Notes on Crepidium (Orchidaceae): Two new combinations, a putative natural hybrid, and four species newly recorded for Thailand

Abstract

In this paper we report various findings from our ongoing studies of Crepidium (Orchidaceae), mainly conducted in preparation of the Flora of Thailand account on the genus: (1) the new combinations C. chamaeorchis (Schltr.) Nuammee, Seelanan, Suddee & H.A.Pedersen and C. szemaoense (Tang & F.T.Wang) Nuammee, Seelanan, Suddee & H.A.Pedersen are made and substantiated; (2) a putative natural hybrid between C. acuminatum (D.Don) Szlach. and C. polyodon (Hook.f.) Szlach. is discussed and highlighted as being the first convincing case of natural hybridization reported for Crepidium; (3) C. bahanense (Hand.-Mazz.) S.C. Chen & J.J. Wood, C. josephianum (Rchb.f.) Marg., C. maximowiczianum (King & Pantl.) Szlach. and C. micranthum (Hook.f.) Szlach. are reported as new national records for Thailand; (4) the name C. josephianum (Rchb.f.) Marg. is lectotypified. All taxa treated in this paper are documented by photos of live plants from their natural habitat in Thailand, and for each newly recorded species we provide a morphological description based on Thai material.

Full text

THAI FOREST BU LL., BOT. 44(1): 35–44. 2016.
DOI: 10.20531/T FB.2016.44.1.08
© 2016 The Forest Herbarium
INTRODUCTION
Seidenfaden (1978) recognized 22 species of
Malaxis Sol. ex Sw. (Orchidaceae) as occurring in
Thailand. Largely adopting the revised generic
classification
of the Malaxideae proposed by
Szlachetko (1995), he later assigned the same pool
of species to Crepidium Blume (19 species), Dienia
Lindl. (1), Glossochilopsis Szlach. (1) and
Oberonioides
Szlach. (1) (Seidenfaden, 1997). These genera are all
accepted in Genera Orchidacearum (Pridgeon et al.,
2005) –
except
Glossochilopsis
which is
considered
congeneric
with the earlier described Crepidium.
In Flora of Thailand, Crepidium is going to be
delimited according to Pridgeon et al. (2005), and in
connection
with our
ongoing
revision of
this
genus
for the Flora, we have
realized
that two new
combina-
tions
need to be published. Furthermore, recent eld
Notes on Crepidium (Orchidaceae): two new combinations, a putative natural hybrid,
and four species newly recorded for Thailand
ANCHALEE NUAMMEE1, TOSAK SEELANAN1,*, SOMRAN SUDDEE2 & HENRIK Æ. PEDERSEN3,*
ABSTRACT
In this paper we report various ndings from our ongoing studies of Crepidium (Orchidaceae), mainly conducted in preparation of the
Flora of Thailand account on the genus: (1) the new combinations C. chamaeorchis (Schltr.) Nuammee, Seelanan, Suddee & H.A.Pedersen
and C. szemaoense (Tang & F.T.Wang) Nuammee, Seelanan, Suddee & H.A.Pedersen are made and substantiated; (2) a putative natural
hybrid
between
C.
acuminatum
(D.Don) Szlach. and C. polyodon (Hook.f.) Szlach. is
discussed
and
highlighted
as being the rst
convincing
case of natural hybridization reported for Crepidium; (3) C. bahanense (Hand.-Mazz.) S.C. Chen & J.J. Wood, C. josephianum (Rchb.f.)
Marg., C. maximowiczianum (King & Pantl.) Szlach. and C. micranthum (Hook.f.) Szlach. are reported as new national records for
Thailand; (4) the name C. josephianum (Rchb.f.) Marg. is lectotypied. All taxa treated in this paper are documented by photos of live
plants from their natural habitat in Thailand, and for each newly recorded species we provide a morphological description based on Thai
material.
KEYWORDS: Crepidium acuminatum × polyodon, Crepidium bahanense, Crepidium chamaeorchis, Crepidium josephianum,
Crepidium maximowiczianum, Crepidium micranthum, Crepidium szemaoense.
Published online: 28 September 2016
1 Plants of Thailand Research Unit, Department of Botany, Faculty of Science, Chulalongkorn University, Pathumwan, Bangkok
10330, Thailand.
