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667
Mycologia,
96(3), 2004, pp. 667–674.
q2004 by The Mycological Society of America, Lawrence, KS 66044-8897
Acanthostigma
and
Tubeufia
species, including
T. claspisphaeria
sp. nov., from
submerged wood in Hong Kong
Rampai Kodsueb
Saisamorn Lumyong
1
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai, Thailand
Pipob Lumyong
Department of Plant Pathology, Faculty of Agriculture,
Chiang Mai University, Chiang Mai, Thailand
Eric H.C. McKenzie
Landcare Research, Private Bag 92170, Aukland,
New Zealand
Wai Hong Ho
Kevin D. Hyde
Centre for Research in Fungal Diversity, Department of
Ecology & Biodiversity, The University of Hong Kong,
Pokfulam Road, Hong Kong
Abstract: Acanthostigma scopulum, Tubeufia claspis-
phaeria
sp. nov. and
T. paludosa
were identified from
submerged wood collected in a small forest stream
on Lantau Island, Hong Kong. The collections of
Acanthostigma scopulum
and
Tubeufia paludosa
dif-
fered slightly from the original descriptions.
Tubeufia
claspisphaeria
differs from previously described spe-
cies in that it has hook-shaped setae that form radi-
ally around the ostiole. This new species is described
and illustrated and compared with the most similar
species. A dichotomous key to the 16 accepted spe-
cies in
Tubeufia
is provided.
Key words:
new species, saprobic fungi, systemat-
ics
INTRODUCTION
There are 22 genera in the Tubeufiaceae (Barr 1980,
Rossman 1987, Kirk et al 2001). Some genera, such
as
Melioliphila
and
Uredinophila,
are hyperparasites
on sooty molds and on rust fungi, while
Podonectria
is parasitic fungi on scale insects.
Acanthostigma
and
Tubeufia
are saprobes, usually found on old, rotten
wood. Previous examinations of freshwater fungi oc-
curring on submerged wood in streams in the tropics
and subtropics have yielded numerous novel fungi,
Accepted for publication October 20, 2003.
1
Corresponding author. E-mail: scboi009@chiangmai.ac.th
including hyphomycetes (Hyde et al 2002, McKenzie
et al 2002), coelomycetes (Hyde 1993) and ascomy-
cetes (Wong and Hyde 1999, Cai et al 2002). During
our survey of fungi occurring on naturally sub-
merged wood, we collected an
Acanthostigma
and two
Tubeufia
species, one being a species new to science,
from a small forest stream in Hong Kong. The three
species are described, illustrated and compared with
known taxa. A key to the 16 accepted species of
Tu-
beufia
is provided.
MATERIALS AND METHODS
Submerged wood was retrieved from a stream near the
Trappist Monastery, Lantau Island, Hong Kong, in Apr 2002
and returned to the laboratory, where it was incubated in
zip-lock plastic bags at room temperature. Humidity was
maintained by adding moistened paper towels. Samples
were examined within 3 d and periodically over 1 mo for
the presence of sporulating structures. Cultures of fungi
were obtained where possible from single spores (Choi et
al 1999). To stimulate sporulation, 1 cm
2
blocks of colo-
nized agar were placed in water in a bubble chamber over-
night.
TAXONOMY
Acanthostigma
De Not. and
Tubeufia
Penz. & Sacc.
Acanthostigma
currently includes six species and re-
cently has been reviewed by Re´blova´ and Barr (2000).
The genus is characterized by vinaceous, reddish-
brown or dark brown ascomata that are covered with
dark brownish-black, often opaque, obtuse or acute
setae. The ascospores are hyaline, multiseptate and
cylindrical-fusiform to elongate fusiform. The ana-
morphs are in
Helicosporium
and
Helicomyces.
In spe-
cies of
Tubeufia
the ascomata are hyaline, whitish or
yellowish to pinkish, but may become dark at matu-
rity. They are smooth, or are covered with protruding
cells, thick-walled hyphal appendages, or short dark
setae (Re´blova´ and Barr 2000).
Acanthostigma scopulum (Cooke & Peck) Peck, Bull.
