Taconomy of Crustoderma (Aphyllophorales, Corticiaceae)
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Mycologia, 76(1), 1984, pp. 40-50. © 1984, by The New York Botanical Garden, Bronx, NY 10458 TAXONOMY OF CRUSTODERMA (APHYLLOPHORALES, CORTICIACEAE) K. K. NAKASONE Center for Forest Mycology Research. Forest Products Laboratory. 1 Forest Service, U.S. Department of Agriculture, P.O. Box 5130, Madison, Wisconsin 53705 ABSTRACT The genus Crustoderma is emended to include nine species. and all, excepty C. longi cystidium, are associated with brown-rot decay of wood. Two new species. C . carolinense and C. marianum, are described and illustrated. New combinations of C. corneum, C. Ion gicystidium, and C. patricium are proposed. Peniophora weiri Bres. is placed in synonymy with C. dryinum. Culture descriptions are included for C. marianum and C. corneum. A key to the species is provided. Key Words: culture descriptions. brown-rot fungi, Corticiaceae, Crustoderma. Crustoderma was proposed by Parmasto (1968) for Corticium dryinum Berk. et Curt. Gilbertson (1981) transferred Peniophora resinosa Jacks. et Deard. to Crustoderma. and C. flavescens Nakas. et Gilbn. and C. opuntiae Nakas. et Gilbn. have been included in the genus (Nakasone and Gilbertson. 1982). Crustoderma is emended to include information on cultural characters, associated brown-rot decay. and the slightly thick-walled basidiospores that are important characters of the genus. A key to the known species is included. Because Crustoderma possesses some characters found in other genera. several species have been placed in Hyphoderma Wallr. and Phlebia Fr. All the species in Crustoderma and many species of Hyphoderma are characterized by well- developed cystidia and large basidiospores. The ceraceous or corneous basidio carps and the dense. thickening subhymenium are characters found in both Crus toderma and Phlebia. A survey indicated that Phlebia longicystidia (Litsch.) Hjortst. et Ryv. and Phlebia cornea (Bourd. et Galz.) Parm. were species of Crustoderma. Cultural and basidiocarp studies, meanwhile, have uncovered two new species. Methods and materials are described in Nakasone and Gilbertson (1982). Sometimes a drop of 10% KOH was used to separate structures in tightly agglu tinated specimens. Herbarium abbreviations are from Holmgren et al. (1981). Changes and additions to the generic characterization are italicized. In the species code, 27 refers to the presence of evenly thick-walled hyphae (Burdsall et al., 1978). Asterisks (*) indicate cultures studied. CRUSTODERMA Parmasto emend. Nomenclatural type: Corticium dryinum Berk. et Curt. Basidiocarps resupinate, effused. ceraceous to crustaceous or corneous when dried, closely adherent to substrate; margins abrupt, narrow or thinning out; fertile areas smooth or with scattered warts. often velutinous because ofnumerous emer gent cystidia, tints of yellow, orange or gray, sometimes olive-colored. Subicular hyphae up to 7 µm diam, thin- to thick-walled, some gelatinized, hyaline or pale yellow, nodose septate, frequently branched. Subhymenium thickening. consisting of mostly vertical. densely agglutinated hyphae. Cystidia cylindrical to clavate, 1 Maintained at Madison, Wisconsin. in cooperation with the University of Wisconsin, 40 NAKASONE: CRUSTODERMA 41 up to 380 µm long, obtuse. often thick-walled in basal part and thinning toward apex or thin-walled throughout. sometimes with adventitious septa, with or with out a crystalline sheath. Basidia narrowly clavate to cylindric, up to 80 µm long, elongated at the base (pedunculate), sometimes thin- or slightly thick-walled, hya line or yellow, clamped at basal septum, 4-sterigmate. Basidiospores subglobose, ellipsoidal or cylindrical, with slightly thickened walls, hyaline or yellow, nonam yloid, acyanophilous but the contents strongly stained in cotton blue. In pure culture on MEA mats white or yellow, appressed, sometimes raised, growth slow to rapid; negative reactions on GAA and TAA; often with a strong, pungent odor. Advancing zone hyphae simple septate or with clamp connections; submerged and surface hyphae nodose septate, thin- walled but often becoming irregularly thick- walled. Associated with a brown-rot decay of dead conifer and/or hardwood logs and branches. Remarks. -BecauseCrustoderma has several characters in common with other genera, it can be difficult to recognize. The critical characters of the genus are the velutinous hymenium. the dense. thickening subhymenium, the large cystidia, the large. slightly thick-walled spores. and association with a brown-rot decay. Chae toderma Parm. (1968), another genus of brown-rot decay fungi, is similar to Crustoderma but typically has longer basidia (up to 100 µm long) and arcuate, fusiform spores. If the associated decay is not known, some