Content uploaded by Leticia Montoya
Author content
All content in this area was uploaded by Leticia Montoya
Content may be subject to copyright.
Crepidotus crocophyllus
found in Costa Rica and Mexico and revision of related
species in subsection
Fulvifibrillosi
Victor M. Bandala
1
Leticia Montoya
Biodiversidad y Sistema´tica, Instituto de Ecologı
´
a, A.C.,
P.O. Box 63, Xalapa, Veracruz 91000, Mexico
Milagro Mata
Instituto Nacional de Biodiversidad, P.O. Box 22-
3100, Santo Domingo de Heredia, Costa Rica
Abstract
: The study of
Crepidotus
specimens collect-
ed in Costa Rica and Mexico revealed that
C.
crocophyllus
occurs in the tropical and subtropical
forests of both countries. Type specimens of seven
species related to
C. crocophyllus
in subsection
Fulvifibrillosi
s. Hesler and Smith were re-examined.
Based on the morphological features, specimens
supporting
C. appalachianensis
,
C. aureifolius
,
C.
distortus
,
C. subaureifolius
and
C. subnidulans
are
interpreted to be
C. crocophyllus
, thus all herein are
proposed as its synonyms. Furthermore
A. nephrodes
is
confirmed as a synonym of
Crepidotus crocophyllus
while
Agaricus malachius
, long considered contaxic
with the former, is proposed as synonym of
Crepidotus
applanatus
. The known records of
C. crocophyllus
indicate a wide but fragmented range of extension of
the taxon throughout the Americas. Description,
illustrations of microscopic features and discussions
are provided.
Key words:
Crepidotaceae, new synonymies, taxo-
nomy, tropical fungi, wood-inhabiting fungi
INTRODUCTION
Subsection
Fulvifibrillosi
Hesler & A.H. Sm. (1965)
consists of an assemblage of up to eight members of
Crepidotus
especially distinguished among the bacu-
late, globose-spored taxa by sharing a pileipellis with
pigmented, often incrusted, thick-walled hyphae that
form macroscopic brown fibrils or squamules on the
pileus surface. Taxonomic distinctions within the
Fulvifibrillosi
are often problematic when attempting
to separate the taxa proposed by the aforementioned
authors based upon the combination of seemingly
variable characters mentioned in the protologs. Fresh
newly discovered specimens related to this group
show transitional stages of variation of the macro-
scopic characters and consequently a high degree of
overlapping when compared on the taxonomic
scheme proposed by Hesler and Smith (op. cit.). In
the process of constructing a stable classification for
Crepidotus
we find it important to take into account
that at least a certain number of type specimens
require additional observation to establish consisten-
cy of described characters. In addition we assume the
hypothesis that many species could have a wider
range of occurrence than that known to date.
For this paper we studied Mexican and Costa Rican
collections related to
Crepidotus crocophyllus
(Berk.)
Sacc., a member of the
Sphaerula s
. Hesler & Smith
(1965), close to the group of species recognized by
these authors in subsection
Fulvifibrillosi
. The mor-
phological variation observed in the samples exam-
ined, as well as some different opinions found in the
literature with regard to the taxonomy of members
belonging to that group, prompted us to make a
critical reappraisal of the taxonomic status of type
collections representing seven species placed by
Hesler and Smith (1965) in the
Fulvifibrillosi
. Six of
these collections are interpreted as
C. crocophyllus
,
while the type collection of
Agaricus malachius
Berk.
& M.A. Curt. is found to represent
Crepidotus
applanatus
(Pers.) P. Kumm.
MATERIALS AND METHODS
Macroscopic characteristics were observed on fresh basi-
diomes collected in Costa Rica and Mexico. Color codes in
descriptions refer either to Kornerup and Wanscher (1967,
e.g. 2A 2–3) or to Munsell (1994, e.g. 2.5Y 8/2–3). Hand
sections of dried specimens were mounted in 3% KOH or
with Congo red. Methods employed in the microscopic
analysis of specimens, including SEM and basidiospore
measurements and their statistics, are those used in
previous works (Bandala et al 1999, 2006, Bandala and
Montoya 2000a, 2004),
x
corresponds to the range of means
of length and width based at least on 50 spores per
collection and
Q
to the range of means of the ratio of
basidiospores length/width of
n
collections. Herbarium
acronyms are according to Holmgren et al (1990).
TAXONOMY
Crepidotus crocophyllus
(Berk.) Sacc. Syll Fung 5:886.
1887. F
IGS. 1–5
Bas.:
Agaricus crocophyllus
Berk. Lond J Bot 6:313. 1847.
Syn.:
Crepidotus applanatus
var.
crocophyllus
(Berk.) Pila´t
Atl Champ Eur 6:35. 1948.
Accepted for publication 17 December 2007.
1
Corresponding author. E-mail: victor.bandala@inecol.edu.mx
Mycologia,
100(2), 2008, pp. 335–346.
#
2008 by The Mycological Society of America, Lawrence, KS 66044-8897
335
C. fulvifibrillosus
Murrill. N Am Fl. 10:153. 1917.
C. applanatus
var.
fulvifibrillosus
(Murrill) Pila´t Atl
Champ Eur 6:35. 1948.
C. appalachianensis
Hesler & A.H. Sm. N. Am. Sp.
Crepidotus
, p 72. 1965.
C. aureifolius
Hesler & A.H. Sm. N. Am. Sp.
Crepidotus
,
p. 75. 1965.
?
C. badiofloccosus
S. Imai. Bot. Mag. 53: 399. 1939. (type
not seen).
C. distortus
Hesler & A.H. Sm. N. Am. Sp.
Crepidotus
,
p. 72. 1965.
Agaricus nephrodes
Berk. & M.A. Curtis. An. Mag. Nat.
Hist. II, 12: 422. 1853.
Crepidotus. nephrodes
(Berk. & M.A. Curtis) Sacc. Syll.
Fung. 5: 882. 1887.
misappl.:
C. nephrodes s.
Horak (1964), Raithelhuber
(1988), Singer (1947, 1949, 1953, 1973, 1986), Singer and
Digilio (1951), Wright and Alberto´ (2002).
C. subaureifolius
Hesler&A.H.Sm.N.Am.Sp.
