Hericium in North America: cultural characteristics and mating behavior

J. GINNS
Biosystematics Research Institute, Agriculture Canada, Ottawa, Ont., Canada KIA OC6
Received January 2, 1985

GINNS,
J. 1985. Hericium in North America: cultural characteristics and mating behavior. Can. J. Bot. 63: 1551- 1563.
Cultural characteristics observed in nearly 75 cultures were the basis for the descriptions of the four North American species
of Hericium. Comparison of the cultural characters showed subtle differences among the species. Similarities in cultural
characters led to the evaluation of nearly 4100 intercollection matings to confirm the identity of the cultures and establish
species circumscriptions. As a result all Canadian cultures labelled H. erinaceus were reassigned to H. americanum. The
presence of H. erinaceus in Canada could not be confirmed. Matings confirmed the conspecificity of North American and
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MICHIGAN on 11/11/14

European cultures of H. coralloides and H. erinaceus. Intracollection matings established or confirmed that H. abietis,
H. americanum, H. coralloides, and H. erinaceus have a bifactorial mating system.

GINNS,
J. 1985. Hericium in North America: cultural characteristics and mating behavior. Can. J. Bot. 63: 1551-1563.
L'auteur a utilisC les caractCristiques culturales de plus de 75 cultures pour dCcrire les quatre especes d'Hericium de
I'AmCrique du Nord. Les comparaisons des caractkres culturaux montrent des diffkrences subtiles entre les espkces. Les
similaritCs dans les caractkres culturaux ont conduit a I'evaluation d'environ 4100 croisements intercollection pour confirmer
I'identitC des cultures et Ctablir la circonscription des especes. Ceci a conduit a reassigner toutes les collection canadiennes
CtiquetCes, H. erinaceus, a H. americanum. La prksence de H. erinaceus au Canada n'a pas pu &treconfirmCe. Les croisements
ont confirm6 la conspecificit6 des cultures nord-amkricaines et europennes de H. coralloides et H. erinaceus. Les croisements
intracollection ont Ctabli ou ont confirm6 que H. abietis, H. americanum, H. coralloides et H. erinaceus sont bipolaires.
[Traduit par le journal]

Introduction (1978) were, no doubt, based upon correctly named cultures.

Four species of Hericium ( H . abietis (Weir ex Hubert) The cultural description for H . erinaceus subsp. erinaceo-
Harrison, H . ramosum (Bull.) Letell., H . coralloides (Scop. : abietis Burdsall, Miller and Nishijima was published when the
For personal use only.

