THAI FOR. BULL. (BOT.) 41: 23–38. 2013.
The genus Leucophanes (Calymperaceae, Bryophyta) in Thailand
CHATCHABA PROMMA1 & SAHUT CHANTANAORRAPINT1
ABSTRACT. A review of the genus Leucophanes Brid. in Thailand is presented based on the study of fresh and herbarium specimens.
Four species are recognized, namely L. angustifolium Renauld & Cardot, L. candidum (Schwägr.) Lindb., L. glaucum (Schwägr.)
Mitt. and L. octoblepharioides Brid. The genus is widely distributed in the southern half of the country. A key to species, descriptions
and line drawings are provided, and notes on the ecology and geographical distribution of the species.
KEY WORDS: bryophytes, Calymperaceae, Leucophanes, mosses, Thailand.
INTRODUCTION
The genus Leucophanes was established by
Bridel (1826), based on a collection of L. octoblepharioides of Nees v. Esenbeck from Java. This genus
is a member of the family Calymperaceae. It includes
12 species worldwide, distributed mainly in tropical
and subtropical regions particularly in Asia and
Africa (Salazar Allen, 1993). Plants of Leucophanes
are characterized by whitish-green leaves growing
in loose to dense tufts or cushions; stems erect,
sparsely branched, branches short; leaves multistratose, narrowly to broadly lanceolate to retuse at
apex, costa percurrent to short excurrent, with a
stereid band.
A monograph off Leucophanes had been
published by Salazar Allen (1993), but there are
few reports of this genus from Thailand, probably
due to the paucity of bryological studies in this
country. This study is based on fresh specimens
from fieldwork as well as herbarium specimens in
BCU, BKF, L and PSU. In addition, the distributions
of each species were assessed by fieldwork throughout
Thailand during 2010−2013, and using the available
taxonomic literature. Distributional and ecological
data were compiled, and detailed descriptions,
illustrations and key to species are provided.
1
Four species of Leucophanes have been
recorded for Thailand: L. angustifolium Renauld &
Cardot, L. candidum (Schwägr.) Lindb., L. glaucum
(Schwägr.) Mitt. and L. octoblepharioides Brid.
Thai species are mostly epiphytes, growing on tree
trunks, usually from ground level to 3 m high but
are also found on decaying wood, soil and rocks,
including limestone and sandstone. They are found
in various forest types, from dry to moist deciduous
or evergreen forests, i.e. coastal, mangrove, spring,
lowland evergreen and montane forests, from lowland
to over 1500 m a.s.l.
Based on geographical localities from the
field surveys, herbarium specimens and data from
previous publications (Brotherus, 1901; Dixon,
1932, 1935; Tixier & Smitinand, 1966; Touw, 1968;
Thaithong, 1984; Sornsamran & Thaitong, 1995),
the genus is widely distributed in the southern half
of the country; it does not seem to occur in the
northern part except for one locality (Fig. 1). The
Southern Peninsular region exhibits great diversity
of species. The most common species off Leucophanes
in Thailand is L. octoblepharioides, distributed in 5
floristic regions including North-Eastern, Eastern,
South-Eastern, South-Western and Peninsular
region (Fig. 1), while L. candidum has a restricted
distribution, found only in the Peninsular region.
Department of Biology Faculty of Science Prince of Songkla University, Hat Yai, Songkhla, 90112, Thailand; e-mail: sahut.c@psu.ac.th
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THAI FOREST BULLETIN (BOTANY) 41
Figure 1. Distribution of the species of Leucophanes in Thailand. Thai Floristic regions N= Northern, NE= North-Eastern, E=
Eastern, SE= South-Eastern, C= Central, SW= South-Western, PEN= Peninsular.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
TAXONOMIC TREATMENT
Leucophanes Brid.
Bryol. Univ. 1: 763. 1826; A. Eddy, Handb. Males.
Mosses 2: 40. 1990; Enroth, Acta Bot. Fenn. 139:
98. 1990. N. Salazar, Bryophyt. Biblioth. 46: 54.
