A REVISION OF WRIGHTIA

(APOCYNACEAE: APOCYNOIDEAE) IN MALESIA

DAVID J. MIDDLETON1

Abstract. The genus Wrightia is revised for the Malesian region. Eight species are recognized, one of which
has two subspecies. One new species, Wrightia palawanensis, is described, and one new combination, Wrightia
novobritannica, is made. A key to the species is given, and the taxa are described. Lectotypes and neotypes are
designated for a number of names.
Keywords: Apocynaceae, Apocynoideae, revision, Wrightia, Malesia.

The genus Wrightia R. Br. was last revised by and thorough assessment of the morphological
Ngan (1965) who recognized 23 species and characters in Wrightia.
several subspecies. Since then a small number of The genus shows some interesting biogeo-
additional species have been described (Bahadur graphic patterns in Malesia. Most curious is the
and Bennet, 1978; Lý, 1985; Middleton and complete absence of any collections of the genus
Santisuk, 2001) from India, Vietnam, and from Borneo, despite the widespread distribu-
Thailand. The genus is here revised for the forth- tion of taxa like Wrightia laevis and Wrightia
coming Flora Malesiana account of the pubescens subsp. lanitii in the surrounding
Apocynaceae, subfamilies Rauvolfioideae and regions. There are two species, Wrightia hanleyi
Apocynoideae (sensu Endress and Bruyns, and Wrightia palawanensis, endemic to Palawan
2000). Eight species are recognized, one of in the Philippines, and another, Wrightia can-
which has two subspecies. A new species from dollei, more widespread but still endemic to the
recently collected material in Palawan is Philippines. Wrightia novobritannica is known
described, and the status of some of Ngan’s sub- only from one collection from the Papua New
species within Wrightia pubescens are Guinea Island of New Britain and would appear
reassessed. Ngan (1965) has given a very good to have similarities to Wrightia candollei.
MATERIALS AND METHODS
Approximately 350 herbarium specimens have been seen unless otherwise indicated.
from Asia and Malesia have been studied. These All dimensions given are for dried specimens
are from the following herbaria: A, AAU, B, except for androecium and gynoecium charac-
BISH, BKF, BM, BO, BRI, C, CANB, E, G, ters, which are from flowers reconstituted by
GH, HM, HN, HNU, K, KEP, L, M, MICH, boiling in water.
MO, NSW, NY, P, PNH, SING, SINU, U, US, Z Author citation follows Brummitt and Powell
(Holmgren et al., 1990). All specimens cited (1992).
TAXONOMIC TREATMENT
Wrightia R. Br., Asclep. 62. 1810; R. Br., Prod. Ngan, Ann. Miss. Bot. Gard. 52: 114. 1965;
467. 1810; Benth. & Hook. f., Gen. Pl. 2: 712. Tsiang & Li, Fl. Reipubl. Popularis Sin. 63:
1876; K. Schum. in Engler & Prantl, Nat. 119. 1977; Lý, Feddes Repert. 97: 619. 1986;
Pflanzenfam. 4, 2: 183. 1895; Pitard in Li et al., Fl. China 16: 174. 1995; Middleton,
Lecomte, Fl. Gén. Indo-Chine 3: 1182. 1933; Fl. Thailand 7: 79. 1999. Wrightia sect.
Pichon, Mém. Mus. Nat. Hist. Nat, sér. B, Bot. Bammatophyton A.DC., Prod. 8: 404. 1844.
1: 73. 1950; Pichon, Not. Syst. 14: 77. 1951; Type species: Wrightia pubescens R. Br.

I would like to thank the curators and staff of the herbaria that loaned material or hosted me on visits (A, AAU, B, BISH,
BKF, BM, BO, BRI, C, CANB, E, G, GH, HM, HN, HNU, K, KEP, L, M, MICH, MO, NSW, NY, P, PNH, SING, SINU,
U, US, Z) and the curatorial staff of the Harvard University Herbaria for their management of the loans.
1 Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Current address: Royal

Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland. E-mail: d.middleton@rbge.ac.uk

Harvard Papers in Botany, Vol. 10, No. 2, 2005, pp. 161– 182.
© President and Fellows of Harvard College, 2005.
162 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

Synonyms: Annasser Blanco, Fl. Filipp. 112 corona of varying degrees of elaboration usu-
(1837). Type species: Anasser lanitii ally present, rarely absent or completely adnate
Blanco (= Wrightia pubescens subsp. to the corolla. Stamens inserted at corolla mouth
lanitii (Blanco) Ngan) or in tube; strongly exserted from or completely
Scleranthera Pichon, Not. Syst. 14: 88. included in corolla tube; filaments usually quite
1951. Type species: Scleranthera cambo- short and wide; anthers narrowly triangular,
diensis (Pierre ex Pitard) Pichon (= apex acuminate, base sagittate, sterile at sides
Wrightia dubia (Sims) Spreng.) and base with fertile locules on the inner upper
half, adnate to the style head, often pubescent
Wrightia sect. Walidda A. DC. in DC., on the surface facing inward and/or the surface
Prod. 8: 407. 1844. Walidda (A. DC.) facing outward. Disk absent. Gynoecium of 2
Pichon, Not. Syst. 14: 87. 1951. Type free carpels united into a common style or of
species: Walidda antidysenterica (L.)
two connate carpels, glabrous, rarely pubescent;
Pichon (= Wrightia antidysenterica L.)
style broadest near top; ovules numerous. Fruit
Shrubs or small trees. Leaves opposite; usu- of paired follicles or connate follicles; fusiform.
ally with glands in the axils. Inflorescence a ter- Seeds linear or narrowly fusiform; with a coma
minal cyme; few to many flowered. Sepals with directed toward the base of the follicle.
large or small colleters in the sinuses inside, the There are ca. 25 species in Africa, Asia, and
larger ones almost covering the inner surface of Australia, and 8 species in Malesia (one of
the sepals. Corolla lobes sinistrorse; mature which may not be native, see note under
flower rotate, salverform or infundibuliform; Wrightia religiosa).