2 The Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand.
3 Natural History Museum of Denmark, University of Copenhagen, Øster Farimagsgade 2C, DK-1353 Copenhagen K, Denmark.
* Corresponding authors: Tosak Seelanan (tosak.s@chula.ac.th), Henrik Æ. Pedersen (henrikp@snm.ku.dk)
inventories have revealed the occurrence in Thailand
of four species not previously recorded for this
country, as well as a putative natural hybrid.
This paper is mainly based on examination of
herbarium specimens (collections from Thailand as
well as type specimens deposited in foreign herbaria)
combined with consultation of the literature and of
digital images of type specimens accessed online.
Based entirely on Thai material, we have prepared
a comprehensive morphological description of each
of the four newly recorded species; and all taxa
treated in this paper are documented by photos of
live plants from their natural habitat in Thailand. All
examined material is cited explicitly in the individual
accounts below.

thai forest bulletin (botany) Vol. 44 no. 1
36
NEW COMBINATIONS
Crepidium
chamaeorchis
(
Schltr
.)
Nuammee
,
Seelanan, Suddee & H.A.Pedersen, comb. nov.—
Microstylis chamaeorchis Schltr., Beibl. Bot. Jahrb.
Syst. 104: 11. 1911.— Malaxis
chamaeorchis
(Schltr.)
Seidenf., Bot. Tidsskr. 65: 316. 1970.—
Glossochilopsis
chamaeorchis (Schltr.) Szlach., Fragm. Florist.
Geobot., Suppl. 3: 123. 1995. Type: Indonesia, West
Sumatra, Gunung Marapi, 24 Jan. 1907, Schlechter
15942 (holotype B, probably destroyed; isotypes
AMES, C!, K!, L!, NSW, P). Fig. 1A–B.
Note.— Glossochilopsis chamaeorchis (Schltr.)
Szlach. (basionym: Microstylis chamaeorchis
Schltr.) is the type species of Glossochilopsis Szlach.
However, the only feature that consistently separates
G. chamaeorchis (and Seidena Szlach.) from all
species of Crepidium Blume s.s. is the lip not being
provided with auricles (Fig. 1); and it remains to be
demonstrated that G. chamaeorchis and Seidena are
not nested in Crepidium s.s. Against this background,
we have decided to follow Pridgeon et al. (2005) in
treating Glossochilopsis and Seidena as synonyms
of Crepidium, and we here provide the necessary
combination under Crepidium. An account on the
occurrence of this species in Thailand can be found
in Seidenfaden (1978: 48–49).
Crepidium szemaoense (Tang & F.T.
Wang
)
Nuammee
,
Seelanan, Suddee & H.A.Pedersen, comb. nov.—
Malaxis
szemaoensis
Tang & F.T.
Wang
,
Acta
Phytotax
.
Sin. 1: 75. 1951. Type: China, Yunnan, Szemao,
“NW Mts”, ca 1525 m, 1901, Henry 13128 (
holotype
K!). Fig. 1C–D.
Notes.— In Flora of China, Chen & Wood
(2009) placed Malaxis szemaoensis in the synonymy
of Crepidium ovalisepalum (J.J.Sm.) Szlach. – a
species originally described as Microstylis ovalisepala
J.J.Sm. from Sumatra (Smith 1928). Only the type
collection of the latter is known from Sumatra [West
Sumatra, Gunung Kuriman, 1000 m, 24 June 1918,
Bünnemeijer 3315 (holotype L!)]; but when publishing
the new combination Malaxis ovalisepala (J.J.Sm.)
Seidenf., Seidenfaden (1978) also referred two col-
lections
from N Thailand, Chiang Mai province,
Doi
Chiang Dao to this species.
Based on comparison of the type specimens
of Microstylis ovalisepala and Malaxis szemaoensis,
and of the descriptions in Smith (1928), Tang & Wang
(1951) and Chen & Wood (2009), we are convinced
that we are dealing with two distinct species that
mainly dier in vegetative characters. Thus, Microstylis
ovalisepala is characterized by a creeping rhizome
and an elongated ascending stem bearing ca 10 foliage
leaves, whereas Malaxis szemaoensis has a short
pseudobulbous stem bearing 2–4 leaves. These dier-
ences
mean that Microstylis ovalisepala probably
shows the same annual stem/rhizome dynamics as
described for Crepidium micranthum below (under
“Ecology”), whereas Malaxis szemaoensis matches
the majority of Crepidium species in producing a new
replacement pseudobulb annually through sympodial
growth (cf. the descriptions of C. bahanense, C.
josephianum and C. maximowiczianum below). Against
this background, we here provide the new combination
Crepidium szemaoense.