1993. Type species: L. octoblepharioides Brid.
Plants epiphytic or terrestrial, small to
moderately large, whitish-green, in loose to dense
tufts or cushions. Stems erect, orange-brown to
dark red, sparsely branched, branches short; stem
in cross section rounded to irregularly triangular;
central strand absent, large central cells surrounded
by one or two layer of small, thick-walled cortical
cells. Rhizoids papillose, orange-red to brownishred, arising from stem and apex of leaves. Leaves
erect to more or less contorted when dry, loosely
overlapping when moist, narrowly to broadly
lanceolate, concave to keeled at base or through most
of their length, plane above; apex acuminate, acute,
obtuse, retuse or bent backwards; costa percurrent
to shortly excurrent, smooth to strongly spinose on
back and apex, convex on the abaxial side, flat on
the adaxial side; margins entire below, serrate
25
above with single or paired teeth; marginal sterome
forming a distinct, sharply differentiated border;
basal hyaline lamina asymmetrical, sheathed at leaf
base, hyaline lamina extending 1/9–1/2 leaf length
on both sides of costa, irregularly bistratose; hyalocysts in surface view rectangular with rectilinear to
sinuose walls, quadrate, irregularly pentagonal to
hexagonal. Leaf in cross section multistratose, a
central layer of quadrangular chlorocysts, surrounded
on each surface by one or more layers of porose
hyalocysts; median strand or sterome composed of
stereids and 2–4 well- developed guide cells, deeply
immersed below the upper surface in limb, exposed
or covered by abaxial hyalocysts. Gemmae on leaf
apex, papillose, fusiform-clavate.
Dioicous or monoicous. Sporophytes rare;
setae 3.1–15 mm long, sinistrorse, smooth; capsules
1.8–3.0 mm long, dark orange, erect to horizontal;
stomata superficial at neck of urn; peristome teeth
16, more or less papillose, lanceolate, with a colorless
prostome, 2–3 cells high; opercula long, subulaterostrate. Calyptrae cucullate. Spores 10–20 μm in
diameter.
KEY TO SPECIES OF LEUCOPHANES IN THAILAND
1. Plant robust; leaf broadly lanceolate, recurved or squarrose, costa smooth on the dorsal side throughout the leaves L. candidum
1. Plant small to medium size leaf narrowly lanceolate to laceolate, erect-spreading to recurved-patent, costa strongly spinose or
smooth on the dorsal side in upper parts of leaves
2. Leaf contorted when dry. Hyalocysts in surface view on abaxial surface at median leaf oblong with sinuose walls and concave
bulging end walls
L. glaucum
2. Leaf little altered when dry, not twisted. Hyalocysts in surface view on abaxial surface not as above
3. Costa strongly spinose on the dorsal side in upper parts of leaves. Leaf in transverse section with 1 or 2 irregular layers of
hyalocysts on the abaxial and adaxial sides of chlorophyllous layer
L. angustifolium
3. Costa smooth on the dorsal side of leaves, rarely spinose at apex. Leaf in transverse section with 1 layer of hyalocysts on
both sides of chlorophyllous layer
L. octoblepharioides
1. Leucophanes angustifolium Renauld & Cardot,
Rev. Bot. Bull. Mens. 9: 395. 1981; Enroth, Acta
Bot. Fenn. 139: 98. Fig. 27a–f. 1990; N. Salazar,
Bryophyt. Biblioth. 46: 69. Figs. 147–173. 1993.
Type: Réunion Island, Rodriguez 74c (lectotype
PC designated by Salazar Allen (1993); isolectotypes PC, S). Figs. 2–3.
Plants small to medium, 0.5–1.2 cm high,
growing in rather loose tufts. Stems erect, simple or
branched. Leaves narrowly lanceolate to linear;
3.6–7.1×0.2–0.4 mm, weakly straight or curved,
straight or curved, keeled, little altered when dry;
apices mostly acute, dentate; margins distinctly
bordered, entire below, serrulate and gradually more
strongly serrate above midleaf; costae strongly
spinose on the dorsal side in the upper parts of leaves,
exposed on abaxial surface; hyaline lamina to 1/5–1/3
the length of leaf; hyalocysts in surface view on
abaxial surface at base quadrate to rectangular, at
median leaf irregularly rectangular or quadrate,
14–44×8–22 μm, with rectilinear walls, at apex
oblong to rounded-quadrate. In cross section of
leaf chlorocysts in a central layer, surrounded by
one or more layers of hyalocysts; near leaf base,
hyalocysts adaxially and abaxially in one layer,
median sterome 14–36 cells, marginal sterome
4–16 cells; at midleaf, hyalocysts in 1–2 layer on
26
THAI FOREST BULLETIN (BOTANY) 41
Figure 2. Leucophanes angustifolium A–B. Habit; C. Cross section of stem; D. Leaves; E. Hyalocysts in surface view on abaxial
surface at midleaf; F–H. Cross sections of leaf, F. near apex, G. midleaf, H. near base. All from Chantanaorrapint & Promma 5
(PSU). Photos by C. Promma.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
27
Figure 3. Leucophanes angustifolium: A. Habit; B–C. Leaf, B. adaxial, C. abaxial; D-F. Leaf apex, D. with gemmae, E. adaxial, F.