Fr.) Pers., and H . erinaceus (Bull.: Fr.) Pers.) were accepted
as part of the North American mycoflora by Harrison (1973).
Recently application of two of these names has been altered.
Hallenberg's (1983) selection of a neotype for H . coralloides
placed H . ramosum, a name frequently seen in the North Amer-
ican literature, in synonymy with H . coralloides. Further,
Ginns (1984) found that the North American fungus tradi-
tionally labelled H . coralloides was not compatible with any
other European or American species and he proposed the name
H . americanum Ginns for it. Thus, the names used hereafter for
the North American species are H . abietis, H . americanum
(formerly H . coralloides of N. Am. auct.), H . coralloides
(sensu neotype = H . ramosum), and H . erinaceus.
Descriptions of cultures of several Hericium species have
been published, but the distinctions among the cultures of
the different species have not been itemized. Some cultures
used in preparing these descriptions were misnamed. Hericium
caput-ursi (Fr.) Banker ( = H . coralloides sensu neotype) was
described by Davidson et al. (1938), but the only culture of
theirs I have seen is H . americanum (see culture 12, below).
Hericium coralloides has been described by Boidin (1958,
p. 241), Stalpers (1978), and Hallenberg (1983). Hallenberg's
(1983) comment that Boidin's (1958) and Stalpers' (1978) de-
scriptions labelled H . coralloides were, in fact, of H . alpestre
is incorrect. Boidin's culture was identified as "CBS (Cart-
wright)" and appeared in the 1968 Centraalbureau voor Schim-
melcultures (CBS) catalog under the accession number 206.29.
I suspect that the culture was from England, outside the range
of H . alpestre Pers., and is H . coralloides sensu neotype.
Stalpers studied the Cartwright culture and a second culture,
493.63 in the 1978 CBS catalog. The latter was from New
York, U.S.A., and is H . americanum (see culture 13, below).
The descriptions of cultures of H . alpestre and H . coral-
loides sensu neotype by Hallenberg (1983) and of H . erinaceus
by Davidson et al. (1942), Boidin (1958, p. 242), and Stalpers
new subspecies was proposed.
The frequency with which Hericium species have been iso-
lated from decay in living trees, especially in Abies spp. and
Tsuga heterophylla (Raf.) Sarg. (Buckland et al. 1949),
Acer saccharum Marsh. (Nordin 1954), and Quercus spp.
(Davidson et al. 1942), and difficulties I had encountered in
assigning names to cultures of Hericium from decay in trees
emphasized the need for a comparative study of the four spe-
cies. In addition, intercollection matings were made to confirm
the identity of the isolates and intracollection matings were
made to determine the mating system of each species.
Materials and methods
In preparing the descriptions, the cultures, studied according to the
procedures of Nobles (1965) supplemented by Boidin (1966), were
grown at 25C in 85 mm diameter Petri plates containing 1.25% malt
agar. Basidiospores were germinated on 1.25% malt agar. Seven
tests were used to determine whether the actively growing mycelium
(3 weeks old) produced extracellular laccase and (or) tyrosinase. In
five tests a drop of reagent was placed at several points on each mat
(i.e., on the margin, the older mycelium, and on an area of interme-
diate age) and any change was recorded in the color of the drop at
3 rnin, 5 min, 1 h, and 3 h. The reagents were alcoholic solutions
of p-cresol, guaiacol, gum guaiacum, syringaldazine, and tyrosine.
Syringaldazine was prepared according to the directions given by
Harkin et al. (1974) and the other four reagents were prepared ac-
cording to Kaarik's (1965, p. 15) instructions. Five cultures of each
species were tested.
Two tests were performed using malt agar with either tannic acid or
gallic acid incorporated into the medium. These media were prepared,
tested, and evaluated following Nobles' (1965, D. 1099) instructions.
Nearly all cultures were tested.
A number of cultures were grown at five different temperatures for
18 days to determine whether growth rate at specified temperatures
could be useful in separating cultures of the different species. The
temperatures were 15, 20, 25, 30, and 35C. At each temperature the
radial growth in millirnetres was measured on five replicates of each
 1552 CAN. I. BOT. \'OL. 63, 1985
culture at 3-day intervals over the 18-day period. The average of the
five replicates was used to represent each culture.
Nearly 4100 intercollection matings were made to confirm the den-
tity of the cultures and to determine the number of intersterile groups
represented. Most matings consisted of a pairing of monokaryotic
mycelia from different individuals and in most instances four mono-
karyotic cultures from one collection werc paired with four monokary-
otic cultures from another collection (i.e., 16 matings were evaluated
to confirm or deny conspecificity). In somc matings it was necessary,
because of a lack of monokaryotic cultures, to pair four monokaryotic
cultures with a dikaryotic culture (e.g., Fig. 30). The matings were set
up by placing 5-mm' agar blocks containing mycelia about 10 mm
apart on malt agar Petri plates. After the mycclia had grown together,
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the mat was examined microscopically for clamp connections. The
presence of clamp connections was judged to be positive evidence
that the two cultures were conspecific. In monokaryon x dikaryon
matings, clamp connections werc sought in the area occupied by the
monokaryotic mycelium.
The mating system of each species was determined by pairing, in
all combinations, 10 or more monokaryotic cultures from one col-
lection of each species. After the mycelia had grown together, the
mat was examined for clamp connections. The grouping in tabular
form of the matings that produced clamp connections showed the
number of mating types (i.e., in this case four, which indicated bi-
factorial heterothallism).
Basidiomes representing each of the species were named by com-
parison with the species concepts in Harrison (1973) and the revisions
of Hallenberg (1983) and Ginns (1984). The cultures from these
basidiomes were used as standards in the mating tests. These standard
specimens and cultures were Her~ciumnbietis, DAOM 22748, culture
50; H. arnericanum, DAOM F2 167, culture 1 ;H. coralloides, DAOM
For personal use only.
2253 I, basidiome produced in culture 40; and H. erirzaceus, FP 58550
(at CFMR), culture 34.
The cultures studied (preserved at DAOM) are listed under the
species names that I have determined to be correct as follows.
Hericium arnericanum
1,' basidiome on Plarnnus, Pennsylvania, U.S.A., 5 November
1931, J. W. Sinden, L. 0 . Overholts Herb. 14844 (DAOM F2167),
culture from the type specimen for H. mmericnnum; 2, decay in live,
320-year-old Acer sacchnrum, central Ontario, V. J. Nordin, M 1784c
(DAOM 21467); 3, basidiome on Fngus, Ottawa, Ontario, 21 Sep-
tember 1936, J. W. Groves (DAOM F6902); 4, decay in Ulmus
thomasi Sarg., near Almonte, Ontario, October 1947, C. W. Fritz,
FPLO A-153 (DAOM 17672); 5, basidiome on wood, town of Maple,
Ontario, 5 July 1946, H. S. Jackson, TRTC 20542 (DAOM 17739);
6, decay in live, 360-year-old Acer saccharum, central Ontario, VJN,
M1728B (DAOM 21468); 7, basidiome on log of ?Acer, Chaffey's
Locks, Ontario, 9 September 1950, M. K. Nobles & C. G. McKeen
50-24 (DAOM 22606); 8, basidiome on Fagus grandifolia Ehr.,
Gatineau Park, QuCbec, 9 October 1957, R. Macrae, culture MKN
57-76 (DAOM 52836); 9, basidiome on Carya log, McDonald Island,
St. Lawrence Islands National Park (SLINP), Ontario, 10 October
1975, P. LeClair (DAOM 152154); 10, basidiome on standing Carya
ovara (Mill.) K. Koch, McDonald I., SLINP, Ontario, I October
1975, P. LeClair & M. Kaufert (DAOM 152155); 1 1, basidiome on
wood, Dorset, Ontario, 4 October 1975, J. Ginns 2970 (DAOM
154251); 12, received from CBS on 21 November 1950 (Originally