KEY TO THE SPECIES OF WRIGHTIA IN MALESIA

1a. Corolla infundibuliform; stamens completely included in the corolla tube . . . . . . . . . . . . . . . . . . . . . 2. W. dubia
1b. Corolla rotate, salverform, or somewhat infundibuliform; stamens clearly exserted from the corolla tube. . . . . . 2
2a. Flowers pendulous; corona absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. W. religiosa
2b. Flowers pendulous or not; corona present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Corona with antepetalous lobes divided more than halfway to base, giving the whole corona a fimbriate
look. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. W. laevis
3b. Corona with antepetalous lobes entire or divided less than halfway to base . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Antepetalous corona lobes deeply divided or at least strongly dentate . . . . . . . . . . . . . . . . . . . . . 7. W. pubescens
4b. Antepetalous corona lobes entire or at most weakly crenate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Gynoecium apocarpus but united into a single style; fruits paired; Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5b. Gynoecium syncarpous; fruits solitary; Philippines and New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6a. Corolla tube 3.8–4.0 mm long; stamens inserted in the upper half of the corolla tube . . . . . . 6. W. palawanensis
6b. Corolla tube 1.8–2.8 mm long; stamens inserted in the lower half or around the middle of the corolla
tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. W. hanleyi
7a. Corolla tube 3.7–10.0 mm long, 1.7–5.5 times as long as sepals; Philippines . . . . . . . . . . . . . . . . 1. W. candollei
7b. Corolla tube ca. 3.5 mm long, ca. 1.6–1.9 times as long as sepals; New Guinea
(New Britain) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. W. novobritannica

1. Wrightia candollei Vidal, Phan. Cuming. midrib or glabrous beneath, 10–16 pairs of sec-
Philipp. 186. 1885 (as cadollei). Fig. 1. ondary veins, tertiary venation reticulate.
Synonym: Wrightia pubescens subsp. candollei Inflorescence 2.5–3.5 cm long, often with leafy
(Vidal) Ngan, Ann. Miss. Bot. Gard. 52: bracts; peduncle 0.2–1.3 cm long, sparsely
153. 1965. TYPE: PHILIPPINES. Luzon, pubescent, pedicels 4–7.5 mm long, pubescent.
Manila, Cuming 1453 (Lectotype: P, des- Sepals ovate, 1.1–4.0 × 1.8–4.5 mm, 0.3–1.1
ignated here; Isotype: BM, K). times as long as wide, apex rounded, sparsely
pubescent, ciliate, colleters large in sinuses.
Branchlets sparsely to densely puberulent. Corolla orange; tube 3.7–10.0 mm long,
Leaves: petiole 4–8 mm long, glabrous to 1.7–5.5 times as long as sepals, 0.4–0.5 times
sparsely puberulent; blade elliptic, 4–20 × as long as lobes, minutely puberulent at top of
3.0–6.5 cm, 1.3–3.7 times as long as wide, apex tube outside, glabrous inside; lobes 6.5–15.0 ×
acuminate, base cuneate to rounded, sparsely 3.3–9.0 mm, 1.7–2.0 times as long as wide,
puberulent to glabrous above, sparsely to minutely puberulent inside and outside; ante-
densely pale puberulent all over or only on petalous corona lobes 4.0–6.5 mm long, adnate
2005 MIDDLETON, A REVISION OF WRIGHTIA 163

FIGURE 1. Wrightia candollei Vidal (Ridsdale 1418, A).

164 HARVARD PAPERS IN BOTANY
to corolla except at apex, apex subentire to
weakly crenate, glabrous inside, 3.8–5.0 times
as long as alternipetalous lobes; alternipetalous
corona lobes simple or slightly bifid 0.7–1.7
mm long; alternating corona lobes lacking.
Stamens inserted at 0.7–0.8 of tube length from
corolla base, exserted from tube; filaments
0.8–1.0 mm long; anthers 5.0–7.8 × 1.1–1.8
mm, pubescent on the outside or only at tips
outside. Gynoecium of two connate carpels,
ovary 1.5–2.6 mm high; style and style head
5–7 mm long. Fruit of a single follicle that
splits lengthwise at maturity as each of the two
component follicles dehisces, 20.5–22.0 cm
long, 1.4–2.2 cm wide, glabrous. Seeds not
observed.
Distribution: Philippines.
Habitat: evergreen or mixed deciduous forest.
Additional material studied: PHILIP-
PINES. Sine loc.: Loher 6519 (M), 6530 (M),
6542 (M); Escritor 21296 (P); McGregor
18579 (P). BOHOL: Ramos 42712 (A).
CAMOTES: Ramos 41607 (BO, L). CEBU:
Lopez & Reyes 27331 (BO, L, P).
GUIMARAS: Gammill 228 (NSW). LEYTE:
Hubasan, Baybay, Gaerlan et al PPI 10603
(GH). LUZON: Bataan: Alongapo, Bartlett
14074 (A). Batangas: Bo. Biga, Lobo, Sulit
PNH 7428 (A, PNH). Benguet: Twin Peaks,
Elmer 6348 (NSW, P). Bulacan: Angat
Watershed area, Norzagaray, Barbon et al PPI
22357 (PNH); near Kay Tianak and Ipo,
Bartlett 14720 (A). Cagayan: Brgy. San Juan,
Sta. Praxedes, Garcia et al PPI 18505 (PNH).
Nueva Ecija: Curran 8442 (SING).
Pangasinan: Alaminos, McGregor 41431
(KEP); Bolinao, Clemens 18171 (BO, C, S);
sine loc., Ramos 8284 (L). Rizal: San Mateo,
Ahern’s collector 1116 (BO, NSW); sine loc.,
Ahern’s collector 3155 (BO, P, SING); sine
loc., Robinson 6757 (BRI, L). Zambales: Acoje
Mine, Santa Cruz, Ridsdale 1418 (A, BO, KEP,
L, MO); San Marcelino, Bgy. Sayasay, Mt.
Mandakogdog, Majaducon & Fernando 8455
(L). – MASBATE: Elmer 3049 (P); Merrill
3082 (BO, L). MINDORO: occidental: Mt.
Gonting, Lubang Island, Romero & Fuentes
PPI 37562 (A, PNH). Oriental: Mabaho Ridge,
Bo. Kabalwa, Mansalay, Sulit PNH 17044 (A,
PNH). NEGROS: Contreras 23404 (BO).
Occidental: Danjugan Island, Bulata, Cauayan,
Madulid & Majaducon PPI 36005 (L).
PALAWAN: Puerto Princesa. Irawan River
Valley, Tatanarom, trail up Mt. Beaufort,
Vol. 10, No. 2
Soejarto et al 6456 (A, MO); Taytay Bay,
Apulit Island, Merrill 9426 (BO, BRI, L, P,
SING); sine loc., Curran 4510 (L); sine loc.,
Fenix 1183 (Z).
This species was included as a subspecies of
Wrightia pubescens by Ngan (1965), but it dif-
fers substantially from that species in the shape
and form of the corolla and corona and in the
flower color. Therefore, it has been raised back
to specific level.
2. Wrightia dubia (Sims) Spreng., Syst. Veg. 1:
638. 1825; King & Gamble, J. As. Soc. Beng.
74, 2: 465. 1907; Ngan, Ann. Miss. Bot. Gard.
52: 172. 1965; Whitmore, Tree Fl. Mal. 2: 24.
1973; Lý, Feddes Repert. 97: 622. 1986;
Turner, Gard. Bull. Sing. 47: 131. 1997;
Middleton, Fl. Thailand 7: 83. 1999. Fig. 2.
Basionym: Cameraria dubia Sims in Curtis,
Bot. Mag. 39: t.1646. 1814. Scleranthera
dubia (Sims) Pichon, Not. Syst. 14: 90.
1951. TYPE: Illustration in plate 1646 in
Curtis, Bot. Mag. 39 (1814).
Synonyms: Strophanthus jackianus Wall. ex
G.Don, Gen. Syst. 4: 85. 1837; Hook. f.,
Fl. Ind. 3: 656. 1882; Ridley, Fl. Mal.
Pen. 2: 355. 1923. TYPE: MALAYSIA.
Penang, Wallich 1643 (Lectotype: K;
Isotype: K-W).
Wrightia dubia var. membranifolia King
& Gamble, J. As. Soc. Beng. 74, 2: 466.
1907. TYPE: THAILAND, Phuket,
Tongkah, February 1893, Curtis 2915
(Lectotype: SING, designated here).
Wrightia cambodiensis Pierre ex Pitard
in Lecomte, Fl. Gén. Indo-Chine 3: 1184.
1933; Kerr in Craib, Fl. Siam. En. 2: 456.
1939. Scleranthera cambodiensis (Pierre
ex Pitard) Pichon, Not. Syst. 14: 89.
1951. TYPE: VIETNAM, Ninh Thuan,
Ca Na, Pierre 4402 p.p. (Lectotype: P;
Isotype: A, P).
Wrightia rubriflora Pitard in Lecomte,
Fl. Gén. Indo-Chine 3: 1185. 1933.
TYPE: VIETNAM, Ninh Thuan, Ca Na,
Poilane 5955 (Lectotype: P; Isotype:
HM, NY, P).
Wrightia kontumensis Lý, Feddes Repert.
96: 173. 1985; Lý, Feddes Repert. 97:
622. 1986. TYPE: Vietnam, Gialai-
Kontum, Loc P 2968 (Holotype: HNU;
Isotype: HN).
2005 MIDDLETON, A REVISION OF WRIGHTIA 165

