The Genera Aganope, Derris, and Paraderris (Fabaceae, Millettieae) in Thailand
Author(s): Y. Sirichamorn, F. A. C. B. Adema and P. C. van Welzen
Source: Systematic Botany, 37(2):404-436. 2012.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364412X635467
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Systematic Botany (2012), 37(2): pp. 404–436
© Copyright 2012 by the American Society of Plant Taxonomists
DOI 10.1600/036364412X635467
The Genera Aganope, Derris, and Paraderris (Fabaceae, Millettieae) in Thailand
Y. Sirichamorn,1 F. A. C. B. Adema, and P. C. van Welzen
Netherlands Centre for Biodiversity Naturalis (section NHN), Leiden University, P. O. Box 9514, 2300 RA
Leiden, The Netherlands.
1
Author for correspondence (sirichamorn@nhn.leidenuniv.nl)
Communicating Editor: Rodger Evans
Abstract—In this revision of the Thai species of the genus Derris and the morphologically similar genera, Aganope and Paraderris, all
currently recognized species are enumerated. Two new species, Derris glabra Sirichamorn and Derris pseudomarginata Sirichamorn, are
described and illustrated. Keys to the genera and species are provided, together with descriptions and notes for all taxa.
Keywords—Aganope, Derris, Fabaceae, Millettieae, Paraderris, Thailand.
some species of Derris and closely related genera in their
analyses. However, the small sample and uncertainties about
the affiliations of the species prevented any final conclusions
for Derris and its allies. Thus, the phylogeny of Derris s. l. is
still far from solved. A more detailed phylogenetic analysis
is necessary, including more morphological, anatomical, and
molecular data. We will perform this in the near future.
Derris s. s. differs from the most closely related Paraderris in
flower and inflorescence structure. The flowers of Paraderris
are relatively larger than those of Derris, and usually possess
basal callosities on the standard petals. The brachyblasts of
Paraderris are long and slender with two or three flowers at
the apex, while those of Derris are knob-like or sometimes
slender with more flowers scattered throughout (Geesink
1984; Adema 2003a). On the other hand, Aganope is morphologically more distinct from these two forementioned genera.
It differs from Derris s. s. and Paraderris by having “true
paniculate” inflorescences. The wing and keel petals are free
from each other and the stamens are distinctly diadelphous
(Polhill 1971; Geesink 1984). The African Ostryocarpus and the
American Lonchocarpus are not included in this paper.
Ostryocarpus is morphologically similar to Aganope, but differs mainly in its pods, which are circular in outline and
wingless. Lonchocarpus is a large, morphologically variable
genus of which the species are usually trees, while most of
Paleotropical Derris species are lianas. Pseudoracemes characterize the genus with nodes usually bearing two flowers
on the top of the brachyblast as found in Paraderris, while the
pods are usually without wings.
Derris is mainly present in Southeast Asia, however, the
type species, D. trifoliata Lour., is also found in Africa, India,
Sri Lanka, and the Southwest Pacific islands (Adema 2003b).
Some species of Derris and its allies are cultivated and used
as an insecticide and fish poison because their roots, leaves,
and stems contain isoflavonoid rotenone. Several species,
for example D. scandens (Roxb.) Benth., D. alborubra Hemsl.
D. reticulata Craib, and D. thorelii (Gagnep.) Craib, are used
as local medicine in Thailand and other Southeast Asian
countries. Some species provide beautiful flowers and some
are fast-growing shade trees, these are occasionally used
as ornamentals.
In this paper, a revision of the Thai species of Derris and
its supposedly closely related genera, Paraderris and Aganope,
is presented. The delimitating characters of the genera are
treated in more detail. A key to all Thai species is presented.
For the revision, material of all species was examined to
evaluate all names.
The genus Derris Lour. s. s. is a small to medium-sized
member of the papilionoid Leguminosae, consisting of approximately 50 species, of which 16 can be found in Thailand
(Adema 2000). On account of its indehiscent pods, the genus
was at first included in the tribe Dalbergieae (Bentham 1860).
Later, Polhill (1981) and Geesink (1981, 1984) transferred
many genera with indehiscent pods, including Derris, from
tribe Dalbergieae to tribe Millettieae, because they showed a
close resemblance and relationship to Millettia and related
genera. Derris was established by De Loureiro (1790), who
described two new species, D. pinnata Lour. and D. trifoliata
Lour. The concept of Derris varied considerably in treatments
by different authors (Table 1). For example, Bentham (1860)
accepted Derris in a wide taxonomic sense and included
some American species, which were later called “American
Derris.” He also divided Derris s. l. into several sections.
Geesink (1984) studied the tribe Millettieae using phenetic
and cladistic methods based on morphological and phytochemical characters. He found some serious disadvantages
of lumping many taxa into a single genus Derris. Thus,
Geesink proposed to raise most of Bentham’s sections to
generic level, i.e. Brachypterum (Wight & Arn.) Benth.,
Paraderris (Miq.) Geesink, Ostryocarpus Hook. f. (including
Aganope Miq., Ostryoderris Dunn, and Xeroderris Roberty),
Lonchocarpus Kunth., and Deguelia Aublet. The last two
genera, Lonchocarpus and Deguelia, are known as American
Derris. They are nowadays mainly included in Lonchocarpus
by most workers on Neotropical floras (Adema 2000). Polhill
(1971) changed the generic delimitations again of Ostryocarpus.
He united Aganope and Ostryoderris together under Aganope,
but still recognized Ostryocarpus and Xeroderris as separate.
Adema (2000) used a morphological cladistic analysis to study
the relationship between these Derris-like genera and reunited
the monotypic Xeroderris with Aganope. Adema accepted the
African genus Ostryocarpus as a good genus separate from,
but closely related to Aganope. In that study he also united
Brachypterum with Derris s. s. Thus, the definition and concept
of the genus Derris and related genera has changed again. In
this account we will adhere to Adema’s concept (2000) and
treat the genera Aganope, Derris s. s., and Paraderris.
Molecular phylogenetic studies have the potential to
resolve some of the controversies surrounding the delimitation of Derris and its related genera. However, those completed to date were not comprehensive enough for this
purpose. For example, Hu et al. (2000, 2002) used chloroplast
trnK/matK and nuclear ITS/5.8S sequences, respectively, to
analyze the phylogeny of the tribe Millettieae. They included
404
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
Table 1. Differences between the three main classification systems
of Derris-like plants proposed by Bentham (1860), Geesink (1984) and
Adema (2000).
Bentham (1860)
Genus Derris s. l.
- sect. Dipteroderris
- sect. Brachypterum
- sect. Paraderris
- sect. Aganope
- sect. Euderris
ser. Asiaticae
ser. Americanae
Genus Lonchocarpus
- sect. Neuroscaphi
- sect. Densiflori
- sect. Eriophylli
- sect. Laxiflori
- sect. Punctati
- sect. Fasiculati
- sect. Paniculati
Geesink (1984)
Adema (2000)
- Genus Derris s. s.
- Genus Brachypterum
- Genus Paraderris
- Genus Ostryocarpus
[= Aganope (Miq.)+
Ostryoderris (Dunn)+
Xeroderris (Roberty)]
- Genus Derris s. s.
- Genus Derris s. s.
- Genus Paraderris
- Genus Ostryocarpus
- Genus Aganope
(including
Ostryoderris and
Xeroderris)
- Genus Derris s. s.
- Genus Deguelia
- Genus Lonchocarpus
- Genus Derris s. s.
- Genus Lonchocarpus
- Genus Lonchocarpus
- Genus Lonchocarpus
- Genus Philenoptera
- Genus Lonchocarpus
- Genus Philenoptera
Materials and Methods
More than 2,000 specimens, mainly Thai, including most types, were
studied morphologically. Their examination was facilitated by L, K, P,
BKF, DNP, BK, BCU, PSU, and QBG herbaria. Herbarium abbreviations
follow Holmgren and Holmgren 2007. Moreover, additional specimens
were collected during flowering and fruiting periods from known natural habitats in Thailand in 2009–2010, using standard procedures for
herbarium material (Bridson and Forman 1999). The field characters
were observed and noted. The specimens were identified with keys to
species from previous taxonomic revisions of Derris and allies and floras or manuals of neighboring countries, for example, the Flore du
Cambodge du Laos et du Viêtnam (Phan and Vidal 2001) and Flora of
China (Wei et al. 2008). The identifications were confirmed by comparison with the type specimens.
Notes on Characters and Morphology—LEAVES—The leaves of Derris,
Paraderris, and Aganope species are imparipinnate. The number of leaflets
varies from three (D. trifoliata) to 41 (D. microphylla (Miq.) B. D. Jacks.).
The shape, size, indumentum, and venation of the leaflets are important
for recognition of Derris species. Many species have reddish or brownish
pigments in their young leaves. These pigments appear not to be related
to the sunlight intensity that the plants receive (pers. obs.). Stipellae can
be found in some species as D. trifoliata, D. scandens, and P. elliptica.
Interestingly, the stipellae are usually found in species which have onewinged pods. This possible correlation needs further study. All species
have reticulate venation. Veins do not usually reach the margin but curve
towards the apex, and sometimes anastomose near the margin. Only in
one species, D. amoena Benth., nerves seemingly reach to and anastomose
with the margin.
INFLORESCENCES—Inflorescence type is one of the main characters
that Geesink (1984) used for generic delimitation within the tribe
Millettieae. For Derris s. s. Geesink only noticed “pseudoracemes and
pseudopanicles” as inflorescences. These inflorescence types are characterized by fascicles of flowers on brachyblasts, the wart-like or shortly
cylindrical structures, which are hypothesized to be reduced lateral
inflorescence axes. However, the term ‘pseudoraceme’ was used loosely
for many taxa and one difficulty is to determine which plants have
pseudoracemes (Tucker 1987: 199). Adema (2003b: 393–394) discovered
that sometimes it is difficult to decide in Derris whether a certain inflorescence is a panicle with short lateral branches or a pseudoraceme with
long basal brachyblasts. As an example, Adema identified the presence
of both types in parts of the same inflorescence of D. rubrocalyx Verdc.
and D. koolgibberah F. M. Bailey. Both species are distinctly paniculate
(with short lateral branches) at the base of the inflorescence, while the
rachis bears only brachyblasts and resembles a pseudoraceme higher up.
In this paper, more species are listed that show the same paniculatepseudoracemoid inflorescences. Similar to D. koolgibberah, specimens of
D. alborubra clearly show elongated lateral branches at the base of the
405
inflorescence and these branches become shorter upward and finally only
brachyblasts are present in the apex of the inflorescences. The original
drawing in the first publication (Hemsley 1905) also shows the same
characteristic. An even further reduction can be found in the limestone
specialist species D. tonkinensis Gagnep., which has pseudopaniculate
inflorescences, but sometimes the brachyblasts disappear near the top of
the inflorescence, so that the flowers become solitary per node.
Although not noted by Geesink (1984), real paniculate inflorescences are
also present in Derris marginata (Roxb.) Benth., a species that can easily be
recognized by its long and slender pedicels, and that has no brachyblasts at
all. All inflorescences in this species are true panicles, with flowers solitary
along the main axis and along the lateral branches. This might be the
reason why Miquel (1855: 152) placed this species in Aganope. Our observation agrees with the study of Polhill (1971), who mentioned paniculate
inflorescences for this species as well as the Taiwanese species, D. laxiflora
Haines, in which flowers are also not fascicled, but are well spaced. These
species are still accepted by us under Derris because of similarities in the
flowers and the monadelphous stamens, by which they more resemble
Derris than Aganope. This means that the generic circumscription of Derris
according to Geesink has to be changed.
Paraderris has only pseudoracemes, which do not often unite into
pseudopanicles. The genus can easily be recognized because of its distinctly long and slender brachyblasts with only two or three flowers at the
apex. No intermediate inflorescences were found in any specimens.
As mentioned above, Aganope has paniculate inflorescences. All flowers are solitary and never fascicled on brachyblasts. However, there
may be an exception. The description of A. polystachya (Benth.) Thoth.
indicates flowers on elongated brachyblasts, up to 17 mm long.
Thothathri (1961) also mentioned the presence of “tertiary branchlets”
which in turn bear six to eight flowers. We were not able to examine
the type specimen for this character and, therefore, for the time being
we refrain from altering the genus description of Aganope.
FLOWERS—Flowers of Derris s. s. are relatively small (less than 1.5 cm
long) when compared to other genera. They are usually more or less
fragrant with whitish, pinkish, or purplish petals. Pedicels are usually
long and distinct. Calyces are often reddish, rarely greenish. The standard petal usually lacks basal callosities but is reflexed at the base and
sometimes shows a basal green-yellowish patch to attract pollinators. The
wings and keel petals adhere either by so-called lateral pockets or are
hooked together by auricles or twisted claws (or any combination). In
general, the wings are not curved or rolled backward to the calyx, except
in D. reticulata and D. tonkinensis, of which the wings are distinctly curved
(Fig. 1). Stamens are always monadelphous with basal fenestrae and
the anthers are glabrous, except in D. koolgibberah and D. rubrocalyx,
which occasionally have hairy anthers. The disk is hardy visible, sometimes annular, tubular or 10-lobed.
Similar to Derris, the flowers of Paraderris are whitish or pinkish and
distinctly pedicellate. The wing petals are adnate to the keel petals. The
stamens are monadelphous with basal fenestrae and the disk is indistinct
and sometimes variable in shape and size as found in Derris. It differs
from Derris in the larger (usually more than 1.5 cm long) flowers with two
large basal callosities on the standard petal. An exception is found in
P. laotica (Gagnep.) Adema, which has small flowers without basal callosities. The anthers usually bear some hairs.
The flowers of Aganope have clearly shorter pedicels than the previous
two genera. The petals are dull white or greenish, and usually become
blackish in dried specimens. Wings and keel petals are free from each
other, although lateral pockets are distinctly present. The stamens are
diadelphous and they have some hairs on the connective of the anthers.
The disk is unique and usually consists of 10 free, finger-shaped glands.
PODS—The pods of all genera are indehiscent, usually thin, flat and
winged except in P. montana (Benth.) Adema, which has pods with thick
woody valves that are sometimes almost wingless. The shape of the pod
varies from relatively round to elongate and strap-like. In A. heptaphylla (L.)
Polhill the pods are also constricted between the seeds. Pod indumentum
is also variable. Velvety pods can be found in D. ferruginea (Roxb.) Benth.
and D. pubipetala Miq., while nearly glabrous pods are present in most
other species. Seed chambers, the hard areas around seeds formed by the
thickening of mesocarp and endocarp, are found in some species which
were previously included in the genus (section) Brachypterum such as
D. scandens, D. microphylla, and D. robusta (Roxb. ex DC.) Benth.
SEEDS—The seeds of Derris and Paraderris vary from kidney-shaped to
almost discoid. They are usually compressed and have a central or
slightly eccentric hilum. In Aganope, the seeds vary also in shape, but the
position of the hilum is more eccentric than that of Derris and Paraderris.
Sometimes, the seeds remain abortive and do not develop during maturation of the pods.
406
SYSTEMATIC BOTANY
[Volume 37
Fig. 1. Wing Petal Variation: a. Straight wings of Paraderris elliptica. (Wall.) Adema. b. Slightly curved wings of Derris trifoliata Lour. c. Revolute
wings of D. reticulata Craib. d. Strongly revolute wings in D. tonkinensis Gagnep.
Taxonomic Treatment
Key to the Genera of Thai DERRIS-like Plants
1.
1.
Flowers solitary per node. Wings free from keel petals. Stamens diadelphous. Disks consisting of 10 free finger-shaped glands.
Seeds with eccentric hilum. Specimens usually turn blackish when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aganope
Flowers clustered on reduced lateral axes (brachyblasts), rarely solitary. Wings adnate with keel petals.
Stamens monadelphous. Disks annular, short tubular or five to 10-lobed, sometimes hardly visible or absent.
Seeds with central hilum. Specimens not turning blackish when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Brachyblasts vary in shape and length, usually with more than three flowers throughout.
Standard petals less than 10 mm long, rarely with basal callosities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Derris
2. Brachyblasts slender with two or three flowers at the apex. Standard petals usually more than 10 mm long,
usually with basal callosities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraderris
Aganope Miq., Fl. Ind. Bat. 1: 151. 1855; Polhill, Kew Bull. 25:
266, 1971; Rudd in Dassan & Fosberg (eds.), Rev. Hand.
Fl. Ceylon 7: 234. 1991. P. K. Lôc & Thuân, Tap Chi Sinh
Hoc (J. Biol.) 16: 11. 1994. Thoth. & Das, Rheedea 2: 62.
1996; P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 45. 2001. Derris Lour. sect. Aganope (Miq.)
Benth., J. Linn. Soc., Bot. 4, Suppl. 103. 1860. Benth. &
Hook. f., Gen. Pl. 549. 1865. Baker in Hook. f., Fl. Brit.
Ind. 2: 246. 1878. Thoth., Bull. Bot. Surv. Ind. 3: 175. 1961.
Derris Lour. subgen. Aganope (Miq.) Kurz., J. Asiat. Soc.
Beng. 45: 277. 1876. Deguelia Lour. sect. Aganope (Miq.)
Taub. in Engl. & Prantl, Nat. Pflanzenfam 3, 2: 345. 1891.—
TYPE. Aganope floribunda Miq. [= Aganope thyrsiflora
(Benth.) Polhill] (lectotype designated by Polhill 1971).
Ostryoderris Dunn, Bull. Misc. Inf. Kew: 363. 1911.—TYPE:
Ostryoderris impressa Dunn [= Aganope impressa (Dunn)
Polhill] (lectotype designated by Polhill 1971).
Xeroderris Roberty, Bull. Inst. Fr.-Afr. Noire, Sér. A, 16: 353.
1954.—TYPE: Xeroderris chevalieri (Dunn) Roberty [= Aganope
stuhlmannii (Taub.) Adema].
Lianas or trees. Leaves imparipinnate; stipules caducous; stipellae absent or present. Leaflets 5–9 per leaf,
opposite, entire. Panicles axillary or terminal. Bracts subtending brachyblasts and flowers persistent or caducous,
shorter than the corresponding flower buds; bracteoles
usually at the apex of the pedicel or at the base of the
calyx. Calyx cup-shaped, bilabiate, usually truncate, upper
lip often indistinctly 2-lobed; lower lip 3-lobed. Corolla
whitish or pale greenish, or with green or brown spots,
usually turning blackish in dry specimens; standard with
or without basal callosities, reflexed at base; wings approximately as long as keel petals, free or hardly adnate to
the keel petals. Stamens 10, diadelphous; anthers all equal,
fertile, hairy. Disc consisting of 9 or 10 free, finger-shaped
glands. Ovary hairy, with 3–9 ovules. Pods indehiscent,
flat, thin, leathery, with a wing along both sutures or
along the upper suture only, seed chamber present or
absent. Seeds (narrowly) bean-shaped, generally 1–6 per pod;
hilum eccentric.
Distribution—The genus is found in Tropical Africa, India,
Sri Lanka, Bangladesh, and SE (Southeast) Asia.
Habitat and Ecology—The genus Aganope can be found in
various types of vegetation, even in disturbed or agricultural
areas. One species, A. stuhlmannii (Taub.) Adema, is also
found in semi-arid areas in tropical Africa.
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
407
Key to the Thai Species of AGANOPE
Trees or treelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. A. balansae
Lianas/woody climbers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Standard petals large, 13–14.5 13–15 mm. Pods sinuate, winged along the upper suture only . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. A. heptaphylla
2. Standard petals smaller, 4.5–6 4.5–5.5 mm. Pods not sinuate, winged along both sutures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. A. thyrsiflora
+ +
1.
1.
Derris exserta Craib, Bull. Misc. Inf. Kew: 383. 1927; Fl. Siam.
Enum. 1: 489. 1928.—TYPE: THAILAND. Muang Loei,
20 Mar 1924, Kerr 8785 (holotype: K!; isotype: BK!).
Representative Specimens Examined—LAOS. Khammouan: Thakhet,
20 Oct 1938, Poilane 28165 (P). THAILAND. Loei: Muang, 20 Mar 1924,
Kerr 8785 (BK). VIETNAM. Thanh Hoa: Hoi Xuan, 22 Aug 1920, Poilane
1758 (P); Lao Cai: Phong Thô, 7 Dec 1937, Poilane 26751 (P); Tonkin: At
the border of Yen-Lang river, August 1987, Balansa 2180 (P).
2. Aganope heptaphylla (L.) Polhill, Kew Bull. 25: 268. 1971.
Sophora heptaphylla L., Sp. Pl.: 373. 1753. Derris heptaphylla
(L.) Merr., Interpret. Herb. Amb. 273. 1917.—TYPE:
CEYLON. Hermann s. n. (holotype: BM, see Polhill 1971).
Pterocarpus diadelphus Blanco, Fl. Filip.: 563. 1837. Derris
diadelpha (Blanco) Merr., Philipp. Journ. Sci. 5, Bot. 103.
1910.—TYPE: PHILIPPINES. Palawan: Taytay, May 1913,
Merrill Species Blancoanea 344 (neotype here designated:
L!, barcode L 0419242).
Derris sinuata Benth. ex Thwaites, Enum. Pl. Zeyl. 93. 1859.
Benth., J. Linn. Soc., Bot. 4 (Suppl.): 113. 1860. Baker in
Hook. f., Fl. Brit. Ind. 2: 246. 1878. Prain, J. Asiat. Soc.
Beng. 66: 98. 1897. Thoth., Bull. Bot. Surv. Ind. 3: 195.
1961. Deguelia sinuata (Benth. ex Thwaites) Taub. in Engl. &
Prantl., Nat. Pflanzenfam. 3, 3: 345. 1891.—TYPE: CEYLON.
Caltura District, Thwaites CP 1491 (holotype: K; isotype:
BM!, photo).
+
+
+
+
+
+
+
+
+
+
+
+
Trees, 4–15 m high. Twigs 4–8 mm diam., lenticellate,
thinly pubescent or glabrescent in the lower part, upper part
pubescent. Stipules caducous. Leaves with 5–9 leaflets.
Petiole 10–17 cm long, striate, glabrous; rachis ca. 20 cm long,
striate, glabrous; pulvinus 1–1.5 mm long, glabrous. Stipellae
absent. Leaflets chartaceous to subcoriaceous, terminal one
elliptic to obovate, 14.5–18 7.5–10.5 cm, length/width ratio
1.7–2, base obtuse to rounded, apex obtuse or shortly acuminate, acumen less than 5 mm long, emarginate, upper surface
glabrous, lower surface glabrous or thinly sericeous, midrib
slightly raised above, sometimes thinly sericeous, raised
below, veins flattened or slightly raised above, raised below,
6–10 per side, 11–30 mm apart, not reaching the margin but
curving towards the apex, sometimes anastomosing near
the margin, venation reticulate; lateral leaflets mostly like
terminal one, ovate to elliptic, 9.5–17 5.3–9.5 cm, length/
width ratio 1.7–1.9; pulvinus 3–6 mm long, glabrous. Panicles
axillary or terminal, 17–35 cm long, peduncle up to 5 cm long,
sericeous; lateral branches 3–6 cm long, sericeous. Bracts subtending lateral branches ovate, 2.5–3 1.5–2 mm, caducous,
outside sericeous, inside glabrous. Brachyblasts absent. Bracts
subtending flowers elliptic or ovate, 1–1.5 0.5–1 mm, outside sericeous, inside glabrous. Pedicels 1–2 mm long,
sericeous. Bracteoles caducous. Calyx cup-shaped, 4–5 mm
high, outside sericeous, inside glabrous, upper lip with
2 short lobes, 0.2–0.25 3.5–4 mm, lateral lobes short and
broadly triangular 0.2–0.5 3–3.3 mm, lower lobe shortly
triangular, 0.2–0.3 3–3.5 mm. Corolla whitish. Standard:
claw 2.5–3.5 mm long; blade ovate or orbicular, 13–15 13–
14 mm, apex slightly emarginate, both sides glabrous, with
basal callosities and thickened above the center of the
petal. Wings: claw 1–2 mm long; blade elliptic, 8.5–9
2.5–2.8 mm, apex rounded, both sides glabrous, upper auricle
ca. 0.5 mm, lower auricle indistinct, lateral pocket 2–4 mm or
sometimes indistinct. Keel petals: claw 3.5–4 mm long; blade
boat-shaped, 10.5–11 ca. 4 mm, apex rounded, both sides
glabrous, upper auricle 0.5–0.7 mm, lateral pocket 1.5–
3 mm long. Stamens diadelphous, 10–11 mm long, free part
4–4.5 mm long, in two lengths, longer ones alternating with
shorter ones, hairy in the upper part near anthers; anthers
1.5–1.7 0.5–0.7 mm, hairy. Disc 9 or 10 glands, up to 1 mm
high, hairy. Ovary 6–8 mm long, stipe indistinct, sericeous;
ovules 6–8; style 5–6 mm long, sericeous throughout. Pods
strap-like, flattened, 9.5–21 3.2–4 cm, glabrous, with wings
along both sutures, upper wing 8–10 mm wide, lower wing
5–8 mm wide; seed chamber absent. Seeds 1–3 per pod,
narrowly bean-shaped, 23–25 10–12 by 1.5–2.5 mm; hilum
eccentric, ca. 21 mm long.
Distribution—The species is found in Vietnam, Laos, and
northern Thailand (Fig. 2).
Habitat and Ecology—This species is found at the edge of
evergreen forest, near rivers, less than 600 m.
Phenology—This species flowers from March to August
and fruits from July to October.
Notes—This species is a small to medium sized tree,
but some labels, e.g. Poilane 28165 and 26751 mention
liana as habit. Morphologically there is no difference
between all specimens, thus we regard them as a single
species. The type of D. exserta from Thailand (Kerr 8785)
has slightly smaller flowers and leaflets, but we agree with
Phan (Phan and Vidal 2001) to regard this name as a synonym of A. balansae, because all other characters are exactly
the same.
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1. Aganope balansae (Gagnep.) P. K. Lôc, J. Biol. (Vietnam)
16: 12. 1994. P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 47. 2001. Derris balansae Gagnep., Notul. Syst.
(Paris) 2: 346. 1913. in Lecomte, Fl. Indo-Chine 2: 462. 1916;
Lock & Heald, Leg. Indoch. 87. 1994.—TYPE: VIETNAM.
Tonkin, Aug 1887, Balansa 2180 (holotype: P!; isotype: P!).
Fig. 2. Distribution of Aganope balansae (Gagnep.) P. K. Lôc (diamond)
and Aganope heptaphylla (L.) Polhill (circle).
408
SYSTEMATIC BOTANY
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Woody climbers. Twigs 6–8 mm diam., lenticellate, striate,
thinly strigose or glabrescent in the lower part. Stipules
caducous. Leaves with 5–7 leaflets. Petiole 6.5–10.5 cm long,
striate, lenticellate, glabrous, articulate at the base; rachis
5–10.5 cm long, striate, glabrous; pulvinus 5–10 mm long,
glabrous. Stipellae absent. Leaflets subcoriaceous, terminal one
elliptic, 9.5–17.5 6–9.5 cm, length/width ratio 1.5–1.9, base
obtuse to rounded, apex slightly acuminate, acumen up to
10 mm long, rounded, upper and lower surface glabrous,
midrib raised in a furrow above, distinctly raised below, veins
raised above and below, 8–10 per side, 8–25 mm apart, not
reaching the margin but curving towards the apex, sometimes
anastomosing near the margin, venation reticulate; lateral
leaflets mostly like terminal one, elliptic to ovate, 9.7–16 6–
7.5 cm, length/width ratio 1.5–2; pulvinus 7–9.5 mm long,
thinly strigose to glabrous. Panicles axillary or terminal,
much branched, up to 30 cm long or more, peduncle 3–7 cm
long, (thinly) sericeous, lenticellate; primary or secondary
lateral branches 1.5–22 cm long, articulating with the inflorescence rachis. Bracts subtending lateral branches ovate to
triangular, 1.1–1.3 0.9–1 mm, caducous, outside (thinly) sericeous, inside glabrous. Bracts subtending flowers ovate, 1–
1.5 0.7–0.8 mm, caducous, outside sericeous, inside glabrous.
