New Elaphomyces species (Elaphomycetaceae, Eurotiales, Ascomycota) from Guyana

Abstract

Elaphomyces compleximurus sp. nov. and E. digitatus sp. nov. are described from the Pakaraima Mountains of Guyana. Macromorphological, micromorphological, habitat and DNA sequence data are provided for each new species. This is the first report of Elaphomyces ascomata associated with ectomycorrhizal members of the Fabaceae and also for the genus from the lowland South American tropics.

Full text

New
Elaphomyces
species (Elaphomycetaceae,
Eurotiales, Ascomycota) from Guyana
Michael A. Castellano
USDA, Forest Service, Northern Research Station,
3200 Jefferson Way, Corvallis, Oregon 97331
Terry W. Henkel
1
Department of Biological Sciences, Humboldt State
University, Arcata, California 95521
Steven L. Miller
Department of Botany, University of Wyoming,
Laramie, Wyoming 82071
Matthew E. Smith
Department of Plant Pathology, University of Florida,
Gainesville, Florida 32611
M. Catherine Aime
Department of Plant Pathology and Crop Physiology,
Louisiana State University Agricultural Center,
Baton Rouge, Louisiana 70803
Abstract
:
Elaphomyces compleximurus
sp. nov. and
E.
digitatus
sp. nov. are described from the Pakaraima
Mountains of Guyana. Macromorphological, micro-
morphological, habitat and DNA sequence data are
provided for each new species. This is the first report
of
Elaphomyces
ascomata associated with ectomycor-
rhizal members of the Fabaceae and also for the
genus from the lowland South American tropics.
Key words: Dicymbe
, Guiana Shield, hypogeous
fungi, sequestrate fungi
INTRODUCTION
The hypogeous false truffle genus
Elaphomyces
Fries
(Elaphomycetaceae, Eurotiales, Ascomycota) is com-
posed of ectomycorrhizal (ECM) fungi with a gleba of
powdery, dark ascospores, a thick peridium and large
cleistothecial ascomata relative to other members of
the Eurotiales. Most of the ,67 species of
Elaphomyces
currently described are from north temperate forests
in association with ECM host plants in the Pinaceae,
Fagaceae and Betulaceae (Castellano et al. 1989).
Cooke (1892) and Rodway (1918) provided the first
reports of
Elaphomyces
for the southern hemisphere,
but they assigned European names to their species
(Castellano et al. 2011). The known
Elaphomyces
diversity in the southern temperate zones was in-
creased substantially by Castellano et al. (2011) who
published 13 new species associated with Myrtaceae
and
Nothofagus
host plants in Australia. Three new
Elaphomyces
species associated with
Nothofagus
recently
were described from New Zealand (Castellano et al.
2012).
While
Elaphomyces
is well represented in the
subtropical
Quercus
forests of Florida and records
exist from montane
Quercus
forests of Costa Rica
(Castellano unpubl), records are exceedingly scarce
from the low-latitude, lowland forests of the tropics.
Corner and Hawker (1955) described the first
Elaphomyces
species from the lowland tropics in
association with Dipterocarpaceae in Singapore, but
to date no species have been described from the
lowland South American tropics.
Over the past 13 y macrofungal collecting expedi-
tions in western-central Guyana have revealed a
wealth of ECM fungi in forests dominated by the
ECM Fabaceae canopy tree species
Dicymbe corymbosa
Spruce ex Benth.,
Dicymbe altsonii
Sandw. (subfam.
Caesalpinioideae; Henkel et al. 2011) and
Aldina
insignis
(Benth.) Endl. (subfam. Papilionoideae;
Smith et al. 2011). Recent molecular studies of ECM
fungal communities in these forests have detected
belowground fungal diversity similar to those revealed
by long-term sporocarp surveys in the region, but the
great majority of taxa are basidiomycetes (Smith et al.
2011). So far, one member of the Elaphomycetaceae,
Pseudotulostoma volvata
O.K. Mill. & T.W. Henkel, has
been described from Guyana (Miller et al. 2001).