FIGURE 2. Wrightia dubia (Sims) Spreng. (Sidek bin Kiah S.151, A).
166 HARVARD PAPERS IN BOTANY
Shrub to 2 m tall. Branchlets glabrous or
sparsely puberulent, becoming glabrous and
lenticellate. Leaves: petiole 2–11 mm long;
blade papery to subcoriaceous, elliptic, oblong
or obovate, 3.6–26.5 × 1.8–9.8 cm, 2.1–5.2
times as long as wide, apex acuminate to sub-
caudate, base cuneate to rounded, puberulent
abaxially, only on midrib and secondary veins
or, rarely, glabrous and then both surfaces papil-
late, 6–14 pairs of secondary veins, ascending,
tertiary venation reticulate. Inflorescence
2.5–4.5 cm long; sparsely puberulent or
glabrous; pedicels 4.0–5.5 mm long, robust.
Flowers inodorous. Sepals ovate, 1.3–3.0 ×
1.3–2.0 mm, 1.1–1.3 times as long as wide, apex
acute to obtuse; sparsely puberulent or glabrous,
ciliate; colleters wide and large. Corolla orange,
pink or reddish; infundibuliform; tube 6.0–15.5
mm long, 3.8–4.7 times as long as sepals,
0.5–0.6 times as long as lobes, glabrous or,
rarely, pubescent-papillose at top of tube out-
side, glabrous inside; lobes 10–28 × 6–9 mm,
1.9–3.2 times as long as wide, ovate, apex
acuminate to obtuse, pubescent-papillose on
lobes outside and slightly so inside; antepetalous
corona lobes adnate to the corolla, 1.5–3.7 mm
long, 1.7–2.5 times as long as alternipetalous
lobes, glabrous; alternipetalous corona lobes
0.6–2.2 mm long; alternating corona lobes lack-
ing. Stamens inserted at 0.3–0.4 of tube length
from corolla base, completely included in the
tube; subsessile; anthers 4.9–5.6 × 1.0–1.2 mm,
pubescent outside. Gynoecium of 2 free carpels
united into a common style, ovaries 1.6–2.5 mm
long, glabrous; style and style head 3.0–4.7 mm
long. Fruit of paired follicles, 13–30 cm long,
3.5–6.0 mm wide, glabrous, not lenticellate.
Seeds linear, 12.5–25.4 × 1.2–2.5 mm; coma
2.6–5.5 cm long.
Distribution: Cambodia, Laos, Vietnam,
Thailand, Peninsular Malaysia, Singapore.
Habitat: growing in wide variety of forest
habitats at lower altitudes.
Additional collections studied: MALAYSIA:
Kedah: Sine loc., Ridley 5521 (SING); Pulau
Langkawi, Telok Apau, Haniff 7086 (SING);
Pulau Langkawi, Stone et al 120 (KLU); Pulau
Langkawi, Pulau Singa Besar, Zainudin et al
4391 (L). Penang: Curtis 188 (SING); Curtis
s.n. (SING); below Waterfall gardens, Sinclair
39023 (E, SING); valley of stream below
waterfall gardens, Sinclair [SING39023] 6627
(E); Butterworth, O’Sullivan s.n. (SING).
Vol. 10, No. 2
Perlis: Kaki Bukit, Burtt & Woods B 1729 (E),
Kiah 35275 (A, BO, L), Kiah s.n. (SING).
Terengganu: Gunong Tebu, Besut, Selvaraj FRI
13006 (KEP, L). SINGAPORE: Botanic
Gardens, Jumali 2866 (SINU), Kiah S 151 (E).
This is the most distinctive species of
Wrightia in Malesia with its pink flowers and
without the anthers exserted.
3. Wrightia hanleyi Elmer, Leafl. Philipp. Bot.
4: 1465. 1912; Ngan, Ann. Miss. Bot. Gard. 52:
168. 1965. TYPE: PHILIPPINES, Palawan,
Puerto Princesa, Mt. Pulgar, Elmer 12873
(Lectotype: GH, designated here; Isotype: A,
BISH, BO, E, K, L, NSW, NY, US, Z). Fig. 3.
Shrub or treelet to 4 m tall. Branchlets
glabrous, sparsely lenticellate with age.
Leaves: petiole 3 mm long, glabrous; blade nar-
rowly ovate to elliptic, 3–11 × 1.3–4.6 cm,
1.6–3.8 times as long as wide, apex acuminate
with a sharp tip, base cuneate to rounded,
glabrous above and beneath, 6–15 pairs of sec-
ondary veins, tertiary venation reticulate.
Inflorescence terminal, 0.8–2.0 cm long, few-
flowered; peduncle 0.1–1.0 cm long, glabrous;
pedicels 1.5–4.0 mm long, glabrous. Sepals
ovate, 1.0–1.5 × 0.9–1.1 mm, 1.1–1.4 times as
long as wide, apex acute to obtuse, glabrous,
ciliate, colleters small in the sinuses. Corolla
greenish-yellow to red; tube with a lower nar-
row part and a campanulate upper part, with a
raised lip inside between the lower and upper
tube, 1.8–2.8 mm long, 1.5–1.9 times as long as
sepals, 0.6–0.7 times as long as lobes, minutely
puberulent at top of tube outside, glabrous
inside; lobes 3.0–3.9 × 1.5–3.0 mm, 1.3–2.0
times as long as wide, apex rounded, minutely
puberulent inside and outside; antepetalous
corona 1.5–2.9 mm long, entire, adnate to
corolla on medial line but with sides free, apex
subentire or slightly crenate, glabrous inside,
4.1–7.5 times as long as alternipetalous lobes;
alternipetalous corona lobes apparently absent
or small and simple, 0.2–0.7 mm long; alternat-
ing corona lobes lacking. Stamens subsessile,
inserted at 0.3–0.5 of tube length from corolla
base, exserted from tube; anthers 2.6–5 ×
0.4–1.0 mm, pubescent only at the top outside.
Gynoecium of 2 free carpels united into a com-
mon style, ovaries 0.7–1.1 mm high, glabrous;
style and style head 1.8–4.0 mm long. Fruit
paired, linear, 6.6–18.0 cm long, 2.5–4.0 mm
wide, glabrous. Seeds 10.0 × 0.6 mm; coma 21
mm long.
2005 MIDDLETON, A REVISION OF WRIGHTIA 167