From Thailand we have studied the two collec-
tions
that Seidenfaden (1978) assigned to Malaxis
ovalisepala as well as more recently collected material
belonging to the complex [NORTHERN: Chiang Mai
province, Doi Chiang Dao, June 1958, Seidenfaden &
Smitinand GT 2643 (C!); 15 July 1958, Smitinand
4693 (BKF!, C!), 20 June 2014, Nuammee 404
(BCU!, BKF!), Nuammee 408 (BCU!, BKF!); Nan
province, Tham Sakoen National Park, 28 July 2011,
La-ongsri et al. 1878 (QBG!)]. All the Thai material
clearly matches Crepidium szemaoensis (Fig.
1C–D).
PUTATIVE NATURAL HYBRID
Crepidium
acuminatum
(D.Don) Szlach.
×
polyodon
(Hook.f.) Szlach.
Crepidium acuminatum (Szlachetko, 1995),
based on Malaxis acuminata D.Don, and C. polyodon
(Szlachetko, 1995), based on Microstylis polyodon
Hook.f., dier in a number of oral characters. Thus,
C. acuminatum has larger owers, fewer teeth in the
distal part of the lip and narrower and dierently
shaped lip auricles – see Table 1 and Figure 2 (A, C).
They both ower in the rainy season, but their pollina-
tion
biology is unknown.
In a mixed colony of the two species at Huai
Yang Waterfall National Park in Prachuap Khiri
Khan province (SW Thailand), the rst author came
across a plant (voucher: 8 May 2015, Nuammee 439

Notes oN Crepidium (orChidaCeae): two New CombiNatioNs, a putative Natural hybrid,
aNd four speCies Newly reCorded for thailaNd (a. Nuammee, t. seelaNaN, s. suddee & h.Æ. pederseN) 37
BCU) that appeared largely intermediary (Table 1;
Fig. 2B). Thus, it had intermediate auricle shape and
an intermediate number of teeth in the distal part of
the lip, whereas ower diameter and auricle width
were within the lower part of the range recorded for
C. acuminatum. Referring to the general trend of
orchid hybrids to be morphologically intermediary
between their parental taxa (e.g. Nilsson, 1985;
Aagaard et al., 2005; Hedrén et al., 2012), we hy-
pothesize that the plant vouchered as Nuammee 439
is a natural hybrid between C. acuminatum and C.
polyodon. At the same time, however, we recognize
that comparative molecular data would be needed
to test our hypothesis (cf. Rieseberg & Carney,
1998).
Although we cannot definitely confirm the
hybrid nature of Nuammee 439, we consider it relevant
to draw attention to this putative case of natural
hybridization in Crepidium in Thailand. Firstly, we
believe that there are no previous reports of natural
interspecic hybrids in Crepidium from Thailand or
elsewhere. Secondly, there is a surprising lack of
records of plant hybrids from Thailand in general, for
which reason a more systematic search was recently
encouraged (Parnell et al., 2013).
Figure 1. A–B: Crepidium chamaeorchis (Schltr.) Nuammee, Seelanan, Suddee & H.A.Pedersen, Nakhon Ratchasima province, Khao
Yai, 11 July 2015 (A: ower in front view, B: ower in side view); C–D: Crepidium szemaoense (Tang & F.T.Wang) Nuammee,
Seelanan, Suddee & H.A.Pedersen (C: habit, D: owers). Photos: A. Nuammee.

thai forest bulletin (botany) Vol. 44 no. 1
38
NEW RECORDS FOR THAILAND
Crepidium bahanense (Hand.-Mazz.) S.C.Chen &
J.J.Wood in Z. Wu et al. (eds), Fl. China 25: 232.
2009.— Microstylis bahanensis Hand.-Mazz.,
Symb. Sin. 7: 1350. 1936.— Malaxis bahanensis
(Hand.-Mazz.) Tang & F.T.Wang. Acta Phytotax.
Sin. 1: 71. 1951. Type: China, Yunnan, sine loco,
21 July 1916, Handel-Mazzetti 9574 [holotype WU
(http://herbarium.univie.ac.at/database/detail.php;
accessed 13 March 2016)]. Fig. 3A–B.