Here we describe two new
Elaphomyces
species
discovered in Guyana’s
Dicymbe
forests. Morphologi-
cal, habitat and DNA sequence data are provided for
each species. These are the first described species of
Elaphomyces
associated with ECM Fabaceae hosts and
the first records for the genus from the lowland South
American tropics.
MATERIALS AND METHODS
Ascomata were collected during the May–July rainy seasons
of 2000–2002, 2004, 2007 and 2009–2010 from the Upper
Potaro River Basin within a 15 km radius of a permanent
base camp at 5u18904.80N, 59u54940.40W, 710 m (Henkel
2003). More were collected during May–June 2011 from the
Upper Demerara River Basin at Mabura Ecological Reserve
within 2 km of a field station at 5u09919.00N; 58u41958.90W,
100 m. At Potaro ascomata were collected from mono-
dominant forests of
D. corymbosa
and other stands
containing
D. corymbosa
,
D. altsonii
and
A. insignis
;at
Submitted 23 Feb 2012; accepted for publication 30 Mar 2012.
1
Corresponding author. E-mail: twh5@humboldt.edu
Mycologia,
104(5), 2012, pp. 1244–1249. DOI: 10.3852/12-061
#2012 by The Mycological Society of America, Lawrence, KS 66044-8897
1244

Mabura collections were made in
D. altsonii
monodominant
stands. Fresh characteristics of ascomata were recorded in
the field. Color was compared with plates from Kornerup
and Wanscher (1978) and is cited in parenthesis (e.g. 5A4).
Specimens were field-dried with silica gel. Dried specimens
were rehydrated and examined in 3%KOH, Melzer’s
reagent and cotton blue. Microscopic descriptions are
based on 3%KOH mounts unless specified. Twenty
ascospores were measured from the holotype collection;
dimensions reported include ornamentation. Dried asco-
spores were mounted on aluminum pegs with double-sided
tape and coated with gold for scanning electron microscopy
(SEM) with an AmRay 3300 FE field emission scanning
electron microscope. Specimens are deposited in these
herbaria: (Holmgren et al. 1990): BRG, University of
Guyana; HSU, Humboldt State University; OSC, Oregon
State University; LSUM, Louisiana State University Mycolo-
gy. Ribosomal DNA sequencing for the ITS and LSU
regions followed the protocols of Aime et al. (2005) and
Smith et al. (2011); GenBank accession numbers are
provided in
Specimens examined
.
TAXONOMY
Elaphomyces compleximurus Castellano, T.W. Hen-
kel et S.L. Mill., sp. nov. FIG.1
MycoBank MB564542
Ascomata 9–15 mm tall 310–20 mm broad, nearly
globose to flattened subovate with a distinct, abruptly
tapered base, with roots, sand and dense mycelium
adherent to the base and upward over lower half of the
ascoma, black when fresh, drying to dark gray (near 10F2–
10F3); peridial surface completely covered with dense black
mycelium visible with a hand lens, beneath this mycelium
verrucose throughout except nearly smooth near base; warts
contiguous, separated by narrow lines, angular-sided, up to
1 mm tall and 0.3 mm broad, with rounded apices.
