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Biodiversidad y Sistemática, Instituto de Ecología, 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
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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The study of Crepidotus specimens collected 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, taxonomy, tropical fungi, wood-inhabiting fungi
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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consists of an assemblage of up to eight members of Crepidotus especially distinguished among the baculate, 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 consistency 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 morphological variation observed in the samples examined, 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.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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, 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 |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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Syn.: Crepidotus applanatus var. crocophyllus (Berk.) Pilát Atl Champ Eur 6:35. 1948.
C. fulvifibrillosus Murrill. N Am Fl. 10:153. 1917.
C. applanatus var. fulvifibrillosus (Murrill) Pilát 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 Albertó (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 circular, 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 incurved 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), yellowish-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 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 distinctive.
Basidiospores (4.5–)5–7.5(–8) x 4.5–7(–8) µm, x = 5.5–6.9 x 5.5–6.8 µm, Q = 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 µm). Basidia 15–45(–48) x 5–9(–10) µm, clavate to narrowly clavate, 4-spored, hyaline, clamped. Pleur-ocystidia absent; among the hymenial elements some collections occasionally present sterile bodies resembling amorphous or abortive basidia or anomalous basidioles appearing as cystidioid-like structures (cf. FIGS. 2e
, 3b, g, 5g). Cheilocystidia (20–)24–75 x 5–10(–12) µm, 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) µm wide, rounded, subcapitate, rarely tapered or branched, hyaline thin-walled, clamped. Pileipellis a cutis consisting of cylindric to somewhat ventricose, septate, pale yellowish, yellowish, pale yellow or pale yellowish-brown hyphae (yellowish-brown to brownish-orange in group) 4–15(–20) µm wide, forming the pigmented fibrils or scales of pileus surface, thick-walled, 0.5–1.0(–1.5) µm, 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 µm wide. Hymenophoral trama subregular to subirregular, hyphae 3–10(–18) µm 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: Área de Conservación La Amistad Pacífico, Z.P. Las Tablas, Cotoncito, 17 Apr 1999, Navarro 962; Finca Cafrosa, Fila Chiquizá, 9 Jul 1999, López 556; Sendero Central, 24 Jun 1999, Oses 346; Sendero Progreso, 30 Sep 2000, Mata 898. GUANACASTE: Área de Conservación Arenal, P.N. Volcán Tenorio, Hacienda Montezuma, 5 May 2000, López 1296-2. ALAJUELA: Área de Conservación Arenal, P.N. Volcán Tenorio, Alto Los Brenes, 19 May 1999, López 370 (all at INB). MEXICO. VERACRUZ: Freeway Fortín-Orizaba, Barranca de Metlác, 26 Aug 1989, Bandala 1854; Km 2.5 old road Xalapa-Coatepec, wooded area of Jardín Botánico Fco. J. Clavijero, 30 Jan 1996, Bandala 2919; 10 Jan 1997, Muñoz 10; 25 Jun 2003, Bandala 3762; Sta. Inés Ranch, 23 May 1991, Tapia 545; Instituto de Ecología, 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; 27 June 2005, Bandala 3963; Los Tuxtlas Region, near Catemaco, 4 Aug 1997, Muñoz 30 (all at XAL).
Other specimens examined: – AUSTRIA. OBERÖ 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 cheilocystidia, and a pileipellis formed of repent, yellow to brown-pigmented, thick-walled, often incrusted hyphae. 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, Lazébni
ek 1970, Moser 1983, Murrill 1917, Pegler 1977 as C. nephrodes; Pereira 1990, Pilát 1950, Phillips 1991, Raithelhuber 1988, Ripková et al 2005, Senn-Irlet 1995, Singer 1973, Wright and Albertó 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. FIG. 1 and Moser and Jülich 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 consequently 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 taxonomically 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 differentiated 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 basidospore 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. subaureifolius) 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) (FIG. 4) also showed such variation. This characteristic, apparently common among Crepidotus species producing 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 pigmented, thick-walled, often incrusted hyphae, cheilocystidia 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 recognized by Hesler and Smith (op. cit.) are not consistent, we concluded therefore that the specimens represent different stages of the macroscopic variation 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 bodies, which seem to be abnormal basidia or basidioles, certainly were encountered at times in the hymenium (FIGS. 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 distinguish taxonomically discrete entities. Singer (1973) and Senn-Irlet and De Meijer (1998: 180) also have expressed their doubts about the taxonomic relevance of this kind of structures called pleurocystidia. Crepidotus is still conceived as a lineage of members lacking pleurocystidia (Aime et al 2005, Nordstein 1990, Pilát 1948, Senn-Irlet 1995, Singer 1973, 1986, Watling and Gregory 1989).
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). Our results additionally provide useful information for taxonomic resolution of the Fulvifibrillosi clade obtained by Ripková 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 characters mentioned in the diagnosis and data afterward provided by Imazeki and Hongo (1987) and Imazeki et al (1988), C. badiofloccosus appears to be indistinguishable 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 Hampshire 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 Albertó 2002).
Crepidotus malachius (Berk. & M.A. Curtis) Sacc., Syll Fung 5:883. 1887. FIG. 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) x (4.5–)5.5–7 (–8) µm, x = 5.5–6.5 x 5.5–6.3 µm, Q = 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 µm thick). Basidia clavate, 4-spored, hyaline, clamped. Pleurocystidia absent. Cheilocystidia although 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 µm diam, terminal elements scarce, undifferentiated. Pileus trama with cylindric to subventricose, thin-walled, hyaline, hyphae. Hymenophoral trama subregular, 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, Pilát 1950, Singer 1947, 1973) who have offered different taxonomic interpretations. Then prevailing concepts of Crepidotus crocophyllus and C. fulvifibrillosus in connection with C. applanatus (cf. the discussions by Hesler and Smith [1965: 79], Pilát [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 Crepidotus applanatus (Horak 1964, 1979, Murrill 1917, Pilát 1950, Raithelhuber 1988, Singer 1947, 1949, 1953, 1973, 1986, Singer and Digilio 1951, Wright and Albertó 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 (Ripková 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 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] basidiospores [FIG. 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 described as devoid of any pigmented covering on pileus surface, and when describing Agaricus malachius Berkeley and Curtis (1859) emphasized its resemblance to A. nephrodes. In the original description by Berkeley and Curtis (1853) Agaricus nephrodes is characterized by its pileus "...pallide flavido-tomentoso...", 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...". Pilát (1948) and Horak (1979) in part suggested the conspecificity between C. nephrodes and C. applanatus. Pilát (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, Ripková 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 Pilát (1948) placed C. fulvifibrillosus 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 Albertó (2002) also treated C. crocophyllus and C. nephrodes as different species. Hesler and Smith (1965) accepted separately C. applanatus, C. crocophyllus, C. malachius and C. nephrodes, using the latter to include Murrill’s C. fulvifibrillosus as a synonym. Therefore the data reveal that one concept of C. nephrodes, in apparent coincidence with the original description, encompasses members devoid 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, Ripková 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) (FIG. 4a–c). Our results strongly support the observations of Hesler and Smith (1965) and part of the arguments of Ripková et al (2005) based on samples from Europe and North America but differ from Pilát (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 basidiospores in the M.A. Curtis 1912 specimen are (5–)6–7(–7.5) x 5.5–7(–7.5) µm, x = 6.6 x 6.5 µm, Q = 1.02 (FIG. 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 morphological 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-examination 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. Führ 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 Ripková 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).
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: victor.bandala{at}inecol.edu.mx
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LITERATURE CITED |
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