µm, clavate somewhat appearing fusoid, bearing four sterigmata c.
5 µm high, clamped at base. Subhymenium rehydrating moderately, cellular up to 24 µm
eschweizerbart_xxx
thick with 3 (somewhat 4) layers of variously inflated cells 10–18 × 7–16 µm, somewhat
broadly clavate or ovoid; wst-near = 10–16 µm; wst-far = 21–25 µm. Lamella trama re-
hydrating moderately well in the specimen, bilateral; wcs= ± 22 µm; abundant filamentous
hyphae 3.5–8 µm frequently branched, with proliferate clamps at the septa gradually di-
verging from the central stratum some more inflating hyphae ranging to 13.5 and infre-
quently up to 26 µm wide also diverging; vascular hyphae absent. Marginal tissue of la-
mellae not studied. Pileus context very thin, slightly satisfactorily rehydrated; acrophysa-
lidioid bodies up to 75 × 40 µm, broadly clavate, inconspicuous; filamentous hyphae
2.5–12 µm, plentiful, very interwoven, commonly branched, septade, clamped; vascular
hyphae up to 5 µm, very occasional. Stipe context moderately well rehydrated, longitudi-
nally acrophysalidic; acrophysalides 140–275 × 42–60 µm, plentiful, very conspicuous,
clavate to subfusoid, somewhat in chains; filamentous hyphae 2.5–10 µm, longitudinally
oriented, rarely branched, septate, clamped; vascular hyphae 2–20 µm wide, unbranched,
common, with longitudinal orientation; stuffed zone of the bulb with hyphae 2.5–6 mm
resem-
very interwoven, somewhat with pale incrustation wall, clamped; inflated cell that resem
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bles acrophysalides up to 175 × 55 µm. Pileipellis a cutis differentiated into two layers
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comprising 180 µm; suprapellis up to 25 µm thick at centre with compressed filamentous
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subpel-
hyphae 3–6 µm wide, non gelatinized, pale or colorless, with clamps in all septa; subpel
lis up to 150 µm thick at centre, with filamentous hyphae 3–5 µm wide, radially oriented,
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most dark colored; vascular hyphae absent. Universal veil: On pileus up to 130 µm thick;
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in the inner layer comprising almost exclusively hyphae 2–6 µm, thickness wall, arising
from the pileipellis which is more or less connected; the upper layer comprising almost
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exclusively of inflated cells globose (e.g. 57 × 57 mm) or ovoid (e.g. 57 × 40 mm), pale
grayish to yellowish brown to brown; immediately bellow having hyphae that somewhat
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range to 12 mm wide, brown colored with slightly incrusted walls. On stipe base: outer
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layer with terminal globose, piriform to broadly ovoid cells 22–57 × 22–55 µm mixed
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with some clavate (e.g. 50 × 20 µm) and some slender ellipsoid (e.g. 50 × 18 µm), yel-
lowish brown to brown and some pale grayish colored, thick walled c. 2 µm; filamentous
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hyphae 2–6 µm, arising from the bulb, somewhat branching, thin to thick walled, hyaline;
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vascular hyphae absent. Partial veil absent.

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Habitat: “in campina vegetation on humus under trees (Humiriaceae, Burseraceae,

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Ochnaceae, Sampindaceae, Sapotaceae and leguminous trees)” (Bas 1978).

Known distribution: State of Amazonas, North Brazil.
Material examined: BRAZIL. Amazonas, road from Manaus to Caracaraí, km 45, 25 Jul
1977, R Singer (& I. Araujo) B 10123 (L 0053722, isotype!).
Observations on exsiccate: The analyzed isotype is well preserved.
Remarks: Bas (1978: 11) referred the remarkable volval limb present only at one side of
bulb and the complete lack of membranous inner limb. He explains that this layer main-
tain its individuality only where the volva loses its contact with the pileipellis in early
stage. In other places where this structure contacts the inner volval layer slightly gelati-
nizing itself, and merges from equally the gelatinized pileipellis. However, this gelatini-
zation was not visible in exsiccate. I also observed that the universal veil hyphae imme-
eschweizerbart_xxx
8 Felipe Wartchow
diately above pileipellis maintain certain connectivity with the pileipellis, but immedi-
ately dissociated when the section is crushed for microscope observation.
Bas (1978) compared A. sulcatissima with A. farinosa Schwein. from USA and A. obsita
Corner & Bas from Malaya which differ mainly in the distinct farinose velar remnants
and lack of basidial clamps (Corner & Bas 1962, Jenkins 1977, Sanmee et al. 2008).
Amanita subvaginata (Cleland & Cheel) E.-J. Gilbert was also used by Bas for compari-
son with the Amazonian taxon, but Reid (1980) did not report if clamps occur or not in
the basidia. Actually, A. sulcatissima might be a rather morphologically isolated taxon if
considering the structure of the universal veil and the conspicuously proliferated clamps
in all examined tissues.
Amanita xerocybe Bas, Persoonia 10: 7. 1978.

MycoBank: MB# 308602
Fig. 2A–E
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From protologue: ‘Basidiomata medium-sized, fairly slender, fleshy, rather fragile, soli
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soli-
tary. Pileus 40–67 mm wide, from conico-convex with broadly rounded apex to convex
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and soon flattened around a low, obtuse umbo, at first sordid whitish, later becoming
ochraceous, at centre brownish ochraceous yellow to ochraceous brown [M&P “Chip “Chip-
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munk” to “Antique Brown”], subfelted-subflocculose; margin widely sulcate-striate (25–
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35% of radius; radial marginal ridges 1.8–2 mm broad) and crenulate edge; context fleshy,
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white to creamy, unchanging; universal veil entirely decorating (densely at centre to

sparsely at margin) as minute gilvous adnate granular-subflocculose dots of volval origin.
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Lamellae distant, free, somewhat intervenose, broad, whitish but soon (particularly on
subflocculose-subgranu-
drying) becoming cream, in dried specimens buff with minutely subflocculose-subgranu
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lamel-
lar concolorous edge; lamellulae very scarce, truncate, sometimes partly adnate to lamel
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lae. Stipe up to 90 × 5 mm, equally thick, gilvous to ochraceous tan (in dried material
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pallid with fine, flocculose brownish buff to ochraceous brown covering); bulb up to
9 mm, enlarged to slenderly fusiform; context narrowly stuffed; partial veil apical, easily
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separating and falling down, thick and fluffy, smooth, concolorous with surface (dried:
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felted-subflocculose at underside; minutely granular at upper side); universal veil as an

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appressed, thin, flocculose (in dried specimens sordid brownish ochraceous) covering
forming a narrow fragmented rim around the upper part of the bulb (hardly visible in
dried state).’ Odor absent and taste not recorded.
Basidiospores [50/2/1] (6.5–)7–9 × (6.5–)7–8.5 µm, (L = 7.8–7.9 µm, L’ = 7.9 µm; W =
7.4–7.5 µm, W’ = 7.5 µm; Q = (1.00–)1.02–1.12(–1.18); Q = 1.05–1.07, Qm = 1.06),
inamyloid, hyaline, colorless, globose to subglobose occasionally broadly ellipsoid,
smooth, moderately thick walled (0.5 µm) to somewhat thickening to c. 1 µm and with
apparent pits (crassospores?); apiculus rather broad or gradually attenuate; contents
showing one large guttula. Basidia difficult to observe e.g. 30–35 × 8–10.5 µm, 4-sterig-
mate, with sterigmata up to 2.5–45 µm long, clamps absent. Subhymenium cellular up
to 31 µm thick, as 2–5 layers of isodiametric cells 14–20 µm diam., some with rather in-
conspicuous peduncles, but difficult to see in other cells; wst-near = ± 30 µm; wst-far =
± 35 µm. Lamella trama indistinctly bilateral, with some narrow elements appearing
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Fig. 2. A–D. Amanita xerocybe (Holotype). A. Basidiospores. B. Basidia and subhymenium. C.

Pileipellis and base of the velar remnants intimately connected. D. Universal veil from centre of
pileus. E. Universal veil of stipe base. F–I. A. coacta (Lectotype). F. Basidiospores. G. Elements of the
Universal veil from pileal patch. H. Hymenium, subhymenium and terminal element of the lamella
trama. I. Elements of universal veil from stipe base. Scale bars are 10 µm.
eschweizerbart_xxx
10 Felipe Wartchow
to gradually diverging from central stratum; wcs = ± 22 µm; filamentous hyphae up to

3–10 µm wide, usually branched; vascular hyphae apparently absent or not observed.
Sterile marginal tissue on lamella edge not seen, but a colorless ellipsoid cell short pedun-
culate with thickened wall up to 52 × 32 µm was observed. Marginal tissue of lamellae
not studied. Pileus context moderately well rehydrated, distinctly acrophysalidic with
acrophysalides up to 112 × 45 µm; filamentous hyphae 1.5–4.5 µm wide, plentiful, com-
monly branched, very interwoven; vascular hyphae very scarce up to 7.5 µm wide. Stipe
context longitudinally acrophysalidic; acrophysalides 200–290 × 22–37 µm, mostly
clavate and some subfusoid; filamentous hyphae 3–10 µm wide, longitudinally oriented,
rarely branched, often turning more or less inflated; vascular hyphae 7.5–13 µm wide,
very scarce, unbranched. Pileipellisa cutis apparently ranging to 110 µm at centre, with-
out suprapellis; filamentous hyphae plentiful up to 3–6 µm wide, frequently with radial
orientation; vascular hyphae 6 µm, very rare; not gelatinized. Universal veil: On pileus:
at upper layers comprising very abundant erect inflated pedicellate cells 37–55(–75) ×
27–60 µm, with pedicel ranging to 9 µm long, yellowish brown to brownish pigmented
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and slightly thickened wall 2.5 µm wide; filamentous hyphae 4–10 µm wide, relatively
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plentiful in lower layer near to pileus surface, intimately connected to pileipellis, where
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they arise turning more scarce in the upper layer, with thin to slightly thickened wall, of
ten colorless. On bulb: similar to that on pileus, in outer layer with inflated pedicellate
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cells 30–50 × 22–45 µm and some inflated clavate elements e.g. 37–50 × 15–22 µm,
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filamen-
mostly colorless with rarely pale gray pigmented, with slightly thickened wall; filamen
tous hyphae 3–10 µm, thin walled, plentiful at internal layer. Partial veil filamentous hyhy-
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phae 3–6 µm, strongly interwoven, branched, plentiful; inflated elements rare or difficult
to observe; elements similar to that of universal veil of bulb 34–50 × 30–40 µm, colorless,
c
thick wall present between the filamentous hyphae.

e
rr
Habitat: “on raw humus under trees in campinarama vegetation, forming ectomycorrhi
ectomycorrhi-
zae with Sapotaceae and/or Leguminosae” ((Bas 1978). Also in “riverine swamp forest
o
and adjacent slope forest dominated by Dycimbe corymbosa (paluwayek) and other
c
sand-
missed hardwoods, on sand soils with thick organic accumulations and on exposed sand
n
stone and cliff boulders” ((Simmons et al. 2002).

u
Known distribution: State of Amazonas, North Brazil (Bas 1978); Pakaraima Moun-
tains, Gyana (Simmons et al. 2002).
Material examined: BRAZIL. Amazonas, road from Manaus to Caracaraí, km 45, 28
Feb 1978, H. St.John & R Singer B 10730 (INPA 73540, holotype!).
Observations on exsiccate: The holotype consists in two basidiomes, but details of la-
mella trama were difficult to observe due to weak rehydration.
Remarks: A. xerocybe is macroscopically characterized by the farinose velar remnants
on pileus and stipe. In this universal veil, the hyphae with more basal localization are in-
timately connected to pileipellis until maturity, given the farinose appearance to pileus.
This species is also characterized by the globose to subglobose basidiospores and clamp-
less basidia (Bas 1978). The species was also collected in Guyana, but with slightly
smaller pileus, relatively longer marginal striations and brighter colored velar remnants
eschweizerbart_xxx
(Simmons et al. 2002). According to Bas (1978), other species with pulverulent velar
remnants on pileus can be compared with A. xerocybe as follows:
Amanita obsita Corner & Bas from Singapore differs in the pallid-whitish pileus and
white, exannulate stipe from early development and basidiospores 5.8–6.7 × 5.2–6.1 µm,
Q = 1.00–1.15, Q = 1.10 (Corner & Bas 1962).
Amanita farinosa Schwein. from the Quercus-Fagus forest in North and Central America,
differs at least in the deeply striate pileus margin, exannulate stipe and broadly ellipsoid to
ellipsoid basidiospores [80/4/4] (6–)6.5–9(–10.5) × (5–)5.5–7.2(–9) µm, L = 7.3–8 µm,
L’ = 7.7 µm, W = 6–6.5 µm, W’ = 6.3 µm, Q = (1.03–)1.08–1.40(–1.47), Q = 1.15–1.31,
Qm = 1.21 (Tulloss et al. 1995, Tulloss & Traverso 2000).
Amanita subvaginata (Cleland & Cheel) E.-J. Gilbert from Australia was not described in
detail, but it differs at least in the grayish pileus and the marginate volva (Gilbert 1940,
Reid 1980).
f
Amanita pulverotecta Bas (Malawi), A. nehuta G.S. Ridl. (New Zealand), and A. siamen-
o
sis R. Sanmee, Zhu L. Yang, P. Lumyong & S. Lumyong (Thailand) also have powdery
ro
universal veil remnants on the pileus, but they also differ from the present species with
regard to pigmentation, habitat and the more elongate basidiospores ((Bas 1982, Ridley
p
1991, Sanmee et al. 2003).
d_
Finally, the Neotropical A. chrysoleuca Pegler from Martinique differs in the floccose-
te
powdery velar remnants on pileus, the small and reduced floccose marginal rim at top of
bulb, universal veil with a fine incrusted pigment and ellipsoid basidiospores 7–9.5 ×
c
4.5–6 µm, L’ = 8 µm, W’ = 5 µm, Q = 1.56 (Pegler 1983).