abaxial; G. Hyalocysts in surface view on abaxial surface at midleaf; H. Cross section of stem; I–K. Cross sections of leaf, I. near
apex, J. midleaf, K. near base. All from Chantanaorrapint & Promma 5 (PSU). Drawn by C. Promma.
28
THAI FOREST BULLETIN (BOTANY) 41
both side, median sterome 20–56 cells, marginal
sterome 7–16 cells; near leaf apex hyalocysts in
two layer on either side of chlorophyllous layer,
median sterome 16–44 cells, marginal sterome 6–14
cells. Gemmae common at the leaf apex. Sporophytes
not seen.
Thailand.— SOUTH-WESTERN: Prachuap Khiri
Khan; SOUTH-EASTERN: Prachin Buri; PENINSULAR:
Krabi, Nakhon Si Thammarat, Satun, Songkhla, Yala.
Distribution.— The Pacific Islands, Australia
(Queenland), New Guinea, Japan (Ryukyu Islands),
SE Asia, India, Sri Lanka, Comoros Islands
(Mayotte), La Réunion, Seychelles (Mahé), tropical
Africa (Salazar Allen 1993).
Habitat and ecology.— On bark at base and
trunk of trees, decaying wood, soil, sandy soil and
rocks, sometime found in association with
L. octoblepharioides. In lowland and montane
forests to ca 1000 m.
2. Leucophanes candidum (Schwägr.) Lindb.,
Oefvers. Förh. Hongl. Svenska Vetensk. Akad. 21:
602. 1865; Enroth, Acta Bot. Fenn. 139:102. Fig.
23a–e. 1990; A. Eddy, Handb. Males. Mosses 2:
42.Fig. 192. 1990; N. Salazar, Bryophyt. Biblioth.
46: 95. Figs. 346–383. 1993.— Syrrhopodon
candidus Schwägr., Spec. Musc. Suppl.1. 2(2): 105.
t. (183)182. Figs. 1–8. 1827. Type: Indonesia,
Reinwardt s.n. (lectotype G, designated by Salazar
Allen (1993); isolectotypes BM, G).— Leucophanes
densifolius Mitt., Bonplandia 9:366.1861; Dixon, J.
Siam Soc., Nat. Hist. Suppl. 9(1): 12. 1932; Tixier,
Ann. Sci. Phom Pehn 4: 91. 1971. Type: Fiji, Milne
s.n. (syntype NY). Figs. 4–5.
Plants medium to moderately large, 1.6–6.2
cm high, growing in dense tufts or cushions. Stems
erect, simple or branched. Leaves crowded, broadly
lanceolate, 2.7–5.5×0.6–1.0 mm, strongly keeled,
recurved, mostly spreading or squarrose sometime
erect, little altered when dry; apices mostly sharply
apiculate; margins bordered, narrow marginal
stereome, entire below, serrulate at least near apex;
costae smooth on the dorsal side throughout the
leaves, exposed on abaxial surface; hyaline lamina
narrow, to 1/8–1/5 the length of leaf; hyalocysts in
surface view on abaxial surface at base rectangular,
short hexagonal to oblong triangular, at median
leaf irregularly pentagonal to hexagonal, oblong
hexagonal with rectilinear walls, 16–44×16–36 μm,
at apex oblong hexagonal. In cross section of leaf
chlorocysts in a central layer, surrounded by one
layer of hyalocysts ; near leaf base hyalocysts
adaxially and abaxially in 1 layer, median sterome
3–15 cells, marginal sterome 4–14 cells; at midleaf
hyalocysts in 1 layer on either side of chlorophyllous
layer, median sterome 6–24 cells, marginal sterome
5–14 cells; near leaf apex hyalocysts in 1 layer on
either side of chlorophyllous layer, median sterome
10–24 cells, marginal sterome 5–14 cells. Gemmae
not seen. Sporophytes 8–10 mm long; setae very
slender, smooth, rugulose just beneath capsule;
capsules cylindrical, 1–2 mm long, erect and
symmetrical; opercula long rostrate; calyptrae
cucullate; peristome teeth of 16 simple teeth,
brownish, articulate, outer surfaces coarsely
papillose.