Crepidotus
, p. 74. 1965.
Claudopus subnidulans
Overh., Ann Mo Bot Gard 3:195.
1916.
Crepidotus subnidulans
(Overh.) Hesler & A.H. Sm. N.
Am. Sp.
Crepidotus
, p. 71. 1965.
For additional synonyms see Hesler and Smith (1965)
and Singer (1973).
Pileus
(9–)11–40 mm broad, somewhat pulvinate in
primordial stages, gradually hemispheric or ungulate,
becoming convex to plano-convex, applanate mainly
in over-mature individuals, circular, irregularly circu-
FIG.1.
Crepidotus crocophyllus
. Basidiocarps. a. Bandala 3908. b. Bandala 3963. Bar: 5 20 mm.
336 MYCOLOGIA
lar, flabelliform or rounded flabelliform, at times
more or less petaloid, spathuliform or reniform, with
or without a slot in the rear portion (seen from the
hymenophore), this forming two short, lobe-like
hemispheres; surface whitish, yellowish-white, cream
yellow (near 4A 2–3) with or without melon shades,
yellowish (near 10YR 8/3–4), pale grayish-yellow
(10YR 7–8/3) or pale yellowish-brown, darker with
age, covered with brown, yellowish-brown, brown-
orange, dark orange or reddish-brown fibrils, then to
the naked eye the pileus appearing rusty (near 2.5YR
4/6–8), reddish-brown (5YR 4/6), orange (7.5YR 6/
8), orange-brown (7.5YR 5/6), pale orange brown (6C
5–7), yellowish brown (5C4, 5C6), brown or brownish
(5C5, 6D5), pale or dark brownish-orange (7.5YR 6/4,
4/6; 6E6, 6D7–8, 6E8), in elements with glabrescent
surface most part of the disk is whitish, white-grayish
or pale grayish-yellow being on or near the rear
portion pale brown pigmented by the relative
abundance of fibrils; the fibrils vary in size and
density on overall surface, then when young more
often fibrillose-scaly or almost as a continuous,
pigmented layer toward the rear portion, becoming
tomentose-fibrillose, scattered fibrillose, densely fi-
brillose, appressedly fibrillose-scaly, minutely scaly or
squamulose, occasionally semiglabrous; dry, rarely
hygrophanous; margin initially involute, soon in-
curved to inflexed, finally straight, more or less plane,
faintly transparently striate in over-mature and wet
samples.
Lamellae
whitish or white-yellowish to pale
yellow (near 2.5Y 8/4) or cream yellow (near 10YR 8/
4; 4A4), becoming grayish-yellow, pinkish-yellow to
melon (5A3), pinkish-orange (7.5YR 7/4–6), yellow-
ish-orange (10YR 8/6), pale orange or orangish
(7.5YR 6/6, 8/6), yellowish-brown, grayish (5C3) or
grayish-brown (pale 4B2), pinkish buff (6B3), pinkish-
brown, pale brown or brownish-orange (7.5YR 6–7/
4), with white or whitish, fimbriate, somewhat
FIG.2.
Crepidotus crocophyllus
. a–c. H. Forstinger 51492 (a. Basidiospores. b. Pileipellis. c. Cheilocystidia). d–g. L.R. Hesler
24851, holotype of
C. appalachianensis
. d. Basidiospores. e. Hymenial elements on lamella face. f. Pileipellis. g. Cheilocystidia.
Bar 5 10
mm, except b and f 5 20 mm.
BANDALA ET AL:
C
REPIDOTUS CROCOPHYLLUS
337
irregular edges; adnexed to narrowly adnate, some
faintly subdecurrent, concurrent to a lateral point,
close to crowded, occasionally subdistant, moderately
broad (#6 mm), more or less subventricose to
ventricose, lamellulae 5–6 different lengths.
Stipe
in
primordial and young stages lateral, rudimentary
(,2 mm long.), whitish, pruinose, at times faintly
fibrillose or villose, then brownish or dark grayish,
with age absent or persisting as a lateral, glabrous
knob (seen from the hymenophore), the pileus
directly attached laterally or almost dorsally to the
substratum; basal mycelium white, present or absent.
Context
white or whitish, brownish or orange-brown
towards pileipellis, moderately thick (0.5–2.5 mm) at
pileus center, up to 5 mm near the attachment, soft,
unchanging on exposure.
Odor
and
taste
not distinc-
tive.
Basidiospores
(4.5–)5–7.5(–8) 3 4.5–7(–8) mm,
x
5
5.5–6.9 3 5.5–6.8
mm,
Q
5 1.00–1.03, globose to
subglobose, spinulose to spinulose-verruculose or at
times verrucose under immersion lens, baculate when
seen under SEM, at times the bacules weakly acute or
in other cases short and somewhat conic; yellowish to
yellowish-brown, moderately thick-walled (#0.5
mm).
Basidia
15–45(–48) 3 5–9(–10) mm, clavate to
narrowly clavate, 4-spored, hyaline, clamped.
Pleur-
ocystidia
absent; among the hymenial elements some
collections occasionally present sterile bodies resem-
bling amorphous or abortive basidia or anomalous
basidioles appearing as cystidioid-like structures (cf.
F
IGS. 2e, 3b, g, 5g).
Cheilocystidia
(20–)24–75 3 5–
10(–12)
mm, numerous, clavate, narrowly clavate or
subclavate to more or less narrowly utriform or
narrowly lageniform, at times subcylindric, somewhat
flexuous or constricted, apex (4–)5–18(–20)
mm wide,
rounded, subcapitate, rarely tapered or branched,
hyaline thin-walled, clamped.
Pileipellis
a cutis con-
sisting of cylindric to somewhat ventricose, septate,
FIG.3.
Crepidotus crocophyllus
. a–d. A.H. Smith 67333, holotype of
C. aureifolius
. a. Basidiospores. b. Hymenial elements on
lamella face. c. Pileipellis. d. Cheilocystidia. e–g. A.H. Smith 67160, holotype of
C. distortus
. e. Basidiospores. f. Pileipellis. g.
Hymenial elements on lamella face. Bar 5 10
mm, except c and f 5 20 mm.