Pedicels 5–6 mm long, sericeous. Bracteoles at the top of
pedicels, minute, ovate, 0.35–0.8 0.3–0.7 mm, caducous, outside sericeous, inside glabrous. Calyx greenish, cup-shaped,
6–6.5 mm high, inserted on a funnel-shaped receptacle, 2–
2.5 mm high; upper lip triangular, 0.7–0.8 2.5–3 mm, sometimes indistinct, lateral lobes short and broadly triangular,
0.5–0.7 4.5–5.5 mm, sometimes indistinct; lower lobe short
triangular, ca. 0.5 2–2.5 mm, outside sericeous, inside glabrous.
Corolla whitish or pale greenish. Standard: claw 2–3.5 mm
long; blade broadly obovate or orbicular, 13–14.5 13–15 mm,
apex slightly emarginate, both sides glabrous, with large
basal callosities and thickened above the center of the petal.
Wings: claw 1.5–2 mm long; blade spathulate, slightly oblique,
9.5–11 3–3.6 mm, apex rounded, curved upward, both sides
glabrous, upper auricle and lower auricle absent, lateral
pocket sometimes present, 2–3 mm long. Keel petals: claw
3–3.2 mm long; blade boat-shaped, 8.6–10.2 4.5–5 mm, apex
rounded, both sides glabrous, upper auricle indistinct, lateral
pocket 4–4.5 mm long. Stamens diadelphous, 10–11.5 mm long,
free part 2.5–4 mm long, in two lengths, longer ones alternating with shorter ones, hairy, usually persistent during pod
maturation; anthers 1.5–2.8
0.4–0.5 mm, hairy. Disc 9 or
10 glands, 0.7–0.8 mm high, hairy. Ovary 9–9.5 mm long, stipe
ca. 0.5 mm long, sericeous; ovules 5–9; style 5–6 mm long,
sericeous throughout. Pods variable in shape but usually
strap-like, flat, 4.5–19.5 2.5–3.8 cm, glabrescent, distinctly
sinuate between the seeds, with wing along the upper suture
only, 2–4 mm wide; seed chamber absent. Seeds narrowly
bean-shaped, 21–22
8–9
ca. 0.5 mm; hilum eccentric,
ca. 2 mm long.
Distribution—This species is found in Sri Lanka, India
(Bengal), Thailand, Malesia: Peninsular Malaysia, Sumatra,
Borneo, Philippines, Papua New Guinea, and Australia (Fig. 2).
Habitat and Ecology—A species is found in primary and
secondary (dipterocarp) rain forest, often along rivers, swamps,
mangrove forest, seashore, and rocky sea-cliffs, up to 120 m.
Phenology—This species flowers from February to
March (Australia and Papua New Guinea), May to October
(SE Asia) and fruits from May to June (Australia and Papua
New Guinea), July to December, January to February (SE Asia).
[Volume 37
Vernacular Names—Brunei: Akar kemedu raung. Indonesia:
Belulu (Sumatra). Papua New Guinea: Waikahiri.
Notes—The holotype “Hermann s. n.” is mentioned in the
study of Polhill (1971) and now deposited in BM. However,
it is not available in this study. Another holotype, “Thwaites
CP 1491” is also in BM but the photo is visible online.
Representative Specimens Examined—BRUNEI. Belait: Sungei Belait,
about 10.5 miles above Bukit Puan, 31 Aug 1960, Sinclair 10521 (L).
INDIA. Andaman and Nicobar Islands: Kamota, near Guest House,
21 May 1977, Bhargava 4687 (L). INDONESIA. Kalimantan Timur: Berau,
INHUTANI I area, near Sg. Sega, 12 Oct 1996, Kessler et al. Berau 223 (L);
Jambi: Teluk Keladi, Puding, 9 Aug 1985, Rahayu 266 (L); Jawa Timur:
Bawean, 24 Nov 1937, Buwalda 3272 (L); Maluku: North Halmahera,
from Limau to Soa, Galela district, 12 Oct 1976, Yoshida 2404 (L);
West Ceram, 10 Feb 1938, Regma 2850 (L); Papua (Irian Jaya): Bomberai
Peninsula, Tangguh survey area, Saengga River, 22 Feb 2002, Takeuchi
et al. 16052 (L); Riau: Kp. Pangalian, 27 Jun 1939, Buwalda 6798 (L);
Sumatera Barat: Road along the coast S. of Padang, near Painan, 11 Aug
1956, Jacobs 4577 (L); Sumatera Utara: Asahan River near Tandjung Balai,
10 Apr 1954, Alston 15368 (L). MALAYSIA. Johore: Sungei Mawai, off
Sedili Besar River, near Mawai Baharu, Kota Tinggi District, 22 May
1978, Maxwell 78–268 (L); Kelantan: Semrak, sandy mangrove coast,
1 Jun 1982, Stone et al. 15223 (L); Pahang: Rompin area, sandy area
next to the swamp forest, 17 Feb 1980, Maxwell 80–28 (L); Perak:
Sungai Krian Estate, 25 Feb 1938, Spare 33277 (L); Sabah: Ulu Segama,
Kalabakan, 15 Dec 1982, Krispinus SAN 95578 (L); Sarawak: Bako
National Park, first Division, 18 Jul 1966, Hou 555 (L); Trengganu:
South bank of Sungei Ibai at ferry, 6 Jul 1953, Sinclair 39829 (L).
MYANMAR. Myebon, 20 Feb 1958, McKee 6101 (K). PAPUA NEW
GUINEA. Gulf: Tributary of Era River, Baimuru subdistrict, 6 Apr 1974,
Croft and Vinas LAE 61317 (L); Manus: Pelikawa, SW Manus, 23 Jun 1971,
Stone and Streimann NGF 10414 (L); Milne Bay: Harada river, below
Waigani Pltn., 8 Jun 1964, Henty NGF 20514 (L); Morobe: Wagan Village,
Lae subdistrict, 26 May 1969, Streimann NGF 39461 (L). PHILIPPINES.
Leyte, 13 Jul 1915, Wenzel 1444 (L). Mindanao: Zamboanga, Nipaan, 30 Nov
1957, Frake PNH 36257 (L); Mindoro: Puerto Galera & vicinity, 12 May
1952, Santos 5339 (L); Negros: Occidental Negros, May 1906, Whitford
1638 (L). SINGAPORE. Tanjong Gul, 28 Oct 1961, Sinclair 10666 (L).
SOLOMON ISLANDS. Choiseul: West of Manggo River, 23 Mar 1970,
Gafui and collectors BSIP 18347 (L); Kolombangara: south of Bambari
harbour, 6 Jan 1968, Gafui and collectors BSIP 7537 (L); Wagina Island,
16 Mar 1964, Whitmore’s collectors BSIP 5457 (L). SRI LANKA. North
Western: Batticaloa Dist, East of Lady Manning Bridge, Kalladi, 11 Feb
1972, Jayasuriya 706 (L). THAILAND. Ranong, 26 Nov 1973, Santisuk 688
(L); Satul: La-ngu, 8 Jan 1928, Kerr 13946 (K); Pangnga: Kaw Sao Sui,
6 Mar 1989, Kerr 17357 (K).
3. Aganope thyrsiflora (Benth.) Polhill, Kew Bull. 25:
268. 1971. Thoth., Rheedea 2: 63. 1992. P. K. Lôc &
J. E. Vidal, Fl. Cambodge, Laos & Vietnam 30: 48.
2001. Millettia thyrsiflora Benth. in Miq., Pl. Jungh. 249.
1852. Derris thyrsiflora (Benth.) Benth., J. Linn. Soc., Bot.
4 (Suppl.): 114. 1860. Baker in Hook. f., Fl. Brit. Ind. 2:
246. 1878. Prain, J. Asiat. Soc. Beng. 66: 99. 1897; Thoth.,
Bull. Bot. Surv. Ind. 3: 195. 1961. Deguelia thyrsiflora
(Benth.) Taub. in Engl. & Prantl., Nat. Pflanzenfam. 3,
3: 345. 1891.—TYPE: MALAYSIA. Malay Peninsula,
Malacca, Griffith KD 1776 (lectotype designated by
Phan in Phan and Vidal 2001: P!; isolectotypes: K!,
CAL, LE).
Aganope floribunda Miq., Fl. Ned. Ind. 1: 151. 1855.—TYPE:
INDONESIA. Java, Blume s. n. (lectotype here designated:
L!, barcode L 0419344).
Aganope macrophylla Miq., Fl. Ind. Bat. 152. 1855.—TYPE:
INDONESIA. Java, Blume s. n. (lectotype here designated: L!, barcode L 0419347).
Derris pyrrothyrsa Miq., Fl. Ind. Bat., Eerste bijv. 297. 1861.—
TYPE: INDONESIA. Sumatra: near Siboga, Teijsmann HB
691 (holotype: U!).
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
Derris wallichii Prain, J. Asiat. Soc. Beng. 66: 99. 1897. Derris
thyrsiflora (Benth.) Benth. var. wallichii (Prain) Thoth.,
Bull. Bot. Surv. Ind. 3: 196. 1961.—TYPE: INDIA.
Andaman Islands, King’s collector s. n. (isotypes: K!,
CAL, see Polhill 1971)
Derris latifolia Prain, J. Asiat. Soc. Beng. 67: 288. 1898.—
TYPE: UPPER MYANMAR. Kachin Mts, Shaik Mokim s. n.
(holotype: P; isotype: K!).
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Woody climbers. Bark smooth, grayish. Twigs 4–8 mm
diam., lenticellate, thinly pubescent or glabrescent in the
lower part, upper part pubescent. Stipules caducous. Leaves
with 5–9 leaflets. Petiole 3.5–10.5 cm long, striate, glabrous,
articulate at the base; rachis 3–19.5 cm long, striate, glabrous
or slightly strigose; pulvinus 5–7 mm long, glabrous.
Stipellae absent. Leaflets subcoriaceous to coriaceous, terminal one elliptic or narrowly elliptic to obovate, 9–17.5 3.5–
7 cm, length/width ratio 1.7–2.6, base obtuse to rounded,
apex acuminate, acumen up to 10 mm long, rounded or
emarginate, upper surface glabrous and shining, lower surface glabrous or thinly strigose along veins, midrib sunken
above, raised below, veins sunken above, raised below, 5–
7 per side, 15–27 mm apart, not reaching the margin but
curving towards the apex, sometimes anastomosing near the
margin, venation reticulate; lateral leaflets mostly like terminal
one, elliptic or narrowly elliptic to ovate, 6–16.5 3.5–6.6 cm,
length/width ratio 1.7–2.7; pulvinus 5–7 mm long, thinly
strigose to glabrous. Panicles axillary or terminal, much
branched, up to 30 cm long or more, peduncle 0.5–5.5 cm long,
sericeous; primary or secondary lateral branches 1.7–25 cm
long, sericeous. Bracts subtending lateral branches ovate to
triangular, 1.5–1.7 0.5–0.6 mm, caducous, outside sericeous,
inside glabrous. Flowers 8–10 mm long; bracts subtending
flowers narrowly ovate, 1–1.5 0.3–0.6 mm, caducous, outside sericeous, inside glabrous. Pedicels 1–2 mm long, sericeous. Bracteoles at the top of the pedicels, narrowly
triangular to acicular, 0.9–1.1 0.1–0.2 mm, outside sericeous,
inside glabrous, usually caducous. Calyx pale yellowish or
pinkish-green, cup-shaped, 3.7–4.2 mm high, outside sericeous, inside glabrous, upper lip shortly triangular, 0.1–
0.25 2–2.5 mm, sometimes indistinct, lateral lobes shortly
and broadly triangular, 0.2–0.3
3–3.3 mm, sometimes
indistinct, lower lobe shortly triangular, 0.1–0.2 1.5–2 mm.
Corolla white or creamy, usually with green and dark brown
spots scattered along the margin of the petals. Standard: claw
1–1.7 mm long; blade obovate or orbicular, 4.5–6
4.5–
5.5 mm, apex slightly emarginate, glabrous, with basal callosities and thickened inside in the middle of the petal.
Wings: claw 1.5–2 mm long; blade elliptic, 6.3– 6.7
3.5–
3.7 mm, apex rounded, both sides glabrous, upper auricle
0.8–1 mm, lower auricle indistinct, lateral pocket indistinct.
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Derris platyptera Baker in Hook. f., Fl. Brit. Ind. 2: 245, 1878.—
TYPE: INDIA. Malabar: near Calicut, Wight KD 836
(holotype: K!).
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Derris eualata Bedd., Ica. Pl. Ind. Or. 42, t 186. 1874. Derris
thyrsiflora (Benth.) Benth. var. eualata (Bedd.) Thoth.,
Bull. Bot. Surv. Ind. 3: 195. 1961.—TYPE: INDIA. S.
Canara, Beddome s. n. (lectotype here designated: K!).
Keel petals: claw 3–3.3 mm long; blade boat-shaped, slightly
oblique, 6–6.5 2.8–3.2 mm, apex rounded, glabrous, upper
auricle 0.5–0.7 mm, lateral pocket 1.5–2 mm long. Stamens
diadelphous, 7.5–8 mm long, free part 3.5–4 mm long, in
two lengths, longer ones alternating with shorter ones, hairy
especially at the upper part near the anthers; anthers 0.6–0.8
0.4–0.5 mm, hairy. Disc 9 or 10 glands, 0.3–0.5 mm high, hairy.
Ovary 5–6 mm long, stipe indistinct, sericeous; ovules 3–6;
style 4.5–5 mm long, sericeous throughout. Pods strap-like,
flat, 5–15 2.5–3.8 cm, glabrescent, not sinuate, with wings
along both sutures, upper wing 3–8 mm wide, lower wing
3–7 mm wide; seed chamber absent. Seeds 1–4 per pod,
narrowly bean-shaped, 11–22
9–13
1.5–2.5 mm; hilum
eccentric, ca. 2 mm long. Figure 3.
Distribution—This species occurs in India, Myanmar,
Vietnam, Laos, Thailand, Malesia: Peninsular Malaysia,
Java, Borneo, and Philippines (Fig. 4).
Habitat and Ecology—A species from dry evergreen forest, mixed dipterocarp forest, or secondary forests, usually
found along roads, in open and highly disturbed areas,
logged over areas, or Hevea rubber plantations, up to 100 m.
Phenology—Flowering is reported from March to May and
June to November; fruiting is from June to December.
Vernacular Names—Thailand: Sae yan, Sae klan (southern), Sae nui (Ranong), Ye ne ko (Songkhla), Tab tao, Tang
mae (Chanthaburi).
+
Aganope subavenis Miq., Fl. Ind. Bat., Eerste bijv. 299. 1861.—
TYPE: INDONESIA. Sumatra: Lampong Province, near
Natar, Teijsmann s. n. (holotype: U!, barcode U 0003515;
isotype: L!, barcode L 0012044).
409
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Representative Specimens Examined—INDIA. Assam: Cherrapunjee,
Khasi Hills, 23 Apr 1952, Chand 5431 (L); Assam: Hmuntha, Lushai
Hills, 7 Apr 1951, Koelz 27595 (L); Kerala: Ponmudi, old sanitorium
Merchison estate, 6 Mar 1980, Ridsdale 463 (L). INDONESIA. Aceh:
Southern part of reserve, Alas River valley, mouth of the Bengkong/
Renum River, ca. 50 km S. of Kutacane, 7 Jul 1979, de Wilde and de
Wilde-Duyfjes 18509 (L); Jawa Barat: Djasinga, Buitenzorg, 23 Nov 1913,
Backer 10178 (L); Jawa Tengah: Semarang, Kedungdjati, 19 Oct 1996,
Koorders 25326 (L); Kalimantan Timur: Along road Labanan-Kelai, off
km 10 PT. Inhutani I Berau area, 15 May 1998, Sidiyasa at al. Berau 1253
(L); Maluku: Ceram, 1859–1860, Teijsmann s. n., barcode L 0419368 (L);
Sulawesi Tengah: Banggai, Feb 1920, Kaudern 511 (L); Sumatera
Selatan: Palembang, Kajoe Agoeng, 1929, de Voogd 281 (L). MALAYSIA.
Johore: fifth mile Johore-Scudai Road, 4 Jul 1965, Sinclair 10828 (L);
Kedah: Pasir Hitam, P. Langkawi, 18 Aug 1972, Soepadmo and Mahmud,
1240 (L); Pahang: Ginting Highlands road, 20 Jun 1974, Stone and
Hattink 12015 (L); Sabah: Kampung Marak Parak, Kota Merudu,
16 Aug 1983, Gibot SAN 99988 (L); Sarawak: Gunong Gading, Lundu
district, first Division, 25 Sep 1974, Mamit S 35176 (L); Selangor: Kuala
Lumpur, 67 m. s. K. L. to Mentatab, 23 Aug 1972, Teo KL 3063 (L).
PHILIPPINES. Luzon: Catanduanes, Virec, Brgy. Kalatagan, 1 Dec
1991, Barbon et al. PPI 5651 (L); Mindanao: Dinagat, Agusan del
Sur, Koligayan, Agusan 3, Rosario (14 km NW of Bunawan), 10 Oct
1991, Sagcal and Fernando PPI 4911 (L). THAILAND. Chanthaburi:
Ma Kam, 25 Nov 1930, Lakshanakara 531 (L); Phangnga: Khlong Nang Yon,
28 Apr 1973, Geesink and Santisuk 5003 (L); Phatthalung: Tamote, near
Tamote village, 9 Aug 1986, Maxwell 86–534 (L); Ranong: Khao Phra
Mi, 5 Jul 1972, K. Larsen et al. 30644 (L); Satun: E. side of Koh Talutao,
17 Jun 1974, Geesink et al. 7313 (L); Songkhla: Rattaphum, Khao Look Lom,
24 Sep 2009, Sirichamorn YSM2009–22 (L). VIETNAM. Quang Ninh:
Sai Wong Mo Shan (Sai Vong Mo Leng), Lung Wan village, Dam-ha,
18 May 1940, Tsang 30074 (L); Ha Tinh: Huong Son Distr., Rao An,
forest trail at Ngam Thep river crossing to the end of the trail on rocky
river bed, 7 km from police station on Highway 8, 27 Nov 1999, Soejarto
et al. 11170 (L).
Derris Lour., Fl. Cochin. 432. 1790, nom. cons. Benth. in Miq.,
Pl. Jungh. 251. 1852. Proca. Linn. Soc. Lond. 4, Suppl.
101. 1860. Merr. Enum. Philip. Pl. 2: 299. 1923. Backer &
Bakh. f., Fl. Java 1: 616. 1964. Verdc., Man. New Guinea
Leg. 314. 1979. Geesink in Polhill & Raven (eds),
Adv. Leg. Syst. 1: 256. 1981. O. N. Allen & E. K. Allen,
Leguminosae 224. 1981. Geesink, Leiden Bot. Series 8: 91.
1984.—TYPE: Derris trifoliata Lour. (Typ. cons.).
410
SYSTEMATIC BOTANY
[Volume 37
Fig. 3. Aganope thyrsiflora (Benth.) Polhill. a. Habit. b. Flower. c. Standard, inside view. d. Wing petal. e. Keel petal. f. Stamens. g. Pistil with disk.
h. Pod [a-g: Maxwell 86–494; h: Maxwell 86–534; all L]. Drawing by Anita Walsmit Sachs (L).
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
411
stipellae absent or present, persistent. Leaflets 3–19 per leaf,
opposite, entire. Inflorescences axillary pseudoracemes,
often combined with terminal or axillary pseudopanicles,
sometimes an inflorescence in the lower part a (pseudo)
panicle and in the upper part a pseudoraceme, rarely true
panicles; flowers usually fascicled on knob-like, club-shaped
to slender brachyblasts. Bracts subtending brachyblasts
and flowers persistent or caducous, shorter than the corresponding flower buds; bracteoles present, usually at the
apex of the pedicel or at the base of the calyx. Calyx
cup-shaped, often maroonish or reddish, bilabiate, upper
lip 2-lobed, often indistinctly so; lower lip 3-lobed. Corolla
whitish, pinkish, or purplish: standard without or rarely with
small basal callosities, reflexed at base; wings approximately
as long as keel petals, in some species curved backward
towards the calyx, wings and keel petals adherent by lateral
pockets or hooked together by auricles or twisted claws (or
any combination). Stamens 10, monadelphous, with basal
fenestrae; anthers all equal, fertile, often glabrous (to hairy).
Disc absent or inconspicuous to tubular or 10-lobed. Ovary
with (1–)2–12 ovules. Pods indehiscent, thin and stiff, with a
wing along both sutures or along the upper suture only,
seed chamber present or absent. Seeds bean-shaped, 1 or
2(–3) to rarely more per pod; hilum usually central.
Distribution—This genus comprises about 50 species
mainly in SE Asia, one species extending from East (E) Africa
to Australia, and W. Pacific (D. trifoliata, a mangrove species),
and one endemic species in N. Australia.
Habitat and Ecology—Species of Derris s. s. are sunlight
and water-preferring plants, usually found in fully exposed
or semi-shaded areas along the streams or rivers. Two species, D. microphylla and D. robusta are deciduous trees and
found in mixed deciduous forests or at the edge of dry
evergreen forests. Two species, D. scandens and D. amoena
are also found in disturbed or degraded areas, D. trifoliata is
typical for the mangrove, and a third, D. tonkinensis, only
occurs on limestone.
Fig. 4. Distribution of Aganope thyrsiflora (Benth.) Polhill.
Dalbergia L. f. subgen. Brachypterum Wight & Arn., Prod. Fl.
Ind. Or. 1: 264. 1834. Brachypterum (Wight & Arn.)
Benth., Comm. Legum. Gen. 37. 1837. Benth. in Miq.,
Pl. Jungh. 253. 1852. Miq., Fl. Ind. Bat. 1: 138. 1855.
Geesink, Leiden Bot. Series 8: 80. 1984. Derris Lour.
sect. Brachypterum (Wight & Arn.) Benth., Proc. Linn.
Soc. Bot. 4, Suppl. 101. 1860. Baker in Hook. f., Fl. Ind. 2:
549. 1878. Thoth., Bull. Bot. Surv. India 3: 175. 1961.
Geesink in Polhill & Raven, Adv. Leg. Syst. 1: 256. 1981.
Thoth., Fasc. Fl. India 8: 1. 1982. Deguelia Aubl. sect.
Brachypterum (Wight & Arn.) Taub. in Engl. & Prantl,
Nat. Pflanzenfam. 3, 3: 345. 1894.—TYPE: Dalbergia
scandens Roxb. [= Derris scandens (Roxb.) Benth.].
Derris Lour. sect. Diperoderris Benth., Proca. Linn. Soc. Lond.
4, Suppl.:102. 1860.—TYPE: Derris ferruginea (Roxb.)
Benth. (lectotype here designated).
Lianas or trees. Twigs usually lenticellate. Leaves
imparipinnate; stipules present, persistent or caducous;
Key to the Thai Species of DERRIS
Trees or treelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaves with 19–41 leaflets. Stipellae 0.4–0.7 mm long. Leaflets 7–30 4–9 mm, apex rounded, emarginate.
Corolla purplish or violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. D. microphylla
2. Leaves with 11–19 leaflets. Stipellae more than 0.7 mm long. Leaflets 14–65 10–25 mm, apex acute, mucronate.
Corolla white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. D. robusta
Lianas/woody climbers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Stipellae present (check for scars when caducous). Pods one-winged or unwinged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaves with up to five leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Leaflets velvety, hirsute or with scattered, appressed hairs (seemingly glabrous) below, base rounded to cuneate
or subpeltate. Calyx sericeous outside. Standard sericeous or at least with some hairs at the apex outside . . . . . . . . . . . . . . 3. D. elegans
5. Leaflets glabrous below, base slightly cordate to subpeltate. Calyx mostly glabrous outside.
Standard glabrous outside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. D. trifoliata
4. Leaves with more than five leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Leaves with nine to 13 leaflets. Leaflets subcoriaceous, elliptic to ovate or obovate, 3–8 1.2–3.5 cm. Stipellae
up to 1.5 mm long. Brachyblasts knob-like, 2–6 mm long, with five to ten flowers. Wing petals distinctly
strigose on both auricles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. D. scandens
6. Leaves with 13–19 leaflets. Leaflets chartaceous to subcoriaceous, narrowly elliptic to narrowly obovate,
4–7.5 1.5–2.3 cm. Stipellae 2.5–3.5 mm long. Brachyblasts small and indistinct, 0.1–0.4 mm long,
with two or three flowers. Wing petals glabrous but thinly ciliate in the basal part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. D. thorelii
3. Stipellae absent. Pods one or two-winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Inflorescences panicles or mixed forms between panicles and pseudoracemes/pseudopanicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Inflorescences panicles. Pedicels 10–12 mm long, glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. D. marginata
8. Inflorescences mixed forms between panicles and pseudoracemes/ pseudopanicles. Pedicels 3–4.5 mm long, sericeous . . . . . . . . . . . . 9
9. Young leaves reddish or brownish. Leaflet apex obtuse and slightly emarginate. Brachyblasts present near apex
of the inflorescence. Wings petals not curved backward to the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. D. alborubra
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SYSTEMATIC BOTANY
9. Young leaves light green. Leaflet apex shortly acuminate. Brachyblasts sometimes absent near apex of the
inflorescence and then flowers solitary. Wings petals curved backward to the calyx . . . . . . . . . . . . . . . . . . . . . . . 15. D. tonkinensis
Inflorescences pseudoracemes or pseudopanicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Leaflet base slightly cordate to subpeltate. Pods one-winged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. D. trifoliata
10. Leaflet base obtuse, (narrowly) cuneate to rounded. Pods two-winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Pods velvety or sericeous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Leaflets lightly strigose to velvety below, apex rounded, obtuse or cuspidate to short acuminate.
Pod elliptic to strap-shaped, 7.8–9.5 3–3.5 cm, upper wing 4–10 mm wide, lower wing 4–7 mm wide.
-Northern and North-eastern Thailand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. D. ferruginea
12. Leaflets usually strigose to almost glabrous below, apex distinctly acuminate.
Pods ellipsoid to almost circular, 3–6.5 1.8–3 cm, upper wing 5–9 mm wide,
lower wing 2–4 mm wide. —Southern Thailand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. D. pubipetala
11. Pods mostly glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Leaflets nine to 11, narrowly obovate, base narrowly cuneate to attenuate. Inflorescences pseudoracemes. . . . 8. D. monticola
13. Leaflets three to seven, sometimes up to 9, elliptic ovate or obovate, base cuneate to obtuse, rarely attenuate.
Inflorescences pseudoracemes or pseudopanicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Inflorescences short, up to 6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. D. reticulata
14. Inflorescences long, usually more than 6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Leaflets sometimes glaucous below. Leaf veins seemingly ending or anastomosing in the margin . . . 2. D. amoena
15. Leaflets never glaucous below. Leaf veins not reaching the leaf margin but curving toward the apex,
sometimes anastomosing near the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Leaves with up to five leaflets. Calyx glabrous outside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. D. glabra
16. Leaves with five or more leaflets. Calyx (thinly) sericeous outside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Leaflets elliptic to obovate, 3–8.5 1.5–3.5 cm, apex acuminate, acumen up to 10 mm long.