Terrestrial, sympodial herb with a strongly
condensed rhizome carrying a fresh, developing
corm distally (from which the aerial shoot is pro-
duced) and 1–2 decaying corms immediately behind;
owering shoots 10–20 cm tall. Roots 3–5, arising
from the basal part of the pseudobulb, 0.5–1 cm
long. Pseudobulbs white, subglobose, 0.6–1.1 cm
Character C. acuminatum1Putative hybrid2C. polyodon3
Width of lip (mm) 4.0–9.5 5.5–6.0 2.5–4.0
Teeth on distal part of lip 2 4–5 12–16
Shape of lip auricles Obliquely triangular
to oblong-ovate
Obliquely
triangular-oblong
Falcately
linear-triangular
Width of lip auricles (mm) 1.5–4.0 2.5 1.0–2.0
Figure 2. A: Crepidium acuminatum (D. Don) Szlach., Prachuap Khiri Khan province, Huai Yang Waterfall National Park, 8 May
2015 (owers); B: Crepidium cf. acuminatum (D. Don) Szlach. × polyodon (Hook.f.) Szlach., Prachuap Khiri Khan province, Huai
Yang Waterfall National Park, 18 May 2015 (ower); C: Crepidium polyodon (Hook.f.) Szlach., Prachuap Khiri Khan province, Huai
Yang Waterfall National Park, 18 May 2015 (ower). Photos: A. Nuammee.
1 Specimens examined: van Beusekom & Phengklai 1282 (L); Garrett 404 (K); Geesink et al. 5896 (L); Kerr 100 (K, 4 specimens),
437 (K), 445 (K), 872 (K); Koyama T-61151 (BKF); Maxwell 88-976 (L); Nuammee 377 (BCU, BKF), 398 (BCU), 449 (BCU),
438 (BCU, BKF); Palee 233 (BKF); Seidenfaden & Smitinand GT 2595 (C).
2 Specimen examined: Nuammee 439 (BCU).
3 Specimens examined: Geesink & Santisuk 5159 (C, L); Kerr 457 (K, 2 specimens); Nuammee 437 (BCU, BKF).
Table 1. Comparison of Crepidium acuminatum (D.Don) Szlach., C. polyodon (Hook.f.) Szlach. and
their putative natural hybrid. All data were scored from material collected in Thailand.

Notes oN Crepidium (orChidaCeae): two New CombiNatioNs, a putative Natural hybrid,
aNd four speCies Newly reCorded for thailaNd (a. Nuammee, t. seelaNaN, s. suddee & h.Æ. pederseN) 39
long, 0.5–1.5 cm in diameter when fully developed,
consisting of 2–4 internodes. Cataphylls few, oblong
to oblong-lanceolate, subacute to acute, 0.8–3 by
0.4–0.7 cm. Foliage leaves 2(–3), sessile, sheathing
at base; lamina green, lanceolate-oblong to ovate-
oblong or ovate with symmetrical base, obtuse to
acute, sometimes slightly apiculate, 3–7.5 by 1–2.5
cm, 1- to 3-veined, with numerous scattered white
papillae on the adaxial side when young (later gla-
brous), margins entire, sometimes slightly undulate.
Inorescence racemose, erect, 9–17 cm long; peduncle
brownish purple, greenish brown or green, 6–9 cm
long, ca 0.1 cm in diameter; rachis laxly 10- to 30-
owered; oral bracts creamy-yellow to yellowish
green, reexed, (linear-)triangular, acute to acuminate,
2–5.5 by 1.1–1.2 mm, equal to or longer than ovaries.
Flowers creamy-yellow with more or less purple lip
mid-lobe, 4–4.5 mm in diameter. Sepals rounded to
obtuse, glabrous, margins revolute; dorsal sepal
ovate to elliptic, ca 2.5 by ca 1.5 mm, 3-veined;
lateral sepals concave, obliquely ovate, ca 2.5 by
1.8–2 mm, 4-veined. Petals revolute, linear-ligulate,
truncate, 2–2.2 by 0.5–0.6 mm, 1-veined, margins
recurved
to
revolute
.
