Vol. XIX, pp. 389-395 January-March 1984

THE (CORTICIACEAE) IN NORTH AMERICA

AND A NOTE ON MEXICANA

By

HAROLD H. BURDSALL, JR.

1/ Center for Forest Mycology Research, Forest Products Laboratory— USDA, Forest Service, Madison, WI 53705, USA

SUMMARY

Studies of wood-rotting Basidiomycetes for the purpose of clarifying taxonomic limits and biological similarities or differences among the taxa led me to investigate the genus Karst., a genus containing numerous white-rot species. During that study, I noted the similarity of Ph. septocystidia (Burt) Eriksson and Ryvarden, Ph. insolita Burds. and Nakas., and Peniophora mexicana Burt to the genus Candelabrochaete Boidin. I then examined the genus Candelabrochaete to establish generic differences from Phanerochaete and to determine Candelabrochaete's specific components. Such know- lege will enable more reliable determination of fungi in these genera, which are used in the study of forest pathology, wood products decay, and wood biodeterioration or bioalteration. As a result of this study, a key to and descriptions of the North American species of Candelabrochaete and a discussion of Pe.mexicana are presented here. Methods followed are those cited in Burdsall and Nakasone (1981).

INTRODUCTION

Candelabrochaete was proposed for the new species, C. africana Boidin, the type species, and Hypochnus langloisii Pat. Both occur on hardwoods in south temperate to subtropical habitats. The genus is similar to Phanerochaete in possessing simple septa in the subicular hyphae and at the base of the basidia, and hyaline, thin- walled, nonamyloid basidiospores. The generic characters of Candelabrochaete differ from Phanerochaete, however, by including small, cylindrical to clavate basidia, septate cystidia, a loosely interwoven subiculum, and a loosely organized hymenium that gives a farinaceous to woolly appearance to the basidiocarps. Eriksson et al. (1981) stated that the palisade arrangement of the hymenium, the clavate basidia, and the lack of percurrent proli- feration of basidia in Ph. septocystidia were more similar to species of Phanerochaete than to those of Candelabrochaete. My studies of several specimens of Ph. septocystidia and the holo- type of C. africana confirm most of the information reported by

1/ Maintained at Madison, Wis., in cooperation with the University — of Wisconsin. 390

Eriksson et al. Although the hymenium is sometimes organized into a palisade in Ph. septocystidia, the basic hymenial structure is similar to that seen in Candelabrochaete. In some areas of some specimens of Ph. septocystidia, the palisade is much less organized than in others. The basidium shape in Ph. septocystidia, however, is frequently some- what cylindrical to constricted, similar to the shape found in C. africana, although in some cases it is clavate. Eriksson et al. (1981) considered percurrent proliferation as a character of generic importance in Candelabrochaete. I did not observe percurrent proliferation in any specimens of Ph. septocystidia, but neither did I observe it in the type specimen of C. africana. In my experience, percurrent proliferation occurs erratically in many species. In Peniophora fuscomarginata Burt, for example, some specimens and parts of other specimens possess striking examples of percurrent proliferation. In other specimens it has been impossible to demonstrate percurrent proliferation. Therefore, the character is certainly not of use as a generic character in Peniophora. I consider it an equally suspect character in the case of Candelabrochaete.

DESCRIPTION OF SPECIES

Three North American species of Candelabrochaete are recognized in this study. A key to the species is provided and Peniophora mexicana, a species with similarities to these three, is included in the key and described.