(Pegler 1983
e
rr
Subgenus Amanita section Vaginatae

o
c
Amanita coacta Bas, Persoonia 10: 15. 1978.

n
MycoBank: MB#308546
u
Typification: BRAZIL. Amazonas, INPA, 27 May 1977, R. Singer B 9728 (INPA 66714
holotypus!; L 0053684 isotypus!– lectotypus hic designatus!).
MycoBank Typification #(MBT) 10016357
Fig. 2F–I
From protologue: ‘Basidiomata solitary, small and slender. Pileus up to 40 mm, concave,
without or with a slight umbo, umber with pale interstriation at margin, matt in dried state
and then with fibrillose structure only visible on dark marginal ridges (under strong lens);
margin broad rather densely sulcate-striate margin (half radius according to field notes;
10–14 grooves per 10 mm of circle half-way sulcate zone); context fleshy at centre the
thinning toward margin, white, unchanging; universal veil as a few irregularly shaped,
pale grey-brown, felted, thin patches with a few very small darker warts (under lens) at
centre of their surface. Lamellae free, crowded, fairly broad, white, probably with con-
eschweizerbart_xxx
12 Felipe Wartchow
colorous edge; lamellulae very scarce, truncate. Stipe up to 70 × 3 mm, tapering upward,
without bulb, white, in dried state sub flocullose (under strong lens); context white, un-
changing; partial veil absent; universal veil at base up to 10 mm high, thin, felted-sub-
membranous, appearing to broken into partly clearly separated, partly still more or less
connected, felted grey patches with their surface appressed paler to nearly whitish short
fibrils (under strong lens).’ Odorless; taste not recorded.
Basidiospores [40/1/1] (6.5–)7.5–10(–10.5) × (4.5–)5.5–7.5(–8) µm, (L = 8.8 µm; W =
6.4 µm; Q = (1.24–)1.28–1.47(–1.50); Q = 1.36), in amyloid, hyaline, colorless, broadly
ellipsoid to ellipsoid, usually at least somewhat adaxially flattened, with broadly rounded
apex, smooth, thin walled; apiculus small sub lateral; contents apparently including one
guttulate (not well observed in the type). Basidia 30–47 × 10–13.5 µm, 4-sterigmate, each
c. 5 µm long, clampless. Subhymenium: cellular, up to 29 µm thick, frequently 3 layers
of isodiametric to almost isodiametric cells 13–23 × 13–20 µm; wst-near = ± 20 µm; wst-
far = ± 35 µm. Lamella trama conspicuously bilateral; wcs= 32 µm; filamentous hyphae
f
up to 5 mm wide somewhat inflating to 15 µm wide, gradually diverging; vascular hyhy
o
phae absent. Marginal tissue of lamellae: sterile; terminal subisodiametric cell e.g. 28 ×
ro
25 µm, hyaline, thin walled and hyphae ranging to 5 µm wide, parallel to edge. Pileus
context apparently acrophysalidic; with acrophysalides-like bodies up to 32 × 15 µm;
p
filamentous hyphae 2.5–5 µm, plentiful, very interwoven and frequently septate; vascular
d_
hyphae somewhat scarce varying 5–11 µm in same hyphae or more or less uniformly
thickened measuring 5 µm. Stipe context longitudinally acrophysalidic; acrophysalides
te
up to 300 × 48 µm; filamentous hyphae 2.5–6 µm wide, branched, with more or less lon lon-
gitudinal orientation; vascular hyphae 2.5–17 µm wide, abundant and irregularly ori- ori
c
ented, very branched or dichotomous. Pileipellisa cutis 25–37 µm thick, with hyphae up
e
to 3 µm wide, more or less radially oriented or interwoven, with scattered universal veil
rr
elements under microscope (but not visible under hand-lens), colorless, not gelatinized.
Universal veil: On pileus: moderately dense tuffs with abundant short-celled hyphae 3–
o
6 µm, slightly thickened and colored walls, indistinctly incrusted, and terminal symmetri-
symmetri
c
cally globose (e.g. 27–40 × 27–37 µm) or rarely ovoid elements (e.g. 25–45 × 17–29 µm)
n
brown to light brown pigmented, moderately thick walled. On stipe: similar to those on
pileus but appearing to be double edged on which the internal layer contain plentiful and
u
interwoven filamentous hyphae 2.5–7.5 µm, mixed to terminal elements symmetrically

globose or nearly so (e.g. 25–37 × 24–37 µm), with some ovoid or ellipsoid (e.g. 35 ×
20 µm) elements, colorless; outer surface contain filamentous hyphae more yellowish and
thick wall distinctly encrusted, and some piriform, terminal elements 50 × 37 µm. Partial
veil: absent.
Habitat: ‘On the ground in a secondary tropical rain forest, all accompanying trees and
shrubs except Neea and Psychotria without ectomycorrhizal associations’ (Bas 1978).
Known distribution: Amazon Forest of the State of Amazonas, North Brazil and Atlan-
tic Forest from the State of São Paulo, Southeast Brazil.
Additional material examined: BRAZIL. São Paulo, Parque Estadual das Fontes do
Ipiranga, 21 Jan 1987, D.N. Pegler 3810 (SP 214459); same place, 19 Nov 2002, M.
eschweizerbart_xxx
Capelari & D.M. Vital MC4189 (SP 393670); same place, 11 Jan 2007, F. Karstedt FK841
(SP 393677); same place, 20 Apr 2007, N. Menolli Jr. et al. NMJ126 (SP 393681).
Observations on exsiccate: The holotype (INPA 66714) is completely destroyed, lacking
all the hymenial surface. Only fragments of pileus and very few basidiospores were
found, turning difficult to interpret the species based in this specimen. On the other hand,
the isotype at L is well preserved. According to code, if the holotype has been lost or de-
stroyed (in this case), the lectotype must be designed (Art. 9.3 and 9.11, Turland et al.
2018). Even so, I choice the isotype deposited in the herbarium L as the lectotype of A.
coacta.
Specimen of A. crebresulcata sensu Pegler (1997) and subsequent collections identified
as A. coacta by Menolli et al. (2009) from the State of São Paulo also were analyzed and
the microscopy is redescribed as follow (Fig. 3A–E):
Basidiospores [142/5/4] (7.5–)8–10(–11) × (5.5–)6–7(–7.5) µm, (L = 8.7–9.1(–9.4) µm;
f
L’ = 8.9 µm; W = 6.5–7 µm W’ = 6.7 µm; Q = (1.14–)1.18–1.56(–1.69); Q = (1.28–)
o
1.31–1.37; Qm = 1.33), inamyloid, colorless, hyaline, smooth, thin walled, broadly el el-
ro
lipsoid to ellipsoid then occasionally subglobose or elongate, usually or at least somewhat
adaxially flattened; apiculus sublateral to subapical; contents with one or two large gut-
gut
tula. Basidia 30–45 × 9–13 µm, 2- or more frequently 4-sterigmate, with sterigmata 3–
p
d_
5 µm long, clampless. Subhymenium moderately rehydrated, cellular, up to 27–32 µm
thick but also with more or less isodiametric cells 16–25 × 14–21 µm somewhat with sub
te
angular appearance, 3–4 layers; wst-near = 45–60 µm; wst-far = 63–73 µm. Lamella
trama rehydrating moderately; bilateral, wcs= 27–33 µm, with plentiful filamentous hy hy-
c
phae ranging to 7 µm and some long-clavate inflated elements up to 60 × 14 µm, gradu-

e
ally diverging from central stratum with angle frequently slightly superior than 45°; vas-
vas
rr
cular hyphae absent. Pileuscontext moderately well rehydrated; acrophysalides e.g. 55–
90 × 20–32 µm, inflate clavate to broadly ellipsoid, somewhat apparently thick walled,
o
common; filamentous hyphae 6–8 µm, very interwoven, common; vascular hyphae up to
c
10–17 µm, common. Stipe context moderately rehydrated, longitudinally acrophysalidic;

acrophysalides 175 × 30 µm longer, long clavate, hyaline, common; filamentous, hyphae
n
3–15 µm, with a longitudinal orientation, somewhat branched, common; vascular hyphae
u
5–16 µm, unbranched, common; partially gelatinized. Pileipellis: up to 120 µm thick with
hyphae more or less radially oriented to somewhat more sparsely at upper surface ca.
6 µm, having pale yellowish brown pigment locally condensed, not or only weakly ge-
latinized; vascular hyphae 6 µm, scarce, with apparently the same orientation of filamen-
tous hyphae or sub perpendicular. Universal veil. On pileus: terminal isodiametric to
subisodiametic but also some ovoid 27–42(–60) × 22–37(–45) µm or less frequently
clavate (e.g. 70 × 20 µm) cells with yellowish brown or slightly paler, slightly thick
walled, very common; filamentous, hyphae 3–6 µm, interwoven and frequently short
celled, somewhat incrusted, common; vascular hyphae 6 µm, unbranched, rare; more or
less gelatinized. On stipe base: plentiful terminal globose, subglobose to ovoid cells 20–
50 × 13–45 µm, with some clavate (e.g. 25 × 12 µm) yellowish brown to paler cells,
somewhat incrusted walls 1 µm thick, colorless or pale yellowish brown; filamentous
hyphae 3–8 µm, interwoven, branched, short celled, common; vascular hyphae absent;
eschweizerbart_xxx
14 Felipe Wartchow
f
o
ro
p
d_
te
c
e
rr
o
c
n
u
Fig. 3. A–E. A. coacta (SP 214459). A. Elements of the universal veil from stipe base. B. Elements of
the Universal veil from pileus. C. Elements of universal veil from the limbus internus. D. Basidiospores.
E. Hymenium, subhymenium and terminal divergent elements of the lamella trama. F–I. A.
craseoderma (Lectotype). F. Basidiospores. G. Hymenium and subhymenium. H. Elements of the
universal veil from white covering of the stipe base. I. Elements of the Universal veil from pileus.
Scale bars are 10 µm.
eschweizerbart_xxx
strongly gelatinized. On stipe 10 mm above base of SP 214459 and SP 393677 (inner

limb): with similar terminal cells, ranging to 27 × 27 µm in SP 214459 and 74 × 59 µm in
SP 393677, yellowish to brownish or paler and wall 1 µm; filamentous hyphae also simi-
lar and somewhat incrusted, common; vascular hyphae up to 23 µm, very rare. Partial veil
absent.
In the materials from São Paulo, a floccose belt above the submembranous cup is fre-
quently observed. It represents the inner limb of the universal veil, i.e. the rest of the
primordial tissue between the lamellae and the stipe after this has been partially trans-
formed into or replaced by the partial veil (Bas 1969: 312). On the other hand, in the type
specimen that structure was not observed, which is suspected to be lost due environmen-
tal factors, handling etc. When partial veil is lacking, as occurs in A. coacta, this tissue is
incoherent and might falls in small pieces during pileus expansion (Bas 1969: 316).
Summarizing, A. coacta is also characterized presenting a small cupulate felted-submem
felted-submem-
branous universal veil on the stipe base that breaks in felted patches. This feature appears
f
to be intermediate between the completely friable and saccate universal veil in sect. Vag-
o
inatae (Bas 1969).
ro
The pileus color of all São Paulo’s materials slightly differs from the protologue, but the
p
microscopic features of the velar remnants on pileus are similar. Bas (1978) clearly stated
d_
that the relevant features for differentiate A. coacta from A. crebresulcata are friable and
the non-saccate universal veil, and the considerable narrower basidiospores (Q’
( = 1.36 in
te
the isotype and Q’ = 1.33 in São Paulo material against Q’ = 1.20 in A. crebresulcata).
Thus, the specimens collected in the Amazon and Atlantic Forest biomes (IBGE 2004,
c
2012) correspond to the same species, which also can be characterized by the following
e
characteristics: (1) the felted velar patches on pileus and stipe near to cupulate universal
rr
veil (see also Bas 1978); ); (2) basidiospores measuring [219/7/6] (6.4–)7.2–11(–11.5) ×
(4.5–)5.3–8.3(–8.5) µm, L = 8.7–9.2(–9.4) µm, L’ = 9 µm, W = 6.4–7.1 µm, W’ = 6.7 µm,
o
Q = (1.14–)1.19–1.56(–1.69), Q = (1.28–)1.31–1.37, Qm = 1.33; (3) the occurrence of

c
frequent (dark colored) short celled hyphae in the universal veil; and (4) the relatively
n
common large subhymenial cells ranging to 25 µm in diam.

u
Remarks: The African A. calopus (Beeli) E.-J. Gilbert might be phenetically similar, due
to the presence of a small cup-like volva on stipe base. However, it differs at least in the
larger and elongate basidiospores [20/1/1] 7.8–13.6(–15.1) × (5.5–)6.0–8.8(–9.0) µm,
L = 11.7 µm, W = 7.7 µm, Q = (1.30–)1.39–1.60(–1.78); Q = 1.53 (Tulloss & Yang
2023). According to Pegler & Shah-Smith (1997), this species was referred with slight
swollen stipe base (see Amanita sp. 1 below), graying lamellae and also more elongate
basidiospores 10–14 × 7–9 µm, L = 11.7 µm, W = 7.7 µm, Q = 1.50.
eschweizerbart_xxx
16 Felipe Wartchow
Amanita craseoderma Bas, Persoonia 10: 20. 1978.