FIGURE 3. Wrightia hanleyi Elmer (Soejarto & Reynoso 6261, A).

168 HARVARD PAPERS IN BOTANY
Distribution: Philippines (Palawan).
Habitat: in evergreen forest.
Additional collections studied: PHILIP-
PINES: PALAWAN: Lipuun, Mendoza 91195
(L); Mt. Cleopatra Ranges, So. Calabayog, Brgy.
Tanabag, Puerto Princesa, Reynoso et al PPI
24249 (PNH); Narra Municip., Bgy Calatigas,
Purok Campsite, Soejarto et al 8602 (L); Puerto
Princesa (Mt. Pulgar); Puerto Princesa Munic.,
Bacungan, Hills above Nagtabon beach,
Soejarto et al 6479 (MO); Puerto Princesa
Munic., Takdua Zigzag, 35–40 km from Puerto
Princesa, Soejarto & Reynoso 6261 (A, L);
Puerto Princesa, Tagburus, Stone et al PPI 262
(PNH); Quezon, Tawa-tawa, Gaerlan et al PPI
13448 (PNH); South of Narra, Bato-Bato river,
Ridsdale 1080 (L); St Paul’s Bay, Mt
Bloomfield, Ridsdale SMHI 1620 (L); Taytay,
Merrill 9373 (L, NSW, P, SING).
4. Wrightia laevis Hook. f., Fl. Brit. Ind. 3: 654.
1882; King & Gamble, J. As. Soc. Beng. 74, 2:
465. 1907; Ridley, Fl. Mal. Pen. 2: 353. 1923;
Kerr, Fl. Siam. En. 2: 457. 1939; Merrill &
Perry, J. Arnold Arb. 24: 215. 1943; Ngan, Ann.
Miss. Bot. Gard. 52: 136. 1965; Whitmore,
Tree Fl. Mal. 2: 24. 1973; Tsiang & Li, Fl.
Reipubl. Popularis Sin. 63: 125. 1977; Lý,
Feddes Repert. 97: 623. 1986; Turner, Gard.
Bull. Sing. 45: 37. 1993; Ba et al., PROSEA
5.2: 513. 1995; Li et al., Fl. China 16: 175.
1995; Turner, Gard. Bull. Sing. 47: 131. 1997;
Middleton, Fl. Thailand 7: 85. 1999.
Basionym: Wrightia tinctoria R. Br. var. laevis
(Hook. f.) Pichon, Not. Syst. 14: 80. 1951.
TYPE: MALAYSIA, Malacca, Mt. Ophir,
Maingay 1065 (Lectotype: K, designated here;
Isotype: K). Fig. 4.
Synonyms: Wrightia millgar F. M. Bailey,
Dept. Agric. Brisbane Bot. Bull. 7: 65.
1893. – Wrightia laevis subsp. millgar (F.
M. Bailey) Ngan, Ann. Miss. Bot. Gard.
52: 137. 1965; Forster, Fl. Aust. 28: 191.
1996. TYPE: AUSTRALIA.
Queensland: Barron River, Cowley 7D
(Holotype: BRI [comprising 2 sheets]).
Wrightia hainanensis Merr., Philipp. J.
Sci. 21: 352. 1922. TYPE: CHINA.
Hainan: Fong Mok Sze, “Chinese
Collector” 425 (Holotype: PNH,
destroyed; photo of holotype: A).
Wrightia balansae Pitard in Lecomte, Fl.
Gén. Indo-Chine 3: 1188. 1933. TYPE:
VIETNAM. Ha Tay, Tu Phap, Balansa
Vol. 10, No. 2
2118 (Lectotype: P; Isotype: P).
Wrightia macrocarpa Pitard in Lecomte,
Fl. Gén. Indo-Chine 3: 1190. 1933.
TYPE: Balansa 2115 (Lectotype: P),
VIETNAM, Ha Tay, Tu Phap.
Wrightia sorsogonensis Elmer, Leafl.
Philipp. Bot. 10: 3698. 1939, nom. nud.
Based on: Elmer 15595 (A, BM, BO, C,
GH, K, L, MO, NSW, NY, P, PNH, S, Z),
PHILIPPINES, Luzon, Sorsogon, Irosin,
Mt. Bulusan.
Wrightia hainanensis var. variabilis
Tsiang, Sunyatsenia 4: 47. 1939. TYPE:
Tsiang 2679 (Lectotype: A, designated
here; Isotype: K, NY), CHINA,
Guangdong, Sunyi District.
Wrightia hainanensis var. chingii Tsiang,
Sunyatsenia 4: 48. 1939. TYPE: Ching
5539 (Holotype: IBSC n.v.; Isotype:
NY), CHINA, Guangxi, Mung Tung
Kon, N of Luchen.
Wrightia laevis subsp. novoguineensis
Ngan, Ann. Miss. Bot. Gard. 52: 138.
1965. TYPE: Brass 5570 (Holotype: A;
Isotype: BM, BO, BRI, K, NY, US),
PAPUA NEW GUINEA, Central
Province, Mt. Tafa.
Usage synonym: Wrightia tinctoria auct. non
R. Br., Pitard, Fl. Gén. Indo-Chine 3
(1933) 1187.
Shrub or tree to 37 m tall and 106 cm dbh, but-
tresses present or absent. Bark smooth to rough
and flakey, gray, grayish-brown or cream-
brown; inner bark white, cream, or pale brown.
Branchlets glabrous, sparsely lenticellate with
age. Leaves: petiole 3–7 mm long; blade subco-
riaceous, elliptic, ovate or obovate, 3.1–17.3 ×
1.4–7.5 cm, 1.3–4.1 times as long as wide, apex
acuminate, base cuneate to obtuse, glabrous,
4–10 pairs of secondary veins, strongly ascend-
ing, curved, tertiary venation mostly paler than
lamina in dried specimens, almost perpendicular
to midrib and oblique to secondary veins and
also reticulate. Inflorescence 1.8–10.0 cm long;
peduncle 1.0–1.6 cm long, glabrous to sparsely
puberulent; pedicels 8–19 mm long, delicate,
glabrous or puberulent. Sepals ovate, 0.7–2.5 ×
0.8–2.3 mm, 0.9–1.2 times as long as wide, apex
obtuse to rounded, glabrous to densely puberu-
lent, ciliate; colleters wide and large. Corolla
white, pale yellowish-white to yellow-orange;
subrotate; tube 1.1–3.0 mm long, 1.5–1.7 times
as long as sepals, 0.2–0.4 times as long as
2005 MIDDLETON, A REVISION OF WRIGHTIA 169

FIGURE 4. Wrightia laevis Hook.f. (Kochummen FRI 16446, A).