Peridium when dried collapsing to form an exterior,
subreticulate pattern of ridges and folds, these irregular in
length, superimposed over surface warts; in section four-
layered; outer first layer 0.03–0.06 mm thick, dark brown, of
closely appressed mycelium, underlain by a second carbo-
naceous layer 0.1–0.15 mm thick, of nearly black, closely
appressed cells, underlain by a third layer 0.5–0.6 mm thick,
mostly white but a thin, pale tan outer zone, cottony, with
embedded, black-mantled ectomycorrhizas scattered
throughout but most numerous near the ascoma base; the
innermost fourth layer 1.3–2 mm thick, pale gray to pale
brownish gray (6F2–6F3), gradually more pale toward the
gleba, similar in texture to the third layer but lacking
embedded ectomycorrhizas. Gleba grayish black (9F2–9F3)
to black, powdery, with fine, pale dissepiments. Odor mildly
of raw potato with fruity element; taste none. Outer first
layer of peridium composed of thick-walled, brown to dark
brown, septate, segmented and irregular (not straight-
walled) hyphae 1–2 mm broad, these somewhat tuberculate,
loosely interwoven and much branched at nearly 90 degree
angles; carbonaceous second layer dense, of compact, dark
brown to nearly black cells with walls ,2mm thick, quickly
grading downward into the third layer of brown, somewhat
puzzle-like textura epidermoidea, grading into the fourth
layer where the pale brown tissue changes quickly to textura
intricata or textura porrecta forming clusters in cross-
hatched formations, this pattern continuing inward with the
hyphae grading from brown to hyaline to brown near the
gleba; hyphae of fourth layer 5–7 mm wide with walls 1–2 mm
thick. Gleba of ascospores and sinuous, hyaline, septate,
acutely branched, loosely interwoven hyphae, these 2–3 mm
wide, with walls ,0.5 mm thick. Asci not observed.
Ascospores globose, brown, 22–25 mm diam (mean 5
23.2 mm) including the reticulate-alveolate ornamentation;
aveolae well defined, 3.5–5.5 mm broad 32–3 mm tall, nearly
round or with subangular sides with irregular to wavy walls;
under SEM the individual alveolar wall consisting of two
layers separated by vertical ribs; these nearly equally spaced
along the wall, with rib ends at the margin of the wall; the
ascospore surface exposed inside the alveolae an irregular,
slightly roughened, central disk with radiating repent rods
and small depressions extending to the base of the
surrounding alveolar wall.
Holotype. Henkel 8880
(BRG; ISOTYPES HSU;
OSC)
Habit, habitat and distribution.
Scattered to gregar-
ious on the forest floor under
D. corymbosa
or
D.
altsonii
on white sand or brown sand soils; May–July
rainy season and also in late August during dry season
transition; known from the type locality in the Upper
Potaro River Basin of Guyana and ,100 km east in the
Mabura Hill region.
Etymology.
Complexus (Latin adj. B) 5complex +
murus (Latin s.m. II) 5wall, referring to the distinctive,
complex structure of the ascospore ornamentation.
Specimens examined
. GUYANA. REGION 8 POTARO-
SIPARUNI: Pakaraima Mountains, Upper Potaro River
Basin, within a 4 km radius of base camp at 5u18904.80N,
59u54940.40W, 710–750 m; vicinity of base camp, 4 Jun 2000,
S.L. Miller 10096
(BRG; HSU; OSC) and 23 Jun 2000,
S.L.
Miller 10157
(BRG; HSU; OSC); ,0.75 km northeast of
base camp on white sand soils in mixed
D. corymbosa-
Micrandra glabra
forest, 25 Aug 2007,
Henkel 8880
(HOLOTYPE BRG; ISOTYPES HSU, OSC; ITS GenBank
JN711441); 1.5 km west of base camp in mixed
D. corymbosa
-
M. glabra
forest on alluvial sand soils, 3 Jun 2010,
Henkel
9254
(BRG, HSU); REGION 10 UPPER DEMERARA-
BERBICE: Mabura Ecological Reserve, field station located
at 5u09919.00N, 58u41958.90W, ,100 m; vicinity of field
station, under
D. altsonii
on brown sand soils, 28 May 2011,
Aime 4331
(BRG; LSUM).
Commentary.
The distinctly reticulate-alveolate asco-
spores of
E. compleximurus
easily distinguish it from
nearly all other described
Elaphomyces
species. The
only known
Elaphomyces
species to combine the
features of a tapered base of the ascoma, black,
verrucose peridium and some degree of reticulate
ascospore ornamentation are the European
E. cyano-
sporus
Tul. & C. Tul. and
E. persoonii
Vittad. Both are
easily separated from
E. compleximurus
by their larger
CASTELLANO ET AL.: NEW SPECIES OF
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LAPHOMYCES
1245

FIG.1.
Elaphomyces compleximurus
(HOLOTYPE;
Henkel 8880
). a. Ascomata in clusters and in longitudinal section. b.