338 MYCOLOGIA
pale yellowish, yellowish, pale yellow or pale yellowish-
brown hyphae (yellowish-brown to brownish-orange
in group) 4–15(–20)
mm wide, forming the pigmented
fibrils or scales of pileus surface, thick-walled, 0.5–
1.0(–1.5)
mm, smooth or finely to coarsely incrusted,
at times the incrustations producing discontinuous
lines more or less transversely or spirally oriented,
with a variable number of terminal elements, these
undifferentiated, often tapering apically or somewhat
narrowly lageniform or narrowly utriform, the layer
varying in thickness and density of hyphae, often
interrupted, at times loosely arranged and in some
areas most hyphae moderately ascending, then almost
as a transition between a loose cutis and a trichoder-
moid pileipellis.
Pileus trama
with hyaline, cylindric to
subventricose, moderately compactly interwoven,
thin-walled hyphae 3–20
mmwide.
Hymenophoral
trama
subregular to subirregular, hyphae 3–10(–18)
mm wide, cylindric to subventricose, thin-walled,
hyaline.
Clamp connections
present in all tissues.
Habitat.
Gregarious, subgregarious, scattered or
solitary, on dead bark, rotten log, rotten branches or
decaying wood, in tropical and subtropical cloud forest
at 350–1700 m alt.
Specimens examined
: COSTA RICA. PUNTARENAS:
A
´
rea de Conservacio´n La Amistad Pacı´fico, Z.P. Las Tablas,
Cotoncito, 17 Apr 1999,
Navarro 962
; Finca Cafrosa, Fila
Chiquiza´, 9 Jul 1999,
Lo´pez 556
; Sendero Central, 24 Jun
1999,
Oses 346
; Sendero Progreso, 30 Sep 2000,
Mata 898
.
GUANACASTE: A
´
rea de Conservacio´n Arenal, P.N. Volca´n
Tenorio, Hacienda Montezuma, 5 May 2000,
Lo´pez 1296-2
.
ALAJUELA: A
´
rea de Conservacio´n Arenal, P.N. Volca´n
Tenorio, Alto Los Brenes, 19 May 1999,
Lo´pez 370
(all at
INB). MEXICO. VERACRUZ: Freeway Fortı´n-Orizaba,
Barranca de Metla´c, 26 Aug 1989,
Bandala 1854
; Km 2.5
FIG.4.
Crepidotus
basidiospores (SEM). a–b.
Crepidotus applanatus
(C.J. Sprague 960, holotype of
Agaricus malachius
). c–l.
Crepidotus crocophyllus
(c. M.A. Curtis 1912, holotype of
Agaricus nephrodes
. d. L.R. Hesler 24851, holotype of
C.
appalachianensis
. e. A.H. Smith 66275. f. A.H. Smith 67333, holotype of
C. aureifolius
. g. A.H. Smith 67160, holotype of
C.
distortus
. h–i. H. Beach 28, holotype of
C. subaureifolius
. j. L.O. Overholts 13045, holotype of
Claudopus subnidulans
. k. T.G.
Lea s.n., holotype of
Agaricus crocophyllus
. l. H. Forstinger 51492). Bar 5 1 mm, except l 5 2 mm.
BANDALA ET AL:
C
REPIDOTUS CROCOPHYLLUS
339
old road Xalapa-Coatepec, wooded area of Jardı´n Bota´nico
Fco. J. Clavijero, 30 Jan 1996,
Bandala 2919
; 10 Jan 1997,
Mun˜oz 10
; 25 Jun 2003,
Bandala 3762
; Sta. Ine´s Ranch, 23
May 1991,
Tapia 545
; Instituto de Ecologı´a, Santuario del
Bosque de Niebla, 3 Mar 2005,
Jarvio 2005
; 3 Nov 2005,
Bandala 4029
; 2 km W of Xalapa, near Coapexpan River,
10 June 2004,
Bandala 3908, 3909-B
;27June2005,
Bandala 3963
; Los Tuxtlas Region, near Catemaco, 4 Aug
1997,
Mun˜oz 30
(all at XAL).
Other specimens examined
: AUSTRIA. OBERO
¨
STER-
REICH: Molln, Iner-Beritenau, 21 Sep 1986,
H. Forstinger
51492
(WU 5628 as
C. crocophyllus
). U.S.A. MICHIGAN:
Chippewa Co., Emerson, 12 Aug 1963,
A.H. Smith 67160
(MICH, Holotype of
C. distortus
); Cheboygan Co., 10 Jul
1946,
T.E. Brooks 1249
(as
C. crocophyllus
); Oakland Co.,
Haven Hill, 1 Sep 1963,
A.H. Smith 67333
(MICH,
Holotype of
C. aureifolius
); Wayne Co., Miller’s Woods, 20
Jun 1978,
A.H. Smith 88575
(as
C. crocophyllus
); Luce-
Chippewa Co., Tahquamenon Falls State Park, 16 Jul 1951,
A.H. Smith 36934
(as paratype of
C. aureifolius
); 6 Jul 1955,
A.H. Smith 49741
; 11 Aug 1955,
H. Beach 28
(Holotype of
C. subaureifolius
),
31
(as paratype of
C. aureifolius
) (all at
MICH). MISSOURI: near St Louis, Jefferson Barracks, 25
Oct 1913,
L.O. Overholts 13045
(BPI, Holotype of
Claudopus subnidulans
). NEW HAMPSHIRE: Hillsburo
Co., Fox Forest, 8 Sep 1959,
O.K. Miller 569
(MICH as
C. nephrodes
). OHIO: Waynesville, 5 Sep 1844,
T.G. Lea
s.n.
(K, Holotype of
Agaricus crocophyllus
). SOUTH
CAROLINA:
M.A. Curtis 1912
(K, Holotype of
Agaricus
nephrodes
). TENNESSEE: The Great Smoky Mountains
National Park, Cades Cove, 2 Jul 1962,
L.R. Hesler 24851
(TENN, Holotype of
C. appalachianensis
). VIRGINIA: Falls
Church, 2–6 Jul 1904,
W.A. Murrill 104
(NY, Holotype of
Crepidotus fulvifibrillosus
).
Notes.