Brachyblasts club-shaped to elongated cylindrical, 2–13 mm long, sericeous,
with 3–8 flowers.—Limestone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. D. tonkinensis
17. Leaflets ovate to elliptic, 6–12 3.3–5.5 cm, apex shortly acuminate,
acumen 4–7 mm long. Brachyblasts knob-like to elongate cylindrical,
sometimes hardly visible, 0.3–5 mm long, thinly sericeous, with 2–3
(rarely up to 5) flowers.—Not on limestone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. D. pseudomarginata
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Lianas, basal diameter 5–10 cm. Twigs 3–4 mm diam.,
glabrous, lenticellate. Leaves with 3–7 leaflets, reddish or
brownish when young. Stipules triangular, ca. 1.5
1.3–
1.5 mm, outside thinly sericeous, inside glabrous. Petiole 2–
4.5 cm long, grooved and striate, thinly sericeous or nearly
glabrous; rachis 3.5–5 cm long, (slightly) grooved above,
sericeous, especially in the groove; pulvinus ca. 5 mm long,
thinly sericeous. Stipellae absent. Leaflets subcoriaceous to
coriaceous; terminal one elliptic, 7–8 ca. 3 cm, length/width
ratio 2.3–2.6, base cuneate, obtuse to rounded, apex obtuse
and slightly emarginate, upper surface shiny, glabrous but
strigose along midrib, lower surface thinly strigose, midrib
slightly raised in a furrow above, veins slightly raised above,
7–10 per side, 5–8 mm apart, not reaching the margin but
curving towards the apex, sometimes anastomosing near
the margin, venation reticulate; lateral ones mostly like terminal one, elliptic, 6.5–8 2–3 cm, length/width ratio 2.6–3;
pulvinus 2–3 mm long, (thinly) sericeous. Inflorescences
axillary or terminal, mixed forms between panicles and
pseudoracemes/ pseudopanicles, up to 30 cm long, peduncle
5–35 mm long, sericeous; lateral branches up to 15 cm long,
becoming shorter upwards and near the apex only
brachyblasts. Bracts subtending brachyblasts (or lateral
branches) triangular to narrowly triangular, 1.3–1.8 0.6–
0.8 mm, outside sericeous, inside glabrous. Brachyblasts
cylindrical, 1.5–3 mm long, with 3–5 flowers, sericeous.
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Derris alborubra Hemsl. var. honheoensis P. K. Lôc in P. K.
Lôc & J. E. Vidal, Fl. Cambodge, Laos & Vietnam 30: 86.
2001.—TYPE: VIETNAM. Khanh Hoa, Nha Trang. Poilane
3134 (holotype: P!; isotype: P!).
Bracts subtending flowers ovate to triangular, 0.7–0.8 0.5–
0.6 mm, outside sericeous, inside glabrous. Pedicels 3–5 mm
long, sericeous. Bracteoles at the top of the pedicels, triangular, ca. 0.7 0.3–0.4 mm, outside sericeous, inside glabrous.
Calyx maroonish, cup-shaped, 4–4.2 mm high, outside sericeous, margin ciliate, inside glabrous, tube 2.7–3 mm high,
upper lip with 2 short triangular lobes, 0.3–0.5 1.5–2 mm;
lateral lobes triangular, 1–1.3 3.5–4 mm; lower lobe triangular, 1–1.3 1.7–2 mm. Corolla pure white. Standard: claw
2.5–3 mm long; blade broadly obovate to orbicular, 8–10
7–8 mm, apex emarginate, basal callosities absent, outside
thinly pubescent at apex, sometimes glabrescent, inside
glabrous. Wings: claw 3–4 mm long; blade elliptic, 8–8.5
1.3–1.5 mm, apex rounded, outside glabrous or thinly ciliate
at apex, inside glabrous, upper auricle 0.7–1 mm long, lower
auricle indistinct, 0.1–0.3 mm long. Keel petals: claw ca. 3 mm
long; blade boat-shaped, 6.5–8 3–3.5 mm, apex rounded,
outside glabrous or thinly ciliate at apex, inside glabrous,
upper auricle 1–1.5 mm long, lateral pocket hardly visible.
Stamens monadelphous, with basal fenestrae, 10.5–11 mm
long, free part 2–3 mm long, glabrous; anthers ca. 0.4
0.3 mm, glabrous. Disc shortly tubular, more or less distinctly
(5- or) 10-lobed, up to 0.6 mm high. Ovary 6.5–7 mm long,
stipe indistinct, sericeous; ovules 4 or 5; style 3–4 mm long,
thinly hairy at base. Pods elliptic to discoid or obovoid, 2–4
2.2–2.5 cm, with few hairs especially at the base and the beak,
sometimes glabrous, winged along both sutures, upper wing
4–5 mm wide, lower wing ca. 2 mm wide; seed chamber
absent. Seeds 1 per fruit, discoid, flat, 7–8 6–7.5 ca. 1 mm;
hilum central, 1–1.3 mm long.
Distribution—This species occurs in China, Vietnam, Laos,
Thailand (northern and north-eastern), and Cambodia (Fig. 5).
Habitat and Ecology—This species is present in evergreen
forests, usually near streams, 100–300 m.
Phenology—In China flowering is reported in June or July.
In Thailand flowering is in September to October. Fruiting
has been reported in August to December.
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1. Derris alborubra Hemsl., Bot. Mag. 131: t. 8008. 1905.
Gagnep. in Lecomte, Fl. Indo-Chine 2: 452. 1916. Craib,
Fl. Siam. Enum. 1: 487. 1928. P. K. Lôc & J. E. Vidal, Fl.
Cambodge, Laos & Vietnam 30: 84. 2001.—TYPE: Curtis
Bot. Mag.: Table 8008, cultivated at K and raised from
seed from Hong Kong (see Note below).
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
413
Benth. var. maingayana (Baker) Prain, J. Asiat. Soc. Beng.
66: 106. 1897. Ridl., Fl. Malay Penins. 1: 597. 1922. Craib,
Fl. Siam. Enum. 1: 487. 1928.—TYPE: MALAYSIA.
Malacca, Maingay KD 1553 (holotype: K!; isotype: CAL).
2. Derris amoena Benth. in Miq., Pl. Junghuhn. 252. 1852.
J. Linn. Soc., Bot. 4 (Suppl.): 110. 1860. Baker in Hook. f.,
Fl. Brit. Ind. 2: 245. 1878. Prain, J. Asiat. Soc. Beng. 66:
105. 1897. Ridl., Fl. Malay Penins. 1: 597. 1922. Craib,
Fl. Siam. Enum. 1: 487. 1928.—TYPE: MYANMAR.
Martaban, Wallich Cat. 5912 (holotype: K!).
Derris maingayana Baker in Hook. f., Fl. Brit. Ind. 2: 245.
1878. Prain, J. Asiat. Soc. Beng. 66: 467. 1897. Derris amoena
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Representative Specimens Examined—CHINA. Canton, 19 June 1887,
Samson s. n. (K); Guangzhou, June 1964, Tang 10026, 10196, 10687 (L).
LAOS. Wiengchan: Borikhan, 29 Apr 1932, Kerr 21303 (K). THAILAND.
Chaiyaphum: Ban Nam Phrom, 24 May 1974, Geesink and Phengklai 6912
(L); Nakhon Nayok: Hell Narok waterfall, Khao Yai National Park, 3 Sep
2009, Sirichamorn YSM2009–14 (L). VIETNAM. Tonkin: Sai Wong Mo Shan
(Sai Vong Mo Leng), Lung Wan Village, Dam-ha, 18 Jul 1940, Tsang 29854,
29138 (L); Tonkin, Aug 1887, Balansa 2297 (L).
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Uses—Stems or roots are sometimes used as local medicine, combined with the stems and roots of D. scandens they
form a treatment for osteoarthritis.
Vernacular Names—Thailand: Tao ta pla (Nakhon
Ratchasima). Vietnam: (Dây) cô mò (Khanh Hoa).
Notes—Vegetatively Derris alborubra is similar to D. scandens
by having glossy, subcoriaceous, elliptic leaflets. Small differences are present, D. alborubra has on average fewer (three to
seven instead of nine to 13) and larger leaflets than D. scandens.
Because of the similarity, the same Thai name is often used for
both species, just as they are also used as medicinal alternatives for treatment of osteoarthritis. Derris alborubra differs from
D. scandens by the different type of inflorescence (see note
on morphological characters), by calyx color and by the twowinged pods without seed chambers. About three or four
specimens at K were mentioned as types of this species, all
with the same information: published in Curtis Bot. Mag.:
Table 8008, but without collecting date, collector, or collector
number. They look relatively new and the first author
assumes that they were collected later, possibly from a plant
cultivated in the Royal Botanic Garden, Kew, which was
grown from seed received from Hong Kong. A living plant
no longer exists in K.
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Fig. 5. Distribution of Derris alborubra Hemsl. (star) and D. amoena
Benth. (circle).
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Lianas. Bark maroonish, lenticellate, glabrous. Twigs 3–
5 mm diam., slightly strigose or glabrescent, lenticellate.
Leaves with 5–9 leaflets, reddish when young. Stipules triangular, ca. 1.5 0.7–1 mm, outside thinly sericeous, inside
glabrous. Petiole 3.5–6 cm long, slightly grooved and striate,
glabrous or with few hairs; rachis 3.5–11 cm long, grooved
and striate, glabrescent; pulvinus 5–9 mm long, thinly strigose. Stipellae absent. Leaflets subcoriaceous; terminal one
elliptic to obovate, 6–9 3–4.5 cm, length/width ratio ca. 2,
base cuneate to rounded, apex acuminate, acumen 2–5 mm
long, rounded, upper surface glabrous or slightly strigose,
especially at the basal part of the midrib, lower surface
glabrous or slightly strigose, usually glaucous, midrib flat
or slightly raised above, veins flat on both sides, 7–13 per
side, 2–10 mm apart, parallel, seemingly ending and anastomosing in the margin, venation reticulate; lateral ones
mostly like terminal one, elliptic to obovate, 4.5–8.5
2–4 cm, length/width ratio 2.1–2.3; pulvinus 5.5–7 mm long,
glabrous. Pseudopanicles axillary, 7–18 cm long, peduncle
0.5–4.5 cm long, striate, glabrous or sometimes slightly
sericeous. Bracts subtending brachyblasts ovate to triangular,
ca. 0.8 0.6–0.7 mm, outside sericeous with ciliate margin,
inside glabrous. Brachyblasts knob-like, 0.5–0.7 mm long,
with 2 or 3 flowers, sericeous. Bracts subtending flowers
ovate, 0.4–0.5 0.2–0.5 mm, outside sericeous, inside glabrous.
Pedicels 3–7 mm long, sericeous. Bracteoles at the top or the
upper part of the pedicels, ovate, 0.3–0.4 0.2–0.3 mm, outside slightly pubescent, inside glabrous. Calyx cup-shaped,
2–3 mm high, maroonish, outside thinly sericeous, margin
ciliate, inside glabrous, tube 2.5–2.8 mm high, upper lip with
2 short lobes, shallowly notched, 0.3–0.4 3–4 mm; lateral and
lower lobes shortly triangular, 0.5–0.6 1.5–2 mm, apex obtuse
or rounded. Corolla purplish. Standard: claw 2.7–3 mm long;
8–8.5 mm, apex
blade broadly elliptic to orbicular, 6–6.5
emarginate, basal callosities absent, outside slightly ciliate at
apex, inside glabrous. Wings: claw 1.7–3 mm long; blade
elliptic, 6–7
1.7–2.3 mm, apex rounded, outside slightly
hairy and ciliate at the apex, inside glabrous, auricle ca.
0.5 mm long. Keel petals: claw ca. 2 mm long; blade boatshaped, 6–6.5 ca. 3 mm, apex rounded, outside thinly sericeous near the apical part and ciliate at apex, inside glabrous,
upper auricle indistinct, 0.1–0.2 mm long, lateral pocket 1.2–
1.7 mm long. Stamens monadelphous, with basal fenestrae,
7–8 mm long, free part 2–3 mm long, glabrous; anthers
ca. 0.5 0.4 mm, glabrous. Disc indistinct. Ovary 4.5–5 mm
long, thinly sericeous; ovules 2 or 3; style 3.5–4 mm long,
slightly hairy at base or sometimes glabrous. Pods broadly
straplike, flat, 4–9.5 1.5–2.6 cm, glabrous, with wing along
both sutures, upper wing 3–4 mm wide, lower wing ca. 2 mm
wide. Seeds unknown.
Distribution—This species is found in Thailand (Peninsular);
Malesia: Malay Peninsula, Sumatra, and Borneo (Fig. 5).
Habitat and Ecology—Derris amoena grows commonly in
edge of evergreen forests, along roadsides or rivers, in disturbed areas, in para-rubber (Hevea) plantations or open
margins of swamp forest, up to 25 m.
Phenology—This species flowers from December or January to March and fruits from February to April.
SYSTEMATIC BOTANY
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Lianas, up to 25 m long. Outer bark green, inner bark
yellow. Twigs 3–12 mm diam., hirsute or thinly strigose to
glabrous. Leaves with 3 or 5 leaflets. Stipules narrowly triangular, 2–5 2–5 mm, caducous, outside strigose or hirsute to
glabrous, inside glabrous. Petiole 1–16 cm long, striate, strigose, or hirsute to glabrous; rachis 1–11 cm long, striate,
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Derris rufula Schum. & Lauterb., Fl. Deutsch. Schutzgeb. 361.
1901.—TYPE: PAPUA NEW GUINEA. Lauterbach 2821
(lectotype designated by Adema 2003a: K!).
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Derris vestita Baker in Hook. f., Fl. Brit. Ind. 2: 242. 1878. Derris
elegans Graham ex Benth. var. vestita (Baker) Prain, J. As.
Soc. Beng. 66: 103. 1879. Ridl., Fl. Mal. Penins. 1: 597.
1922. Craib, Fl. Siam. Enum. 1: 488. 1928. Verdc., Kew
Bull. 32: 469. 1978.—TYPE: MALAYSIA. Malacca,
Maingay KD 608 (1633) (holotype: K!, isotype: L!).
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3. Derris elegans Graham ex Benth. in Miq., Pl. Jungh. 252.
1852. Ridl., Fl. Mal. Penins 1: 596. 1922. Merr. Enum.
Philip. Fl. Pl. 2: 299. 1923. Craib, Fl. Siam. Enum. 1: 488.
1928. Burkill, Dict. 1: 784. 1935. Verdc., Kew Bull. 32: 468.
1978. Adema, Blumea 48: 396. 2003. Derris elegans var.
elegans, Adema, Blumea 48: 397. 2003.—TYPE: MYANMAR. Martaban, Wallich 5883 (holotype: K!).
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Representative Specimens Examined—BRUNEI. Tutong: near KPG,
Danale, 10 Jun 1967, van Niel 4386 (L). INDONESIA. Borneo: Lingkas,
Tarakan, Kalimantan, Timur, 8 Mar 1981, Wiriadinata and Pawiroatmodjo
128 (L); Sumatra: East coast, subdivision Laboehan Batoe, District Kota
Pinang, 15 Apr-16 May 1933, Rahmat Si Toroes 4072 (L). MALAYSIA.
Johore: Kg. Hubang Development area, 100 m. s. Endau Road, Lowland
forest, 14 Jul 1959, Burkill HMB 1902 (L); Kelantan: Bukit Baka forest
Reserve, 2 Jun 1982, Stone, Chin et al. 15235 (L); Trengganu: Kemaman,
Hill side, 13 Jul 1931, Osman and Tachon 27014 (L). SINGAPORE. Nee
Soon: open margins of the swamp forest, 14 Feb 1983, Maxwell 83–
11 (L). THAILAND. Ranong: entrance to Ngao Falls Forest Reserve,
18 Jan 1987, Maxwell 87–94 (L); Satun: Edge of evergreen forest,
28 Dec 1927, Kerr 13700 (L); Surat Thani: Roadside, on Motorway No. 44
(from Phun Phin to Kiansa), km 92, 22 Sep 2009, Sirichamorn YSM
2009–20 (L).
strigose or hirsute to glabrous; pulvinus 4–10 mm long,
(thinly) hirsute. Stipellae acicular, 1.5–5 mm long. Leaflets
chartaceous; terminal one elliptic to obovate, 6.5–20.5 2.5–
9 cm, length/width ratio 1.6–2.8, base rounded to cuneate
or subpeltate, rarely cuneate, apex obtuse to rounded or
shortly and broadly acuminate, acumen 2–3 mm long,
rounded or emarginate, upper surface glabrous or more or
less hirsute on midrib and nerves in lower part, lower surface hirsute to velvety, in older leaflets often much less hairy,
then mostly along midrib and nerves, midrib and nerves
raised above, nerves 6–10 per side, 6–29 mm apart, not
reaching the margin but curving towards the apex, sometimes anastomosing near the margin, venation reticulate;
lateral ones mostly like terminal one, (broadly) ovate to
elliptic, 4.5–16.5
3–7.5 cm, length/width ratio 1.5–2.9;
pulvinus 2–9 mm long, hirsute to glabrous. Pseudoracemes
axillary and ramiflorous, sometimes several together forming
pseudopanicles, 1.5–23 cm long; peduncle 0–20 cm long,
thinly hirsute, sometimes strigose. Bracts subtending brachyblasts narrowly triangular or narrowly elliptic to ovate,
1–5.5 0.3–0.8 mm, outside sericeous, inside glabrous. Brachyblasts knob-like, shortly cylindrical, up to 2.5 mm long,
with 2 or 3 flowers. Bracts subtending flowers (narrowly)
0.2–6 mm,
ovate to triangular to elliptic or linear, 0.5–4
outside sericeous, inside glabrous. Pedicels 2.5–9 mm long,
hirsute or strigose. Bracteoles in the upper part of the pedicels, often at different heights, narrowly ovate or narrowly
triangular to elliptic or linear, 0.5–1.7
0.2–0.5 mm, outside sericeous, inside glabrous. Calyx 2.5–4.5 mm high,
brownish to purple or red, rarely green, outside sericeous,
inside glabrous, tube 2–3.5 mm high; upper lip 0.5–1
1.5–4.5 mm, shallowly notched; lateral lobes triangular or
semicircular, 0.5–1
1.5–3 mm; lower lobe triangular,
0.5–1.5 1.5–2.5 mm. Corolla pink to white. Standard: claw
1.5–3 mm long; blade orbicular, rarely obovate, 8–9.5 9.5–
10.5 mm, apex emarginate, without basal callosities, outside sericeous or at least with some hairs at the apex,
inside glabrous. Wings: claw 2.5–4 mm long; blade elliptic,
7–9
3–3.5 mm, apex rounded, upper auricle 0.5–1 mm
long, lower auricle indistinct, lateral pocket indistinct,
outside with some hairs at the ventral side at the apex,
inside glabrous. Keel petals: claw 2–4 mm long; blade
more or less boat-shaped, 7–8.5 2.5–4 mm, apex rounded,
auricle 0.2–1 mm long, lateral pocket 1–2.5 mm long, outside sericeous at the upper part, at least at the ventral side,
inside glabrous, rarely with some hairs at the apex. Stamens
monadelphous, with basal fenestrae, 8–12 mm long, free
part 1–4 mm long, alternately longer and shorter, glabrous;
anthers 0.3–0.6 0.2–0.4 mm, glabrous. Disc hardly visible.
Ovary 3.5–6 mm long, sericeous; stipe 0.5–1 mm long,
sericeous; ovules 2–5; style 4–8 mm long, sericeous at base.
Pods more or less oblong to orbicular, 3–6 1.5–3 cm, only a
single wing along the upper suture, sometimes hardly visible,
0–3 mm wide, (thinly) hirsute; seed chamber absent. Seeds
discoid or bean-shaped, compressed, 9–14 7–13 1–4 mm;
hilum central, 1–2 mm long.
Distribution—This species is found in Myanmar,
Andaman Islands, Indochina, Thailand, Malesia: Peninsular
Malaysia, Sumatra, Java, Borneo, Philippines (Mindanao,
Palawan), New Guinea, Solomon Islands, and Vanuatu
(Fig. 6).
Habitat and Ecology—This species grows in primary or
secondary forest along rivers, up to 300 m.
+
Uses—The stems are ocassionally used as rope.
Vernacular Names—Thailand (Peninsular): Yan Sao Kam,
Yan Sao Dam, Sao Dam, Ya Na Lae
Notes—Derris amoena and D. maingayana share similar
morphological characters. Some authors (Baker 1878; Prain
1897; Adema 2003b) accepted D. maingayana as a separate
species. While others (Prain 1897; Ridley 1922; Craib 1928)
placed it as a variety of D. amoena. Craib (1928: 487) mentioned that the varieties do not obviously differ in most
characters, only the whitish waxy coating on the lower surface of the leaflets and the width of the wings along the two
sutures are used as characters to differentiate this species
from D. amoena (Baker 1878: 245). Field observations show
that the plant has reddish or brownish young leaves and the
degree of waxy coating increases during the maturation
of the leaves, thus younger leaves are less glaucous than
the older ones. However, there does not seem to be any real
difference between both species/varieties based on the
degree of glaucous coloration. We also found that specimens
identified as D. amoena usually had young leaves. The type
specimens of D. amoena and D. maingayana are also similar.
The leaflets of both taxa share the unique characteristic of
secondary veins, which are parallel, seemingly ending and
anastomosing in the margin. Moreover, the distributions of
both taxa are entirely the same. Thus, here we treat D. amoena
and D. maingayana as one species.
[Volume 37
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SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
415
rachis 5–7 cm long, slightly grooved above, sericeous; pulvinus 5–10 mm long. Stipellae absent. Leaflets subcoriaceous;
terminal one elliptic or obovate, 6.5–12 3–5 cm, length/
width ratio 2.1–3, base cuneate, apex acuminate or obtuse to
rounded, acumen up to 15 mm long, rounded, upper surface
glabrous but hairy along midrib and lateral main veins,
lower surface slightly strigose to velvety, midrib and veins
flat or slightly raised above, veins 7–9 per side, 5–10 mm
apart, not reaching the margin but curving towards the apex,
sometimes anastomosing near the margin, venation usually
reticulate but sometimes slightly scalariform near midrib;
lateral ones (narrowly) elliptic, ovate or obovate, 4–15 2–
4.5 cm, length/width ratio 2–3.3, base cuneate, apex acuminate or obtuse to rounded, acumen up to 1.5 cm long,
rounded; pulvinus 5–6 mm long, sericeous. Pseudoracemes/
pseudopanicles axillary or terminal, 9–23.5 cm long, peduncle 1–1.5 cm long, sericeous, branches 6–11 cm long. Bracts
subtending brachyblasts ovate-triangular, 1.5–2.5 0.8–1.2 mm,
outside sericeous, inside glabrous. Brachyblasts knob-like
to cylindrical, 1–3.5 mm long, with 3–7 flowers. Bracts
subtending flowers ovate, 1–1.8 0.5–0.8 mm, outside velvety, inside glabrous. Pedicels 6–7 mm long, sericeous. Bracteoles in the middle or in the upper part of pedicels nearly at
the top, sometimes caducous, ovate or narrowly ovate, 0.8–
1.2
0.2–0.5 mm, outside velvety, inside glabrous. Calyx
cup-shaped, dull maroonish, 2–2.5 mm high, outside thinly
sericeous, inside glabrous or sericeous along the margin, tube
ca. 2 mm high; upper lip with 2 short lobes, 0.1–0.2 2–
2.5 mm; lateral lobes short triangular, 0.7–1 2–2.5 mm, apex
obtuse; lower lobe shortly triangular, 0.6–0.8 2–2.5 mm,
apex obtuse. Corolla white. Standard: claw 2.7–2.8 mm long;
blade orbicular, ca. 9 7 mm, apex emarginate, without auricle or basal callosities, outside thinly sericeous, especially at
the apex, inside glabrous. Wings: claw 3–3.2 mm long; blade
elliptic, 6–7
0.7–1 mm, apex rounded, outside hairy at
the apex, inside glabrous, upper auricle ca. 1 mm long.
Keel petals: claw ca. 2.8 mm long; blade boat-shaped, ca.
7.5 2.5 mm, apex rounded, outside hairy at the apex, inside
glabrous, lateral pocket small or indistinct, 0.5–1 mm long,
auricle ca. 1 mm long. Stamens monadelphous, with basal
fenestrae, 9–10 mm long, free part of filaments 3–4 mm long,
glabrous; anthers ca. 0.5 0.4 mm, glabrous. Disc indistinct.
Ovary sericeous to velvety, 7–7.5 mm long, stipe sericeous, 1–
1.2 mm long; ovules 3–5; style 5–6 mm long, glabrous but
slightly sericeous at base. Pods elliptic to strap-like, 7.8–9.5
3–3.5 cm, flat, thin, velvety, leathery with a wing along both
sutures, flat, upper wing 4–10 mm wide, lower wing 4–7 mm
wide; seed chamber hardly visible. Seeds 1 or 2 per pod,
bean-shaped, flat, 18–20 12–15 mm; hilum central.
Distribution—Derris ferruginea is known from India,
Myanmar, south China, Laos, Thailand (northern and northeastern), and Vietnam (Fig. 7).
Habitat and Ecology—This species grows in open areas
along streams, in the forest margin or on hill slopes with
granite bedrock, 450–1,200 m.
Phenology—Flowering is from February to March or from
July to August. Fruiting is from August to November.
Uses—The roots are known as “Indian tiba root” used in
the preparation of insecticides; but when compared with
Paraderris elliptica then the rotenone content is less (0.1–4.3%)
(Thothathri 1982: 19).
Vernacular Names—Vietnam: Bö gan (Khanh Hoa), Cóc
kèn set (Pham Hoang Hô).
+
Lianas, basal diameter 10–15 cm. Bark smooth, greenish
and aging to more pale grayish-brown. Twigs densely covered with golden-brown hairs. Leaves with 7–9 leaflets, occasionally reddish or brownish when young. Stipules ovate to
triangular, 3–4 1–4 mm, outside velvety, inside glabrous.
Petiole 3.5–6.5 cm long, slightly grooved above, sericeous;
+
+
+
+
+
+
+
+
4. Derris ferruginea (Roxb.) Benth. in Miq., Pl. Jungh. 252.
1852. J. Linn. Soc., Bot. 4 (Suppl.): 109. 1860. Baker in
Hook. f., Fl. Brit. Ind. 2: 245. 1878. Craib, Fl. Siam.
Enum. 1: 490. 1928. Thoth., Bull. Bot. Surv. Ind. 3: 189.
1961. P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 56. 2001. Robinia ferruginea Roxb., Fl. Ind,
Ed. 2, 3: 329. 1832.—TYPE: INDIA. Silhet, Wallich 5885 A
(lectotype designated by Thothathri 1982: K!; isolectotypes: CAL, E, LE).
+
Representative Specimens Examined—INDONESIA. Sumatra: Aceh,
Middle Alas River (Lae Sauraya) area, c 15 km N. of Gelombang, S. of
Bengkong River, mainly plateau forest. 26 Jul 1985, de Wilde & de WildeDuyfjes 20359A (L). MALAYSIA. Perak: Sungai Krain Estate, 10 Jan 1939,
Spare SFN 36208 (L); Sabah: Bodukan, Ranau dist., 22 July 1986, Amin
and Francis SAN 116078 (L); Selangor: Ulu Gombak: 25 Feb 1971, Teo and
Pachiappan KL 2959 (L); Ulu Kelang, Klang Gate, 11 Apr 1974, Kasim
and Zainudin 01749 (L); Ulu Langat: above Pansoon, 7 Feb 1975, van
Balgooy and Stone 2257 (L). PAPUA NEW GUINEA. Central Province: 6 miles
south-west of Karema, Brown river, 30 Jul 1962, Schodde 2667 (L); Madang
Province: Enecil village, near Gogol River, Madang sub-province,
25 Nov 1969, Katik NGF 46616 (L); West Sepik Province: Gorge Nth of
Meinat flood plains, Bewani Subprov., 19 Sep 1982, Kerenga LAE 56523
(L). THAILAND. Narathiwat: Sungei Kolok, Nikom Waeng, 2 Mar 1974,
K. Larsen and S. S. Larsen 32828 (L); Songkhla: Klong Lahm Tahp, Saba
Yoi, 22 Feb 1987, Maxwell 87–183 (L); Satun: Thung Nui, 13 Feb 1961,
Suvarnakoses 38131 (L).