MycoBank: MB#308547
Typification: BRAZIL. Amazonas, 30 km North of Manaus, 30 Sep 1977, R. Singer B
10160 (L 0053685, isotype – lectotypus hic designatus!).
Fig. 3F–I
From protologue: Basidiomata medium-sized, solitary. Pileus up to 60 mm diam. plano-
convex with subumbilicate centre, very dark brownish gray [M&P 8A8] with pale mar-
ginal grooves; margin broadly sulcate striate (50–60% of radius); context relatively thin
and very fragile, white, unchanging; universal veil as very few, small, dark gray-brown,
irregularly shaped wart-like remnants, in dried state shiny and without fibrillose structure
visible under lens. Lamellae free, fairly crowded, very pale grayish; lamelullae very rare,
(obliquely?) truncate. Stipec. 110 × 7 mm, subcylindric, without basal bulb, fuscidulous
gray, subglabrous to glabrous but under lens in dried specimens darkly punctuate on up up-
f
per part and whit very fine short dark fibrils on lower part; context white, unchanging;
o
partial veil absent; universal veil forming one nearly complete narrow, dark gray-brown,
ro
subfloccose belt up to 6 mm above base and a whitish (near upper margin somewhat gray gray-
ish), adnate subfloccose layer on lower 3 mm of stipe. Odor not distinguishes and taste
p
not recovered.
d_
Basidiospores [50/1/1] (8–)8.3–10(–10.5) × (6.5–)7–8.5(–9.5) µm, ((L = 8.8 µm; W =

te
7.7 µm; Q = (1.00–)1.03–1.31(–1.38); Q = 1.14), inamyloid, hyaline, colorless, globose

to subglobose then broadly ellipsoid occasionally ellipsoid, smooth, thin-walled; apiculus
c
rounded/truncate, subapical to sublateral; contents including a large guttule. Basidia dif-

ficult to localize, up to 27 × 11 µm, clavate, 4-sterigmate, 4.5 µm high; apparently clamp
clamp-
e
less. Subhymenium: unsatisfactory rehydrated, up to 30 µm thick, with 3–4 layers of

rr
isodiametric cells up to 8–17 µm diam.; wst-near = ± 18 µm; wst-far = ± 25 µm. Lamella

o
trama apparently bilateral, with wcs= ± 30 µm (weakly rehydrated), with common fila-
mentous hyphae ± 6 µm, unbranched and common fusoid inflated elements ranging up to
c
80 × 32 µm gradually diverging from central stratum; vascular hyphae scarce 6–10 µm

n
wide, more or less longitudinal to lamella edge. Marginal tissue of lamellae

lamellae: not studied.
u
Pileus context very thin, difficult to delimitate in the type, but distinctly acrophysalidic;
acrophysalides up 100 × 40 µm, inflated clavate, common; filamentous hyphae 3–6 µm
wide, very common, interwoven, forming a loose matrix were other elements occurs;
vascular hyphae ± 22 µm wide, very scarce, appearing only near to pileipellis. Stipe con-
text longitudinally acrophysalidic, with acrophysalides up to 284 × 450 µm, very com-
mon, mostly clavate; filamentous hyphae 2–6 µm wide and having a longitudinal orienta-
tion, unbranching; vascular hyphae 3–20 µm wide, plentiful, unbranched with some
branched or dichotomous and having more or less longitudinal orientation to infrequently
transposing transversally tha stipe. Pileipellis up to 50 µm thick composed by hyphae up
to 5–24 µm, apparently with a periclinal orientation, mostly with brownish pigment; vas-
cular hyphae 5–9 µm wide, abundant; not gelatinized. Universal veil on pileus: with lo-
cally plentiful hyphae 4–12 µm wide, short celled, moderately thick walled (somewhat
ranging to 1 µm thick) or with incrustation, clampless, and globose, subglobose piriform
eschweizerbart_xxx
to ovoid (22–65 × 18–56 µm) cells, brownish colored to somewhat paler, pigment equally
dissolved, walls up to 2 µm thick. Whitish covering of extreme base: abundant filamen-
tous hyphae 3–9 µm, moderately thick walled, common, and fairly abundant inflated
subglobose, ovoid to piriform (30–45 × 30–42 µm), colorless, thick walled (somewhat
ranging to 2 µm thick), very common, organized in tuffs at outer layer, vascular hyphae
absent. Partial veil absent.
Habitat: On yellow-soil hylea in primary “terra firme” forest, with Lecythidaceae, Legu-
minosae and other dicotyledonous trees, few Palmae (Bas 1978).
Known distribution: Amazon Forest of the States of Amazonas, North Brazil.
Observations on exsiccate: The holotype (INPA 280290) is destroyed (Dirce L. Komura,
pers. corresp.). On the other hand, the isotype at L is well preserved and designed here as
lectotype (Art. 9.11, Turland et al. 2018) of A. craseoderma.
Remarks: The most distinct feature of A. craseoderma is the occurrence of inflated ele
ele-
f
ments to 25 µm wide in the pileipellis (Bas 1978).
o
ro
Amanita crebresulcata Bas, Persoonia 10: 18. 1978.
MycoBank: MB#308549
p
d_
Typification: BRAZIL. Amazonas, Manaus, INPA, 19 May 1977, R. Singer B 9680
(INPA 66710 holotype!; L 0053686 isotype – lectotypus hic designatus!
designatus!);
te

Figs. 4A–I, 5A–B
c
Basidiomata medium-sized, slender, solitary or sub gregarious. Pileus 19–70 mm, in the
e
young specimen campanulate with rather acute umbo but somewhat hemispheric with
rr
rounded obtuse apex (then without umbo), later plano-convex to flat or plane-concave
o
with low obtuse umbo or depressed at centre, dark umber to fuscous sepia or at umbo or
dark grayish brown [M&P 16A3 “Phantom”, OAC 836 or darker than OAC 832] centre
c
(in one young specimen fuliginous [W ‘cigar brown 16’], somewhat pale around umbo
n
but with narrow darker fuscous zone (in dried specimens examined this zone has disap disap-
u
peared) at margin dark grayish brown [OAC 832, ‘W sepia 26’] in one young specimen
or slightly paler sulcate zone, glabrous, viscid, in dried specimens matt to somewhat
shiny (particularly at centre) and without or with very fine innate fibrillose structure un-
der lens; margin broad densely sulcate (in dried specimens 14–16 grooves per 10 mm of
circle half way sulcate zone); context white to whitish; universal veil absent in the great
majority of basidiomata, but in one (INPA 82341) as a single patch up to 3 mm in diam.,
concolorous with the universal veil at stipe base. Lamellae free, crowded, narrow or up to
5 mm, white, with concolorous edge or (more often?) with a very thin brown line along
the entire edge or only near margin cap; lamellulae rather scarce, uncommon truncate.
Stipe 50–125 × 5–10 mm, tapering upward or more or less cylindrical, hollow, with white
base and often paler apex or pale cream [M&P 10B1 Oyster White], glabrous and smooth,
at upper half to apex sometimes minutely innately punctulate or covered with brownish/
fuliginous fibrils; context hollow or stuffed, white to whitish, unchanging, central cylin-
eschweizerbart_xxx
18 Felipe Wartchow
f
o
ro
p
d_
te
c
e
rr
o
c
n
u
Fig. 4. A–C. A. crebresulcata (lectoype). A. Hymenium and subhymenium. B. Basidiospores. C.

Elements of the universal veil from stipe base. D. A. crebresulcata (INPA 82341). Hymenium,
subhymenium and divergent elements of the lamella trama. E–F. A. crebresulcata (URM 80122). E.
Hymenium, subhymenium and divergent elements of the lamella trama of the immature basidiome.
F. Hymenium, subhymenium and inflated cells of mature basidiome. G–I. A. crebresulcata (INPA
82341). G. Himênio e subhimênio. H. Basidiosporos. I. Elements of the universal veil from pileal
patch. Scale bars are 10 µm.
eschweizerbart_xxx
der up to 5 mm in diam.; partial veil absent; universal veil saccate at stipe base, thin or
membranous, white to cream grey or cream [M&P 15C7 Racquet] or in places pale buff,
narrowly sheathing, higher than broad up to 10–25 × 14–15 mm, felted (not all powdery
or flocculose), under lens locally with minute buff patches or glabrous, volva in dried
specimens adnate only to lower 5–10 mm of stem and (when visible) with a narrow,
fringed internal limb on inner surface 2–3 mm high above place of attachment to stem,
but much more visible in young fresh specimen. Odor unpleasant and taste not recovered.
Basidiospores [213/9/8] (7.3–)7.7–11(–11.5) × (6.5–)6.8–9(–9.5) µm, (L = 8.7–9.4(–9.6)
µm, L’ = 9.1 µm; W = 7.6–7.9(–8.2) µm, W’ = 7.8 µm; Q = (1.05–)1.06–1.34(–1.40); Q
= 1.14–1.20, Qm = 1.16), inamyloid, hyaline, colorless, subglobose to broadly ellipsoid
to ellipsoid, thin-walled, smooth; apiculus sublateral to subapical; contents as large gut-
tule. Basidia 35–40 × 10–11.5 µm, clavate, bearing four sterigmata, each 5–8 µm high;
clampless. Subhymenium When immature: about 18 µm thick, with exclusively unin-
flated elements somewhat slender clavate e.g. 12 × 9 µm, ramose to inflated ramose.
f
When mature: very weakly rehydrated, impossible to study in some exsiccates, but in
o
recent collections obviously cellular, up to 23 µm thick, with 3–4 layers of isodiametric
ro
elements 13–15 µm diam.; wst-near = 20–37 µm; wst-far = 24–45 µm. Lamella trama
bilateral: when immature wcs = 17 µm.; when mature with wcs = 23–34 µm (difficult to
rehydrate in older exsiccates), inflated divergent terminal elements up to 50 × 21 µm wide
p
d_
gradually diverging, frequent; filamentous hyphae 2.5–9 µm, straight, and somewhat di- di
chotomous branched and also gradually diverge from central stratum; vascular hyphae
te
moderately scarce 4.2–5.4 µm, rarely branched. Marginal tissue of lamellae sterile; com-
posed of common filamentous hyphae 3.5–7 µm wide, collectively parallel to lamella
c
edge or somewhat interwoven; inflated cells hyaline, colorless, inamyloid, terminal cells
e
22 × 15 µm. Pileus context very difficult to rehydrate in older exsiccates, but appearing
to be distinctly acrophysalidic; acrophysalides 84–100 × 23–33 µm; filamentous hyphae
rr
2.5–12 µm wide, plentiful; vascular hyphae range 9–20 µm wide, moderately frequent.
o
Stipe context longitudinally acrophysalidic, with acrophysalides up to 200–370 × 35–

c
75 µm; filamentous hyphae 2–5 µm wide, having a longitudinal orientation, unbranching;

n
vascular hyphae 5–15 µm wide, moderately abundant, unbranched with some branched
or dichotomous and having more or less longitudinal orientation. Pileipellis a cutis 120–
u
180 µm on which appears to range to 195 µm at centre in some cases, difficult to differ-
entiate the two layers in older collection, but in some points easily to observe, mostly at
centre (in the isotype it is well differentiated); suprapellis as less pigmented ixocutis ap-
parently ranging up to 10–15 µm in some collection but to 40 µm in the isotype, mainly
at centre composed by distant filamentous hyphae up to 3 µm, radially oriented immerse
in dense gelatinous matrix; subpellis 110–180 µm thick at centre; compressed filamen-
tous hyphae up to 3 µm with periclinal orientation dark brown pigmented but somewhat
palling to yellowish brown; vascular hyphae 5–17 µm wide, common, not branching,
transposing the all the thickness. Universal veil on pileus: absent in the majority of the
specimens examined, but only one as abundant filamentous hyphae 2–5 µm wide, very
interwoven, with some incrustation and mixed with some soon collapsing colorless sub
globose cells up to 42 × 37 µm; On stipe base: at internal limb filamentous hyphae 5–
8 µm, sometimes with incrustation, loosely interwoven and mixed with fairly abundant
eschweizerbart_xxx
20 Felipe Wartchow
f
o
ro
p
d_
te
c
e
rr
o
c
n
u
Fig. 5. Amanita crebresulcata (URM 80122). A. Basidiomes. B. Longitudinal section of the stipe base
of the young basidiome showing the limbus internus. Scale bar is 10 mm.
eschweizerbart_xxx
inflated cells sub globose, ovoid and piriform (e.g. 43–80 × 32–63 µm) or rarely broadly
clavate cells (e.g. 72–81 × 40–42 µm), colorless, frequently immerse in a gelatinous ma-
trix (not very evident in the isotype) with some patches of yellowish brown extracellular
granular matter, mainly at upper part of sac; in outer surface the elements appears to be
organized in tuffs. Partial veil absent.
Habitat: ‘fairly common on the soil in mixed secondary tropical rain forest, growing near
Neea (Nyctanginaceae) with one case of mycorrhizal connection definitely established
with Psychotria (Rubiaceae) in Amazon’ (Bas 1978); in mature Atlantic Forest fragments
among trees of Euphorbiaceae, Leguminosae, Mimosaceae, Flacourtiaceae, Sapotaceae
and others (Alves-Araújo et al. 2008); in “capoeiras” i.e. secondary vegetation of about
12 years regeneration (Kimmel et al. 2010) which are more sensitive to anthropogenic
influence and originated from mature forests (Trindade et al. 2008, Schessl et al. 2008).
Known distribution: Amazon Forest, State of Amazonas and Atlantic Forest of Pernam
Pernam-
buco (IBGE 2004, 2012).
f
o
Additional material examined: BRAZIL. Amazonas, Manaus, 27 May 1977, R. Singer
ro
B 9729 (INPA 66711, paratype!); ); in same place, 1977, R. Singer B 9796 (INPA 66712,
paratype!);); Aleixo’s Road, INPA Campus, 9 Jan 1978, R. Singer B 10477 (INPA 77199
p
as “cf. Amanita dunicola Guzmán”); INPA, 25 Nov 1978, R. Singer B 11395 (INPA
d_
82341); in same place, 14 Oct 1978, R. Singer B 11415 (INPA 82350); Pernambuco,
Araçoiaba, Usina São José, Engenho Campinas, 05 Jun 2007, A.S. Santana & D.K.A.
te
Silva s.n. (URM 80122); Araçoiaba, Usina São José, Engenho Cumbé de Baixo, 05 Jun
2007, A.S Santana & D.K.A. Silva s.n. (URM 80123); Igarassu, Usina São José, Mata dos
c
Macacos, 29 Jul 2005, F. Wartchow 27/2005 (URM 78690); Recife, Jardim Zôo-Botânico
e
Dois Irmãos, 31 Aug 1951, J. Nery s.n. (IPA 2874, as ‘Amanitopsis