species of Crustoderma may appear to belong in Hyphoderma or Phlebia. However, the slightly thick- walled basidiospores can be used to identify a species of Crustoderma. Addition ally, in pure culture Crustoderma species do not react on gallic acid or tannic acid agar while Hyphoderma and Phlebia species react strongly. DESCRIPTIONS OF NEW SPEClES Crustoderma carolinense Nakas., sp. nov. FIG. 1 Differt a Crustodermatis cornei hyphidiis 30-45 × 23 µm: basidiosporis 8-9 × (5-)6-7µm, late ellipsoideis vel subglobosis. hyalinis, laevis, leviter crassitunicis. Holotypus: HHB 2479. ad lignum Castaneum sp., Shope Fork, Coweeta Experimental Forest. Macon County, North Carolina. U.S.A., 28 July 1969, legit H. H. Burdsall, Jr., in herb. CFMR. ETYMOLOGY: From carolinensis (L., adj.), referring to the state in which the holotype was found. Basidiocarp annual. effused up to 100 × 40 mm, up to 500 µm thick, texture when fresh not known. when dried corneous. adherent except for cut edges which curl up and loosen from substrate: fertile area continuous. smooth with scattered warts, projecting cystidia. easily seen under a 30× lens, numerous and well-spaced, in fresh specimens Pale Smoke Gray with Pale Olive-Buff to Olive-Buff spots, thicker and well-developed areas Olive-Gray. thinner and younger areas Avel laneous, when dried projecting cystidia giving a glaucous cast over fertile areas. some fertile areas almost invisible. very thin. hyaline, smooth and noticeable only because of the projecting cystidia: margins abrupt. fimbriate or punctiform. Light Ochraceous-Buff to pale Cinnamon-Buff. Hyphal system monomitic. Subiculum up to 120 µm thick. composed of tightly agglutinated hyphae arranged parallel to substrate: subicular hyphae (3-)5-7 µm diam, hyaline. smooth. thin-walled. nodose septate. infrequently branched. Sub hymenium thickening, up to 200 µm thick. composed of compact. agglutinated. vertical hyphae: subhymenial hyphae 2.5-5 µm diam. hyaline. smooth. thin- or thick-walled, nodose septate. moderately branched. agglutinated but easily sep arated in 10% KOH. Hyphidia cylindrical. often branched at apex. 30-45 × 2-4 42 MYCOLOGIA FIG. 1. Crustoderma carolinense (HHB 2479, holotype). a. Cystidia (note scale). b. Subhymenial hyphae. c. Basidia. d. Hyphidia. e. Basidiospores. FIG. 2. Crustoderma marianum (FP 105991, ho lotype). a. Cystidia. b. Subicular hyphae. c. Basidia. d. Basidiospores. µm, hyaline, thin-walled, clamped at basal septum, smooth, arising from hy menium. Cystidia cylindrical to clavate, 120-380 × 14-22 µm, hyaline, protrud ing up to 230 µm, walls slightly thickened or thick-walled at base then gradually thinning toward apex, clamped at basal septum, smooth or with scattered crys talline matter, arising from subhymenium or hymenium. Basidia clavate, some- NAKASONE: CRUSTODERMA 43 times stalked, 52-68 × 8-9 µm, hyaline, smooth, thin-walled, clamped at basal septum, 4-sterigmate, sterigmata up to 6 µm long. Basidiospores broadly ellipsoid to subglobose, 8-9 × (5-)6-6.5(-7)µm, hyaline, walls slightly thickened, smooth, nonamyloid, nondextrinoid, acyanophilous. Remarks. - Crustoderma carolinense, associated with a brown-rot of dead chest nut, is known only from the type locality. Its drab, dark color and horny texture are useful characters to differentiate it from most other species in the genus. It is closely related to C. corneum, which also has a horny texture. However, the broadly ellipsoid to subglobose spores, hyphidia, and very large cystidia of C. carolinense can be used to distinguish it from C. corneum. The latter species appears to grow exclusively on Pinus spp. Crustoderma marianum Nakas., sp. nov. FIG. 2 Differt a speciebus Crustodermatum basidiis 30-35 × 5.5-7.5 µm; basidiosporis (9-)10-12 (-13) × 5.5-6.5(-7)µm, ellipsoideis vel late cylindraceis, hyalinis, laevis, leviter crassitunicis. Holo typus: FP 105991, ad lignum Pinus sp., Telegraph Road vicinia, Beltsville Forest Disease Laboratory, Laurel, Maryland, USA., 28 Jan. 1966, legit 0. K. Miller. Jr., et R. L. Gilbertson, in herb. CFMR. ETYMOLOGY: From marianus (L., adj.), referring to Maryland, the state in which the holotype was found. Basidiocarp annual. effused up to 100 × 50 cm, up to 300 µm thick, texture when fresh not known. when dried crustaceous, adherent; fertile area Pale Ochra ceous-Buff to Light Ochraceous-Buff. continuous. smooth, thinner areas floccose. projecting cystidia easily visible at 30×; margins gradually thinning out and prui nose or abrupt, concolorous with fertile areas; subiculum white and cottony when exposed. Hyphal system monomitic. Subiculum up to 100 µm thick, composed oftightly woven hyphae: subicular hyphae 4-5 µm diam, hyaline, smooth, slightly thick- to thick-walled, nodose septate, moderately branched. easily separated in 10% KOH. Subhymenium thickening,