New York State Mus. 1:22 (1887) (F
IGS
. 1–6)
Ascomata 145–250 mm diam, superficial, globose, solitary
or gregarious, dark brown to black, membranous, ostiolate,
with setae; setae (56–)80–90(–99) mm long, 4–6 mm wide
at base, 1.5–2 mm wide at apex, dark brown, 1–2-septate,
thick-walled, straight. Asci (64–)90–120(–130) 38–10 mm
668 M
YCOLOGIA
F
IGS
. 1–6.
Acanthostigma scopulum
(from HKU[M] 17121). 1. Ascomata on natural substratum. 2. Dark brown, septate
setae with acute apex. 3–4. A cluster of mature and immature asci. 5. Ascus. 6. Ascospore. Pseudoparaphyses in F
IG
.3–4
(arrowed). Bars: 1 5200 mm; 2–5 530 mm; 6 525 mm. F
IGS
. 2–3 mounted in water, other figures mounted in lactophenol.
(
x¯
5103 39.2 mm, n 520), 8–spored, cylindric-clavate,
bitunicate, short stalked, pseudoparaphyses. Ascospores
(66–)70–85(–90) 33–3.5(– 4) mm(
x¯
578 33.4 mm, n 5
30), fasciculate, long-fusiform to cylindrical-fusiform, hya-
line, straight or slightly curved, (5–)9–12-septate, smooth-
walled, guttulate, lacking appendages and sheaths.
Anamorph. Helicosporium aureum
(Corda) Linder,
Ann. Missouri Bot. Gard. 16:279, 1929, from Re´blova´
and Barr (2000).
Colonies on potato-dextrose agar olive-colored,
reaching 1 cm diam in 10 d at room temperature
(
;
28 C), no pigment diffusing into agar, not sporu-
lating even after submergence overnight in a bubble
chamber.
Substratum.
Wood submerged in streams, decaying
wood (
Fagus sylvatica, Pinus
sp. and
Tsuga
sp.)
Known distribution.
Australia, Europe, Hong Kong
and U.S.A.
Specimen examined.
HONG KONG. Lantau Island, Trap-
pist Monastery, small stream in forest, on wood partially sub-
merged, 21 Apr 2002,
K.D. Hyde
(HKU[M] 17121; living
culture HKUCC 9117).
Notes.
A key to species of
Acanthostigma
was pro-
vided by Re´blova´ and Barr (2000). This collection
differs from the description for
A. scopulum
provided
by Re´blova´ and Barr (2000) in that it has septate
setae and wider ascospores 3–3.5(–4)
m
m versus
(2–)2.5–3(–3.5)
m
m but in other aspects it is similar.
Tubeufia claspisphaeria Kodsueb, sp. nov.
(F
IGS
. 7–16)
Ascomata 185–330(–350) mm diametro, superficialia, glo-
bosa, solitaria vel aggregata, hyalina vel pallide brunnea ubi
immatura, atrobunnea vel atris ubi maturus, membranea,
ostiolata, setosa; setae usque 47 mm longa, 10 mm lata,
(0–)1(–2)-septata, atrobrunnea, hamatus. Peries ascomati
usque 60 mm lata crassus, vinacea-brunnea, e cellulis pseu-
doparenchymatis compositum, 4–5 stratosus, textura glob-
ulosa. Asci (93–)105–120(–127) 3(12–)14–16(–18) mm, oc-
tospori, cylindrico-clavati, bitunicati, pedicellati. Ascosporae
(34–)45–60(–64) 35–7(–8) mm, fasciculatae, elongatae-fu-
siformis, (3–)5–6(–8)-septatae, hyalinae, curvatae, guttula-
tae.
Etymology.
In reference to the clasp-like setae on
the ascomata. Ascomata 185–330(–350)
m
m diam, su-
perficial, globose, solitar y or grouped, hyaline to pale
brown when immature, dark brown becoming black
when mature, membranous, ostiolate, with setae that
form around ostiole. Setae from surface of ascomata,
up to 47
m
m long and 10
m
m wide, (0–)1(–2)-septate,
dark brown, thick-walled, hook-shaped. Peridium up
to 60
m
m wide, vinaceous brown, comprising 4–5 lay-
669K
ODSUEB ET AL
:
A
CANTHOSTIGMA AND
T
UBEUFIA FROM
H
ONG
K
ONG
F
IGS
. 7–16.
