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Instituto de Ecología, Biodiversidad y Sistemática, P.O. Box 63, Xalapa, Veracruz, Mexico
Egon Horak
Geobotanical Institute, Herbarium, Zollikerstrasse 107, CH-Zurich, Switzerland
| ABSTRACT |
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Two species of Crepidotus are recorded from cloud forest in the central region of Veracruz State (eastern Mexico): Crepidotus rubrovinosus sp. nov. and Crepidotus septicoides. The latter species was known previously only from the type locality in Brazil and from one record in tropical rain forest in southern Veracruz (as C. longicystis s. str. Singer). Descriptions, illustrations and discussions for both taxa are provided. A type study of C. fusisporus var. longicystis from USA is included, and it is concluded that the collection supporting this variety belongs to C. luteolus.
Key words: Agaricales, new synonymies, taxonomy, wood inhabiting fungi
| INTRODUCTION |
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In this paper two species of Crepidotus are presented that occupy different microhabitats in the cloud forests near Xalapa. Crepidotus rubrovinosus, proposed here as new, was found exclusively inside the forest growing on bark of a dead, unidentified hardwood tree. Collections of C. septicoides (Singer) Singer, however, seems to be restricted to the forest margin, where it was found colonizing dead leaves of Platanus mexicana Moric. In Mexico C. septicoides previously was known from tropical rain forest in southern Veracruz (Singer 1973
as "C. longicystis", Bandala et al 1999
). The present fresh specimens gathered in Xalapa inform about the taxons range of extension and revealed additional data especially on its macromorphologic variation.
| MATERIALS AND METHODS |
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corresponds to the range of means of length and width of 35 spores per collection. Q corresponds to the range of the average value of the ratio of basidiospores length/breadth computed per collection examined. The acronyms for the herbaria follow Holmgren et al (1990)| TAXONOMY |
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Etymology: rubrovinosus (Lat.) refers to the reddish, red or red-wine color of pileus and lamellar edges.
Pileus 520 mm broad, almost circular in primordial stages, becoming flabelliform to rounded flabelliform or semicircular, some more or less spathuli-form or petaloid, somewhat pulvinate when young, gradually convex, becoming plano-convex, dry, weakly hygrophanous, colored in various shades of red: vivid red or reddish (11E7) to dark or pale wine-red (11D67, 12C612DE67, 12D8), depending on age, desiccation or exposure the range of color can vary between red, pale vinaceous (11BC5) or pinkish (11B34), near the point of attachment white, yellowish or pinkish; surface villose-strigose, pubescent to tomentose-villose, somewhat felty, in old specimens at times glabrescent; margin in younger stages involute or in some elements at times attached to the incipient stipe, becoming inflexed to moderately straight, not striate, villose-strigose or tomentose-villose, then more or less appendiculate by such fibrillosity. Lamellae concurrent at point of attachment, adnate to narrowly sinuate, at times subdecurrent, subdistant to close, subventricose to more or less linear, 12(3) mm broad, rarely furcate, lamelullae of 23(4) different lengths; whitish to yellowish (near 7.5 yr 8/23, 7.5 yr 8/4), finally pale brown (10 yr 8/3; 2.5 yr 6/3), edges persistently red or reddish, distinctly fimbriate. Stipe absent in mature specimens, but in primordial stages a short, central, ventricose pseudostipe is present, 15 x 13 mm, with age a persisting, more or less lateral, irregular knob can be observed (as seen from hymenophore) but the pileus is directly attached to the substrate, strigose or villose both in young and old basidiomes, white, pinkish or pale reddish. Context white to yellowish, reddish or pinkish in or beneath pileipellis, thin (0.51 mm thick) to moderately thick (up to 2 mm) near point of attachment, soft, unchanging on exposure. Odor and taste not distinctive.
Spore print (seen on pileus surface) yellow-brown.