170 HARVARD PAPERS IN BOTANY
lobes, glabrous or minutely puberulent at top of
tube outside, glabrous inside; lobes 6.1–14.0 ×
2.6–5.2 mm, 1.6–2.8 times as long as wide,
elliptic to slightly obovate, apex rounded,
minutely puberulent outside and inside; corona
of antepetalous, alternipetalous, and alternate
lobes; antepetalous lobes adnate to the corolla
at base, fimbriate, 2.8–9.4 mm long, 0.9–1.8
times as long as alternipetalous lobes, glabrous;
alternipetalous lobes free, simple, deeply bifid,
or sometimes compound, 2.4–6.5 mm long;
alternating lobes 0.6–5.8 mm long, simple,
shorter than the other two whorls. Stamens
inserted at 0.8–0.9 of tube length from corolla
base, completely exserted; filaments 0.2–1.7
mm long; anthers 3.9–5.6 × 0.7–1.6 mm,
pubescent outside, sometimes only sparsely so
at base. Gynoecium of 2 free carpels united into
a common style, ovaries 0.7–1.8 mm long,
glabrous; style and style head 3.5–5.5 mm
long. Fruit of paired follicles; 10–50 cm long,
5–17 mm wide, glabrous, lenticellate or not
(see note). Seeds linear; 18–22 x 1.7–2.5 mm;
coma 3.3–4.0 cm long.
Distribution: southern China and continental
Southeast Asia through Malesia to Queensland.
Habitat: in a wide variety of forest habitats.
Additional collections studied: Sine loc.:
Forbes 2802 (L). EAST TIMOR. Bobonaro,
Balibo: Widjaja 1213 (L). INDONESIA:
WEST PAPUA: Bernhard Camp, Idenburg
River, Brass 13893 (BRI, L); Kebar, Asiti, Vink
BW 11420 (A, CANB, L, SING); Manokwari,
Kostermans 256 (BO, L, SING); Ransiki,
Kostermans 100 (BO, L); Masni Plain, c.25 km
W of Manokwari, Koster BW 11046 (L);
Oemboei, near Andai, SW of Manokwari,
Koster BW 11804 (L); Pulau Japen, Schram BW
10592 (BO, CANB, L), van Dijk 309 (BO);
Pulau Japen, Mariarotu, van Dijk 168 (BO),
196 (BO), 226 (BO); Pulau Japen, Wasabori
near Serui, Aet & Idjan 419 (BO, L); Pulau
Japen, Seroei, Neth. Ind. For. Serv. bb 30530
(L, SING), bb 30391 (L, NY, SING), bb 30417
(L, SING), bb 30447 (SING); Plateau N of
River Pami, 8 km NW of Manokwari, Koster
BW 4349 (CANB, L), Koster BW 4355
(CANB, L); Sausapor, Sorong, Versteegh BW
3991 (CANB, L), BW 4628 (L); Sekoli Plain,
near Jayapura, Iwanggin BW 9136 (BO, BRI,
L); Tiporra, Rijklof van Goensbaai, Stefels BW
5105 (L); Warsamson valley, E of Sorong, Moll
BW 11560 (BO, CANB, L); Wersar, 6 km S of
Vol. 10, No. 2
Teminaboean, Versteegh BW 4971 (L). SUMA-
TRA. Bt. Gajabuih, Ulu Gadut, 15 km E of
Padang City, Hotta 25889 (BO); Palembang,
Unknown 1078 (L); Tanjong Ning, Palembang,
Forbes 2766 (L, SING); near Sigirik mountain
within Sebelah Nature Reserve, Laumonier
TFB 4464 (L); Gunung Leuser NP, Ketambe,
valley of Lau Alas, ca. 35 km NW of Kutatjane,
de Wilde & de Wilde-Duyfjes 12369 (BO, KEP,
L, US); Gunung Leuser NP, Middle Alas River,
15 km N of Galombang, de Wilde & de Wilde-
Duyfjes 20199 (BO, L, US, Z); vicinity of
Loemban, Asahan, Rahmat Si Boeea 7533 (S,
SING). MALAYSIA. Pulau Pomgoh, Curtis
2943 (SING); Johor: Kluang, Labis FR, Moton
KEP 94083 (KEP, L, SING), Kochummen FRI
16446 (A, KEP, L, SING); Gunong Besar, Ulu
Sungei Segamat, Samsuri & Ahmad SA 691 (C,
G, KEP); Near Ulu Sungei Redong, Jumali 875
(SING); Ulu Sungei Redong, Kg. Tenang,
Labis, Jumali 4339 (SINU). Kelantan: Gua
Panjong at Gua Ninik, Henderson 19599
(SING); Kuala Mersing, Sg. Brok, Ng FRI
5392 (KEP); SE, Ulu S. Labir Kecil, Whitmore
FRI 4405 (KEP, L, SING). Kuala Lumpur: Ulu
Gombak FR, Kochummen KEP 99607 (KEP).
Negri Sembilan: Johol, Ridley s.n. (SING); Ulu
Pedas, Nur 11728 (SING). Pahang: Jalan
Simpon, FR, Taman Negara, Ang FRI 23347
(KEP, L), Teo & P. 548 (L, SING); Krau Game
reserve, Kuala Lompat, Saw FRI 36282 (KEP);
Lanchang, Zainudin 3326 (L); Lesong Forest
Reserve, Rompin Dist., Maxwell s.n. (L,
SINU); Nr Batu Balai, Burkill & Haniff 15826
(SING); S. Tembeling nr K. Keniyum S.
Redab, Whitmore FRI 8568 (KEP, L, SING);
Sungei Lenget, Rumping, Yeob 3205 (KEP);
Taman Negara, 3 km from Kuala Tahan, van
Balgooy 2436 (L, MO, Z), 2488 (L, Z), Chin &
Mahmud 1289 (KLU). Perak: sine loc., King’s
Collector 8614 (BO, L), Scortechini 77 b
(SING), Ridley 14949 (SING); Padang Batang,
Tapah Hills FR, Loh FRI 17384 (KEP, L,
SING); Ulu Temango, Ridley 14605 (SING).
Selangor: Bukit Endong FR, Symington 57059
(KEP), 57060 (KEP), 57061 (KEP); Bukit
Lagong FR, Kochummen FRI 2973 (KEP, L,
SING), Sakat KEP 116122 (KEP), Chan FRI
17654 (KEP, L), Sinclair SFN 40103 (BO, E,
KEP, L, SING), Suppiah FRI 108885 (KEP, L),
Pagi KEP 98711 (KEP, L); Dusun Tua, Ridley
s.n. (SING); Gaiting Bulai, Ridley 7566
(SING);Gunung Nuang Ulu Langat, Symington
