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19
October
1992
AUS~.
syst. BO~.,
5,
639-43
Australasian Truffle-like Fungi.
VI*.
Gigasperma
(Basidiomycotina, Tricholomataceae) and
Horakiella
gen. nov. (Basidiomycotina, Sclerodermataceae)
Michael
A.
Castellano* and James M. TrappeB
AUnited States Department of Agriculture, Forest Service,
Pacific Northwest Research Station, Forestry Sciences Laboratory,
Corvallis, OR 97331, U.S.A.
BDepartment of Forest Science, Oregon State University,
Corvallis, OR 97331, U.S.A.
Abstract
Re-examination of both species assigned too the genus
Gigasperma
reveals significant differences between
the two taxa that warrant separation at the order level.
Gigasperma cryptica
is characterised by large,
smooth, extremely thick-walled, reddish brown spores borne on 4-spored basidia as in some Agaricales.
Horakiella clelandii
gen. et comb. nov. differs in its ornamented, hyaline to pale brown spores enclosed
by nurse hyphae as in the Sclerodermatales.
Introduction
Hydnangium clelandii
was erected by Rodway (1924) to accommodate a truffle-
like fungus characterised by pale yellow, globose, smooth to slightly rough-walled spores.
Hydnangium
Wallroth is based on
H.
carneum
Wallroth, which is related to the agaric
genus
Laccaria
Berk.
&
Br. and is characterised by hyaline to pale pink, subglobose
echinulate spores.
Gigasperma
was erected by Horak (1971) to accommodate a new
species from New Zealand which is similar to
H. clelandii
in the possession of subglobose,
extremely thick-walled, more or less smooth spores. At the time, Horak recombined
H. clelandii
as
G. clelandii
after studying a portion of the type. In the course of fungal
collecting in Tasmania, we collected a fungus that from field observations appeared
to be a hypogeous
Scleroderma
sp. Further examination revealed it to belong to the
Sclerodermatales based on spore ontogeny and it became apparent that it was conspecific
with
G. clelandii.
We present here a redescription of the two species assigned to the
genus
Gigasperma,
comment on placement and relatedness of these two species and
erect a new genus for
G. clelandii.
Methods of collection and macroscopic and microscopic procedures were generally
those of Smith and Zeller (1966). Herbarium names are abbreviated according to
Holmgren
et al.
(1990). Colours of fresh specimens are in general terms of the authors.
Hand-cut sections of both fresh and dried material were mounted in
5%
KOH or Melzer's
reagent for standard light microscopy. Spore dimensions are based on the measurement
of at least 50 randomly selected spores; spore lengths do not include sterigmal attachment.
Measurements of sterile tissues
(e.g. peridium) and spores are from mature sporocarps.
Light photomicrographs are from material mounted in
5%
KOH.
Taxonomy
Gigasperma
Horak,
N.
Zeal.
J.
Bot.
9:
49 1.
197
1.
Etymology:
Latin
giga,
large, and
sperma,
seed, for the large size of the spores.
*
Part
V,
Aust. Syst. Bot.,
1992,
5,
631-8.
1030-1887/92/050639%05.00
640
M.
A. Castellano and
J.
M.
Trappe
Distribution and Hosts
Known so far only from New Zealand. Probable mycorrhizal associate of
Nothofagus
SPP.
Discussion
Gigasperma
is characterised by the large, subglobose to globose, smooth, extremely
thick-walled spores that often have a distinct sterigmal attachment. Currently it is
restricted in distribution to New
Zealand and found exclusively under
Nothofagus
spp.
with which it presumably forms mycorrhizae.
Gigasperma
is considered to be monotypic
and of unknown relation to other truffle-like fungi or other members of the Agaricales.
Gigasperma cryptica
Horak,
N. Zeal.
J.
Bot.
9:
491. 197 1.
Basidiomata
hypogeous, pyriform or irregularly tuberiform, occasionally subglobose,
5-25mm in diam., pale brownish yellow as dried, surface sectored into more or less
uniform polygons, occasional adherent concolourous hyphae attached to slightly tomen-
tose surface.
Peridium in section
one layered, 0.3-0.4 mm thick, concolorous with with
surface.
Gleba
never exposed, friable, not gelatinised, pale reddish brown to reddish
brown marbled with tramal tissue that is concolorous with peridium and similar in
texture; locules round to elliptical, empty and lined with spores; sterile base distinct,
conical, and of similar structure and colour (slightly paler) to peridium, projecting 1-5 mm
from flattened base; columella absent. KOH on peridium: yellow-brown.