Thailand.— PENINSULAR: Surat Thani,
Phangnga, Krabi, Nakhon Si Thammarat, Trang,
Satun, Narathiwat.
Distribution.— The Pacific Islands, Australia
(Queenland), New Guinea, Philippines, Malaysia,
Indonesia, Thailand (Salazar Allen 1993).
Habitat and ecology.— On bark of trees and
rocks, in lowland forests to ca 300 m.
3. Leucophanes glaucum (Schwägr.) Mitt., J.
Linn. Soc., Bot. Suppl. 1: 25. 1859; Enroth, Acta
Bot. Fenn. 139: 104. Fig. 24a–f. 1990; A. Eddy,
Handb. Males. Mosses 2: 45. Figs. 193–195. 1990;
N. Salazar, Bryophyt. Biblioth. 46: 101. Figs. 384–
445. 1993.— Syrrhopodon glaucus Schwägr., Spec.
Musc. Suppl. 2(2): 103. 181. 1827. Type: Indonesia.
Moluccas, Gaudichaud 10 (lectotype G; isolectotypes G, L!, BM).— Leucophanes glaucescens
Müll. Hal. ex M. Fleisch., Musci Fl. Buitenzorg. 1:
178. 1904; Broth. in J. Schmidt, Fl. Koh Chang Pt.
3. 1: 116. 1900–1961; Dixon, J. Siam Soc., Nat.
Hist. Suppl. 9(1): 12. 1932; Tixier. Ann. Sci. Phom
Pehn. 4: 91–166. 1971. Type: Indonesia. Java,
Fleischer s.n. (syntype FH); Sumatra, Fleischer
s.n. (syntype FH), Andaman Islands, Kurzz (syntype
FH).— Leucophanes albescens Müll. Hal., Bot.
Zeitung. 22: 347. 1864; Dixon, J. Siam Soc., Nat.
Hist. Suppl. 8(1): 19. 1929; Dixon, J. Siam Soc.,
Nat. Hist. Suppl. 9(1): 12. 1932; Touw, Nat. Hist.
Bull. Siam Soc. 22 (3–4): 228. 1968; Tixier, Nat.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
29
Figure 4. Leucophanes candidum: A–B. Habit; C. Cross section of stem; D. Leaves; E. Hyalocysts in surface view on abaxial surface
at midleaf; F–H. Cross sections of leaf, F. near apex, G. midleaf, H. near base. All from Chantanaorrapint 2442 (PSU). Photos by
C. Promma.
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THAI FOREST BULLETIN (BOTANY) 41
Figure 5. Leucophanes candidum: A. Habit; B–C. Leaf, B. adaxial, C. abaxial; D–E. Leaf apex, D. adaxial, E. abaxial; F. Hyalocysts
in surface view on abaxial surface at midleaf; G. Cross section of stem; H–J. Cross sections of leaf, H. near apex, I. midleaf, J. near
base; K. Capsule; L. Peristome teeth. A., K–L. From Smitinand & Sleumer 1258 (BKF), B-J. From Chantanaorrapint 2442 (PSU).