Key to species of Candelabrochaete and Peniophora mexicana

1. Basidiospores thick walled, broadly ellipsoid to subglobose...... P. mexicana 1. Basidiospores thin walled, ellipsoid to allantoid ...... 2 2. Basidiospores ellipsoid ...... C. magnahypha 2. Basidiospores allantoid to broadly allantoid...... 3 3. Basidiospores broadly allantoid to allantoid, 7-9.5 x 3-4 µm ...... C. langloisii 3. Basidiospores allantoid 4.5-6.5 x 1.5-2 µm . . . . C. septocystidia

CANDELABROCHAETE LANGLOISII (Pat.) Boidin, Cah. Maboké 8:24. 1970. Fig. 1. = Hypochnus langloisii Pat., Bull. Soc. Mycol. France 24:3. 1908. = Pellicularia langloisii (Pat.) Rogers, Farlowia 1:101. 1943. s Botryobasidium langloisii (Pat.) Gilbn. et Budington, J. Arizona Acad. Sci. 6:92. 1970. = Phanerochaete insolita Burds. et Nakas., Mycologia 73:467. 1981. Basidiocarp annual, effused in small patches, up to 0.25 mm thick, farinaceous to hypochnoid, adnate but easily separable; fertile area discontinuous, farinaceous to pubescent, "greyish orange" (near 5B4 2/ or 6B4– ), staining red with 2% KOH; subiculum very thin, byssoid, con- colorous with hymenium; margin not differentiated, abrupt, irregular in outline.

2/ Burdsall and Nakasone (1981). – 391

Hyphal system monomitic. Subiculum a loose textura intricata, hyphae 4-9(-12) µm diam, thick walled (walls up to 5 µm thick), hyaline to pale yellow, encrusted with hyaline crystals and yellow to orange granules that dissolve in KOH producing a red solution, regularly branched, long celled, simple septate; subhymenium a textura intricata, hyphae firm walled, pale yellow, short celled, frequently branched in candelabrum pattern, encrusted with orange granules that dissolve in KOH turning solution pink; pseudocystidia cylindrical. 75-200 x 9-15(-21) µm, protruding up to 150 µm, with numerous simple septa, often constricted at septa, thin walled or with slight wall thickening, encrusted at first with a nearly translucent sheath up to 4 µm thick, later developing into a thin, orange granular coating; basidia broadly clavate, 15-18 x 6.5-8 µm, hyaline, thin walled, simple septate at base, 4 sterigmate, sterigmata up to 8 µm long; basidiospores ellipsoid to broadly allantoid, 7-9.5 x 3-4 µm, hyaline, thin walled, smooth, Melzer's -, acyanophilous. Habitat--On fallen well-decayed hardwood branches; associated with a white rot. Distribution--Known from Florida and Louisiana. Specimens examined--FLORIDA--HHB 9561 on Liquidambar styraciflua L., Leon County, holotype of Phanerochaete insolita (CFMR). LOUISIANA--Langlois 2968, sur bois pourri, St. Martinville, holotype of H. langloisii (FH). Discussion--Because of the poor condition of the cystidia and basidia in the type specimen of H. langloisii, the description and illustration are from the type of P. insolita.

CANDELABROCHAETE MAGNAHYPHA (Burt) Burds., comb. nov. Fig. 2. z Peniophora magnahypha Burt, Ann. Missouri Bot. Gard. 12:238. 1925 [1926]. Basidiocarp annual, effused in small patches up to 2 cm diam, appressed farinaceous to hypochnoid, adherent but easily separable; fertile area continuous, white to pale yellowish gray (near 3B2); margin abrupt, not differentiated; subiculum thin, not distinguishable. Hyphal system monomitic. Abhymenial surface not differentiated; subiculum a loose textura intricata, hyphae (6-)9-12(-15) µm diam, thin walled to firm walled, walls occasionally up to 1 µm thick, hya- line to pale yellow, with right-angle branching, simple septate or rarely nodose septate, smooth or encrusted with small brownish yellow granules and crystals; subhymenium hyphae much branched at right angles, 3.5-4.5 µm diam, thin walled, hyaline, smooth or encrusted with brownish yellow granules and crystals, simple septate; pseudocystidia arising from subiculum, cylindrical, with frequent simple septa, 100-150 x 9-12 µm, thin walled to firm walled, walls occasionally up to 1 µm thick, usually encrusted with brownish yellow granules and crys- tals, sometimes smooth, embedded in subiculum and protruding up to 40 µm above basidia; basidia broadly clavate, 12-15 x 5.5-6 µm, hya- line, thin walled, simple septate at base, 4-sterigmate, sterigmata up to 5 µm long; basidiospores ellipsoid, 6-7(-8) x 3-4 µm, hyaline, thin walled, smooth, Melzer's -, acyanophilous. Habitat-- On well-decayed hardwood; apparently associated with a white rot. Distribution--Known only from the type collection from Florida. Specimens examined--FLORIDA--R. Thaxter 57, on wood, Coconut Grove (FH). 392