Distinctive features of
C. crocophyllus
include
a pileus variably covered with brown-pigmented fibrils
or squamules, moderately large, globose and baculate
(spinulose or spinulose-verruculose under oil lens)
basidiospores, variably clavate or subclavate cheilocys-
tidia, and a pileipellis formed of repent, yellow to
brown-pigmented, thick-walled, often incrusted hy-
FIG.5.
Crepidotus crocophyllus
. a–d. H. Beach 28, holotype of
C. subaureifolius
a. Basidiospores. b. Hymenial elements on
lamella face. c. Pileipellis. d. Cheilocystidia. e–g. L.O. Overholts 13045, holotype of
C. subnidulans
e. Basidiospores. f.
Pileipellis. g. Hymenial elements on lamella face. Bar 5 10
mm, except c & f 5 20 mm.
340 MYCOLOGIA
phae. Several descriptions of this species inform about
the plasticity of specimens regarding the size and
variation in color of pileus and hymenophore through
its different developmental stages (Barron 1999,
Berkeley 1847, Bon and Massart 1996 as
C. nephrodes
;
Eryssartier 1997, Hesler and Smith 1965, Horak 1964,
Laze´bnicˇek 1970, Moser 1983, Murrill 1917, Pegler
1977 as
C. nephrodes
; Pereira 1990, Pila´t 1950, Phillips
1991, Raithelhuber 1988, Ripkova´ et al 2005, Senn-
Irlet 1995, Singer 1973, Wright and Alberto´2002).
Crepidotus crocophyllus
embraces transitional forms of
individuals at times almost glabrous in appearance,
with scattered, innate fibrils that may or may not be
conspicuous to the naked eye or form distinctly
pigmented squamules, hence the pale ground surface
along with the pigmented fibrils producing differences
in the overall aspect of pileus surface (pigmentation
and texture) (cf. F
IG. 1 and Moser and Ju¨ lich 1986).
Even the dried specimen from Ohio
T.G. Lea s.n.
,
holotype of
Agaricus crocophyllus
, is scarcely fibrillose,
showing sparsely brown-pigmented fibrils (under
lens). Microscopically the pileipellis varies consequent-
ly in the density of the most superficial hyphae but
these are consistently pigmented, often incrusted and
thin- to thick-walled, varying therefore the color
intensity of the complete layer (the type specimen of
C. fulvifibrillosus
, for example has a thin, interrupted
or at times poorly represented layer of yellow to pale
yellowish, thick-walled hyphae). The type of cutis
composed of yellow to brown pigmented, often
incrusted, thick-walled hyphae is regarded in different
lineages of
Crepidotus
suggesting that it is a taxonom-
ically significant and distinctive feature (e.g. as found
in specimens of
C. calolepis
[Fr.] P. Karst.,
C. kaufmanii
Hesler & A.H. Sm. or
C. rainierensis
Hesler & A.H. Sm.
[Bandala and Montoya 2004, Gonou-Zagou and
Delivorias 2005, Hesler and Smith 1965, Senn-Irlet
1995, Singer 1973]). It is important to note, however,
that the interpretation of the pileipellis hyphae, as
pigmented or not, in any specimen examined varies
depending on the pileus area studied and their
relative abundance in such points (this is due to the
pattern of occurrence of the hyphae on the surface).
In specimens of
C. applanatus
parts of the pileipellis
bear at times a mixture of hyaline and yellow or pale
yellow hyphae (cf. Senn-Irlet 1995, Senn-Irlet and De
Meijer 1998, pers. obs.) which however, are distinct
from those even forming the most poorly differentiat-
ed pileipellis of specimens of
C. crocophyllus
.Among
the collections examined we did not find any with a
pileipellis made of exclusively pale pigmented or even
hyaline hyphae. Variation of the pattern of basidos-
pore ornamentation can be observed when it is
analyzed under SEM. In a single sample for example
some or several spores can exhibit short, rounded or
truncate projections, these can even be interconnected
forming more or less coarse bulges and so appearing
warty or verrucose instead of decidedly baculate.
Under light microscope the spore then can appear
smooth to punctate or verrucose more than spinulose
or spinulose-verrucose. We have observed this situation
in some specimens examined, for example the
collections of
H. Beach 28
(holotype of
C. subaur-
eifolius
)and
H. Forstinger 51492
(FIG. 4), or many
others specimens related to
C. applanatus
or
C.
aquosus
Murrill. A number of spores in the sample
of
L.O. Overholts 13045
(holotype of
C. subnidulans
)
(F
IG. 4) also showed such variation. This characteristic,
apparently common among
Crepidotus
species pro-
ducing globose, baculate basidiospores, also has been
reported by Bigelow (1980) for
C. nyssicola
(Murrill)
Singer and it may be inferred from the information
given by Pegler and Young (1972) and Senn-Irlet
(1995) on basidiospores of
C. applanatus
.Such
variation seems to be attributable to the individual
differentiation during spore maturation, hence care
should be taken in considering it of taxonomic value
among collections showing differences in the process
of ornamentation development. Another characteristic
apparently common among taxa exhibiting globose,
baculate basidiospores is the presence of basal ridges
interconnecting the rod-like protuberances of the
basidiospores, then several spores in a single collection
at times appear more or less spinulose-rugulose as also
observed by Pegler and Young (1972) and Senn-Irlet
(1995) on basidiospores of
C. applanatus
.
Owing to subtle differences in color (of lamellae
and pileus) and the presence of the so-called
pleurocystidia, Hesler and Smith (1965) recognized
C. appalachianensis
,
C. aureifolius
,
C. distortus
,
C.
subaureifolius
and
C. subnidulans
as taxonomically
distinct from
C. crocophyllus
. The re-examination of
type materials of these species revealed that all share a
consistent combination of taxonomically important
micromorphological characters (globose, baculate
basidiospores, pileipellis as a cutis bearing pigment-
ed, thick-walled, often incrusted hyphae, cheilocysti-
dia commonly clavate, clamped hyphae) that fall
however within variation seen in
C. crocophyllus
(FIGS.2–5, TABLE I). As observed in our fresh
collections and according to different descriptions
of this taxon, the apparently discrete stages recog-
nized by Hesler and Smith (op. cit.) are not consis-
tent, we concluded therefore that the specimens
represent different stages of the macroscopic varia-
tion of
C. crocophyllus
. A close similarity between
C.
aureifolius
and
C. subaureifolius
in fact was indirectly
stressed by Hesler and Smith (1965: 75–76), when
under both epithets they cited the same specimen
H.