+
Phenology—Flowering has been reported from January to
March and from July to September. Fruiting has been
reported from March to April and from July to August.
Uses—Derris elegans is used as a fish poison in New
Guinea and possibly Peninsular Malaysia. A solution of
crushed leaves in (green) coconut water is used in Papua
New Guinea to wash out snake bites.
Vernacular Names—Malaysia: Aka tuba (Peninsular).
Papua New Guinea: Imora (Kabali, Central Province).
Thailand: Oud nam, Oud laud (Surat Thani)
Notes—According to Adema (2003b), D. elegans is a variable, widespread species with four varieties: D. elegans var.
elegans, D. elegans var. celebica Adema, D. elegans var.
gracillima (Hemsl.) Verdc., and D. elegans var. korthalsiana
(Blume ex Miq.) Adema. The description provided here pertains to var. elegans, the only variety found in Thailand. This
variety is recognized by its denser indumentum than the
other varieties.
+
Fig. 6. Distribution of Derris elegans Graham ex Benth.
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2012]
SYSTEMATIC BOTANY
[Volume 37
Derris spanogheanae similissima, foliolis floribusque
paucioribus (foliolis 3–5, floribus 2–5 in brachyblastis, versus
plus quam 5 foliolis floribusque), axe communi ramis
lateralibus pedicellis glabrioribus differt.
Lianas, basal diameter ca. 5–10 cm. Twigs 3–8 mm diam.,
lenticellate, glabrous, striate. Stipules triangular, 1–1.1 9–
1.1 mm, caducous, outside thinly sericeous, margin ciliate,
inside glabrous. Leaves with 3–5 leaflets. Petiole 2.5–5 cm
long, striate, glabrous; rachis 0.7–2.5 cm long, striate, glabrous; pulvinus 4–6 mm long, glabrous. Stipellae absent.
Leaflets subcoriaceous; terminal one (broadly) elliptic, 6.5–
9.5 3.4–5.8 cm, length/width ratio 1.6–1.9, base obtuse, apex
acuminate, acumen 5–6 mm long, rounded or slightly emarginate, upper and lower surface glabrous, midrib distinct,
pale, flat above, distinctly raised below, veins slightly raised
above and below, 4–6 per side, 7–15 mm apart, not reaching
the margin but curving towards the apex, sometimes anastomosing near the margin, venation reticulate; lateral ones
mostly like terminal one but usually smaller, elliptic to ovate,
4.5–5.5 2.5–3.8 cm, length/width ratio 1.4–1.8; pulvinus 4–
6 mm long, glabrous. Pseudoracemes/ pseudopanicles axillary or terminal, 12.5–28 cm long; peduncle 10–15 mm long,
glabrous; lateral branches 2.5–17 cm long. Bracts subtending
brachyblasts ovate, 0.9–1 0.4–0.5 mm, outside glabrous or
with some hairs at the base, margin ciliate, inside glabrous.
Brachyblasts knob-like to elongated cylindrical, 1–11 mm
long with 2–5 flowers throughout, glabrous or with few
scattered hairs. Bracts subtending flowers ovate, 0.6–
0.7
0.4–0.5 mm, glabrous on both sides, margin ciliate.
Pedicels 3–4 mm long, glabrous or with some scattered hairs.
Bracteoles at the apex of pedicels, elliptic to ovate, 0.5–
0.6
0.2–0.4 mm, glabrous on both sides, margin ciliate.
Flowers slightly fragrant. Calyx reddish-green, cup-shaped,
2.7–3.2 mm high, outside glabrous, margin ciliate, inside glabrous or thinly sericeous in the apical part, tube 2.5–2.7 mm
high, upper lip, 0.3–0.5 2.3–2.5 mm; lateral lobes triangular
or semi-circular, 0.3–0.5 1.7–2 mm; lower lobe triangular or
semi-circular, 0.3–0.5 1.5–1.6 mm. Corolla whitish or with
pale pinkish hue at the top. Standard: claw 2.7–3 mm long;
blade broadly obovate to orbicular, 6.5–7 7–8 mm, apex
emarginate, basal callosities absent, glabrous. Wings: claw
2.7–3 mm long; blade elliptic, 5.6–6
1.8–2 mm, apex
rounded, upper auricle 0.5–1.2 mm long, lower auricle indistinct, both sides glabrous. Keel petals: claw 2.7–3 mm long;
blade boat-shaped, 5.5–6 3–3.5 mm, apex rounded, upper
auricle 0.4–0.7 mm long, lateral pocket ca. 1.5 mm long,
sometimes hardly visible, both sides glabrous or with some
hairs at apex. Stamens 10, monadelphous with basal fenestrae, 8–9 mm long, free part 2–2.5 mm long, glabrous; anthers
ca. 0.5 0.4–0.6 mm, glabrous. Disc annular or indistinct, 0.2–
0.5 mm high, glabrous. Ovary 5.5–6 mm long, thinly strigose,
stipe indistinct; ovules usually 3; style 3.5–4 mm long, glabrous. Immature pods strap-like, 7–8 1–1.5 cm, glabrous,
with a wing along both sutures, upper wing 3–3.3 mm wide,
lower wing 2.7–3 mm wide; seed chamber absent. Seeds
unknown. Figure 8 .
Distribution—This species is endemic to Thailand (Peninsular) (Fig. 7).
Habitat and Ecology—This species grows in evergreen
forests and in fully exposed to semi-shaded areas near
roadsides and on hill slopes. Soil: Laterite, 300–400 m.
Phenology—Flowering is from August to September and
fruiting is in October.
+
+
416
Representative Specimens Examined—CHINA. Hainan: Hung Mo
Shan, Lai [Loi] area, 21 July 1929, Tsang and Fung 544 (K). INDIA. Assam:
Jenkins, 1845, Hooker s. n. (K); Silhet: Cachar, June 1874, Keenan s. n. (K).
THAILAND. Chiang Rai: Mae Chan, Ban Mai Pattana, Nawng Pah Gaw
Subdistrict, 16 Apr 1989, Maxwell 89–467 (L); Udon Thani: Phu Foi Lom,
Nong Sang, 1 Sept 2009, Sirichamorn YSM2009–13 (L). VIETNAM. Hatinh:
Hong Son Distr., Rao An, Ngam Thep River about 4–5 km from Police
station on Highway 8, 26 Nov 1999, Soejarto et al. 11141 (L).
5. Derris glabra Sirichamorn, spec. nov.—TYPE: THAILAND.
Songkhla: Rattaphum, Khao Luk Lom, 24 Sep 2009,
Sirichamorn YSM 2009–23 (holotype: L!; isotypes: L!, BK!).
+
+
+
+
+
+ +
+
+
+
Notes—This species is similar in many characters to D.
pubipetala. They have different, disjunct distributions. Derris
ferruginea has a northern distribution, ranging from India to
southern China, Laos, northern-northeastern Thailand and
northern Vietnam, whereas D. pubipetala is found in the
southern part, from southern Thailand to Peninsular Malaysia,
Borneo, Celebes, and the Philippines (Luzon, Mindoro,
Mindanao, Palawan). Morphologically there are some differences. Adema (2003b: 401) noted differences in pod wings
and the apex of the leaflets. Derris ferruginea has pods with
more unequal wings, the lower wing distinctly narrower
than the upper one; in D. pubipetala the wings almost have
the same width. In D. ferruginea the apex of the leaflets varies
from cuspidate to shortly acuminate or rounded, but they
are distinctly acuminate in D. pubipetala. However, in the
first author’s opinion these characters are too variable to
be useful. Some specimens of D. ferruginea, i.e. Sirichamorn
YSM2009–13 from northeastern Thailand and Soejarto et al.
11141 from northern Vietnam, also show more equal wings
of the pods and leaflets with an acuminate apex (acumen
up to 1.5 cm long) as found in specimens of D. pubipetala.
With regards to floral color, Thothathri (1982) also mentioned
that D. ferruginea has purple flowers, while D. pubipetala has
whitish or greenish ones. In this study, some specimens identified as D. ferruginea (for example Maxwell 89–467) also have
white flowers. Although both species are morphologically
similar, preliminary molecular results of cpDNA markers,
trnK-matK, trnL-F IGS (intergenic spacer), and psbA-trnH, as
well as one Nuclear marker, ITS, showed that they are not
closely related, and as their distributions are non-overlapping,
we still recognize both species as separate taxa.
+
+
Fig. 7. Distribution of Derris ferruginea (Roxb.) Benth. (circle),
D. marginata (Roxb.) Benth. (inverted triangle), D. monticola (diamond),
D. thorelii (triangle), D. pseudomarginata Sirichamorn (square), D. glabra
Sirichamorn (star).
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
417
Fig. 8. Derris glabra Sirichamorn. a. Habit. b. Brachyblast with flower buds. c. Flower. d. Standard, inside view. e. Wing petal. f. Keel petal.
g. Stamens. h. Pistil. i. Pod [a-i: Sirichamorn YSM 2009–23; all L]. Drawing by Anita Walsmit Sachs (L).
SYSTEMATIC BOTANY
6. Derris marginata (Roxb.) Benth. in Miq., Pl. Jungh. 1:
252. 1852. Benth., J. Linn. Soc. 4, Suppl.: 111. 1860. Baker
in Hook. f., Fl. Brit. Ind. 2: 245. 1878. Gagnep. in
Lecomte, Fl. Indo-Chine 2: 453. 1916. Craib, Fl. Siam.
Enum. 1: 490. 1928. Thoth., Bull. Bot. Surv. Ind. 3: 187.
1961. P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 81. 2001. Dalbergia marginata Roxb., Fl.
Ind. ed. 2, 3: 230. 1832. Aganope marginata (Roxb.) Miq.,
Fl. Ned. Ind. 1: 152. 1855. Deguelia marginata (Benth.)
Taub. in Engl. & Prantl., Nat. Pflanzenfam. 3, 2: 345.
1891.—TYPE: BANGLADESH. Silhet, Wallich 5909
(holotype: K!; isotypes: CAL, LE, P!).
+
+
+
+
+
+
+
+
Lianas. Twigs 4–6 mm diam., glabrous, lenticellate. Stipules triangular, 2–2.5 ca. 2 mm, outside thinly sericeous,
inside glabrous. Leaves with 5–7 leaflets. Petiole 2.5–8 cm
long, striate, glabrous; rachis 5.5–11 cm long, striate, glabrous; pulvinus 8–10 mm long, glabrous. Stipellae absent.
Leaflets subcoriaceous to coriaceous; terminal one elliptic, 6–
14.5 3.5–6.5 cm, length/width ratio 1.7–2.5, base obtuse to
rounded, apex acuminate, acumen 8–14 mm long, rounded,
upper and lower surface glabrous, midrib slightly raised in a
furrow above, distinctly raised below, veins raised on both
sides, 9–12 per side, 7–20 mm apart, not reaching the margin
but curving upwards, sometimes anastomosing near the
margin, venation reticulate; lateral ones mostly like terminal one, elliptic to ovate, 6.8–13.5 3.2–6 cm, length/width
ratio 2.1–2.2; pulvinus 6–11 mm long. Panicles axillary,
rarely terminal, 12–22 cm long, peduncle 10–35 mm long,
glabrous or thinly strigose, striate; lateral branches 2.5–
14 cm long. Bracts subtending lateral branches triangular,
1.5–1.6 1.3–1.4 mm, outside glabrous to thinly sericeous,
inside glabrous but with some hairs at base. Brachyblasts
absent. Bracts subtending flowers elliptic to ovate, 0.9–1
0.4–0.6 mm, both sides glabrous or with few hairs,
margin ciliate. Pedicels slender, 10–12 mm long, glabrous.
Bracteoles at the upper part of pedicels, narrowly elliptic to
ovate, 0.7–0.9 0.2–0.3 mm, both sides glabrous, margin
slightly ciliate. Calyx reddish-green, cup-shaped, 2.5–3.7 mm
high, both sides glabrous, margin ciliate, tube 2–3 mm high,
upper lip indistinct or with 2 short lobes, 0.2–0.3
1.5–
3 mm; lateral lobes triangular or semi-circular, 0.7–1 2–
+
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2.7 mm; lower lobe triangular or semi-circular, ca. 0.7
1.5–1.6 mm. Corolla whitish or pinkish. Standard: claw 3.3–
3.5 mm long; blade broadly obovate to orbicular, basal
callosities absent, 8–9 7–7.5 mm, apex emarginate, outside
glabrous or with few hairs at the apex, inside glabrous.
Wings: claw 3–3.5 mm long; blade elliptic, 8.3–9
1.8–
2.2 mm, apex rounded, upper auricle 1–1.7 mm long, lower
auricle indistinct, both sides glabrous. Keel petals: claw 2.5–
3 mm long; blade boat-shaped, 7–7.3 2.7–3.1 mm, apex
rounded, glabrous, upper auricle indistinct, lateral pocket
1.5–2 mm long. Stamens monadelphous, with basal fenestrae, 10–10.5 mm long, free part 1.5–2 mm long, glabrous;
anthers ca. 0.3 0.2–0.3 mm, glabrous. Disc annular, more
or less distinctly (5–)10-lobed, up to 0.3 mm high. Ovary 4–
5 mm long, stipe indistinct, glabrous to thinly sericeous;
ovules 4 or 5; style 3–3.5 mm long, glabrous. Pods elliptic
to strap-like, 7–10 2.5–3.5 cm, glabrous, with wing along
both sutures, upper wing 3–10 mm wide, lower wing 2–5 mm
wide, seed chamber absent. Seeds 1 per fruit or sometimes
12–13
1–2 mm; hilum
immature, discoid, flat, 13–14
central, 1–1.4 mm long.
Distribution—This species is distributed from India
through Nepal, Bangladesh, Myanmar, south China, Laos,
Thailand, and Vietnam (Thothathri 1961; Phan and Vidal
2001) (Fig. 7).
Habitat and Ecology—This species is found growing in
evergreen forests, rarely in coastal forest, 3–800 m.
Phenology—Flowering season has been reported in May
to July and fruiting is from July to September (Phan and
Vidal 2001).
Vernacular Names—Vietnam: cóc kèn bı̀a (Pham Hoang
Hô). Thailand: Tang mae.
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Representative Specimens Examined—INDIA. Assam, Jenkins s. n. (L);
Calcutta, Hortus Botanicus (cultivated), Pierre s. n. (P). THAILAND.
Chanthaburi: Khao Soi Dao Wildlife Sanctuary, in front of the park
headquarters, 29 Aug 2009, Sirichamorn YSM2009–12 (L).
7. Derris microphylla (Miq.) B. D. Jacks., Index Kewensis
1: 332. 1895. Heyne, Nutt. Pl. 2: 307. 1916. Backer &
Bakh.f., Fl. Java 1: 618. 1964. P. K. Lôc & J. E. Vidal, Fl.
Cambodge, Laos & Vietnam 30: 72. 2001. Brachypterum
microphyllum Miq., Fl. Ned. Ind., Eerste bijv. 296. 1860.—
TYPE: INDONESIA. Sumatra: Palembang, Teysmann s. n.
(holotype: L!, barcode L 0475651; isotype: U!, barcode
U 0003512).
Derris dalbergioides Baker in Hook. f., Fl. Brit Ind. 2: 241. 1878.
Prain, J. As. Soc. Beng. 66: 101. 1897. Gagnep. in Lecomte,
Fl. Indo-Chine 2: 456. 1916. Ridl., Fl. Malay Penins. 1:
595. 1922. Craib, Fl. Siam. Enum. 1: 487. 1928.—TYPE:
MALAYSIA. Malacca, Maingay s. n. (lectotype here
designated: K!).
Deciduous trees 5–20 m high, dbh up to 25 cm. Bark
pale grayish-brown. Twigs ca. 4 mm diam., lenticellate,
velutinous. Stipules narrowly triangular, 4–5 0.5–0.7 mm,
both sides sericeous, hairs shorter towards the base. Leaves
with 19–41 leaflets, golden-brownish-green when young.
Petiole 9–17 mm long, grooved above, pubescent; rachis 5–
15 cm long, grooved above, pubescent; pulvinus up to 5 mm
long. Stipellae acicular, 0.4–0.7 mm long, sericeous. Leaflets
chartaceous, terminal one (narrowly) elliptic to obovate, 14–
7–9 mm, length/width ratio 2.1–3.3, base cuneate to
30
broadly cuneate, apex rounded, emarginate, upper and
lower surface thinly sericeous, midrib slightly sunken above,
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Representative Specimens Examined—Known only from the type.
[Volume 37
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Note—This species is similar to D. spanogheana Blume ex
Miq. in most morphological characters. It differs only by
having fewer leaflets and fewer flowers per brachyblast
(D. spanogheana usually has more than five leaflets and more
than five flowers per brachyblast). Both species are mostly
glabrous, however, pedicels, lateral branches, and main axes
of the inflorescences of D. glabra are more glabrous than those
of D. spanogheana. The first Thai specimen we have seen
(Maxwell 86–674) was identified as D. affinis Benth., but this
name is probably a synonym of D. trifoliata. Additional collections with flowers and young pods from the same locality
of Maxwell’s specimen were obtained during a field trip.
Adema (2003b: 402) tried to clarify the “mostly glabrous”
species of Derris and he united the specimens with 2-winged
pods into D. spanogheana. However, the habitat of the Thai
specimens is slightly different, they are collected in wet
areas, while most specimens of D. spanogheana are from drier
areas in the Philippines e.g. Luzon, Mindanao, Mindora,
Leyte. Cebu, Panay, Alubat, Celebes, and the Lesser Sunda
Islands (Adema pers. comm.).
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SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
Fig. 9.
Distribution of Derris microphylla (Miq.) B. D. Jacks.
8. Derris monticola (Kurz) Prain, J. Asiat. Soc. Beng. 66:
361. 1897. Dunn, J. Linn. Soc. 41: 237. 1912. Craib, Fl.
Siam. Enum. 1: 491. 1928. Thoth., Bull. Bot. Surv. Ind. 3:
185. 1961. Grierson & Long, Fl. Bhutan 1: 657. 1987.
Millettia monticola Kurz, J. Asiat. Soc. Beng. 42: 67. 1873.
Baker in Hook. f., Fl. Brit. Ind. 2: 106. 1876.—TYPE:
MYANMAR. Martaban, Kurz 1776 (holotype: CAL, see
Thothathri 1961).
Derris acuminata Benth. var. sikkimensis Thoth., J. Jap. Bot.
51(5): 143. 1976.—TYPE: INDIA. Sikkim, Hooker s. n.
(holotype: K!).
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Lianas. Twigs 3–5 mm diam., striate, glabrous, lenticellate.
Leaves with 9–11 leaflets. Stipules ovate-triangular, 2–2.2
1.5–1.8 mm, outside sericeous, inside glabrous. Petiole ca.
9.5 cm long, thinly hirsute; rachis ca. 8.5 cm long, thinly
hirsute; pulvinus 9–10 mm long. Stipellae absent. Leaflets
subcoriaceous; terminal one narrowly obovate, ca. 8.5
2.7 cm, length/width ratio ca. 3.2, base narrowly cuneate to
attenuate, apex cuspidate, acumen 3–5 mm long, more or
less rounded, upper surface glabrous, lower surface thinly
sericeous and sometimes glaucous, midrib and veins flat to
slightly raised above, raised below, veins 15–18 per side, 3.5–
6 mm apart, reaching to near the margin, then curving
towards the apex, sometimes anastomosing near the margin,
venation reticulate; lateral ones mostly like terminal one,
elliptic to narrowly obovate, 6 –9 2.3–2.5 cm, length/width
ratio 2.6–3.6; pulvinus 3–5 mm long. Pseudoracemes axillary,
12–17 cm long, peduncle 1.7–2.5 cm long, thinly strigose.
Bracts subtending brachyblasts ovate-triangular, 1–1.2 ca.
0.9 mm, outside sericeous, inside glabrous. Brachyblasts
knob-like, 1–1.5 mm long with 3–5 flowers throughout,
pubescent. Bracts subtending flowers narrowly ovate, 1–1.2
0.4–0.7 mm, outside sericeous, inside glabrous. Pedicels
4–4.5 mm long, sericeous. Bracteoles at the base of the calyx,
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Representative Specimens Examined—CHINA. Guangdong: Guangzhou,
15 Jun 1981, Yip 286 (L). INDIA. Assam: Kohima, Naga Hills, 27 Jun 1950,
Koelz 25355 (L). INDONESIA. Celebes: 22 Sep 1938, bb 25550 (L); Java:
Bandoeng, 20 Oct 1948, Popta 0036 (L); Sumatra: 16 Apr 1932, Lörzing
16522 (L). MADAGASCAR. Fianarantsoa: 5 Dec 1993, Dan Turk 590 (K).
MALAYSIA. Negeri Sembilan: Jempol, Serting Tengah F. R., 25 May 1993,
Saw FRI 39802 (L); Penang: Sg. Tembeling, King George Nat. Park, 17 Apr
1954, Wyatt-Smith KEP 71947 (L). Selangor: Ulu Gombak, University of
Malaya Field Study Center, near the Lab, 3 May 1968, Mahmud 827 (L).
SRI LANKA Peradenija, 3 May 1955, Worthington 6733 (L). THAILAND.
Chumphon: 35 km North of Chumphon, 16 Jul 1963, King 5593 (L);
Nakhon Si Thammarat: Khao Lawng, 10 May 1955, Snan 298 (L);
Pangnga: Nai Chong, 9 May 1973, Geesink and Santisuk 5274 (L); Surat
Thani: Khao Sok, 30 km E. of Takua Pa, 9 Jun 1992, Larsen et al. 42735 (P).
UGANDA. Kampala: Mengo District, 24 Sept 1952, Dale U802 (K).
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veins flat above, slightly raised below, 5–8 per side, 1–5 mm
apart, becoming hardly visible near the margin, venation
reticulate; lateral ones mostly like terminal one, (narrowly)
elliptic, 7–29 4–8 mm, length/width ratio 1.6–3.7; pulvinus
1–2 mm long. Pseudoracemes axillary, 8.5–16.5 cm long,
peduncle 1.5–5 cm long, pubescent. Bracts subtending
brachyblasts narrowly triangular to acicular, 2.2–2.5 0.3–
0.4 mm, both sides sericeous, hairs shorter and more dense
in lower part. Brachyblasts knob-like or shortly cylindrical,
more or less curved upwards, 1–3 mm long, with 5–8 flowers.
Bracts subtending flowers narrowly ovate to acicular, 0.7–
1 0.2–0.3 mm, outside sericeous, inside glabrous or with
some hairs. Pedicels 1–3 mm long, sericeous. Bracteoles at
the base of the calyx, narrowly ovate, 0.6–0.8 ca. 0.3 mm,
outside sericeous, inside glabrous. Calyx brownish-green,
cup-shaped, ca. 2.5 mm high, tube 1.5–2 mm high; upper lip
2–2.5 mm;
with 2 short lobes, shallowly notched, 0.5–1
lateral lobes triangular, 0.5–1 1–1.5 mm; lower lobe triangular, 0.5–1 ca. 1.5 mm, outside sericeous, inside glabrous.
Corolla purplish Standard: claw 1.5–1.7 mm long; blade
broadly obovate to orbicular, 7–7.7 5.5–6 mm, apex emarginate, with 2 small basal callosities, outside with some hairs
at the apex and along the upper part of the midrib or
glabrescent, inside with some hairs at the apex. Wings: claw
ca. 2 mm long; blade elliptic, 6.5–7 2–2.5 mm, apex slightly
curved upward, apex rounded, auricles and lateral pocket
indistinct, both sides glabrous. Keel petals: claw ca. 2.5 mm
long; blade more or less boat-shaped, 6–6.5 ca. 3 mm, apex
rounded, lateral pocket 1–2 mm long, outside with some
hairs to thinly sericeous on major veins at ventral side and
apex, inside glabrous. Stamens monadelphous, with basal
fenestrae, 8–8.5 mm long, free for 2–3.5 mm, glabrous;
anthers ca. 0.4 0.2 mm, glabrous. Disc tubular, ca. 0.3 mm
high. Ovary 5–6 mm long, sericeous, stipe 0.5–1 mm long;
ovules 11 or 12; style 2–3.5 mm long, sericeous at base. Pods
more or less falcate, flattened, bulging over the seeds, 3.5–7
1.3–1.8 cm, stipe 2–3 mm long, upper wing 2–4 mm wide,
outside thinly sericeous, glabrescent, seed chamber visible from outside, dark, veined. Seeds bean-shaped to
transverse-ellipsoid, 4.5–7.5 3–5 mm; hilum central, ca.
1 mm long.
Distribution—Derris microphylla is present in from
Uganda, Madagascar, India, Sri Lanka, south China, Indochina, Thailand, Malesia: Peninsular Malaysia, Sumatra,
Java and Celebes (Fig. 9).
Habitat and Ecology—This species is found in deciduous
forest, road and riversides, up to 1,200 m.
Phenology—Flowering season is from April to August.
Fruiting is from May to September.
Uses—This species is used as timber and shade trees. In
Malaysia a poultice of roots or bark is used to treat itch,
and sometimes for ornamental purposes (Thothathri and
Rugayah 1997: 116).
Vernacular Names—Indonesia: Kayu retak (Palembang).
Malaysia: Daun berayai, batai betek (Peninsular). Thailand:
Lak keay, khang ten (Surat Thani and South-eastern), Prang
pun, ma naam chaai (Chumphon). Di ngu, fantae (Peninsular). Vietnam: (Cây) tràch (Dông Nai), dau dau (Ninh Thuân),
tri-et (Lâm Dông).
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SYSTEMATIC BOTANY
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Comparatione D. marginata foliorum venis secondariis vix
visibilibus. Inflorescentiae pseudopaniculatae. Brachyblasti
variabiles inter relative longi ad brevissimi interdum vix
visibiles praecipue prope inflorescentiae apicem, in D.
marginata absentes. Pedicelli distincte breviores crassiores
pilis numeriosioribus quam in D. marginata. Flores paulo
minores eis D. marginata.
Lianas, basal diameter ca. 5 cm. Twigs 5–8 mm diam.,
lenticellate, the lower part thinly sericeous to glabrous, the
upper part sericeous. Stipules ovate-triangular, 1.7–1.8
1.6–1.7 mm, outside thinly sericeous, margin ciliate, inside
glabrous. Leaves with 5–7 leaflets, subcoriaceous. Petiole 4–
6 cm long, striate, glabrous to thinly strigose; rachis 5–8 cm
long, striate, glabrous to thinly strigose; pulvinus 4–8 mm
long, thinly strigose. Stipellae absent. Leaflets: terminal one
elliptic to obovate, 4.5–12 2.3–5.5 cm, length/width ratio
1.8–2.2, base obtuse, apex shortly acuminate, acumen 4–7 mm
long, rounded, upper and lower surface glabrous, midrib
raised in a furrow above, distinctly raised below, veins flat or
slightly sunken above, raised below, 6–9 per side, 6–17 mm
apart, not reaching the margin but curving towards the
apex, anastomosing near the margin, venation reticulate; lateral ones mostly like terminal one, ovate to elliptic, 5–10.5
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9. Derris pseudomarginata Sirichamorn, spec. nov.—
TYPE: THAILAND. Chonburi: Sriracha district, Collins
275 (holotype: K!).