Tubeufia claspisphaeria
(from holotype). 7. Ascomata on natural substratum. 8. Longitudinal section through
an ascoma. 9. Setae. 10. A cluster of mature and immature asci. 11. Asci. 12–16. Ascospores. Pseudoparaphyses in F
IG
.10
(arrowed). Bars: 7 5250 mm; 8 550 mm; 9 525 mm; 10–11 530 mm; 12 515 mm; 13–16 510 mm. F
IGS
. 8–10 and 12
mounted in water, other figures mounted in lactophenol.
F
IGS
. 17–23.
Tubeufia paludosa
(from HKU[M] 17122). 17. Ascomata on natural substratum (arrowed). 18. Anamorph
(
Helicosporium
sp.). 19. A cluster of mature and immature asci. 20–21. Asci. 22–23. Ascospores. Pseudoparaphyses in F
IG
.19
(arrowed), Mucilaginous pads in F
IG
. 22–23 (arrowed). Bars: 17 5200 mm; 18–21 530 mm; 22–23 525 mm. F
IGS
. 18, 20–
21 and 23 mounted in lactophenol, other figures mounted in water.
670 M
YCOLOGIA
T
ABLE
I.
Tubeufia
species and their present synonyms (accepted name in bold)
Taxa Synonym Notes
T. acaciae
Tilak & S. B. Kale None Although not examined this taxon may
not be a
Tubeufia
with its violet to red
ascomata and multiseptate ascospores
T. aciculospora
Katum. & Y. Hara-
da
None This species is unusual as it has asco-
spores with pointed ends Katumoto
and Harada (1979)
T. albo-ostiolata
Rossman
Thaxteriella alboostiolata
Transferred in Crane et al (1998)
T. alpina
L. Holm & Nograsek
Acanthostigmina longisporum
This was considered to be synonymous
with
A. longisporum
by Re´blova´ and
Barr (2000)
T. amazonensis
Samuels, Rossman
&E.Mu¨ll.
Thaxteriella amazonensis
Transferred in Crane et al (1998)
T. anceps
Penz. & Sacc.
Tubeufia paludosa
Synonymized by Rossman (1977)
T. asclepiadis
Bat. & Garnier
Saccardomyces socius
Henn. Rossman (1979)
T. aurantiella
(Penz. & Sacc.) Ross-
man
Section Nectrioidea (5
Calonec-
tria aurantiella
)
Mentioned in Crane et al (1998), trans-
ferred from
Calonectria
(Rossman
1979)
T. brevispina
(M. E. Barr & Roger-
son) J. L. Crane, Shearer & M.
E. Barr)
(5
Acanthostigmella brevispina
) Crane et al (1998)
T. cerea
(Berk. M. A. Curtis)
Ho¨hn.
Section Necrioidea Mentioned in Crane et al (1998), for de-
scription see Sivanesan (1984)
T. clintonii
(Peck) M. E. Barr
Acanthostigma perpusillum
This was considered to be synonymous
with
A. perpusillum
by Re´blova´ and
Barr (2000)
T. coccicola
(Ellis & Everh.)
Podonectria coccicola
Petch Rossman (1987)
T. coronata
Penz. & Sacc.
Tubeufia paludosa
Synonymized by Rossman (1977)
T. cor ynespora
Munk
Thaxteriella corynespora
Transferred in Crane et al (1998)
T. cylindrothecia
(Seaver) Ho¨hn.
Tubeufia paludosa
Synonymized by Rossman (1977)
T. dactylariae
Chang None Chang (2003)
T. eriodermae
Etayo None Etayo (2002)—possibly
Chaetosphaerulina
T. genuflexa
Ho¨hn.
Acanthostigmella genuflexa
Barr (1977)
T. hebridensis
Dennis
Taphrophila hebridensis
Re´blova´ and Barr (2000)
T. helicoma
(W. Phillips & Plowr.)
Piroz.
Thaxteriella helicoma
Transferred in Crane et al (1998)
T. helicomyces
Ho¨hn.
Tubeufia paludosa
Synonymized by Rossman (1977)
T. indica
(Dharne & E. Mu¨ll.)
Deoray & Ujjainkar
Thaxteriella indica
Mentioned in Crane et al (1998)
T. javanica
Penz. & Sacc.