labelled H. caput-ursi. Culture from R. W. Davidson, U.S.A., ac-
cording to CBS 1950 catalog. No further data available. Apparently
same culture now CBS 129.40 (DAOM 22552)); 13, basidiome on
Tsuga, Leon, New York, coll. M. N. Schwalb, det. R. H. Petersen
'Numbers were assigned to simplify citation of cultures in the
text, tables, and figures. Each number represents a culture of distinct
origin, whether from a basidiome or decayed wood, despite some
similarities in collection data. Cultures incorporated when the study
was nearing completion were designated by a number and letter code
(e.g., 44a).
(CBS 493.63); 14, decay in Betuln alleghaniensis Britt., Swan Lake,
Ontario, J. Basham 467 KO; 15, decay in Fnglcs grnndifolin, Lindsay,
Ontario, JB, TA 8305(1); 16, same data as 15 except number TA
8420B; 17, basidiome on Fngus grnndifolin, Kejimkujik National
Park, Nova Scotia, 1 l September 1973, K. A. Harrison 11378; 18,
basidiome on Carya, State College, Pennsylvania, 26 October 1932,
L. L. Sluzalis, FP 56482-S; 19, basidiome on Fngus, near East Smith-
field, Pennsylvania, 14 October 1936, W. A. Campbell, FP 71370-S;
20, decay in Betuln pnpyrifera Marsh., Ontario, 1947, CWF (FPLO
A- 154); 2 1, decay in Fagus grnndifolin, Ontario, 1947, CWF (FPLO
A-155); 22, samc data as 21 except number FPLO A-156; 23, decay
in Betuln alleghaniensis, Ontario, 1930, CWF (FPLO A-157); 24,
decay in Populus tremuloides Michx., Ontario, 1954 (FPLO A-232);
25, decay in Larix lr~ricinn(Duroi) K. Koch, Ontario, 1958, J. K.
Shields (FPLO A-283); 26, basidiome on Acer sncchnrum, Ontario,
1958, JKS (FPLO S-489).
Hericium erirzaceus
28, decay in Qrcercus velutinn Lam., Mt. Alto, Pennsylvania,
4 January 1928, R. M. Nelson et nl., FP 53360-R; 29, basidiome on
live Quercus nlbn L., near Camden, New Jersey, 2 December 1930,
L. W. R. Jackson, FP 55175-S; 30, basidiome on live Fngus grnndi-
folin, near Bromall, Pennsylvania, 20 May 1932, LWRJ, FP 55 1 7 8 4 ;
31, basidiome on Quercus nlbn, near Camden, New Jersey, 23 Scp-
tember 193 1, LWRJ, FP 55 192-S; 32, basidiome on Acer sacchn-
rinum L., near Camden, New Jersey, 1 1 September 1931, LWRJ,
FP 55195-S; 33, basidiome on Quercus albn, State College, Penn-
sylvania, 8 November 1932, L. L. Sluzalis, FP 56471; 34, basidiomc
on Quercus rubra L., near Fairfax, Virginia, 2 November 1933,
R. W. Davidson, FP 58550-S; 35, basidiome on wood, ?Georgia,
February 1934, J. D. Diller & M. M. Lahman, FP 58570-S; 36,
basidiome on Quercus trunk, Laurel, Maryland, 21 November 1965,
0 . K. Miller 3766-S; 37, basidiome on standing Quercus, Laurel,
Maryland, 14 October 1966, J. Lindsay, OKM 4950-S; 38, pre-
sumably from basidiome on wood, Nunspeet, Netherlands, October
1931, J. Hoogland (CBS 202.31); 39, basidiome on Fagus, Burnham
Beeches, Bucks., England, October 1938, Dr. Ernest (FPRL 277).
Hericium cornlloides
40, basidiome on Fngus or Carpinus, Epping Forest (NE of
London), England, 9 November 1928, W. P. K. Findlay, FPRL 60
(DAOM 2253 1); 41, decay in Populus, Ontario, 1951, JB, T-30571-
40 (FPLO A- 182); 42, basidiome on Acer or Fngus, Fairfax, Virginia,
28 August 1934, R. W. Davidson, FP 59062-S; 43, basidiome on
rotting Beruln, Huron Mt. Club, Marquette Co., Michigan, September
1971, RHP (TENN 36357); 44, basidiome on decaying trunk of
Quercus robur L., Boda forest, land, Sweden, 14 October 1973,
A. Kaarik 73.150; 44a, basidiome on Populus, Palgrave, Peel Co.,
Ontario, 17 September 1946, R. F. Cain, TRTC 21219 (DAOM
17666); 44b, decay in Populus rrichocarpa T. & G., near Moose
Heights, British Columbia, G. P. Thomas er a/., DAVFP 2725
(DAOM 21 112); 44c, basidiome on decorticated windfall of Populus
rrichocarpa, Cinema, British Columbia, 1 1 August 1949, W. G.
Ziller, DAVFP 5080 (DAOM 22310); 44d, basidiome on Salix,
Crystal Bay, Ontario, 16 September 1956, J. W. Groves, culture
MKN 56-48 (DAOM 53898); 44e, basidiome on decorticated log of
?Populus, Cinema, British Columbia, I I August 1949, MKN,
DAVFP 5094 (DAOM 72184); 44f, basidiome on scar on ?Acer,
Dorset, Ontario, 6 October 1954, MKN 54-78 (DAOM 73315); 44g,
basidiome on Populus log, Black Sturgeon lake, Ontario, 6 September
1973, JG 2385 (DAOM 144673); 44h, same data as 44g except JG
2335 (DAOM 144680); 44i, received from RHP, 23 April 1976 as
Dodd 54 (no further data available); 44j, basidiome on rotting Beruln,
Big Bay, Marquette Co., Michigan, 2 September 1971, RHP 3631 1
(DAOM 191 156, TENN); 44k, basidiomc on Quercus robur, Boda,
Oland, Sweden, I I October 1972, AK 72.272; 441, basidiome on
Beruln alba s. Coste, Ullisjoki, Norrbotten, Sweden, 7 September
1974, AK 74.194-2; a m , basidiome on Populus tremula L., Ullis-
joki, 7 September 1974, AK 74.201-1; 44n, basidiome on Fagus,
Fanefjord skov, Moen, Denmark, 18 September 1980, N. Hallenberg,
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MICHIGAN on 11/11/14
For personal use only.
FIGS.1-4. Hericium nbietis. Variation in gross morphology of 6-week-old mats. Figs. 1 and 3 are replicates of culture 46. Figs. 2 and 4
were from cultures 47 and 45, respectively. Petri dishes 85 mm diam.
GB-145; 440, basidiome on fallen trunk of Fagus, Herrfallsang,
Hallsberg par., Narke, Sweden, 24 Septembcr 1981, NH, GB-331;
44p, basidiome from vicinity of Marquette, Michigan, 30 August
1971, RHP 36290 (DAOM 191 157, TENN).
Hericium cibietis
45, decay in Tsuga heterophylln, Cranbeny River, Kitwanga, Brit-
ish Columbia, Canadian Forestry Service, Victoria, VC 63-12; 46,
basidiome on large, dead, decayed conifer, Cameron, Vancouver
Island, B.C., 25 October 1952, A. T. Foster, A-131 (DAVFP 8276);
47, basidiome on old, large windfall of Tsugn heterophylln, Cameron,
V.I., B.C., 23 October 1952, D. J. MacPherson, A-98 (DAVFP
8278); 48, basidiome on broken Tsugn heterophylla, Cameron, V.I.,
B.C., 25 October 1952, ATF, A-134 (DAVFP 8280); 49, decay in
standing Abies or Tsugn, near Alberni, V.I., B.C., June 1945, R. E.
Foster et al., BSW 127 (DAOM 16601); 50, basidiome on break on
old windfall on Abies or Tsugn, Hope, B.C., 7 September 1950,
W. G. Ziller, DAVFP 6712 (DAOM 22748); 51, decay in Abies
nobilis Lindl., ?Idaho, E. Wright 58, received from R. W. Davidson,
6 June I944 (DAOM 1 1760); 52, decay in Tszcgn heterophylln, British
Columbia, P. J. Salisbury, V1984 (DAOM 16153); 53, decay in Abies
grandis Lindl., near Alberni, V .I., B .C., June 1945, REF et al., BSW
660 (DAOM 16602); 54, decay in Abies, same data as 53 except
number BSW 561-1 (DAOM 16676); 55, decay in Abies nmnbilis
(Dougl.) Forb., near Alberni, V.I., B.C., Summer 1946, D. C.
Buckland et al., BSW 1303 (DAOM 17054); 56, data as 55 except
number BSW 1257A-1 (DAOM 17183); 57, basidiome on Tsuga
heterophylla, Queen Charlotte Islands, B.C., 20 Aug. 1947. REF,
V3336 (DAOM 175 17); 58, decay in Tsiiga heterophylla, Q.C.I.,
B.C., summer 1947, REF et a/., H 21 06 (DAOM 21 107); 59, basid-
iome on Tsugci heterophylln, Cameron, V.I., B.C., 25 October 1952,
ATF, A- 135 (DAVFP 8279); 60, basidiome on Abies grnnclis, Copper
Canyon, V.I., B.C., 5 October 1949, ATF (DAVFP 5417).
Herici~crncilpestre
61, basidiome on Abies nlba Mill., Kalbenwald, Koralpe, Steier-
mark. Austria, 3 November 1981, N. Hallcnberg, culture 392; 62,
data as 61 except number 407.
Hericium erinncens subsp. erinnceo-nbietis
63, basidiome on live Quercus sp., Carvins Covc, Hollins,
Roanoke County, Virginia, U.S.A., 4 October 1975, J. Deutsch,
OKM 15159.
Results
Descriptions of cultural characteristics
Hericium abietis Figs. 1-4
Cultures growing slowly, up to 40 mm in radius in 6 weeks.
Mats in 2 weeks white to tan, appressed, an occasional culture
densely felty or sparsely woolly, submerged growth common
and in a fan-shaped pattern, margin even; mats at 4 weeks
white to yellow brown or white mottled with yellow brown,
submerged growth predominating, aerial growth appressed,
sparsely downy or sparsely cottony, with dense mycelial
growth on the inoculum; margin even to uneven, appressed
 1554 CAN. J. BOT. VOL. 63, 1985