Basidiospores (5.5)68(8.5) x 4.56 µm,
= 6.57.5 x 4.95.7 µm, Q = 1.321.39, ovoid in lateral view, oblong or ellipsoid in dorso-ventral view, suprahilar depression (in lateral view) weakly developed, commonly with a distinctive, plage-like, smooth suprahilar area; strongly rugose, the ornamentation consisting of short, broad and sinuous, blunt bulges and ridges, often anastomosed and forming a subreticulate pattern; bright yellow to yellow, with darker ornamentation (then yellow-brown) in KOH, lacking reddish tints, inamyloid; in SEM the ornamentation consists of distinctive broad bands, arranged in a subreticulate to reticulate or more or less labyrinthiform pattern, the suprahilar area is faintly verrucose. Basidia (15)1826(30) x (5)67(8) µm, 4-spored, rarely 2-spored, clavate to subcylindric, thin-walled, hyaline, clamped. Pleurocystidia absent. Cheilocystidia 35138(147) x 37 (8) µm, numerous, cylindric-clavate to clavate, rarely narrowly utriform, occasionally septate, sinuous toward base, apex rounded or subcapitate, rarely attenuated, (4)510 µm wide, at times bifurcate or lobate, thin-walled, clamped; when observed in water with a conspicuous, reddish or vinaceous, cytoplasmic pigment becoming yellowish, in KOH the pigment is dissolved immediately and then hyaline or yellowish, without intraparietal or encrusting pigments, occasionally covered with resinous, hyaline material. Pileipellis a cutis or a transition between a cutis and a loose trichoderm, composed of cylindric, interwoven hyphae, 39(10) µm wide, thin-walled or moderaterly thick-walled (<1 µm thick), clamped, the layer is variable in depth and with a variable number of undifferentiated, rarely subcapitate, straight or slightly curved terminal elements, which are abundant and densely packed at pileus margin (resembling a prostrated turf ); near pileus trama the hyphae are more compactly arranged, often sinuous or curved, simple or bifurcate or with short lateral projections; in water most hyphae contain a more or less granulose pigment, which depending on age varies from pinkish to wine-red color, after 12 min the pigment is dissolved or bleached and in KOH almost instantaneously is lost, then the cytoplasmic content turns colorless or pale yellowish (similar to the pileus and hymenophoral trama). Pileus trama composed of cylindric to more or less ventricose hyphae 515(18) µm wide, thin-walled or slightly thick-walled (<1 µm thick), hyaline or pale yellowish, compactly interwoven. Hymenophoral trama subregular, but irregular toward the lamellar edge, hyphae (4)612(14) µm wide, cylindric to more or less ventricose, thin-walled or slightly thick-walled (<1 µm thick), hyaline or yellowish. Clamp connections abundant.
Habitat. Subgregarious, on bark (covered with mosses) of a dead trunk (probably of a hardwood tree), in cloud forest.
Specimens examined. MEXICO. VERACRUZ: km 2.5, old road Xalapa-Coatepec, Instituto de Ecología, Santuario del Bosque de Niebla, 1300 m alt., 22 Jan 2004, Bandala 3882; 30 Jan 2004, Bandala 3885; 18 Feb 2004, Bandala 3889 (HOLOTYPE, XAL; ISOTYPE, ZT); 10 Mar 2004, Bandala 3890; 18 Mar 2004, Montoya 4092, 25 March 2004, Montoya 4093; 22 Apr 2004, Montoya 4101 (all at XAL).
Commentary.
Both macroscopically and microscopically, Crepidotus rubrovinosus recalls C. subpur-pureus S. Ito & S. Imai (1940)
, originally described from Bonin Islands ( Japan) and subsequently gathered also in Papua New Guinea (Horak and Desjardin 2004
). The type material of this taxon (SAP, TMI 5210: Bonin Islands, Hahajima, Kitamura, Sekimonzan, 20 Nov 1936, S. Ito, S. Imai & K. Hino, s.n.), is in fragmentary condition and several taxonomically important microscopic characters were not mentioned in the protologue. A study of the small fragments revealed that the hyphae are clamped and, comparing the Japanese and the Mexican specimens, minor differences in basidiospore size were observed (viz. 5.57[7.5] x 45 [5.5] µm,
= 6.5 x 4.6 µm, Q = 1.42) for C. subpurpureus. However these two taxa can be distinguished easily in the pattern of the spore ornamentation, among other features mentioned below. In C. subpurpureus the ornamentation consists of coarse, isolated verrucae, occasionally interconnected by short and irregular ridges (FIGS. 2e
, 3ef
), then distinctively strongly verrucose when observed in both immersion oil and SEM. The basidiospores of C. rubrovinosus are characterized by broad and sinuous, blunt bulges and ridges, interconnected forming a subreticulate pattern (FIGS. 1b
, 2b
, 3ad
).