Drawn by C. Promma.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
Hist. Bull. Siam Soc. 23(4–5): 542. 1970; Thaithong,
J. Hattori Bot. Lab. 56: 86. 1984. Type: Malaysia,
J. Lange s.n. (isotype BM, FH, S). Figs. 6–7.
Plants small to medium, 0.3–1.0 cm high,
growing in rather dense to very dense tufts or
cushions. Stems erect, simple or branched, loosely
foliate below, densely foliate above. Leaves narrowly
lanceolate to linear lanceolate, 1.8–3.3×0.3–0.5 mm,
spreading to curved or nearly erect, sheathing base,
upper parts keeled to channeled, often twisted when
dry; apices mostly acute to apiculate; margins
distinctly bordered, entire below, serrulate and
gradually more strongly serrate above midleaf;
costae strongly spinose on the dorsal side in upper
parts of leaves, exposed on abaxial surface; hyaline
lamina to 1/3–1/2 the length of leaf; hyalocysts in
surface view on abaxial surface at base quadrate to
rectangular, at midleaf 18–52×10–26 μm, oblong
with sinuose walls and concave bulging end wall,
at apex irregularly quadrate oblong. In cross section
of leaff chlorocysts in a central layer, surrounded by
one or more layers of hyalocysts, near leaf base
hyalocysts in 1 layer adaxially and in 2–3 layer
abaxially, median sterome 14–30 cells, marginal
sterome 3–8 cells; at midleaf hyalocysts in 1 layer
on either side of chlorophyllous layer, median
sterome 22–58 cells, marginal 8–18 sterome cells;
near leaf apex hyalocysts in 1 layer on either side
of chlorophyllous layer, median 16–58 sterome
cells, marginal sterome 8–18 cells. Gemmae at leaf
apex. Sporophyte not seen.
Thailand.— NORTH-EASTERN: Loei; EASTERN:
Nakhon Ratchasima; SOUTH-WESTERN: Prachuap
Khiri Khan; SOUTH-EASTERN: Prachin Buri, Trat;
PENINSULAR: Ranong, Surat Thani, Phangnga,
Phuket, Krabi, Nakhon Si Thammarat, Phatthalung,
Trang, Satun, Songkhla.
Distribution.— The Pacific Islands, Australia
(Queenland), New Guinea, Japan, Taiwan, SE Asia,
India, Sri Lanka, Seychelles (Salazar Allen 1993).
Habitat and ecology.— On bark at base and
tree trunks, occasionally on decaying wood, soil
and sandy soil, sometime found in association with
L. octoblepharioides, common in lowland forests,
but also in mangroves, below 750 m.
4. Leucophanes octoblepharioides Brid., Bryol.
Univ. 1: 763. 1827; Enroth, Acta Bot. Fenn. 139:
31
96. Fig. 19a–g. 1990; A. Eddy, Handb. Males.
Mosses 2: 46. Fig. 192. 1990; N. Salazar, Bryophyt.
Biblioth. 46: 62. Figs. 90–112. 1993. Type:
Indonesia, Java, Nees v. Esenbeckk (lectotype B,
designated by Salazar Allen (1993); isolectotype
BM); Nepal, De Candolle (syntype B). Figs. 8–9.
Plants small to medium, 0.5–1.2 cm high,
growing in loose to dense tufts or cushions. Stems
erect, simple or branched. Leaves linear-lanceolate,
2.9–4.8×0.2–0.5 mm, crowded, erect-spreading to
recurved-patent, flat above, more or less keeled
below, little altered when dry; apices mostly obtuse
to acute; margins distinctly bordered, entire below,
serrulate at least near apex; costae smooth on the
dorsal side, rarely spinose at apex, usually exposed
on abaxial surface, rarely covered by abaxial
hyalocysts; hyaline lamina to 1/4–1/3 the length of
leaf; hyalocysts in surface view on abaxial surface
at base rectangular, rhombic-elongated, or quadrate, at midleaf quadrate to irregularly rectangular,
16–44×10–22 μm, with rectilinear walls, at apex
oblong, rounded-quadrate to rectangular. In cross
section of leaff chlorocysts in a central layer, surrounded by one layer of hyalocysts, near leaf base
hyalocysts adaxially and abaxially in 1 layer, median
sterome 10–20 cells, marginal sterome 3–8 cells; at
midleaf hyalocysts in 1 layer on either side, median
sterome 16–38 cells, marginal sterome 8–16 cells;
near leaf apex hyalocysts in 1 layer on either side,
median sterome 16–30 cells, marginal sterome 6–14
cells. Gemmae common at leaf apex. Sporophytes
not seen.
Thailand:— NORTH-EASTERN: Loei; EASTERN:
Nakhon Ratchasima; SOUTH-WESTERN: Prachuap
Khiri Khan; SOUTH-EASTERN: Prachin Buri,
Chanthaburi, Trat; PENINSULAR: Ranong, Phuket,
Krabi, Nakhon Si Thammarat, Phatthalung, Trang,
Satun, Songkhla, Yala.