‘ volvata’)
rr
Observations on exsiccate: The holotype (INPA 66710) is destroyed, remaining small

fragments of pileus and stipe (https://specieslink.net/rec/126/280290), while the isotype
o
is well preserved. According to code, if the holotype has been lost or destroyed (in this
c
case), the isotype must be designed as lectotype (Art. 9.11, Turland et al. 2018).
n
heav-
In almost all collection any trace of velar remnants on pileus was observed due the heav
u
ily gelatinized pileipellis (Bas 1978), except the single velar patch found in the exsiccate
‘R. Singer B 11395’ (INPA 82341). It might occur due the insolation or dry climate condi-
tions, decreasing the gelatinous suprapellis and making the universal veil more adherent
to the adjacent pileipellis (Bas 1969: 310).
The material ‘R. Singer B 10477’ identified as “cf. Amanita dunicola Guzmán” lacks
remnants of universal veil on pileus, presents broadly sulcate striate (more than 50% of
radius) pileus margin, sac-like universal veil at stipe base and basidiospores [15/1/1]
(7.5–)8–10 × (6.5–)7.5–8.5 µm, (L = 9.1 µm; W = 7.6 µm; Q = 1.12–1.28(–1.30); Q =
1.20). Amanita dunicola was described with smooth to short sulcate marginal zone and
presence of small velar patches scattered on pileus surface, infrequent clamps in the ba-
sidia, and basidiospores [60/3/1] (8.5–)10–13.9(–15) × (6–)7.2–9.5(–11.2) µm, L = 11–
eschweizerbart_xxx
22 Felipe Wartchow
12.8 µm, L’ = 11.8 µm; W = 8–9.1 µm, W’ = 8.6 µm; Q = (1.17–)1.25–1.56(–1.67); Q =

1.35–1.41, Qm = 1.38 (Guzmán 1982, Tulloss 1994).
Remarks: This species is well characterized by the strongly gelatinized pileipellis since
the beginning of the basidioma expansion (Bas 1978, Wartchow & Maia 2007).
Amanita sp. 1
Fig. 6A–D
Basidiomata small, apparently gregarious. Pileus ranging to 20 mm in exsiccate (20–
30 mm in the drawings), gray, plane-depressed; margin pectinate about 1/3 ratius;
universal veil as few small wart-like (floccose patches) remnants scattered on surface,
with color Chicle to Picassimu. Lamellae free but not distant from stipe, white to whit-
ish, crowded to subcrowded; lamellulae rare, attenuate. Stipe ranging to 35 × 2.5 mm in
exsiccate (31–43 × 3–4 mm in the drawings), distinctly tapering upward, mostly inflate
but not truly bulbous at base; universal veil as floccose belt 7 mm above base,
base but not
f
distinguished at exsiccate and apparently a small cup like structure at very base.
o
ro
Basidiospores [70/3/1] (8.5–)9–12(–12.5) × (8–)8.5–10.5(–11) µm, ((L = 9.9–11 µm, L’ =
10.4 µm; W = 8.8–10.1 µm, W’ = 9.3 µm; Q = (1.00–)1.04–1.37(–1.47); Q = 1.09–1.14,
p
Qm = 1.13), inamyloid, hyaline, colorless, subglobose to broadly ellipsoid only occasion
occasion-
d_
ally ellipsoid, smooth, thin-walled; apiculus sublateral to lateral; contents difficult to
locate. Basidia 38–50 × 15–16.5 µm, 2 or 4 sterigmata, each ranging to 7 µm high, clamp
clamp-
te
less. Subhymenium weakly rehydrated difficult to analyze, apparently cellular; appearing

to range 30 µm thick with 3 layers of cells measuring up to 14 × 13 µm; wst-near =
c
± 30 µm; wst-far = ± 55 µm. Lamella trama difficult to rehydrate; wcs= ± 30 µm wide;

e
filamentous hyphae 3.5–7(–9) µm not fully longitudinal to lamella; clavate elements

rr
± 27 × 13 µm, rare. Pileus context difficult to rehydrate; acrophysalides approx. 70 ×

30 µm, difficult to distinguish; filamentous, hyphae 3–6 µm, interwoven, common; vascu-
vascu
o
lar hyphae 4–15 µm, frequent; slightly gelatinized. Stipe context moderately well rehy-
rehy
c
drated; acrophysalides 270 × 35 µm; filamentous hyphae 3–7 µm, frequent; vascular hy-
n
phae 3–8 µm, very scarce. Pileipellis: up to 80 µ at centre; differentiated in two layers;
u
filamen
subpellis a ixocutis up to 15 µm thick; comprising on radially arranged very thin filamen-
tous hyphae immersed withing a extracellular (gelatinous?) matrix; suprapellis up to
70 µm with plentiful interwoven filamentous hyphae ranging to 3–15 µm wide, infre-
quently inflating to 20–30 µm, radially arranged, pale brown to colorless; vascular hyphae
rare, up to 5 µm. Universal veil from stipe base: formed by scarce interwoven tuffs of
globose, subglobose to piriform (e.g. 27–40 × 20–32 µm) and somewhat more or less el-
lipsoid (e.g. 37 × 25 µm) or clavate (e.g. 45 × 23 µm) cells, pale grayish or colorless,
abundant; filamentous hyphae 4–8 µm somewhat slightly colorless incrustations. On pi-
leus: abundant more or less isodiametric to piriform terminal disordered cells ranging to
20–50 µm, mostly with yellowish brown to brown pigment but somewhat paler, wall c.
1 µm thick; abundant; filamentous hyphae 3-6 µm, mostly colorless, common; vascular
hyphae absent. Partial veil absent.
eschweizerbart_xxx
f
o
ro
p
d_
te
c
e
rr
o
c
n
u
Fig. 6. A–D. Amanita sp.1. A. Basidiospores. B. Hymenium and subhymenium. C. Elements of the
universal veil from pileal wart. D. Elements of the universal veil from stipe. E–G. Amanita sp.2. E.
Basidium and subhymenial cell. F. Basidiospores. G. Elements of the universal veil from pileus. Scale
bars are 10 µm.
eschweizerbart_xxx
24 Felipe Wartchow
Habitat: On soil in igapó type Amazonian forest, a black-water inundated type forest that
has ectotrophic trees (Singer & Aguiar 1986).
Known distribution: Amazon Forest of the State of Amazonas, North Brazil.
Material examined: BRAZIL. Amazonas, Manaus, Igarapé do Tarumãzinho, on soil,
7 Jan 1979, R Singer B 11436 (INPA 82932).
Remarks: These materials macroscopically resemble A. craseoderma due to the pres-
ence of small, friable wart-like remnants of universal veil on pileus, and the floccose belt
universal veil elements in stipe base. However, Amanita sp. 1 differs in the slightly in-
flated stipe base, larger basidiospores, exclusively narrow hyphae in the pileipellis and
presence of 2-spored basidia. According to Bas (Bas 1969: 320), basidiospores from 1- to
3-spored basidia tend to be larger and more voluminous than in 4-spores basidia. The
presence of two sterigmata in some basidia might be good taxonomic feature of Amanita
sp. 1. This entity probably represents an undescribed taxon, but fresh well preserved adad-
f
ditional specimens are needed before to describe it as new species.
o
The size of the basidiospores of this exsiccate suggest phenetically similarity with A.
ro
craseoderma sensu Pegler (1983)) from Martinique, who reported basidiospores 8.5–12 ×
7.5–8.5 µm, L = 9.7 µm; W = 8 µm; Q = 1.20. p
d_
Amanita sororocula Tulloss, Ovrebo & Halling from Colombia also has globose to
subglobose basidiospores (8.8–)9.7–12.8(–16.8) × (8–)8.8–12(–15.5) µm, L = 10.2–
te
12.1 µm, L’ = 11 µm; W = 9.3–11.5 µm, W’ = 10.2 µm; Q = (1.00–)1.01–1.16(–1.25);

Q = 1.05–1.10, Qm = 1.07, but differs at least in the normal pileipellis and common vas
vas-
c
cular hyphae at universal veil (Tulloss 1992).

Tulloss et al. 1992
e
Amanita calopus also presents a small swollen or “small but distinct marginate” base
rr
(Pegler
Pegler & Shah-Smith 1997)
1997) at the base of the stipe, but differs in the slightly larger and
longer basidiospores [20/1/1] 7.8–13.6(–15.1) × (5.5–)6.0–8.8(–9.0) µm, L = 11.7 µm,
o
W = 7.7 µm, Q = (1.30–)1.39–1.60(–1.78); Q = 1.53, and pileipellis with narrower hy-

c
phae (Tulloss 2023).

Tulloss & Yang 2023
n
u
Amanita sp. 2
Fig. 6E–G
Basidiome solitary, slender. Pileus up to 20 mm, plano-umbonate, gray to blackish
brown (‘atrobrunneus’) at centre, viscid; margin pectinate at 1/3 of ratius; universal
veil absent. Lamellae free, white, crowded; lamellulae present, but apparently rare, at-
tenuate. Stipe ranging to 80 × 2 mm, indistinctly tapering upward, without bulb, white,
glabrous; partial veil absent; universal veil saccate, white, up to 10 mm high, strongly
incrusted by soil particles.
Basidiospores [30/1/1] (9–)9.5–12.5(–13) × (6.5–)8–10.5(–11) µm, (L = 10.6 µm; W =
9.2 µm; Q = (1.05–)1.07–1.30(–1.44); Q = 1.15), inamyloid, hyaline, colorless, subglo-
bose to broadly ellipsoid only occasionally ellipsoid, thin-walled, smooth, apiculus sub-
lateral to lateral. Basidia up to 33 × 12.5 µm, clavate, 3-sterigmata each ± 2 µm high,
eschweizerbart_xxx
probably clampless. Subhymenium difficult to rehydrate, but cellular bearing subisodia-

metric cells ± 20 × 17 µm; wst-near = not recorded; wst-far = ± 20 µm. Lamella trama
unsatisfactorily rehydrated; wcs = ± 28 µm thick; filamentous hyphae ± 6 µm. Pileus con-
text poorly rehydrated; acrophysalides difficult to distinguish; filamentous hyphae 3–
6 µm, interwoven, common; vascular hyphae not observed. Stipe context moderately well
rehydrated; acrophysalides up to 300 × 45 µm; filamentous hyphae 3–6 µm, frequent;
vascular hyphae 3–7 µm, very scarce. Pileipellis up to 60 µm at centre, with filamentous
hyphae about 3–5 µm wide, radially arranged, colorless; vascular hyphae not observed;
non- or only weakly gelatinized. Universal veil on pileus: absent. On stipe base: difficult
to rehydrate due to soil incrustation; scarce interwoven globose, subglobose to piriform
cells e.g. 42 × 37 µm, hyaline, difficult to observe; filamentous hyphae 2–5 µm, very
common. Partial veil absent.
Habitat: Apparently solitary on soil in an igapó Amazonian forest.
Known distribution: Amazon Forest of the State of Amazonas, North Brazil.
f
o
Material examined: BRAZIL. Amazonas, Manaus, Igarapé do Tarumãzinho, 7 Jan 1979,
ro
R. Singer B 11482 (INPA 82936).
Observations on exsiccate: Due to the poor preservation of the specimen, details of the
p
hymenium, subhymenium and lamellatrama were impossible to observe. As seen with
d_
Amanita sp. 1, Amanita sp. 2 also might correspond to an undescribed species, but this
hypothesis will be confirmed only by additional collections.
te
Remarks: Among the Neotropical Amanita with saccate volva at stipe base, this material
c
can be compared with A. crebresulcata and A. dunicola,

dunicola differing at least in the blackish
e
brown pileus (according to Singer’s field in the herbarium F), larger basidiospores and the
rr
lack of distinctly gelatinized suprapellis.

Another South American taxon with dark brown to blackish pileus centre is A. fuligineo-
o
disca Tulloss, Ovrebo & Halling from the Andean Colombia, but it differs in the globose
c
to subglobose basidiospores [585/29/12] (7.5–)9–12(–15.5) × (6.5–)8.2–11.2(–15) µm,

n
L = (9.5–)9.6–11.4(–11.5) µm, L’ = 10.6 µm; W = (8.6–)8.9–10.6(–10.7) µm, W’ =

u
9.9 µm; Q = (1.00–)1.02–1.18(–1.40); Q = (1.05–)1.06–1.12(–1.15), Qm = 1.08, absence

of gray tones on pileus and the association with Quercus (Tulloss et al. 1992).
Amanita sp. 1 and Amanita sp. 2 were collected in an “Igapó Forest”, a periodically inun-
dated area in Amazonia, dominated by leguminous trees, particularly Aldinia latifolia and
Swartzia cf. polyphylla in soil with low mineral nutrient content (Singer & Aguiar 1986,
Parolin et al. 2004).
eschweizerbart_xxx
26 Felipe Wartchow
Subgenus Lepidella section Lepidella
Amanita grallipes Bas & de Meijer, Persoonia 15: 345. 1993.

