Australian species of Elaphomyces (Elaphomycetaceae, Eurotiales, Ascomycota)

Abstract

The sequestrate ascomycete genus Elaphomyces is described and illustrated from Australia. The following thirteen new species are described: Elaphomyces aurantias, E. austrogranulatus, E. chlorocarpus, E. cooloolanus, E. coralloideus, E. laetiluteus, E. nothofagi, E. pedicellaris, E. queenslandicus, E. rugosisporus, E. suejoyceae, E. symeae, and E. timgroveii. A key is provided to all Elaphomyces species from Australia.

Full text

Australian species of Elaphomyces (Elaphomycetaceae,
Eurotiales, Ascomycota)
Michael A. Castellano
A,D
, James M. Trappe
B
and Karl Vernes
C
A
US Department of Agriculture, Forest Service, Northern Research Station, 3200 SW Jefferson Way,
Corvallis, Oregon 97331, USA.
B
Department of Forest Science, Oregon State University, Corvallis, Oregon 97331-5752, USA.
C
Department of Ecosystem Management, The University of New England, Armidale, NSW 2351, Australia.
D
Corresponding author. Email: mcastellano@fs.fed.us
Abstract. The sequestrate ascomycete genus Elaphomyces is described and illustrated from Australia. The following
thirteen new species are described: Elaphomyces aurantias,E. austrogranulatus,E. chlorocarpus,E. cooloolanus,
E. coralloideus,E. laetiluteus,E. nothofagi,E. pedicellaris,E. queenslandicus,E. rugosisporus,E. suejoyceae,
E. symeae, and E. timgroveii. A key is provided to all Elaphomyces species from Australia.
Introduction
Elaphomyces is a sequestrate genus of hypogeous Ascomycota
occurring across the northern hemisphere and Australasia. The
first published report of Elaphomyces is as Tubera cervina
(de L’Obel 1591). The genus name Elaphomyces was first
proposed by Nees von Esenbeck (1820) and sanctioned by Fries
(1825). Many authors contributed new species through the
years, the only comprehensive work being by Dodge (1929),
now outdated. Elaphomyces was first reported from the southern
hemisphere by Cooke (1892), who listed E. leveillei Tul.
(described originally from France) as occurring in Queensland,
Australia. Later, Rodway (1918) listed E. citrinus Vittad.
(described originally from Italy) from Tasmania. These reports
have subsequently proven to be erroneous as both collections
represent undescribed species that we treat below.
In the course of extensive collecting expeditions in forested
regions of Australia over the past 25 years (Lebel and Castellano
1999), we have unearthed over 100 collections of Elaphomyces.
In comparing Australian Elaphomyces specimens with
previously described Elaphomyces species from the northern
hemisphere, we found no overlap in distributions. All
Australian Elaphomyces collections belong to endemic
undescribed species. On the basis of peridial characteristics,
the Australian Elaphomyces species can be separated into two
broad groups, similar to those from the northern hemisphere.
Most Australian species have a very dark brown or black,
carbonaceous outer peridial layer, similar to that of many
species in the northern hemisphere, i.e. E. anthracinus Vittad.,
E. leveillei Tul. or E. viridiseptum Trappe & Kimbrough.
One species, E. austrogranulatus Castellano, Trappe &
Vernes, has a thick, brown, leathery peridium, similar to that
of E. granulatus Fries.
In the northern hemisphere, Vittadini (1831) divided
Elaphomyces into the following two subgenera: Malacoderma
typified by E. papillatus Vittad., characterised by species with
small (<15 mm) spores and a more or less soft and thin peridium
that is furrowed to wrinkled on drying, and subgenus
Scleroderma, characterised by species with large (>14 mm)
spores and a hard, brittle peridium unchanged when dried.
Vittadini (1831) further divided subgenus Scleroderma into two
unnamed groups, including (1) a group typified by E. leucosporus
Vittad., possessing a smooth outer peridium, and (2) a group
typified by E. reticulates Vittad., possessing a warty outer
peridium. Fries (1849) named the section in subgenus
Scleroderma that is represented by E. leucosporus,Ceratogaster
(Corda) Fr. and the section represented by E. reticulatus as
Ceraunion (Wall.) Fr. Subsequently, Dodge (1929) divided
subgenus Scleroderma section Ceraunion into two subsections,
mostly on the basis of the absence (subsection Hypogeum (Pers.)
Dodge) or presence (Phlyctospora (Zobel) Dodge) of a rooting
base to the ascoma.
The Australian Elaphomyces species have representatives that
can be placed into all these groups except subgenus Scleroderma
section Ceraunion subsection Phlyctospora, as none of the
Australian Elaphomyces species possesses a rooting base. The
use of spore size as a delimiting character among subgenera is not
supportable when placing Australian Elaphomyces species into
this scheme, because several small-spored species have a smooth,
hard, carbonaceous peridium and spores <15 mm in diameter
(i.e. E. chlorocarpus,E. pedicellaris, and E. rugosisporus).
Vittadini’s(1831) subgenus Scleroderma, if it were valid, would
now be subgenus Elaphomyces typified by E. granulatus
according to the International Code of Botanical Nomenclature.
In any event, we prefer to await the results of future molecular
analysis before recognising groups below the genus level.
The patterns of spore ornamentation vary markedly among
Australian Elaphomyces species, as do those of the northern
hemisphere species. Spore ornamentation ranges from
individual rods or spines to a complete reticulum or labyrinth,
or even an amorphous, plate-like structure. Scanning electron
CSIRO 29 April 2011 10.1071/SB10012 1030-1887/11/010032
CSIRO PUBLISHING
www.publish.csiro.au/journals/asb Australian Systematic Botany,24,32–57

microscopy (SEM) proved critical to elucidate the exact nature of
spore ornamentation to help distinguish among some species.
Correlating ornamentation structure that is observed with light
microscopy with what is shown with SEM helped sort out taxa
when observed with a light microscope at 1000. Molecular
information is presently lacking because extracting useable DNA
for sequencing has been difficult for most Elaphomyces
collections (Kerry O’Donnell, USDA, Agricultural Research
Service, pers. comm.). Until comprehensive molecular work is
accomplished, we hypothesise that the Australian Elaphomyces
species form a distinct clade or clades within the genus, as has
been demonstrated for many other sequestrate genera from
Australia (Hosaka et al.2006).
Elaphomyces does not so far overlap in species between
western and eastern portions of Australia, although some
have a broad latitudinal distribution. In all, 12 of the 13
Elaphomyces species described here occur with Myrtaceae or
Casuarinaceae, whereas only one, E. nothofagi, is associated
with Nothofagus (Nothofagaceae). E. nothofagi also possesses a
spore ornamentation pattern unique within the genus, with an
irregular reticulum beneath yarn-like hyphae.
Below we describe 13 new Elaphomyces species from
Australia and provide a key to them for identification and
classification purposes.
Materials and methods
Methods of collection and the macroscopic and microscopic
study were generally as described in Castellano et al.(1989).
Colours of fresh specimens are described in general terms.
Hand-cut sections of both fresh and dried material were
mounted in 5% KOH or water for standard light microscopy.
Spore dimensions include ornamentation and are based on the
measurement of 20 randomly selected spores, plus the largest
and smallest seen. Measurements of sterile tissues and spores
are from mature ascomata. Dried spores were mounted on 10-mm
diameter aluminium support stubs using double-sided tape,
rimmed with colloidal carbon and coated with 15 nm gold in
a Hummer 6.2 sputtering unit (Anatech, Springfield, VA).
SEM performed by Jim Ehrman used a JEOL JSM-5600 SEM
(JEOL USA, Peabody, MA) operating at 10 kV and 8–48 mm
working distance at the Digital Microscopy Facility, Mount
Allison University. SEM performed by Michael Castellano
used an AmRay 3300 FE SEM (AmRay, Bedford, MA) under
similar kV and working conditions as above. Holotype and
paratype specimens are deposited in appropriate mycological
herbaria in Australia depending on the origin of the collection
(CANB, DAR, HO, MEL or PERTH) and in the mycological
herbarium of Oregon State University (OSC) or Landcare
Research, New Zealand (PDD). Abbreviations are according
to Index Herbariorum (http://sciweb.nybg.org/science2/Index
Herbariorum.asp).
Taxonomy
Key to Australian Elaphomyces species (spore size
includes ornamentation)
1. Spores >40 mm in diameter; associated with Nothofagus .......................
........................................................................................ E. nothofagi
1. Spores <31 mm in diameter; associated with Acacia or Myrtaceae ...... 2
2. Spores 8–17 mm in diameter ................................................................. 3
2. Spores 18 mm or larger in diamete ........................................................ 8
3. Spores 8–9mm in diameter; Western Australia .................E. timgroveii
3. Spores 10–16 mm in diameter ............................................................... 4
4. Spores 14–16 mm in diameter; ornamentation of rods and spines ..........
........................................................................................ E. aurantius
4. Spores 10–13 mm in diameter; with at least a partial reticulum............ 5
5. Ascoma enclosed in deep orange, orangy-red, or red mycelium..........6
5. Ascoma enclosed in white or yellow or green mycelium..................... 7
6. Spores 11–12 mm in diameter (mean 11.1 mm); ornamentation a
pedicellate, fine reticulum; eastern Australia...............................
.................................................................................E. rugosisporus
6. Spores 12–13 mm in diameter (mean 12.8 mm); ornamentation a
pedicellate, irregular reticulum; Western Australia............ E. symeae
7. Spores 10–11 mm in diameter (mean 10.9 mm); glebal spore mass dark
greyish-blue; ascoma enclosed in bright yellow, yellowish-green to
green mycelium; Western Australia......................... E. chlorocarpus
7. Spores 11–13 mm in diameter; glebal spore mass black; ascoma enclosed
in white or yellow mycelium; eastern Australia ...............................8
8. Spore ornamentation of rods and spines to form a partial reticulum,
ascomata enclosed in white mycelium, Queensland...........................
.................................................................................... E. cooloolanus
8. Spore ornamentation of pedicellate, irregular plates forming a labyrinth,
ascomata enclosed in yellow mycelium, Victoria...............................
..................................................................................... E. pedicellaris
8. Spores 26–30 mm in diameter ............................................................... 9
8. Spores 18–25 mm in diameter ............................................................. 11
9. Peridium brown, leathery, with crowded pyramidal warts .....................
............................................................................E. austrogranulatus
9. Peridium black, carbonaceous, smooth or wrinkled...........................10
10. Spores 26–30 mm in diameter (mean 27.5 mm), ornamentation of tall
(2–5mm) labyrinthine ridges; ascomata enclosed in brown mycelium
...................................................................................... E. suejoyceae
10. Spores 23–26 mm in diameter (mean 24.5 mm), ornamentation of dense
rods that appear punctate when young, knobby and somewhat
amorphous when mature; ascomata enclosed in yellow mycelium ....
....................................................................................... E. laetiluteus
11. Spores 24–25 mm in diameter (mean 24.5 mm), ornamentation of
pedicellate, labyrinthine ridges, 3mm tall; glebal spore mass slate
grey to bluish-black at maturity ............................E. queenslandicus
11. Spores 18–20 mm in diameter (mean 19.1 mm), ornamentation a fine
coral-like labyrinth of narrow ridges; glebal spore mass dark brown
to black at maturity .................................................... E. coralloideus
Elaphomyces aurantius Castellano, Trappe & Vernes,
sp. nov. (Figs 1, 14a)
Ascomata in mycelio aurantio vel flavissimo omnino inclusa.
Peridium strati duorum compositum: stratum exterius 0.1 mm
crassum, anthracinum, carbonaceum, pagina levis vel parum
foveata; stratum interius 1.5–3 mm crassum, pallide griseum
vel griseum maculosum, venis albis marmoratum, interdum
erubescens. Gleba maturitate pulvereis griseo-caeruleis vel
fuscis sporis repleta. Asci globosi, 40 mm lati, parietibus
1–2mm crassis, 8-spori. Sporae globosae, 14–16 mm latae
virgasque spinas ornamenta 1–1.5 mm longas inclusae, in
maturitate in KOH viridinigrae vel obscure porphyreae.
Holotypus: Australia: Tasmania, Cradle Mountain National
Park, Waldheim, 13.v.1991, M. Castellano H5045 (HO, isotypes
PERTH, OSC).
Ascomata up to 10–22 11–30 mm, globose to subglobose or
irregular, completely enclosed in bright orange to occasionally
Australian species of Elaphomyces Australian Systematic Botany 33