TABLE1 . Radial growth (mm) of mycelial mats of Hericium species after 18 days at each of five temperatures

Incubation temperature ("C)

Total no. of
Species cultures tested 15 20 25 30 35
-
abietis 16 (8-)lo- 18(-20)* (10-)15-22(-30) (6-)lo- 19(-22) 0(-5) 0
coralloides 3 8-24 15-36 38-61 28-69 8- 17
americanum 1I (8-) 14-23(-49) (14-)25-46(-69) (17-)32-59(-83) (18-)3 1-39(-47) 2(-7)
erinaceus 12 (12-)18-25(-29) (12-)16-35(-39) 16-26(-49) (10-)27-31(-47) 0-9(- 15)
*Figures in parentheses are the minimum and maximum growth measurements. In H. coralloides only three measurements were available

TABLE2. Coding of the cultural characters in species of Hericirttn

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Species Code symbols from Nobles (1965)

- - - - - - - -

abietis 2a.*3.15. 34. 36.(38).(39). 47. (48). 51. (53). 55.60

americanum 2a. 3.15. 34. 36.(38). (44).46.(47).(48).(51).(53).54.55.60
coralloides 2a. 3.15. 32. 36. 38. 43-45-47. (48).(51). 53. 54. 60
erinaceus 2a. 3.15.(32).(34).36.(38).(39). 45-46-47. (48).(51).(53).54. 60
- - - -
*The "a" is a modification proposed by Boidin (1966, p. 6). It indicales that only laccase was detected. Boidin used
" b to indicate the presence of tyrosinase.

and in part submerged, growing in a radiating, fanlike pattern;
at 6 weeks (Figs. 1-4) the mats sodden with sparse aerial
growth, white, some pale yellow brown to pale brown, the
advancing zone submerged; the reverse side unchanged or
yellow brown to brown, the odor varying from indistinct to
very musty; basidiospores produced before 6 weeks by two of
the isolates.
For personal use only.
The results of pairing in all combinations of 11 single-spore
cultures from one basidiome (No. 46) showed there were four
mating types and the cultures were assigned the factors AIB,,
1 and 2; A,B2, 4 and 5; A2B2:3,6,8, and 9; and A2Bl, 10,11,
and 12. There were no illegitimate matings, and five pairings
were not made (i.e., 3 X 4, 3 X 5, 3 X 6, 4 x 6, 5 X 6).
Hericium abietis is bifactorially heterothallic.

Mycelial growth at five temperatures showed considerable
variation between cultures, e.g., after 18 days at 25C the
radial growth of the slowest culture was 6 mm, whereas that of
the fastest was 22 mm (Table 1). The growth rates at 15, 20,
and 25C were essentially the same, but at 30 and 35C only
two cultures showed a trace of growth. The others did not
grow.
Cultures numbered 45-60 were used in the preparation of
the description and the temperature - growth ratestudies.
Seven tests were performed to detect extracellular oxidases
on cultures 45, 46,47, 49, and 50. No tyrosinase was detected
with p-cresol and tyrosine. The test results for laccase were
variable. Guaicol was negative; syringaldazine was positive
but of weak intensity except in No. 47, where it was negative;
gum guaiacum was positive but weak.
On tannic acid agar only three cultures (51, 54, and 58) of
16 tested gave a positive reaction and then the reaction was
weak. On gallic acid agar the oxidase reaction was positive but
weak in intensity.
Hyphae of the advancing zone hyaline, thin walled,
(2-)2.4-3.2(-5) pm diam with relatively large clamp con-
nections, branches generally arising between the septa or
occasionally at a clamp; appressed and submerged hyphae es-
sentially as in the advancing zone but (i) some cells or seg-
ments slightly swollen, (ii) some narrow hyphae with numer-
ous refractive spots on the inside of the wall, (iii) some hyphal
tips with numerous short branches and nodules (resembling
staghorn branching) (Fig. 5), and (iv) some broad hyphae con-
torted with refractive areas and notable swellings. Gloeocys-
tidia (Fig. 6) usually common, up to 330 x 4-6 pm, terminal,
sometimes branched, contents oily and yellow, unchanged in
sulphobenzaldehyde, the apex often moniliform. Chlamydo-
spores (Fig. 7) rare to common, terminal or intercalary, ellip-
tical, 7- 14 X 4-8 pm, the wall hyaline, smooth, thin io
slightly thickened.
The species code (Nobles 1965) for H. abietis is compared
with the codes for the other species in Table 2.
Hericium americanum Figs. 5, 17-20
Cultures growing at a moderate to slow rate, the Petri plates
covered in 4-6 weeks or the mats 25-80 mm in radius at
6 weeks. Mats (Fig. 20) in 2 weeks white, 15-30(-55) mm
in radius, appressed (to 1 mm thick), texture variable, typically
dense and felty around the inoculum, often floccose, becoming
sparse toward the margin, with appressed, radiating, thin
strands bearing minute mycelial tufts in beadlike chains, some
mats producing sectors in which the growth was in a tree-
like pattern, some mats concentrically zonate, some cultures
with distinct submerged sectors; margin raised 1-2 mm,
1-2(-5) mm wide, even or irregular, and then with more
rapidly growing sectors which have segments branching in a
treelike pattern, some sectors submerged; reverse unchanged
except in a few cultures where a yellow or yellow brown
discoloration occurred beneath the inoculum; odor nil to musty;
at 6 weeks mats (Figs. 17- 19) white, most covering the dish,
the density of aerial growth varied, typically floccose, cottony
or felty on the older growth, the younger areas similar or with
sparser aerial growth which overlaid appressed strands with
minute tufts in beadlike chains, some concentrically zonate;
reverse unchanged or some mats with slight yellow discol-
oration; odor in most isolates a strong musty smell, in some
cultures only slight; fruiting in some isolates.
Measurement of mycelial growth at five temperatures
(Table 1) showed considerable variation among the cultures,
e.g., at 25C there was a 66-mm difference after 18 days
between the fastest and slowest growing cultures.
Cultures studied to prepare the description and in the tem-
perature - growth rate studies were Nos. 1-8 and 12- 19.
Seven tests were performed to detect extracellular oxidases
on cultures 1,6,8, 12, and 13. No tyrosinase was detected with
 GINNS 1555

p-cresol and tyrosine. Laccase was detected with guaiacol,

gum guaiacum, and syringaldazine, but the intensity was weak
to moderate.
On tannic acid agar the oxidase reaction was positive but
weak, except in two cultures, where it was moderate. On gallic
acid agar the oxidase reaction was positive but weak to mod-
erate. Cultures tested were Nos. 1-8, 12-16, 18, and 19.
Hyphae of the advancing zone hyaline, thin walled,
(2-)2.4-4.4(-6) pm diam, with clamp connections, with
branches arising at or between septa; aerial and submerged
hyphae as in the advancing zone, except the broad hyphae
4-6.4 p m diam, most cultures with noticeable numbers of
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narrow hyphae 1.6-3.2 pm diam with short branches, some

hyphae had irregularly swollen segments which produced a
few short branches, other hyphae, usually terminal segments,
only had short branches and resembled staghorns (Fig. 5).
Gloeocystidia terminal, cylindrical or slightly swollen, often
weakly branched, typically with moniliform apices, 100-
120 X 2.8-7 p m , the contents oily, refractive, pale yellow,
unchanged in sulphobenzaldehyde. Chlamydospores scattered
and uncommon, intercalary and terminal, fusoid to broadly
ellipsoid, 8.8- 12 X 4-8 pm, the wall hyaline, smooth, thin
to slightly thickened.
Mating of 10 monokaryotic cultures from culture No. 14 in
all combinations showed there were four mating types and the
cultures were assigned the factors A , B I , 4 and 13; A,B2, 10;
A2B2, 5,9, and 12; and A 2 B l , 6,7,8, and 1 1 . 'There were no
illegitimate matings. Hericium americanum is bifactorially het-
For personal use only.

erothallic.
The species code (Nobles 1965) for H . americanum is com-
pared with those for the other Hericium species in Table 2.