MycoBank: MB#361022
Fig. 7A–C
From protologue: ‘Basidiomata large and slender, solitary to subgregarious. Pileus 22–
90 mm, from hemispherical or conico-convex to plano-convex or flat without umbo,
uniformly dark brown to somewhat paler grayish brown, dry; margin entire, smooth, ap-
pendiculate when young; context fleshy, up to 10 mm thick near centre and up to 6 mm
thick above midpoint of lamellae, white, unchanging; universal veil as scattered, dark
brown to paler grayish brown, adnate, pyramidal warts. Lamellae free, very crowded,
fairly broad (up to 11 mm), first pure white then cream and finally yellow to golden yel-
low, with concolorous, even edge; lamellulae attenuate. Stipe 50–120 × 6–11 mm (includ-
ing bulb) then 10–18 mm wide at broadest part just above the soil, white, solid, dry; bulb
rooting 20–45 × 10–18 mm tapering downwards; context fleshy, white, unchanging ex ex-
f
o
cept for a slight yellowing on the base of stipe after bruising; partial veil present at first,
apical, pendulous, rather thick felted-membranous, white and smooth below and above,
ro
at margin with brown pyramidal warts similar to those on pileus, fugacious and soon
disappearing; universal veil as small, erect to appressed, pale yellowish brown, floccose
p
d_
scales. Odor not distinguished. Taste mild’.
Basidiospores [20/1/1] (6.5–)7–8(–8.5) × (6–)6.5–7.5(–8) µm, ((L = 7.9 µm, W = 5.7 µm,
te
Q = (1.24–)1.26–1.40(–1.49); Q = 1.34), broadly ellipsoid to ellipsoid, smooth, thin

walled, usually or at least somewhat adaxially flattened; apiculus sublateral, small, ob ob-
c
tuse; somewhat with contents guttulate, amyloid, colorless, hyaline. Basidia very scarce
e
up to 32 × 11 µm, 4-sterigmate, each up to 3 µm long, clamps probably present at base,

rr
but difficult to observe. Subhymenium poorly rehydrated, up to 32 µm thick, ranging to

3 layers more or less isodiametric cells (e.g., 9 × 8 µm), but not rehydrating very well;
o
wst-near = approx. 30–50 µm; wst-far = 45–68 µm. Lamella trama moderately rehy-
c
drated, but distinctlybilateral; wcs = 45 µm; filamentous hyphae up to 2.5–5 µm wide, with
n
elements gradually diverging from central stratum and somewhat rather abrupt and more
u
or less perpendicular to hymenium, inflating as clavate cells up to 40 × 18 µm or some-

what ranging to 25 µm wide; vascular hyphae absent. Pileus context moderately rehy-
drated, apparently acrophisalidic; acrophysalides 75 × 30 µm, inflated clavate, thin
walled; filamentous hyphae 2.5–4 µm wide, appearing to be branched, interwoven, form-
ing a loose matrix where other elements occur; vascular hyphae not seen; moderately
gelatinized. Stipe context moderately rehydrated, longitudinally acrophysalidic; acrophy-
salides 150–260 × 25–60 µm, clavate to somewhat subfusiform, plentiful; filamentous
hyphae 3–15 µm wide, longitudinally oriented, somewhat branched; vascular hyphae 15
or 35 µm wide, very scarce, unbranched; clamps somewhat present at septa. Pileipellis
difficult to differentiate. Universal veil on pileus: absent in material studied, but a single
rounded brown colored and thick walled rounded cell 14 × 12 µm observed. On stipe:
very scarce, with elements apparently arising from context, long clavate to subfusiform
e.g. 85–150 × 20–30 µm, colorless; filamentous hyphae up to 5 µm wide, very scarce,
eschweizerbart_xxx
f
o
ro
p
d_
te
c
e
rr
o
c
n
u
Fig. 7. A–C. A. grallipes (Meijer 2921). A. Basidiospores. B. Basidium and subhymenium. C. Universal
veil from stipe. D. Amanita strobiliformis (PACA 12221). Basidiospores. E–H. A. campinaranae
(Holotype). E. Universal veil from stipe. F. Basidiospores. G. Hymenium, subhymenium and divergent
elements from central stratum. H. 53. Universal veil from pileus. I. Amanita phaea. Universal veil from
pileus. Scale bar is 10 µm.
dichotomously branched, thin walled; vascular hyphae absent. Partial veil absent in the
studied material.
Habitat: Found on humus in mixed in mixed ombrophilous forest with or without Arau-
caria angustifolia, at 900 m altitude.
eschweizerbart_xxx
28 Felipe Wartchow
Known distribution: Known only from the State of Paraná, South Brazil.
Material examined: BRAZIL. Paraná, Curitiba, Bariqui Park, 6 Nov 1994, A.A.R de
Meijer 2921 (MBM).
Observations: Although de Meijer (2001) reported that A. grallipes grows in large num-
ber of basidiomes somewhat forming a conspicuous fairy ring, only a single very poor
preserved material was sent for analysis.
Remarks: Amanita grallipes is known only from the type locality, growing gregariously
sometimes forming a large fairy ring (Bas & de Meijer 1993, de Meijer 2001, 2008). It
belongs to stirps Vittadiniae due to the presence of clamp-connection in the basidia,
broadly ellipsoid to ellipsoid basidiospores and the velar remnants evenly distributed over
all stipe below the partial veil (Bas 1969, Bas & de Meijer 1993). The species readily dif-
fers from the other species of this group in the dark brown pileus, distinctly yellow lamel-
lae and rooting stipe (Bas & de Meijer 1993). ). They reported that A. silvifuga Bas from
f
North America is the most phenetically similar taxon, differing in the paler pileus, non-
o
rooting stipe and lack of yellow lamellae.
ro
Amanita savannae Tulloss & Franco-Mol. from Colombiia also has brownish tints, but
clearly differs in the considerably slender basidiome with pileus 20–30 mm in diam. and
p
stipe 13–30 × 4–7 mm, white lamellae, and slightly larger basidiospores [140/7/1] (6.5–)
d_
7.2–10.8(–13.5) × (6.0–)6.2–8.5(–9.5) μm (L L = 7.6–9.2(–10.2) μm; L’ = 8.6 μm; W =
6.6–7.6(–8.2) μm; W’ = 7.2 μm; Q = (1.02–)1.08–1.36(–1.73); Q = 1.14–1.22(–1.27);
te
Q’ = 1.19) (Tulloss
Tulloss & Franco-Molano 2008).
2008).
c
Amanita spissa var. laeta Rick,, Broteria Ser. Bot. 5: 25. 1906.
e
rr
Observation: The specimen description and the neotypification status can be found in
Wartchow (2021).
o
Additional remarks: Amanita spissa (Fr.) Bertill. belongs to subg. Amanitina (E.-J. Gil-
c
bert) E.-J. Gilbert sect. Validae (Fr.) Quél. (Cui et al. 2018) and corresponds to a synonym
n
of A. excelsa (Fr.) Bertill. ((Tulloss & Yang 2023). It is characterized by the non-appendic-
u
ulate pileus margin, globose to emarginated stipe base and universal veil composed of
globose inflated elements (Bas 1969, Neville & Poumarat 2004, Cui et al. 2018).
The neotype of A. spissa var. laeta, on the other hand, is a specimen of the subg. Lepidella
subsect. Vittadiniae Bas emend. Tulloss et al. stirps Vittadiniae (Bas 1969, Tulloss et al.
2016, Cui et al. 2018). It is characterized as having non-striate pileus margin, amyloid
basidiospores, lack of membranous volval limb at stipe base, the non-bulbous stipe, the
dominance of elongate-fusiform cells on universal veil remnants on pileus, the apparent
presence of clamp connections (although difficult to locate in the basidia due to poorly
preserved hymenium), and the apparent distribution of the velar remnants over all stipe
length (Wartchow 2021).
Regarding the basidiome color, Bas (1969) suspected that Rick’s description is close to
A. singeri Bas. However, the thin and almost inconspicuous universal veil on stipe, and
eschweizerbart_xxx
the distinctly narrower basidiospores [178/10/7] (6–)7.5–11.0(–15.4) × (4.5–)4.9–7.5

(–9.5) µm, L = (7.1–)7.6–10.0 µm; L’ = 9.1 µm; W = (4.9–)5.6–7.2 µm; W’ = 6.6 µm;
Q = (1.12–)1.21–1.60(–2.00); Q = 1.23–1.42(–1.60); Q’ = 1.39 (Bas 1969, Tulloss &
Yang 2023) separate A. singeri from the holotype designated by Wartchow (2021).
Although lacking information regarding to collection and ecology, Rick’s material is
probably related to A. vittadinii (Moretti) Vittad. (Wartchow 2021). The indistinct pileipel-
lis and the concentrically arranged appressed to recurved membranous scales on stipe
below the partial veil (Rick 1906) are similar as observed in A. vittadinii. On the other
hand, A. vittadinii differs in the white and distinctly larger basidiomes and subcylindrical
stipe (Moretti 1826: 66-67, Bas 1969, Vizzini et al. 2017). Also, A. vittadinii is an Euro-
pean taxon occurring in field open woods and parkshaving medium-sized to large pileus
75–170 mm, absence of filamentous hyphae in the universal veil (Bas 1969) and slightly
larger basidiospores [20/1/1] 9.5–12(–14) × (7.2–)7.5–9.3(–10) μm, L = 11 μm; W =
8.2 μm; Q = (1.17–)1.19–1.44(–1.47); Qm = 1.34 (TullossTulloss & Yang 2023).
2023).
f
Amanita prairiicola Peckfrom USA and Argentina primarily differs in the distinctly
o
longer and more elongated basidiospores [700/34/11] (8.0–)10–14(–19.2) × (5.2–)
ro
6.4–10(–12.2) µm, L = (10.4–)11–13.3(–13.7) µm; L’ = 12 µm; W = (6.5–)6.6–9.6
(–9.7) µm; W’ = 8 µm; Q = (1.09–)1.26–1.82(–2.31); Q = (1.30–)1.33–1.75(–1.85); Qm =
p
1.51 (Tulloss & Yang 2023)) and the presence of rather distinct pileipellis (Bas
( 1969).
d_
Amanita grallipes was suspected to be conspecific with A. spissa var. laeta (Bas & de
te
Meijer 1993),), but readily differs in the brown pileus and distinctly smaller basidiospores
[50/4/2] (7.2–)7.5–9.6 × 5.6–6.9(–7.5) µm (Q = (1.15–)1.20–1.50(–1.60); Q = 1.25–1.40)
c
(Bas & de Meijer 1993, de Meijer 2008)) segregate both taxa.

e
Amanita lilloi Singer, known from Argentina ((Singer & Digilio 1951, Bas 1969) and
rr
Brazil (Wartchow et al. 2007)) shares in the presence of pileal pyramidal wart-like velar
remnants and the narrowing downward stipe. But these similarities are only superficial
o
since it presents whitish basidiomes and considerably smaller basidiospores [100/5/2]

c
(7–)7.5–9.5(–10.5) × (6–)6.5–7.5(–8) µm, L = 8–8.6 µm, L’ = 8.3 µm, W = 6.8–7 µm,

n
W’ = 7 µm, Q = (1.11–)1.14–1.36(–1.43); Q = 1.16–1.22, Qm = 1.20 (Wartchow et al.

u
2007).
Even so, basidiospore size helps to segregate A. spissa var. laeta from A. grallipes, A. lil-
loi, A. vittadinii and A. prairiicola. This material probably corresponds to an undescribed
species among subsect. Vittadiniae, but additional collections are needed before describ-
ing it as a new taxon.
eschweizerbart_xxx
30 Felipe Wartchow
Subgenus Amanitina section Validae
Amanita campinaranae Bas, Persoonia 10: 4. 1978.

MycoBank: MB#308543
Fig. 7E–H
From protologue: ‘Basidioma medium-sized, solitary, rather slender. Pileus about 60 mm
wide in mature specimens, convex, later with depressed centre, white to pale grayish,
viscid, without distinct fibrillose structure; margin smooth non-appendiculate; context
white, thin, unchanging; universal veil as dense, gray, sub felted crust-like remnants at
centre but forming isolates patches on limb. Lamellae free, crowded, often forked in both
directions, narrow, white turning cream to yellow when freshly dried, in exsiccates with
entire more or less concolorous edges; lamellulae attenuate. Stipe 77 × 8 mm at apex to
10 mm (above bulb), slightly tapering upward, pale gray, glabrous; bulb rather abrupt,
globose, 20 mm wide, dirty ochre and white; context white, unchanging, stuffed in bulb;
f
partial veil apical, thin-membranous, pendulous, with white and smooth upper side and
o
very pale gray underside; universal veil as grayish subflocculose-subverrucose rim on
ro
transitional zone between bulb and stipe, giving bulb a submarginate appearance. Odor
lacking and taste not recovered’. p
Basidiospores [50/1/1] (5–)5.5–7(–7.5) × (4.5–)5–6.5(–7) µm, ((L = 6.2 µm; W = 5.8 µm;
d_
Q = (1.00–)1.05–1.16(–1.18); Q = 1.08), amyloid, hyaline, colorless, globose to subglosubglo-
bose only occasionally broadly ellipsoid, smooth, thin walled; apiculus rather broad then
te
attenuate at apex, subapical or sublateral; contents as one large guttula. Basidia 15.5–
23 × 6.5–9 µm, clavate, bearing four sterigmata 0.5–1.5 µm high, clampless. Subhyme Subhyme-
c
nium cellular, moderately well rehydrated, up to 33 µm thick, as 2–4 layers of more or