Odour
like
garlic.
Taste
acidulous.
Peridium
of hyaline to pale yellow-brown, non-gelatinised, thin-walled, subparallel
to closely interwoven hyphae, 3-6 pm in diam., hyphae more loosely interwoven at
surface; pale yellow pigment soluble from outer portion of
cutis; clamp connections
common.
Trama
thin, of hyaline, thin-walled, compact interwoven hyphae with many scattered
inflated cells.
Basidia
not crowded, hyaline, thin-walled, pear-shaped or peanut-shaped,
45-60
x
15-30 pm, 4-spored; sterigma stout, hyaline, 7-8
x
3-4 pm; basidia separated
by hyaline, thin-walled, inflated cells up to 25-30 pm in diam.; clamp connections
not seen.
Cystidia
absent.
Spores
smooth, globose to subglobose, 29-36 (-38) pm diam., sterigmal attachment
often conspicuous, 4
x
10 pm; spore wall 5-8 pm thick, sometimes somewhat thicker
at base than apex, penetrated by basal pore; in KOH reddish brown to orange-brown
singly, reddish brown in mass; no reaction in Melzer's reagent (Fig.
1.).
Etymology:
Greek
crypto,
'hidden' in reference to the sporocarps that were hypogeous
and thus hidden from view.
Distribution and Hosts
New Zealand; presumptive mycorrhiza formers with
Nothofagus fusca
and
N.
menziesii.
Collections examined
NEW ZEALAND: Nelson: Lake Rotoiti, track to St Arnaud Range,
E.
Horak,
23
.v.
1968
(holotype
PDD
27002,
isotype
ZT
681504);
Canterbury: Craigieburn State Forest Park, Cave Stream,
R.
E.
Beever
522,
19
.iv.
1986
(PDD
48944,
OSC).
Discussion
Gigasperma
differs from all known truffle-like fungi by virtue of its large, smooth,
extremely thick-walled, reddish brown spores that have a distinct basal pore.
Australasian Truffle-like Fungi.
VI
64 1
Figs
1-3.1,
basidiospores of
Gigasperma cryptica, Beever
522,
scale: 15 pm;
2,
basidiomata
of
Horakiella
clelandii,
H1556 (left and centre), HI557 (right), scale: 1Omm;
3,
basidiospores
of
H.
clelandii,
HO
113739, scale: 20 pm.
Horakiella
Castellano
&
Trappe, gen. nov.
A
Gigasperma
differt sporis ornamentalis et nonporis. A
Scleroderma
differt sporis
pallidis parietibus crassis e loculis persistentibus.
Etymology:
In honour of Dr Egon Horak, eminent mycologist of southern hemisphere
fungi.
Typus:
Horakiella clelandii
(Rodway) Castellano
&
Trappe
(E
Hydnangium clelandii
Rodway).
Distribution and Hosts
Known so far only from Tasmania, Australia. Probable mycorrhizal associate of
Myrtaceae.
642
M.
A. Castellano and
J.
M.
Trappe
Discussion
The genus
Horakiella
is erected to accommodate a species that has been placed
in four widely different truffle-like genera,
Hydnangium, Leucophlebs
Harkness,
Octa-
viania
Vitt., and
Gigasperma.
The spores are surrounded by nurse hyphae and the
sporocarp when collected has many macroscopic field characters in common with the
genus
Scleroderma
s.1.
Horakiella clelandii
(Rodway) Castellano
&
Trappe, comb. nov.
Hydnangium clelandii
Rodway,
Pap. Proc. Roy. Soc. Tasmania
1923, 108.
1924.
Leucophlebs
clelandi(Rodway)
Zeller
&
Dodge in Dodge
&
Zeller,
Ann. Missouri
Bot. Gard.
23, 596. 1936.
Octaviania clelandii
(Rodway) Cunningham,
Trans. Proc. Roy. Soc.
N.
Zeal.
67,
408. 1938.
Gigasperma clelandii
(Rodway) Horak,
N.
Zeal.
J.
Bot.
9,493. 1971.
Basidiomata
hypogeous, subglobose to slightly irregular, 11-14
x
11-20 mm, brownish
yellow, surface tomentose with patches of encrusted soil (Fig. 2).
Peridium
in section
one layered,
f
0.5 mm thick, dirty pale yellow at first, slowly staining red.
