Table Of ContentTaxonomic Studies on Asplenium Thamnopteris
sect.
(Aspleniaceae) Cytological Observations
I:
Dong
Shi-Yong
South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China;
Asplenium L. a natural genus with about 700 or more species in various
is
regions all over the world (Kramer and Viane, 1990; Bellefroid et ah, 2010).
Though many
widely accepted monophyletic, Asplenium, with
as so species,
subdivide because
difficult to of a lack of suitable distinguishing characters
is
the intrageneric Asplenium Thamnopteris
at level. sect. C. Presl (1836), or
Wu,
genus
treated as a separate Neottopteris (Smith, 1841; 1999), however,
is
one of the very few groups which can be clearly recognized. characterized
It is
by and
simple, entire fronds intramarginal veins in morphology, usually
is
on and
epiphytic tree trunks or rocks in forests with bird-nest-like appearance,
and Old
distributed in tropics subtropics of the World.
is
Despite being readily recognized as a morphologically distinct group, the
circumscription of species is confusing and the identification is very difficult
within Asplenium Thamnopteris. Since Linnaeus's A. nidus the
sect. (1753),
member
first in this group, 33 taxa have been published (Tardieu-Blot, 1933;
Ching and Wang, Wu,
1964; Holttum, 1974; Miao, 1980; Jones, 1988; 1989;
Murakami The was
et al, 1999). establishment of these taxa mostly based on a
minor variation of one or two characters, especially the shape of fronds, which
Though
results in the obscure delimitation of species. Holttum (1974)
presented a taxonomic revision on the section Thamnopteris, his treatment
was based on herbarium specimens time and many problems
at his in
nomenclature and taxonomy remained. Holttum no
After
(1974),
still
taxonomic work was carried out on this group other than that of Seto (1979)
and Murakami et (1999) who studied the morphology and the molecular
al.
systematics of Thamnopteris from Japan, respectively. Lacking a comprehen-
sive comparison of characters, the taxonomic confusion of this section partly
is
many
by named
reflected the present situation that specimens were as
DONG: CYTOLOGY OF ASPLENIUM
THAMNOPTERIS
SECT.
"Asplenium
nidus
L. s.l." or "A. phyllitidis sensu Holttum" and more than 20
new
species were proposed but not formally published based on
materials in
Aiming
taxonomy
to revise the of this difficult group, the present author
undertook examining
type collections, studying materials in large herbaria of
southeastern Asia and Europe, field surveying in China, Philippines and
chromosome
Indonesia, counting, and gametophyte
observations
in last six
years. Cytological data are necessary and important understand
to the concepts
and
and
relationships of species to date only A. antiquum Makino, A.
australasicum, A. nidus, and A. Don have been documented
phyllitidis D. with
chromosome numbers n = and =
72 2n 144 Abraham
or 1960;
/ {Bir, et al,
Kawakami,
1962; 1970, 1997; Tsai and Shieh, 1983; Kato and Nakato,
1999;
Yatabe and
et 2001; Tindale Roy, These
al., 2002). cytological observations
are
my
reported as the first part of serial studies on the taxonomy of Asplenium
Thamnopteris.
sect.
Material and Methods
morphology and
Gross name
identification.-—In order to correctly the living
materials involved morphology was
in this research, gross studied based on
herbarium specimens and wild and The
cultivated living plants. present
author examined specimens Asplenium
of Thamnopteris
deposited
sect. in the
BM, GAUA,
following herbaria: BO, HITBC, IBK, IBSC, KUN, KYO,
K, PE,
L, P,
PNH, PYU, and SING
The
before the year 2009. specimens
of the section of
Thamnopteris examined
being above amount
in herbaria about 2500
to
To
collections. understand the living state and variation of morphological
made
characters, field observations in 17 trips to southern and southwestern
I
China
since 2005, one trip to Mindanao, Philippines in 2007, one
trip to Java,
Indonesia in 2009, and one trip to West Papua and Kalimantan, Indonesia
in
2010. In each field trip, the state of following characters was determined or
measured:
distribution, shape, color, and size of scales; outline, size, and basal
shape of fronds; prominent extent and transaction of frond midribs;
and
distribution, length, density of sori; color of stipes; spreading angle of
and
fronds; habitat. In addition, one or two living plants were introduced
to
my
green house South China
in Botanical Garden, Guangzhou, morpho-
for
and chromosome
logical observation
counts.