e
less isodiametric cells up to 9–21 × 9–18 µm, colorless, thin walled; wst-near = 21–
rr
36 µm; wst-far = 32–50 µm; with basidia arising only one layer before the terminal ele ele-
ment. Lamella trama bilateral, perfectly rehydrated, with wct= 36 µm, plentiful terminal
o
inflated clavate elements 45–55 × 18–26 µm, gradually then more or less abruptly diverg-
c
ing for central stratum, becoming an angle higher than 60°, mixed with common filamen-
filamen
n
tous hyphae 5–9 µm; vascular hyphae absent. Marginal tissue of lamellae sterile; consist
consist-
u
ing of scattered, usually solitary layer of globose cells e.g. 15–23 × 14–21 µm, colorless,
slightly thickness walls. Pileus context weakly rehydrated, with plentiful acrophysalides
e.g. 21 × 11 µm mostly inflated clavate but somewhat clavate; filamentous hyphae 2.5–
8 µm, plentiful, strongly interwoven, branching. Stipe context longitudinally acrophysa-
lidic, with acrophysalides 300 × 42 µm, common; filamentous hyphae 3–10 µm, longitu-
dinally oriented or somewhat branching and colorless then ranging to 10–13 µm near the
surface, commonly with condensed brownish pigment; vascular hyphae 5.5–11 µm,
scarce pileipellis a cutis up to 75 µm thick at centre, easily differentiated into two layers;
suprapellis an ixocutis approx. 23 µm at centre with plentiful filamentous hyphae 2–5 µm
wide, mostly radially oriented turning more distant at upper surface, colorless; subpellis
approx. 45 µm thick at centre with very condensed hyphae 2.5–4 µm, pigment appearing
difficult to localize, more or less radially oriented; vascular hyphae common, scattered
5–15 µm; strongly gelatinous. Universal veil on pileus: plentiful subglobose, sphaerope-
eschweizerbart_xxx
dunculate, ovoid to broadly clavate or piriform 30–82 × 23–62 µm or some clavate termi-
nal cells (e.g. 75 × 32 µm), with brownish vacuolar pigment and slightly thickness walled,
with erect or suberect orientation; abundant filamentous then briefly inflated hyphae
ranging to 20 µm; vascular hyphae up to 10 µm, unbranched, straight, uncommon; im-
mersed on a dense gelatinous matrix. Between stipe base and bulb: plentiful ovoid to al-
most isodiametric terminal cells 30–90 × 23–75 µm, somewhat clavate (e.g. 60 × 25 µm)
or ellipsoid (e.g. 80 × 37 µm), with pale brown or pale gray vacuolar pigment and thick
walled (up to 2 µm); weakly incrusted hyphae 3–10 µm, mostly pale gray, plentiful, im-
mediately below the cells, somewhat branched; filamentous hyphae 2–5 µm, also plenti-
ful, mixed with incrusted hyphae, thin walled, colorless. Partial veil not seen.
Habitat: Terrestrial in campinaranae vegetation under Leguminosae and Sapotaceae in
Amazonas, forming ectomycorrhizae (Bas 1978); in monodominant Dicymbe forest from
Guyana (Henkel et al. 2002).
Known distribution: State of Amazonas, North Brazil and Guyana (Henkel 2002).
(Henkel et al. 2002
f
o
Material examined: BRAZIL. Amazonas, road from Manaus to Caracaraí km 45, INPA,
ro
3 Feb 1977, R. Singer B 10602 (INPA 73538, holotype!).
Observation: The holotype is satisfactorily well preserved and consist half basidioma,
p
but the partial veil probably leaved out.
d_
Remarks: This species probably presents one of the smallest spores and basidia among
te
the genus Amanita.. The subabrupt and submarginate bulb of A. campinaranae is similar
to A. guyanensis Mighell & T.W. Henkel. However, this latter species differs in the dark
c
gray-brown to grayish brown with darker pileus centre, larger basidiospores 5.0–9.0
(–11.0) × (4.0–)5.0–9.0(–10.0) μm, Qr = (0.94–)1.0–1.27(–1.33), Qm = 1.09 and larger
e
basidia 22–38 × 5.5–13 μm (Mighell

rr
2019).
Mighell et al. 2019
Amanita karea G.S. Ridl. from New Zealand also has a considerable distinct “marginate
o
to marginate-depressed” bulb with a “rim of pulverulent dark grayish sepia to fuscous

c
velar remnants”. However A. karea differs by the grayish to mouse gray pileus with dark
n
grayish sepia to mouse gray velar remnants, and larger ellipsoid basidiospores [223/21]
u
6.5–9 × 5.5–6.5(–8) µm, L = 8.2–8.7 µm, Q = (1.00–)1.5–1.54; Q = 1.32 (Ridley 1991).

The African A. echinulata Beeli shares in the proportionally small basidiospores (5–)5.5–
7(–9) × (4–)4.5–6(–7) μm, Qr = (1.00–)1.20–1.45(–1.50), Qm = 1.26, but differs at least
in the grayish brown pileus with conical to pyramidal velar warts, the less abrupt but
globose to rarely subclavate bulb, and the two broadly ellipsoid to ellipsoid basidiospores
(Beeli 1927, Mighell et al. 2021)
Amanita grisella E.-J. Gilbert & Cleland sensu Wood (1997) from Australia shares in the
grayish pileus with gray “flat, thin membranous velar scales” (although reported as paler
than pileus surface) and the “rounded bulb, rarely more turbinate, usually with some mar-
ginate ridges on the upper portion”. However, the Australian entity differs at least in the
considerable larger and broadly ellipsoid to ellipsoid basidiospores (7.5–)8.4–9.9(–12.3)
× (5.4–)6–7.5(–9.9) µm, Q = 1.19–1.35 and the presence of slightly inflated “subbasidial”
eschweizerbart_xxx
32 Felipe Wartchow
cells. This last sentence suggests that the material does not present a truly cellular subhy-
menium. Actually, A. grisella represents a confused species originally described as hav-
ing convex campanulate pileus without remnants of universal veil, slightly inflated stipe
base and larger basidiospores 8.5–10(–12.3) × 7–8.5 µm (Gilbert 1940).
Amanita phaea Bas nom. prov.

Fig. 7I
From original description: ‘Basidiomata (pileus just opened) large, terrestrial, solitary.
Pileus 65 mm wide and 45 mm high, conical with broadly rounded apex uniformly spadi-
ceous-sepia or very dark slightly purplish-brownish grey (“Hudson seal” M&P), some-
what lubricous in dried specimen shiny and very dark with only at margin a slight fibril-
lose structure visible; margin inflexed, smooth; context fleshy, white unchanging; univer-
sal veil as scattered small (in dried specimen up to 3 mm wide) grayish-whitish volval
warts and patches. Lamellae free, crowded, moderately broad, white (in dried specimen
f
sordid ochraceous with pallid subflocculose edge under lens); lamellulae subtruncate
o
(very short ones) to attenuate. Stipe 135 × 7 mm, sub cylindrical with pointed sub bulbous
ro
base, solid, white above and grey bellow ring but white at base, fibrillose all over but
particularly in upper half, glabrescent bellow; context white, unchanging; partial veil api-
p
cal, ample, pendulous, thin-membranous (in dried specimen entirely sticking to stipe),
d_
white, smooth, with somewhat lacerate-fimbriate edge; universal veil as scattered small
whitish scales on lower part of sub bulbous base. Odor unpleasant and taste not recorded’.
te
Basidiospores absent. Basidia not recovered, probably collapsed. Subhymenium un un-

c
known, collapsed. Lamella trama unknown. Pileus context poorly rehydrated; indistinctly
acrophysalidic; acrophysalides appearing to be 75 × 25 µm, difficult to observe; filamen-
e
tous hyphae 7–10 µm, entirely interwoven and some brownish pigmented ranging to
rr
10–12.5 µm; vascular hyphae up to 7–12 µm, very scarce. Stipe context longitudinally
acrophysalidic, with acrophysalides up to 145–210 × 20 µm; filamentous hyphae 3–
o
15 µm, hyaline with longitudinal orientation and brownish pigmented hyphae near the
c
surface, somewhat ranging to 20 µm wide with a slight attenuate or with branched infla
infla-
n
tions on some with terminal position; vascular hyphae 7–13 µm, uncommon. Pileipellis
u
very difficult to distinguish, but numerous filamentous hyphae 3–7 µm in the upper side
of context distinctly interwoven brownish pigmented (somewhat condensate), clampless;
vascular hyphae not seen; strongly gelatinous. Universal veil On pileus: abundant fila-
mentous hyphae 3–7 µm branched and more or less erect, frequently brownish pigmented,
clampless, and inflate ellipsoid, subglobose or ovoid terminal cells with approx. 75 ×
70 µm, 75 × 45 µm or 37 × 25 µm, difficult to observe in the exsiccate, apparently with
anticlinal orientation, colorless. On bulb: not recovered, probably collapsed. Partial veil
filamentous hyphae 3.5–7 µm, strongly interwoven, branched, common, somewhat with
condensed brownish or slightly paler pigments; inflated elements difficult to observe;
vascular hyphae very scarce up to 5 µm wide.
Habitat: Solitary on soil in secondary tropical forest (found only once).
Known distribution: State of Amazonas, North Brazil.
eschweizerbart_xxx
Material examined: BRAZIL. Amazonas, Manaus, INPA, 1977, R. Singer B 9791 (INPA
66709).
Observation: The material is poorly preserved, with the hymenium completely destroyed
by insects.
Remarks: The pigmented pileus, non-appendiculate margin and the absence of membra-
nous, limbate or saccate volva place A. phaea in the subg. Amanitina sect. Validae (Cui et
al. 2018). This material presents filamentous hyphae frequently containing condensed
brownish vacuolar pigments in pileus trama tissues studied, a feature not mentioned by
Bas (1978). It might be a helpful feature in future species description.
Amanita perphaea C. Simmons, T.W. Henkel & Bas described from Guyana is very sim-
ilar. However it primarily differs in the presence of filamentous hyphae with incrusted
brown pigment with terminal inflated elements ranging to 24 µm wide in the pileipellis
(Simmons et al. 2002).
). The Guyanese species is used as food by indigenous people ((Hen-
f
kel et al. 2004).
o
Species with occurrence in Brazil not confirmed
ro
p
d_
‘Amanita strobiliformis Vitt.’
Fig. 7D
te
Material examined:: BRAZIL. Rio Grande do Sul, São Leopoldo, 1929, J. Rick s.n.
c
(PACA 12221).
e
Remarks:: The correct citation of this name is A. strobiliformis (Paul. ex Vitt.) Bertil.
rr
(Neville & Poumarat 2004). Rick (1938, 1961) reported this material as having (trans-
(1938, 1961
lated from Latin) “Pileus 200–240 mm, convex expanded, white, skinned, margin ex ex-
o
ceeding, awkwardly (irregular?), warts hard, angulated, adnate, contextcompact, white,

c
probably unchanging. Lamellae free, edge smooth. Stipe 160–190 × 40 mm, solid, floc-
n
cose-squamose, bulbous marginate, incrusted with soil; ring ‘lacerate’; basidiospores

u
13–14 × 8–9 µm.”

Singer (1953) and Bas (1969) did not analyse this specimen, and this exsiccate contains
two different basidiomata belonging to two different genera:
One species contains apparently a single smooth pileus and two detachable stipes. They
bear strongly incrusted with soil emarginated bulb as mentioned by Rick (1938, 1961).
Also, the stipe was referred as having “floccose-squamose” surface (Rick 1938, 1961).
Although the lamella trama was impossible to rehydrate due to poorly preserved material,
the stipe tissue was relatively satisfactorily rehydrated and no acrophysalides were found.
I also found thick walled hyaline basidiospores, with conspicuous germ pore covered by
hyaline cap with weak dextrinoid reaction, characteristics of Macrolepiota Singer (Vel-
linga 2003).
eschweizerbart_xxx
34 Felipe Wartchow
The second basidiome really corresponds to a specimen of poorly preserved Amanita.

The stipe is also separate from pileus but no bulb was found, and an apparent pendant and
incomplete ring-like structure is observed, and the pileus surface is verrucose. The stipe
tissue showed very poor and unsatisfactory rehydration, but very few apparent acrophy-
salides were observed; also very few basidiospores were in condition to measure: [6/1/1]
(8–)9–11(–12.5) × (7–)7.5–8 µm, (L = 10.2 µm; W = 7.6 µm; Q = (1.20–)1.24–1.37
(–1.55); Qm = 1.34), amyloid, hyaline, thin walled, broadly ellipsoid to ellipsoid, with a
sublateral apiculus. These features are similar to A. spissa var. laeta (Wartchow 2021),
but other anatomical structures could not be observed.
Amanita strobiliformis was studied by Bas (1969), and was also referred with pileus hav-
ing “…thick, wart- to patch- or crust-like volval remnants…”. This feature can be analo-
gous to Rick’s “hard, adnate warts”. When using “adnate” in his description, Rick (1938,
1961) probably tried meant that the warts are adhering to the pileus like a patch or crust,
as it occurs in the European species. This mistake sounds reasonable when considering
f
the scarcity of publications and the primitive conditions for microscopic studies of aga
aga-
o
rics during the early Twentieth Century
ro
Amanita ameghinoi (Speg.) Singer p
MycoBank: MB#292438
d_
Remarks:: The specimen was described from the Argentinean Pampa region as Armilla-
te
ria ameghinoi Speg. and transferred to Amanita by Singer (1952). The species belongs to
subsect. Vittadinii stirps Vittadinii,, and the holotype is poorly preserved ((Bas 1969).
c
Pegler (1997)) reported a specimen with collector number ‘Puiggari 202’ from São Paulo
and I contacted the herbaria SP, LPS and BAFC asking for this exsiccate. They responded
e
that do not exist material assigned under this collection number.

rr
o
c
Excluded species
n
u
Amanitopsis plumbea Rick and Collybia sulcatissima Rick were previously interpreted as
doubtful taxa and excluded by Bas (1978). Six more exsiccates identified as Amanita
must also be excluded.
“Amanita porphyria Fr.”