bright yellow mycelium with much soil, roots and debris that
form a crust, the fresh yellow hyphae sometimes turning orange
over the course of the day after collection. Peridium 2-layered;
outer layer dark brown–black, carbonaceous, smooth to
somewhat pitted; inner layer pale grey mottled with grey and
marbled with mostly periclinal white veins, occasionally
Fig. 1. Elaphomyces aurantius.(a) Ascoma in cross-section. (b) Structure of the inner peridial layer. (c) Bundled hyphae. (d) Asci
with developing spores. (e) Spores viewed under light microscopy. ( f) Scanning electron micrograph of a spore. Scale bars = 3 mm
(a), 50 mm(b,c), 20 mm(d), 15 mm(e) and 5 mm(f).
34 Australian Systematic Botany M. A. Castellano et al.

blushing faintly pink near gleba where cut. Gleba initially hollow,
then filling with white, cottony-membranaceus, ascogenous
tissue that blushes slightly pink where cut; spore mass
powdery, greyish-blue in youth, becoming brownish-
black when mature, mixed with arachnoid hyphae that are
greyish-blue to olive, finally dark brown at maturity. Odour
mild. Taste not recorded.
Outer layer of the peridium approximately 100 mm thick, of
bundled hyphae 15–20 mm thick, arranged perpendicularly to
each other and composed of numerous individual brownish-
black, septate hyphae 3–4mm in diameter, with walls
approximately 1 mm thick; inner layer 1.6–3.1 mm thick at
maturity, of bundled hyphae 40–70 mm thick arranged
perpendicularly to each other and composed of numerous
individual, interwoven to subparallel, hyaline hyphae up to
7mm broad and with walls 1.5 mm thick. Gleba of spores
and intermingling, hyaline, septate, acutely branched, sinuous to
somewhat knobby, loosely interwoven hyphae 2–3mm broad
and with walls <0.5 mm thick. Asci globose, 40 mm broad,
pedicellate, hyaline, the walls 1–2mm thick, 8-spored, arising
from scattered, knot-like clusters of pale green (in KOH)
ascogenous hyphae 5–6mm broad. Spores globose, 14–16 mm
broad (mean = 15.0 mm) including the ornamentation of rods and
spines 1–1.5 mm tall, in clumps or bundles, giving the spore a
somewhat coarse appearance under light microscopy; SEM
reveals that single rods or spines often anastomose at their tips
to form clumps; spore walls 1mm thick, in KOH hyaline singly
and in mass in youth, at maturity greenish-black to dark reddish-
brown both singly and in mass.
Etymology
Latin, aurantius (orange), in reference to the colour of the
ascoma-enclosing mycelium.
Distribution, habitat and season
Queensland, south through New South Wales to Victoria
and Tasmania; sandy, coastal lowlands to tablelands and
mountains, hypogeous under various combinations of Acacia
longifolia,A. terminalis,A. verticillata,Allocasuarina
littoralis,A. torulosa,Corymbia intermedia,C. maculata,
Eucalyptus baxteri,E. botryoides,E. muelleriana,E. paniculata,
E. pilularis,E. resinifera,E. sieberi,E. tenuiramis,E. umbra,
Leptospermum sp. and Melaleuca linariifolia; March through
June.
Material examined
NEW SOUTH WALES: Mimosa Rocks NP, track leading to Gillards,
1.3 km E of junction with Tanja–Tathra road, 4.vi.1999, G. Mires
AWC3134 (DAR, OSC). QUEENSLAND: Atherton Tablelands,
Mothar Mountain, 9.v.1988, N. Malajczuk, M. Amaranthus &
M. Castellano H4091 (PERTH, OSC, PDD); 12 km W of Mt Glorius,
4.v.1992, J. Trappe H5971 (PERTH, OSC); Cooloola, 10.v.1988,
P. Reddell H4142 (PERTH, OSC); 10.v.1988, M. Amaranthus,
N. Malajczuk & M. Castellano H4147 (PERTH, OSC); 10.v.1988,
N. Malajczuk, M. Amaranthus & M. Castellano H4148 (PERTH,
OSC, PDD); and 10.v.1988, R. Young H4135 (PERTH,
OSC). TASMANIA: Cradle Mountain NP, along creek, 13.v.1991,
N. Bougher H5030 (HO, PERTH, OSC); Waldheim,
13.v.1991, N. Malajczuk H4884 (HO, PERTH, OSC) and 13.v.1991,
N. Malajczuk H4885 (HO, PERTH, OSC); Tasman Peninsula, vi.1916,
L. Rodway (HO, OSC). Tasman Peninsula, Wedge Bay, vi.1916,
L. Rodway (HO, OSC); near Colebrook, 30.ix.1991, C. Johnson
T31759 (HO, OSC, PERTH 07508042); VICTORIA: Cape
Conran–Sydenham Inlet Coastal Park (all following collections): car
park at ocean beach along Pearl Point Rd, 25.v.2001, J. Trappe,
A. Giachini & A. Jumpponen AWC4136 (MEL, OSC) and 1.vi.2003,
W. Colgan III & A. Claridge AWC5238 (MEL, OSC). Fletch’s Stretch,
0.2 km N of Pearl Point Rd, 27.v.2001, A. Jumpponen AWC4191
(MEL, OSC). Fletch’s Stretch, 0.6 km N of Pearl Point Rd,
28.v.1999, J. Trappe AWC2719 (MEL, OSC) and 28.v.2001,
A. Giachini AWC4140 (MEL, OSC); Yeerung River crossing, 29.
v.1999, J. Trappe AWC2780 (MEL, OSC) and A. Jumpponen
AWC2781 (MEL, OSC) and 27.v.2001, A. Giachini AWC4116
(MEL, OSC); 0.6 km E of Yeerung River crossing, 29.v.1996,
W. Colgan III T18498 (MEL, OSC) and 27.v.2001, A. Jumpponen
AWC4124 & AWC4119 (MEL, OSC); Alpine NP, Benambraorryong
Rd, 2 km S of junction with Sipthorp Track, 10.xi.1996, D. Mills
T20127 (OSC).
Discussion
Elaphomyces aurantius resembles E. rugosisporus in the
colour of the mycelium enclosing the ascoma and the black,
carbonaceous, smooth or somewhat smooth outer peridial
surface. The spores of E. auranticus are larger (14–16 mm,
mean = 15.0 mm) and more coarsely ornamented than those of
E. rugosisporus ((10–)11–12 mm, mean = 11.1 mm). The spores
of E. aurantius have spines and rods that are clumped to give a
coarse appearance, whereas E. rugosisporus has a fine, low,
complete reticulum that appears similar to the dimpling of a
golf ball when observed under light microscopy. Both species
occur in similar habit and have overlapping distributions
from Queensland south to Tasmania. E. aurantius is also
similar to E. cooloolanus in the appearance and size of their
spores under light microscopy. The spores of E. aurantius are
larger (14–16 mm, mean = 15.0 mm) and have a more clumped
ornamentation than those of E. cooloolanus (11–13 mm,
mean = 12.0 mm). E. aurantius is one of the more commonly
encountered Elaphomyces species of eastern Australia.
Rodway (1918) erroneously cited his collections as E. citrinus
Vittad.
Elaphomyces austrogranulatus Castellano, Trappe & Vernes,
sp. nov. (Figs 2,14b)
Ascomata in mycelio luteobrunneo vel aurantiobrunneo omnino
inclusa. Peridium strati trium compositum: stratum externum
verrucis pyramidalibus congestis usque ad 0.5 mm crassum,,;
stratum medium angustum, obscurum; stratum intimum 1–3mm
crassum, griseum, pallide violaceobrunneum, rhodobrunneum
vel fuscobrunneum. Gleba maturitate pulvereis nigris sporis
repleta. Asci globosi, 20–25 mm lati, parietibus maturitate
1–2mm crassis, 8-spori. Sporae globosae, (22–)24–28 (–32) mm
latae verrucas applanatas 4–5mm longas ornamentum inclusae,
in maturitate in KOH fuscorufae.
Holotypus: Australia: Queensland, Atherton Tablelands,
Davies Creek National Park, Davies Creek Road, 1.7 km from
end of road, 6.iii.1994, D. Arora, S. Joyce, & M. Castellano;
T13487 (CANB, isotype OSC).
Australian species of Elaphomyces Australian Systematic Botany 35