As pointed out by Horak and Desjardin (2004)
, the Japanese C. subpurpureus also occurs in Papua New Guinea. Based upon the features observed on the Papuan specimens, these authors presented a complete description and proposed its transfer from Crepidotus to Pyrrhoglossum (P. subpurpureum [S. Ito & S. Imai] E. Horak & Desjardin). As mentioned above the Mexican C. rubrovinosus and the Japanese-Papuan P. subpurpureum share important characters. The most noteworthy features in common are the colors (vivid red, wine-red or deep lilac to purple) rarely observed in other representatives of Crepidotus and Pyrrhoglossum, the colored gill edges, the conspicuously ornamented basidiospores and with a plage-like, supra-apicular spot, the plasmatic pigment in both the pileipellis hyphae and cheilocystidia (rapidly dissolving in KOH in C. rubrovinosus), and finally the presence of clamps on the septa.
Crepidotus rubrovinosus and Pyrrhoglossum subpurpureum appear to be morphotaxonomically intermediate between two established crepidotoid genera, Crepidotus and Pyrrhoglossum. However these two taxa can be separated by the general structure of the pileipellis, the different color of both the hymenophore and spore print, as well as the basidiospore ornamentation (and perhaps size). Crepidotus rubrovinosus is characterized by yellow or yellow-brown basidiospores correlated with a yellow-brown spore print and yellowish to pale brown lamellae, contrary to Pyrrhoglossum subpurpureus which has a rust brown spore print and basidiospores of the same color correlated with the pale gray and gradually pinkish lamellae. Furthermore the two taxa are separated by the color of the pileus context (gray for P. subpurpureus; white to yellowish for C. rubrovinosus) and the subtle but distinctive differences of the plasmatic pigments localized both in the pileipellis hyphae and in the cheilocystidia (i.e. the cells filled with wine-red to purple plasmatic pigments persistent in KOH in P. subpurpureus). Based on results of pending DNA analysis the present taxonomic position of both Crepidotus rubrovinosus and Pyrrhoglossum subpurpureum could be re-evaluated later. The authors for the time being prefer to describe the new Mexican species as a member of Crepidotus.
On account of its ovoid basidiospores bearing a rugose-verrucose pattern of ornamentation and the presence of clamp connections, C. rubrovinosus probably is related to the group of species around C. subverrucisporus Pilát in sect. Crepidotellae Hesler & A.H. Sm. (ss. Senn-Irlet 1995
). In comparison to the species close to this group (cf. C. icterinus Singer, C. lundelli Pilát, C. subverrucisporus, C. xanthocephalus Singer, C. rubriceps Singer, among others), the coarse ornamentation of the basidiospores of C. rubrovinosus is remarkable even when seen under immersion oil. The color of both pileus and lamellar edges, together with pileal structure, in combination with the shape and ornamentation of the basidiospores, and the distinctive cheilocystidia, are a set of features that taxonomically distinguish C. rubrovinosus from related taxa. Aged or overmature basidiomes of C. rubrovinosus macroscopically could recall C. cinnabarinus Peck or C. rubriceps which also possess predominantly red basidiomes. Microscopic analysis demonstrates, however, that they are distinctly separated: C. cinnabarinus has clampless hyphae, more or less lageniform cheilocystidia, and broadly ellipsoid basidiospores (Sect. Sphaerula Hesler & A.H. Sm., ss. Senn-Irlet & De Meijer 1998
). Crepidotus rubriceps differs in its ellipsoid to subamygdaliform, verrucose basidiospores (with more or less broad, isolated warts), 67.5 x 45 µm, the claviform cheilocystidia and the pileipellis hyphae with distinctly swollen, pigmented terminal cells (after type studies and data provided by Luther and Redhead 1981
, Singer 1973
, Senn-Irlet and De Meijer 1998
).
Crepidotus septicoides (Singer) Singer, Beih. Nova Hedwigia 44:399. 1973. FIGS. 2f
, 4b
, 5ad
Basionym: Marasmiellus septicoides Singer. Mycologia 47:773. 1955.