Material examined: BRAZIL. Pernambuco, Recife, “Jardim Zôo-Botânico de Dois Ir-
mãos”, 03 Jul 1951, C. Ferreira s.n. (IPA 2695).
The correct name is Amanita porphyria Alb. & Schwein.: Fr., originally described from
Germany (Albertini & Schweinitz 1805). The poorly preserved exsiccate analyzed here
presents thick walled basidiospores with a truncate germ pore, and might corresponds to
a species of Chlorophyllum Massee (Vellinga 2003).
eschweizerbart_xxx
“Amanita sp.”
Material examined: BRAZIL. Paraíba, João Pessoa, Campus I da UFPB, 30 May 1990,
I.C. Oliveira 278 (JPB 17567).
Remarks: This material is poorly preserved, and examination of the exsiccatum revealed
the pileus to 70 mm in diam., stipe 130 × 11 mm with a thick annulus and abrupt bulb to
25 mm wide. Microscopically the basidiospores are hyaline and colorless without germ
pore, thick-walled to 1 µm thick, slightly dextrinoid and metachromatic in cresyl blue; the
stipe lack acrophysalides; pileus covering if formed by with erect hyphae. These features
are characteristic of some species of Chlorophyllum with basidiospores lacking germ
pore, as for example, C. demangei (Pat.) Z.W. Ge & Zhu L. Yang, C. hortense (Murrill)
Vellinga, and other (Ge et al. 2018, Loizides et al. 2020, Sysouphanthong et al. 2021).
“Amanita porphyria Fr.”
f
Material examined: BRAZIL. Pernambuco, Recife, “Jardim Zôo-Botânico de Dois Ir Ir-
o
mãos”, 06 Jul 1951, C. Ferreira s.n. (IPA 2697, “Amanita
Amanita porphyria Fr.”); same place, 03
ro
Jul 1951, C. Ferreira s.n. (IPA 2705).
These exsiccates are poorly preserved, but direct observations on basidiomata reveal a
p
distinctly marginate bulb and free lamellae. Under microscope only a single thick-walled
d_
colorless weakly dextrinoid, slightly metachromatic, and cyanophilic basidiospore lack-
lack
ing germ pore measuring 9.5 × 5 µm was found. Although stipe context very weakly re
te
re-
hydrated, no acrophysalides were observed. These exsiccates probably correspond to
c
genus Leucoagaricus (Vellinga 2004).

e
“Amanita sp.”
rr
Material examined:: BRAZIL. Bahia, Jussari, Serra do Teimoso, 28 Jul 2002, J.L. Be-
o
zerra JLB 159 (CEPEC 238).

c
Remarks:: This exsiccatum consists in a single basidiome without field notes, but with
n
small fibrils over pileus, and apparent free lamellae. Microscopic analyses reveal lack of
u
acrophysalides in stipe context, and hyaline dextrinoid thick-walled basidiospores with

reticulate ornamentation and absence of germ pore. It certainly represents a species of
Rugosospora Heinem. (Heinemann 1973), more probably R. pseudorubiginosa (Ci-
fuentes & Guzmán) Guzmán & Bandala (Guzmán et al. 1989, Franco-Molano 1995).
“Amanita sp.”
Material examined: BRAZIL. Paraíba, João Pessoa, Campus I da UFPB, 26 Aug 1991,
I.C. Oliveira 358 (JPB 17585).
Remarks: No collector notes accompanying exsiccate. This material is poorly preserved,
on which critical points, as for example the lamella trama, do not rehydrate satisfactory.
Obviously it does not represent a specimen of Amanita mainly due the lack of acrophy-
salides in the stipe tissue.
eschweizerbart_xxx
36 Felipe Wartchow
‘Amanitopsis regalis Berk.’

Material examined: BRAZIL. Pernambuco, Recife, “Jardim Zôo-Botânico de Dois Ir-
mãos”, 13 Jun 1952, J.N. Silva s.n. (URM 704).
Remarks: ‘Amanitopsis regalis’ was cited by Maia et al. (2002), and represents a single
poorly preserved basidiome without any field notes. Direct examination of the exsiccate
reveals in the slightly inflated stipe base a sac-like volva fragments. The hymenium is
completely collapsed and was impossible to observe the lamella trama. The basidiospores
are not hyaline, but more or less stramineous with somewhat pinkish thick walls and no
reaction with Melzer’s reagent was seen. They resemble the ones of and the stipe tissue
does not present any inflated elements that resemble acrophysalides.This material does
not represent a specimen of Amanita but an unidentified Volvariella Speg. (Singer 1986),
and must be excluded from the list of taxa known from Pernambuco and Brazil.
Originally described as Agaricus regalis Berk., Amanitopsis regalis (Berk.)Sacc. repre-
f
sents a synonym of Amanita berkeleyi (Hook. f. in Berk.) Bas, a species of subg. Amani-
o
tina subsect. Solitariae Bas stirps Eriophora originally described from India ((Bas 1969,
ro
Cui et al. 2018).
p
d_
Acknowledgements
te
I would like show all my gratitude to Dr. Rodham E. Tulloss for teaching several aspects
of anatomy, morphology and taxonomy of Amanita
Amanita, and first of all, by his great friendship
c
during my visiting to Roosevelt-NJ. Dr. Maria Auxiliadora de Queiroz Cavalcanti and Dr.
e
Leonor Costa Maia are kindly acknowledged by the supervision during my Ph.D. thesis.
rr
Dr. Dirce L. Komura (INPA, Manaus) is very thankfull by sharing information about the
preservation of the exsiccates deposited in INPA. The following herbaria curators and
o
responsible are acknowledged for answering author questionings and, when possible,
c
loan of exsiccate used in this study: Maria S. Marchioretto (PACA), Carlos Franciscon
n
(INPA), Ronaldo Kummrow (MBM), Dr. Adriana M. Gugliotta (SP), José L. Bezerra
u
(CEPEC), Jair Putzke (HCB), Machiel E. Noordeloos (L), Mara R. Ritter (ICN), Luciano
Q. Paganucci (HUEFS), Clarice Loguercio-Leite (FLOR), Vilma Rosato (LPS), James
M. Trappe (OSC), Taciana B. Cavalcanti (CEN), Maria Regina V. Barbosa (JPB), Antô-
nio F. Carpanezzi and Cristiane V. Helm (HFC), Élen N. Garcia (PEL), Paulo Labiak
(UPCB), Vera Martins (R), Andrés de Errasti (BAFC), Maria S. Marchioretto (PACA),
Viviane Scalon (OUPR), Ubiratã Jacobi (HURG), Hélen Sotão (MG), Lana S. Sylvestre
(RBR), Margarida F. Ito (IACM), Ronaldo Wasum (HUCS), André Jasper (HVAT), Paulo
S.T. Brioso (UFRJ) and Liliana Dalavale (HERBARA). The following collegues are ack-
nowledged for helping with collecting materials: Ângelo Souto, Daniela K.A. da Silva
and Nicácio O. Freitas. Dr. Geoffrey R. Ridley is thanked for sent literature. I also ack-
nowledge CNPq for providing PhD scholarship (PROTAX/CNPq/MCT Proc. 563969/05-
9), funding the project ‘Fungos agaricoides em áreas de Mata Atlântica e Caatinga no
Estado da Paraíba’ (Edital Universal Proc. 420.448/2016-0) and providing ‘Produtividade
eschweizerbart_xxx
em Pesquisa’ (Proc. 307922/2014-6, Proc. 307947/2017-3 and Proc. 309652/2020-0)

grant to F. Wartchow.
References
Albertini, J.B., & Schweinitz, L.D. (1805). Conspectus Fungorum in Lusatiae Superioris Agro Ni-
skiensi Crescentium e Methodo Persooniana. Lipsiae: Sumtibus Kummerianis. https://doi.
org/10.5962/bhl.title.3601
Alves-Araújo, A., Araújo, D., Marques, J., Melo, A., Maciel, J. R., Uirapuã, J., . . . Alves, M. (2008).
Diversity of angiosperms in fragments of Atlantic Forest in the State of Pernambuco, Northeast-
ern Brazil. Bioremediation, Biodiversity & Bioavailability, 2, 14–26.
Bas, C. (1969). Morphology and subdivision of Amanita and a monograph on its section Lepidella.
Persoonia, 5, 285–579.
Bas, C. (1978). Studies in Amanita. I. Some Amazonian species. Persoonia, 10, 10, 1–22.
Bas, C. (1982). Studies in Amanita. II. Persoonia, 11, 429–442.
f
Bas, C., & de Meijer, A. A. R. (1993). Amanita grallipes,, a new species in Amanita subsection Vit-
o
tadiniae from southern Brazil. Persoonia, 15, 345–350.
ro
Beeli, M. (1927). Contribution a l’étude de La flore mycologique du Congo. Bulletin de la Socété
Royale de Botanique de Belgique, 59, 101–112. p
Capelari, M., & Maziero, R. (1988). Fungos macroscópicos do estado de Rondônia região dos Rios
d_
Jaru e Ji-Paraná. Hoehnea, 15, 28–36.
Corner, E. J. H., & Bas, C. (1962). The genus Amanita in Singapore and Malaya. Persoonia, 2,
te
241–304.
molecu
Cui, Y.-Y., Cai, Q., Tang, L.-P., Liu, J.-W., & Yang, Z.-L. (2018). The family Amanitaceae: molecu-
lar phylogeny, higher-rank taxonomy and the species in China. Fungal Diversity, 91(1), 5–230.
c
https://doi.org/10.1007/s13225-018-0405-9
e
De Meijer, A. A. R. (2001). Mycological work in the Brazilian state of Paraná. Nova Hedwigia,
rr
72(1-2), 105–159. https://doi.org/10.1127/nova.hedwigia/72/2001/105

De Meijer, A. A. R. (2008). Notable Macrofungi from Brazil’s Paraná Pine Forest/ Macrofungos
o
Notáveis das Florestas de Pinheiro-do Paraná. Colombo: Embrapa Florestas.

Franco-Molano, A. E. (1995). Observations on Rugosospora Heinemann in the Neotropics. Myco-
c
logia, 87(4), 574–578. https://doi.org/10.1080/00275514.1995.12026570

n
Ge, Z.-W., Jacobs, A., Vellinga, E. C., Sysouphanthong, P., van der Walt, R., Lavorato, C., . . . Yang,
u
Z. L. (2018). A multi-gene phylogeny of Chlorophyllum (Agaricaceae, Basidiomycota): New

species, new combinations and infrageneric classification. MycoKeys, 32, 65–90. https://doi.
org/10.3897/mycokeys.32.23831
Gilbert, E.J. (1940). Amanitaceae. In J. Bresadola, Iconographia Mycologica, Vol. 27 (Suppl. I),
Milano: Società Botanica Italiana.
Guzmán, G. (1982). New species of fungi from the Yucatán Peninsula. Mycotaxon, 16, 249–261.
Guzmán, G., Bandala, V. M., Montoya, L., & Saldarriaga, Y. (1989). Nuevas evidencias sobre las
relaciones micoflorísticas ente África y el neotrópico. Brenesia, 32, 107–112.
Heinemann, P. (1973). Leucocoprinées nouvelles d’Afrique centrale. Bulletin du Jardin Botanique
National de Belgique, 43(1/2), 7–13. https://doi.org/10.2307/3667558
Henkel, T. W., Terborgh, J., & Vilgalys, R. J. (2002). Ectomycorrhizal fungi and their leguminous
hosts in the Pakaraima Mountains of Guyana. Mycological Research, 106(5), 515–531. https://
doi.org/10.1017/S0953756202005919
Henkel, T. W., Aime, M. C., Chin, M., & Andrew, C. (2004). Edible mushrooms from Guyana. The
Mycologist, 18(3), 104–111. https://doi.org/10.1017/S0269915X04003027
eschweizerbart_xxx
38 Felipe Wartchow
IBGE (2004). Mapas de Cobertura Vegetal dos Biomas Brasileiros. Primeira Aproximação. Bra-
sília: Ministério do Meio Ambiente, Ministério do Planejamento, Orçamento e Gestão Instituto
Brasileiro de Geografia e Estatística, Diretoria de Geociências.
IBGE (2012). Mapa da Área de Aplicação da lei Nº 11.428 de 2006. 2a edição. Brasília: Ministério
do Meio Ambiente; Instituto Brasileiro de Geografia e Estatística; Diretoria de Geociências.
Jenkins, D. T. (1977). A taxonomic and nomenclatural study of the genus Amanita section Amanita
for North America. Bibliotheca Mycologica, 57, 1–126.
Kimmel, T. M., do Nascimento, L. M., Piechowski, D., Sampaio, E. V. S. B., Nogueira Rodal, M.
J., & Gottsberger, G. (2010). Pollination and seed dispersal modes of woody species of 12-year-
old secondary forest in the Atlantic Forest region of Pernambuco NE Brazil. Flora (Jena),
205(8), 540–547. https://doi.org/10.1016/j.flora.2009.12.022
Kramer, L. A. (2004). The Online Auction Color Chart. Stanford: Online Auction Color Co.
Loizides, M., Alvarado, P., Polemis, E., Dimou, D. M., Zervakis, G. I., Thines, M., . . . Gube, M.
(2020). Multiple evolutionary origins of sequestrate species in the agaricoid genus Chlorophyl-
lum. Mycologia, 112(2), 400–422. https://doi.org/10.1080/00275514.2020.1712179
Maerz, A. J., & Paul, M. R. (1950). A dictionary of color.. New York: McGraw-Hill Book Company,
Inc.
f
Maia, L. C., Melo, A. M. Y., & Cavalcanti, M. A. (2002). Diversidade de fungos no estado de Per Per-
o
nambuco. In M. Tabarelli & J. M. C. da Silva (Eds.), Diagnóstico da Biodiversidade de Per Per-
ro
nambuco. Recife: Ed. Massangana.
Menolli, N., Jr., Asai, T., & Capelari, M. (2009). Amanita coacta ((Amanitaceae, Agaricales) with
key to Amanita species occuring in Brazil. Mycotaxon, 107(1), 419–430. https://doi.
p
d_
org/10.5248/107.419
Mighell, K. S., Henkel, T. W., Koch, R. A., Goss, A., & Aime, M. C. (2019). New species of Ama-
nita subgen. Lepidella from Guyana. Fungal Systematics and Evolution, 3(1), 1–12. https://doi.
te
org/10.3114/fuse.2019.03.01
Mighell, K. S., Henkel, T. W., Koch, R. A., Chin, M. L., Brann, M. A., & Aime, M. C. (2021). Am-
c
anita in the Guineo-Congolian rainforest: Epitypes and new species from the Dja Biosphere
e
Reserve, Cameroon. Mycologia, 113 113(1),