Hericium coralloides Figs. 12-16

Cultures growing at an intermediate to slow rate, the Petri
plates covered in 3-5 weeks or the mats 30-80 mm in radius
at 6 weeks. Mats in 2 weeks 12-25 mm in radius, white, of
varying morphology, typically appressed but sections distinctly
submerged with scattered aerial cottony tufts, downy to
cottony-woolly or felty, with a low ridge of raised mycelium
toward the margin, margin even, downy, appressed or with
submerged segments, 1- 2(- 5) mm wide; reverse unchanged;
at 6 weeks mats (Figs. 12- 15) white or with some cream color
around the inoculum, some cultures with concentric zones of
cottony mycelium, between the zones the mat appressed and
sodden, other cultures with mats downy-cottony, translucent,
toward the margin the hyphae appressed and often in narrow,
radiating strands bearing minute mycelial tufts in beadlike
chains (cf. Fig. 20); reverse unchanged to weakly yellow under FIGS.5- 11. Microscopic features from cultures of Hericiurn.
the mat; odor a strong musty or earthy smell; most cultures Fig. 5. Staghorn branching. Fig. 6. Gloeocystidia. Fig. 7. Chlamy-
fruiting, some after only 4 weeks incubation (Fig. 16). dospores. Fig. 8. Segments of a broad hypha. Fig. 9. Segment of a
Measurement of mycelial growth at five temperatures narrow hypha. Fig. 10. Segment of a narrow hypha with small knobs.
showed considerable variation among the three cultures, e.g., Fig. 11. Highly branched hyphal segment. All from 4-week-old
at 30C the difference in mat radius between the fastest and culture of H. erinaceus, culture 34, except Fig. 5, which is from
slowest growing isolates after 18 days was 41 mm (Table 1). H. americanum, culture 13. Scale equals 10 km.
Cultures used in preparation of description and tempera-
ture - growth rate studies were Nos. 40, 42, 44. reaction in No. 40. On gallic acid agar the oxidase reaction in
Seven tests were performed to detect extracellular oxidases the three cultures was positive but of weak to moderate
on cultures 40-44. No tyrosinase was detected with p-cresol intensity.
and tyrosine. Laccase was detected with guaiacol, gum guaia- Hyphae of the advancing zone hyaline, thin walled,
cum, and syringaldazine, but the intensity of the reaction was 2-3.3 p m diam with a relatively large clamp connection at
weak to moderate. each septum, branches arising at and between septa; aerial
On tannic acid agar no oxidases were detected in the three and submerged hyphae essentially as in the advancing zone
cultures (40, 42, 44) tested, except for a very weakly positive but some narrow (1.5-3 p m diam) (cf. Fig. 9) and others
 1556 CAN. 1. BOT. VOL. 63, 1985
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RGS. 12- 15. Hericium coralloides. Variability in the gross morphology of 6-week-old mats. Figs. 12 and 13 are replicates of culture 40.
Figs. 14 and 15 were from cultures 42 and 44, respectively. Petri dishes 85 mm diam.

4(-6) p m diam (cf. Fig. 8), frequently branched, the branches

FIG. 16. Hericium coralloides. Basidiome, I I mm broad, pro-
duced on 4-week-old culture 43.
often short. Gloeocystidia (Fig. 6) scattered at 2 weeks, more
common at 6 weeks, with refractive, oily, yellowish contents,
unchanged in sulphobenzaldehyde, about 130 x 3-4 pm, ter-
minal, often with some short branches, in some the apex
narrowing or weakly moniliform. Chlamydospores lacking.
The results of mating 10 monokaryotic cultures from one
basidiome (No. 40) showed four groups: A I B , ;1,3,17, and 19;
A2B2, 12 and 23; A,B2, 2,20, and 22; and A2BI, 18. In two
matings, 1 X 2 and 2 X 3, clamps were observed, but when 2
was subsequently mated with 1, 3, 18, 20, and 22, all five
were negative. An additional 13 monokaryotic cultures were
mated with one culture of each of the compatibility types. Each
of the 13 cultures formed clamp connections with only one
mating type and the 13 monokaryon cultures were assigned as
A I B , ,7 and 15; A,B2, 9,12,13, and 21; AIB2,6,10,11,14, and
16; and A,Bl, 4 and 8. Hericium coralloides is bifactorially
heterothallic.
The species code (Nobles 1965) for H . coralloides is com-
pared with the codes for other Hericium species in Table 2.
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For personal use only.
FIGS. 17-20. Mycelial mats of Hericium americanutn 6 weeks old, except 2 weeks old in Fig. 20. Fig. 17. Floccose aerial mycelium (culture
13). Fig. 18. Treelike or fanlike pattern of sectors (culture 4). Fig. 19. Extensive aerial mycelium (culture 1). Fig. 20. Mycelial strands with
minute, beadlike tufts (culture 14). Petri dishes 85 mm diam.
Hericium erinaceus Figs. 6- 11, 21 -28
Cultures growing at an intermediate to slow rate, the Petri
plates covered in 5-6 weeks or the mat only 4.5-8.5 cm in
radius at 6 weeks. Mats at 2 weeks white, growth typically
1-3.5 cm radius by varying from a trace to 4.5 cm; sub-
merged growth typically extensive and in a treelike pattern
(cf. Fig. 22), aerial growth appressed, sparsely cottony but
toward the margin downy and raised 2 mm, most cultures had
rapidly growing sectors which were characterized by a central
core of branching strands which were covered with minute
mycelial tufts in beadlike chains (cf. Fig. 22); margin sparse,
downy to submerged, even or irregularly undulating; at 6
weeks the mats (Figs. 21 -28) white, downy to cottony, some
with floccose patches,gome sodden, typically with some
strands which branch to form a treelike pattern and have minute
mycelial tufts in beadlike chains, in some cultures the myce-
lium principally submerged and usually developed in a treelike
pattern; reverse typically discolored yellow to brown but in a
few cultures unchanged; odor, if present, musty; fruiting on a
few plates.
Measurement of mycelial growth at five temperatures
(Table 1) showed some variation, but the rate of growth of
different cultures was more uniform at 15, 20, and 25C than
it was in the other Hericium species.
Cultures studied in preparing the description and in the tem-
perature - growth rate studies were 28 - 32 and 34- 39.
Seven tests were performed to detect extracellular oxidases
on cultures 28, 34, and 37-39. No tyrosinase was detected
with p-cresol and tyrosine. Laccase was detected with guaiacol,
gum guaiacum, and syringaldazine, but the intensity varied
from weak to moderate.
On tannic acid agar the oxidase reaction was positive, typi-
cally of weak or moderate intensity, but a few cultures showed
a moderate to strong reaction. On gallic acid agar the oxidase
reaction was positive, typically moderate to strong. Cultures
tested were Nos. 28-32 and 34-39.
Hyphae of the advancing zone hyaline, thin walled
2.4-4.4(-5.2) pm diam, with relatively large clamp con-
nections, branches usually arising between septa or sometimes
at a clamp connection; aerial and submerged hyphae as in the
advancing zone except narrow hyphae, 2-3 pm diam, more
frequently branched and some terminal segments of one or
several cells with numerous short branches which were often
weakly branched giving a staghorn appearance, and broad
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CAN. J . BOT. VOL 63. 1985