Tubeufia paludosa
Synonymized by Rossman (1977)
T. minuta
Munk
Herpotrichiella
Barr (1980)
T. miscanthi
W. H. Hsieh, C. Y.
Chen & Sivan.
Taphrophila miscanthi
Re´blova´ and Barr (2000)
T. nigrotuberculata
T. Hino & Ka-
tum.
Herpotrichia nigrotuberculata
Pirozynski (1972)
T. ovatum
Rossman
Thaxteriela ovata
Transferred in Crane et al (1998)
T. pachythrix
(Rehm) Rossman None For discussion see Rossman (1979)
T. palmarum
(Torrend) Samuels,
Rossman & E. Mu¨ll.
Section Nectrioidea Samuels et al (1979); mentioned in
Crane et al (1998)
T. paludosa
(P. Crouan & H.
Crouan) Rossman
Section Tubeufia Samuels et al (1979); mentioned in
Crane et al (1998)
T. pannariae
Etayo None Etayo (2002)—possibly
Chaetosphaerulina
T. parvula
Dennis None Drawn in Ellis and Ellis (1985)
T. pezizula
(Berk. & M. A. Curtis)
M. E. Barr
Thaxteriella pezizula
Type species of
Thaxteriela
(Sivanesan
1984, Barr 1980)
T. roraimensis
(Samuels & E.
Mu¨ll.)
Thaxteriella roraimensis
Samuels and Mu¨ller (1978); also men-
tioned in Crane et al (1998)
671K
ODSUEB ET AL
:
A
CANTHOSTIGMA AND
T
UBEUFIA FROM
H
ONG
K
ONG
T
ABLE
I. Continued
Taxa Synonym Notes
T. rugosa
C. Booth
Tubeufia helicoma
Barr (1980)
T. scopula
(Cooke & Peck), M. E.
Barr
Acanthostigma scopulum
Barr (1980), Re´blova´ and Barr (2000)
T. setosa
Sivan. & W. H. Hsieh
Acanthostigma minutum
Re´blova´ and Barr (2000)
T. stromaticola
(Henn.) Rossman Transferred from
Calonectria
Rossman (1979)
T. trichella
(Sacc., E. Bommer &
M. Rousseau) Scheuer
Taphrophila trichela
Re´blova´ and Barr (2000)
T. trichospora
(Berk. & Broome)
Petch
Ophionectria trichospora
Rossman (1977)
T. vermicularispora
(T. Hino & Ka-
tum.) Sivan.
Chaetosphaerulina vermicularis-
pora
This has black carbonaceous ascomata
and is probably not a
Tubeufia.
It was
described as a
Chaetosphaerulina
(Hino
and Katumoto 1954)
T. yasudae
(T. Hino) Sivan.
Chaetosphaerulina yasudae
Mentioned in Crane et al (1998)
ers of thick-walled textura globulosa, outer cells
brown-walled, inner cells with hyaline walls. Pseudo-
paraphyses 1.5–2.5
m
m wide, hypha-like, numerous,
cellular, unbranched. Asci (93–)105–120(–127)
3
(12–)14–16(–18)
m
m(
x¯
5
109
3
15.1
m
m, n
5
15),
8-spored, cylindric-clavate, bitunicate, with a small oc-
ular chamber, persistently pedicellate. Ascospores
(34–)45–60(–64)
3
5–7(–8)
m
m(
x¯
5
49
3
5.9
m
m,
n
5
35), fasciculate, elongate-fusiform, (3–)5–6(–8)-
septate, hyaline, sometimes slightly curved, smooth-
walled, guttulate, lacking appendages or sheaths.
Anamorph.
Unknown.
Colonies on potato-dextrose agar dark green to
black, 1 cm diam in 1 wk at room temperature (
;
28
C). Mycelium mostly immersed, aerial mycelium vel-
vety to fluffy, no pigment diffusing into agar, not
sporulating even after submergence overnight in a
bubble chamber. Mycelium less dense at the outer
edge than in center, with branching mycelial strands
extending from edge of colony.
Substratum.
Wood submerged in streams.
Known distribution.
Hong Kong.
Holotype.
HONG KONG. Lantau Island, Trappist
Monastery, small stream in forest, on wood partially
submerged, 21 Apr 2002,
K.D. Hyde
(HKU[M]
17123). Living cultures ex holotype HKUCC 9116,
HKUCC 9119.