Longitudinal section of ascoma showing the powdery gleba (gl) and the embedded ectomycorrhizas (ec) within the peridium. c.
Peridial layer 2 with carbonaceous cells (ep). d. Mature ascospores under bright field microscopy showing ornamentation in outline
and surface view of aveolae. e. Ascospores under scanning electron microscopy (SEM) showing the irregularly sided aveolae. f. Aveolae
under SEM showing complex ornamentation of ribs between alveolar wall layers. Bars: a 515 mm; b 52mm;c,d,e510 mm; f 51mm.
1246 MYCOLOGIA

ascospores; in
E. cyanosporus
the ascospore diameter
ranges from 27–30 mm (mean 528 mm), for
E.
persoonii
29–33 mm(mean531.1 mm), vs. 22–25 mm
(mean 523.2 mm) for
E. compleximurus
.Both
E.
compleximurus
and
E. persoonii
have ectomycorrhizas
embedded within the outer layers of the peridium, a
condition otherwise highly unusual in sequestrate
fungi.
This is the first report of
Elaphomyces
ascomata
associated with fabaceous hosts. However, an ITS
rDNA sequence of another
Elaphomyces
species
(ECM1108, GenBank JN168718) as yet undiscovered
as ascomata recently was reported from ectomycor-
rhizas of
D. corymbosa
(Smith et al. 2011) and also
found on
D. jenmanii
(Smith and Henkel unpubl),
indicating that taxa beyond those described here
exist in the region. In Guyana
Elaphomyces complex-
imurus
has been encountered less frequently than
E.
digitatus
(described below), but the former is harder
to find in the field because of its dark color.
Pseudotulostoma volvata
, another sympatric taxon
from the Elaphomycetaceae, is common in
Dicymbe
-
dominated forests both as ascomata and on ECM
roots and also has the unusual peridium-embedded
ectomycorrhizas (Miller et al. 2001; Henkel et al.
2006, 2011).
Elaphomyces digitatus Castellano, T.W. Henkel et
S.L. Mill., sp. nov. FIG.2
MycoBank MB564543
Ascomata 4–11 mm tall 34–31 mm broad, nearly globose
to flattened or ellipsoidal, occasionally furrowed, when
immature cream (5A2–5A3) to light orange (5A5), light
yellow (4A5) or grayish orange (5B3), at maturity brownish
orange (5B3–5B8–5C8) to occasionally darker brown (6E8–
6F8), with fine surface tomentum throughout that is
occasionally absent at maturity; ectomycorrhizas with
adherent soil irregularly covering the lower one-quarter to
one-half of the ascoma, forming a loose exterior agglomer-
ation, not embedded within the peridial structure, occa-
sionally attached to more extensive fine roots extending
into soil; peridial surface beneath the tomentum finely
roughened macroscopically, under hand lens verrucose,
consisting of fine, contiguous concolorous warts except
where damaged; warts ,0.13 mm tall and 0.25 mm broad,
pyramidal with obtuse apices, with 4–6 sides of irregular
dimensions. Peridium when fresh leathery and tough, 0.9–
1.8 mm thick overall; in dried section 0.5–0.75 mm thick
overall, three-layered; outer layer of brownish orange
pyramidal, blunt warts up to 0.13 mm tall, underlain by a
dull creamy white to off-white layer 0.1–0.15 mm thick, this
grading into an innermost, gray to reddish gray layer 0.25–
0.3 mm thick. Gleba dark brown (7F8) to dark rusty brown
(8F8) when mature, powdery, with numerous, fine, white
dissepiments throughout. Odor musky or skunky or of latex
when sectioned; taste none. Outer layer of peridium of
small pyramidal warts up to 125 mm tall, red-brown at apex
and yellow-brown near base; cells short, pigmented, with
walls 1–2 mm thick; the pyramids intraspace 50–60 mm deep
of hyaline, more or less parallel, inflated cells that are 5–
10 mm broad; outer layer grading inward into a disorganized
textura epidermoidea of interwoven, hyaline hyphae 10–
12 mm wide with walls 2–4 mm thick, grading quickly inward
into a darker layer tinted yellow-brown from scattered dark
concolorous granules; hyphae immediately adjacent to the
gleba occasionally inflated up to 35 mm broad. Mature gleba
of ascospores and sinuous, interwoven, hyaline, septate
hyphae, these 2–3 mm broad with walls ,0.5 mm thick,
intermixed with inflated hyphae up to 6 mm broad and
encrusted with brown, amorphous deposits. Asci present in
immature gleba, globose, 42–65 mm diam, walls 1–2 mm
thick, containing 2–8 ascospores (mostly 8), with a
constricted base up to 12 37mm attached to a knot of
granulated, pale brown, thick-walled hyphae 67mm wide
with walls 2–3 mm thick. Ascospores globose, dark red-
brown, (16)20–23(24) mm diam (mean 521.9 mm) includ-
ing the irregular, labyrinthine-ridged ornamentation that
forms a partial reticulum of dark, crowded ridges 3–4 mm
tall; under SEM reticulum walls wavy, fully fused near
margins but separating downward into columnar basal
elements near ascospore surface; margins with numerous
short, irregular, digitate projections.