 GINNS
hyphae (Fig. 8) 5-7 pm diam, branched infrequently. Gloeo-
cystidia (Fig. 6) 60-400 x 4-9 pm, terminal, often weakly
branched, cylindrical to fusoid, typically moniliform at the
apex, contents refractive, oily, pale yellow, unchanged or
slightly darker in sulphobenzaldehyde. Chlamydospores
(Fig. 7) lacking to common, terminal or intercalary, fusoid to
subglobose, 8- lo(- 14) X 5-7(-8) pm, the wall smooth,
thin to slightly thickened, hyaline.
Mating of 10 monokaryotic cultures from one collection
(No. 34) showed the four groups A , B , , 5 and 12; A,B,, 6,8,
and 14; A2B2, 10,11, and 13; and A,B,, 2 and 3. One illegiti-
mate mating, 2 x 5, was seen where clamp connections were
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formed on the mycelium from isolate culture 5. Hericium
erinaceus is bifactorially heterothallic.
The species code (Nobles 1965) for H. erinaceus subsp.
erinaceus is compared with those for other Hericium species in
Table 2. Hericium erinaceus subsp. erinaceo-abietis Burdsall
et al. was not included in this phase of the study.
Intercollection matings
The results were distributed into five groups (Figs. 29 and
30). Within four groups (Fig. 29) the cultures were intrafertile
(i.e., clamp connections were produced in matings) but inter-
sterile (i.e., no clamps were produced in matings between
cultures from different groups). Of about 4100 matings sum-
marized in Fig. 29, about 3050 were monokaryon x mono-
karyon and abiut 1080 were dikaryon X monokaryon. Subse-
quently each group was assigned a species name based on the
For personal use only.
features of the basidiomes from which some cultures had been
derived. Thus each dikaryon or monokaryon culture was com-
patible only with others of the same species and incompatible
with collections of the other three species. In Figs. 29, 30, and
31 the numbers 1 through 26 represent H. americanum, num-
bers 28 through 39 are H. erinaceus, numbers 40 through 44p
are H. coralloides, and numbers 45 through 60 are H. abietis.
The fifth group (Fig. 30) consisted of 15 dikaryon cultures
which did not produce basidiospores; thus no monokaryon
cultures were available from these isolates and the nearly
380 matings using these cultures all had to be dikaryon X
monokaryon. In all these matings the monokaryons failed to
be dikaryotized.
To confirm the identity of these 15 cultures the basidiomes
which had given rise to nine of the cultures were studied. The
basidiomes for cultures M a , 4 4 - 4 4 h , 44j, and 44p had the
small spores and arrangement of hymenial spines along the
branches typical of H. coralloides (cf. Hallenberg 1983). The
basidiomes for the other cultures were not available. Since
H. coralloides and H. erinaceus are the only Hericium species
in Denmark and Sweden (J. Ginns, unpublished observations)
and because basidiomes of these species are readily distin-
guished, cultures 4 4 k , 441, 44m, and 440 were, no doubt,
correctly named H. coralloides.
The results from the intercollection rnatings showed that a
number of cultures had been misnamed. Of the five cultures
determined by matings to be H. coralloides (Nos. 40-44),
cultures 40 and 42 had been labelled H. coralloides N. Am.
aucts. (=H. americanum) when 1 received them. More sur-
prising were the 16 cultures (No. 1, 2, 4, 6 , 14- 16, 18-26)
originally labelled H. erinaceus that did not mate with other
F r ~ s 21-28.
. Hericium erinaceus. Variability in gross morphology of 6-week-old mats. Fig. 21. Extensive aerial growth (culture 31).
Fig. 22. Prominent aerial development of hyphal fans (culture 34). Fig. 23. Moderate aerial growth (culture 28). Figs. 24-26. Extensive
mycelial development within the agar which emerged in scattered spots (cultures 37, 33, 36, respectively). Fig. 27. Slow growth of aerial and
submerged hyphae (culture 39). Fig. 28. Very slow growth (culture 29). Petri dishes 85 mm diam.
1559
H. erinaceus cultures and were redetermined, using the mating
tests results, to be H. americanum. The only culture originally
labelled H. caput-ursi (No. 12) mated with H. americanum.
The cultures determined by matings to be either H. erinaceus
(Nos. 28-39) or H. abietis (Nos. 45-50) had been correctly
labelled when received.
Compatible matings (Fig. 29) resulted when H. erinaceus
from England (No. 39) and 'The Netherlands (No. 38) was
paired with a culture from the eastern United States (No. 34).
Also compatible were H. coralloides matings of cultures from
England (No. 40) and Sweden (No. 44) paired with North
American cultures (Nos. 41, 42, 43).
Matings between North American and European monospor-
ous cultures labelled H. coralloides sensu aucts. were one of
the final phases of this study because European cultures had
been lacking. All matings were incompatible and Ginns (1984)
concluded that the North American fungus was distinct from
the other species and proposed that it be called H. americanum.
The European fungus is now called H. alpestre (Hallenberg
1983). Monokaryon cultures of H. alpestre were not com-
patible with monokaryon cultures of H. abietis or H. erinaceus
subsp. erinaceo-abietis (Fig. 31). Further, subsp. erinaceo-
abietis was compatible with H. abietis (No. 46) and H. eri-
naceus (No. 34) but was not tested against H. americanum or
H. coralloides.
Discussion
Initially the purpose of this study was to describe the cultural
features of the North American species of Hericium and to
determine the distinguishing cultural features of each species.
When some cultures produced basidiospores, the study was
expanded to include intracollection monokaryon x mono-
karyon matings. The results showed all four species to have a
bifactorial mating system. Previously, H. cllpestre (Hallenberg
1983), H. coralloides (Fries 1941; Hallenberg 1983), and
H. erinaceus subsp. erinaceo-abietis (Burdsall et al. 1978) had
been reported to have a bifactorial system.
At the end of the initial study of the cultures no characters
had been found which obviously distinguished the species.
Perhaps some cultures had been misnamed and they were re-
sponsible for the apparent overlapping between species or
perhaps the species were so similar that few, if any, taxo-
nomically significant cultural characters existed. If the latter
were true, the validity of recognizing four species whose sepa-
ration had been based on small differences in spore size, a few
somewhat variable basidiome features, and apparent ecological
associations (Harrison 1973) would be questionable.
To identify misnamed cultures and to assess the validity of
the species concepts, the results of nearly 4100 intercollection
matings were used to determine conspecificity of the cultures.
The mating results fell into five groups. Four groups were
intracornpatible and intersterile (Fig. 29). Each of these groups
was assigned a species name. The fifth group (Fig. 30) con-
sisted of 15 dikaryon cultures which had not dikaryotized
monospore cultures representing the four North American
species. These cultures were assigned to H. coralloides based
on the study of some basidiomes or other available data (see
Results).
 1560 CAN. J . BOT. VOL. 63, 1985
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For personal use only.

AMERICANUM ERINACEUS ABlETlS

CORALLOIDES
FIG.29. Results of interspecific and intraspecific matings in Hericium. Numbers on the axes are numbers assigned to cultures described in
the Materials and methods section. Compatible matings produced clamp connections and are represented by a +; incompatible matings did not
produce clamp connections and are represented by a -. Typically each + or - symbol represents the results of mating four monospore cultures
from each of the partners, i.e., 16 matings.