Beach 28
(the type of
C. subaureifolius
). Sterile
B
ANDALA ET AL:
C
REPIDOTUS CROCOPHYLLUS
341
bodies, which seem to be abnormal basidia or
basidioles, certainly were encountered at times in
the hymenium (F
IGS. 2e; 3b, g; 5b, g). Such structures
in our opinion could hardly be recognized as true
cystidia (pleurocystidia) (i.e. as a reliable taxonomic
character to separate the samples). In the hymenium
of isolated samples representing different
Crepidotus
species, we and other authors (Bandala and Montoya
2000a, Gonou-Zagou and Delivorias 2005, Senn-Irlet
1995) have detected similar sterile elements that
however seem to reflect at most an extreme in a
continuous range of variation rather than to distin-
guish taxonomically discrete entities. Singer (1973)
and Senn-Irlet and De Meijer (1998: 180) also have
expressed their doubts about the taxonomic rele-
vance of this kind of structures called pleurocystidia.
Crepidotus
is still conceived as a lineage of members
lacking pleurocystidia (Aime et al 2005, Nordstein
1990, Pila´t 1948, Senn-Irlet 1995, Singer 1973, 1986,
Watling and Gregory 1989).
In the absence of one or more distinctive taxo-
nomic features no arguments support the taxonomi-
cal segregation of collections phenotypically identical.
Furthermore there is evidence of intercompatibility
between samples identified under some of the
aforementioned epithets (Aime 2004). Our results
additionally provide useful information for taxonom-
ic resolution of the
Fulvifibrillosi
clade obtained by
Ripkova´ et al (2005) where specimens treated under
different names (but belonging indeed to the same
taxon) are tightly clustered and key out with
C.
crocophyllus
as herein interpreted.
Agaricus malachius
and
Agaricus nephrodes
, two species long related to
Crepidotus crocophyllus
and cited in such clade,
deserve to be discussed separately (see below under
C. malachius
).
Crepidotus badiofloccosus
S. Imai (1939)
from Japan has been described as differing from
C.
rubriflavus
Murrill (1917) (a new name for
C. dorsalis
Peck [non Bosc] [Halling 1986], this latter being a
synonym of
C. crocophyllus
[Singer 1947, Hesler and
Smith 1965]) by subtle differences in basidiospore
ornamentation (asperulate). On account of charac-
ters mentioned in the diagnosis and data afterward
provided by Imazeki and Hongo (1987) and Imazeki
et al (1988),
C. badiofloccosus
appears to be indistin-
guishable from
C. crocophyllus
.
On the basis of the present information it can be
recognized that
C. crocophyllus
has a wide but
fragmented range of distribution throughout the
Americas. In the USA the species has been recorded
mainly in eastern states (in the west it is known in
Oregon and Idaho) almost coastal from New Hamp-
shire to the Gulf of Mexico and now southward to
Central America. In South America
C. crocophyllus
is
known from Bolivia, Brasil and Argentina (Hesler and
Smith 1965, Horak 1964, Pereira 1990, Raithelhuber
1988, Singer and Digilio 1951, Singer 1973, Wright
and Alberto´ 2002).
Crepidotus malachius
(Berk. & M.A. Curtis) Sacc., Syll
Fung 5:883. 1887. F
IG. 4a–b
Bas.:
Agaricus malachius
Berk. & M.A. Curtis, An Mag Nat
Hist III, 4:291. 1859.
HOLOTYPE. U.S.A. NEW ENGLAND: Aug 1856,
C.J.
Sprague 960
(K, Herb. Curtis 5730; Isotype FH).
Basidiospores
(4.5–)5.5–7(–8) 3 (4.5–)5.5–7 (–8)
mm,
x
5 5.5–6.5 3 5.5–6.3 mm,
Q
5 1.00–1.03, globose
to subglobose, spinulose to spinulose-verruculose
seen under light microscope, baculate when observed
under SEM, the bacules at times more or less conic,
small verrucae can be present among the bacules;
yellowish to yellowish-brown, moderately thick-walled
(#0.5
mm thick).
Basidia
clavate, 4-spored, hyaline,
clamped.
Pleurocystidia
absent.
Cheilocystidia
al-
though not completely recovered by the condition
of the dried specimen, proved to be clavate, hyaline,
thin-walled, clamped.
Pileipellis
a cutis composed of
more or less compactly arranged, hyaline, thin-walled,
cylindric or somewhat ventricose hyphae 5–15
mm
diam, terminal elements scarce, undifferentiated.
Pileus trama
with cylindric to subventricose, thin-
walled, hyaline, hyphae.
Hymenophoral trama
subre-
gular, hyphae similar to the pileus trama.
Clamp
connections
present in all tissues.
Notes.
The type specimen of this species as well as
that of
Agaricus nephrodes
have been re-examined by
different authors (Hesler and Smith 1965, Pegler 1977,
Pila´t 1950, Singer 1947, 1973) who have offered
different taxonomic interpretations. Then prevailing
concepts of
Crepidotus crocophyllus
and
C. fulvifibrillo-
sus
in connection with
C. applanatus
(cf. the
discussions by Hesler and Smith [1965: 79], Pila´t
[1948] and Singer [1947, 1973: 374, 382]) influenced
early taxonomic decissions. Two hypotheses remain in
the literature. One proposes
Agaricus nephrodes
as a
name having priority and as a species embracing
A.
malachius
, hence placing it closely related to
Crepido-
tus applanatus
(Horak 1964, 1979, Murrill 1917, Pila´t
1950, Raithelhuber 1988, Singer 1947, 1949, 1953,
1973, 1986, Singer and Digilio 1951, Wright and
Alberto´ 2002). The other explores the inclusion of
C.
nephrodes
in the group of
C. crocophyllus
(i.e. within
the
Fulvifibrillosi
s. Hesler and Smith above discussed),
either as a taxon conspecific with
C. crocophyllus
(Ripkova´ et al 2005) or as a separate entity (Bon and
Massart 1996, Grgurinovic 1997, Hesler and Smith
1965, Natarajan and Raman 1983, Pegler 1977, Ueki
and Smith 1973). Our revision of type specimens and
fresh and herbaria samples related to
C. applanatus
,
C.