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Representative Specimens Examined—MYANMAR. Amherst, Mulayit
ridge, 4 Feb 1927, Parkinson 5164 (K). THAILAND. Chiang Mai: 29 Mar
1911, Kerr 1731 (L, P).
3–5.5 cm, length/width ratio 1.8–1.9, base obtuse to rounded;
pulvinus 4–7 mm long, thinly strigose. Pseudoracemes/
pseudopanicles axillary or terminal, 5–17 cm long, peduncle
1–5 cm long; lateral branches 4.5–15 cm long, thinly sericeous, glabrescent. Bracts subtending brachyblasts ovate or
triangular, 0.7–0.8 0.5–0.7 mm, outside thinly sericeous or
glabrous, inside glabrous. Brachyblasts knob-like to elongate
cylindrical, sometimes hardly visible, 0.3–3 mm long, thinly
sericeous, bearing 2 or 3(-5) flowers. Bracts subtending
flowers ovate, 0.6–0.8 0.4–0.6 mm, outside thinly sericeous
or glabrous, inside glabrous. Pedicels 3–5.5 mm long, thinly
sericeous. Bracteoles at the top of the pedicels, ovate, ca.
0.6 0.4 mm, outside glabrous or thinly sericeous, inside
glabrous, margin ciliate. Calyx maroonish, cup-shaped, 2.5–
3.5 mm high, outside thinly sericeous, margin ciliate, inside
thinly sericeous in the upper part, tube 2–2.5 mm high, upper
lip indistinct or with 2 short lobes, 0.2–0.4 2–2.5 mm; lateral lobes triangular or semi-circular, 0.3–0.5 1.2–1.5 mm;
lower lobe triangular or semi-circular, 0.4–0.5 1.5–1.6 mm.
Corolla whitish. Standard: claw 3–3.5 mm long; blade
broadly obovate to orbicular, 4.5–5.5
4.5–5 mm, apex
slightly emarginate, basal callosities absent, both sides glabrous. Wings: claw 3–3.5 mm long; blade elliptic, 4–4.5
1.5–1.7 mm, apex rounded, upper auricle 0.7–0.8 mm long,
lower auricle indistinct, both sides glabrous. Keel petals: claw
2.7–3 mm long; blade boat-shaped, 4.7–5 1.6–2 mm, apex
rounded, upper auricle indistinct, lateral pocket ca. 1 mm
long, sometimes hardly visible, both sides glabrous or sometimes with scattered hairs at the apical part of the petal.
Stamens monadelphous, with basal fenestrae, 8–8.5 mm long,
free part 2–2.5 mm long, glabrous; anthers 0.3–0.4
ca.
0.3 mm, glabrous. Disc annular, more or less distinctly (5-)10lobed, up to 0.2 mm high, hairy. Ovary 7–8 mm long, stipe
indistinct, (thinly) sericeous; ovules 3 or 4; style 3–4 mm long,
thinly sericeous at base, upwards glabrous. Pods elliptic to
strap-like, 3.3–8 2.5–3.7 cm, glabrous, with a wing along
both sutures, upper wing 4–8 mm wide, lower wing 2–5 mm
wide; seed chamber absent. Seeds 1 or 2 per fruit, broadly
bean-shaped, flat, 15–17 13–14 0.5–1 mm; hilum subcentral, ca. 2 mm long. Figure 10.
Distribution—This species is endemic in Thailand (southeastern) (Fig. 7).
Habitat and Ecology—This species grows in (dry) evergreen forests or open thickets in the forest, up to 300 m.
Phenology—Flowering is from August to October.
Fruitingis from September to January
Note—The specimens, Maxwell 76–31 and Maxwell 73–415,
were originally indentified as Aganope thrysiflora, because of
the highly branching inflorescences and two-winged pods.
Another two specimens from Kew, Collins 275 and Collins
279 (see also Craib 1927) were identified as D. marginata.
These four specimens were collected from the same locality
(Sriracha district, Chon Buri province) and they show the
same morphological characters. The first author studied
these specimens in more detail. However, characters such as
leaflets, inflorescences and flowers are all completely unlike
A. thrysiflora and D. marginata. The inflorescences of this
species are pseudopanicles, the brachyblasts vary between
relatively long to short to knob-like, and sometimes they are
hardly visible. In A. thrysiflora and D. marginata, there are
no brachyblasts and, therefore, the flowers are all solitary.
Pedicels of this species are distinctly longer than those of
A. thrysiflora, but distinctly shorter, thicker, and with more
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elliptic or ovate, 0.5–0.7
0.4–0.5 mm, outside sericeous,
inside glabrous. Calyx cup-shaped, 3–3.5 mm high, outside
sericeous, inside glabrous, tube ca. 3 mm high; upper lip with
2 short lobes, 0.3–0.4 1.8–2 mm; lateral lobes triangular,
0.5–0.7 1–1.5 mm; lower lobe triangular, ca. 1 1.5–2 mm.
Corolla pink or purple (blue according to Thothathri 1982).
Standard: claw 1.5–2 mm long; blade orbicular or ovate, 10–
12 8.5–10 mm, apex emarginate, basal callosities indistinct,
both sides glabrous. Wings: claw 2.5–2.8 mm long; blade
2.5–2.7 mm, apex rounded,
narrowly obovate, 7.3–7.5
upper auricle 0.5–1 mm long, lateral pocket 1–1.5 mm long,
sometimes inconspicuous, both sides slightly sericeous at the
apex. Keel petals: claw 2.5–2.8 mm long; blade boat-shaped,
3.4–3.5 mm, apex rounded, auricle hardly visible,
7–7.2
lateral pocket 1.5–2 mm long, glabrous. Stamens monadelphous, with basal fenestrae, 9–10 mm long, free part of filaments 1–3 mm long, glabrous; anthers 0.5–0.9 0.3–0.5 mm,
glabrous. Disc annular, 0.2–0.3 mm high, sometimes hardly
visible. Ovary hairy, 6–7.5 mm long, stipe indistinct; ovules 4
or 5; style 2.5–4.5 mm long, glabrous but slightly hairy at
base. Pods (Thothathri 1961) narrowly elliptic or strap-like,
flat, glabrous, 7–10 1.5–2.5 cm, with a wing along both
sutures, upper wing wider than lower. Seeds unknown.
Distribution—This species occurs in India, Bangladesh,
Myanmar and northern Thailand (Fig. 7).
Habitat and Ecology—A species from evergreen forests,
720–2,000 m.
Phenology—In Thailand, flowering has been reported in
March. Fruiting is still unknown.
Notes—This species is rare in Thailand. Only one flowering
specimen was collected by A. F. G. Kerr in Chiang Mai, northern Thailand. Unfortunately, we were not able to study the
holotype of Millettia monticola Kurz deposited in CAL.
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SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
421
Fig. 10. Derris pseudomarginata Sirichamorn. a. Habit. b1. Longer brachyblast with flower buds, usually on the basal position of the inflorescence. b2.
Short or hardly visible brachyblast on the apical position of the inflorescence. c. Flower. d. Standard, inside view. e. Wing petal. f. Keel petal. g. Stamens.
h. Pistil. i. Pod [a-i: Collins 275, K]. Drawing by Anita Walsmit Sachs (L).
SYSTEMATIC BOTANY
Derris mindorensis Perkins, Fragm. Fl. Philip. 2: 82. 1904.
Merr., Enum. Philip. 2: 300. 1923.—TYPE: PHILIPPINES.
Mindoro, Palauan, Merrill 953 (holotype: B or PNH, lost;
isotype: K!).
Derris micans Perkins, Fragm. Fl. Philip. 2: 82. 1904. Merr.,
Enum. Philip. 2: 300. 1923.—TYPE: PHILIPPINES.
Luzon, Tanay, Merrill 2284 (holotype: B or PNH, lost;
isotypes: K!, US!).
Derris ferruginoides Quisumb., Philip. J. Sca. 41: 324. 1930.—
TYPE: PHILIPPINES. Luzon, Clemens 7406a (holotype: UC!).
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Representative
Specimens
Examined—INDONESIA.
Celebes:
Minahassa (Menado), 1897, Koorders 17668 (L); Lesser Sunda Is.:
W. Flores, 30 May 1975, Schmutz 3804 (L); Moluccas: Pulau Trangan,
Kerei, 29 Oct 1994, van Balgooy 6722 (L); Sumatra: Palembang, Bosch
Talangbobang, 29 Oct 1929, van Steenis 3382 (L). MALAYSIA. Perak:
Mount of Phang Nga, near Tapah, 26 Jul 1941, Ward SFN 37585 (L); Sabah:
Lahad Datu, Takun, Kennedy Bay, section 42, 5 Sep 1961, Chai SAN 26076
(L); Tambunan, 15 Mar 1969, Nooteboom 1290 (L); Ulu Biah, Keningau
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Lianas. Bark slightly pinkish. Twigs 3–5 mm diam., sericeous or glabrescent. Leaves with 5–9 leaflets. Stipules triangular, 1.5–2 1.5–2 mm, outside sericeous, inside glabrous.
Petiole 1.5–6 cm long, striate, hirsute or thinly hirsute; rachis
3.5–9 cm long, striate, (thinly) hirsute; pulvinus 5–10 mm
long. Stipellae absent. Leaflets subcoriaceous; terminal one
elliptic to obovate, 5.5–12.5 2.2–5 cm, length/width ratio
ca. 2.5, base cuneate, obtuse to rounded, apex acuminate,
acumen 2–13 mm long, rounded, upper surface glabrous to
slightly hairy, lower surface strigose to almost glabrous, midrib slightly sunken above, veins slightly raised or flat above,
raised below, 4–9 per side, 5–18 mm apart, not reaching the
margin but curving towards the apex, sometimes anastomosing near the margin, venation usually reticulate but
sometimes slightly scalariform; lateral ones mostly as terminal one, elliptic to ovate, 3.5–11
1.7–4.5 cm, length/
width ratio 2.1–2.4; pulvinus 3–6 mm long. Pseudoracemes/
pseudopanicles axillary or terminal, 5–28 cm long, peduncle
1.5–5 cm long, sericeous, branches up to 19 cm long. Bracts
subtending brachyblasts ovate-triangular, 0.8–1.5 0.6–1.0 mm,
outside sericeous, inside glabrous. Brachyblasts knob-like
to slenderly cylindrical, 2–25 mm long with 5–7 flowers
throughout, pubescent. Bracts subtending flowers elliptic to
ovate or triangular, 0.6–0.7 0.3–0.5 mm, outside pubescent,
inside glabrous. Pedicels 2–6 mm long, pubescent. Bracteoles
at the base of the calyx, elliptic, 0.5–0.8 0.3–0.5 mm, outside pubescent, inside glabrous. Calyx cup-shaped, reddish
brown to purplish, 3.5–4 mm high, outside sericeous, inside
sericeous at the apex, tube ca. 3 mm high; upper lip with
2 short lobes, shallowly notched, 1–2 2–5 mm; lateral lobes
semicircular to rounded triangular, 0.7–1 2–3 mm, lower
lobe triangular, ca. 1 1.5–3 mm. Corolla white or pale green.
Standard: claw 3–4 mm long; blade orbicular or ovate, 6–8
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Derris acuminata Miq., Fl. Ned. Ind. 1: 144. 1855. Benth., Proc.
Soc. Lond. 4 (Suppl.): 115. 1860. Backer & Bakh. f., Fl.
Java 1: 619. 1964.—TYPE: INDONESIA. Sumatra,
Korthals s. n. (lectotype here designated: L!, barcode
L0476235; isolectotype: U!, barcode U 0139545).
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10. Derris pubipetala Miq., Fl. Ned. Ind. 1: 145. 1855. Benth.,
Proc. Linn. Soc. Lond. 4 (Suppl.): 115. 1860. Adema,
Blumea 48: 401. 2003.—TYPE: INDONESIA. Java, Blume
1286 (holotype: L!, barcode L 0476233; isotypes: A, K, L!).
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Representative Specimens Examined——THAILAND. Chon Buri:
Sriracha, Khao Khiew, 21 Jan 1976, Maxwell 76–31 (L).
8–9 mm, apex emarginate, without auricle or basal callosities, outside sericeous except for the margin and base,
mostly hairy in the middle and apical part, inside glabrous
but sericeous at the apex. Wings: claw 3–3.5 mm long; blade
elliptic to semi-hastate, 7–8.5 1.5–2.5 mm, apex rounded,
upper auricle 1–2.5 mm long, lower auricle rarely present,
0.4–0.5 mm long, outside hairy at the apex, inside glabrous,
lateral pocket inconspicuous. Keel petals: claw 2.3–2.5 mm
long; blade boat-shaped, 6–6.5 2.3–2.5 mm, apex rounded,
auricle 0.5–1 mm long, outside hairy at the apex, inside glabrous, lateral pocket 1.5–2.7 mm long, sometimes hardly visible. Stamens monadelphous, with basal fenestrae, 9–10 mm
long, free part of filaments 2–3.5 mm long, glabrous; anthers
0.3–0.5 0.3–0.4 mm, glabrous. Disc annular, 0.1–0.3 mm high.
Ovary hairy, 5–6.5 mm long, stipe hairy, ca. 1 mm long; ovules
3–5; style 5–8 mm long, glabrous but slightly hairy at base.
Pods ellipsoid to almost circular, 3–6.5 1.8–3 cm, leathery
with wing along both sutures, flat, upper wing wider than
lower, 5–9 mm wide, lower wing 2–4 mm wide, pubescent,
seed chamber hardly visible. Seeds 1 or 2 per pod, discoid
or broadly bean-shaped, 7–8 5–6.5 1–2 mm; hilum central,
0.4–0.5 mm long.
Distribution—This species occurs in Thailand (Peninsular); Malesia: Peninsular Malaysia (Perak), Sumatra, Borneo,
Philippines (Luzon, Mindanao, Mindoro, Palawan), Celebes,
and Moluccas (Fig. 11).
Habitat and Ecology—A species from primary, either disturbed or secondary wet evergreen forest, near river banks or
seashore. Soil: Limestone, clay or sandstone, up to 1,000 m.
Phenology—Flowering is from May to October. Fruiting is
from April to July, or November.
Uses—Roots said to be used as fish poison.
Notes—The species is variable in the density of its
indumentum, and similar in most characters to the more
northern species, D. ferruginea (see also the note under
D. ferruginea).
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hairs than those of D. marginata. The flowers are slightly
smaller than those of A. thrysiflora and D. marginata, and are
more similar to Derris flowers. The morphological characters
of this species are unique and, therefore, it is to describe them
as a new species. Future molecular phylogenetic evidence
will have to prove the taxonomic status of this taxon.
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Fig. 11. Distribution of Derris pubipetala Miq.
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
423
District, 9 Oct 1976, Sundaling SAN 83906 (L). PHILIPPINES. Luzon:
Dipaguidin, near Palanan Point, 18 Apr 1991, Ridsdale et al. ISU 54 (L);
Laguna, Feb-Mar 1917, Catalan 26463 (L); Aurora, Baler, N. of Gupa
Beach, 3 May 1992, Stone et al. 5492 (L); Mindanoa, Sep 1909, Elmer 11738
(L); Palawan: Hinabaan, Guinayangan, Quezon, 27 Oct 1967, Mendoza
PNH 97918 (L); Mindoro: E of Mindoro, beside quarry track, Mt.
Talipana, 7 km SW of Puerto Galera, 29 Apr 1987, Burley 193 (L); Paluan,
Apr 1921, Ramos 39669 (L); Puerto Galera, 12 Jul 1955, Sulit 32817 (L);
Palawan, Apr 1911, Elmer 13078 (L). THAILAND. Phuket: Hill near
Thalang, E. of the road to Phuket, 8 May 1968, van Beusekom and
Phengkhlai 665 (L); Pattani: Koke Po, Sai Khao Falls Forest Park, 5 Apr
1985, Maxwell 85–370 (L).
11. Derris reticulata Craib, Kew Bull. 384. 1927. Fl. Siam.
Enum. 1: 435, 493. 1928.—TYPE: THAILAND. Kanburi
(= Kanchanaburi), 4 Jan 1926, Kerr 10142 (holotype: K!;
isotype: BK!).
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Lianas, basal diameter 3–10 cm. Bark thin, smooth, light
brown. Twigs 3–5 mm diam., glabrous, lenticellate. Leaves
usually with 5(–7) leaflets Stipules caducous, triangular,
1.2–1.5 1.4–1.5 mm, outside sericeous, inside glabrous. Petiole 2–6 cm long, grooved above, glabrous or with some
scattered hairs, rachis 2.5–5 cm long, pulvinus 4–6 mm long.
Stipellae absent or reduced to hairy knob-like structures.
Leaflets: subcoriaceous, terminal one (narrowly) elliptic to
(narrowly) obovate, 6.5–13 2.5–4 cm, length/width ratio
2.6–3.3, base cuneate to obtuse, apex acuminate, acumen ca.
9–15 mm long, rounded, upper surface glabrous, lower surface glabrous except along the midrib with some scattered
hairs, midrib flat or slightly raised above, distinctly raised
below, veins flat above, raised below, 6–11 per side, 8–13 mm
apart, not reaching the margin but curving towards the
apex, sometimes anastomosing near the margin, venation
reticulate; lateral ones (narrowly) elliptic to (narrowly) ovate,
5.5–12 1.5–4 cm, length/width ratio 2.2–3.6, base broadly
cuneate to obtuse, apex acuminate, acumen 5–7 mm long,
rounded, above glabrous, below glabrous except along the
midrib with some scattered hairs; pulvinus 3–4 mm long.
Pseudoracemes/ pseudopanicles axillary or (rarely) terminal, 2.5–5.5 cm long, striate, sericeous; peduncle up to
5 mm long, striate, sericeous; lateral branches up to 3.5 cm
long. Flowers (known only from some photographs therefore floral information partly available) white. Brachyblasts
knob-like to elongate and cylindrical, ca. 9 mm long, (thinly)
sericeous, with 2–4 flowers throughout, if 2-flowered, then
flowers sometimes seemingly at apex of the brachyblast.
Pedicels ca. 7 mm long, (thinly) sericeous. Bracteoles in
the middle or upper part of the pedicels. Calyx reddish or
greenish-red, cup-shaped, ca. 2.5 mm high, outside (thinly)
sericeous or glabrescent. Standard: blade obovate, ca. 9
7 mm, apex emarginate, without basal callosities, both sides
glabrous. Wings curved backward to the calyx. Keels
unknown. Stamens monadelphous, with basal fenestrae, ca.
10 mm long. Ovaries unknown. Pods strap-like or sometimes ellipsoid, 5–9.5 1.5–2.5 cm, thin, with a wing along
both sutures, upper wing 4–7 mm wide, lower wing 2–3 mm
wide, with some scattered short hairs, especially on both
wings, or sometimes glabrescent, seed chamber indistinct,
but usually slightly thickened and with reticulate veins
around the seed. Seeds unknown.
Distribution—This species is endemic to Thailand (Fig. 12).
Habitat and Ecology—This species is found in semishaded area of dry evergreen forests, edge of evergreen
mixed (dipterocarp) forest, in bamboo forest, or along
streams, 50–450 m.
Fig. 12. Distribution of Derris reticulata Craib (circle) and D. tonkinensis
Gagnep. (square).
Phenology—Flowering is from August to September.
Fruiting is from September to January.
Uses—The stems contain some sweet chemical substance
and are used as a sweetener in local medicine; also used
as a laxative.
Vernacular Names—Thailand: Cha aim nua, Oi sam suan.
Notes—Derris reticulata is a mysterious species. It was first
described in 1927 (Craib 1927: 384) from a single specimen
with short inflorescences and a 2-winged pod, which is reticulate around the seeds, while some leaflets are narrowly
elliptic and almost glabrous. The Thai name “Cha aim nua”
was mentioned on labels of type specimens by Kerr, who
collected the type specimen in Kanchanaburi and which
means “Licorice of the North.” The Thai name implies that
the local people knew the sweetening quality of the stems
and that they used them in local medicine for a long time.
Later, the use has been forgotten, perhaps because the species became scarce in Kanchanaburi or was difficult to find,
as no more collections were found. From field expedition in
2009 the first author found some unidentified Thai specimens (see below), which all have short inflorescences with
2-winged pods which are reticulate around seeds, and variously shaped, almost glabrous leaflets. These specimens
were collected from central and eastern Thailand, which is
far away from the type locality. However, the fruiting time
is almost exactly the same. Moreover, these unidentified
specimens were called “Oi sam suan,” which is Thai for
“Three fields sugarcane.” It is possible that these specimens
also contain sweet substances in their stems. In September
more fruiting specimens were collected together with data
from local people and a researcher at Sakaerat Environmental Research Station in Nakhon Ratchasima province. Unfortunately, it was too late to collect flowering specimens.
However, some photographs of the flowers were taken by a
researcher of this environmental research station, Mr. Buasai
Somsung. The photos were here used for a qualitative
description. This species has as typical character of the
wing petals curved backwards to the calyx (see note on
morphological characters). This character is also found in
SYSTEMATIC BOTANY
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Brachypterum polyphyllum Miq., Fl. Ned. Ind. 1: 139. 1855.
Derris polyphylla (Miq.) Benth., Proc. Linn. Soc. 4 (Suppl.):
104. 1860. Koord. & Valeton, Boomsoort. Java 2: 89. 1895.
Koord., Exkursionsfl. Java 2: 393. 1912. Backer & Bakh.
f., Fl. Java 1: 618. 1964.—TYPE: INDONESIA. Java,
Soerakarta, Horsfield s. n. (L16) (holotype: U!, barcode
U 0103949; isotypes: CAL, K!,).
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Deciduous trees, 9–20 m high, dbh 20–25 cm. Bark smooth,
pale grayish-brown. Twigs terete, 1–5 mm diam., strigose.
Stipules acicular, 4–5.5 0.3–0.5 mm, sericeous. Leaves with
11–19 leaflets. Petiole 2.5–3 cm long, striate, grooved above,
sericeous; rachis 10–13 cm long, striate, grooved above, sericeous; pulvinus 3–7 mm long. Stipellae acicular, 0.7–2.5 mm
long, sericeous. Leaflets subcoriaceous; terminal one elliptic,
2.6–6 1–2.5 cm, length/width ratio ca. 2.5, base cuneate,
apex acute, mucronate, upper surface thinly sericeous or
nearly glabrescent, lower surface sericeous, midrib and veins
slightly raised or flat above, raised below, veins 4–6 per side,
3.5–7 mm apart, curving towards the apex, anastomosing
near or sometimes at the margin, venation reticulate; lateral
ones mostly as terminal one, ovate to elliptic, 1.4–6.5 1–1.7 cm,
usually oblique, basiscopic side narrower; pulvinus 2–3 mm
long. Pseudoracemes axillary, 5.5–27.5 cm long, peduncle
2–5.5 cm long, sericeous. Bracts subtending brachyblasts narrowly ovate, ca. 0.5 1–2.2 mm, outside sericeous, inside glabrous, sometimes caducous. Brachyblasts slender 2–3.5 mm
long, 3–7-flowered, if 3-flowered, then all flowers usually at
the apex of the brachyblast, if more than 3 flowers or if
the brachyblasts short, then the flowers seemingly more
0.3–
scattered. Bracts subtending flowers elliptic, 0.7–0.8
0.4 mm, caducous, outside sericeous, inside glabrous. Pedicels 3–4.5 mm long. Bracteoles at the base of the calyx, elliptic,
ca. 0.3 mm, outside sericeous, inside glabrous.
0.6–0.7
Calyx 2.5–3 mm high, outside sericeous, inside glabrous,
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12. Derris robusta (Roxb. ex DC.) Benth., Proc. Linn. Soc.
4 (Suppl.): 104. 1860. Baker in Hook. f., Fl. Brit. Ind. 2:
241. 1878. Craib, Fl. Siam. Enum. 1: 435, 493. 1928. Backer &
Bakh. f., Fl. Java 1: 618. 1964. Thoth., Fasc. Fl. India 8:
26. 1982. P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 70. 2001. Dalbergia robusta Roxb. [Hort. Beng.
53. 1814, nom. nud.] ex DC., Prod. 2: 417. 1825. Deguelia
robusta (Roxb. ex DC.) Taub. in Engl. & Prantl, Nat.
Pflanzenfam. 3, 2: 345. 1891.—TYPE: INDIA. Silhet,
Wallich 5849B (lectotype designated by Thothathri 1982:
CAL; isolectotype: K!).
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Representative Specimens Examined—THAILAND. Chaiyaphum:
Dat Don, 13 Aug 1972, K. Larsen et al. 31804 (L); Nakhon Ratchasima:
Wang Nam Kheaw, Sakaerat Environmental research station, 17 Sep
2009, Sirichamorn YSM 2009–18 (L); Saraburi: Muak Lek, 31 Aug 1924,
Kerr 9084 (L).
tube 2–2.5 mm high; upper lobes triangular, 0.5–0.7
2–
2.5 mm; lateral lobes triangular, 0.5–0.7 1.5–1.7 mm; lower
lobe triangular, ca. 0.5 1.5–1.7 mm. Corolla white. Standard:
claw 1.5–2 mm long; blade orbicular, 6–7.5 5.5–5.7 mm, apex
emarginate, basal callosities small or indistinct, both sides
sericeous at apex. Wings: claw 2–2.5 mm long; blade elliptic,
5–5.5 1.5–1.7 mm, apex rounded, lateral pocket up to 1 mm
long, sometimes hardly visible, auricle indistinct, both sides
glabrous or with few hairs at apex. Keel petals: claw 2–3 mm
long; blade boat-shaped, 5–6.5 2–2.5 mm, apex rounded,
lateral pocket 1–1.5 mm long, glabrous or with few hairs at
apex. Stamens monadelphous, with basal fenestrae, 7–8 mm
long, free for ca. 2 mm, glabrous; anther 0.3–0.4 mm long,
glabrous. Disc tubular, 0.3–0.4 mm high. Ovary ca. 5 mm long,
stipe ca. 0.5 mm long, sericeous; ovules 7–11; style 2–2.5 mm
long, glabrous and slightly sericeous at base. Pods strap-like,
1.5–8.5 0.7–1.1 mm, flat, sericeous, 4–8 seeds per pod, with a
wing along the upper suture, ca. 2 mm wide; seed chamber
slightly solid, visible from outside. Seeds ellipsoid, 4–4.5 3–
5 0.6–1.5 mm; hilum central, 0.4–0.5 mm long.
Distribution—This species occurs in India, Myanmar,
China, Laos, Thailand, and southward though Java, Sulawesi,
and Sumatra (Fig. 13).
Habitat and Ecology—A species from disturbed, degraded,
fire-prone areas, deciduous forest with much bamboo, or
dry evergreen forest, up to 1,000 m.
Phenology—In Thailand flowering is reported from March
to May, from July to October for other regions. Fruiting is
from August to November.
Uses—Wood is used for making tea boxes, shade trees and
green manure (Hamid 1999: 241; Thothathri 1982: 27).
Vernacular Names—Indonesia: Bekel, Kedusan, Wedusan
(Javanese). Thailand: Khang sai chang (Pitsanulok), Khee mord,
kang khee mord (Saraburi), Haang kow, Pee chan (Northern),
Lawit tua mea (Chanthaburi). Vietnam: Cóc kèn, kra mui.