The inability of the dikaryon cultures to dikaryotize the con-
specific monokaryon cultures (Fig. 30) is perplexing because
dikaryon-monokaryon matings using different cultures of
H . coralloides resulted in the formation of clamp connections.
Such incompatibility between different collections of one
species is not unknown. Mounce and Macrae (1938) found that
in Fomes pinicola there were two almost completely intersterile
groups in North America. Harmsen (1960) found some collec-
tions of Merulius himantioides to be compatible with a third
collection but not with each other. Boidin (1980, p. 53) dis-
cussed additional comparable examples. These examples indi-
cate that negative dikaryon-monokaryon mating results do not
necessarily mean that the paired cultures are distinct species;
alternatively examples of compatible matings between differ-
 MONOKARYON
AMEPICANUM EPINACEUS CORALLOIDES ABIETIS
The matings confirmed that there are definitely four species
I* 2 14 34 40 43 45 49 of Hcricium in North America and three in Canada. The geo-
DIKARYON 4 ' 5 2 4 6 1 0 2 3 3 5 6 8 1 0 2 3 4 2 1 2 3 1 2 3 4 2 3 4 5 6 2 3 9

I :-
44 p*
--- ------------ -- - --- graphic distribution of these species is generally known
---- ----------- ---- - ---
44 a
44 b
--- ----------- ---- - - (Harrison 1973; Smith and Smith 1973), but a more precise
44 C
--- ----------- ---- - definition of their ranges is needed. The present study some-
-- ---------- ---- -
44 d
44 e -- ---------- ---- --- what refines and supplements Harrison's (1973) compilation of
-- - - - - - - - - - - - ---- -----
the hosts and geographic distribution of the species in North
44 f
_ --------___ ---- - ---I
44 9
--_ -___--_-_-_ --- _ _--I America. Harrison noted H . american~itn(as H . coralloides)
44 h
44 1
- - - -- ----- __ - - -/
in North America as limited to east of the Great Plains and
--- ------------ -- -----
I1-
I --- --- --- -- north of North Carolina and Tennessee, where it is found on
44 k
--- --- --- 1- --
M I
44 rn
- -- -- --- I--- live trees and logs, etc., of Acer, Carya, Fagus, and Quercus.
P --- ---- ---
I The cultures studied herein were from Fagus in Nova Scotia
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'STUDY NUMBER
'MONOKARYON
ASSIGNED
CULTURE
CULTURE
NUMBER
and Quebec and from Acer, Betula, Carya, Fagus, Larix,
Populus, and Ulmus in Ontario, and from Carya, Fagus, and
FIG. 30. Results of intercollection rnatings between rnonokaryon Platanus in Pennsylvania and Tsuga in New York. The cultures
cultures representing the four North American species of Hericium from Larix and Tsuga are of interest because H . americatzum
and 15 dikaryon cultures. has been reported infrequently on coniferous wood in North
America. Harrison noted that "Populus has been a common

i i ERINACEUS
ERINACEO-ABIETIS
63"
SSP.
1 ABIETIS
46 49
I substrate," but he found no specimen on that host and neither
did Pomerleau (1980) in southern Quebec, and I had only
ALPESTRE lt204751 1 3 4 1 0 2 3 9
one confirmed culture from Populus. All of this suggests that
prior reports were based upon one of the growth forms of
H . coralloides that resembles H . americanum.
Hericium abietis causes a white pocket rot of Abies, Picen,
Pseudotsuga, and Tsuga from Alaska into California (Harrison
1973). All cultures studied were from Abies or Tsuga in British
Columbia, except No. 5 1, which was presumably from Idaho.
The only report of H . abietis from east of the Rocky Mountains
For personal use only.