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Pileus 0.517 mm broad, subglobose to hemispheric in primordial stages, occasionally cupuloid, becoming ungulate, convex or plano-convex, rarely plane, at times slightly umbonate near point of attachment, circular, subcircular or rounded flabelliform, commonly with a slot in the rear portion (when seen from the hymenophore), this at times produces two lobe-like hemispheres; white to almost pure white, later pale yellowish on disk, dry, weakly hygrophanous, densely fibrillose to somewhat villose, becoming woolly to tomentose-fibrillose or tomentose, pubescent-woolly toward point of attachment; margin incurved, later straight, smooth or weakly sulcate, at times (mainly when fully expanded) wavy or more or less crenate, occasionally translucent striate; dorsally or laterally attached to substrate. Lamellae free to adnexed, rarely narrowly adnate, in several stages of development leaving a minute smooth area around of the rudimentary stem or knob, concurrent to an eccentric or lateral point, subclose, subdistant or distant, broad to medium broad (
1.5 mm broad), subventricose, lamelullae of 23(4) different lengths; white, becoming brownish-yellow (10 yr 8/ 34) or darker (10 yr 7/4), at times with pinkish shades, white edges fimbriate. Stipe in primodial or young stages central, rudimentary (up to 1 mm long.), with age persisting as an eccentric or lateral, cylindric knob (when seen from the hymenophore) but the pileus is directly attached to the substratum; the rudimentary stem white, pruinose and finaly as a glabrous, withish knob; basal mycelium white, present or absent. Context white to whitish, thin (<1 mm thick), hygrophanous, soft, unchanging on exposure. Odor and taste not distinctive.
Spore print (seen on pileus surface) ochraceous to pale brownish.
Basidiospores (6.5)710(11) x 4.56.5(7) µm,
= 7.38.9 x 5.15.5 µm; Q = 1.411.64, ellipsoid to more or less elongate, often with weak suprahilar depression and slightly attenuated at apex, in lateral view more or less amygdaliform or rarely broadly subfusiform (apex not beaked), weakly rugulose, pale yellowish to yellowish in KOH, inamyloid, thin-walled. Basidia 2035 x 79(10) µm, 4-spored, rarely 2-spored, clavate or subcylindric, thin-walled, hyaline, clamped. Pleurocystidia absent. Cheilocystidia 3580 x (3)47(8) µm, numerous, subcylindric or more or less narrowly sublageniform, flexuous, commonly with constrictions, apex rounded or sligthly tapering and rounded, occasionally with irregular or bifurcate outgrowths, hyaline, thin-walled, clamped. Pileipellis a loose cutis or a loose trichoderm, composed of cylindric, interwoven, hyphae (2)37(10) µm wide, thin-walled or moderately thick-walled (<0.5 µm), hyaline to pale yellowish, nongelatinized, with a variable number of terminal elements which are undifferentiated, rarely narrowly lageniform, sinuous or curved, occasionally coiled. Pileus trama composed of cylindric to more or less ventricose hyphae 612 (15) µm wide, thin-walled, hyaline, nongelatinized, compactly arranged. Hymenophoral trama subregular to subirregular, hyphae 410 µm wide, cylindric to more or less ventricose, hyaline, thin-walled. Clamp connections abundant.
Habitat. Densely gregarious or scattered on dead leaves of Platanus mexicana Moric, at times on the rachis or debris of dead ferns, more rarely on dead mosses and debris covering rocks. Found at margin of a cloud forest.
Specimens examined. MEXICO. VERACRUZ: km 2.5, old road Xalapa-Coatepec, Instituto de Ecología, Santuario del Bosque de Niebla, 1300 m alt., 24 Jun 2002 Bandala 3592, 3593, 3594, 3595; 6 Jun 2003, Bandala 3736; 10 Jun 2003, Bandala 3740 ; 22 Jun 2003, Montoya 3995 (all at XAL).
Commentary.
As proposed in a previous paper (Bandala et al 1999
) the specimens reported by Singer (1973)
as "C. longicystis" (gathered in a tropical forest south of Veracruz) are conspecific with C. septicoides. The present collections indicate a broad range with regard to the macroscopic variation of the basidiomes of this species. The size of the basidiospores in the type material of Marasmiellus septicoides from Brazil (1012 [12.5] µm length) differs only slightly from the Mexican specimens of this species we have recognized (Bandala et al 1999
). The remarkably wide variation of basidiospore shape in single collections of C. septicoides (Singer 1973
, Bandala et al 1999
) again was confirmed in the specimens from Xalapa.
Crepidotus septicoides is a striking species distinguished by its small, white basidiomes, and especially by having pale, very weakly rugulose, ellipsoid basidiospores, long, flexuous, subcylindric cheilocystidia, and clamped hyphae. Microscopic characters (basidiospores, cheilocystidia) shown by the type collection of Crepidotus fusisporus var. longicystis Hesler & A.H. Sm. (1965) indicate that this taxon is clearly different from C. septicoides (i.e. C. longicystis s. str. Singer) and taxonomically different indeed from the group of taxa around C. fusisporus Hesler & A.H. Sm. (cf. the type study of var. longicystis presented below).