2005 MIDDLETON, A REVISION OF WRIGHTIA
51751 (KEP); Semempih, Hume 8352 (SING);
Ulu Gombak FR, Kuala Lumpur, Kochummen
99607 (BO, L, SING); Ulu Gombak, Univ.
Malaya Field Studies Centre, Soepadmo &
Students FSC 832 (KLU, L). PHILIPPINES.
Sine loc.: Mendoza 42092 (L, SING). LUZON:
Sorsogon: Irosin, Mt Bulusan. MINDANAO,
Davao: de Mesa 27538 (BO, L, SING).
PALAWAN: Mt Cleopatra, Reynoso et al PPI
24423 (L); Mt.Cleopatra Ranges, So.
Calabayog, Brgy. Tanabag, Puerto Princesa,
Reynoso et al PPI 24244 (PNH); So.
Calabayog, San Rafael, Puerto Princesa,
Barbon et al PPI 18700 (PNH). SAMAR:
Loquilocon, Wright, Sulit 6055 (BO, L, SING);
Mt Cansayao, Catarman, Sulit 14455 (BO, L).
PAPUA NEW GUINEA. Buna Hinterland, 7
miles NW of Embi Lakes, Smith NGF 1272
(BRI, CANB, L). Central: Brown River forest
station, Eddowes & Maru NGF 13112 (A, BO,
BRI, CANB, L, NSW, SING); near Maipa
airstrip, Kairuku subprov., Saunders 1108 (L).
Gulf: near Putei, junction of Tauri & Kapau
rivers, Craven & Schodde 964 (A, BRI, CANB,
L). Madang: Brahman, Bundi subdist, Henty
NGF 49267 (BO, CANB), NGF 49267 (A, BRI,
E, L, M, NSW, SING); Gogol River, Katik NGF
46662 (A, BRI, L, NSW, SING); Utu village,
Eddowes & Maru 13057 (A, BRI, E, NSW,
SING, US); Ramu Valley, ca. 5 miles SE Faita
airstrip, Saunders 484 (BRI, CANB, L), 505
(BRI, L); Utu village, near Madang, Eddowes &
Maru NGF 13057 (BO, CANB, L). Milne Bay:
E of Lake Lavu, Fergusson Island, Croft et al
LAE 68795 (BRI, CANB, L), LAE 68795 (A, E,
M, SING); near Ailuluai village, Esa’ala sub-
dist, Lelean & Katik NGF 49910 (A, BRI, E, L,
US). Morobe: Bulolo, Havel & Kairo NGF
17009 (BO, BRI, CANB, CBG, SING, US);
Bulolo, Streiman & Kairo NGF 26050 A (A,
BO, BRI, CANB, L, M, NSW, SING, US);
Oomsis near Lae, White NGF 10479 (A, BO,
BRI, CANB, L, SING). Northern: near Koreaf
village, Tufi subdist., Hoogland 4840 (BO, BRI,
CANB, L, MEL, NSW). Admiralty Islands,
Manus Island, Unknown NGF 543 (L). SINGA-
PORE: Bukit Timah, Ridley s.n. (SING).
Wrightia laevis is an extremely widespread
and variable species. Ngan (1965) recognized
three subspecies across this range on the basis of
corona and inflorescence pubescence charac-
ters, for which I have been unable to find clear
demarcations. I am not, therefore, recognizing
171
the subspecies. However, the specimens from
New Guinea in fruit show a shorter and wider
fruit type than is found in the other parts of the
range, and the trees are reported as much taller.
In flower they are unremarkable except for the
extremely long and fine corona lobes of the for-
mer subsp. novoguineensis. This variation does
not, however, correspond to the limits of any of
the former subspecies and is not the basis for
recognizing any formal ranks.
5. Wrightia novobritannica (Ngan) D. J.
Middleton, stat. nov.
Basionym: Wrightia pubescens subsp. novobri-
tannica Ngan, Ann. Miss. Bot. Gard. 52:
153. 1965. TYPE: PAPUA NEW
GUINEA. New Britain, Nodup,
Waterhouse 270 (Holotype: NY; Isotype:
A, F n.v., US).
Tree. Branchlets puberulent, glabrescent with
age. Leaves: petiole 4–6 mm long, puberulent;
blade elliptic to oblong, 12.8–19.0 × 3.6–6.2
cm, 3.3–3.7 times as long as wide, apex acumi-
nate, base cuneate, very sparsely puberulent all
over above, at least when young, puberulent
beneath, more densely so on midrib and vena-
tion, 13–15 pairs of secondary veins, tertiary
venation reticulate. Inflorescence terminal,
many-flowered, 2.5–4.5 cm long; peduncle
0.6–0.9 cm long, densely puberulent; pedicels
6–9 mm long, densely puberulent. Sepals
ovate, 1.8–2.2 × 1.6–1.8 mm, 1.0–1.4 times as
long as wide, apex obtuse to rounded, densely
puberulent, ciliate. Corolla brick red; tube ca.
3.5 mm long, ca. 1.6–1.9 times as long as
sepals, ca. 0.3 times as long as lobes, glabrous
or with a few hairs at top of tube outside,
glabrous inside; lobes ca.10.5 × 4.7 mm, 2.2
times as long as wide, elliptic, apex rounded,
minutely pubescent outside and inside; ante-
petalous corona adnate to the corolla lobe for
half of length, free at the margins, ca. 3.5 mm
long, apex entire, glabrous inside, ca. 2.3 times
as long as alternipetalous lobes; alternipetalous
corona lobes simple, ca. 1.5 mm long; alternat-
ing corona lobes lacking. Stamens inserted at
ca. 0.8 of tube length from corolla base,
strongly exserted from tube; filament ca. 1 mm
long; anthers 6.5 × 1.0 mm, pubescent outside.
Gynoecium of two connate carpels, ovary ca. 2
mm high, glabrous; style and style head ca. 5
mm long. Fruit of connate follicles, splitting
into two at dehiscence, ca. 17.5 cm long, 2.4