Ascomata 8–25 mm in diameter, subglobose, slightly flattened
to irregular, completely enclosed in golden brown, yellowish-
brown to orangy-brown mycelium with much intermixed soil and
debris, peridial surface yellowish-brown to brown, of crowded,
pyramidal warts, 0.5 mm tall. Peridium 3-layered; warts of outer
layer with a golden brown interior in cross-section and a thin
brown outer surface; middle layer a more or less uniform dark-
coloured line directly beneath the outer layer; innermost layer
Fig. 2. Elaphomycesaustrogranulatus.(a) Ascoma in cross-section. (b) Stacked hyphae found between peridial warts. (c) Thick-walled hyphae,
with scattered crystals found in the middle peridial layer. (d) Opaque mature spores. (e) Maturing spores, showing surface ornamentation.
(f) Scanning electron micrograph of a spore. Scale bars = 1 mm (a), 25 mm(b), 10 mm(c), 25 mm(d,e) and 10 mm(f).
36 Australian Systematic Botany M. A. Castellano et al.

grey to pale violet brown or rosy-brown to dark brown. Gleba
initially hollow, filling with hyphae then spores at maturity, spore
mass powdery, black, combination of hyphae and spores. Odour
mildly pleasant or indistinct. Taste not recorded.
Outer layer of the peridium 0.5 mm thick, warts composed of
yellowish–reddish-brown, irregularly shaped cells, 3–4mm thick,
with walls 1–2mm thick, and the hyphae between warts hyaline,
periclinal, septate and compact, with walls 2mm thick; middle
layer 100 mm thick, of interwoven hyphal bundles composed of
parallel, hyaline hyphae 5mm across, with walls 1mm thick;
the layer has a high concentration of dark reddish-brown pigment
dispersed across some hyphae, appearing at first as pigment
particles but when closely examined, is shown to be coloration
of parts of hyphal walls; inner layer up to 3 mm thick, of similar
structure as middle layer but with a significantly lower
concentration of the pigment across the hyphae. Gleba of
spores and thin-walled, hyaline to reddish-brown, highly
branched (90-degree angle), interwoven hyphae with cells
35 mm long and 1.5–2.5 mm broad, the hyphae becoming
dark reddish-brown and ornamented with knobby, coarse
crystals in age. Asci globose 20–25 mm in diameter,
pedicellate, hyaline, with walls 3–6mm thick when spores start
to develop, then thinner (1–2mm) when spores mature, 8-spored;
pedicel hyaline, 10 2–3mm; asci enlarging as spores develop
when mature; ascogenous hyphae not seen. Spores globose,
(22–)24–28(–32) mm broad (mean = 25.8 mm) including the
ornamentation of coarse, flattened, irregularly shaped warts,
4–5mm tall under light microscopy; SEM reveals an absence
of rods or spines; as spores mature, they become very dark and
large and the ornamentation is difficult to distinguish because
it becomes more dense with amorphous granules and hyphae
adhering to the spore surface, making the spore outline
subglobose to irregular in optical section; spore walls 1–2mm
thick, in KOH hyaline singly and in mass and unornamented or
slightly ornamented when immature, soon darkening to finally
dark reddish-brown to opaque when mature.
Etymology
Latin, austro- (southern) + -granulatus (granulate); hence,
‘southern granulate’in reference to its similarity to the
northern hemisphere E. granulatus complex.
Distribution, habitat and season
Northern Queensland, Victoria and Tasmania; tropical tablelands
to mountains, hypogeous under various combinations of
Acacia flavescens,A. melanoxylon,Allocasuarina littoralis,
Corymbia citriodora,C. clarksoniana,Eucalyptus
dalrympleana,E. fastigata,E. phaeotricha, and E. radiata;
December through May.
Material examined
QUEENSLAND: Atherton Tablelands, Davies Ck NP, Davies Ck Rd,
3.v.1988, C. Theodorou, J. Trappe & M. Castellano H4029 (CANB,
PERTH, OSC); 3.ii.1992, M. Castellano H5473 (OSC) and H5471
(CANB, PERTH, OSC); 19.iv.1996, K. Vernes T22544 (CANB,
OSC); 10.xii.1996, K. Vernes T22533 (CANB, OSC); Kennedy Rd,
5.v.1988, C. Theodorou, J. Trappe & M. Castellano H4059 (CANB,
PERTH, OSC); Mt Windsor Tablelands, 2.ii.1992, M. Castellano &
P. Reddell H5458 (CANB, PERTH, OSC); TASMANIA; Cascades,
Old Fern Rd, 2.v.1990, N. Malajczuk H1294 (HO, PERTH, OSC);
VICTORIA, Errinundra NP, Gunmark Rd, 1.6 km SE of Survey
Rd, 26.v.2001, A. Jumpponen AWC4100 (MEL, OSC); 27.v.1999,
A. Jumpponen AWC2695 (MEL, OSC).
Discussion
Elaphomyces austrogranulatus resembles E. granulatus Fries
from the northern hemisphere, particularly in macroscopic
peridial characters. They both have a distinctly warty, brown,
leathery outer peridium, with a non-marbled inner peridium. The
mature spores of E. granulatus are usually 39–40 mm in diameter
and coarsely ornamented with irregularly clumped rods as
opposed to the much smaller (24–28 mm in diameter) spores
of E. austrogranulatus that have coarse, flattened, irregularly
shaped warts. Some collections are packed with dark tissue that
becomes hard when dried and lacks spores. Whether this is a
parasite or aberrant ascomatal development is unknown.
Elaphomyces austrogranulatus is widely distributed,
although apparently uncommon. It is the only known
Elaphomyces from the southern hemisphere with a brown,
leathery peridium.
Elaphomyces chlorocarpus Castellano, Trappe & Lebel,
sp. nov. (Figs 3,14c)
Ascomata in mycelio laete luteo, luteo vel chlorino omnino
inclusa. Peridium strati trium compositum: stratum externum
tenue, anthracinum, carbonaceum, pagina verruculosa vel
rugulosa; stratum medium griseum; stratum intimum album.
Gleba maturitate pulvereis obscuris griseo-caeruleis sporis
repleta. Sporae globosae, 10–11(–12) mm latae ornamentum
clathratum punctatuminclusae, in maturitate in KOH
viridinigrae vel obscurae porphyreae.
Holotypus: Australia, Western Australia, 10 km west of
Albany, Howell Road, just off South Coast Highway,
25.vii.2001, K. Syme, M. Hart & R. van der Waag KS1157/01
(MEL 2233805; isotypes OSC, PERTH).
Ascomata 8–30 mm in diameter, globose to subglobose,
slightly flattened to irregularly lobed, completely enclosed in
bright yellow, yellowish-green to green mycelium with soil and
ectomycorrhizal roots that form a crust up to 5(–8) mm thick.
Peridium 3-layered; surface black, carbonaceous, smooth and not
wrinkled but with small, irregular bumps and folds, brittle and
easily cracked and chipped, roots adhering to peridial surface
forming a cross-hatched husk; interior 2–5 mm thick, 2-layered,
with an outer pale grey layer and an inner white layer. Gleba
initially hollow, then with white, cottony mycelium, at maturity
filled with dark greyish-blue spore powder and scattered
arachnoid hyphae. Odour indistinct to mild. Taste not recorded.
Outer layer of the peridium 50–75 mm thick at maturity, of
compact, short, hyphal bundles 3–4mm in diameter, with walls
1mm thick, somewhat branched, abruptly transitioning into the
layer beneath; middle layer of irregularly interwoven hyphal
bundles, sometimes these bundles are perpendicular to each
other, composed of pale grey, slightly wavy, compact hyphae
5–7mm across, with walls 2mm thick; innermost layer of similar
structure as middle layer but hyphae hyaline. Gleba of spores
and intermingled, hyaline, somewhat twisted or curled, septate,
elongate, interwoven hyphae 2mm in diameter, with walls
<0.5 mm thick. Asci hyaline, globose with a tapered base,
Australian species of Elaphomyces Australian Systematic Botany 37

Fig. 3. Elaphomyces chlorocarpus.(a) Surface of ascoma, without adherent crust. (b) Inner peridium. (c) Short, dark, thick-walled hyphae of the outer
peridial layer. (d) Asci arising from hyphal knot. (e) Maturing spores, showing surface ornamentation. ( f) Scanning electron micrograph of a spore. Scale
bars = 0.5 mm (a), 75 mm(b), 25 mm(c), 10 mm(d,e) and 5 mm(f).
38 Australian Systematic Botany M. A. Castellano et al.