Discussion-- Candelabrochaete magnahypha is not nonspecific with Candelabrochaete langloisii as suggested in previous studies (Rogers, 1943; Boidin, 1970). The basidiocarp of C. magnahypha is nearly white, not grayish orange as in C. langloisii. In addition, C. magnahypha possesses subiculum hyphae that are broader, much thinner walled, and with paler crystals. Finally, the ellipsoid basidiospores distinguish C. magnahypha from other Candelabrochaete species.

CANDELABROCHAETE SEPTOCYSTIDIA (Burt) Burds., comb. nov. Fig. 3. ~ Phanerochaete septocystidia (Burt) Erikss. et Ryv. in Erikss. et al., Cort. North Europe 5:1021. 1978. % Peniophora septocystidia Burt, Ann. Missouri Bot. Gard. 12:260. 1925 [1926]. s Scopuloides septocystidiata (Burt) Jülich, Persoonia 11:422. 1982. = Odonticium raitviirii Parm., Consp. Syst. Cort. p. 218. 1968 (fide Erikss. et al. 1978).

Basidiocarps broadly effuse, extending up to 8 x 4 cm, hypochnoid, thin, somewhat discontinuous; fertile area pubescent to farinaceous, pale yellowish tan, not changing color in KOH; subiculum concolorous; margin thin, white, pruinose to farinaceous. Hyphal system monomitic. Subiculum a loose textura intricata, poorly developed or up to 250 µm thick, hyphae 4-6(-9) µm diam, thin to thick walled (walls up to 1.5 µm thick), hyaline, simple septate, noticeably constricted at septa, branching at wide angles, smooth or irregularly coated with yellow-brown granular to globular material; subhymenium a dense textura intricata, hyphae short celled, 2.5-4 µm diam, frequently branched, densely encrusted with yellow-brown granular to globular material; pseudocystidia cylindrical, arising in subiculum at all levels, 60-150 x 5-9(-12) µm, protruding up to 35 µm beyond basidia, simple septate, with many septa, short celled, constricted at septa, firm walled to thick walled (walls up to 1.5 µm), apical cell with thinner walls, covered with large orange-brown granular and globular material, simple septate at base; basidia cylindrical to clavate, sometimes constricted, 12-18 x 4.5-5.5 µm, hyaline, thin walled, simple septate at base, 4-sterigmate, sterigmata 3-4 µm long; basidiospores allantoid, 4.5-6.5 x 1.5-2 µm, thin walled, hyaline, Melzer’s -, ~acyanophilous. Habitat--On angiospermous or occasionally gymnospermous wood; associated with a white rot. Distribution--Uncommon in eastern North America, Jamaica, and North Europe. Specimens examined--JAMAICA--Murrill and Harris 840, HMBG 61490, Troy and Tyre, Cockpit Country (BPI), holotype of Pe. septocystidia. NORWAY--Ryvarden 8182, on wood, Aust-Adger, Landvik:Skiftenes (GB, CFMR). U.S.A.: NORTH CAROLINA--RHB 4207 and 4208, on Liriodendron tulipifera L., Haywood County (CFMR); NEW YORK--HMBG 55968, on coniferous wood, Westport County (BPI); NEW JERSEY--HMBG 63455, on Pinus sp., Newfield County (BPI); MICHIGAN--HHB 10319, on Acer saccharum Marsh., Marquette County (CFMR); MINNESOTA--RLG 9759, on Populus balsamifera L., (ARIZ, CFWR) and FP 100695 on Populus sp., both Clearwater County (CFMR); TENNESSEE--HHB 3884, on Acer sp., Sevier County (CFMR); WISCONSIN--FP 101895, on hardwood log, Door County (CFNR).