Lip
ca 4.5 by 2.5–3 mm (
maximum
dimensions), oblong with a widened, strongly sagittate
-auriculate base, apically bilobed with obtuse to
retuse lobes, basal part forming a cavity that is
broadly ovate to suborbicular in outline and bordered
by a more or less horseshoe-shaped, distally prolonged,
glabrous callus; auricles obliquely triangular to ovate,
obtuse, ca 1.5 by ca 1 mm. Gynostemium yellow,
erect, stout, 1–1.5 mm long, ca 1 mm in diameter;
staminodes eshy, oblong, truncate; anther orange-
yellow, broadly ovate in upper view, ca 0.3 mm long,
pollinia 4 in 2 pairs, obliquely clavate, without
caudiculae; rostellum transversely rectangular,
truncate; fertile part of stigma small. Ovary (including
pedicel) greenish yellow,
fusiform
-
cylindrical
, with 6
longitudinal ridges, 2–2.5 mm long, glabrous. Capsule
ellipsoid, 4–5 cm long, 0.2–0.3 cm in diameter; fruit
pedicel 0.2–0.3 cm long.
Thailand.— NORTH-EASTERN: Loei [Phu Ruea
National Park, trail to Suan Hin Phali, 20 July 2013,
Nuammee 379 (BKF!), Nuammee 380 (BCU!, BKF!),
Nuammee 383 (BCU!);
4 May 2014
[in bud], Nuammee
397
(
BKF!
); 2
July
2014,
Nuammee
415
(
BCU
!)].
Distribution.— China (Yunnan).
Vernacular
.
— Hu suea chio bai tang ()
(here proposed).
Ecology.—
Crepidium
bahanense grows in sandy
soil along streams and shallow moist depressions in
open grassland. It owers in the rainy season (June
to July) and survives the dry season as a leafless
corm.
Note.— During field surveys in Phu Ruea
National Park, a large population of this species was
discovered along the trail to Suan Hin Phali.
Crepidium josephianum (Rchb.f.) Marg., Ann.
Bot. Fenn. 39: 65. 2002.— Microstylis josephiana
Rchb.f., Bot. Mag. 103: t. 6325. 1877.— Malaxis
josephiana (Rchb.f.) Kuntze, Revis. Gen. Pl. 2: 673.
1891. Type: Drawing by W. Fitch from a plant sent
by Gammie from Sikkim (India) and owering in
Royal Bot. Gard., Kew (lectotype in the library of
K!, designated here; see reproduction in Bot. Mag.
103: t. 6325. 1877). Fig. 3C–D.
Terrestrial, sympodial herb with a strongly
condensed rhizome carrying a fresh, developing
pseudobulb distally (from which the aerial shoot is
produced) and 1 decaying pseudobulb immediately
behind; flowering shoots up to 15 cm tall. Roots
several, arising from the basal part of the pseudobulb,
1–3 cm long. Pseudobulbs green, oblong-fusiform, 5–6
cm long, 1–1.5 cm in diameter when fully developed,
consisting
of
4–5
internodes
.
Cataphylls
2–3,
triangular
-ovate to lanceolate-oblong (the lowermost tubular),
acute 1.5–5 by 0.8–1 cm. Foliage leaves 3–5, sessile,
sheathing at base; lamina pale purplish brown,
purple-veined beneath, ovate to elliptic-
lanceolate
with slightly asymmetrical base, acuminate,
6–7 by
3.3–3.8 cm, 5- to 6-veined,
glabrous
,
margins
undulate
.
Inorescence racemose, erect, 8–10 cm long; peduncle
purplish brown, 5–6 cm long, 0.2–0.3 cm in diameter,
with 6 longitudinal ridges at base;
rachis densely
6- to 12-owered; oral bracts purplish
brown, patent
to reexed, triangular to triangular-ovate, acuminate,
3.5–4 by 1.5–2 mm, shorter than ovaries. Flowers
light brownish yellow with red-brown or purplish
red
markings
on the
basal
part of the lip, 10–12 mm in
diameter. Sepals obtuse, glabrous, margins recurved;
dorsal sepal revolute, ovate, 10–12 by 4–5.5 mm,
4-veined; lateral sepals broadly elliptic, recurved,

thai forest bulletin (botany) Vol. 44 no. 1
40
Figure 3. A–B: Crepidium bahanense (Hand.-Mazz.) S.C.Chen & J.J.Wood, Loei province, Phu Ruea National Park, 20 June 2013
(A: habit, B: owers); C–D: Crepidium josephianum (Rchb.f.) Marg., Loei province, Phu Luang Wildlife Sanctuary, 1 April 2015
(C: habit, D: owers). Photos: A. Nuammee.