25–26 mm in diameter, with walls 1mm thick, 8-spored; arising
from hyphal knots composed of distorted, hyaline, short in length
ascogenous hyphae 7mm in diameter, with walls up to 2 mm
thick. Spores globose, 10–11(–12) mm broad (mean = 10.9 mm)
including the ornamentation of an irregularly multidimensional
lattice that appears punctate under light microscopy; SEM reveals
an irregular reticulum-like structure that is several layers deep,
with fine holes in the lattice; spore walls 1mm thick, in KOH
yellowish-brown singly and dark yellowish-brown in mass when
mature, with spores appearing nearly uniform in size and
ornamentation.
Etymology
Latin, chloro- (yellowish-green) + -carpus (sporocarp), hence
‘yellowish-green sporocarp’, in reference to the colour of the
enclosing mycelium completely encasing the ascoma.
Distribution, habitat and season
Western Australia; hypogeous, gregarious, usually in deep litter
under various combinations of Allocasuarina fraseriana,
Corymbia ficifolia or Eucalyptus diversicolor; July, October
through December.
Material examined
WESTERN AUSTRALIA: E of Albany, Two Peoples Bay Nature
Reserve, 12.xii.1998, K. Syme, G. Evans, & M. Hart KS1040/98
(MEL 2105018, OSC); 15.xi.1998, G. Evans & M. Barrett KS1038/
98 (MEL 2105014); 22.x.1998, K. Syme & G. Evans KS1030/98
(MEL 2105009); Manjimup, Scatter Rd, 1.xii.1981, N. Malajczuk
(PERTH 07508093); Walpole–Nornalup NP, junction of Ficifolia
Rd and Nut Rd, 22.vii.1993, M. Castellano & J. Trappe T14919
(PERTH 07508123, OSC).
Discussion
Elaphomyces chlorocarpus has small spores similar to those
of E. cooloolanus,E. pedicellaris,E. rugosisporus and
E. symeae. E. chlorocarpus has the smallest spores of all
Australian Elaphomyces species and the yellowish-green
mycelium enclosing the ascoma easily distinguishes it from
all other small-spored species except E. pedicellaris.
E. chlorocarpus is restricted to Western Australia and
possesses a fine, punctate spore ornamentation, whereas
E. pedicellaris occurs in Victoria and has coarse spore
ornamentation. The appearance of the spore ornamentation
under light microscopy is closest to that of E. rugosisporus.
The multi-dimensional lattice structure of the ornamentation of
E. chlorocarpus is more complex than the fine reticulate
ornamentation of E. rugosisporus spores. The spore structure
is unusual in Elaphomyces, although it is somewhat similar
to that found in E. anthracinus from Europe, except in
E. anthracinus the fine reticulate spore ornamentation is
overlain with irregularly shaped plates to form a widely spaced
patchwork. This is one of three known Elaphomyces species
from Western Australia. These species can be distinguished
from each other in the field by the colour of the ascoma-
enclosing mycelium, which is black for E. timgroveii and red
for E. symeae.
Elaphomyces cooloolanus Castellano, Trappe & Vernes,
sp. nov. (Fig. 4)
Ascomata in mycelio pallide albo vel brunneolo inclusa. Peridium
strati duorum compositum: stratum externum tenue, nigrum,
laeve; stratum intimum 1–1.2 mm crassum, in juventute
lavandulum, in maturitate brunneolum. Gleba in maturitate
pulvereis nigris sporis repleta. Sporae globosae, 11–13 mm
latae ornamentum spinisque, virgisque cristiscongestis 1.0 mm
altum labyrinthum irregularis facientibus inclusae, in maturitate
in KOH fuscorufae.
Holotypus: Australia, Queensland, Cooloola, Rainbow Road,
10.v.1988, M. Castellano H4139 (CANB, isotypes PERTH &
OSC).
Ascomata 7–20 mm in diameter, globose to subglobose,
completely enclosed in white to pale tan mycelium,
ectomycorrhizal roots and sand that form a crust up to 7 mm
thick. Peridium 2-layered; surface black, smooth, rubbery; inner
layer rubbery, lavender when young, then pale brown at maturity.
Gleba initially hollow, becoming filled with black spore powder
at maturity. Odour not determined. Taste not determined.
Outer layer of the peridium 50–80 mm thick, of interwoven
bundles of hyphae 20–30 mm in diameter, composed of dark
brown to reddish-brown or nearly black, parallel at times,
agglutinated, interwoven hyphae 2–3mm in diameter with
walls 1mm thick; subcutis 1–1.2 mm thick, of interwoven
bundles of hyphae 12–15 mm in diameter, composed of pale
brown, interwoven hyphae 4–5mm in diameter with walls
1mm thick; peridial tissues much more disorganised than in
other species. Gleba composed of spores and thin-walled, septate,
hyaline, loosely interwoven hyphae 2–3mm in diameter, some
hyphae withered, some robust, some curly. Asci hyaline, globose
with a tapered base, 28–30 mm in diameter, collapsing around the
spores when mature to become irregular in outline, the walls
1.0–1.5 mm thick, 8-spored; tapered base hyaline, 4 mm wide by
4mm long, somewhat tapered to 3 mm wide at base, arising from
knots or clusters of hyaline, contorted ascogenous hyphae
5mm in diameter, scattered throughout the gleba. Spores
globose, 11–13 mm (mean = 12.0 mm) including ornamentation
that appears as coarse, crowded spines, rods and short ridges,
1mm tall, forming an irregular-looking labyrinth under light
microscopy; SEM reveals the ornamentation as individual rods or
spines that anastomose at their tips in irregular clumps, these
clumps are crowded to form almost a complete covering at times;
spore walls 0.5 mm thick, in KOH initially hyaline soon dark
green at maturity then reddish-brown singly and in mass.
Etymology
The place name, cooloola +-anus (relating to), hence relating to
Cooloola, a sand dune area along the southern Queensland coast.
Distribution, habitat and season
Queensland, known only from the Cooloola coastal dune area in
southern Queensland; hypogeous under various combinations
of Allocasuarina littoralis,Corymbia intermedia,
Eucalyptus pilularis,E. resinifera,E. umbra,Leptospermum
spp. and Melaleuca linariifolia; May.
Australian species of Elaphomyces Australian Systematic Botany 39

Fig. 4. Elaphomyces cooloolanus.(a) Ascoma in cross-section, with adherent crust. (b) Hyphal bundle of the outer peridial layer. (c) Asci with
developing spores. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of spores. ( f) SEM of a spore, showing the spines
and rods that anastomose at their tips. Scale bars = 1 mm (a), 20 mm(b,c), 10 mm(d), 5 mm(e) and 1 mm(f).
40 Australian Systematic Botany M. A. Castellano et al.

Material examined
Known only from the type collections.
Discussion
Elaphomyces cooloolanus has small spores, similar in size to
those of E. chlorocarpus,E. rugosisporus and E. timgroveii. The
deep orange to reddish-orange mycelium enclosing the ascoma of
E. rugosisporus easily distinguishes it from E. cooloolanus which
has white to pale tan mycelium on the ascoma surface and from
E. chlorocarpus which has yellowish-green mycelium enclosing
the ascoma. E. timgroveii has smaller and more finely ornamented
spores (8–9mm, mean = 8.7 mm) than those of E. cooloolanus
(11–13 mm, mean = 12.0 mm).
The spore ornamentation appears to be 1mm tall when
observed in cross-section under a light microscope and much
lower than that observed at the surface view or by SEM. This
is because the ornamentation of rods or spines anastomose
at their tips to form a mostly intact outer shell. This also
occurs in E. pedicellaris,E. queenslandicus,E. symeae and
E. timgroveii.
Elaphomyces coralloideus Castellano, Trappe & Vernes,
sp. nov. (Fig. 5)
Ascomata in mycelio fusco omnino inclusa. Peridium strati
duorum compositum: stratum externum usque ad 0.5 mm
crassum, anthracinum, carbonaceum, verruculosum; stratum
intimum usque ad 1 mm crassum, albidum. Gleba in maturitate
sporis pulvereis fuscis vel nigris repleta. Sporae globosae,
18–20(–23) mm latae ornamentum 2mm altum cristis
angustis, labyrinthum, coralloides facientibus inclusae, in
maturitate in KOH atrocastaneae.
Holotypus: Australia, Queensland, Atherton Tablelands,
Davies Creek National Park, Davies Creek Rd, 19.iv.1996,
K. Vernes T22547 (CANB, isotype OSC).
Ascomata up to 1.8 2.5 cm in diameter, subglobose,
completely enclosed in dark brown mycelium with much
soil, ectomycorrhizal roots and debris, when dried the ascoma
collapses and forms a large reticulate pattern to the surface.
Peridium 2-layered; surface black, carbonaceous, with low,
rounded, circular warts 0.4 mm in diam.; inner layer up off-
white, more or less uniform. Gleba powdery, dark brown to black
when mature. Odour not determined. Taste not determined.
Outer layer of the peridium up to 0.5 mm thick, of dark
reddish-brown to black near the surface to dark reddish-brown
to reddish-brown as it nears the inner layer, thick-walled, short
segmented or bead-like, parallel hyphae 3mm in diameter;
a distinct demarcation between layers in colour but not in
structure; inner layer up to 1 mm thick, of hyaline to pale
reddish-brown, thick-walled (3mm thick), interwoven
hyphae with enlarged ends, hyphae 8–11 mm in diameter, ends
enlarged up to 10–20 mm in diameter. Gleba composed
of spores and thin-walled, hyaline, somewhat encrusted,
sinuous, loosely interwoven hyphae, 2mm in diameter. Asci
subglobose and shortly pedicellate, up to 37 42 mm, the walls
2mm thick, hyaline at first then pale greenish-black from
maturing spores, 8-spored; ascogenous hyphae not seen.
Spores globose, 18–20(–23) mm (mean = 19.1 mm) including
ornamentation of anastomosed rods that form a fine, coral-like
labyrinth of narrow ridges, 2mm tall; SEM reveals the ridges
to be composed of anastomosed rods that form sinuous,
undulating lines; spore walls 1–2mm thick, in KOH initially
hyaline singly and in mass, soon pale reddish-brown then
finally dark reddish-brown when mature.
Etymology
Latin, coralloideus (coral-like), in reference to the coral-like
spore ornamentation as observed by SEM.
Distribution, habitat and season
Known only from northern Queensland; hypogeous under
various combinations of Acacia flavescens,Allocasuarina
littoralis,Corymbia citriodora,C. clarksoniana or Eucalyptus
phaeotricha; September and December.
Material examined
QUEENSLAND: Atherton Tablelands, Davies Creek NP, Davies Ck Rd,
10.xii.1995, K. Vernes T22565 (CANB, OSC); 29.ix.1996, K. Vernes
T22567 (CANB, OSC).
Discussion
Elaphomyces coralloideus resembles E. cooloolanus,
E. queenslandicus and E. suejoyceae in the black,
carbonaceous outer peridium enclosed by tan to brown
mycelium. The spores of E. coralloideus (18–20(–23) mm,
mean = 19.1 mm) are much larger than those of E. cooloolanus
(11–13 mm, mean = 12 mm) and have a distinctly different
ornamentation. The spores of E. suejoyceae are larger
((24–)26–30(–31) mm, mean = 27.6 mm) than the spores of
E. coralloideus and the labyrinthine ornamentation is
much finer in appearance. The spores of E. queenslandicus
and E. coralloideus nearly overlap in size; however, the
ornamentation of E. queenslandicus is much coarser in
appearance.
Elaphomyces coralloideus is restricted to northern
Queensland and seems to fruit irregularly, but this may be an
artefact of its being underrepresented in herbaria.
Elaphomyces laetiluteus Castellano, Trappe & Vernes,
sp. nov. (Figs 6,14d)
Ascomata in mycelio laete luteo omnino inclusa. Peridium strati
duorum compositum: stratum externum 50–65 mm crassum,
brunneonigrum vel anthracinum, carbonaceum, laeve; stratum
intimum 1 mm crassum, in juventute laete flavovirens, in
maturitate fumeum vel zonatum. Gleba in maturitate sporis
pulvereis nigris repleta. Sporae subglobosae vel globosae,
(22–)23–26(–28) mm latae ornamentum virgas dispersas vel
congestas inclusae, in maturitate in KOH purpureonigrae.
Holotypus: Australia, Queensland, Atherton Tablelands,
Davies Creek National Park, Davies Creek Road, 5.iii.1994,
D. Arora & M. Castellano T13452 (CANB, isotype OSC).
Ascomata up to 18 25 mm, subglobose to irregular,
completely enclosed in bright yellow mycelium with much
soil, ectomycorrhizal roots and debris forming a distinct
Australian species of Elaphomyces Australian Systematic Botany 41