Gleba
never
exposed, solid, firm, dry, dark brown to brownish black, with white, pale yellow-white
to pale brown tramal plates; locules irregular to isodiametric, persistent, completely
filled with hyaline spores and some hyphae; sterile base and columella absent.
Odour
strongly of fresh peanut butter.
Taste
mild.
Peridium
200-300 pm thick, of hyaline to pale yellow, subgelatinised, thick-walled
(1-2 pm), closely interwoven hyphae 4-7 (-8) pm in diam.; yellow pigment instantly
soluble in KOH; tomentum covering the peridium of nontapered, subgelatinised, loose,
crystal encrusted, thick-walled
(f
2
pm), pale yellow hyphae
f
4 pm in diam.; clamp
connections common.
Trama
thin, of hyaline, thin-walled, compactly interwoven hyphae.
Basidia
not seen.
Cystidia
absent.
Spores
ornamented with a low to moderate reticulum, some spores enclosed within
nurse hyphae; subglobose to irregularly shaped, 25-30
x
30-35 pm, sterigmal attachment
inconspicuous; spore wall 4-7 pm thick, basal pore absent; in KOH hyaline to pale
brown singly, pale yellow-brown in mass; no reaction in Melzer's reagent (Fig. 3);
many aborted spores are small
(f
10 pm) and highly pigmented and scattered across
hymenium giving rise to the dark glebal colour.
Etymology:
In reference to Dr
J.
B. Cleland (1878-1971), eminent mycologist from
South Australia.
Distribution and Hosts
Tasmania, Australia; under
Acacia melanoxylon
and
Eucalyptus globulus.
Collections Examined
AUSTRALIA: Tasmania: Hobart,
L.
Rodway
(holotype HO 113739, isotype
PDD
8358); Cascades,
L.
Rodway 300,
v. 1925 (HO 113738); Cascades,
L.
Rodway,
ix. 1923 (HO 113737); Cascades,
L.
Rodway
319,
v. 1925 (HO); Huon Valley, south of Geeveston, Hermons Rd,
M.
Castellano,
H1556,
7 .v. 1990 (OSC); same locality and date,
M.
Castellano,
HI557 (HO, OSC); same locality and date,
M.
Amaranthus,
H1558 (HO, OSC).
Discussion
Horakiella clelandii
has had a much travelled history, having been placed in four
other genera,
Hydnangium, Leucophlebs, Octaviania
and
Gigasperma.
By virtue of
its nurse hyphae surrounding some spores and the characters of the sporocarp, it is
now readily apparent that
H. clelandii
is a member of the Sclerodermatales and is
Australasian Truffle-like Fungi.
VI
643
closely related to Scleroderma. The strong odour of fresh peanut butter (paste) is
unknown from any other truffle-like fungus. The glebal colour seems to be due more
to the colour of the small, aborted, highly pigmented spores than to the more hyaline
to pale brown mature spores. In fact, one specimen was collected that was almost
black in glebal colour and it was found to have an extremely high portion of highly
pigmented, aborted spores. Upon closer examination, we discovered that the other
Hydnangium species (H.
mcalpinei Rodway) described by Rodway
(1924)
is also a
member of the Sclerodermatales but is not congeneric with Horakiella clelandii.
Acknowledgments
Partial funding for this work was provided by CSIRO through ACIAR project
8736.
We appreciate the loan of material from the following herbaria: DSIR Plant Protection,
Auckland, New Zealand (PDD), and the Tasmanian Herbarium, University of Tasmania,
Hobart
(HO).
The senior author appreciates the opportunity to study fungi in the
private herbarium of Dr Horak.
References
Holmgren, P.
K.,
Holmgren, N. H., and Barnett,
L.
C.
(1990). Index Herbariorum. Edition 8. Part
I. The Herbaria of the World.
Regnum Vegetabile
120,
1-693.
Horak, E. (1971). Contributions to the knowledge of the Agaricales s.1. (fungi) of New Zealand.
New
Zealand Journal of Botany
9,491-3.
Rodway,
L.
(1924). Description of two underground fungi.
Papers and Proceedings of the Royal Society
of Tasmania
1923, 108.
Smith, A. H., and Zeller, S. M. (1966). A preliminary account of the North American species of
Rhizopogon. Memoirs of the New York Botanical Garden
14,
1-178.
Manuscript received 26 November 1991, accepted 21 May 1992