Cytology.—Root tips of sporophytes or prothalli in gametophyte phase were
chromosome
used
for counts. Living plants (sporophytes) or fresh spores were
me
collected by from natural populations in various localities or from Chinese
my
gardens and
(Table cultivated green house South China
1) in in Botanical
Garden. Fresh immature
root tips or prothalli (nearly cordate in shape, before
sex organs formed) raised from spores were pretreated 3-6 hours with
for
mol/L
0.002 8-hydroxyquinoline at about 25 C, then fixed for hour in
1
volume
Carnoy's fluid of pure acetic acid and volumes 95%
(1 3 of ethanol)
at
N
about 4 C. The tips were hydrolyzed 10 minutes HCL
for in 1 at 60 C,
macerated for 10 min in 45% acetic acid, and then stained in 2% aceto-orcein
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3 (2011)
2%
for 1-2 hours. Finally, the root tips or prothalli were squashed in aceto-
The chromosomes were counted and photographed
orcein. using
a light
microscope (Olympus BX41).
—
Reproductive The number sporangium
characteristics. of spores in a
mode
indicates the possible reproduction most leptosporangiate
for ferns,
ferns with 64 spores per sporangium reproduce sexually while those with
i.e.,
32 spores per sporangium reproduce apogamously (Manton, 1950; Lovis, 1977;
DONG: CYTOLOGY OF ASPLENIUM THAMNOPTERIS
SECT.
When
Walker, Kato and number
1979; Nakato, checking
1999). the of spores, a
well-developed, intact sporangium was selected and moved to a drop of water
on a microscope slide with a needle. Using the needle break the sporangium,
to
spores were freed from the sporangium, and then number was
the of spores
counted with microscope (Olympus
a light BX41). For one given specimen,
the
spores of at least five sporangia were counted.
The
on morphology and
data
distribution given in the following section
is
entirely based on herbarium specimens examined and
recent collections
gathered by
the present author.
Results and
Discussion
As shown
in Table the somatic chromosome number 216 Asplenium
1, is for
humhertii and 144 for other species. As the basal number of chromosome
(x) is
Asplenium,
36 in A. humhertii hexaploid and
is others are all tetraploid. In
my
examination showed
addition,
there are 64 spores in each sporangium and
the spores can normally develop and produce new
sporophytes
the
for
all
specimens examined
cytologically in this study. Therefore, the eight species
with chromosomes
counted
here are of sexual reproduction mode.
Chromosome
NUMBER, nomenclature, and
identification, distribution
Asplenium antiquum Makino
in Jap. Bot. 6: 32. 1929. Neottopteris antiqua
J.
Masam.
(Makino) TYPE.—
in Trans. Nat. Hist. Soc. Taiwan. 22: 215. 1932.
Kagoshima: Yakusima,
Japan. 30 1961, H. 289 (neotype! designated
Jul Ito
SING).
here,
Cytology—The
plants of this species cultivated in two botanical gardens of
=
China were examined and
determined be with n and
to tetraploid 72
(Fig.
1)
=
2n The
144. origin of the plants examined unknown.
Previously,
a
is
population of this species from Tokunoshima was reported with
IsL, Japan, the
same chromosome number =
(2n (Kawakami,
144) 1970).
Diagnostic characters.—Stipe scales broadly lanceolate, 15-30 x 2-6
(8)
mm;
fronds narrowly lanceolate; midribs prominent on both
slightly surfaces;
sori long and sparse, occupying 2/3-3/4 length of veins, 6-10 every cm
sori 2
length along midribs.
Distribution—China (common
(Fujian), Japan in southern Korea
islands),
The
(Cheju Island). misreport of this species in Hainan Island, China (Wu,
1999),
due
to the misidentification of A. phylUtidis subsp. malesicum Holttum.
is
Representatives of 32 specimens examined.—China. Fujian: Luoyuan, 25"07'N,
430 m, Dong
123 32'E, 12 2010, 3403 without
Jul (IBSC); specific 20
locality,
Aug
1943, Lin s.n. (IBSC, herb no. 295306). Taiwan: Nantou, 1300 m, 28 Aug
2002, Lu s.n. (PYU); Taitung, 23 Jan 1940, Tagawa 3071
(L).
May
Korea. Cheju: without specific locality, 29 1908, Taquet 2369
(P).
Hizen: without
Japan. specific locality, 16 Jun 1909, Mistuo (KYO).
s.n.