Distribution.— The Pacific Islands, Australia
(Queenland), New Guinea, Japan (Ryukyu Islands),
Taiwan, SE Asia, India, Sri Lanka, Mauritus,
Seychelles (Salazar Allen 1993).
Habitat and ecology.— On bark at base and
tree trunks, roots, trunks of tree fern, decaying
wood, soil, and sandy soil, sometime found in
association with L. glaucum and L. angustifolium.
Common in lowland and montane forests to ca
1000 m.
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THAI FOREST BULLETIN (BOTANY) 41
Figure 6. Leucophanes glaucum: A–B. Habit; C. Cross section of stem; D. Leaves; E. Hyalocysts in surface view on abaxial surface
at midleaf; F–H. Cross sections of leaf, F. near apex, G. midleaf, H. near base. All from Chantanaorrapint & Promma 1 (PSU).
Photos by C. Promma.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
33
Figure 7. Leucophanes glaucum: A. Habit; B–E. Leaf, B. adaxial, C. abaxial, D.lateral, E. when dry; F–G. Leaf apex, F. adaxial, G.
abaxial; H. Hyalocysts in surface view on abaxial surface at midleaf; I. Cross section of stem; J–L. Cross sections of leaf, J. near
apex, K. midleaf, L. near base. All from Chantanaorrapintt & Promma 1 (PSU). Drawn by C. Promma.
34
THAI FOREST BULLETIN (BOTANY) 41
Figure 8. Leucophanes octoblepharioides: A–B. Habit; C. Cross section of stem; D. Leaves; E. Hyalocysts in surface view on abaxial
surface at midleaf; F–H. Cross sections of leaf, F. near apex, G. midleaf, H. near base. All from Chantanaorrapint 2353 (PSU).
Photos by C. Promma.
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
35
Figure 9. Leucophanes octoblepharioides: A. Habit; B–D. Leaf, B. adaxial, C. abaxial, D. lateral; E–G. Leaf apices, E. with gemmae,
F. adaxial, G. abaxial; H. Hyalocysts in surface view on abaxial surface at midleaf; I. Cross section of stem; J–M. Cross sections of
leaf, J–K. near apex, L. midleaf, M. near base. All from Chantanaorrapint 2353 (PSU). Drawn by C. Promma.
36
THAI FOREST BULLETIN (BOTANY) 41
Note.— Salazar Allen (1993) had divided L.
octoblepharioides into three subspecies, based on
leaf structures, peristome and spore ornamentation.
She reported 2 subspecies, L. octoblepharioides
subsp. octoblepharioides and L. octoblepharioides
subsp. meijeri Salazar Allen, in Thailand. The first
subspecies is rather common in this country, while
the later is known only from a single collection
from Khao Yai national park (Touw 123966 (BKF,
BM, L)). According to Salazar Allen (1993), subsp.
meijeri differs from subsp. octoblepharioides in leaf
costa completely covered by hyalocysts on the abaxial side from base to apex. However, an investigation of Touw 123966 (sterile specimens) revealed
that a leaf costa in selected leaves of the same plant
has no hyalocysts on the abaxial surface. As this
character may vary even in the same plant and has
not been frequently recorded from other Thai
collections, we are hesitant to assign Thai material
to subspecific level due to the lack of collections with
sporophytes.
ACKNOWLEDGEMENTS
The authors would like to thank Assoc. Prof.
Dr Obchant Thaithong (BCU) and Assoc. Prof. Dr
Kitichate Sridith (PSU) for their valuable comments
on the first draft of the manuscript. Sincere thank to
the anonymous readers who reviewed this manuscript.
Also to the curators and staffs of BCU, BKF and L.
This work was supported by the Faculty of Science,
Prince of Songkla University. The first author
would like to express her sincere appreciation to
the Development and Promotion of Science and
Technology Talent Project (DPST) for funding.