FIG.31. Results of intercollection rnatings between rnonokaryon
cultures of Hericium alpestre and the taxa H. abietis and H . erinaceus
ssp. erinaceo-abietis.
ent species are lacking.
The intercollection matings reaffirmed Harrison's (1973)
recognition of H . abietis as a distinct species based on subtle
features of the basidiomes. Maas Geesteranus (1960) had
suggested that H . abietis was a synonym of H . coralloides
aucts. (including H . alpestre and H . americanum). Matings
between H. alpestre from Austria and H . abietis were incom-
patible (Fig. 31), as were numerous matings (Fig. 29) where
H . abietis was paired with H . americanum, H . coralloides, and
H . erinaceus.
The intercollection matings confirmed, on a genetic basis,
the long-accepted premise based on basidiome features that
H . coralloides and H . erinaceus were present in North
America, as well as Europe. However, cultures from North
America previously labelled H . coralloides N. Am. aucts. did
not mate with any European or North American species. They
have been renamed H . americanum (Ginns 1984).
The description of the cultural features of H . caput-ursi in
Davidson et al. (1938) may have been of H . americanum
because one culture (No. 12) from Davidson and labelled
H . ca~ut-ursimated with the four collections of H . ameri-
canum with which it was paired.
This study was concerned with cultures of Hericium species
and dealt with basidiomes only where it was necessary to con-
firm the name to be assigned to a culture. The reader should be
aware that considerable variation in the gross morphology of
the basidiomes exists and few microscopic features have been
found which distinguish the species. Although this variation
has been acknowledged (Harrison 1973; Maas Geesteranus
1959), the range within each species should be determined
by intercollection matings.
(Pomerleau 1980, from New Hampshire on a conifer trunk)
probably was based on a form of H . americanum.
Hericium coralloides is widespread in North America and
is the most commonly collected North American Hericilim.
Harrison (1973) reported the range of H . coralloides (as H .
ramosum) to be from Nova Scotia to Alaska and south to
Michigan and California. The cultures 1 studied were from
British Columbia, Ontario, Virginia, and Michigan. The spores
of some basidiomes did not germinate readily (J. Ginns, unpub-
lished observations), but this does not explain the relatively
infrequent records of H . coralloides from decay in live trees
in Canada. Harrison (1973) recorded the basidiomes only "on
decaying logs of frondose species" and perhaps H . coralloides
does not often cause decay of living trees. Basham (1958,
p. 496) reported only one culture out of 272 isolations from
live Populus tremuloides and culture No. 41 herein may be that
one. Basham and Morawski (1 964, p. 4 l), as a result of inter-
collection matings, labelled one culture from a live tree
H . coralloides (as H . laciniatum). Some of the cultures that
they recorded as "Hericiurn spp." from live Acer and Fagus
may have been H . coralloides. Most of the North American
cultures that I studied came from amentiferous trees, espe-
cially Populus, and a few from Betula, Fagus, and Salix. 1 am
not aware of any confirmed reports of H. coralloides from
conifer hosts.
The H . erinaceus cultures were from the U.S.A. (Maryland,
Pennsylvania, New Jersey, Virginia, and ?Georgia), and one
each was from The Netherlands and England. Harrison (1973)
reported H . erinaceus "from Florida to California in the south,
and in the north from Washington, southern Michigan and
New York State." Smith and Smith (1973) summarized its
distribution as "widespread in eastern and central U.S. and the
Pacific Coast where oak is found, . . ."
The 26 cultures herein named H . americanum were origi-
nally labelled H . coralloides aucts. (nine cultures), H . eri-
 1562 CAN. J. BOT. VOL. 63. 1985
naceus, and one was H. caput-ursi. In the intercollection
matings 16 (13 from Ontario and 3 from Pennsylvania) labelled
H. erinaceus mated with H. americanum. The 13 were all the
available cultures of H. erinaceus from Canada and all had
been isolated from rotten wood. These misidentifications pre-
sumably arose when Canadian cultures from decayed wood
were matched with a culture (No. 1, herein) labelled H. eri-
naceus that had been received by Mildred K. Nobles of Ottawa
from the laboratory of L. 0. Overholts of Pennsylvania. This
culture in the intercollection matings (Fig. 29) was not com-
patible with H. erinaceus but was compatible with cultures
finally assigned to H. americanum. The basidiome (Overholts
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Herb. 14844 at PAC) which had yielded this culture was
studied. It had spores 6.5 -7 X 6-7 pm and although
tuberculate had the spines borne in terminal clusters on short
branches. This collection was designated as holotype for
H. americanum (Ginns 1984). Most reports of H. erinaceus in
Canada have been based on cultures isolated from rotted wood.
Two studies (Basham and Morawski 1964; Nordin 1954) have
recorded H. erinaceus from decay in live trees in Canada.
Some of the 13 cultures initially labelled H. erinaceus were
from Basham and Nordin. One culture that Basham and
Morawski (1964, p. 41) labelled H. erinaceus from decay in
Betula had been named following matings, but I suspect the
tester strain may have been misnamed. However, it was not
possible to establish definitely that the previous reports of
H. erinaceus were in error. The occurrence of H . erinaceus
in Canada remains unconfirmed. Neither Harrison (1973) nor
For personal use only.
Pomerleau (1980) knew of any basidiomes from Canada. How-
ever, based on some basidiomes which were tuberculate and
had spines 2-4 cm long, Pomerleau (1984) accepted this spe-
cies as part of the QuCbec mycoflora. Hence, I examined two
basidiomes, QFB 2632 and 16392, labelled H. erinaceus, from
QuCbec from the herbarium where Pomerleau often deposits
collections, but I was not convinced that they were H. eri-
naceus. The specimen specifically mentioned by Pomerleau as
collected in September 1982 in the environs of QuCbec City
does not seem to have been preserved.
The differences between the species code for H . coralloides
in Hallenberg (1983) and in Table 2, herein, requires expla-
nation. The principal difference is that what I call gloeo-
cystidia, Hallenberg labelled gloeoplerous hyphae. Thus the
symbol 15 appears in Table 2 and the symbols 26 and 37 are
seen in Hallenberg's code. Presumably he was using the sym-
bols 26 and 37 to describe the sometimes inflated apices and the
yellowish contents of the gloeocystidia.
The code (Burdsall et al. 1978) for H. erinaceus subsp.
erinaceo-abietis is essentially the same as the code for H.
erinaceus, but Burdsall et al. used symbol 27 to represent "the
presence of normal cylindrical thick-walled hyphae with clamp
connections at septa." 1believe they meant to say "with a clamp
connection at each septum."
Cultures of the four species were tested for the presence of
tyrosinase, using p-cresol and tyrosine. All tests were nega-
tive, a result which contrasts with the results on Hericium
coralloides and H. erinaceus in some earlier studies. For
H . coralloides a positive test was reported with p-cresol by
Kakirik (1965, p. 66) and Stalpers (1978), whereas Boidin
(1958, p. 241) and Hallenberg (1983) reported negative tests.
By using tyrosine, both Boidin and Kaarik recorded positive
reactions, but Hallenberg reported a negative reaction. For
H . erinaceus a positive result was reported with p-cresol by
Boidin (1958), Kaarik (1965), and Stalpers (1978) and with
tyrosine by Boidin and by Kaarik. These authors tested few
isolates, often only one, whereas I have tested five of each
species. Regardless, the discrepancy exists and I have no
explanation for it.
The cultural similarities between the four species reinforced
the conclusion from studies on basidiomes (e.g., Harrison
1973; Maas Geesteranus 1959, 1971) that these species were
congeneric. The coding of the cultural features enables rapid
assessment of differences (Table 2). The first code numbers
for the Hericium species were typically 2a.3.15. Nobles'
(1965) key did not have any species code with this combina-
tion. Thus the series 2a.3.15 can be said to characterize the
genus Hericium. These three features are the presence of
laccase, clamp connections, and gloeocystidia. In addition, a
distinctive feature not accommodated in the code was the Dro-
duction of conspicuous beaded stands with the beads made
up of minute tufts of hyphae (Fig. 20). However, species of
Gloeocystidiellum Donk and Lentinellus Karst. and allied
genera related to Hericium may have similar codes.
Distinctions between the cultures of the North American
species of Hericium are subtle. The pertinent features can be
summarized as follows: H. abietis, mat pale brown, relatively
slow growth, no growth at 30 and 35"C, restricted to conif-
erous wood in western North America, and some hyphae have
refractive patches on the inner surface of the wall; H . ameri-
canum, mat typically dense, felty around the inoculum, often
floccose, becoming sparse toward the margin; reverse un-
changed, rarely slightly yellow; restricted to the eastern half of
North America; H. coralloides, the only species of the four
which lacks chlamydospores and does not stain tannic acid
agar; and H . erinaceus, mat typically with extensive sub-
merged growth, aerial growth appressed, sparsely cottony; re-
verse typically discolored yellow to brown; from eastern U.S.
and the Pacific Coast where oak occurs, not known in Canada.
In searching for additional features which would aid in sep-
arating cultures of Hericium species, the growth rate at five
temperatures was measured for 3 to 17 isolates of each species
(Table 1). The radial growth of the mycelium within species
often varied considerably (e.g., at 25OC the fastest and slowest
growing isolates of H. americanum had grown 83 and 17 mm,
respectively, after 18 days). At each temperature H . abietis
grew slower than the other species and at 30 and 35C a11
but 3 of the 17 isolates of H. abietis failed to grow. The average
growth of H. americanum, H. coralloides, and H. erinaceus
was about the same except at 25 and 30C, where H . erinaceus
grew slower. The growth rates were so variable that it doesn't
appear that they alone would be of value in distinguishing
cultures of the Hericium species.
Acknowledgements
Modra Kaufert, Christine Shugar, and Louise Lefebvre were
responsible, at different times, for preparing and examining the
matings. Several colleagues supplied many of the cultures or
items of information, in particular J. Boidin of Lyon, H. H.
Burdsall and Frances Lombard of Madison, Dorothy Chu and
Daphyne Lowe of Victoria, C. L. Fergus of University Park,
N. Hallenberg of Goteborg, K. A. Harrison of Wolfville, Anio
Kaarik of Uppsala, L. Magasi of Fredericton, F. Oberwinkler
of Tubingen, R. H. Petersen of Knoxville, J. Savory of Princes
Risborough, J. K. Shields of Ottawa, and J. Stalpers of Baarn.
A. Strid was most helpful during my 1982 visit to Naturiska-
historiska Riksmuseet, Stockholm.
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