crust. Peridium 2-layered; surface brownish-black to black,
carbonaceous, smooth, somewhat wrinkled when dried; inner
layer bright yellowish-green when young, then evenly smoky or
zonate with a pale zone near the surface layer, very dark when
dried. Gleba powdery, black when mature. Odour indistinct.
Taste (of peridium) nutty.
Fig. 5. Elaphomyces coralloideus.(a) Surface view of the ascoma, with crust removed. (b) Large ends cells within the inner peridial layer.
(c) Asci with developing spores. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of a spore. ( f) SEM of
a spore, showing the construction of the narrow ridges. Scale bars = 1 mm (a), 20 mm(b–d), 10 mm(e) and 2.5 mm(f).
42 Australian Systematic Botany M. A. Castellano et al.

Peridial tissues leaching a dark greenish-black pigment when
mounted in 5% KOH, outer layer of the peridium 50–65 mm thick,
of subhyaline to pale green, thick-walled (1–2mm thick) and thin-
walled hyphal elements 4–6mm in diameter; inner layer 1mm
thick, of hyaline, thick-walled, septate, capitate, interwoven
hyphae, up to 10 mm in diameter, with the hyphae spotted with
Fig. 6. Elaphomyces laetiluteus.(a) Surface view of the ascoma, with crust removed. (b) Capitate cells found with the inner peridial layer.
(c) Asci with developing spores. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of spores. ( f) SEM of
a spore, showing individual rods and clumps of rods. Scale bars = 3 mm (a), = 20 mm(b,c), = 10 mm(d,e) and 2 mm(f).
Australian species of Elaphomyces Australian Systematic Botany 43

numerous tiny to medium-sized occlusions. Gleba composed of
spores and thick-walled (0.5–1.0 mm diameter), hyaline, loosely
interwoven, septate, thread-like hyphae, 3mm in diameter. Asci
globose with a tapered base, 50–52 mm in diameter, the walls
2mm thick, hyaline at first, then smoky grey from maturing
spores, 8-spored arising from knots or clusters of pale green,
ascogenous hyphae 5mm broad, scattered throughout the gleba.
Spores subglobose to appearing globose, (22–)23–26(–28) mm
(mean = 24.6 mm) including ornamentation of scattered and
irregularly clustered spines or rods, 2mm tall, that give a
punctate appearance under light microscopy; SEM reveals the
clustered spines anastomose to form a mosaic pattern of clumps
and individual spines; spore walls 5mm thick when young,
thinning at maturity, in KOH initially hyaline singly and in mass
then dark greenish-black to pale violet to finally dark purplish-
black when mature.
Etymology
Latin, laeti- (bright) + -luteus (yellow), in reference to the bright
yellow enclosing mycelium.
Distribution, habitat and season
Northern Queensland, New South Wales and Victoria;
hypogeous under various combinations of Allocasuarina
littoralis,Eucalyptus cypellocarpa,E. dalrympleana,
E. fastigata,E. pauciflora,E. stellulata or E. tereticornis;
March, May, June and September.
Material examined
NEW SOUTH WALES, Coolangubra State Forest, Coolangubra Forest
Way, 1 km N of the junction with Wog Way, vi.1992, A. Claridge T17601
(DAR, OSC); QUEENSLAND, Davies Ck NP, Davies Ck Rd,
28.ix.1996, K. Vernes T22562 (CANB, OSC); VICTORIA, Alpine
NP, Native Cat Track, 0.3 km S of the junction with Black Mountain
Rd, 16.v.2001, A. Claridge AWC3476 (MEL, OSC).
Discussion
Elaphomyces laetiluteus can resemble E. aurantius in the colour
of the mycelium enclosing the ascoma. Usually, the enclosing
mycelium on E. aurantius ascomata is bright orange but
sometimes it is bright yellow when fresh and turns orange
over time after collection. The spores of E. aurantius are
distinctly smaller (14–16 mm, mean = 15.0 mm) and differ in
colour (dark reddish-brown) from the dark purplish-black
spores of E. laetiluteus ((22–)23–26(–28) mm, mean = 24.6 mm).
Elaphomyces laetiluteus is widely distributed in eastern
Australia, although rather infrequently encountered.
Elaphomyces nothofagi Castellano, Trappe & Vernes,
sp. nov. (Figs 7,14e)
Ascomata in mycelio cremeo omnino inclusa. Peridium strati
duorum compositum: stratum externum 70–80 mm crassum,
nigrum, carbonaceum, laeve; stratum intimum 1–2mm
crassum, in juventute viridoluteum, in maturitate olivaceum
vel prasinum. Gleba in maturitate sporis pulvereis nigris
repleta. Sporae globosae, 43–55 mm latae ornamentum cristis
implicatis, labyrinthinis inclusae, in maturitate in KOH
atroporphyreae.
Holotypus: Australia, Victoria, Tarra-Bulga National Park,
Tarra Valley picnic area, 17.vi.1994, J. Trappe H6811 (MEL,
isotypes PERTH, OSC).
Ascomata up to 12 18 mm, irregularly flattened, completely
enclosed in cream-coloured mycelium, ectomycorrhizal roots
and soil that form a crust. Peridium 2-layered; surface black,
carbonaceous, smooth; inner layer olive green near outer layer
and pale to bright green underneath when mature, greenish-
yellow when young. Gleba initially hollow, lined with brown,
wet hyphae, at maturity with a wet, black spore mass. Odour not
recorded. Taste not recorded.
Outer layer of the peridium 70–80 mm thick, of green to olive,
compact, interwoven hyphae up to 10 mm in diameter, with
walls 0.5–1.0 mm thick, encrusted with debris and fine crystals,
particularly the outer hyphae; inner layer 1–2 mm thick,
rehydrated, of hyaline, compact, interwoven to parallel hyphae
4–5mm in diameter; the demarcation between layers distinct.
Gleba composed of spores and septate, thread-like, curly,
hyaline, loosely interwoven hyphae 3mm in diameter, with
walls 0.5 mm thick. Asci not seen but knots or clusters of
pale green, contorted, ascogenous hyphae 5–6mm in diameter
scattered throughout the gleba. Spores globose, 43–55 mm
(mean = 47.9 mm) including ornamentation of infolded
labyrinthine ridges 5–7mm tall under light microscopy; SEM
reveals the folds to be entangled hyphae emanating from the
spore surface and these hyphae are sometimes flask-shaped
and appear yarn-like; walls 5–6mm thick, in KOH dark
reddish-brown singly and in mass.
Etymology
Latin, nothofagi, in reference to the association of this species
with the tree genus Nothofagus (Nothofagaceae).
Distribution, habitat and season
Victoria; hypogeous under Nothofagus cunninghamii; June.
Material examined
Known only from the type collections.
Discussion
Elaphomyces nothofagi is the only Australian Elaphomyces
species discovered to be associated with Nothofagus. It is also
unique amongst Australian Elaphomyces species in that it has
very large spores (43–55 mm, mean = 47.9 mm) and is covered by
entangled hyphae, similar to nurse hyphae of some Scleroderma
spp. It is known only from the holotype collection.
Elaphomyces pedicellaris Castellano, Trappe & Vernes,
sp. nov. (Fig. 8)
Ascomata in mycelio luteolo omnino inclusa. Peridium
strati trium compositum: stratum externum 100–125 mm
crassum, nigrum, carbonaceum, laeve vel prope laeve; stratum
medium 2 mm crassum, album, succulentum; stratum intimum
0.7–1 mm crassum, albidum, cavitatibus irregularibus. Gleba
in maturitate sporis pulvereis nigris repleta. Sporae globosae,
11–13(–14) mm latae ornamentum catillis irregularibus,
pedicellatis inclusae, in maturitate in KOH luteobrunneae.
44 Australian Systematic Botany M. A. Castellano et al.

Fig. 7. Elaphomyces nothofagi.(a) Interface between the outer and inner peridial layers. (b) Sinuous hyphae within the gleba. (c) Surface
view of spores under light microscopy. (d) Cross-sectional view of spores under light microscopy, showing enclosing hyphae. (e) Scanning
electron micrograph (SEM) of spores. ( f) SEM of a spore, showing the entangled hyphal cells. Scale bars = 10 mm(a,b), 20 mm(c–e)
and 10 mm(f).
Australian species of Elaphomyces Australian Systematic Botany 45

Fig. 8. Elaphomyces pedicellaris.(a) Ascoma in cross-section. (b) Surface view of the ascoma, with crust removed. (c) Interface of outer and
middle peridial layers. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of a spore. ( f) SEM of spore surface,
showing the pedicellate plates with rugose surface. Scale bars = 3 mm (a), 2 mm (b,c), 10 mm(d), 5 mm(e) and 2 mm(f).
46 Australian Systematic Botany M. A. Castellano et al.