(1), 168–190. https://doi.org/10.1080/00275514.2020.181
6386
rr
Moretti, G. (1826). Il botanico Italiano ossia discussioni sulla Flora Italica. Giornale di Fisica,
Chimica, Storia Naturale, Medicina ed Arti
Arti, 2(9), 65–85.
o
Neville, P., & Poumarat, S. (2004). Amaniteae: Amanita, Limacella & Torrendia. Fungi Europaei,
c
9, 1–1120.
n
Parolin, P., Adis, J., Rodrigues, W. A., Amaral, I., & Piedade, M. T. F. (2004). Floristic study of an
u
igapó floodplain forest in Central Amazonia, Brazil (Tarumã-Mirim, Rio Negro). Amazoniana,
18, 29–47.
Pegler, D. N. (1983). Agaric Flora of the Lesser Antilles. Kew Bulletin Additional Series, 9, 1–668.
Pegler, D. N. (1997). The Agarics of São Paulo, Brazil. Kew: Royal Botanical Gardens.
Pegler, D. N., & Shah-Smith, D. (1997). The genus Amanita (Amanitaceae, Agaricales) in Zambia.
Mycotaxon, 61, 389–417.
Raithelhuber, J.H. (2004). Nueva flora mycológica Argentina. Stuttgart: Published by the author.
Reid, D. A. (1980). A monograph of the Australian species of Amanita Pers. ex. Hook. (Fungi).
Australian Journal of Botany. Supplementary Series, 8, 1–97.
Rick, J. (1906). Pilze aus Rio Grande do Sul. Brotéria. Série Botânica, 5, 5–53.
Rick, J. (1938). Agarici Riograndensis. II. Lilloa, 2, 251–316.
Rick, J. (1961). Basidiomycetes Eubasidii in Rio Grande do Sul – Brasilia. 5. Iheringia. Série
Botânica, 8, 296–450.
Ridley, G. S. (1991). The New Zealand species of Amanita (Fungi: Agaricales). Australian System-
atic Botany, 4(2), 325–354. https://doi.org/10.1071/SB9910325
eschweizerbart_xxx
Sanmee, R., Yang, Z.-L., Lumyong, P., & Lumyong, S. (2003). Amanita siamensis, a new species
of Amanita from Thailand. Mycotaxon, 88, 225–228.
Sanmee, R., Tulloss, R. E., Lumyong, P., Dell, B., & Lumyong, S. (2008). Studies on Amanita
(Basidiomycetes: Amanitaceae) in Northern Thailand. Fungal Diversity, 32, 97–123.
Schessl, M., Silva, W. L., & Gottsberger, G. (2008). Effects of fragmentation on forest structure and
litter dynamics in Atlantic rainforest in Pernambuco, Brazil. Flora (Jena), 203(3), 215–228.
https://doi.org/10.1016/j.flora.2007.03.004
Simmons, C., Henkel, T. W., & Bas, C. (2002). The genus Amanita in the Pakaraima Mountains of
Guyana. Persoonia, 17, 563–582.
Singer, R. (1952). Type studies on Basidiomycetes. VIII. Armillaria-types from Instituto de Bo-
tánica “C. Spegazzini”. Sydowia, 6, 344–351.
Singer, R. (1953). Type studies on Basidiomycetes. VI. Lilloa, 26, 57–159.
Singer, R. (1986). The Agaricales in modern taxonomy. Koenigstein: Koeltz Scientific Books.
Singer, R., & Aguiar, I. A. (1986). Litter decomposing and ectomycorrhizal Basidiomycetes in an
igapó forest. Plant Systematics and Evolution, 153, 107–117. https://doi.org/10.1007/
BF00989420
Singer, R., & Araújo, I. J. (1979). Litter decomposition and Ectomycorrhiza in Amazonian forests.
f
1. A comparison of litter decomposing and ectomycorrhizal Basidiomycetes in latosol-terra-
o
firme rain forest and white podzol campinarana. Acta Amazonica, 99(1), (1), 25–41. https://doi.
ro
org/10.1590/1809-43921979091025
Singer, R., & Digilio, A. P. L. (1951). Pódromo de la flora agaricina Argentina. Lilloa, 25, 5–461.
Sysouphanthong, P., Thongklang, N., Liu, J.-K., & Vellinga, E. C. (2021). Description of Lepiota
Lepiota-
p
d_
ceous fungal species of the genera Chlorophyllum, Clarkeinda, Macrolepiota, Pseudolepiota
Pseudolepiota,
and Xanthagaricus from Laos and Thailand. Diversity, 13 13(12),
(12), 666. https://doi.org/10.3390/
d13120666
te
Thiers, B. (2023). Index Herbariorum: A global directory of public herbaria and associated staff.
New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/
c
Trindade, M. B., Lins-e-Silva, A. C. B., Silva, H. P., Figueira, S. B., & Schessl, M. (2008). FragFrag-
e
mentation of the Atlantic rainforest in the northern coastal region of Pernambuco, Brazil: Re Re-
cent changes and implications for conservation. Bioremediation, Biodiversity & Bioavailability,
rr
2, 5–13.
Tulloss, R. E. (1994). Type studies in Amanita section Vaginatae I: Some taxa described in this
o
century (studies 1–23) with notes on description on spore and refractive hyphae in Amanita.
c
(1), 305–396.
Mycotaxon, 52(1),
n
Tulloss, R. E. (2000). Note sulla metodologia per lo studio del genere Amanita (Agaricales). Bolle-
u
tino del Gruppo Micologico G. Bresadola

Bresadola. Nuova serie, 43, 41–58.
Tulloss, R. E., & Franco-Molano, A. E. (2008). Studies in Amanita subsection Vittadiniae I – a new
species from Colombian savanna. Mycotaxon, 105, 317–323.
Tulloss, R. E., & Halling, R. E. (1997). Type studies of Amanita morenoi and Amanita pseudospreta
and a reinterpretation of crassospores in Amanita. Mycologia, 89(2), 278–288. https://doi.org/1
0.1080/00275514.1997.12026782
Tulloss, R. E., & Lindgren, J. E. (2005). Amanita aprica – a new toxic species from western North
America. Mycotaxon, 91, 193–205.
Tulloss, R. E., & Traverso, M. (2000). Amanita basiana – a new species from pure Pinus forest and
resembling the Alnus-associated species Amanita friabilis. Mycotaxon, 77, 47–55.
Tulloss, R. E., & Yang, Z.-L. (2023). Studies in Amanitaceae. www.amanitaceae.org
Tulloss, R. E., Stephenson, S. L., Bhatt, R. P., & Kumar, A. (1995). Studies on Amanita (Amanita-
ceae) in West Virginia and adjacent areas of the Mid-Appalachians. Preliminary results. Myco-
taxon, 56, 243–293.
Tulloss, R. E., Kuyper, T. W., Vellinga, E. C., Yang, Z.-L., Halling, R. E., Geml, J., . . . Pringle, A.
(2016). The genus Amanita should not be split. Amanitaceae, 1(3), 1–16.
eschweizerbart_xxx
40 Felipe Wartchow
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., . . .
Smith, G.F. (2018). International Code of Nomenclature for algae, fungi, and plants (Shenzhen
Code). Regnum Vegetabile, 159. Glashütten: Koeltz Botanical Books.
Vellinga, E. (2003). Chlorophyllum and Macrolepiota (Agaricaceae) in Australia. Australian Sys-
tematic Botany, 16(3), 361–370. https://doi.org/10.1071/SB02013
Vellinga, E. C. (2004). Genera in the family Agaricaceae: Evidence from nrITS and nrLSU se-
quences. Mycological Research, 108(4), 354–377. https://doi.org/10.1017/S0953756204009700
Vizzini, A., Redhead, S. A., & Dovana, F. (2017). Epitypification of Agaricus vittadinii (Basidio-
mycota, Amanitaceae). Phytotaxa, 326(3), 230–234. https://doi.org/10.11646/phytotaxa.326.3.8
Wartchow, F. (2017). O gênero Amanita (Fungi): Sistemática e distribuição no Brasil. Pesquisa e
Ensino em Ciências Exatas e da Natureza, 1(1), 28–44. https://doi.org/10.29215/pecen.v1i1.165
Wartchow, F. (2021). Neotype of Amanita spissa var. laeta (Basidiomycota, Amanitaceae). Anales
del Jardin Botánico de Madrid, 78(2), e118. https://doi.org/10.3989/ajbm.2598
Wartchow, F., & Gamboa-Trujillo, J. P. (2013). Amanita chocoana – a new species from Ecuador.
Mycotaxon, 121(1), 405–412. https://doi.org/10.5248/121.405
Wartchow, F., Tulloss, R. E., & Cavalcanti, M. A. Q. (2007). The discovery of Amanita lilloi in
Brazil. Mycotaxon, 99, 167–174.
f
Wartchow, F., Maia, L. C., & Cavalcanti, M. A. Q. (2012). Studies on Amanita (Agaricomycetidae,
o
Amanitaceae) in Brazil: two yellow gemmatoid taxa. Nova Hedwigia, 96(1-2), 61–71. https://
ro
doi.org/10.1127/0029-5035/2012/0053
Wood, A. E. (1997). Studies in the genus Amanita (Agaricales) in Australia. Australian Systematic
Botany, 10(5), 723–854. https://doi.org/10.1071/SB95049
p
d_
Yang, Z.-L. (2000). Revision of the Chinese Amanita collections deposited in BPI and CUP. Myco-
taxon, 75, 117–130.
Yang, Z.-L., & Oberwinkler, F. (1999). Die Fruchtkörperentwicklung von Amanita muscaria (Ba-
te
sidiomycetes). Nova Hedwigia, 68(3-4), (3-4), 441–468. https://doi.org/10.1127/nova.hedwi-

gia/68/1999/441
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Manuscript received: September 25, 2023

Revisions requested: October 26, 2023
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Revised version received: November 2, 2023

Manuscript accepted: December 23, 2023
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Responsible editor: G. Kost

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C Nova Hedwigia, Vol.

000 (2024), Issue 0-0, 000–000

Studies on types and additional materials of Amanita

Keywords: Agaricomycetes; Agaricomycetidae; Amanitaceae; the Neotropics; taxonomy

© 2024 J. Cramer in Gebrüder Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de

Material and methods

al. (2016) and Cui et al. (2018).

were analyzed: Amanita lanivolva

Subgenus Amanita section Amanita

Amanita lanivolva Bas, Persoonia 10: 13. 1978.

and stipe (“unitangent volva”).’ Odorless; taste not checked by Bas.

(1.21–)1.29–1.46(–1.49), Q = 1.34), inamyloid, hyaline, colorless, broadly ellipsoid to

= up to 25 µm; wst-far = up to 36 µm. Lamella trama difficult to rehydrating in the

9897 (INPA 66716, paratype!).

Observation on exsiccate: The exsiccate consists in a well preserved specimen with

Amanita sulcatissima Bas, Persoonia 10: 9. 1978.

fibrillose structure; margin densely sulcate-striate (14–18 grooves per 10 mm of circle

vascular hyphae absent. Partial veil absent.

Habitat: “in campina vegetation on humus under trees (Humiriaceae, Burseraceae,

Ochnaceae, Sampindaceae, Sapotaceae and leguminous trees)” (Bas 1978).

Amanita xerocybe Bas, Persoonia 10: 7. 1978.

white to creamy, unchanging; universal veil entirely decorating (densely at centre to

felted-subflocculose at underside; minutely granular at upper side); universal veil as an

Fig. 2. A–D. Amanita xerocybe (Holotype). A. Basidiospores. B. Basidia and subhymenium. C.

to gradually diverging from central stratum; wcs = ± 22 µm; filamentous hyphae up to

thick wall present between the filamentous hyphae.

stone and cliff boulders” ((Simmons et al. 2002).

4.5–6 µm, L’ = 8 µm, W’ = 5 µm, Q = 1.56 (Pegler 1983).

Subgenus Amanita section Vaginatae

Amanita coacta Bas, Persoonia 10: 15. 1978.

interwoven filamentous hyphae 2.5–7.5 µm, mixed to terminal elements symmetrically

phae ranging to 7 µm and some long-clavate inflated elements up to 60 × 14 µm, gradu-

10–17 µm, common. Stipe context moderately rehydrated, longitudinally acrophysalidic;

strongly gelatinized. On stipe 10 mm above base of SP 214459 and SP 393677 (inner

Q = (1.14–)1.19–1.56(–1.69), Q = (1.28–)1.31–1.37, Qm = 1.33; (3) the occurrence of

common large subhymenial cells ranging to 25 µm in diam.

Amanita craseoderma Bas, Persoonia 10: 20. 1978.

Basidiospores [50/1/1] (8–)8.3–10(–10.5) × (6.5–)7–8.5(–9.5) µm, ((L = 8.8 µm; W =

7.7 µm; Q = (1.00–)1.03–1.31(–1.38); Q = 1.14), inamyloid, hyaline, colorless, globose

rounded/truncate, subapical to sublateral; contents including a large guttule. Basidia dif-

less. Subhymenium: unsatisfactory rehydrated, up to 30 µm thick, with 3–4 layers of

isodiametric cells up to 8–17 µm diam.; wst-near = ± 18 µm; wst-far = ± 25 µm. Lamella

80 × 32 µm gradually diverging from central stratum; vascular hyphae scarce 6–10 µm

wide, more or less longitudinal to lamella edge. Marginal tissue of lamellae

MycoBank Typification #(MBT) 10016355

Fig. 4. A–C. A. crebresulcata (lectoype). A. Hymenium and subhymenium. B. Basidiospores. C.

Stipe context longitudinally acrophysalidic, with acrophysalides up to 200–370 × 35–

75 µm; filamentous hyphae 2–5 µm wide, having a longitudinal orientation, unbranching;

Dois Irmãos, 31 Aug 1951, J. Nery s.n. (IPA 2874, as ‘Amanitopsis

Observations on exsiccate: The holotype (INPA 66710) is destroyed, remaining small

12.8 µm, L’ = 11.8 µm; W = 8–9.1 µm, W’ = 8.6 µm; Q = (1.17–)1.25–1.56(–1.67); Q =

less. Subhymenium weakly rehydrated difficult to analyze, apparently cellular; appearing

± 30 µm; wst-far = ± 55 µm. Lamella trama difficult to rehydrate; wcs= ± 30 µm wide;

filamentous hyphae 3.5–7(–9) µm not fully longitudinal to lamella; clavate elements

± 27 × 13 µm, rare. Pileus context difficult to rehydrate; acrophysalides approx. 70 ×

12.1 µm, L’ = 11 µm; W = 9.3–11.5 µm, W’ = 10.2 µm; Q = (1.00–)1.01–1.16(–1.25);

cular hyphae at universal veil (Tulloss 1992).

W = 7.7 µm, Q = (1.30–)1.39–1.60(–1.78); Q = 1.53, and pileipellis with narrower hy-

phae (Tulloss 2023).

probably clampless. Subhymenium difficult to rehydrate, but cellular bearing subisodia-

can be compared with A. crebresulcata and A. dunicola,

lack of distinctly gelatinized suprapellis.