Kagoshima: Cape Sata, 19 Oct 1924, Tashiro (KYO); Oosumi, 26
s.n. 1913,
Jul
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asi;. 1. /l.s/i/.;.M,;;; i. irr^zhou, China [Dong 3360 2. A. antrophyoides
,. ' );
: i
m
Guangxi, Cliiria iisicnm cultivated in Cxuangzhou. China [Dong
I/'
A. cymbifoliuni Philippines [Dong
2610].
90); 4.
-ii.io,
DONG: CYTOLOGY OF ASPLENIUM THAMNOPTERIS
SECT.
Aug
Tashiro (KYO);
s.n. ibid., 12 1916, Tashiro s.n. (KYO); ibid., 100 m, 24 Jan
1965, Togashietal. 10071 Yakushima, Aug 1907, Kudo 5 (KYO); Sep
(L); ibid., 1921,
Koidzumi s.n. (KYO, PE); ibid., 13 Dec 1956, Togasi 30-50 m, 14 Nov
s.n. (P); ibid.,
&Nagamasu
1983, Mitsuta 275 (PYU); 30-100 m, 16 1979, Yamazaki
ibid., Jul et
al.
2330 (KUNJ; 30 Jul 1961, Ito 289 (SING). Nagasaki: Nam, 28 Sep 1910, Tashiro
s.n.
(KYO). Nagono: without specific location, 1866-74, Savatier 1551 Okinawa:
(P).
Iriomote, 200 m, 19 Mar 1982, Saiki 2050 (PYU). Wakayama: Susami, 10 Aug 1913.
Ui s.n. (KYO); Katsuura, 5 Jun 1951, Kodama s.n. (PE); Wakayama, Aug 1901,
Nov
Kinashi s.n. (KYO). Without specific locality: 11 1881, Dickins 1877
(P).
In nomenclature, Asplenium antiquum was invalidly published by Makino
because no was
(1929) type designated. tried to access the original material
I
by Makino
studied but designated neotype
failed. a for this species to validate
I
name
Asplenium antiquum
the Makino.
Asplenium
antrophyoides
H. Christ in Bull. Acad. Int. Geogr. Bot. 20: 170.
Mem.
1909. Neottopteris antrophyoides Ching Fan
(H. Christ) in Bull.
Inst.
TYPE.—
Biol., Bot. 10: 1940. China. Guizhou: Lofu, Sep 1907, Cavalerie
7.
1877
(holotype, isotype, BM!).
P!;
—
The chromosome number
Cytology. somatic 144 population from
for a
is
Guangxi, China
(Fig. 2). This is the first cytological record for A. antro-
phyoides.
X mm;
Diagnostic characters.— broadly 15-20 3-5
SXi^ie scales lanceolate,
more
fronds or less spathulate the upper part the broadest; midribs
v\^ith
obviously keeled on abaxial surface with the transection long and
deltoid; sori
cm
usually occupying 2/3-3/4 7-10
sparse, length of veins, every
(12) sori 2
length along midribs.
Distribution.—China (Guangdong, Guangxi, Guizhou, Hunan,
Sichuan,
Yunnan), Thailand (Chiang Vietnam
Mai), (Tonkin).
Representatives of 160 examined.— Guangdong:
ca. specimens China. Liannan,
Wang
150 m, 16 Jan 2005, et 725 (IBSC); Lianxian, 21 Nov 1930, Ko 50962
al.
(PE). Guangxi: Huangjiang, 25°07'N, 107°58'E, 600 m, 13 Aug 2005, Dong 1976
&
(IBSC); Lingyun, 11 Dec 1933, Steward Cheo 7 (PE); Longzhou (formerly
Lungchow), Morse
1901, 63 (K); Napo, 23°00'N, 105°51'E, 1000 m, 13 Jun 2009,
Dong
2956 Dong
(IBSC); Tianlin, 24°29'N, 106°21'E, 1150 m, 20 Jun 2009, 3001
(IBSC). Guizhou: Libo, 25"17'N, 107°56'E, 600 m, 30 Dong
2010,
Jul et al.
GZE52
(IBSC); Pingchow, 350 m, 15 Sep 1930, Tsiang 7141 (BM, Hunan:
P).
Nov
Xinning, 370 m, 14 1962, Liu 15545 (IBSC, KUN). Sichuan: Qianwei,
Aug
380 m, 12 1984, Xing 05235 Yunnan: Hekou,
et (PE). 22 40'N,
al.