Notes—This is a deciduous species that is present in many
vegetation types. In Thailand, it is mainly found in the north
and less commonly so in other parts of the country. However, some specimens are found in the southeast of Thailand,
e.g. Chanthaburi province. Specimens from these two areas
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D. tonkinensis. It is reported that the flowering period was
short, between August (or late July) to September, after that
the flowers easily drop. Because of its short inflorescences
and brief, unpredictable flowering period, it may be hard to
find, which explains why there are no flowering specimens in
herbaria and why local people use the plant less in medicine.
We decided use the name D. reticulata for all unidentified
specimens with 2-winged pods that are thickened and reticulate at the seeds, and which have almost glabrous 5-foliolate
leaves, short pseudoraceme or pseudopanicle inflorescences
(less than six cm long), sweet stems, and flowers in August
or September.
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Fig. 13. Distribution of Derris robusta (Roxb. ex DC.) Benth.
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
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Dalbergia timorensis DC., Prod. 2: 417. 1825. Brachypterum
timorensis (DC.) Benth. ex Miq., Fl. Ned. Ind. 1: 138.
1855. Derris timorensis (DC.) Pittier, Contr. U. S. Nat.
Herb. 20: 41. 1917.—TYPE: not indicated (P or G-DC).
Dalbergia venusta Zipp. ex Span., Linnaea 15: 197. 1841.—
TYPE: LESSER SUNDA ISL. Timor, Zippelius s. n. (holotype: L!, barcode L 0475849; isotype: L!, barcode L
0475873).
Millettia litoralis Dunn, Philip. J. Sca., Bot. 6: 316. 1912. Linn.
Soc. Bot. 41: 181. 1912. —TYPE: PHILIPPINES. Mindanao,
St. Cruz, DeVore and Hoove 250 (holotype: K!).
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Lianas, basal diameter 2–5 cm. Twigs 3–5 mm diam., thinly
strigose to glabrous, lenticels throughout. Leaves with 9–13
leaflets. Stipules triangular, 1–1.5 0.9–1.3 mm, outside sericeous, inside glabrous. Petiole 2–5 cm long, grooved above,
thinly hirsute or sometimes glabrescent; rachis 5.5–10 cm
long, grooved above, (thinly) hirsute to glabrous; pulvinus
3–7 mm long. Stipellae present or sometimes inconspicuous,
acicular, up to 1.5 mm long. Leaflets subcoriaceous; terminal
one elliptic to obovate, 4.5–8 1.5–3.2 cm, length/width ratio
2.5–3, base cuneate to broadly cuneate or rounded, apex
obtuse, usually emarginate or shortly obtusely acuminate,
acumen 2–6 mm long, emarginate to rarely rounded, upper
surface shiny, glabrous to slightly strigose along midrib,
lower surface thinly strigose, midrib and veins slightly raised
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13. Derris scandens (Roxb.) Benth., Proc. Linn. Soc. 4,
(Suppl.): 103. 1860. Gagnep. in Lecomte, Fl. Indo-Chine
2: 415. 1916. Ridl., Fl. Malay Penins. 1: 595. 1922. Merr.,
Enum. Philipp. 2: 301. 1923. Craib, Fl. Siam. Enum. 1:
492. 1928. Thoth., Bull. Bot. Surv. India 3: 177. 1961.
Backer & Bakh. f., Fl. Java 1: 618. 1964. Thoth., Fasc. Fl.
India 8: 27. 1982. Dalbergia scandens Roxb., Pl. Coromandel 2: 49, pl. 192. 1805. Wight & Arnott, Prod. 1: 264.
1834. Brachypterum scandens (Roxb.) Benth., Ann. Wien.
Mus. 2: 101. 1837. Miq., Fl. Ned. Ind. 1: 138. 1855.—
TYPE: INDIA. Roxburgh, Pl. Coromandel 2: 49, pl. 192.
1805 (lectotype designated by Thothathri 1982: CAL).
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Representative Specimens Examined—INDIA. Assam: Takubama,
Naga Hills, 1–9 Sep 1950, Chand 3618 (L); Cherrapunjee, Khasi Hills,
16 Jul 1952, Koelz 30289 (L). INDONESIA. Celebes: 8 Apr 1896, de Wit
I. B. 43 (L); Java: Semarang, Kedoengdjati, 2 Nov 1896, Koorders 25510 (L);
Sumatra: Palembang, 19 May 1941, de Wit I. F. 3 (L). LAOS. Pakse: North of
Paksong, Bassac, 26 Sep 1928, Poilane 15732 (L); Sayaboury: Phinag,
Nampouy village area, near Nam Pouy River, 29 Aug 1999, Maxwell 99–
236 (L). THAILAND. Chanthaburi: Pong Namrawn, Khao Soidao Tai,
12 Feb 1959, Smitinand 5496 (L); Chiang Mai: Doi Sutep-Pui National
Park, south side, Mae Heeyah Nai Village area, Muang district, 13 Feb
1990, Maxwell 90–198 (L); Chiang Rai: Doi Laung, Roadside to Ban Mai
Pattana, 27 Aug 2009, Sirichamorn YSM 2009–10 (L); Lampang: Muang
Pan, Jae Sawn National Park, park headquarters, 30 May 1996, Maxwell
96–752 (L); Mae Hong Son: 18 Jun 1973, Geesink et al. 5951 (K);
Phitsanulok: Kaeng Sopha, 28 Jun 1967, Phusomsaeng 265 (L); Phrae,
16 Mar 1961, Phengkhlai 18 (L); Uttaradit: Huay Hin, 15 Apr 1961,
Phengkhlai 119 (K).
or flat above, raised below, veins 7–12 per side, 3–5 mm
apart, curving towards the apex and sometimes reaching the
margin, sometimes anastomosing near the margin, venation
reticulate; lateral ones mostly like terminal, elliptic to ovate,
rarely obovate, 3–6.8 1.2–3.5 cm, length/width ratio 1.9–
2.5; pulvinus 1.5–5 mm long. Pseudoracemes axillary, rarely
several together as pseudopanicles, 5–28 cm long, peduncle
1.5–5 cm long, thinly strigose. Bracts subtending brachyblasts
ovate-triangular, 0.7–1 0.5– 0.7 mm, outside sericeous, inside
glabrous. Brachyblasts knob-like, 2–6 mm long, 5–10-flowered,
sericeous. Bracts subtending flowers elliptic to ovate, 0.6–
0.7 0.3–0.5 mm, outside sericeous, inside glabrous. Pedicels 5.5–9 mm long, sericeous. Bracteoles at the base of the
calyx, elliptic or ovate, 0.5–0.8 0.3–0.7 mm, outside pubescent, inside glabrous. Calyx reddish or greenish-red, cupshaped, 2.5–2.7 mm high, outside sericeous, inside glabrous,
tube ca. 2 mm high; upper lip and lateral lobes hardly visible;
lower lobe triangular, ca. 0.7
1.5–2 mm. Corolla white
or pale pink. Standard: claw 2–3 mm long; blade orbicular or
obovate, 5–8 6–8 mm, apex emarginate, without auricle
or basal callosities, outside glabrous or sometimes slightly
sericeous at apex, ciliate in lower part to up to halfway,
inside glabrous. Wings: claw 2–2.5 mm long; blade narrowly obovate, 4.5–6.5 1–1.5 mm, apex rounded, outside
slightly sericeous at the apex and distinctly strigose on both
auricles, inside glabrous, upper auricle ca. 0.5 mm long, lower
auricle hardly visible, lateral pocket inconspicuous. Keel
petals: claw 2.3–2.5 mm long; blade boat-shaped, 6–6.5 2.3–
2.5 mm, apex rounded, outside hairy at the apex, sometimes
also along the major veins ventrally, inside glabrous, auricle
and lateral pocket hardly visible. Stamens monadelphous, with
basal fenestrae, white, 9–12 mm long, free part of filaments
1.5–3.5 mm long, glabrous; anthers 0.3–0.5
0.3–0.4 mm,
glabrous. Disc short-tubular, more or less distinctly (5-)10lobed, up to 0.6 mm high. Ovary hairy, 4–5 mm long, stipe
hairy, ca. 1 mm long; ovules 8–10; style 4–5.5 mm long,
glabrous but slightly hairy at base. Pods strap-like, 4.5–9
0.7–1.5 cm, flat, bulging around the seeds, with a wing along
the upper suture only, wing 1–2 mm wide, thinly sericeous;
seed chamber hard, usually visible from outside. Seeds beanshaped, flat, ca. 7 5 1 mm; hilum central, ca. 1 mm long.
Figure 14.
Distribution—Derris scandens is widespread species ranging from Reunion island to India, Sri Lanka, Bangladesh,
Myanmar, thoughout Indochina, Thailand, Malay Peninsular, Sumatra, Borneo, Java, Philippines, Celebes, Lesser
Sunda Islands, Papua New Guinea (Morobe Prov.), and
Australia (Fig. 15).
Habitat and Ecology—This common species grows in
various types of vegetation: open and disturbed or abandoned cultivated areas, shrubbery, agricultural areas, edges
of rice fields, grasslands, or beach or mangrove forest, up
to 750 m.
Phenology—Flowering is from July to September, or
throughout the year. Fruiting is from December to August.
Uses—Derris scandens is used as a fish poison. In Thailand
stems or roots are used as local medicine for the treatment of
osteoarthritis, common cold or backache, also as a diuretic,
laxative, expectorant and emmenagogue. Sometimes it is cultivated as an ornamental plant (Hamid 1999: 241).
Vernacular Names—Cambodia: Priang (Kampot). Indonesia:
Bendan, Gobul (Javanese), Sobi (Madura). Laos: Ta pa
(Savannakhet). Philippines: Malasaga (Tagalog), Lapak
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are morphologically similar and treated as the same species.
A disjunct northern – southeastern distribution is not uncommon (van Welzen pers. comm.).
When the brachyblasts are short or when there are more
than three flowers on a brachyblast, then the flowers are
seemingly more scattered over the brachyblast. In some rare
cases, when there are three flowers at the apex of the
brachyblasts, then the specimens may easily be mistaken for
the genus Paraderris. However, the three-flowered condition
has never been found in Thailand so far.
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SYSTEMATIC BOTANY
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Fig. 14. Derris scandens (Roxb.) Benth. a. Habit. b. Brachyblast with flower buds. c. Flower. d. Standard, inside view. e. Wing petal. f. Keel petal.
g. Stamens. h. Pistil with disk. i. Pod [a-h: Maxwell s. n., barcode L 0475773; i: Maxwell 87–15; all L]. Drawing by Anita Walsmit Sachs (L).
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
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nate, both sides slightly sericeous. Leaves with 13–19 leaflets.
Petiole 3.5–4.5 cm long, grooved above, sericeous to glabrescent, rachis 17–20 cm long, sericeous to glabrescent, pulvinus 4–6 mm long. Stipellae acicular, 2.5–3.5 mm long,
glabrous or with some hairs. Leaflets chartaceous to subcoriaceous; terminal one narrowly obovate, 6–6.5 2–2.3 cm,
length/width ratio 2.8–3, base cuneate, apex obtuse to slightly
acuminate, acumen 0.2–0.5 mm long, mucronate, upper
surface slightly sericeous or sometimes glabrescent, lower
surface sericeous, midrib slightly sunken above, veins flat
above, raised below, 9–12 per side, 3–5.5 mm apart, curving
towards the apex, anastomosing near or sometimes at the
margin, venation reticulate; lateral leaflets narrowly obovate
or narrowly elliptic, 4–7.5 1.5–2.3 cm, length/width ratio
2.6–3.3, base rounded to cordate, apex obtuse to slightly
acuminate with mucronate tip, above slightly sericeous or
sometimes glabrescent, below sericeous, venation reticulate;
pulvinus ca. 2 mm long. Pseudoracemes axillary, rarely
form pseudopanicles, 3–6 cm long, striate, (thinly) sericeous;
peduncle up to 1.5 cm long, striate, (thinly) sericeous. Bracts
subtending brachyblasts narrowly ovate, 1.5–1.6 0.3–0.5 mm,
outside sericeous, inside glabrous. Brachyblasts knob-like,
indistinct, 0.1–0.4 mm long, with 2 or 3 flowers. Bracts
subtending flowers narrowly ovate to acicular, 1.5–4 0.2–
0.4 mm, outside pubescent, inside glabrous. Pedicels 4.5–
5 mm long, sericeous. Bracteoles on the upper part or at
the top of the pedicels, narrowly ovate, 1.3–1.5 0.3–0.4 mm,
outside pubescent, inside glabrous. Calyx greenish, cupshaped, 3–3.5 mm high, outside thinly sericeous, inside
glabrous, tube 2.8–3 mm high; upper lobes triangular, 0.5–
0.9
1.3–1.5 mm; lateral lobes triangular, 0.4–0.5
1.2–
1.7 mm; lower lobe triangular, 0.7–0.8 1.5–1.7 mm. Corolla
pale greenish-white. Standard: claw 1.5–1.7 mm long; blade
elliptic or ovate, 11–12 9–11 mm, apex emarginate, basal
callosities indistinct, both sides glabrous. Wings: claw 1.8–
2 mm long; blade narrowly elliptic, 7.8–8 1.8–2 mm, apex
rounded, auricle indistinct, lateral pocket ca. 0.5 mm long,
both sides glabrous but thinly ciliate at the basal part. Keel
petals: claw 2.3–2.5 mm long; blade boat-shaped, apex
rounded, 6.3–6.5 2.5–2.7 mm, both sides glabrous, lateral
pocket 1–2 mm long, auricle indistinct. Stamens monadelphous, with basal fenestrae, 9–10 mm long, free part of filaments 1.5–2 mm long, glabrous; anthers ca. 0.8 0.35 mm,
glabrous. Disc annular, more or less distinctly 5-lobed, up to
0.5 mm high. Ovary hairy, 6–6.5 mm long, stipe ca. 1 mm long;
ovules 7–12; style 2.5–3 mm long, hairy at base. Pods narrowly
ovate to strap-like, 5–11
ca. 2 cm, with thinly scattered
appressed hairs especially along suture, thin, veined, wing
along the upper suture only, ca. 2 mm wide; seed chamber usually visible from outside, membranous. Seeds ellipsoid, slightly flattened, 7.4–7.7 4–6 mm; hilum central,
1.5–1.7 mm long.
Distribution—This species is found in Laos and northern
Thailand (Fig. 7).
Habitat and Ecology—A species from open, fire prone
areas or hill slopes in deciduous, mostly dipterocarp forest
with bamboo, shale bedrock, 200–300 m.
Phenology—Flowering is from June to September. Fruiting
is from August to November
Vernacular Names—Thailand: Tao tab pla, Khruea tab pla,
Khruea ta pla, Khee chang tao, Khruea lai
14. Derris thorelii (Gagnep.) Craib, Fl. Siam. Enum. 1: 435,
493. 1928. P. K. Lôc & J. E. Vidal, Fl. Cambodge, Laos &
Vietnam 30: 56. 2001. Millettia thorelii Gagnep., Not. Syst.
(Paris) 2: 365. 1913.—TYPE: LAOS. De Xieng Khouang à
Pak Lai, 1866–1868, Thorel s. n. (lectyotype designated
by Phan in Phan and Vidal 2001: P!; isolectotype: P!).
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Lianas, basal diameter 2–4 cm. Bark thin, smooth, brown.
Stipules narrowly triangular, 9–10 1–2 mm, apex acumi-
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Representative Specimens Examined—AUSTRALIA. Queensland:
near Boonal, between Terriotville and Anthony, 30 Nov 1930, Hubbard
5355 (L). BRUNEI. Belait: Seria, 7 Aug 1965, van Niel 4065 (L). INDIA.
Andaman and Nicobar Islands: Middle Andaman, from Rangat to logging area ca. 6 km to the N.E. Much, 7 Jun 1977, Bhargara et al. 6438 (L);
Tamil Nadu: Kodamady, 23 Jul 1966, Shetty 27910 (L). INDONESIA. Java:
Besuki, SW. foot of Mt Baluran, 0–2 km from the road between
Banjuwangi (38.3 km) and Situbondo, 25 May 1957, Jacobs 4916 (L); Nusa
Tenggara Barat: Along dry riverbed, 8 km east of Sumbawa Besar, 9 Apr
1961, Kostermans 18047 (L); Nusa Tenggara Timur: Flores, 24 Dec 1974,
Verheijen 4538 (L); Sumatra: Serdang by Rantaripandjang, 13 Aug 1922,
Lörzing 9128 (L). LAOS. Champasak: Khong, Sahong Island, east side,
along the bank of the Maekong River, 15 Oct 1997, Maxwell 97–1186 (L);
Vientiane: Hatxiafong, Ban Khuay Daeng, forest 2–3 km from village,
7 Jul 1999, Soejarto and Sydara 10831 (L). MALAYSIA. Johore: Bukit
Kluang, Besut, 11 Sep 1955, Sinclair and Kiah bin Salleh S. F. 40815 (L);
Kedah: Pulau, Langkawi, opposite Timun, 19 Aug 1972, Chin 1836 (L).
PHILIPPINES. Leyte: Municip. Ormoc, Bgy. Danao, Lake Danao National
Park, Forest along unpaved road surrounding the lake, southest side
of the lake, 26 Jun 1993, Soejarto et al. 8063 (L); Luzon: W slope of
Mt. Malinaoc, near Buhi, Camarines Sur., 7 Apr 1987, Burley 88 (L);
Mindoro: Base SE slope of Mt. Yagaw, 16 Aug 1953, Conklin PNH 18726
(L); Palawan: Jul-Aug 1925, Cenabre FB 29993 (L). REUNION ISLAND.
9 Mar 1977, Cadet 5731 (K). SRI LANKA. Anuradhapura: Hinnukkiriyawa
off Kekirawa, 7 Aug 1973, Jayasuriya 1247 (L); Central Province: Matale
district, Ereula Tank ca. 5 miles ESE of Dambulla, 11 Oct 1974, Davidse
7401 (L). THAILAND. Ang Thong: Wat Giang, 15 Aug 1971, Maxwell 71–
481 (L); Chon Buri: Siracha, Si Chang island, east side of Kow Kwang,
9 Sep 1993, Maxwell 93–1023 (L); Khon Kaen, 24 Jul 1967, Bunma 37910 (L);
Nakhon Ratchasima: Pak Chawng, 4 Sep 1958, Smitinand 4849 (L); Pattani:
Muang, Ban Boo Boh, 6 Jan 1987, Maxwell 87–15 (L); Phetchaburi: Kaeng
Krachan, Kaeng Krachan National Park, along the road toward national
park headquarters, 9 Aug 2002, Middleton et al. 858 (L); Phuket: Ao Sane
beach, 10 July 1979, Niyomdham et al. 259 (L); Prachuap Khiri Khan: Pran
Buri, Kui Buri National Park, trail from park headquarters, 19 Aug 2002,
Middleton et al. 1214 (L); Ratchaburi: Khao Nam Tok, west of Huai Yang,
10 Aug 1966, K. Larsen et al. 1352 (P); Sakon Nakhon: Panna Ni Kom,
28 Jun 1932, Lakshnakara 1023 (L); Trat: between Trat and Roem Ngop,
wayside, 1 Aug 1973, Murata et al. T-17394 (L). EAST TIMOR. E. of Laleia
river, N. Timor., 13 Dec 1953, Van Steenis 18009 (L).
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(Bikol). Thailand: Tao wan priang (Central), Ta pla, Tao ta
pla, Khruea khao nang (Nakhon Ratchasima), Pan sanai
(Peninsular). Vietnam: (Dây) Thùc binh (Ninh Thuân)
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Fig. 15. Distribution of Derris scandens (Roxb.) Benth.
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2012]
Representative Specimens Examined—THAILAND. Chiang Mai: Doi
Suthep, 9 Apr 1909, Kerr 588 (K); Chiang Rai: Hui Pa Lai Doi Duan,
SYSTEMATIC BOTANY
15. Derris tonkinensis Gagnep., Not. Syst. (Paris) 2: 349.
1913; in Lecomte, Fl. Indo-Chine 2: 458, Fig. 44. 1916. F. C.
How, Acta Phytotax. Sin. 3: 220. 1954. P. K. Lôc & J. E.
Vidal, Fl. Cambodge, Laos & Vietnam 30: 77. 2001.—TYPE:
VIETNAM. Quang Ninh, rochers calcaires de la Baie de
Ha Long, 3 Jul 1885, Balansa 1189 (lectotype designated
by Phan in Phan and Vidal 2001: P!; isolectotype: K!).
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Lianas, basal diameter 2–4 cm. Bark thin, smooth, light
brown-gray, lenticellate. Stipules triangular, ca. 1.5
1–
1.5 mm, outside slightly hairy, glabrescent, inside glabrous.
Leaves with 5–7 leaflets. Petiole 1.8–4 cm long, grooved
above, glabrous, rachis 1.7–4 cm long, pulvinus 2–4.5 mm
long. Stipellae absent. Leaflets: subcoriaceous; terminal one
elliptic to obovate, 4.5–8.5 1.7–3.5 cm, length/width ratio
2.5–2.6, base broadly cuneate, apex acuminate, acumen up to
10 mm long, rounded, upper and lower surface glabrous or
thinly sericeous, midrib raised above, veins flat or slightly
sunken above, slightly raised below, 7–14 per side, 3.5–
10 mm apart, not reaching the margin but curving towards the
apex, venation reticulate; lateral ones elliptic to narrowly
1.5–2.7 cm, length/width ratio 2–2.6, base
elliptic, 3–7.2
broadly cuneate to rounded, apex acuminate, acumen 5–
7 mm long, rounded, above and below glabrous or thinly
sericeous, not glaucous; pulvinus 1.5–3 mm long. Pseudoracemes/pseudopanicles axillary or terminal, 6.5–20 cm
long, striate, (thinly) sericeous, glabrescent; peduncle 0.5–
2 cm long, striate, (thinly) sericeous, glabrescent; lateral
branches up to 11 cm long. Bracts subtending brachyblasts
ovate to triangular, 0.6–1 0.4–0.8 mm, outside pubescent,
inside glabrous. Brachyblasts club-shaped to elongate cylindrical, 2–13 mm long, with 3–8 flowers, sometimes
disappearing near the top of the inflorescence and then
flowers solitary. Bracts subtending flowers ovate with acute
0.3–0.4 mm, outside pubescent, inside glaapex, 0.4–0.7
brous. Pedicels 2–5 mm long, sericeous. Bracteoles at the top
of the pedicels, ovate, ca. 0.6 0.4 mm, outside pubescent,
inside glabrous. Calyx maroonish, cup-shaped, 2–3 mm high,
outside thinly sericeous, inside glabrous or sericeous along
the margin, tube 2–2.5 mm high; upper lip with 2 short lobes,
0.2–0.3 2–2.5 mm; lateral and lower lobes shortly triangular, ca. 0.5
1.5 mm, apex obtuse or sometimes rounded.
Flowers slightly fragrant. Corolla white with pinkish hue.
Standard: claw 0.7–1 mm long; blade orbicular 6–7 ca. 6 mm,
apex emarginate, without auricles or basal callosities, both
sides glabrous. Wings usually curved backward; claw ca.
3 mm long; blade narrowly elliptic, 7–8 0.7–1 mm, apex
rounded, auricle ca. 1 mm long, both sides glabrous. Keel
petals: claw ca. 2 mm long; blade boat-shaped, 7–7.5 2–
2.5 mm, apex rounded, both sides glabrous, lateral pocket
up to 1.5 mm long or sometime indistinct, auricle 0.8–1 mm
long. Stamens monadelphous, with basal fenestrae, 10–
12 mm long, free part of filaments 1.5–2 mm long, glabrous;
0.2 mm, glabrous. Disc annular, 0.2–
anthers ca. 0.5
0.3 mm high. Ovary 6.5–7 mm long, hairy, stipe indistinct;
style 5–6 mm long, hairy at base; ovules 4 or 5. Pods straplike or sometime elliptic, 5–12 1.5–3 cm, thin, glabrescent,
with wings along both sutures, upper wing 4–7 mm wide,
lower wing 1–3 mm wide; seed chamber hardly visible. Seeds
1 or 2 per pod, ellipsoid, flat, ca. 8 19 1–2 mm; hilum
central, 1–4 mm long.
Distribution—Derris tonkinensis occurs in south China,
northern Vietnam, and Thailand (Fig. 12).
Habitat and Ecology—This is a species that grows on rugged limestone mountain, in mixed evergreen or deciduous
hardwood forest, light shade to open exposed areas damaged
seasonally by fire, 550–1,625 m.
Phenology—Flowering is from August to September.
Fruiting is from October to December.
Notes—Derris tonkinensis is a limestone species (Phan and
Vidal 2001: 78) known from mountains in Laos, Vietnam, and
northern Thailand. The first author also found specimens
from limestone mountains in southwestern and Peninsular
Thailand, that are similar to the northern specimens in many
characters. However, they have not been included in
D. tonkinensis yet, awaiting further molecular evidence.
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12 Jul 1926, Garrett 299 (L); Loei: Wang Sapung, 8 Mar 1924, Kerr 8629
(K); Nan: 18 Jul 1926, Winit 1758 (K); Phrae: Song, Mae Yom National
Park, west side of Mae Yom River, Mae Den area, 12 Nov 1991, Maxwell
91–1024 (L).
[Volume 37
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Representative Specimens Examined—THAILAND. Chiang Mai: Chiang
Dao, Doi Chiang Dao Animal Sanctuary, Doi Luang valley, 6 Nov 1995,
Maxwell 95–1085 (L); Lampang: Wahng Nua, Jae Sawn National Park,
northern part, Wahng Die Subdistrict; Pah Ngam (Nahn Kaht) limestone
mountain; summit above Maw Cave, 3 Oct 1996, Maxwell 96–1282 (L).
VIETNAM. Quang Ninh, 3 Jul 1885, Balansa 1189 (P).
16. Derris trifoliata Lour., Fl. Cochin. 433. 1790. Miq., Fl.
Ned. Ind. 1: 144. 1855. Craib, Fl. Siam. Enum. 1: 494. 1928.
Thoth., Bull. Bot. Surv. India 3: 181. 1961; P. K. Lôc &
J. E. Vidal, Fl. Cambodge, Laos & Vietnam 30: 74. 2001.
Deguelia trifoliata (Lour.) Taub. in Engl. & Prantl, Nat.
Pflanzenfam. 3: 345. 1894.—TYPE: CHINA. Guangdong,
Loureiro s. n. (holotype: P!).
Dalbergia heterophylla Willd., Sp. Pl. 3: 901. 1803. Derris
heterophylla (Willd.) Backer ex Heyne, Nutt. Pl. ed. 2: 806.
1927. Backer & Bakh. f., Fl. Java 1: 619. 1964. Rudd, Rev.
Hand. Fl. Ceylon 7: 232. 1991.—TYPE: INDIA, Herbarium
Willdenow 13073 (Roxburgh?, see Note) (lectotype designated by Rudd 1991: B!, photo).
Robinia uliginosa Willd., Sp. Pl. 3: 1133. 1803. Galedupa
uliginosa (Willd.) Roxb., Fl. Ind. ed.2, 3: 243. 1832. Derris
uliginosa (Willd.) Benth. in Miq., Pl. Junghuhn. 252.
1852. Miq., Fl. Ned. Ind. 1: 141, Table 3, fig. H. 1855.
Baker in Hook. f., Fl. Brit. Ind. 2: 241. 1878. Heyne,
Nutt. Pl. 2: 308. 1916. Gagnep. in Lecomte, Fl. IndoChine 2: 453. 1916. Ridl., Fl. Mal. Pensins. 1: 595,
Fig. 54. 1922.—TYPE: INDIA. Roxburgh s. n. (holotype:
B, lost, isotype: BM!, photo).