Notes oN Crepidium (orChidaCeae): two New CombiNatioNs, a putative Natural hybrid,
aNd four speCies Newly reCorded for thailaNd (a. Nuammee, t. seelaNaN, s. suddee & h.Æ. pederseN) 41
10–11 mm by 6–7 mm, 5-veined. Petals linear, rounded,
10–11 by ca 2 mm, 3-veined, margins revolute.
Lip
12–14 by 14–16.5 mm
(maximum dimensions), bowl-
shaped with a
strongly cordate-auriculate base,
apically emarginate;
basal part forming a cavity that
is narrowly rectangular in outline, with 2 chambers,
small; auricles broadly and obliquely elliptic-oblong,
rounded, 4–5 mm by 6–7 mm. Gynostemium green,
erect, very stout, ca 2 mm long, ca 2.2 mm in diameter;
staminodes eshy, oblong, truncate; anther orange-
yellow, broadly ovate in upper view, ca 1 mm long;
pollinia 4 in 2 pairs, obliquely clavate, without
caudiculae; rostellum transversely rectangular,
truncate; fertile part of stigma small. Ovary (including
pedicel)
greenish
cream, tinged with purple, cylindrical,
with 6 longitudinal ridges, 3–5 mm long, glabrous.
Capsule not seen.
Thailand.— NORTH-EASTERN: Loei [Phu
Luang Wildlife Sanctuary, near Khok Nokkaba
Forest Protection Unit, 1 Apr. 2015 [in bud], Nuammee
435 (BCU!, BKF!)].
Distribution.— Nepal, NE India.
Vernacular.— Haeo mu pa sikkhim (
)(here proposed).
Ecology.— Crepidium josephianum grows in
humus-rich soil in hill evergreen forest. It owers
early in the rainy season (May) and survives the dry
season as a leaess pseudobulb.
Note.— During eld surveys in Phu Luang
Wildlife Sanctuary, a small population of this species
was discovered in a shaded area near Khok Nokkaba
Forest Protection Unit.
Crepidium maximowiczianum (King & Pantl.)
Szlach., Fragm. Florist. Geobot., Suppl. 3: 129.
1995.— Microstylis maximowicziana King & Pantl.,
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 64: 329.
1895.— Malaxis maximowicziana (King & Pantl.)
Tang & F.T.Wang, Acta Phytotax. Sin. 1: 72. 1951.
Type: India, West Bengal, Mungpoo Cinchona
Plantation, July 1892, Pantling 226 [holotype BM
(http://data.nhm.ac.uk/
dataset
/collection
-specimens;
accessed 13 March 2016), isotype K!]. Fig. 4A–B.
Terrestrial, sympodial herb with a strongly
condensed rhizome carrying a fresh, developing
pseudobulb distally (from which the aerial shoot is
produced) and 1 decaying pseudobulb immediately
behind; owering shoots up to 30 cm tall, robust.
Roots several, arising from the basal part of the
pseudobulb, 1–7 cm long. Pseudobulbs green, terete
from a fusiform base, ca 10 cm long, 0.7–0.8 cm in
diameter, consisting of 5–7 internodes. Cataphylls
2–3, lanceolate-oblong (the lowermost tubular),
acute to acuminate, 3–7 by 0.7–1.4 cm. Foliage
leaves 5–6, petiolate, sheathing at base; petiole 2–3
cm long; lamina green, elliptic to elliptic-lanceolate
or ovate with slightly asymmetrical base, acuminate,
9–17 by 4.8–6.5 cm, 7- to 9-veined, glabrous, margins
undulate. Inorescence racemose, erect, ca 20 cm
long; peduncle green, 8–9 cm long, 0.2–0.3 cm in
diameter, with several longitudinal ridges; rachis
densely many-owered; oral bracts green, strongly
recurved, linear-triangular, acuminate, 3.5–8 by
0.7–1.2 mm, shorter to longer than ovaries. Flowers
green, 2–3 mm in diameter. Sepals rounded to acute,
glabrous, margins revolute; dorsal sepal lanceolate
to oblong-lanceolate, 4.2–4.3 by ca 1.5 mm,
3-veined; lateral sepals concave, broadly and
obliquely elliptic, 3–3.5 by 1.9–2 mm, 3-veined.