U
103=56'E, 150 m, 6 Oct 2009, Song 116 (IBSC); Jingdong, 1350 m, 7 Dec 1961,
3740 (KUN); Jinping, 850 m, 17 Jan 1959, Chu 4704 (PYU); Maguan, 650 m, 23
Nov Chu
1982, et 15340 (PYU); Mengla, 700-800 m, 25 1984, Sino-Japan
al.
Jul
Nov Wang
Exped. 104 (KUN); Yanshan, 1100 m, 15 84955 (KUN,
1939, PE).
Thailand. Chiang Mai: Chiang Dao, 19 25'N, 98 1500-1900 m, Dec
55'E, 7
Hennipman
3256 1600-2100 m, Tagawa
1965, (L); ibid., 4 Jan 1966, et
al.
T4210 1600 m, Mar Maxwell 95-176
(IBSC,
L); ibid., 2 1995,
(L)
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Cao Nguyen
Vietnam. Bang: Binh, 22°39'N, 500-550 m, Apr
105°57'E, 21
1999, Loc et al. 1609 (P); without specific locality, Feb 1925, Colani 1982
(P).
Lai Chau: Ngai Thau, 1000-1200 m, 7 Apr 1936, Poilane 25554 without
(P);
specific locality, 800-900 m, 6 Jan 1938, Poilane 27030 Thanhoa: Lung
(P).
Van, 1000-1200 m, 26 Poilane 18889
Jan 1931,
(P).
Asplenium
australasicum Sm]
Hook.,
Fil. Exot. 88. 1859. Neottopteris
t.
(J.
TYPE.—
australasica]. Sm., CuU. Ferns 49. 1857. Cult. Hort. Bot. Kew., origin
BM?
Australia (lectotype designated by Holttum in 1974, isolectotype,
K!).
Cyto7ogy.—Cultivated plants from South China Botanical Garden were
A
examined. of 144 chromosomes were counted sporophytes
total
for
(Fig.
3)
my
and
about 72 for gametophytes. Before examination, this species was
reported to be tetraploid with 144 chromosomes based on from
root-tips
Howe
Middle
Brother and Lord and
Island, Australia (Tindale Roy,
2002].
Diagnostic characters.— Stipe scales narrowly lanceolate, 10-20 X 1-1.5 mm;
fronds lanceolate; midribs distinctly keeled on abaxial surface with the
transection deltoid; sori short and dense, occupying less than 1/2 length of
cm
veins, (10)13-18 sori every 2 length along midribs.
New
Howe
Distribution. —Austvalia (Queensland, South Wales, Lord
Island], Fiji, Samoa, Tonga, Vanuatu (New Hebrides], Polynesia (Tahiti] and
other
Pacific islands.
Howe
Representatives of ca. 110 specimens examined.—Australia. Lord Island:
without specific locality, Sep 1853, Horald 696 Dec 1869 More 76
(K]; ibid.,
(?],
New
May
(K]. South Wales: Coffs Harbour, 150 m, 2 1956, Constable P7404
(K);
Newell Falls Creek, 3000 Werner 18 Queensland: Cape York, 420 m, 17
(K].
ft,
Aug
Mt
1948, Brass 19879 (K]; Lewis, 16=29'S, 145 14'W, 21 Sep 1975, Coveny
Mt
Nov
7242 (K, L]; Mistake, 24 1930, Hubbard 5202 (K]; Richmond River,
Anonymous
1885, s.n. (SING, herb no. 0079510].
Pacific Islands. Christmas: 1 52'N, 157 20'W, 4 Dec 1987, Maclean
s.n. (K]. Fiji:
May
Ovalau, 17 42'S, 178°48'W, 100-300 m, 8-11 1953, Smith 7322
(K, L); Viti
Levu, 17°58'S, 178°06'W, 150-250 m, 17 Oct Smith 8602 Guam:
1953,
(K, L].
Finegayan, 29 Jul 1987, Flis 15398 (K]. Henderson: North end, 24°22'S, 128°19'W,
5 m, 17 June 1934, John &Fosberg 15088 Mangaia: 21 157°55'W, Oct
(K). 55'S, 1972,
New
25 54 Nov Anonymous
Dickie (K]. Caledonia: 21 26'S, 165 'W, 1911, 147
(K);
Mar
Hennecart
v^rithout specific locality. 1889, s.n. (K]. Nine: 19°01'S, 169"55'W,
m, Yuncher9590
20 11 Jan 1940, Oeno: 23 130 44'W, Quayle 409
56'S,
(K]. 1921(?],
Samoa: Laugapapa, Sep 1877, Powell 164 without
(K]. (K]; specific locality, Jun
Mt May
1872(?], Powell 164 [K]. Tahiti: Marau, 17=37'S, 149 32'W, 1250 m, 30 1985,
New
Fosberg 64558 Tonga: Tavau, 18 174=00'W, Crosby 277 Vanuatu:
(K]. 40'S,
[K.].