malachius
,
C. nephrodes
and
C. crocophyllus
reveal that
342 M
YCOLOGIA
TABLE I. Comparison of microscopic characters among type collections of taxa related to
Crepidotus crocophyllus
C. appalachianensis C. aureifolius C. crocophyllus C. distortus C. fulvifibrillosus C. subaureifolius C. subnidulans
Basidiospores
size (
mm) 4.5–6(–6.5) (4.5–)5–8 5.5–6.5 5.5–7 (5–)5.5–6.5 5–6.5 (5.5–) 6–7.5
SL 6 SD
a
5.5 6 0.48 6.5 6 0.70 6.0 6 0.38 6.4 6 0.46 6.0 6 0.39 5.7 6 0.45 6.8 6 0.53
SW 6 SD 5.5 6 0.49 6.5 6 0.70 6.0 6 0.32 6.3 6 0.43 5.9 6 0.34 5.7 6 0.45 6.7 6 0.52
Cheilocystidia
size (
mm) 20–43 3 4–7 21–44 (–56) 3 5–9 34–55 3 5–8 * (28–)37–55 3
(4–)5–8(–10)
(25–)27–54 3
5–10(–12)
*
Ch-apex (
mm)
b
4–9 6–10 5–9(–10) * (6–)7–9.5(–11) (6–)7–11 *
Hymenium
Sterile bodies on
lamellae side
c
occasional
inconspicuous
sporadic 6
conspicuous
not
differentiated
scattered
conspicuous
not differentiated occasional
inconspicuous
scattered 6
conspicuous
Pileipellis
hyphae (diam
mm) (5–)6–15 6–18 4–12 8–14 5–12 (7–)8–15(–17) 7–13 (–18)
finely incrusting
pigment
often present present often present present present present present
hyphae color yellowish-brown to
brownish-orange
yellow or yellowish
to yellow-brown
yellowish to
pale yellowish
yellow to
yellowish-brown
yellow to pale
yellowish
yellow to yellowish-
brown
yellow to
yellowish-brown
a
SL and SW refers to the means of both spore length and width respectively 6 standard deviation based on 50 spores per collection.
b
Ch-apex refers to the cheilocystidia apex width.
c
refers to the apparently abnormal basidia which resemble cystidioid-like structures (so-called pleurocystidia).
*
the lamellae edges of
C. distortus
type specimen are collapsed whereas the type of
C. subnidulans
is in a fragmentary condition, in both cases preventing to recover the
cheilocystidia.
BANDALA ET AL:
C
REPIDOTUS CROCOPHYLLUS
343
in some point of the macroscopic variation (size, shape
and color of the basidiomes) even among the elements
of a single collection there can be transitional stages.
Except for the presence or absence of pigmented
hyphae in the pileipellis the aforementioned
taxa certainly share the same distinctive pattern
of microscopic characteristics that often makes
their differentiation difficult (i.e. pileipellis in
cutis, globose, baculate [punctate to spinulose or
spinulose-verrucose seen under oil lens] basidio-
spores [F
IG. 4]), cheilocystidia often clavate and
clamped hyphae (the set of features in fact of a
number of taxa around
C. applanatus
within the
Sphaerula
) (Hesler and Smith 1965, Senn-Irlet 1995,
Singer 1973).
Both
C. applanatus
and
C. malachius
were de-
scribed as devoid of any pigmented covering on pileus
surface, and when describing
Agaricus malachius
Berkeley and Curtis (1859) emphasized its resem-
blance to
A. nephrodes
. In the original description by
Berkeley and Curtis (1853)
Agaricus nephrodes
is
characterized by its pileus ‘‘…pallide flavido-tomen-
toso…’’, hence, ‘‘…clothed with dingy yellowish-white
down…’’. It apparently differs from fibrillose or
squamulose pigmented forms such as
C. crocophyllus
which Berkeley (1847) previously described (under
Agaricus
) with pileus ‘‘…ochraceo-fusco adpresse
squamoso…’’, then, ‘‘…clothed with minute ad-
pressed scales…’’. Pila´t (1948) and Horak (1979) in
part suggested the conspecificity between
C. nephrodes
and
C. applanatus
.Pila´t (1950) interpreted
C.
nephrodes
as the same as
C. applanatus
, whereas
Singer (1947, 1949, 1953, 1973, 1986) related it to
C.
malachius
. Based on the presence or absence of
brown pigmented fibrils or squamules on pileus
surface, Singer (1947) was the first to suggest the
differences between
C. applanatus
and the North
American
C. fulvifibrillosus
, which years later was
recognized as a synonym of
C. crocophyllus
(Eryssartier
1997, Ripkova´ et al 2005, Senn-Irlet 1995, Singer
1973, also hereafter type study). Murrill (1917) clearly
separated his species from
C. nephrodes
, interpreting
this as having a yellowish pileus with a tomentose
surface versus the uniformly dull-white pileus with a
tawny, fibrillose-scaly surface of
C. fulvifibrillosus
.
Dissimilarity in such pileal characteristics perhaps
was the reason why Pila´t (1948) placed
C. fulvifi-
brillosus
as a variety of
C. applanatus
. Singer (1949,
1986) confirmed his previous interpretation of
C.
applanatus
,
C. crocophyllus
,
C. fulvifibrillosus
and
C.
nephrodes
(this including
C. malachius
as its synonym)
as separate entities. Horak (1964), Raithelhuber
(1988), Singer and Digilio (1951) and Wright and
Alberto´ (2002) also treated
C. crocophyllus
and
C.
nephrodes
as different species. Hesler and Smith
(1965) accepted separately
C. applanatus
,
C. croco-
phyllus
,
C. malachius
and
C. nephrodes
, using the
latter to include Murrill’s
C. fulvifibrillosus
as a
synonym. Therefore the data reveal that one con-
cept of
C. nephrodes
, in apparent coincidence with
the original description, encompasses members de-
void of any pigmented fibrillosity on pileus surface
hence embracing
C. malachius
and close to
C.
applanatus
. Another circumscribes a taxon with
fibrillose to squamulose pileus, then close to
C.
crocophyllus
and excluding
C. malachius
.This
latter criterion introduced by Hesler and Smith
(1965) was followed by other authors (Bon and
Massart 1996, Grgurinovic 1997, Natarajan and
Raman 1983, Pegler 1977, Ripkova´ et al 2005, Ueki
and Smith 1973).