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SPECIMENS EXAMINED
Leucophanes angustifolium Renauld & Cardot
Promma 7, 11, 23 July 2011, Nakhon Si Thammarat,
Ron Phibun, Khao Ram Rome (PSU);
Chantanaorrapint & Promma 5, 22 Oct. 2011,
Krabi, Klong Thom, Sra Morakot (PSU);
Chantanaorrapint 1302, 24 Dec. 2006, Nakhon Si
Thammarat, Nopphitam, Khao Nan National Park,
Klong Kan Station (PSU); Chantanaorrapint 1409,
22 Feb. 2007, Nakhon Si Thammarat, Ron Phibun,
Khao Ram Rome (PSU); Chantanaorrapint 1453,
1457, 17 April 2007, Nakhon Si Thammarat,
Nopphitam, Khao Nan National Park (PSU);
Chantanaorrapint 2369B, 16 May 2011, Prachuap
Khiri Khan, Thap Sakae, Huai Yang, Khao Luang
(PSU); Chantanaorrapint 2437, 28 May 2011,
Yala, Betong, Ban Chulabhorn Pattana 10 (PSU);
THE GENUS LEUCOPHANES (CALYMPERACEAE, BRYOPHYTA) IN THAILAND (C. PROMMA & S. CHANTANAORRAPINT)
Touw 11414, 11498, 2 Feb. 1966, Nakhon Si
Thammarat, granitic massive Khao Luang (BKF,
L); Chantanaorrapint, Inuthai & Promma 499, 19
Nov. 2011, Prachin Buri, Khao Yai National Park,
(PSU); Chantanaorrapint, Inuthai & Promma 606,
4 Dec. 2011, Yala, Betong, Ban Chulabhorn Pattana
10 (PSU); Chantanaorrapint, Inuthai & Promma
646, 5 Dec. 2011, Yala, Betong, Ban Piyamit 2
(PSU); Chantanaorrapint, Inuthai & Promma 732,
14 Dec. 2011, Songkhla, Khlong Hoi Khong, Pa
Dum Waterfall (PSU); Chantanaorrapint, Inuthai
& Promma 771, 16 Dec. 2011, Satun, Thungwa,
Than Plio Waterfall (PSU).
Leucophanes candidum (Schwägr.) Lindb.
Smitinand & Sleumerr 1258, 24 Sept. 1963, Surat,
Khao Na Daeng (BKF); Chantanaorrapint 2348,
17 Feb. 2011, Phang Nga, Thai Muang, Khao Lam
Pee-Had Thai Muang National Park (PSU);
Chantanaorrapint 2442, 10 Aug. 2011, Satun,
Thungwa, Than Plio Waterfall (BKF, PSU);
Chantanaorrapint, Inuthai & Promma 772, 16
Dec. 2011, Satun, Thungwa, Than Plio Waterfall
(BKF, PSU); Chantanaorrapint, Inuthai & Promma
797, 798, 11 Jan. 2012, Trang, Palian, Chao Pa
Waterfall (BKF, PSU); Chantanaorrapint, Inuthai
& Promma 1800, 9 Dec. 2012, Nakhon Si
Thammarat, Nopphitam, Krung Ching Waterfall
(PSU).
Leucophanes glaucum (Schwägr.) Mitt.
Promma 2, 3, 22 July 2011, Nakhon Si Thammarat,
Thung Song, Nam Tok Yong National Park (PSU);
Promma 13, 14, 16, 2 Aug. 2011, Songkhla, Hat
Yai, Ton Nga Chang Wildlife Sanctuary, Ton Nga
Chang Waterfall (PSU); Promma 17B, 13 Aug.
2011, Krabi, Klong Thom, Sra Morakot (PSU);
Inuthai 732, 28 Sept. 2011, Satun, Thungwa, Than
Plio Waterfall (PSU); Inuthai 747,750, 23 Oct.
2011, Trang, Yan Ta Khao, Peninsular Botanic
Garden (Thung Khai) (PSU); Chantanaorrapint &
Promma 1, 22 Oct. 2011, Krabi, Klong Thom, Sra
Morakot (BKF, PSU); Chantanaorrapint &
Promma 19, 20, 23, 23 Oct. 2011, Trang, Na Yong,
Khao Chong Wildlife Conservation Development
and Extension Center (BKF, PSU); Thaithong
37
1004, 25 April 1979, Ranong, Ban-Dan (BCU);
Thaithong 1124, 26 April 1979, Phangnga, mangrove forest site 3 (BCU); Boonkerd 11, 24 April
1980, Phangnga, on the way to Rue-sri Sawan cave
(BCU); Porn-sookk sawang 5, 5 May 1981, Ranong,
Ka-pure (BCU); Porn-sookk sawang 50-51, sd.,
Satun, Tarutao Island, Ta-lo-wow Bay (BCU);
Touw 11129, 11145, 11147, 24 Jan. 1966, Phuket
Island, near Thalang, Khao Kluai (BKF, L); Touw
11334, 29 Jan. 1966, Mainland Phuket, near Tong
Lang village (BKF, L); Touw 12288, 18 Feb. 1966,
Prachin Buri, Khao Yai National Park (BKF, L);
Chantanaorrapint, Inuthai & Promma 471, 19 Nov.