Petals reexed, linear, rounded to obtuse, 3.6–4 by
0.5–0.7 mm, 1-veined, margins revolute. Lip 2.7–3
by 3–3.7 mm (maximum dimensions), bowl-shaped,
semicircular to transversely rectangular in outline
(when spread) with a sagittate-auriculate base, apically
(truncate-)rounded with a slightly thickened, recurved
and subcrenate apiculum; basal/central part forming
a cavity that is ovate to elliptic in outline, subdivided
by a longitudinal keel and bordered by a more or
less horseshoe-shaped, glabrous callus; auricles
obliquely to subfalcately triangular, acute, 0.6–1 by
0.5–1 mm. Gynostemium green, erect, 1.5–1.8 mm
long, ca 1 mm in diameter; staminodes at, broadly
and obliquely oblong, obtuse to truncate, overarching
and partly hiding the anther; anther light yellow,
elliptic in upper view, ca 0.5 mm long, pollinia 4 in
2 pairs,
narrowly
clavate,
without
caudiculae
;
rostellum
transversely rectangular, truncate; fertile part of stigma
large. Ovary (including pedicel) green, fusiform-
cylindrical, with 6 longitudinal ridges, 3–4 mm long,
glabrous. Capsule ellipsoid, 0.5–0.7 cm long, 0.2–
0.4 cm in diameter; fruit pedicel 0.1–0.2 cm long.
Thailand.— NORTHERN: Chiang Mai [Doi Ang
Khang, Pongkai 100 (BCU!, BKF!)].
Distribution.— NE India.
Vernacular.— Hu suea morakot ()
(here proposed).

thai forest bulletin (botany) Vol. 44 no. 1
42
Ecology.— Crepidium maximowiczianum
grows in humus-rich soil in shaded areas in pine
forest, ca 1700 m alt. It owers in the rainy season
(June to July) and survives the dry season as a leaess
pseudobulb.
Note.— During eld surveys in Doi Ang Khang,
a small population of this species was discovered at
National Highway 1249, ca 5 km south of Royal
Agricultural Station Ang Khang.
Crepidium micranthum (Hook.f.) Szlach., Fragm.
Florist. Geobot., Suppl. 3: 129. 1995.— Microstylis
micrantha Hook.f., Hooker’s Icon. Pl. 19: t. 1834.
1889.— Malaxis micrantha (Hook.f.) Kuntze,
Revis. Gen. Pl. 2: 673. 1891. Type: Malaysia,
Peninsular Malaysia, Perak, “Taiping Hills”, sine
anno, Scortechini 581 [holotype K!]. Fig. 4C–D.
— Microstylis avoviridis Ridl., J. Straits Branch
Roy. Asiat. Soc. 61: 37. 1912. Type: Malaysia,
Peninsular Malaysia, Perak, Gunung Korbu, sine
anno, Hani 3982 [holotype K!].
— Microstylis trinervia Ridl., Bull. Misc. Inform.
Kew 1926: 84. 1926. Type: Indonesia, West Sumatra,
Mentawi Islands, Sipura, sine anno, Boden Kloss
14655 [holotype K!].