Near the locality where C. septicoides was collected in Xalapa, Bandala et al (1999)
recorded C. epibryus (Fr. : Fr.) Quél., growing also on dead leaves of Platanus. In the field the basidiomes of this taxon do recall those of C. septicoides, but microscopically C. epibryus is readily separated due to its smooth, slender (cylindric) basidiospores, more or less whip-like cheilocystidia and clampless hyphae.
Crepidotus fusisporus var. longicystis Hesler & A.H. Sm., North Amer. Sp. Crepidotus p. 93. 1965.
FIGS. 5ef
, 6a, b
HOLOTYPE. U.S.A. MICHIGAN: Cheboygan Co., 7 Jul 1949, A.H. Smith 32464 (MICH).
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= 8.6 x 4.7 µm; Q = 1.84, ellipsoid, often apically attenuated, then more or less amygdaliform or even subfusoid (apex not beaked), when seen under immersion oil roughened-punctate to faintly rugulose, yellowish to pale yellowish, thin-walled, inamyloid; under SEM the pattern of ornamentation consists of low bulges often interconnected, producing a rugulose or rugulose-verruculose ornamentation. Basidia 2530(35) x 57 µm, 4-spored, occasionally 2-spored, clavate, thin-walled, hyaline, clamped. Pleurocystidia absent. Cheilocystidia 2155 x 38 µm, numerous, subcylindric-clavate or more or less narrowly utriform, rarely clavate, apex rounded, 510(12) µm wide, commonly sinuous, hyaline, thin-walled. Pileipellis a loosely interwoven cutis of hyphae 36(8) µm wide, cylindric, hyaline or yellowish in some segments, nongelatinized, thin-walled, clamped, terminal hyphae undifferentiated and variable in number, in some parts more or less semierect, then the layer appears in transition between a cutis and a trichoderm. Pileus trama with cylindric to more or less ventricose hyphae 310 µm wide, thin-walled, hyaline, more or less compactly arranged. Hymenophoral trama subregular to subirregular, hyphae 38 µm wide, cylindric to more or less ventricose, hyaline, thin-walled. Clamp connections abundant. Commentary. The specimen supporting the var. longicystis is not conspecific with C. fusisporus and in our opinion, the collection of A.H. Smith 32464 fits the concept of C. luteolus (Lambotte) Sacc. These observations lead us to consider that this variety should be reduced to a synonym:
The characteristics described above for the type collection of C. fusisporus var. longicystis (especially the basidiospores and cheilocystidia) indicate that this taxon has no taxonomic relationship with C. fusisporus (var. fusisporus and allied varieties). Although some of the basidiospores of the specimen of A.H. Smith 32464 are apically tapered they are not comparable in shape with the remarkable fusoid basidiospores (with extremes almost beaked) of C. fusisporus. The basidiospores of C. fusisporus var. fusisporus (and allied varieties) also differ by having an ornamentation consisting of scattered, low, isolated, conical or hemispherical verrucae (under SEM), which are distinctly punctate when observed under immersion oil (Bandala and Montoya 2000a
, Senn-Irlet 1991
). This type of ornamentation differs sharply from the rugulose or rugulose-verruculose type of ornamentation of C. fusisporus var. longicystis. The more conspicuous suprahilar depression and the pale yellowish to almost hyaline yellowish color of the spore wall, as well as the variably shaped cheilocystidia (e.g. narrowly lageniform, contorted or sinuous) are also distinctive for C. fusisporus. It should be pointed out that an additional sample kept at MICH under C. fusisporus var. fusisporus (Michigan: Luce Co., Upper Falls, Tahquamenon St. Park, 28 Jun 1956, Thiers 2849) was found to be identical with C. fusisporus var. longicystis (FIG. 6c, d
). These collections (32464 and 2849) were included in the group of C. fusisporus by Hesler and Smith (1965)
, probably due to the shape of the basidiospores.