Holotypus: Australia, Victoria, Gippsland Plain, Cranbourne,
Royal Botanic Gardens, near works department on hill above
woodlands near boundary fence, 11 Aug. 2000, T. Lebel TL233
(MEL 2063443, isotype OSC).
Ascomata 8–28 8–25 mm in diameter, globose to
subglobose, completely enclosed in pale yellow to yellow
mycelium, sand and ectomycorrhizae that forms a tightly
adherent mesh enclosing the ascoma. Peridium 3-layered;
surface black, carbonaceous, smooth or nearly so, becoming
somewhat irregularly wrinkled when dried, firm; inner peridium
white and fleshy towards the surface, with a thin, off-white layer
of small, irregular cavities. Gleba at maturity filled with black,
powdery spore mass. Odour disagreeable. Taste not recorded.
Peridium is overlain by cottony, hyaline, somewhat encrusted
hyphae. Outer layer 100–125 mm thick, of compact, smooth, dark
brown, shortly segmented, highly branched (90-degree angle)
interwoven hyphae 5mm in diameter and with walls 1mm
thick, outer 50 mm dark then gradually paler towards middle layer;
middle layer 2000 mm thick, of hyaline, compact interwoven
hyphae 4mm in diameter and with walls 2mm thick,
occasional hyphae bundled in 2’s, 3’sor4’s; inner layer
700–1000 mm in diameter, hyaline, interwoven hyphae 5–6mm
in diameter and with walls 2mm thick, mostly in bundles of
many hyphae, with the bundles loosely interwoven in a cris-cross
pattern. Gleba of spores and septate, elongate, curly, hyaline,
loosely interwoven hyphae, 2–3mm in diameter and with walls
0.5 mm thick, mostly smooth but some bumpy or kinked.
Asci and ascogenous hyphae not seen. Spores globose,
11–13(–14) mm broad (mean = 12.2 mm) including the
ornamentation of irregularly shaped, pedicellate plates that
appear somewhat warty under light microscopy; SEM reveals
that the plates are supported by rods and spines 1–2mm tall, and
the surface of the plates is finely rugose; spore walls 1mm thick,
in KOH pale yellowish-brown to yellowish-brown singly and
in mass when mature.
Etymology
Latin, pedicellaris (pedicellate), in reference to the pedicellate
spore ornamentation.
Distribution, habitat and season
Victoria; hypogeous under trees in open woodland; August.
Material examined
Known only from the type collections..
Discussion
Elaphomyces pedicellaris is similar to E. cooloolanus and
E. rugosisporus in spore size, but differs in spore
ornamentation, as revealed by SEM, in the colour of the
enclosing mycelium and distribution. The type collection of
E. pedicellaris was associated with wombat diggings.
Elaphomyces queenslandicus Castellano, Trappe & Vernes,
sp. nov. (Figs 9,14f)
Ascomata in mycelio fusco omnino inclusa. Peridium strati
duorum compositum: stratum externum 100–200 mm crassum,
nigrum, carbonaceum, verrucis irregularibus, applanatis;
stratum intimum 1–2 mm crassum, plus minusve pallide
luteobrunneum vel brunneum, columbrinum. Gleba in
maturitate sporis pulvereis schistaceis vel atramentariis repleta.
Sporae globosae, (23–)24–25(–26) mm latae ornamentum
1–3mm altum cristis labyrinthinis, pedicellatis inclusae,
maturitate in KOH atrovirides vel brunneae.
Holotypus: Australia, Queensland, Atherton Tablelands,
Davies Creek National Park, Davies Creek Road, 5.iii.1994,
D. Arora & M. Castellano T13457 (CANB, isotype OSC).
Ascomata up to 22 mm in diameter, subglobose, completely
enclosed in dark brown mycelium, soil and ectomycorrhizae.
Peridium 1–1.5 mm thick, 2-layered; surface black,
carbonaceous, of irregularly sided, flattened warts up to
0.5 mm in diameter; inner layer rose-tinged directly beneath
the carbonaceous layer, then mottled pale tan and off-white
but basically pale tan towards the surface, grading to more
brown toned near the gleba; when dried, ascoma collapses to
form large wrinkles on the surface. Gleba initially stuffed with
white cottony mycelium, at maturity becoming powdery with
slate grey to dark bluish-black spores. Odour indistinct. Taste not
determined.
Peridium overlain by a layer of bristly, dark reddish-brown,
thin-walled, septate, somewhat knobby, extended hyphae that
are slightly encrusted; outer layer 100–200 mm thick, dark
reddish-brown to black textura angularis near the surface to
dark reddish-brown to reddish-brown to red near the subcutis,
the cells 10–18 mm in diameter; inner layer 1.0–1.4 mm thick, of
hyaline to pale tan, septate, compact, interwoven hyphae
2–5mm in diameter and with walls 0.5 mm thick. Gleba
composed of spores and hyaline to pale green, sinuous,
loosely interwoven hyphae 4–5mm in diameter and walls
0.5 mm thick. Asci subglobose to globose, with a short tapered
base, 45–55(–70) mm in diameter and walls 1–1.5 mm thick,
hyaline at first, then greenish-black from maturing spores,
8-spored, the tapered base 5mm long 4mm wide, arising
from knots or clusters of pale green, short-segmented,
contorted, ascogenous hyphae 5mm in diameter scattered
throughout the gleba. Spores globose, (23–)24–25(–26) mm
(mean = 24.5 mm) including ornamentation of pedicellate,
labyrinthine, rounded ridges 3mm tall under light
microscopy, smaller when immature; SEM reveals that the
labyrinthine ridges are supported by spines or rods and the
surface of the plate-like structures is finely pitted or scored;
occasional irregular patches of reddish-brown amorphous
tissue on many mature spores; spore walls 1.5–2mm thick, in
KOH in youth hyaline singly and in mass, then dark green, by
maturity reddish-brown to brown.
Etymology
In reference to its occurrence in Queensland, Australia.
Distribution, habitat and season
Northern Queensland; hypogeous under various combinations
of Acacia flavescens,Allocasuarina littoralis,A. torulosa,
Corymbia citriodora,C. clarksoniana,Eucalyptus grandis,
E. phaeotricha or Syncarpia hillii; September through March.
Australian species of Elaphomyces Australian Systematic Botany 47

Fig. 9. Elaphomyces queenslandicus.(a) Cell structure of the outer peridial layer. (b) Ascus with developing spores. (c,d) Spores viewed under
light microscopy. (e) Spores in ascus, a scanning electron micrograph (SEM). ( f) SEM of a spore surface, showing the etchings of the labyrinth.
Scale bars = 20 mm(a), 10 mm(b), 25 mm(c,d), 10 mm(e) and 5 mm(f).
48 Australian Systematic Botany M. A. Castellano et al.

Material examined
QUEENSLAND: type locality, 30.i.1997, K. Vernes T22577 (OSC);
5.iii.1994, D. Arora & M. Castellano T13453 (OSC); 5.iii.1994,
M. Castellano T13448 (OSC); 9.xii.1996, K. Vernes T22525 (OSC);
21.xi.1996, K. Vernes T22529 (OSC); 27.ix.1996, K. Vernes T22585
(OSC); 27.ix.1996, K. Vernes T22588 (OSC); near Paluma, 17.ii.1992,
R. Halling H5663 (OSC).
Discussion
The black, carbonaceous peridium with dark brown enclosing
mycelium of E. queenslandicus resembles E. cooloolanus,
E. coralloideus and E. suejoyceae in appearance. The spores
of E. cooloolanus (11–13 mm, mean = 12.0 mm) and
E. coralloideus (18–20 mm, mean = 19.1 mm) are significantly
smaller. Both E. cooloolanus and E. coralloideus spores differ
distinctly in ornamentation from E. queenslandicus. The spores of
E. suejoyceae ((24–)26–30(–31) mm, mean = 27.6 mm) are in size
similar to those of E. queenslandicus ((23–)24–25(–26) mm,
mean = 24.5 mm); however, the ornamentation is distinctly
different.
In a surface view under a light microscope, the spores
of E. queenslandicus appear to have shallow ornamentation.
SEM and cross-sectional viewing under light microscopy
reveal the spines or rods beneath the rounded pitted
plate-like structures and a total height to the ornamentation of
3mm.
Elaphomyces rugosisporus Castellano, Trappe & Vernes,
sp. nov. (Figs 10,15a)
Ascomata in mycelio flammeo, miniato vel puniceo omnino
inclusa. Peridium strati trium compositum: stratum externum
60 mm crassum, nigrum, carbonaceum, laeve; stratum
medium 170–180(–500) mm crassum; strato interiore „1mm
crasso aliquantum transienti, uterque albo vel albido, saepe
areo carneo, rubro vel magenteo. Gleba in maturitate sporis
pulvereis atrocaesiis vel atramentariis repleta. Sporae globosae,
(10–)11–12 mm latae ornamentum reticulo tenui, punctato vel
rugoso inclusae, in maturitate in KOH pallide olivaceae.
Holotypus: Australia, Queensland, Atherton Tablelands,
Russell Pocket Road, 7.1 km from Gilles Highway, 8.iii.1995,
M. Castellano & J. Trappe T15300 (CANB, isotype OSC).
Ascomata 6–20 8–28 mm, ellipsoid to subglobose,
completely enclosed in deep orange to reddish-orange to red
mycelium, ectomycorrhizae and soil that form a crust 3–10 mm
thick or thicker. Peridium 3-layered; surface black, carbonaceous,
smooth, brittle; inner layers somewhat merged, white to off-
white, often developing erratic areas of pink to deep red to
magenta, especially on well matured specimens; these layers
are shrinking significantly on drying. Gleba initially stuffed with
white cottony mycelium, becoming filled with dark greyish-blue
to greyish-black to bluish-black spore powder, the adjacent
peridial tissue pink. Odour mild or indistinct. Taste not
determined.
Peridium in youth 2–5 mm thick, narrowing to as little as
1–3 mm by maturity, in KOH with a distinct rose blush to
the extreme outer surface of the carbonaceous layer, with outer
layer 60 mm thick, of dense, dark brown, grading into pale
brown textura epidermoidea that grades into hyaline, more or less
periclinal hyphae 170–180 mm thick, occasionally up to 500 mm
thick; middle (transitional) layer of perpendicularly stacked
bundles of hyaline hyphae 5 mm in diameter, inner layer
1.5 mm thick, of pale rose-tinted, inflated cells, 15 mmin
diameter. Gleba composed of spores and thin-walled, hyaline,
highly branched (90-degree angle), septate, loosely interwoven
hyphae 2mm in diameter. Asci globose and with a short tapered
base, 22–25 mm in diameter, the walls 2mm thick when young
and thinning down to 0.5 mm thick at maturity, hyaline,
8-spored; tapered base hyaline, 5mm broad; asci enlarging
as spores develop and the walls thinning by maturity, arising
from knots or clusters of pale green, contorted, ascogenous
hyphae 5mm in diameter scattered throughout the gleba.
Spores globose, (10–)11–12 mm (mean = 11.1 mm) including
ornamentation of a fine reticulum 2mm tall that appears
somewhat rugose to punctate under light microscopy; SEM
reveals a fine lattice work; spore walls 1mm thick, in KOH
pale greenish-brown singly and in mass.
Etymology
Latin, rugose (wrinkled) + sporus (spore), in reference to the
wrinkled appearance of the spore surface.
Distribution, habitat and season
Northern Queensland, New South Wales and Tasmania;
hypogeous under various combinations of Acacia flavescens,
A. falciformis,A. longifolia,A. mearnsii,Allocasuarina
littoralis,Corymbia citriodora,C. clarksoniana,
C. gummifera,Eucalyptus cypellocarpa,E. dalrympleana,
E. globulus,E. grandis,E. obliqua,E. phaeotrichaor
E. sieberi; November through June.
Material examined
NEW SOUTH WALES: Tantawangalo NP, New Line Rd, 0.7 km N of
the junction with Postman’s Track, 1.vi.1999, S. Cork AWC2953 (DAR,
OSC); QUEENSLAND: Atherton Tablelands, Davies Ck NP, Davies Ck
Rd, 18.iv.1996, K Vernes, T22523 (CANB, OSC); 23.v.1996, K. Vernes,
T22522 (CANB, OSC); 12.vi.1996, K. Vernes, T22546 (CANB, OSC);
7.xii.1996, K. Vernes, T22548 (CANB, OSC); 4.ii.1997, K. Vernes,
T22566 (CANB, OSC); 16.iv.1996, K. Vernes, T22564 (CANB, OSC);
5.ii.1992, M. Castellano & R. Young H5534 (PERTH, OSC); 6.iii.1994,
M. Castellano, T13485 & T13486 (CANB, OSC); Kirrama, 9.v.1991,
N. Malajczuk & M. Castellano H4736 (PERTH, OSC); Kenilworth,
12.v.1988, R. Young H4165 (PERTH, OSC); near Atherton, Rifle
Range Rd, 7.v.1991, P. Reddell & M. Castellano H4671 (PERTH,
OSC); Paluma, 9.iii.1990, P. Reddell NQ7 (CANB, OSC); SOUTH
AUSTRALIA: Kuitpo Forest, 25.xi.1971, J. Warcup (MEL 2024731);
TASMANIA: S of Geeveston, Huon Valley, Hermons Rd, 7.v.1990,
J. Trappe & N. Malajczuk H1563 (HO, PERTH, OSC); VICTORIA:
Croajingalong NP, Genoa Park Rd, 3.3 km N of the junction
with Hard-to-Seek Track, 29.v.2001, A. Jumpponen AWC4275 (MEL,
OSC).
Discussion
Elaphomyces rugosisporus resembles E. auranticus in having a
black, carbonaceous peridium with orange-coloured mycelium
enclosing the ascoma. See the discussion under E. auranticus
for differences.
Australian species of Elaphomyces Australian Systematic Botany 49