172 HARVARD PAPERS IN BOTANY
cm wide, sparsely puberulent, lenticellate.
Seeds 13.3–14.8 × 1.5–2.3 mm; coma 3.4–3.8
cm long.
Distribution: only from from the type col-
lection from New Britain, Papua New Guinea.
Habitat: not known.
6. Wrightia palawanensis D. J. Middleton, sp.
nov. TYPE: PHILIPPINES. Palawan: Irawan
Valley head on the lower slopes of Mt.
Beaufort, riverine forest, 150 m altitude, 29
March 1984: Podzorski SMHI 698 (Holotype:
A; Isotypes: BO, BRI, KEP, L). Fig. 5.
Frutex. Folia elliptica, apice acuminata, basi
cuneata, 2.6–11.1 cm longa, 1.0–3.3 cm lata.
Inflorescentia terminalis, 2–5-flora, 1–4 cm
longa, glabra. Calyx lobis 1.9–2.1 mm longis,
2.4–2.9 mm latis. Corolla tubus 3.8–4.0 mm
longus; lobi 8.5–8.9 mm longi, 7.0–7.7 mm lati,
alabastro sinistrosum obtegentes.
Shrub or small tree to 3 m tall. Branchlets
glabrous, sparsely lenticellate. Leaves: petiole
5–7 mm long, glabrous; blade elliptic, 2.6–11.1
× 1.0–3.3 cm, 2.4–4.3 times as long as wide,
apex acuminate, base cuneate, glabrous above
and beneath, 10–17 pairs of secondary veins,
tertiary venation reticulate. Inflorescence termi-
nal, 2- to 5-flowered, 1–4 cm long; peduncle
0.2–1.4 cm long, glabrous; pedicels 5.5–7.0 mm
long, glabrous. Sepals ovate, 1.9–2.1 × 2.4–2.9
mm, 0.7–0.8 times as long as wide, apex obtuse
to rounded, glabrous, ciliate. Corolla salmon
pink or yellow, infundibuliform; tube 3.8–4.0
mm long, widening at stamen insertion, 1.8–2.1
times as long as sepals, 0.4–0.5 times as long as
lobes, sparsely and minutely puberulent at top
of tube outside, glabrous inside; lobes 8.5–8.9 ×
7.0–7.7 mm, 1.1–1.3 times as long as wide,
elliptic, apex rounded, minutely puberulent out-
side and inside; antepetalous corona lobes
adnate to the corolla lobe for entire length but
free at margins, 4.3–4.5 mm long, apex suben-
tire to weakly crenate, glabrous inside, ca. 6
times as long as alternipetalous lobes;
alternipetalous corona lobes free but often so
small as to be difficult to see, simple, 0.7–0.8
mm long; alternating corona lobes lacking.
KEY TO SUBSPECIES OF WRIGHTIA PUBESCENS
1a. Corolla tube 1.2–1.5 times as long as sepals; sepal apex obtuse to rounded, margins and apex often slightly
reflexed; antepetalous corona lobes glabrous inside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7a. ssp. pubescens
1b. Corolla tube 2.0–3.9 times as long as sepals; sepal apex acute to obtuse, margins not reflexed; antepetalous
corona lobes mostly slightly pubescent inside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7b. ssp. lanitii
Vol. 10, No. 2
Stamens inserted at ca. 0.7 of tube length from
corolla base, exserted from tube; filament ca. 1
mm long; anthers 5.0–7.1 × 1.1–1.7 mm, sagit-
tate, adnate to the style head, pubescent on the
outside only at the very tip. Gynoecium of 2
free carpels united into a common style, ovaries
1.7–2.2 mm high, glabrous; style and style head
3.0–5.5 mm long. Fruit unknown (see notes).
Distribution: Philippines (Palawan).
Habitat: reported from riverine and mixed
forest and from ultramafic soils at 100–200 m
altitude.
Etymology: the specific epithet refers to the
distribution of this species in Palawan.
Additional collections studied: PHILIP-
PINES. PALAWAN: Bataraza Munic., Bgy Rio
Tuba, Belanjao Range, Gamayon Settlement,
Soejarto et al 8402 (L); Brookes Point munici-
pality, ca. 15 miles N, Infanta mining area,
Podzorski SMHI 2156 (A, L).
This new species would appear to have
affinities to Wrightia hanleyi Elmer, but it has
larger flowers and the stamens are inserted
higher in the corolla tube. It differs from
Wrightia candollei Vidal in the free carpels and
the lack of pubescence on the outside of the
anthers. None of the specimens report the pres-
ence of white latex, but it would be surprising
if this were not present. I have not seen any
fruit, but they are reported on one specimen to
be paired.
7. Wrightia pubescens R. Br., Prod. 467. 1810;
R. Br., Asclep. 64. 1810; Miq., Fl. Ned. Ind. 2:
432. 1857; Pichon, Not. Syst. 14: 83. 1951;
Ngan, Ann. Miss. Bot. Gard. 52: 150. 1965;
Backer & Bakh.f., Fl. Java 2: 241. 1965; Tsiang
& P. T. Li, Fl. Reip. Pop. Sin. 63: 123. 1977;
Lý, Fedd. Rep. 97: 623. 1986; Li et al., Fl.
China 16: 174. 1995; Ba et al., PROSEA 5.2:
513. 1995; Forster, Fl. Australia 28: 191. 1996;
Middleton, Fl. Thailand 7: 86. 1999; Kessler et
al., Blumea suppl. 14: 15. 2002. TYPE:
AUSTRALIA. North Coast (= Northern
Territories), Brown Iter Australiense 2861
(Lectotype: BM).
2005 MIDDLETON, A REVISION OF WRIGHTIA 173