Hebrides, 17 168 21'W, 580 2002
31'S, 2 1971, Braithwaite
Jul (K].
ft,
This species morphologically characterized by having fronds ascending
is
and
steeply midribs keeled on Nakato
distinctly abaxial surface. (1987]
reported a population of "A. australasicum" from Iriomotejima, with
Japan,
144 somatic chromosomes. However, known
A. australasicum so be
is far to
Queensland and nearby The
restricted to Pacific islands. so-called A.
DONG: CYTOLOGY OF ASPLENIUM THAMNOPTERIS
SECT.
163
australasicum from southern Japan proved to be A. setoi N. Murak. & Seriz.
based on molecular (Murakami
data
et al, 1999].
Asplenium cymbifolium lingganum Alderw.
in Bull. Bot. Buitenzorg
Jard.
f.
TYPE.—
Aug
184. 1922. Indonesia. Lingga Arch., 8 1919, Bunnemeijer
III, 5:
7388
(holotype, K!, isotype,
L).
—
Both and
Cytology. the root tips prothalli originally from a Philippine
population were examined. The chromosome number
cytologically 144
for
is
and young
root tips (Fig. 4) ca. 72 for prothalli. This taxon confirmed to be a
is
sexual tetraploid for the time.
first
X mm;
Diagnostic characters.—Stipe scales narrowly lanceolate, 10 1-2
ca.
and widened
fronds lanceolate at base; midribs prominent on abaxial surface
with and
the transection elliptic; sori long sparse, occupying 2/3-3/4 length of
cm
4-7
veins, sori every 2 length along midribs.
(9)
New
Distribution.—Indonesia (Lingga Arch., Sumatra), Papua Guinea,
Philippines (Mindanao).
Additional specimens examined.—Indonesia. Jappen-Biak: Wasabori near Seroei,
Aug
12 1939, Aet &- Idjam 400 (BO). Sumatra: Sikundar Nature Conservations,
100-250 m, Apr
14 1971, Iwatsuki S361 without
et Ditschi: specific
al. (K).
Mayr
locality, 1200 m, Jun 1928, 183a (BO).
New Mt
Papua Guinea. East Sepik: Samai, 4°29'S, 142°41'E, 450 m, 27 Apr
Takeuchi 6216A
1991,
(K).
Mindanao: Nov Dong
Philippines. Marilog, Davao, 1200 m, 17 2007, 2610
(IBSC).
In comparison with A. cymbifolium with wide fronds (18-20 cm), the form
lingganum was by much cm
characterized smaller fronds wide). Besides the
(6
remarkable difference in frond size the form lingganum from the form
differs
cymbifolium in lacking dense scales on abaxial surface of midrib towards the
base. So seems more suitable to treat the form lingganum as a variety or a
it
separate species from A. cymbifolium.
Asplenium
humbertii Tardieu, Asplen. Tonkin 1-2. 1932.
25, pi. 2,
f.
Tagawa
Neottopteris humbertii (Tardieu) in Bot. 22: 161. 1948.
Jap.
J.
TYPE.—
Vietnam. Tonkin, Jan 1886, Balansa 68 (holotype,
P!).
—
A
Cytology. population from Hainan Island and other one from Guangxi,
=
China proved be sexual hexaploids with chromosome number 2n 216
to
(Fig. 5). This species is the only hexaploid so far found in Asplenium sect.
Thamnopteris.
—
X mm;
Diagnostic characters. Stipe scales broadly lanceolate, 8-10 2-3
more
fronds or less spathulate with long and narrow stipes; midribs slightly
prominent with the transection long and usually occupying
elliptic; sori sparse,
cm
more 8-13
than 2/3 length of veins, sori every 2 length along midribs.
—
Distribution. China (Hainan, Guangxi, Yunnan), Laos (Cammon), Thailand
Vietnam Asplenium
(Chiang Mai), (Tonkin). humbertii only occurs
in
limestone within these regions.
DONG: CYTOLOGY
OF ASPLENIUM
THAMNOPTERIS
SECT.
Additional SPECIMENS EXAMINED.— Hainan: 19=01
China. Changjiang,
'N, 109"06'E,
1000 m, 20 Oct 2004, Dong 1234 (IBSC); ibid. 5 Apr 1987, Chen 703 (IBSC).