Fig. 10. Elaphomyces rugosisporus.(a) Interface between outer and middle peridial layers. (b) Bundled hyphae of the middle peridial layer. (c) Hyphal
cluster, giving rise to asci. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of spores. ( f) SEM of a spore surface, showing
the complex lattice structure of the ornamentation. Scale bars = 25 mm(a,b), 10 mm(c,d), 5 mm(e) and 2 mm(f).
50 Australian Systematic Botany M. A. Castellano et al.

Fig. 11. Elaphomyces suejoyceae.(a) Interface between outer and inner peridial layers. (b) Inner peridial layer, showing the scattered,
small crystals. (c) Asci with developing spores. (d) Spores viewed under light microscopy. (e) Scanning electron micrograph (SEM) of a spore.
(f) SEM of spore ornamentation, showing the complex construction of anastomosed rods. Scale bars = 30 mm(a,b), 20 mm(c,d), 5 mm(e) and
2mm(f).
Australian species of Elaphomyces Australian Systematic Botany 51

Elaphomyces suejoyceae Castellano, Trappe & Vernes,
sp. nov. (Figs 11,15b)
Ascomata in mycelio albo, brunneolo vel fusco omnino inclusa.
Peridium strati duorum compositum: stratum externum
170–180 mm crassum, fuligineum vel nigrum, carbonaceum,
laeve; stratum intimum 1–4 mm crassum, album vel prope
glebam brunneum. Gleba in maturitate sporis pulvereis
brunneonigris, fuscis, atroporphyreis vel atroferrugineis
repleta. Sporae globosae, (24–)26–30(–31) mm latae
ornamentum 2–4(–5) mm altum cristis labyrinthinis inclusae, in
maturitate in KOH porphyreae.
Holotypus: Australia, Queensland, Atherton Tablelands,
Davies Creek National Park, Davies Creek Road, 3.v.1988,
M. Trappe & N. Malajczuk H4031 (CANB, isotypes PERTH,
OSC).
Ascomata 12–35 23–50 mm, subglobose, with scant to
abundant, white or near white to pale brown to dark brown
mycelium enclosing the ascoma. Peridium 2-layered; surface
brownish-black to black, carbonaceous, smooth; inner layer
white and grading to brown near gleba, then with brown areas
marbled pale brown and dark brown clear in some places,
obscure in others, fleshy leathery. Gleba at maturity powdery
or sometimes wet, brownish-black to rich blackish-brown or dark
reddish-brown or even rusty dark brown. Odour mild or
indistinct. Taste not determined.
Outer layer of the peridium 170–180 mm thick, of dark brown
to reddish-brown textura epidermoidea, hyphae 4–5mmin
diameter, with walls 0.5 mm thick: inner layer 1–4 mm thick,
of thin-walled, hyaline to pale tan (in patches), closely interwoven
hyphae, some patches pale brown with small brown crystals
scattered across the layer; hyphae with walls up to 3 mm
thick near the gleba and inflated up to 10 mm in diameter,
thin-walled hyphal layer thin and only adjacent to dark-
coloured outer hyphae. Gleba composed of spores and thin-
walled, hyaline, slightly encrusted, branched, septate, loosely
interwoven hyphae, 3–4mm in diameter. Asci globose with a
short tapered base, 50 mm in diameter, with walls 2–4mm
thick, hyaline, 8-spored; tapered base hyaline, 4 mm wide by
3mm long, arising from knots or clusters of pale green
(in KOH), contorted, ascogenous hyphae 5mm in diameter
scattered throughout the gleba. Spores globose, (24–)26–30(–31) mm
(mean = 27.6 mm) including ornamentation of labyrinthine
ridges 2–4(–5) mm tall, ridges irregular in thickness under light
microscopy; SEM reveals the ridges to be composed of
anastomosed spines or rods, sometimes in ranks of 2 or 3 or
more; walls 1–2mm thick, in KOH initially hyaline, soon
golden reddish-brown to reddish-brown singly and in mass,
typically when mature the ridges are dark against the paler
valleys between them.
Etymology
In honour of Ms Sue Joyce, an accomplished collector of
sequestrate fungi in Queensland and a gracious hostess to
numerous mycological expeditions.
Distribution, habitat and season
Northern Queensland and New South Wales; hypogeous under
various combinations of Acacia flavescens,Allocasuarina
littoralis,A. torulosa,Corymbia citriodora,C. clarksoniana,
C. intermedia,Eucalyptus grandis,E. phaeotricha,
E. pilularis,E. propinqua,E. tereticornis or Syncarpia hillii;
November through July.
Material examined
QUEENSLAND: Atherton Tablelands, Davies Ck NP, Davies Ck Rd,
5.iii.1994, M. Castellano, T13437, T13446, T13455, T13456 (all CANB
& OSC); 18.ii.1996, K. Vernes, T22524 (CANB & OSC); 1.ii.1997,
T22528 (CANB & OSC); 27.ix.1996, T22530 (CANB & OSC);
7.xii.1995, T22572 (CANB & OSC); 30.i.1997, T22574 (CANB &
OSC); 31.i.1997, T22581 (CANB & OSC); 19.xi.1996, T22582
(CANB & OSC); 9.xii.1996, T22583 (CANB & OSC); 1.ii.1997,
T22584 (CANB & OSC); 27.vii.1996, T22586 (CANB & OSC);
5.iv.1989, N. Malajczuk H4379 (PERTH & OSC); near Atherton,
Rifle Range Rd, 3.iv.1989, M. Castellano H4321 (PERTH & OSC);
Mt Glorius, 16.v.1988, M. Amaranthus H4268 (PERTH & OSC); Mothar
Mountain, 9.v.1988, R. Young H4086 (PERTH & OSC); Tully Falls NP,
Red Rd, 4.v.1988, J. Trappe H4047 (CANB & OSC); near Paluma,
26.iv.1990, N. Malajczuk H4524 (PERTH & OSC); 9 km W of Mt
Glorius, 4.v.1992, R. Schaefer H5990 (PERTH & OSC); Mt Windsor
Tablelands, 2.ii.1992, M. Castellano H5460 (PERTH & OSC);
VICTORIA: East Gippsland, Kuark Forest Management Block,
Fred’s Track, 0.7 km NW of junction with Little Arte Trail,
28.vii.1996, A. Claridge, T19845 (MEL & OSC); 0.3 km off Omeo
Hwy, 2.1 km S of junction with Ash Range Rd, 11.xi.1996, E. Cázares,
T20145 (MEL & OSC).
Discussion
The black, carbonaceous peridium and the brown enclosing
mycelium of E. suejoyceae resembles that of
E. queenslandicus,E. cooloolanus and E. coralloideus. The
spores of E. cooloolanus (11–13 mm, mean = 12.0 mm) and
E. coralloideus (18–20 mm, mean = 19.1 mm) are significantly
smaller than and both have ornamentation that differs
distinctly from those of E. suejoyceae. The spores of
E. suejoyceae ((24–)26–30(–31) mm, mean = 27.6 mm) are
similar in size to those of E. queenslandicus ((23–)24–25(–26) mm,
mean = 24.5 mm); however, the ornamentation is distinctly
different.
Elaphomyces symeae Castellano, Trappe & Lebel, sp. nov.
(Figs 12,15c)
Ascomata in mycelio saturate mineato, saturate cinnamomeo
vel ferrugineo omnino inclusa. Peridium strati duorum
compositum: stratum externum 120–160 mm crassum, nigrum,
carbonaceum, irregulariter asprum; stratum intimum 1mm
crassum, album, interdum caerulescens. Gleba in maturitate
sporis pulvereis viridigriseis vel carboneis repleta. Sporae
globosae, 12–13(–14) mm latae ornamentum 1mm altum
reticulo pedicellato inclusae, in maturitate in KOH
auratobrunneae.
Holotypus: Australia, Western Australia, east of Albany,
Two Peoples Bay Nature Reserve, opposite the track to heath
site, 30.vi.1998, K. Syme KS1005/98 (MEL 2104387, isotype
OSC).
Ascomata 7–17 mm in diameter, globose, completely
enclosed in deep orangy-red, deep orangy-brown to rusty red
mycelium, ectomycorrhizal roots and sand that form a crust.
Peridium 2-layered; surface black, carbonaceous with some
incorporated sand granules, irregularly roughened, brittle;
inner layer white to off-white, sometimes bruising blue, firm.
52 Australian Systematic Botany M. A. Castellano et al.