In an attempt to determine the application of the
name
Agaricus malachius
we re-examined its type
collection as well as that of
Agaricus nephrodes
(
M.A.
Curtis 1912
)(see
Crepidotus crocophyllus
above)
(F
IG. 4a–c). Our results strongly support the observa-
tions of Hesler and Smith (1965) and part of the
arguments of Ripkova´ et al (2005) based on samples
from Europe and North America but differ from Pila´t
(1948, 1950), Horak (1964, 1979) and Singer (1947,
1949, 1953, 1973) with regard to the description of
the gross morphology of pileus surface. We found
that macro- and microscopically the specimen of
C.J.
Sprague 960
supporting
Crepidotus malachius
is
morphologically different from
C. nephrodes
,the
former being phenotypically more similar to
C.
applanatus
than to
C. crocophyllus
. The pileus of the
Agaricus nephrodes
type collection when analyzed
under magnification possesses fine, tiny, weakly
pigmented and separately spaced fibrils making the
surface appear almost glabrous (almost in conformity
with that formerly described by Berkeley and Curtis
1853). Although its tissues are not easily recovered in
KOH, there is evidence of the presence of pale
yellowish, yellow or pale brownish (yellowish-brown in
whole), thick-walled hyphae that occasionally are
found in the pileipellis (relatively more frequent
toward the point of pileus attachment). The basidio-
spores in the
M.A. Curtis 1912
specimen are (5–)6–
7(–7.5) 3 5.5–7(–7.5)
mm,
x
5 6.6 3 6.5 mm,
Q
5 1.02
(F
IG. 4c). The lamellae edges are collapsed and little
information is obtained about the cheilocystidia, they
are however clavate, hyaline, thin-walled, clamped.
The pileipellis hyphae are comparatively similar to
those found in the type (
T.G. Lea s.n.
) and other
examined samples of
C. crocophyllus
. Based on their
pattern of occurrence on the pileus surface, as well as
correlating this feature with the basidiospore size, we
must recognize that the specimen of
M.A. Curtis
1912
represents an extreme stage of the morpholog-
344 M
YCOLOGIA
ical variation of
C. crocophyllus
(i.e. a member with
barely fibrillose pileus). On the other hand the
Agaricus malachius
type collection
C.J. Sprague 960
(holotype and isotype) differs from
A. nephrodes
mainly in having a less pigmented pileus, with a
pileipellis composed of hyaline, thin-walled hyphae
(i.e. pileus without [pigmented] fibrils or squamules
already described by Berkeley and Curtis [1859] and
basidiospores slightly smaller in average).
Taking into account all the aforementioned
information and available data after our re-examina-
tion of type material,
Crepidotus crocophyllus
and
Agaricus nephrodes s. str.
Berkeley are recognized
as taxonomically identical also including other
synonyms cited in the literature (see
C. crocophyllus
above). We interpret Berkeley’s taxon as excluding
Agaricus malachius
, which on account of its smooth,
white, cuneiform, subflabellate pileus, gills at
first white then yellow-brown (Berkeley and Curtis
1859), in combination with the above mentioned
microscopic characteristics seen on type material,
support that it should be considered conspecific with
Crepidotus applanatus
, and therefore we propose its
synonymy:
Crepidotus applanatus
(Pers.) P. Kumm. Fu¨hr Pilzk
p. 74. 1871.
Bas.:
Agaricus applanatus
Pers. Obs Mycol 1:8. 1796.
Syn.:
A. malachius
Berk. & M.A. Curtis, An Mag Nat Hist
III, 4:291. 1859.
Crepidotus malachius
(Berk. & M.A. Curtis) Sacc. Syll
Fung 5:883. 1887.
In fact
C. malachius
(i.e.
C. applanatus
) appears to
occupy a distinct clade in suggested groupings by
Ripkova´ et al (2005) where
C. crocophyllus
(there
including
C. appalachianensis
,
C. distortus
and
C.
nephrodes s.
Hesler and Smith) represents a sister
group. For further differences between
C. applanatus
and
C. crocophyllus
see Senn-Irlet (1995).
ACKNOWLEDGMENTS
We appreciate the loan of specimens from herbaria FH, K,
MICH, NY, TENN and WU. We thanks Dr Z. Gonou-Zagou
(University of Athens) for reviewing the manuscript and
providing valuable comments. Part of this work was
supported by CONACYT (Project 52364-Q to V.M. Ban-
dala). Two anonymous reviewers provided critical observa-
tions to improve the manuscript.
LITERATURE CITED
Aime MC. 2004. Intercompatibility tests and phylogenetic
analysis in the
Crepidotus
Sphaerula group complex:
concordance between ICGs and nuclear fDNA sequenc-
es highlight phenotypic plasticity within Appalachians
species. In: Cripps C, ed. Fungi in forest ecosystems:
systematics, diversity and ecology. New York: New York
Botanical Garden. 363 p.
———, Vilgalys R, Miller OK. 2005. The Crepidotaceae
(Basidiomycota, Agaricales): phylogeny and taxonomy
of the genera and revision of the family based on
molecular evidence. Am J Bot 92:74–82.
Bandala VM, Montoya L. 2000a. A taxonomic revision of
some American
Crepidotus
. Mycologia 92:341–353.
———, ———. 2004.
Crepidotus
from Mexico: new records
and type studies. Mycotaxon 89:1–31.
———, ———, Moreno G. 1999. Two
Crepidotus
from
Mexico with notes on selected type collections.
Mycotaxon 72:403–416.
———, ———, Horak E. 2006.
Crepidotus rubrovinosus
sp.
nov. and
Crepidotus septicoides
, found in the cloud
forest of eastern Mexico, with notes on
Crepidotus
fusisporus
var.
longicystis
. Mycologia 98:137–146.
Barron G. 1999. Mushrooms of northeast North America.
Canada: Lone Pine. 336 p.
Berkeley MJ. 1847. Decades of fungi XII–XIV. London J Bot
6:312–326.