Guangxi: Longzhou,
22=34'N, 106°48'E, 200 m, 14 Mar Dong 2287B
2007,
Yunnan:
(IBSC). Hekou, 22°40'N, 103°56'E, 150 m, 11 Aug Dong 3248
2010,
(IBSC); ibid., 400 m. Song 119 (IBSC); ibid., 400 m, Feb 1959, Chu 3922 (PYU).
Jinping, 23°02'N, 103 24'E, 500 m, 12 Aug Dong 3432
2010,
(IBSC);
ibid.,
600 m, 10 Oct 2009, Song 133
(IBSC).
Gammon:
Tham, Nov
Laos. 160 m, 1930, Colani 4098 Thok, 160 m, Nov
(P);
1930, Colani 4091
(P).
Thailand. Chiang Mai: Chiang Dao, 1100 m, 16 Feb 1958/59, Sorensen ah
et
1237
(K); ibid., 550 m, 15 Jan 1989, Maxwell 89-59 Chiang name
(L). Rai: local
unknown, May
500 m, Muang
27 1926, Garrett 289 Kwai Noi
(K). Sing: Basin
150 m, 3 Jun 1946, Hoed 920
(L).
Bac May
Vietnam. Giang: Lang Met,
1929, Colani 1982 Lang Son: Lang
(P).
Mac, Feb
1929, Colani Without
260-390
s.n. (P). specific locality, m, 22 Dec
1964, Sino-Vietnam Exped. 699
(PE).
Asplenium
nidus
L., Sp. PI. 1079. 1753. Neottopteris nidus Sm.
in Bot.
(L.)
J. J.
TYPE.—
409. 1841. Osbeck
3: Indonesia. Java, 49 (lectotype designated by
Holttum
in 1974,
L).
Cytology.—Wild
plants of species from Hainan, Yunnan,
this Guangxi,
or
cultivated plants (Table proved to be sexual tetraploid with chromosome
1)
number = =
2n 144 n
(Fig. 6) or 72. This species has been reported as tetraploid
from
northern India southern (Abraham
(Bir, 1960), India Kagoshima,
etal., 1962),
Japan (Kawakami, 1970, 1997), Taiwan (Tsai and Hsieh, Hainan
1983), Island,
China and
(Kato Nakato, and from
1999), Indonesia
Java, (Yatabe et al, 2001).
—
Diagnostic characters. Stipe scales narrowly lanceolate, 15-20 x 1-2 mm;
fronds narrowly
lanceolate; midribs obviously prominent on
adaxial surface
and on
nearly
abaxial and
flat surface; sori short dense, occupying usually
less
than cm
1/2 length of veins, 12-18
(10) sori every 2 length along midribs.
Distribution.—The whole range Asplenium
of Thamnopteris, mainly on
sect.
Asia and
tropical Pacific islands, west to Africa and northeast Hawaii.
to
Representatives of ca. 1220 specimens examined.—Australia. Queensland:
fron
Range, Cape York
Peninsula, 20 m,
6 Jun 1948, Brass 19049
(K).
China. Xizang: Motuo, 800 m, Aug
11 1974, Qinghai-Xizang Exped. 74-4215
(KUN, Mt
Hainan:
PE). Yinggeling, 680 m, Dec Dong
3 2005, 1689 (IBSC).
Guangxi: Napo, 51
23 OO'N, 105 'E, 1000 m, 13 Jun 2009, Dong 2939 (IBSC).
Yunnan.
Malipo, 23 Ol'N, 104 46'E, 780 m, 17 Jun 2009, Dong 2980
(IBSC).
Andamans: Mt
India. Harriet, 3 Jan 1998, Balachandra 0879
(K).
Kalimantan:
Indonesia. Balikapapan, 01 09' 116 50' 40 m, 16 Dec
S, E, 2010,
Dong
3470
(IBSC); Samarinda, 01 07' S, 117 12' E, 20 m, 18 Dec 2010, Dong
3474
(IBSC). Irian Jaya: Manokwari, 00 59' 133 58' 40 m, Dec
S, E, 12 2010,
Dong &
3455 3456 (IBSC). Java: Bogor, 280 m, 25 Oct 2009, Dong 3362
(IBSC).
Seram: Manusela
National Park, 3 08-09'S, 129 29'E, 1290-2000 m,
3 Jan
1985, Kato C350his
et (BO).
al.