Pterocarpus frutescens Blanco, Fl. Filip.: 562. 1837. ed. 2: 392.
1845. ed. 3, 2: 356. 1879.—TYPE: PHILIPPINES. Luzon,
Bulacan Prov., Obando, Merrill Species Blancoanea 541
(neotype here designated: L!; isoneotype: US).
Derris multiflora Benth. in Miq., Pl. Jungh. 253. 1852. Benth.,
Proc. Linn. Soc. 4 (Suppl.): 108. 1860.—TYPE: INDONESIA. Java, Anjol, Mt. Blitaran, Junghuhn 47 (holotype: K!).
Brachypterum floribundum Miq., Fl. Ned. Ind. 1: 139. 1855.
Derris floribunda (Miq.) Benth., Proc. Linn. Soc. 4
(Suppl.): 105. 1860. King, J. Asiat. Soc. Beng. 66: 105.
1897. Deguelia floribunda (Miq.) Taub. in Engl. & Prantl.
Nat. Pflanzenfam. 3, 3: 345. 1894.—TYPE: INDONESIA.
Java, near Samarang, Horsfield s. n. (L20) (holotype: K!;
isotypes: CAL, U!, barcode U 0003511).
Derris forsteriana Blume ex Miq., Fl. Ned. Ind. 1: 144, t. 3, fig. G.
1855.—TYPE: INDONESIA. Celebes, 8 Sep 1813, Forsten
s. n. (lectotype here designated: L!, barcode L 0476090).
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
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Lianas, sometimes more or less creeping, up to 8 m long,
up to 5 cm diam. Twigs hollow, 4–7 mm diam., glabrous,
lenticellate. Leaves with 3–7 leaflets, rarely unifoliolate. Stipules triangular, 1.5–2 ca. 2 mm, glabrous, caducous. Petiole
1.5–10 cm long, grooved above, striate, glabrous; rachis 1.5–
6 cm long, grooved above, striate, glabrous; pulvinus 3–
10 mm long, glabrous. Stipellae sometimes present, acicular,
0.5–1(–1.5) mm long, glabrous. Leaflets subcoriaceous to coriaceous; terminal one ovate to elliptic, 3.5–16
1.5–8.5 cm,
length/width ratio 1.8–3, base slightly cordate to (sub)peltate,
rarely rounded, apex emarginate, rounded or acuminate, acumen 3–25 mm long, upper and lower surface glabrous, midrib
and nerves flat or slightly raised, veins 5–11 per side, 3–20 mm
apart, not reaching the margin but curving towards the
apex, anastomosing before the margin, venation reticulate;
lateral ones mostly like the terminal one, narrowly elliptic
to ovate, 2.5–12 1–5.5 cm, length/width ratio 1.7–3.6; pulvinus 4–7 mm long. Pseudoracemes axillary or ramiflorous
(then often on creeping branches), rarely pseudo-panicles,
1–18 cm long, peduncle 0.2–4.5 cm long, thinly strigose.
Bracts subtending brachyblasts triangular, 0.6 –1.4
0.5 –
1 mm, outside glabrous or thinly sericeous, margin ciliate,
inside glabrous. Brachyblasts knob-like, up to 5 mm long,
with 2 or 3(–7) flowers. Bracts subtending flowers elliptic to
triangular, orbicular or cordate, 0.5–0.8 0.3–0.6 mm, both
sides glabrous, margin ciliate. Pedicels 2–6 mm long, glabrous or with few hairs. Bracteoles at the apex or the upper
part of the pedicels, elliptic, triangular, orbicular or cordate,
0.5–0.7 0.4–0.6 mm, both sides glabrous, margin ciliate.
Calyx cup-shaped, 2.5–4 mm long, outside glabrous, teeth
ciliate, inside sericeous in upper part, tube 2–3 mm long;
upper lip usually shallowly notched, 0.25–0.5 2.5–4 mm; lateral lobes triangular or semi-circular, ca. 0.5 1–2 mm; lower
lobe triangular, ca. 0.5 1.5–2 mm. Corolla white to pale
pink, rarely purple or pale blue. Standard with a green
spot at the base; claw 2–3 mm long; blade orbicular, 7–
9.5 7.5–10 mm, apex slightly emarginate, basal callosities
absent, both sides glabrous. Wings: claw 2.5–4 mm long;
blade elliptic, 6–9 2–3 mm, apex rounded, both sides glabrous, lateral pocket indistinct, upper auricle 0.5–1 mm
long. Keel petals: claw 2.5–3.5 mm long; blade boat-shaped,
6–8 3–4 mm, apex rounded, both sides glabrous, lateral
pocket ca. 2 mm long, upper auricle indistinct or sometimes
up to 1 mm long. Stamens monadelphous, with basal fenestrae, 8–12 mm long, free part 1.5–4 mm long, glabrous;
anthers 0.6–0.8 0.3–0.5 mm, glabrous. Disc absent. Ovary
3–5 mm long, stipe indistinct, sericeous; ovules 3–8; style
6–9 mm long, glabrous or sometimes sericeous for up to 2/3.
Pods elliptic to more or less discoid or obovoid, 30–55 20–
37 mm, with few hairs especially basally and on the beak,
upper wing only, 1–5 mm wide; seed chamber absent. Seeds
1–3 per fruit, flat, bean-shaped, 13–23 12–21 2–5 mm;
hilum central, ca. 1 mm long.
Distribution—Derris trifoliata is the most widespread species of the genus Derris, occurring along southern and eastern
coast of Africa, Madagascar, Reunion, Seychelles, India and
Sri Lanka to China and the Ryukyus, southward to SE Asia,
Thailand and throughout Malesia up toward Australia and
the west Pacific (Fig. 16).
Habitat and Ecology—This is a species that grows in primary and secondary forests along coasts, on beaches, in mangrove, along dikes, river banks, and brackish swamps, up to
400 m.
429
Distribution of Derris trifoliata Lour.
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Fig. 16.
Phenology—Flowering and fruiting are throughout the year.
Uses—It is used as a fish poison. Also used in local medicine as a stimulant, antispasmodic, and counter-irritant, and
against rheumatism, chronic paralysis, and dysmenorrhoea.
In Papua New Guinea a decoction of the roots is used externally against fever and internally against sores. Thai traditional doctors used roots or stems as a laxative, carminative,
and anti-arthritis treatment.
Vernacular Names—Indonesia: Areuy ki tonggeret, Tuwa
areuy (Sundanese); Gadal (Javanese). Malaysia: Tuba bekut
(Peninsular). Papua New Guinea: Gamo (Zimanki), Woifatum
(Wanigela). Philippines: Silasila, Asiasimanan (Tagalog);
Butong (Bisaya). Thailand: Tob tab nam, Khweap ta-le,
Phak tab (Central), Top tab ta-le (South-western). Vietnam:
(Dây) Cóc kèn, (Dây) Ngái.
Notes—Types of Dalbergia heterophylla and Robinia
uliginosa were studied from microfilm deposited in L. The
first one belongs to Herbarium Willdenow 13073 without
name or number of the collector on the specimen’s label.
However, Roxburgh is possibly the collector of this specimen because his name (and the same handwriting) is present on other related specimens.
Representative Specimens Examined—AUSTRALIA. Northern Territory: Oenpelli, on muddy bank of tidal East Alligator River, 14 Oct
1948, Specht 1199 (L); Queensland: McIvor River, 25 Jul 1972, Stocker 895
(L). BRUNEI. Belait: Kula Belait, 7 Aug 1965, van Niel 4064 (L); BruneiMuara: Sengkurong, Jerudong, 26 Jan 1994, Said BRUN 16702 (L). FIJI. Viti
Levu: Serua, Flat coastal strip in vicinity of Ngaloa, 26 Dec 1953, Smith
9335 (L). GUAM Riverine Forest at Mouth of Talofofo River, 3 Jun 1962,
Stone 4180 (L). INDIA. Andaman and Nicobar Islands: Jula, Katchal
Island, roadside, 27 Dec 1974, Chakrabarty 2238 (L). INDONESIA. Bali:
Beroa Bay, 25 Sep 1992, Gordon and Bougher M34 (L); Celebes: Along South
shore of Lake Matano, West of Soroaka, 14 Jun 1979, Hennipman 5810 (L);
Java: Pantai Damas, Prigi, 12 Jun 1983, Afriastini 1229 (L); Kalimantan
Timur: PT. Inhutani I, Berau, Labanan area beside Labanan river, 29 May
1997, Ambriansyah Berau 555a (L); Moluccas: Halmahera, Babano igo,
2 Nov 1951, Idjan and Mochtar 378 (L); Nusa Tenggara Timur: Flores,
13 Apr 1973, Verheijen 3385 (L); Papua: Batana Island, Amdoei, distr.
Radjah Ampat, 2 Apr 1954, van Royen 3301 (L); Sumatra: Atjeh, Kampong
Sigleng, N. from Troemon, 29 Aug 1941, Asdat 152a (L); Riau, Lingga
Arch, Pulau Lingga, Kampong Daik, 14 Jul 1919, Beumée 1925 (L). JAPAN.
Ryukyu Islands: Yonaguni Island, east of Sonai, 27 Aug 1951, Walker and
Tawada 6796 (L). MADAGASCAR. Toamasina: N. end of Bay of Antongil,
c 2 km east of Maroansetra, 24 Apr 1989, D. J. and B. P. Du Puy M 211 (K).
MALAYSIA. Johore: Sedili Kecil, 25 Sep 1971, Res 00420 (L); Perak: Kuala
Sepetang near Port Weld mangrove zone, 9 Jun 1983, Stone 15558 (L);
Negeri Sembilan: Port Dickson, 22 Feb 1946, Monod de Froideville 800 (L);
Selangor: 13 Sep 1967, Carrick 1634 (L). Sabah: Kunak, Road to oil palm,
Kunak town, 10 Jul 1979, Madani SAN 90782 (L); Sarawak: Gunong Api,
ulu Sg. Melinau, Tutoh, Baram District. NE flank of mountain, 27 Sep
1971, Anderson S 30762 (L); Trengganu: Kemaman, 13 Jul 1931, Osman
430
SYSTEMATIC BOTANY
and Tachon KEP 27013 (L). MICRONESIA. Chuuk: Moen Island, south
of Moen Village, 15 Nov 1949, Anderson 738 (L); Kosrae: Lela (Lele)
Island, Lela (Lele) Harbor, 19–21 Aug 1946, Fosberg 26544 (L); Pohnpei:
Nanmatol Islet, Matalanim District, 11 Aug 1946, Fosburg 26384 (L). NEW
CALEDONIA. Ponérihouen: Monéo, 1 Jan 1961, Mckee 7879 (L). PALAU
ISLANDS. Babeldaob (Babelthuap) Island: Garamiscan Colony, Garamiscan
(Almiokan) River, 20 Jul 1946, Fosberg 25739 (L). PAPUA NEW GUINEA.
Madang: Laing Island, Hansa Bay near Bogia, 28 Jun 1980, Goetghebeur 3509
(L); Manus: Lou Island, area of hot spring and mud field, close to the south
coast, 4 Dec 1975, Sands et al. 3027 (L); Milne Bay: Goodenough island, Bolu
Bolu, 29 Sep 1953, Brass 24434 (L); West New Britain: Fullerborn Harbour,
Gasmata, 7 May 1973, Womersley LAE 55338 (L); West Sepik: Creek between
Wara Col and Musu river along Vanimo-Wutung road, Vanimo, 29 Sep
1977, Wiakabu LAE 73335 (L). PHILIPPINES. Luzon: La Union, Sta. Rita,
19 Oct. 1992, Barbon et al. PPI 8968 (L); Mindanao: Mati, Davao, Mar-Apr
1927, Ramos and Edano PNH 49040 (L); Mindanao: Agusan Norte, Tungao,
NALCO, 19 May 1991, Barton et al. PPI 1892 (L); Mindoro: Mt. Yagaw,
25 Mar 1958, Conklin PNH 37861 (L); Palawan: Sidanow, 25 Apr 1964,
Mendoza and Espiritu PNH 91257 (L). SINGAPORE. Changi Point, 18 Sep
1979, Maxwell 79–16 (L). SOLOMON ISLANDS. Santa Isabel: Binusa, N.W.
Santa Isabel, Beach side, 18 Jan 1966, Beer’s collectors 6793 (L); Ulawa Island:
Olosu’u, beach side, 4 Feb 1965, Teona 6225 (L). SOUTH AFRICA. Natal:
Mtunzini, 4 Mar 1967, Strey 7396 (K). SRI LANKA. Eastern Province:
Trincomalee District, ca. 5 miles due S of Tamaivillu, 14 Oct 1974, Davidse
7583 (L); Western Province: Thalahena, Colombo District, 10 Aug 1974, Waas
723 (L). THAILAND. Kanchanaburi: 3 Jan 1929, Kerr 10130 (L); Narathiwat:
Ban Ka Pong Baa Ruu, Ra Ngea, 21 Aug 1988, Niyomdham and Ueachirakan
1895 (L); Pangnga: Kraburi, 8 May 1973, Geesink and Santisuk 5249 (L);
Ranong: Ban Tha Mai, 6 km from the coast, 10 Jul 1968, van der Kevie 37 (L);
Samut Prakan: Sawangkaniwas (Thai red cross society), Tai Ban, Muang,
20 Aug 2009, Sirichamorn YSM2009–06 (L); Samut Songkhram: Bahn Koon
Chong, Muang District, 23 Aug 1975, Maxwell 75–916 (L); Songkhla: Yaw
Island (Ko Yao), south side, 27 Aug 1986, Maxwell 86–621 (L). EAST TIMOR.
Weluli, 21 Jul 1966, Friedberg 398 (L).
Paraderris (Miq.) Geesink, Leiden Bot. Ser. 8: 109. 1984.
Derris Lour. sect. Paraderris Miq., Fl. Ind. Bat. 1: 145.
1855. Benth., Proc. Linn. Soc. Lond. 4, Suppl.: 111. 1860.
Geesink, Adv. Leg. Syst. 1: 256. 1981. Deguelia Aubl. Sect.
[Volume 37
Paradeguelia Taub. in Engl. & Prantl., Nat. Pflanzenfam.
3: 345. 1894, nom illeg.—TYPE: Derris cuneifolia Benth.
(lectotype designated by Geesink 1984). [= Paraderris
cuneifolia (Benth.) Geesink].
Lianas. Leaves imparipinnate; stipules present, sometimes
caducous; stipellae absent or occasionally present but then much
reduced. Leaflets 3–15 per leaf, opposite, entire, often obovate.
Pseudoracemes axillary, sometimes terminal, rarely pseudopanicles; flowers usually 2 or 3 on the top of long and slender
brachyblasts. Bracts subtending brachyblasts and flowers persistent or caducous, shorter than the corresponding flower
buds; bracteoles usually in the upper part of the pedicel or at
the base of the calyx. Calyx cup-shaped, bilabiate, upper lip
2-lobed, often indistinct; lower lip 3-lobed, inside usually sericeous in upper part. Corolla whitish or pinkish; standard
with 2 large basal callosities, in 1 or 2 species without any
basal callosities, reflexed at base; wings approximately as
long as keel petals, wings and keel petals adherent by lateral
pockets or hooked together via auricles or twisted claws (or
any combination). Stamens 10, monadelphous, with basal
fenestrae; anthers all fertile, usually hairy. Disc inconspicuous,
annular. Ovary with 1–7 ovules. Pods indehiscent to rarely
tardily dehiscent, leathery, rarely woody, with a wing along
both sutures or along the upper suture only, rarely unwinged,
seed chamber present or absent. Seeds 1–3, bean-shaped, often
flat; hilum usually central.
Distribution—The genus is from SE Asia, from India,
Myanmar, throughout Malesia up to New Guinea.
Habitat and Ecology—Like Derris, Paraderris prefers light
and water. The genus is usually found in fully exposed to
semi-shaded areas along streams or rivers.
Key to the Thai Species of Paraderris
Leaflets (thinly) sericeous below. Calyx sericeous outside. Standard sericeous except for the rim and claw . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. elliptica
Leaflets mostly glabrous below. Calyx glabrous or thinly sericeous outside. Standard glabrous, sometimes thinly ciliate at apex . . . . . . . . . . . . . . . 2
2. Blade of standard 8–8.5 mm long, basal callosities small, indistinct or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. laotica
2. Blade of standard more than 10 mm long, basal callosities large and distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaflets 5–9 3.3–5 cm, apex rounded to shortly acuminate, acumen 3–4 mm long.—N. India, Nepal, Myanmar, S. China,
and N. Thailand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. cuneifolia
3. Leaflets 6.5–18.5 3.3–9 cm, apex acuminate, acumen 5–20 mm long.—Thailand, throughout Malesia . . . . . . . . . . . . . . . . . . . . . 4. P. montana
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1.
Galedupa marginata Roxb. [Hort. Bengal. 53. 1814 (nom.
nud.)], Fl. Ind. 3: 241. 1832.—TYPE: INDIA. Silhet,
Wallich Cat. 5896A (holotype: K-W; isotype: P!).
Derris discolor Benth., Proca. Linn. Soc. 4 (Suppl.): 111. 1860.—
TYPE: INDIA. Sikkim: Terai, lower hill, J. D. Hooker s. n.
(holotype: K!; isotypes: K!, P!).
Derris hancei Hemsl., Bot. Mag. t. 8008 (after D. alborubra
Hemsl.): 131. 1905.—TYPE: CHINA. Canton: along the
river, Sampson in herb. Hance 9920 (holotype: K!).
Derris glauca Merr. & Chun, Sunyattsenia 2: 246. 1953.—
TYPE: CHINA. Hainan: Ngai Yuen, 4 Jun 1933, F. C.
How 70860 (holotype: NY!, photo; isotypes: US!, A).
Derris truncata Craib, Kew Bull. 385. 1927.—TYPE: THAILAND.
Pitsanulok: Nakhawn Tai, 3 Apr 1924, Kerr 8898 (holotype:
K!; isotype: BK!).
Derris cuneifolia Benth. var. longipedicellata Thoth., Bull. Bot.
Surv. India 3: 191. 1961. Fasc. Fl. Ind 8: 15. 1982.—TYPE:
INDIA. Sikkim: Sivoke, Ribu 760 (holotype: CAL, see
Thothathri 1961).
Derris cuneifolia Benth. forma assamica Thoth., Bull. Bot. Surv.
India 3: 192. 1961. Fasc. Fl. Ind 8: 15. 1982.—TYPE:
INDIA. Assam: Cachar, Bishnupur, U. Kanjilal 4835
(holotype: CAL, see Thothathri 1961).
Lianas. Twigs 3–6 mm diam., smooth or slightly
lenticellate, thinly hirsute in young parts, older parts
glabrescent. Leaves with 5–9 leaflets, rarely up to 15. Stipules
ovate, 1.5–2 2–2.5 mm, outside thinly sericeous, margin
ciliate, inside glabrous. Petiole 5–7.5 cm long, striate; rachis
4–8 cm long, striate, glabrous to thinly hirsute at the upper
side; pulvinus 4–8 mm long, hirsute, especially at the base.
Stipellae absent. Leaflets chartaceous, terminal one obovate,
5–9 3.4–5 cm, length/width ratio 1.5–1.8, base cuneate, apex
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Paraderris cuneifolia (Benth.) Geesink, Scala
Millettiearum: 109. 1984. Derris cuneifolia Benth. in Miq.,
Pl. Jungh. 253. 1852. Proc. Linn. Soc. 4 (Suppl.): 112. 1860.
Prain, J. Asiat. Soc. Beng. 66, 2: 459. 1897. Thoth., Fasc. Fl.
India 8: 13. 1982. Deguelia cuneifolia (Benth.) Taub. in
Engl. & Prantl, Nat. Pflanzenfam. 3: 345. 1894.—TYPE:
NEPAL. Nookate, Wallich Cat. 5887 (lectotype designated
by Thothathri 1982: CAL; isolectotypes: K!; BM, LE).
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SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
Thomson s. n. (P). THAILAND. Pitsanulok: Nakhawn Tai, 3 Apr 1924,
Kerr 8898 (BK).
2. Paraderris elliptica (Wall.) Adema, Thai. For. Bull., Bot.
28: 11. 2001. Pongamia elliptica Wall., Pl. Asiat. Rar. 3: 20,
t. 237. 1832. Derris elliptica (Wall.) Benth., Proc. Linn.
Soc. 4 (Suppl.): 111. 1860. Prain, J. Asiat. Soc. Beng. 66, 2:
460. 1897. Gagnep. in Lecomte, Fl. Indo-Chine 2: 461.
1916. Thoth., Bull. Bot. Surv. India 3: 194. 1961. Ridl., Fl.
Malay Penins. 1: 597. 1922. Craib, Fl. Siam. Enum. 1: 488.
1928. Backer & Bakh. f., Fl. Java 1: 617. 1964. P. K. Lôc &
J. E. Vidal, Fl. Cambodge, Laos & Vietnam 30: 60. 2001.—
TYPE: INDIA. Hort. Bot. Calcutta (originally from
Ambon), Wallich 5881A (holotype: K!; isotype: CAL,
fragments).
Dalbergia purpurea Reinw. ex Hassk., Flora 25, Beibl. 2: 53.
1842.—TYPE: INDONESIA. Java, Blume s. n. (holotype:
L!, barcode L 0475217).
Pongamia hypoleuca Miq., Fl. Ned. Ind. 1: 148. 1855.—TYPE:
BORNEO. Korthals s. n. (lectotype drsignated by Adema
2003a: L!, barcode L 0475263).
Pongamia volubilis Zoll. & Mor. var glaucophylla Miq., Fl. Ned.
Ind. 1: 149. 1855. Derris elliptica (Wall.) Benth var
glaucophylla (Miq.) Kaneh. & Hatus., Bot. Mag. Tokyo
56: 364. 1942.—TYPE: INDONESIA. Java, Anonymous
s. n. (lectotype designated by Adema 2003a: L!, barcode
L 0475218).
Derris elliptica (Wall.) Benth. var. chittagongensis Thoth., Bull.
Bot. Surv. India 3: 195. 1961. Fasc. Fl. India 8: 18. 1982.
Paraderris elliptica (Wall.) Adema var. chittagongensis
(Thoth.) Adema, Blumea 48: 137. 2003.—TYPE: INDIA.
Chittagong: Kodla, Baldal Khan 450 (C) (holotype: CAL!;
isotype: CAL, photo K!).
Lianas, up to 5 m high. Bark dull greenish to brown. Twigs
4–15 mm diam., hirsute, glabrescent in older parts,
lenticellate. Leaves with 7–15 leaflets. Stipules triangular or
ovate, 3–3.5 1.7–3 mm, outside sericeous, inside glabrous.
Petiole 4–14.5 cm long, striate, (thinly) hirsute; rachis 6–20.5 cm
long, striate, (thinly) hirsute; pulvinus 5–11 mm long, (thinly)
hirsute. Stipellae rarely present, acicular, ca. 1.5 mm long.
Leaflets chartaceous to subcoriaceous, terminal one obovate to
narrowly obovate, 7–16 2–5 cm, length/width ratio 3.2–4.3,
base cuneate, apex acuminate, acumen 3–20 mm long,
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Representative Specimens Examined— CHINA. Guangzhou: along
the river, 1870, Sampson 9920 (P); Hainan: Ku Tung Village, Ching
Mai District, 2 May 1933, Lei 612 (L). INDIA. Silhet: s. d., Hooker and
Fig. 17. Distribution of Paraderris cuneifolia (Benth.) Geesink (circle)
and P. laotica (Gagnep.) Adema (diamond).
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shortly acuminate or obtuse, acumen 3–4 mm long, rounded,
upper surface glabrous, lower surface with widely scattered
appressed hairs, especially along the midrib and veins, midrib flat to slightly sunken above, raised below, nerves flat or
slightly raised above, raised below, 8–11 per side, 6–10 mm
apart, curving towards the apex and almost reaching the leaf
margin, anastomosing near or close to the margin, venation
reticulate; lateral leaflets mostly like terminal one, elliptic or
obovate, 6–7.5 3.3–4 cm, length/width ratio 1.8–1.9; pulvinus ca. 4.5 mm long, thinly sericeous. Pseudoracemes axillary or rarely terminal, rarely combined to pseudopanicles,
7–16 cm long; peduncle 1–4.5 cm long, glabrous to thinly
strigose; lateral branches up to 7 cm long. Bracts subtending
brachyblasts ovate to triangular, 1.7–1.8 1.5–1.6 mm, outside sericeous, margin ciliate, inside glabrous. Brachyblasts
slender, cylindrical, 5–9 mm long, with 2 or 3 flowers at the
apex. Bracts subtending flowers broadly elliptic to ovate, 1–
1.1 0.7–0.8 mm, outside glabrous or thinly sericeous, margin ciliate, inside glabrous. Pedicels 3–9.5 mm long, glabrous
or thinly sericeous. Bracteoles at the top of the pedicels, elliptic to ovate or orbicular, ca. 1 0.7–0.8 mm, outside glabrous
or thinly sericeous, margin ciliate, inside glabrous or thinly
sericeous at the apex. Calyx 3–4.5 mm high, reddish, outside
glabrous to thinly sericeous, margin ciliate, inside glabrous,
tube 2.5–4 mm high; upper lip indistinct or sometimes with
2 short semicircular lobes; lateral lobes (broadly) triangular,
0.3–0.5 2–2.5 mm; lower lobe triangular, 0.4–0.5 2–3 mm.
Corolla white or pink. Standard: claw 2–2.5 mm long; blade
broadly ovate to orbicular, 11.5–12 10–13 mm, apex rounded,
glabrous, basal callosities present. Wings: claw 2.5–4 mm long;
blade elliptic, 10–11 3.8–4 mm, apex rounded, both sides
glabrous or thinly ciliate at apex, upper auricle 0.5–1 mm long,
lower auricle 0.2–0.4 mm long, lateral pocket 2.5–3.5 mm long,
Keel petals: claw 4–4.5 mm long; blade boat-shaped, 9.6–10
3.5–3.7 mm, apex rounded, both sides glabrous or with some
hairs at the apex, auricle up to 0.7 mm long or sometimes
indistinct, lateral pocket 3–4 mm long. Stamens monadelphous
with basal fenestrae, 9–11 mm long, free part 3–4 mm long,
glabrous; anthers 0.6–0.9 0.3–0.5 mm, glabrous. Disc annular
or with (up to) 10 lobes, 0.2–0.5 mm high, with some hairs.
Ovary 3.5–7 mm long, sericeous; stipe usually indistinct but
sometimes up to 1 mm long; ovules 3 or 4; style 6–7 mm long,
sericeous in lower part. Pods elliptic or ovate, 40–50 20–
23 mm, glabrous, with wing along both sutures, upper wing
3–4.5 mm wide, lower wing 3–4.5 mm wide, sometimes indistinct; seed chamber absent. Seeds unknown.
Distribution—This species occurs in India, Nepal,
Bangladesh, Myanmar, south China, Laos, Vietnam, Thailand,
Peninsular Malaysia, Celebes (Fig. 17).
Habitat and Ecology—This is a species from primary and
secondary dry evergreen forests, often along rivers.
Phenology—Flowering is from March to May. Fruiting is
from June to July.
Notes—Pods of P. cuneifolia are flatter and more distinctly
winged than those of P. montana. Without pods, the species
are similar. However, we still consider them as separate
species, because they also have a different distribution.
P. cuneifolia has a more northern distribution, while P. montana
has a southern distribution. Type specimens deposited in
CAL could not be examined.