———, Curtis MA. 1853. Centuries of North American
fungi XLI. Ann Mag Nat Hist ser II 12:417–435.
———, ———. 1859. Centuries of North American fungi
XXXI. Ann Mag Nat Hist ser III 4:284–296.
Bigelow HE. 1980.
Crepidotus nyssicola
. Mycologia 72:1227–
1231.
Bon M, Massart F. 1996. Deux espe`ces Americaines
decouvertes dans le sud-ouest de la France. Doc Mycol
26(103):29–32.
Eryssartier G. 1997. Quelques taxones interessants recoltes
en Dordogne. Doc Mycol 26(104):1–6.
Gonou-Zagou Z, Delivorias P. 2005. Studies on Basidiomy-
cetes in Greece 1: the genus
Crepidotus
. Mycotaxon 94:
15–42.
Grgurinovic CA. 1997. Larger fungi of South Australia.
Adelaide: Botanic Garden Adelaide & St. Herb. & flora
and fauna of South Australia Handbooks Committee.
725 p.
Halling RE. 1986. Index to species and infraspecific taxa of
agarics and boletes described by William A. Murrill.
Mem NY Bot Gard 40:1–120.
Hesler LR, Smith AH. 1965. North American species of
Crepidotus
. New York: Hafner Publishing. 168 p, 33 figs.
Holmgren PK, Holmgren NH, Barnett LC. 1990. Index
Herbariorum. Part I. The herbaria of the world. 8th ed.
New York. 693 p.
Horak E. 1964. Fungi Austroamericani XI. Nov Hedwig 8:
333–346.
———. 1979. Fungi, Basidiomycetes Agaricales y Gaster-
omycetes Secotioides. In: Guarrera SA, Gamundi I,
Rabonovich D, eds. Flora Criptogamica de Tierra de
Fuego. 11(6). Buenos Aires. 523 p.
Imai S. 1939. Studia Agaricacearum Japonicarum I. Bot Mag
53:392–399.
Imazeki R, Hongo T. 1987. Colored illustrations of
mushrooms of Japan. I. Osaka: Hoikusha. 325 p.
———, Otani Y, Hongo T. 1988. Fungi of Japan. Tokyo:
Yama-kei. 326 p.
BANDALA ET AL:
C
REPIDOTUS CROCOPHYLLUS
345
Kornerup A, Wanscher JH. 1967. Methuen handbook of
colour. 2nd ed. London: Methuen.
Laze´bnicˇek J. 1970.
Crepidotus crocophyllus
(Berk.) Sacc., a
new species for the European mycological flora. C
ˇ
eska´
Mikol 24:78–85.
Moser M. 1983. Key to agarics and boleti (Polyporales,
Boletales, Agaricales, Russulales). London: Roger
Phillips. 670 p.
———, Ju¨ lich W. 1986. Farbatlas der Basidiomyceten III.
Stuttgart: Gustav Fischer. Munsell Soil Colour Charts
1994. New Windsor: Macbeth, 4 p, 9 charts.
Murrill WA. 1917. Agaricales. Agaricaceae. Agariceae. N Am
Fl 10(3):145–226.
Natarajan K, Raman N. 1983. South Indian Agaricales a
preliminary study of some dark spored species. Bibl
Mycol 89:7–203.
Nordstein S. 1990. The genus
Crepidotus
(Basidiomycotina,
Agaricales) in Norway. Synopsis Fungorum 2. Oslo:
Fungiflora. 115 p.
Pegler DN. 1977. A preliminary agaric flora of East Africa.
Kew Bull. Add. Ser. VI. London: HMSO. 615 p.
———, Young TWK. 1972. Basidiospore form in the British
species of
Crepidotus
. Kew Bull 27:311–323.
Pereira AB. 1990. O genero
Crepidotus
no Rio Grande do
Sul, Brasil. Cad Pesquisa (Bot) 2:65–85.
Pila´t A. 1948. Monographie des espe`ces europe´enes du
genre
Crepidotus
Fr. In: Pila´t A, Kavina K, eds. Atlas des
champignons de l’Europe 6. Praha. 84 p, 24 figs.
———. 1950. Revision of the types of some extra-European
species of the genus
Crepidotus
Fr. Trans Brit Mycol Soc
37:215–249.
Phillips R. 1991. Mushrooms of North America. Canada:
Little Brown. 319 p.
Raithelhuber J. 1988. Flora mycologica Argentina II.
Stuttgart: Mycosur. 500 p.
Ripkova´ S, Aime MC, Lizonˇ P. 2005.
Crepidotus crocophyllus
includes
C. nephrodes
. Mycotaxon 91:397–403.
Senn-Irlet B. 1995. The genus
Crepidotus
(Fr.) Staude in
Europe. Persoonia 16:1–80.
———, De Meijer A. 1998. The genus
Crepidotus
from the
state of Parana´, Brazil. Mycotaxon 66:165–199.
Singer R. 1947. Monograph of the genus
Crepidotus
. Lilloa
13:59–95.
———. 1949. The Agaricales (mushrooms) in modern
taxonomy. Lilloa 22:5–832.
———. 1953. The Agaricales of the Argentine sector of
Tierra del Fuego and limitrophous regions of the
Magallanes area. Sydowia 7:206–265.
———. 1973. The genera
Marasmiellus
,
Crepidotus
and
Simocybe
in the neotropics. Beih. Nov Hedwig 44:1–484.
———. 1986. The Agaricales in modern taxonomy. 4th ed.
Koeningstein: Koeltz Science Books. 981 p, 88 plates.
———, Digilio APL. 1951. Pro´dromo de la flora Argentina.
Lilloa 25:5–461.
Ueki R, Smith CW. 1973. The genus
Crepidotus
in Hawaii.
Can J Bot 51:1251–1254.
Watling R, Gregory NM. 1989. Crepridotaceae, Pleurotaceae
and other pleurotoid agarics. Brit Fung Fl 6. Edin-
burgh: Royal Botanic Garden. 157 p.
Wright JE, Alberto´ E. 2002. Hongos, guı´a de la Regio´n
Pampeana 1. Buenos Aires: Literature of Latin Amer-
ica. 279 p.
346 MYCOLOGIA