Tubeufia paludosa (P. Crouan & H. Crouan) Ross-
man, Mycologia 69:383 (1977) (F
IGS
. 17–23)
Ascomata 185–350
m
m diam, superficial, globose,
brown to dark-brown, solitary, sparse, surface rough-
ened, membranous. Asci (162–)170–180
3
(14–)15–
18(–19)
m
m(
x¯
5
170
3
16.1
m
m; n
5
17), numer-
ous, 8-spored, cylindric-oblong, bitunicate, hyaline,
with a subapical ring, rounded at apex, persistently
pedicellate, pseudoparaphyses. Ascospores (77–)120–
135(–155)
3
4–5(–6)
m
m(
x¯
5
127
3
4.3
m
m, n
5
30),
fasciculate, narrowly elongate, cylindric or filiform,
hyaline, (18–)20 –25-septate, straight or slightly
curved, smooth-walled, guttulate, with small mucilag-
inous pads at each end.
Anamorph. Helicosporium
sp.
Colonies on potato-dextrose agar 3.5 cm diam in
3 wk. Mycelium mostly immersed, no pigment diffus-
ing into agar, hyphae less dense in outer zone, spor-
ulating, forming a brownish-gray turf of upright se-
tiform conidiophores, becoming ochraceous or
brownish with age; conidiogenous cells produced lat-
erally as thin-walled pegs. Conidia coiled two or three
times, multiseptate, coils 37–86.4
m
m diam, cells 4.6–
5.4
m
m wide.
Substratum.
Wood submerged in streams, decaying
woody fruit, palm fruit peduncle and bamboo.
Known distribution.
Bermuda, Brazil, Columbia,
Europe, Hong Kong, India, Indonesia, Panama, Trin-
idad, U.S.A. and Venezuela.
Specimen examined.
HONG KONG. Lantau Island, Trap-
pist Monastery, small stream in forest, on wood partially sub-
merged, 21 Apr 2002,
K.D. Hyde
(HKU[M]17122; living cul-
ture HKUCC 9118).
The genus
Tubeufia
has included at least 40 epi-
thets as listed in IndexFungorum (http://www.
indexfungorum.org/Names/Names.asp). Many of
these taxa have been transferred to other genera
(Barr 1980, Crane et al 1998) as summarized in T
A
-
BLE
I. Our two species are typical of
Tubeufia
be-
cause the ascomata are initially pale and have either
672 M
YCOLOGIA
F
IG
. 24. Diagrammatic representation of ascospores of
Tubeufia
species drawn to the same scale for comparison. A.
T.
acaciae
(from Tilak and Kale 1970). B.
T. aciculospora
(from Katumoto and Harada 1979). C.
T. aurantiella
(from Rossman
1987). D.
T. brevispina
(from Barr and Rogerson 1983). E.
T. cerea
(from Barr 1980). F.
T. claspisphaeria
(this paper). G.
T.
cylindrothecia
(from Barr 1980). H.
T. dactylariae
(from Chang 2003). I.
T. eriodermae
(from Etayo 2002). J.
T. helicomyces
(from Ellis and Ellis 1985). K.
T. palmarum
(from Samuels et al 1979). L.
T. paludosa
(from Sivanesan 1984). M.
T. pannariae
(from Etayo 2002). N.
T. par vula
(from Ellis and Ellis 1985). Bar 520 mm.
673K
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A
CANTHOSTIGMA AND
T
UBEUFIA FROM
H
ONG
K
ONG
a glabrous surface or only short setae (F
IG
. 24). In
ascospore size and morphology
Tubeufia claspis-
phaeria
is most similar to
T. acaciae, T. pachythrix
and
T. stromaticola.
It differs from these species in
that it has a row of hook-like setae that form radially
around the ostiole. The ascospores of
T. claspis-
phaeria, T. pachythrix
and
T. stromaticola
are elon-
gate-fusiform, while those of
T. acaciae
are cylindri-
cal or vermiform. Ascospores of
T. claspisphaeria
are
wider than those of the other three species. Our col-
lection of
Tubeufia paludosa
is similar to the descrip-
tion given by Barr (1980) but differs in that it has
globose ascomata and mucilaginous pads at both
ends of the ascospore.