Holotype. Henkel 8887
(BRG; ISOTYPES HSU;
OSC)
Habit, habitat and distribution.
Usually in clusters of
2–5 ascomata in hilly country on red lateritic soils at
,710 m in
D. corymbosa
monodominant forest, also in
similar terrain in mixed
D. altsonii
,
D. corymbosa
and
A. insignis
forest, infrequently in sandy, alluvial soil
partially embedded in humic material on forest floor
under
D. corymbosa
, rarely elevated 1–2 m up the bole
of a
D. altsonii
in mixed
Dicymbe
forest, or in
D.
altsonii
-dominated stands at low elevation on brown
sand soils; May–July and December–January rainy
seasons; known from the type locality in the Upper
Potaro River Basin of Guyana and ,100 km east in the
Mabura Hill region.
Etymology.
Digitatus (Latin adj. A) 5digitate,
referring to the small, finger-like projections emanat-
ing from the outer ascospore ornamentation.
Specimens examined
. GUYANA. REGION 8 POTARO-
SIPARUNI: Pakaraima Mountains, Upper Potaro River
Basin, within a 15 km radius of Potaro base camp at
5u18904.80N, 59u54940.40W, 710–750 m; old Ayanganna
airstrip, 21 May 2000,
S.L. Miller 10041
(BRG; HSU;
OSC) and 21 May 2000,
Aime 995
(BRG; LSUM); ,200 m
southwest of base camp at
D. altsonii
site, 22 May 2000,
Henkel 7448
(BRG; HSU; OSC) and
S.L. Miller 10053
(BRG; HSU; OSC);
Dicymbe
plot 1, 28 May 2000,
Aime 1047
(BRG; OSC; LSUM) and 14 May 2010,
Aime 3914
(BRG;
LSUM); vicinity of base camp, 30 May 2000,
Aime 1079
(BRG; OSC; LSUM) and 5 Jul 2004,
Henkel 8744
(BRG;
HSU; OSC) and 26 Aug 2007,
Henkel 8887
(HOLOTYPE
BRG; ISOTYPE HSU, OSC; ITS GenBank JQ657705);
Benny’s ridge under
D. corymbosa
on lateritic ironstone
CASTELLANO ET AL.: NEW SPECIES OF
E
LAPHOMYCES
1247

FIG.2.
Elaphomyces digitatus
(HOLOTYPE;
Henkel 8887
). a. Fresh ascomata including transverse section. b. Outermost
peridium layer showing stacked hyaline cells in of space between warts. c. Interior peridium layer with scattered dark granules (gr)
interspersed with hyaline hyphae. d. Mature ascospores under bright field microscopy with ornamentation in outline. e. Ascospores
under scanning electron microscopy (SEM) with the labyrinthine-ridged ornamentation. f. Individual ascospore under SEM with
the digitate protrusions on the wall margins of ornamentation. Bars: a 510 mm; b 550 mm; c 520 mm; d, e 510 mm; f 55mm.