2011, Prachin Buri, Khao Yai National Park (PSU);
Chantanaorrapint, Inuthai & Promma 503B, 19
Nov. 2011, Nakhon Ratchasima, Khao Yai National
Park, Pha Kluai Mai Waterfall (PSU);
Chantanaorrapint, Inuthai & Promma 697, 13
Dec. 2011, Phatthalung, Tamot, Lan Mom Jui
Waterfall (BKF, PSU); Chantanaorrapint, Inuthai
& Promma 718, 721, 13 Dec. 2011, Phatthalung,
Kong La, Manora Waterfall (PSU).
Leucophanes octoblepharioides Brid.
Promma 1, 22 July 2011, Nakhon Si Thammarat,
Tung Song, Nam Tok Yong National Park (PSU);
Promma 5, 6, 8, 9, 10, 23 July 2011, Nakhon Si
Thammarat, Ron Phibun, Khao Ram Rome (PSU);
Promma 15, 2 Aug. 2011, Songkhla, Hat Yai,Ton
Nga Chang Wildlife Sanctuary, Ton Nga Chang
Waterfall (PSU); Promma 17A, 13 Aug. 2011,
Krabi, Klong Thom, Sra Morakot (PSU);
Chantanaorrapint 1465, 18 April 2007, Nakhon Si
Thammarat, Nopphitam, Khao Nan National Park,
San yen (PSU); Chantanaorrapint 1588, 20 April
2007, Nakhon Si Thammarat, Nopphitam, Khao
Nan National Park, San yen (PSU);
Chantanaorrapint 2353, 2354, 22 April 2011,
Satun, La Ngu, Tarutao National Park, Tarutao
Island (PSU); Chantanaorrapint 2361, 2369A, 16
May 2011, Prachuap Khiri Khan, Thap Sakae,
Huai Yang, Khao Luang (PSU); Chantanaorrapint
2436, 28 May 2011, Yala, Betong, Ban Chulabhorn
Pattana 10 (PSU); Thaithong s.n., 24 March 1975,
Satun, La Ngu, Tarutao National Park, Tarutao
Island (BCU); Touw 11196, 24 Jan. 1966, Phuket
Island, near Thalang, Khao Kluai (BKF, L); Touw
38
THAI FOREST BULLETIN (BOTANY) 41
11212, 11230, 25 Jan. 1966, Phuket Island, near
Thalang, Khao Phara (BKF, L); Touw 12396, 19
Feb. 1966, Prachin Buri, Khao Yai National Park,
Kluai Mai Waterfall (BKF, L); Touw 12334, 18
Feb. 1966, Prachin Buri, Khao Yai National Park
(BKF, L); Phloechit 2630, sd., Nakhon Si
Thammarat, Khiriwong, Thap Chang (BKF);
Chantanaorrapint, Inuthai & Promma 493, 19
Nov. 2011, Prachin Buri, Khao Yai National Park
(PSU); Chantanaorrapint, Inuthai & Promma 502,
503A, 19 Nov. 2011, Nakhon Ratchasima, Khao
Yai National Park, Pa Kluai Mai Waterfall (PSU);
Chantanaorrapint, Inuthai & Promma 519, 520, 531,
541, 1 Dec. 2011, Songkhla, Khlong Hoi Khong,
Pa Dum Waterfall (BKF, PSU); Chantanaorrapint,
Inuthai & Promma 629, 4 Dec. 2011, Yala, Betong,
Ban Chulabhorn Pattana 10 (PSU);Chantanaorrapint,
Inuthai & Promma 649, 653, 667, 4 Dec. 2011, Yala,
Betong, Ban Piyamit 2 (PSU); Chantanaorrapint,
Inuthai & Promma 716, 13 Dec. 2011, Phatthalung,
Kong La, Manora Waterfall (BKF, PSU);
Chantanaorrapint, Inuthai & Promma 731, 14
Dec. 2011, Songkhla, Khlong Hoi Khong, Pa Dum
Waterfall (BKF, PSU).