394

Discussion-- Candelabrochaete septocystidia is distinguished from the other Candelabrochaete species by its allantoid basidiospores measuring 4.5-6.5 x 1.5-2 µm. Its distribution into the north temper- ate areas of North America and Europe is unusual for a species of Candelabrochaete because all the other species are from subtropical or south temperate regions. Jülich’s (1982) transfer of Peniophora septocystidia (as P. septocystidiata) to Scopuloides (Massee) Hjortst. et Ryv. (1979) because it possesses septate cystidia, is unwarranted. The basidio- carp structure and the type of cystidia of C. septocystidia are entirely unlike those found in species of Scopuloides. The septa of hyphae in the axes of the teeth of species such as Phanerochaete (Scopuloides) rimosa (Cooke) Burds. are not primary septa such as those found in C. septocystidia. Candelabrochaete verruculosa Hjortstam (1983), not known from North America, differs from C. langloisii and C. magnahypha in possessing shorter basidiospores, although the description is similar in other characteristics. It differs from C. septocystidia in possessing ovoid rather than allantoid spores.

PENIOPHORA MEXICANA Burt, Ann. Missouri Bot. Gard. 12:243. 1925 [1926]. Fig. 4. Basidiocarp broadly effuse, up to 0.25 mm thick, woolly, adherent; fertile area continuous, pubescent, "pale yellow" (4A2), not changing color in KOH; subiculum concolorous with fertile area; margin abrupt, pubescent, irregular in outline, concolorous. Hyphal system monomitic. Subiculum a loose textura intricata, hyphae 6-12 µm diam, walls up to 3.5 µm–thick, pale yellow to pale brownish yellow, smooth branching at right angles, long celled, simple septate; subhymenium compact, hyphae 4-6 µm diam, firm walled, pale yellow, smooth, with frequent branching, short celled, simple septate; pseudocystidia poorly differentiated, cylindrical, 90-150 x 9-20 µm, hyaline, aseptate, with walls up to 3 µm thick heavily encrusted with hyaline crystals, protruding up to 50 µm; basidia poorly preserved, not measurable, 4 sterigmate [... subcylindrical to short clavate, 9-11.5 x 6.5-7 µm, ... fide Liberta, 1968]; basidiospores ellipsoid to broadly ellipsoid, 7-10(-12) x (4)5-6 µm, hyaline, thick walled, smooth, Melzer's -, cyanophilous. Habitat--On hardwood slash and debris; associated with a white rot. Distribution--Known only from type locality. Specimens examined--MEXICO--Murrill 773, HMBG 54633, on wood, January 10-14, 1910, Veracruz, Orizaba (Lectotype and isotype BPI, FH). Discussion-- The specimens on deposit at BPI and that at FH are portions of a single collection; however the BPI specimen cannot be located at this time. The microscopic characters of P. mexicana suggest a relationship to Phanerochaete and Candelabrochaete (with broad simple septate hyphae and pseudocystidia) and to Hypochnicium (with thick walled spores). Because of the discrepancies, it is maintained in Peniophora until an appropriate generic placement is determined. It is discussed here only because of the superficial similarities (i.e. the septate cystidia). 395

LITERATURE CITED

ACKNOWLEDGMENTS

Mss. F. F. Lombard and K. K. Nakasone and Drs. R. L. Gilberton, R. Halling, and W. J. Sandberg are all thanked for critically reading the manuscript.