1248 MYCOLOGIA

soils, 1 Jun 2000,
S.L. Miller 10088
(BRG; HSU; OSC) and 4
Jun 2000,
S.L. Miller 10105
(BRG; HSU; OSC) and 10 Jun
2000,
S.L. Miller 10124
(BRG; HSU; OSC);
Dicymbe
plot 3,
18 May 2001,
Aime 1512
(BRG; OSC; LSUM; LSU GenBank
JN713147) and 8 Jun 2002,
Aime 1923
(BRG; OSC; LSUM;
LSU GenBank JN713148); near Tadang base camp in mixed
Dicymbe–Aldina
forest on lateritic soils, 27 Dec 2009,
Henkel
9161
(BRG; HSU) and attached directly to roots of
A.
insignis
, 29 Dec 2009,
Henkel 9166
(BRG; HSU); REGION
10 UPPER DEMERARA-BERBICE: Mabura Ecological Re-
serve, field station at 5u09919.00N, 58u41958.90W, ,100 m;
vicinity of field station, under
D. altsonii
on brown sand
soils, 24 May 2011,
Aime 4284
(BRG; LSUM).
Commentary.
Theorangeperidiumof
E. digitatus
is
distinctive in the genus, and this along with the sub-
epigeous presentation of ascomata in clusters makes
them relatively easy to find in the field. The European
species
E. aculeatus
Vittad.,
E. anthracinus
Vittad.,
E.
echinatus
Vittad.,
E. leveillei
Tul. & C. Tul. and
E.
leucosporus
Vittad. have ascospores of similar size as
E.
digitatus
, but each has a black carbonaceous peridium
and a different pattern of ascospore ornamentation.
Elaphomyces cyanosporus
and
E. persoonii
have similarly
reticulate ascospore ornamentations with short, finger-
like projections from the wall margin, but both have
significantly larger ascospores than
E. digitatus
(28.0 mm
and 31.3 mm vs. 21.9 mmmeandiamrespectively).
Elaphomyces austrogranulatus
Castellano, Trappe &
Vernes from Australia has a brown peridium and
relatively small ascospores like
E. digitatus
but its
ascospore ornamentation is irregularly verrucose (Cas-
tellano et al. 2011). Other Australian
Elaphomyces
species
with ascospores of similar size have a black peridium.
Elaphomyces digitatus
was found fairly frequently in
long-term
D. corymbosa
macrofungal study plots in the
Upper Potaro Basin, with ascomata recorded in 16.3%
of 630, 100 m
2
sampling quadrats over 7 y (Henkel et al.
2011). The congener
E. compleximurus
(described
above) was never found in the study plots but recovered
sporadically in ‘‘off-plot’’ collecting forays. In addition
to
Dicymbe
spp.,
E. digitatus
appears to form mycorrhi-
zae with the papilionoid leguminous tree
A. insignis
.In
one collection of
E. digitatus
(
Henkel 9166
) the
ascomata were found directly attached to ECM roots
of this confirmed ECM host (Smith et al. 2011).
ACKNOWLEDGMENTS
This research was made possible by grants to TWH from the
National Science Foundation grant DEB-0918591, National
Geographic Society’s Committee for Research and Explora-
tion, Smithsonian Institution’s Biological Diversity of the
Guiana Shield Program, Linnaean Society of London, Hum-
boldt State University Foundation and to MCA from the
Explorer’s Club, Washington Group and NSF DEB-0732968.
Additional support was provided by a grant to SLM from NSF
DEB-1050292. Dillon Husbands functioned as Guyanese field
counterpart for several of the expeditions and assisted with
collection and description of specimens. Additional field
assistance in Guyana was provided by Mimi Chin, Christopher
Andrew, Leonard Williams, Valentino Joseph, Francino Ed-
mond and Luciano Edmond. Research permits were granted
by the Guyana Environmental Protection Agency. This paper is
No. 182 in the Smithsonian Institution’s Biological Diversity of
the Guiana Shield Program publication series.
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