Fig. 12. Elaphomyces symeae.(a) Ascoma in cross-section. (b) Hyphal structure of the inner peridial layer. (c) Cross-section view of
spores viewed under light microscopy. (d) Surface view of spores under light microscopy. (e) Scanning electron micrograph (SEM) of spores.
(f) SEM of spore ornamentation, showing the supporting rods to the honey-comb spore surface. Scale bars = 0.5 mm (a), 20 mm(b), 10 mm(c,d)
and 5 mm(e,f).
Australian species of Elaphomyces Australian Systematic Botany 53

Fig. 13. Elaphomyces timgroveii.(a) Ascoma in cross-section. (b) Hyphae of the inner peridial layer. (c) Cross-section view of spores under light
microscopy. (d) Surface view of spores under light microscopy. (e) Scanning electron micrograph (SEM) of spores. ( f) SEM of spores, showing the low
irregular reticulum. Scale bars = 0.5 mm (a), 15 mm(b), 10 mm(c,d), 5 mm(e) and 2 mm(f).
54 Australian Systematic Botany M. A. Castellano et al.

Fig. 14. Elaphomyces ascomata. (a)E. aurantius.(b)E. austrogranulatus.(c)E. chlorocarpus.(d)E. laetiluteus.(e)E. nothofagi.(f)E. queenslandicus.
Scale bars = 10 mm.
Australian species of Elaphomyces Australian Systematic Botany 55

Gleba initially hollow, then with white, cottony mycelium that
sometimes bruises pink or blue, at maturity filled with dark
greenish-grey to charcoal-coloured spore powder and scattered
thread-like hyphae. Odour indistinct. Taste not recorded.
Outer layer of the peridium 120–160 mm thick at maturity, of
compact, smooth, dark reddish-brown to nearly black,
agglutinated, short hyphae of 2–3mm in diameter and with
walls 1mm thick; inner layer 1 mm thick, somewhat off-
white to pale brown but hyaline as it nears gleba, of irregularly
interwoven (perpendicular at times) hyphal bundles composed of
hyaline, agglutinated, compact hyphae up to 6 mm in diameter
and with walls 2mm thick. Gleba composed of spores and
thin-walled, hyaline, septate, smooth, sinuous hyphae 2–5mm
in diameter. Asci globose, 20–22 mm in diameter, hyaline,
the walls 1mm thick, 8-spored; arising from knots of
hyaline, short-celled, contorted, hyaline, ascogenous hyphae
5–10 mm in diameter, with walls up to 2 mm thick. Spores
globose, 12–13(–14) mm broad (mean = 12.8 mm) including the
ornamentation of an irregular, pedicellate reticulum 1mm
tall that appears as a fine to coarse punctation or honey-comb
under light microscopy; SEM reveals an irregular honey-comb
surface that is supported by individual rods or spines; spore
walls 1mm thick, in KOH golden yellowish-brown singly
and dark golden yellowish-brown in mass when mature;
spores appearing nearly uniform in size and ornamentation at
maturity.
Etymology
In honour of Ms Karen Syme, accomplished mycological artist
and collector of sequestrate fungi in Western Australia.
Distribution, habitat and season
Western Australia; hypogeous, usually in deep litter under
various combinations of Allocasuarina fraseriana,Eucalyptus
conferruminata,E. marginata,E. megacarpa or Gastrolobium
bilobum; June, August or October.
Materials examined
WESTERN AUSTRALIA: E of Albany, Two Peoples Bay Nature
Reserve, west of Firebreak Valley Track, 21.x.1998, K. Syme &
G. Evans KS1029/98 (MEL 2104377); 22.x.1998, KS1031/98 (MEL
2105010); Bold Island Nature Reserve, 15.viii.2007, L. Bell, T. Friend &
N. Bougher 387 (PERTH).
Fig. 15. Elaphomyces ascomata. (a)E. rugosisporus.(b)E. suejoyceae.(c)E. symeae.(d)E. timgroveii. Scale bars = 10 mm (a,c,d) and 20 mm (b).
56 Australian Systematic Botany M. A. Castellano et al.

Discussion
Elaphomyces symeae resembles E. rugosisporus with its small
spores and orange–red mycelium enclosing the ascoma.
E. symeae is known only from Western Australia and has
slightly larger spores than does E. rugosisporus which is
found in eastern Australia. E. timgroveii also occurs in
Western Australia and has black mycelium enclosing the
sporocarp surface and spore ornamentation constructed
differently from that of E. symeae, as revealed by SEM
(however, they may appear similar under light microscopy).
Elaphomyces symeae is sometimes found in pink-stained sand
that turns orange near the ascoma surface.
Elaphomyces timgroveii Castellano, Trappe & Vernes,
sp. nov. (Figs 13,15d)
Ascomata in mycelio atro omnino inclusa. Peridium strati duorum
compositum: stratum externum 35–50 mm crassum, fumosum,
laeve; stratum intimum 1 mm pallide olivaceum. Gleba in
maturitate sporis pulvereis pallide viridis vel nigris repleta.
Sporae globosae, 8–9mm latae ornamentum 1mm altum
reticulo pedicellato, irregulari, tenui inclusae, in maturitate in
KOH olivaceae.
Holotypus: Australia, Western Australia, no locality, xi.1978,
T5550 (PERTH, isotype OSC).
Ascomata 8–20 12–23 mm, subglobose, completely
enclosed in black mycelium, ectomycorrhizal roots and sand
that form a thick crust. Peridium 2-layered; surface greyish-
brown, smooth; inner layer pale olive, rubbery. Gleba of a
pale green to black spore powder when mature. Odour musty.
Taste not recorded.
Outer layer of the peridium 35–50 mm thick, of hyaline to
pale green, thin-walled, septate, compact, interwoven hyphae
2–3mm in diameter, with much amorphous material incorporated
throughout the layer; inner layer 1 mm thick, of pale yellowish-
green, coarse, granulate-encrusted, compact, interwoven hyphae
7–9mm in diameter and with walls 2mm thick. Gleba of
spores, no hyphae seen. Asci and ascogenous hyphae not
seen. Spores globose, 8–9mm (mean = 8.7 mm) including
ornamentation of shallow, short, meandering ridges in an
irregular, fine reticulum 1mm tall, that appears punctate
under light microscopy; SEM reveals the ornamentation to be
an irregular reticulum, with some patchy amorphous material
adhering at various spots on the spore; walls 1mm thick, in KOH
olive singly and in mass.
Etymology
In honour of Dr Tim Grove, esteemed colleague and
accomplished collector of sequestrate taxa in Australia.
Distribution, habitat and season
Western Australia; hypogeous under Eucalyptus spp.; July,
November or December.
Material examined
WESTERN AUSTRALIA: no locality, xi.1978, T5534 (PERTH &
OSC); Pemberton, vii.1982, N. Malajczuk (PERTH 00746169);
Manjimup, Pine Ck Rd, 1.xii.1981, N. Malajczuk (PERTH 07507992).
Discussion
The very small spores of E. timgroveii are similar in size to
those of E. cooloolanus,E. rugosisporus and E. symeae. The
spore ornamentation of E. cooloolanus is made up of close rods
or spines that form short ridges, the spore ornamentation of
E. rugosisporus is a tall (2mm) fine reticulum, the
ornamentation of E. timgroveii is also a fine reticulum but not
as tall in stature (1mm). The spore size of E. timgroveii (8–9mm,
mean = 8.7 mm) does not overlap with the that of the other three
species (E. cooloolanus spores 11–13 mm, mean = 12.0 mm;
E. rugosisporus spores (10–)11–12 mm, mean = 11.1 mm;
E. symeae spores 12–12(–14) mm, mean = 12.8 mm).
SEM confirms that an outer shell of fine ornamentation is
supported underneath by rods or spines.
Acknowledgements
This research was supported in part by Australian Biological Resources Study
(ABRS) Grant No. 95130, by Australian Centre for International Agricultural
Research Grant No. 8736, by National Science Foundation Grants DEB
9007186 and 9400545, and by CSIRO, Forestry/Forest Products and
Sustainable Ecosystems. We appreciate the review of this manuscript by
A. Claridge and D. Luoma. We appreciate the expert assistance in the field
by M. Amaranthus, D. Arora, N. Bougher, E. Cázares,A. Claridge, W. Colgan
III, S. Cork, A. Giachini, Dr T. Grove, R. Halling, S. Joyce, A. Jumpponen,
T. Lebel, N. Malajczuk, D. Mills, G. Mires, P. Reddell, R. Schaefer, K. Syme,
C. Theodorou and R. Young. We appreciate the contribution of some of the
SEM images by Jim Ehrman of Mount Allison University, Canada. We also
appreciate the loan of specimens from CANB, HO, MEL, OSC, PDD, and
PERTH.
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Australian species of Elaphomyces Australian Systematic Botany 57
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