431
SYSTEMATIC BOTANY
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rounded, upper surface glabrous to thinly sericeous, lower
surface more or less sericeous, older leaflets often less hairy,
midrib flat above, rarely slightly sunken, raised below, nerves
flat above, raised below, 7–14 per side, 3–16 mm apart, curving
towards the apex and almost reaching the margin, sometimes
anastomosing near the margin, venation reticulate; lateral
leaflets mostly like terminal one, narrowly elliptic to obovate,
4–16 2–5.5 cm, length/width ratio 2–4; pulvinus 3–7 mm
long, (thinly) sericeous. Pseudoracemes/ pseudopanicles axillary, rarely terminal, up to 30 cm long; peduncle 0.5–5.5 cm
long, sericeous; lateral branches 5–20 cm long. Bracts subtending brachyblasts ovate to triangular, 1.5–2.5 1–1.5 mm,
outside sericeous, inside glabrous. Brachyblasts long cylindrical, 10–25 mm long, with 2 or 3 flowers at the apex. Bracts
subtending flowers ovate, elliptic, triangular or cordate, 1.5–
2.5 1–2 mm, outside sericeous, inside glabrous. Pedicels 5–
11 mm long, sericeous. Bracteoles at the top of the pedicels,
(narrowly) ovate or triangular, 1–2.7 1–2 mm, outside sericeous, inside glabrous. Calyx 5.5–7 mm high, brown to dark
maroonish, outside sericeous, inside sericeous at apex, tube
4–6 mm high; upper lip hardly visible or sometimes with
2 short semicircular lobes; lateral lobes (broadly) triangular, 1–
1.5 3–5 mm; lower lobe triangular, 1.5–2.5 3–5 mm. Corolla
white to pink, or violet, standard with green blotch at base.
Standard: claw 3–4 mm long; blade orbicular, 12–17 10–
17 mm, apex emarginate, basal callosities large, outside sericeous, except for the margin and claw, inside sericeous at
apex. Wings: claw 4.5–5 mm long; blade obliquely elliptic,
10 –13 4.5–5 mm, apex rounded, upper auricle 0.5–1 mm long,
lower auricle absent, lateral pocket 2–5 mm long, sometimes
hardly visible, outside with some hairs at the ventral side
part of the apex, inside sericeous at apex. Keel petals: claw
4.8–5 mm long; blade boat-shaped, 10–11 5–6 mm, apex
rounded, auricle indistinct, lateral pocket large, 4–5 mm long,
outside with some hairs at apex, inside sericeous at apex,
rarely with some hairs at the apex. Stamens monadelphous, with basal fenestrae, 14–23 mm long, free part 3–7 mm
long, glabrous, rarely with some hairs; anthers 0.3–0.6 0.2–
0.4 mm, with some hairs outside. Disc indistinct, annular, usually with some hairs. Ovary 6–10 mm long, outside sericeous,
inside with a row of hairs along the lower suture; stipe 1–
1.7 mm long, sericeous; ovules 3–5; style 8–14 mm long,
sericeous in lower part. Pods elliptic or strap-like, flat, 4–
11 1.5–2 cm, thinly sericeous, sometimes also glaucous,
with wing along the upper suture only or rarely winged on
both sutures, upper wing 1.5–2 mm wide, lower wing usually undeveloped, up to 1.5 mm wide; seed chamber absent.
Seeds seen immature in well developed pods, bean-shaped,
flat, ca. 13 8–8.5 ca. 3 mm; hilum central, 2.5–3 mm long.
Figure 18.
Distribution—This species occurs in India, Myanmar,
Laos, Vietnam, Cambodia, throughout Thailand, Malesia:
Malay Peninsula, Philippines (Palawan); Caroline Islands,
and probably introduced in Hawaii (Fig. 19).
Habitat and Ecology—This is a species from primary and
secondary forest, often along riverbanks, up to 1,450 m.
Phenology—Flowering is from December to April. Fruiting
is from March to May, October, or December.
Uses—The roots commonly known as “derris or tuba root
of commerce,” are the source of rotenone whose content
varies from 1.6–10.0% (Thothathri 1982: 17). The plants are
cultivated for fish poison and insecticide.
[Volume 37
Vernacular Names—Brunei: Tuba (Dusun, Malay).
Cambodia: Ca bia. Indonesia: Tuba, Oyod tungkul (Javanese),
Tuwa leteng (Sundanese). Malaysia: Tuba, Akar tuba. Philippines:
tubli, Tugling-pula (Tagalog), Upei (Bontok). Myanmar: Hon.
Thailand: Hang lai daeng, Lai nam, Lo tin (Northern, Central),
Kalamphoh (Phetchaburi), Oud nam (Surat Thani). Veitnam:
Khê’ dai (Northern), (Dây) Cô˜ rùa (Southern).
˙
Notes—This
species is variable in the size of the leaflets,
color and density of the indumentum, shape of the pods,
and the presence of a wing along the lower suture of the
pod. The leaflets from most specimens are obovate, but vary
in size. Hairs are usually colorless, whitish or silvery, but
some specimens, including those formerly recognized as
var. chittagongensis, have yellowish or golden-brown hairs.
The density of hairs also varies between thinly and densely
sericeous. The size of pods is ca. 7 2 cm, but some specimens
from Myanmar, northwestern and western Thailand have
narrowly strap-like pods, which are more than 7 cm long
and less than 1.7 cm wide (Fig. 18h2). A narrow lower
wing along the lower suture of the pod is also occasionally observed. This lower wing is also reported for var.
chittagongensis (Adema 2003a). The variability between the
specimens is without gaps and, therefore, taxonomic recognition of the different forms is not warranted. At the moment,
there is no reason to distinguish entities at the infraspecific
level and, therefore, all names are still treated as synonyms.
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Representative Specimens Examined—BRUNEI. Belait: Sungai Liang,
2 Mar 1965, van Niel 3721 (L). INDONESIA. Jawa Barat: Isthmus of
Udjung Kulon, 31 Dec 1961, Kostermans 19337 (L); Jambi, 28 Aug 1945,
Posthumus 818 (L); Jawa Tengah: Semarang, Kedoengdjati, 18 Sep 1896,
Koorders 40930 (L); Kalimantan Timur: W. Koetai no.1, near Kp.
Djembajan, 16 Jun 1925, Endert 1452 (L); KPC area beside river Bengalon
to Tepian Lang, 1 Apr 1996, Ambri et al. AA1620 (L); Maluku: Tobelo, Pos
12, Eil. Raoh., 23 Mar 1938, bb 24549 (L); Nusa Tenggara Timur: Flores,
1 Nov 1980, Schmutz 4706 (L); Papua (Irian Jaya): Bird’s head Penin.,
Surrounding of Ayawasi, 31 Oct 1995, Ridsdale 2137 (L); Sulawesi Tengah:
Subdiv. Poso. Bemba, secondary vegetation, 8 Aug 1937, Eyma 1587 (L);
Sumatra Utara: Asahan River, near Tandjung Balai, 10 Apr 1954, Alston
15365a (L). MALAYSIA. Pahang: Jeram Panjang: 22 Jul 1970, Shah and
Noor MS 2044 (L). MICRONESIA. Chuuk: Lukunor Atoll, Oneop Islet,
7 Jan 1950, Anderson 2120 (L); Satawan Atoll: Satawan Islet, 2 Jan 1950,
Anderson 1078 (L); Kosrae: Lela (Lele) Island, Lela (Lele) Harbor, around
village and ruins of ancient temple, 19–21 Aug 1946, Fosberg 26547 (L).
MYANMAR. Mandalay: Inkadi, 10 May 1928, Khant 2480 (K). PAPUA
NEW GUINEA. East Sepik: Angoram, upper Karawar river, 2 Sep 1968,
Dornstreich NGF MDD-9 (L); Morobe: Markham Valley, Markham road
near Lae, Oct 1959, Henty NGF 11574 (L); Nomad: Nomad river Patral
Post, 21 Jun 1974, Luoma 3 (L). PHILIPPINES. Negros Island: Negros
Occidental, Patag Protected forest, 27 Mar 1992, Stone et al. PPI 6414 (L).
SINGAPORE. 13 3/4 mile Hurong Road, 9 Apr 1951, Sinclair s. n. (L).
THAILAND, Chiang Rai: Mae Chan, Nawng Pah Gaw subdistrict, Ban
Mai Pattana, 13 Apr 1988, Maxwell 88–467 (L); Kanchanaburi: Pompee
village near Khwae Noi river, E. of Sangkhla, 25 Mar 1968, van Beusekom
and Phengkhlai 102 (P); Nakhon Si Thammarat: Khao Lawng, 26 Mar 1958,
Snan 1016 (L); Nong Khai: Ho Kam community forest, Pak Khat-Bueng
Kan, 4 May 2002, Pooma et al. 3459 (L); Surat Thani: Ban La, 19 Feb 1930,
Kerr 18127 (L); Surat Thani: Phun Phin, Pum Duang, along the canal,
22 Sep 2009, Sirichamorn YSM2009–19 (L); Tak: Tha Songyang Mae Song,
Ban Maesong, km 107–108, near Moei river, 23 Jun 2005, Pooma et al. 5445
(L). USA, Hawaii: 5 miles west of Hana on main Road, 13 Apr 1974,
Fosburg 55438 (L).
3. Paraderris laotica (Gagnep.) Adema, Blumea 48: 138.
2003. Derris laotica Gagnep., Not. Syst. 2: 348. 1911. Craib,
Fl. Siam. Enum. 1: 490. 1928. P. K. Lôc & J. E. Vidal,
Fl. Cambodge, Laos & Vietnam 30: 60. 2001.—TYPE:
CAMBODIA. Bhâklon, 20 Oct 1909, Magnen et al. s. n.
(lectotype designated by Phan in Phan and Vidal 2001:
P!; isolectotype: K, fragments).
2012]
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
433
Fig. 18. Paraderris elliptica (Wall.) Adema. a. Habit. b. Flower. c. Standard, inside view. d. Wing petal. e. Keel petal. f. Stamens. g. Pistil. h. Pods
showing the variation of shape and pod wing: h1. Broadly elliptic pod with one wing (commonly found), h2. Narrowly strap-like pod with two indistinct
wings (occasional) [a-g: van Beusekom and Phengkhlai 102; h1: Pooma et al. 3459, h2: Pooma et al. 5445; all L]. Drawing by Anita Walsmit Sachs (L).
SYSTEMATIC BOTANY
[Volume 37
apex emarginate, basal callosities indistinct or absent (in
var. virens), both sides glabrous. Wings: claw 2.8–3 mm
long; blade elliptic, 6.5–7 1.8–2 mm, apex rounded, upper
auricle 0.5–0.6 mm long, both sides glabrous, lower auricle
indistinct, lateral pocket indistinct or absent. Keel petals:
claw 2.9–3 mm long; blade boat-shaped, 7–9 3.5–3.7 mm,
apex rounded, both sides glabrous, upper auricle ca. 0.5 mm
long, sometimes indistinct, lateral pocket 1.8–3 mm long.
Stamens monadelphous, with basal fenestrae, 11–13 mm
long, free part 2.5–4.5 mm long, glabrous or with some hairs
at apex; anthers 0.5–0.6 0.3–0.4 mm, glabrous. Disc annular
or indistinct, 0.5–1 mm high, glabrous. Ovary 5.8–7 mm long,
outside sericeous, inside glabrous; stipe up to 0.5 mm long,
sometimes indistinct; ovules 4 or 5; style 6–8 mm long, sericeous at lower part. Pods and seeds unknown.
Distribution—According to Phan and Vidal (2001), this
species is found in Vietnam, Laos, Thailand, and Cambodia
(Fig. 17).
Habitat and Ecology—This is a species from dense evergreen forest on humid ground, 600–800 m.
Phenology—Flowering is from July to August. Fruiting is
in October.
Notes—This species is different from all other Paraderris
species. Leaflets and flowers are smaller and the standard
petals lack basal callosities. However, it still has slender
brachyblasts with the flowers more or less apically (The
specimen, Maxwell 07–470, has more scattered flowers on
brachyblast). A species of Paraderris, P. lushaiensis Thoth.
was also reported to lack basal callosities on the standard
petals (Thothathri 1982). However, like other species of
Paraderris, the size of the flowers is relatively larger than
those of P. laotica. The specimens studied are only the types
and one specimen from Cambodia (Maxwell 07–470). None
have fruits. The first author’s preliminary results of a molecular analysis of three chloroplasts markers, trnL-F IGS psbAtrnH, and trnK-matK, including one nuclear marker, ITS,
indicate that this species is not closely related to other species
of Paraderris. In contrast, it is closer to species of Derris,
especially the type species, D. trifoliata. However, in this
paper this species is still recognized as a member of Paraderris
due to its distinct characteristic brachyblast and flower
position. Further analyses of more molecular nuclear and
chloroplast markers will be carried out in the near future to
verify its taxonomic status.
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Fig. 19.
Distribution of Paraderris elliptica (Wall.) Adema.
Derris laotica Gagnep var. virens Gagnep, Not. Syst. 2: 348.
1911. Craib, Fl. Siam. Enum. 1: 490. 1928.—TYPE:
CAMBODIA. Siem-reap, 15 Jul 1875, Godefroi 686 (lectotype designated by Phan in Phan and Vidal 2001: P!;
isolectotype: P!).
Representative Specimens Examined—CAMBODIA. s. d., Magnen et al.
s. n. (P); Kratie: Mekong River, Svan Island, north-western side, 1 Aug 2007,
Maxwell 07–470 (L); Siem Reap: 15 Jul 1875, Harmand 686 (L).
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Lianas, 6–10 m high. Twigs 3–4 mm diam., striate,
lenticellate, glabrous. Leaves usually trifoliate. Stipules
ovate-triangular, 1.3–1.5 0.9–1.3 mm, outside glabrous to
thinly sericeous, especially at the base, margin ciliate, inside
glabrous. Petiole 1.5–5 cm long, striate, glabrous or with few
hairs; rachis 0.6–1.3 cm long, striate, glabrous or with few
hairs; pulvinus 2–5 mm long, glabrous to thinly hirsute,
especially at the base. Stipellae absent or rarely present,
acicular, 0.5–1 0.2 mm, glabrous. Leaflets subcoriaceous;
terminal one elliptic to obovate, 5.5–6 2–3.5 cm, length/
width ratio 1.7–2.7, base rounded to subcordate (cuneate in
former var. virens), apex slightly acuminate, acumen 2–5 mm
long, rounded, upper surface glabrous, lower surface glabrous or with widely scattered appressed hairs, midrib flat
to slightly raised in a furrow above, nerves slightly raised
above, 6–8 per side, 3–6.5 mm apart, not reaching the margin
but curving towards the apex, anastomosing near the margin, venation reticulate; lateral leaflets mostly like terminal
one, (narrowly) elliptic to ovate, 3.3–7.5
0.9–3.2 cm,
length/width ratio 2.3–3.6; pulvinus 1–2 mm long, glabrous.
Pseudoraceme/pseudopanicle axillary, rarely terminal, 6.5–
20 cm long; peduncle 0–1 cm long, thinly strigose; lateral
branches up to 18.5 cm long. Bracts subtending brachyblasts
narrowly elliptic to ovate, 0.7–1 0.3–0.4 mm, outside glabrous, margin ciliate to fimbriate, inside glabrous except for
some hairs at the base. Brachyblasts slender, cylindrical, 5–
10 mm long, with 2 (–3) flowers at the apex. Bracts subtending
flowers ovate to triangular, 0.6–0.7
0.4–0.5 mm, outside
glabrous, margin ciliate to fimbriate, inside sericeous. Pedicels
3–5 mm long, glabrous or thinly sericeous. Bracteoles at the
upper part of the pedicels, (narrowly) ovate, 0.5–0.6 0.2–
0.4 mm, outside glabrous or thinly sericeous, margin ciliate to
fimbriate, inside glabrous or thinly sericeous. Calyx cupshaped, 2.5–2.7 mm high, outside glabrous, margin ciliate,
inside sericeous at apex, tube 2–2.2 mm high; upper lip
indistinct or sometimes with 2 shallow notches; lateral lobes
triangular, ca. 0.5 1.9–2.1 mm; lower lobe triangular, 0.5–
0.7
1–1.5 mm. Corolla pink. Standard: claw 2.5–2.8 mm
long; blade broadly ovate to orbicular, 8–8.5
7–7.8 mm,
4. Paraderris montana (Benth.) Adema, Blumea 48: 139. 2003.
Derris montana Benth. in Miq., Pl. Jungh. 253. 1852; Proc.
Linn. Soc. 4 (Suppl.): 113. 1860.—TYPE: INDONESIA. Java,
Dieng, Junghunh s. n. (holotype: BO, see Adema 2003a).
Derris cuneifolia Benth. var. malaccensis Benth., Proc. Linn.
Soc. 4, (Suppl.): 112. 1860. Derris malaccensis (Benth.)
Prain, J. Asiat. Soc. Beng. 66, 2: 107. 1897. Paraderris
malaccensis (Benth.) Adema, Thai For. Bull., Bot. 28: 11.
2001.—TYPE: MALAYSIA. Malacca, Griffith KD 1774
(holotype: K!; isotype: CAL).
Derris malaccensis (Benth.) Prain var. aptera Prain, J. Asiat. Soc.
Beng. 66, 2: 108. 1897.—TYPE: MALAYSIA. Perak, King’s
coll. 4518 (lectotype designated by Adema 2003a: K!;
isolectotypes: A, CAL).
SIRICHAMORN et al.: AGANOPE, DERRIS, AND PARADERRIS IN THAILAND
Derris malaccensis (Benth.) Prain var. millettiodes Prain, J.
Asiat. Soc. Beng. 66, 2: 108. 1897.—TYPE: MALAYSIA.
Perak, King’s coll. 10696 (holotype: K!; isotype: CAL).
monadelphous, with basal fenestrae, 11–13 mm long, free
part 3–5 mm long, glabrous; anthers 0.9–1
0.3–0.5 mm,
glabrous, rarely with some hairs. Disc with up to 10-lobes,
0.5–1 mm high, glabrous. Ovary 9–10.5 mm long, outside
sericeous, inside glabrous; stipe indistinct; ovules 1–4; style
6–8 mm long, sericeous in lower part. Pods woody, variable
in shape and thickness, ellipsoid, ovoid or strap-like, 2–9.5
2–4 0.5–1.4 cm, glabrous, wings absent or obscurely present
on both sutures, upper wing 1–2 mm wide, lower wing up to
1 mm wide; seed chamber absent. Seeds bean-shaped, 16–
19 12 4–5 mm; hilum central, 2.5–3 mm long.
Distribution—This species occurs in Bangladesh, Thailand,
Malesia: Peninsular Malaysia, Sumatra, Borneo, Java, Celebes,
and New Guinea (Fig. 20).
Habitat and Ecology—This is a species from primary and
secondary forests, often along rivers, forest margins, roads and
railroads, or in para rubber (Hevea) plantations, up to 1,000 m.
Phenology—Flowering has been reported from February
to May, or August to December. Fruiting is from March or
May to September, or November.
Uses—It is used as insecticide and fish poison.
Vernacular Names—Malaysia: Tuba merah (Peninsular).
Thailand: Hang lai khao.
Notes—The characters of the pod are different from Derrislike plants. They are often inflated and unwinged. However,
in some specimens the pods are slightly winged along the
upper suture or both sutures. The pods appear to be tardily
dehiscent. The type of Derris montana, Junghunh s.n, is possibly deposited at BO and not available.
Representative Specimens Examined—BANGLADESH. Rangamati:
Kaptai, Sitap(a)har west, Jarultala (Silchari), 24 Apr 1997, Huq and A. I.
10463 (L). BRUNEI. Belait: 27 Apr 1969, van Niel 4638 (L); Tutong:
between Belabau and Benutan on Tutong R., riverside forest, 31 Mar
1990, Coode 6436 (L). INDONESIA. Aceh: Sekundur Forest Reserve,
E. side of Gunung Leuser National Park, Langkat, Base camp at Aras
Napal, upper Besitang river area, 1 Aug 1991, de Wilde and de WildeDuyfjes 21227 (L); Jawa Barat: Bantam, Menés, 13 Mar 1913, Backer 7061
(L); Jawa Timur: Besuki, Mount Puger, 29 Feb 1940, Buwalda 7283 (L);
Kalimantan Tengah: Sampit, Along Sg. Penjahuan, N. of Kuala Kuajan,
17 Mar 1948, Kostermans 4656 (L); Kalimantan Timur: along Sg. Lunok
near camp Malinau, 23 Dec 1982, Axelius 329 (L); Wanariset I area, off km
5.5, Wanariset-Semoi Road, 29 Oct 1991, Ambriansyah and Arifin AA243
(L); Sulawesi Selatan: Samalea Hill, Tompokbalang, Maros, 30 Sep 1975,
Soenarko 365 (L); Sulawesi Tengah: Sopu Valley, ca. 80 km SSE of Palu,
7 May 1979, de Vogel 5251 (L). MALAYSIA. Johore: Sungai Bahan East
Coast of Jahore, 23 Jun 1934, Corner SFN. 28638 (L); Negeri Sembilan:
+
+
+
+
+
+
+
+
+
+
+
Lianas, up to 20 m high. Bark (gray) brown, inner bark
green. Twigs 3–6 mm diam., lenticellate, thinly hirsute or
thinly strigose in young parts, glabrescent. Leaves with
5–13 leaflets. Stipules ovate-triangular, 1.5–3
1.5–3 mm,
outside glabrous to thinly sericeous, inside glabrous. Petiole
3.5–11 cm long, striate, glabrous to thinly hirsute at the upper
side; rachis 4.5–14.5 cm long, striate, glabrous to thinly
hirsute at the upper side; pulvinus 5–11 mm long, hirsute,
especially at the base. Stipellae rarely present, acicular, ca.
1 mm long. Leaflets chartaceous to subcoriaceous; terminal
one elliptic to obovate, 7.5–18.5 4–9 cm, length/width ratio
1.8–2.3, base obtuse to rounded, apex acuminate, acumen 5–
20 mm long, rounded, upper surface glabrous or with widely
scattered appressed hairs, lower surface glabrous to thinly
strigose, midrib flat to slightly sunken above, nerves flat
above, 6–11 per side, 10–16 mm apart, curving towards the
apex and almost reaching the leaf margin, venation reticulate; lateral leaflets mostly like terminal one, ovate, elliptic or
obovate, 6.5–15.5 3.3–7.5 cm, length/width ratio 1.8–2.7;
pulvinus 5–11 mm long, sericeous at base. Pseudoracemes/
pseudopanicles axillary, rarely terminal, 8–22 cm long;
peduncle 1–3 cm long, glabrous to thinly strigose; lateral
branches up to 20 cm long. Bracts subtending brachyblasts
ovate to triangular, 1–1.7 0.7–1.1 mm, outside glabrous to
sericeous, ciliate, inside glabrous or with some hairs at the
base. Brachyblasts slender, cylindrical, 10–18 mm long, with
2 or 3 flowers at the apex. Bracts subtending flowers broadly
ovate to triangular, 0.5–1.1 0.4–1.3 mm, outside glabrous or
thinly sericeous, margin ciliate, inside glabrous. Pedicels 4–
9 mm long, glabrous or thinly sericeous. Bracteoles at the top
of the pedicels, elliptic to ovate or orbicular, 0.7–1.4 0.6–
1.3 mm, outside glabrous or thinly sericeous, margin ciliate,
inside glabrous. Calyx 3.5–5 mm high, pink, maroonish or
brown, outside glabrous to thinly sericeous, margin ciliate,
inside glabrous, sometimes sericeous at apex; tube 3–3.5 mm
high; upper lip indistinct or sometimes with 2 short semicircular lobes; lateral lobes (broadly) triangular, 0.5–1
2–3.5 mm; lower lobe triangular, 1–1.5 2–3 mm. Corolla
white or pink to purplish, standard with green blotch at base.
Standard: claw 2.5–3 mm long; blade broadly ovate to orbicular, 10–12 10–13 mm, apex emarginate, both sides glabrous or thinly ciliate at apex, basal callosities large. Wings:
claw 3.5–4 mm long; blade elliptic, 8–11 2.5–3.3 mm, apex
rounded, both sides glabrous or thinly ciliate at apex, upper
auricle 0.5–1 mm long, lower auricle absent, lateral pocket
1.5–2.5 mm long, sometimes hardly visible. Keel petals: claw
4.5–5 mm long; blade boat-shaped, 8–11 3–3.6 mm, apex
rounded, both sides glabrous or with some hairs at apex,
auricle indistinct, lateral pocket 3–5 mm long. Stamens
+
Derris danauensis Backer, Blumea 5: 513. 1945.—TYPE:
INDONESIA. Java: Rawah Danau, 11 Aug 1937, Van
Steenis 10539 (holotype: BO?; isotype: L!).
+
Derris caudata Backer, Blumea 5: 513. 1945.—TYPE: INDONESIA.
Java: Bantam, Menés, 13 Mar 1913, Backer 7061 (holotype:
BO?; isotype: L!).
+
+
Derris pachycarpa Merr., J. Straits Branch Roy. Asiat. Soc. 86:
310. 1922.—TYPE: MALAYSIA. Borneo: Sabah, Batu
Lima, BS (Ramos) 1250 (holotype: PNH, lost; isotype: K!).
435
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2012]
Fig. 20.
Distribution of Paraderris montana (Benth.) Adema.
436
SYSTEMATIC BOTANY
Jelebu district, Pasoh Forest Reserve, 10 Jul 1996, Gardette 1997 (L);
Pahang: Jenderak Halt, Railway Track, 28 Mar 1959, Kadim and Mahmood,
KM 48 (L); Perak: Maxwell’s Hill-Tea Station, 4 Mar 1965, Hardial and
Samsuri 265 (L); Sabah: Ranau, roadside, 27 Apr 1988, Amin et al. SAN
121505 (L); Sarawak: By rail to Matang dam., 29 Sep 1963, Smythies
S 14074 (L); Selangor: Ulu Selangor, 19 Jul 1969, Chan FRI 11246 (L).
PAPUA NEW GUINEA. Central Prov: near Mori river, ca. 20 km NE
of Cape Rodney, 26 Sep 1968, Paijmans 439 (L); East Sepik: Angoram
subdistrict, upper Karawar river, 2 Sep 1968, Dornstreich NGF MDD-10
(L); Gulf: Kikori river, 4 miles downstream from Kikori station, 30 Mar
1974, Womersley NGF 46495 (L); Morobe: Markham bridge, Lae subdistrict,
20 Sep 1977, Katik LAE 70799 (L); Northern Prov: Subdist. Kokoda, 0.5 km
east of Hanjiri village, on the Luwuni River bank, 27 Sep 1975, Wiakabu and
Kairo LAE 70237 (L); West New Britain: Kandrian subdist., Unai River,
Fullerborn Harbour, 24 Feb 1965, Sayers NGF 21735 (L). THAILAND.
Nakhon Si Thammarat: Lansagah, Gahrome Falls, Khao Luang National
Park, 4 Mar 1986, Maxwell 86–140 (L); Pangnga: Bang To, 24 Feb 1929,
Kerr 17228 (L); Pattani: Khao Kalalwiti, 1 Apr 1928, Kerr 14963 (L);
Songkhla: Khao Klong Rhang, Na Mom district, 24 Sep 2009, Sirichamorn
YSM2009–21 (L); Trang: Muang District, Khao Chong Forest Park, 10 Mar
1985, Maxwell 85–283 (L).
Acknowledgments. We wish to express our sincere thanks to all curators and staff of BCU, BK, BKF, CMU, K, L, and P for providing laboratory
facilities, specimen loans, and various kinds of support. The first author is
also indebted to the Royal Thai Government for funding his PhD study. Jan
Frits Veldkamp kindly translated the diagnosis of two new species into Latin.
Ms. Anita Walsmit Sachs skillfully prepared the beautiful illustrations.
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