Tubeufia
species have rarely been reported from
freshwater habitats (Shearer 1993) and are more
commonly found on rotting vegetation (Barr
1980).
Tubeufia paludosa
previously has been re-
ported from submerged grasses (Shearer 1993),
T.
cylindrothecia
and
T. palmarum
from submerged
wood (Ho et al 2001, Sivichai et al 2002). The an-
amorphs, however, commonly are reported from
submerged freshwater litter (Ho et al 2002, Sivichai
et al 2002).
DICHOTOMOUS KEY TO THE SPECIES OF
TUBEUFIA
1. Lichenicolous fungi
............................................................................
2
1. Not lichenicolous fungi
.........................................................................
3
2. Ascomata white, ascospores 105–145 34–5 mm, acicular, 17–19-septate
.......................
T. eriodermae
2. Ascomata orange-yellow to grayish-yellow, ascospores 40–50 3(3.5– 4.5 mm, fusiform, 5–8-septate
....
T. pannariae
3. Ascospores with ,10 septa
.......................................................................
4
3. Ascospores with .10 septa
......................................................................
14
4. Ascospores shorter than 21 mm
................................................................
5
4. Ascospores longer than 21 mm
................................................................
6
5. Ascomata reddish-brown, ascospores 14–15 33.5–4 mm, oblong-elliptical, 3-septate
...................
T. par vula
5. Ascomata hyaline to pale yellow, ascospores (13–)14–18(–21) 3(3.5–)4–5(–7) mm, fusoid, 3-septate
....
T. brevispina
6. Ascospores up to 7 septa
.....................................................................
7
6. Ascospores .7 septa
.......................................................................
11
7. Ascospore 48–56 36–7 mm, cylindrical or worm-like, often curved, pointed of both ends, 5–7-septate
.....
T. acaciae
7. Ascospores narrower than 6 mm
..................................................................
8
8. Ascospores less than 3.5 mm wide
..............................................................
9
8. Ascospores more than 3.5 mm wide
............................................................
10
9. Ascospores 65–75 32.5–3.5 mm, acicular, 4–7-septate
........................................
T. aciculospora
9. Ascospores 66–150 32–3 mm, filiform, 5–7-septate
..........................................
T. helicomyces
10. Ascospores 40–57 34–5 mm, long-fusiform, multiseptate (4-celled according to Rehm 1907)
.......
T. pachythrix
10. Ascospores 50–70 34–5 mm, long-fusoid, 5–7-septate
....................................
T. stromaticola
11. Ascomata with a row of hook-like setae, form radially around the ostiole, ascospores (34–)45–60(–64) 35–7(–8) mm,
elongate-fusiform, (3–)5–6(–7)-septate
..................................................
T. claspisphaeria
11. Ascomata not as above
.........................................................................
12
12. Ascospores 32–36 36–7 mm, fusiform, 7–8-septate
.......................................
T. dactylariae
12. Ascospores narrower than 6 mm
..............................................................
13
13. Ascospores 30–40(–50) 33– 4 mm, fusiform with subacute to round ends, 5–7(–9)-septate
...........
T. palmarum
13. Ascospores 45–66 33.5–5 mm, narrowly fusiform to cylindric, 7–9-septate
........................
T. aurantiella
14. Ascospores shorter than 70 mm
...............................................................
15
14. Ascospores (70–)100 –200(–230) 3(2–)3.5–7(–8) mm, fusiform with acute ends, up to 35 septa
......
T. paludosa
15. Ascospores 40–55(–65) 3(2.5–)3–5 mm, elongate clavate or fusoid, (5–)7–9(–13)-septate
..........
T. cylindrothecia
15. Ascospores (27–)30–52 32.5–3.5(– 4.5) mm, elongate fusoid, often curved, (5–)7–10(–13)-septate
..........
T. cerea
ACKNOWLEDGMENTS
The senior author would like to thank Department of Ecol-
ogy & Biodiversity, The University of Hong Kong, for facil-
itating the work at the Centre for Research in Fungal Di-
versity. Thanks to Chiang Mai University Graduate School
for partial support to the first author’s doctoral study.
Thanks also are extended to B. Bussaban, H.Y.M. Leung,
M.H.W. Ng, I. Promputtha and S. Thongkantha for various
help.
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