Terrestrial, sympodial herb with a creeping
rhizome; owering shoot erect from a decumbent
base, 15–37 cm tall. Rhizome green to green-brown,
terete, 8–20 cm long, 0.3–0.8 cm in diameter, internodes
ca 1 cm long. Roots several, 1–3 arising from each
node of the rhizome and from the lower nodes of
the stem, 1–7 cm long. Foliage leaves 6–14, petiolate,
sheathing at base; petiole light green to greenish
purple, 1–2.5 cm long; lamina green to purplish
brown, lanceolate to ovate-lanceolate or elliptic with
asymmetrical base, acuminate, 5–12 by 2–4.5 cm, 3-
to
5-veined, glabrous, margins undulate. Inorescence
racemose, erect, 15–30 cm long; peduncle green to
purplish brown, 8–10 cm long, 0.2–0.3 cm in diameter,
with several longitudinal ridges, bearing a few lan-
ceolate sterile bracts; rachis laxly many-owered;
oral bracts green to purplish brown, reexed, tri-
angular, acuminate, 4–12.5 by 1–2.5 mm, longer
than ovaries. Flowers greenish yellow, more or less
flushed with purple, 4–5 mm in diameter. Sepals
rounded to acute, glabrous, margins revolute; dorsal
sepal ovate, 2.5–4 by 1.5–2 mm, 3-veined; lateral
sepals broadly and obliquely ovate, 2.5–3.5 by
1.5–2.5 mm, 3-veined. Petals spreading, linear,
obtuse to rounded, 2.5–3 by 0.5–1 mm, 1-veined,
margins revolute. Lip 4–5 by 4.5–6 mm (maximum
dimensions), reniform to nearly semicircular in
outline with a strongly sagittate-auriculate base,
apically rounded with coarsely and unevenly 6- to 8-
dentate margin and a large, triangular, bid, usually
incurved apiculum; basal/central part forming a
small, shallow cavity that is narrowly triangular to
rectangular in outline; auricles obliquely (ovate-)
triangular, obtuse, 1–2 by 1–2 mm. Gynostemium
greenish yellow, erect, stout, 1–1.5 mm long, ca 1
mm in diameter; staminodes flat, broadly and
obliquely triangular-oblong, rounded to obtuse;
anther orange-yellow, ovate in upper view, ca 0.5 mm
long; pollinia 4 in 2 pairs, narrowly clavate, without
caudiculae; rostellum transversely rectangular,
truncate; fertile part of stigma small. Ovary (including
pedicel) green to purplish brown, cylindrical, with
6 longitudinal ridges, 3–5 mm long, glabrous.
Capsule oblongoid-ellipsoid, 0.7–0.9 cm long,
0.3–0.5 cm in diameter; fruit pedicel 0.2–0.3 cm
long.
Thailand.— PENINSULAR: Nakhon Si
Thammarat [Khao Luang National Park, trail from
Ai Khiao Waterfall to Noen Lom Fon, sine anno,
Chantharaprasong H 9040 (BCU!); 21 Sept. 2015,
Nuammee 444 (BCU!, BKF!), Nuammee 445
(BCU!, BKF!)].
Distribution.— Peninsular Malaysia, Sumatra,
Borneo.
Vernacular.— Mangkon lueang thin tai (
)(here proposed).
Ecology.— Crepidium micranthum grows in
humus-rich soil in evergreen forest, ca 800–1200 m
alt. It owers late in the rainy season (September).
After owering, the inorescence decays and the
stem gradually becomes prostrate. However, at least
the proximal part of the stem – and its leaves – survive
the dry season and do not disappear until a new,
young, erect, potentially owering shoot has been
produced from one of the lower leaf axils.
Note.— During field surveys in peninsular
Thailand, a large population of this species was
discovered in a shaded area nearby along a stream
in Khao Luang National Park.

Notes oN Crepidium (orChidaCeae): two New CombiNatioNs, a putative Natural hybrid,
aNd four speCies Newly reCorded for thailaNd (a. Nuammee, t. seelaNaN, s. suddee & h.Æ. pederseN) 43
Figure 4. A–B: Crepidium maximowiczianum (King & Pantl.) Szlach., Chiang Mai province, Doi Ang Khang, 23 June 2014 (A: habit,
B: owers); C–D: Crepidium micranthum (Hook.f.) Szlach., Nakhon Si Thammarat province, Khao Luang National Park, 19 September
2015 (C: habit, D: owers). Photos: A. Nuammee.

thai forest bulletin (botany) Vol. 44 no. 1
44
ACKNOWLEDGEMENTS
Anchalee Nuammee is grateful for nancial
support from the Human Resource Development in
Science Project (Science Achievement Scholarship
of Thailand, SAST) 2013–2015. We are grateful for
the permission of Department of National Parks,
Wildlife and Plant Conservation under the research
project name “Taxonomic Revision and Pollination
Biology of Orchid Genera Crepidium Blume and
Dienia Lindl. (Malaxidinae, Orchidaceae) in
Thailand”.
In addition, the curators and staff of
herbaria K, L and QBG are acknowledged for their
help and hospitality during the visits of A. Nuammee,
and H.Æ. Pedersen thanks the Augustinus Foundation
for nancially supporting his participation in this
study. Finally, we are grateful for suggestions of
André Schuiteman on the lectotypification of
Crepidium josephianum.
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