The basidiospore shape and ornamentation of the C. fusisporus var. longicystis type collection relate it to members of Sect. Crepidotellae (Senn-Irlet 1995
) which possess ellipsoid or amygdaliform basidiospores bearing a rugulose to rugulose-verruculose ornamentation pattern (cf. FIG. 6
). Macro- and microscopic features suggest that the North American type collection of this variety seems to represent extreme variation of C. luteolus, a member of that section. Except by the whitish pileus ("light buff when dried") mentioned in the protologue (Hesler and Smith 1965
), the material of A.H. Smith 32464 fits that concept well. Crepidotus luteolus indeed embraces specimens with yellow to whitish basidiomes (Breitenbach and Kränzlin 2000
, Lonati 2000
, Malençon and Bertault 1975
). The pileipellis structure and the basidiospores match the neotype of C. luteolus (Switzerland. Dorfwald ob Schwarzenburg, Kt. Bern, 12 Oct 1989, Senn-Irlet 89/243, G) (FIG. 6e, f
) although the cheilocystidia in the North American collection are less polymorphic (i.e. ± long-cylindric, narrowly sublageniform, often sinuous and mostly with outgrowths or apically branched in the neotype). A similar type of spore ornamentation as that exhibited by the spores of C. luteolus is found in C. lundellii. This species differs by its white basidiomes, clavate, branched-clavate or subutriform cheilocystidia, and although it is reported with a wide range of variation of shape and size of basidiospores (Senn-Irlet 1993
, 1995
; Nordstein 1990
) these latter are consistently oblong to ovoid (with rounded extremes) and less frequently amygdaliform as observed also in the lectotype (Fungi Exsiccati Suecici 220, 15 Sep 1935, leg. Lundell & Åberg, kept in K) (FIG. 6g, h
).
On account of the aforementioned features we concluded that C. fusisporus var. longicystis should be considered a later synonym of C. luteolus. Therefore this synonymy is proposed:
Crepidotus luteolus (Lambotte) Sacc., Sylloge Fungorum 5:888. 1887.
Basionym: Agaricus luteolus Lambotte, Fl. Mycol. Belg. 1:181. 1880.
Synonym: Crepidotus fusisporus var. longicystis Hesler & A.H. Sm., North Amer. Sp. Crepidotus p 93. 1965.
| ACKNOWLEDGMENTS |
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| FOOTNOTES |
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1 Corresponding author. E-mail: bandala{at}ecologia.edu.mx
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, , Moreno G. 1999. Two Crepidotus from Mexico with notes on selected type collections. Mycotaxon 72:403416.
Breitenbach J, Kränzlin F. 2000. Fungi of Switzerland. 5. Agarics part 3. Cortinariaceae. Lucerne: Verlag Mykologia. 338 p.
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Holmgren PK, Holmgren NH, Barnett LC, eds. 1990. Index herbariorum. I. The herbaria of the world. 8th ed. New York: New York Botanical Garden. 693 p.
Horak E, Desjardin DN. 2004. Two remarkable taxa of Crepidotus from Bonin Islands (Japan) and new records from the Hawaiian Islands and Papua New Guinea. In: Cripps CL, ed. Fungi in forest ecosystems, diversity and ecology. New York: The New York Botanical Garden. p 6170.
Ito S, Imai S. 1940. Fungi of the Bonin Island, IV. Trans Sapporo Nat Hist Soc 16:4556.
Kornerup A, Wanscher JH. 1967. Methuen handbook of colour. 2nd ed. London: Methuen & Co. Ltd. 243 p.
Krisai-Greilhuber I, Senn-Irlet B, Voglmayr H. 2002. Notes on Crepidotus from Mexico and the southeastern USA. Persoonia 17:515539.
Lonati G. 2000. Il genere Crepidotus in area mediterranea. Mic Veg Med 15:161186.
Luther BS, Redhead SA. 1981. Crepidotus cinnabarinus in North America. Mycotaxon 12:417430.
Malençon G, Bertault R. 1975. Flore des champignons superieurs du Maroc, II. Rabat: Institut Scientifique Cherifien et Faculte des Sciences de Rabat. 536 p.
Munsell soil color charts. 1994. New York: Macbeth. 4 p. + 9 color charts.
Nordstein S. 1990. The genus Crepidotus (Basidiomycotina, Agaricales) in Norway. Sinopsis Fungorum 2:1115.
Senn-Irlet B. 1991. Crepidotus, Pellidiscus and Ramicola in Greenland (Agaricales: Crepidotaceae). Nord J Bot 11:587597.[CrossRef]
. 1993. Type studies in Crepidotus II. Persoonia 15:155167.
. 1995. The genus Crepidotus (Fr.) Staude in Europe. Persoonia 16:180.
, De Meijer A. 1998. The genus Crepidotus from the state of Paraná, Brazil. Mycotaxon 66:165199.
Singer R. 1973. The genera Marasmiellus, Crepidotus and Simocybe in the neotropics. Beih. Nova Hedwigia 44:1484.
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