Vol. XIX, pp.

389-395 January-March 1984

THE GENUS CANDELABROCHAETE (CORTICIACEAE) IN NORTH AMERICA

AND A NOTE ON PENIOPHORA MEXICANA

By

HAROLD H. BURDSALL, JR.

Center for Forest Mycology Research, Forest Products Laboratory1— /

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 Phanerochaete 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.
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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.