FIGURE 5. Wrightia palawanensis D.J.Middleton (Podzorski SMHI 698, holotype, A).

174 HARVARD PAPERS IN BOTANY
7a. Wrightia pubescens ssp. pubescens. Fig. 6.
Synonyms: Wrightia calycina A. DC., Prod. 8:
406. 1844; Back. & Bakh. f., Fl. Java 2:
242. 1965. TYPE: TIMOR. Unknown
collector in Museum de Paris 1821
(Holotype: G-DC).
Wrightia spanogheana Miq., Fl. Ned.
Ind. 2: 434. 1857. Nerium macrocarpum
Span., Linnaea 15: 325. 1841, nom. nud.
TYPE: TIMOR. Spanoghe s.n.
(Lectotype: L [herb. no. 898,112–487],
designated here; Isotype: L).
Wrightia multiflora Zipp. ex Span.,
Linnaea 15: 325. 1841, nom. nud. Based
on TIMOR. Zippelius s.n. (L [herb. no.
898,112–449]).
Usage synonym: Wrightia tinctoria auct. non
R. Br.: Spanoghe, Linnaea 15: 325. 1841.
Branchlets glabrous to sparsely puberulent.
Leaves: petioles 3–6 mm long, sparsely pubes-
cent; blade elliptic, 3.8–11.8 × 1.6–5.6 cm,
1.8–3.8 times as long as wide, apex acuminate,
base cuneate, 8–11 pairs of secondary veins,
tertiary venation reticulate, sparsely puberulent
on midrib to sparsely pubescent all over above,
sparsely puberulent on midrib and secondary
veins to densely rusty pubescent all over
beneath. Inflorescence terminal, 2.5–6.0 cm
long; peduncle 0.2–0.8 cm long, sparsely to
densely pubescent; pedicels 4.3–9.5 mm long,
sparsely to densely pubescent. Sepals ovate,
2.8–5.0 × 2.2–3.0 mm, 1.2–1.8 times as long as
wide, apex obtuse to rounded and slightly
reflexed or not, margins slightly reflexed,
sparsely to densely pubescent, colleters large
and wide in sinuses. Corolla tube 3.6–6.0 mm
long, 1.2–1.5 times as long as sepals, 0.5–0.7
times as long as lobes, minutely pubescent at
top of tube outside, glabrous to sparsely
puberulent inside; lobes 6.7–11.0 × 2.8–4.7
mm, 1.8–2.8 times as long as wide, elliptic,
apex obtuse; antepetalous corona lobes 3.5–5.2
mm long, denate at apex, adnate to corolla lobe
for most of length, glabrous inside, 1.3–2.6
times as long as alternipetalous lobes;
alternipetalous corona lobes 1.5–3.0 mm long,
narrow, bifid; alternating lobes lacking.
Stamens inserted at 0.7–0.9 of tube length from
corolla base, strongly exserted from tube; fila-
ment 1 mm long; anthers 5.6–6.6 × 1.3–1.7
mm, densely pubescent outside. Gynoecium of
two connate carpels, ovary 1.3–2.0 mm high,
glabrous; style and style head 6.7–8.2 mm
Vol. 10, No. 2
long. Fruit of connate follicles, splitting into
two at dehiscence, 6.6–29.5 cm long, 1.2–1.6
cm wide (see note), glabrous, lenticellate or
not. Seeds 12.3–17 × 1.3–2.3 mm; coma
3.2–4.7 cm long.
Distribution: Australia (Northern
Territories), Eastern Java, Lesser Sunda
Islands, Moluccas, New Guinea.
Habitat: in dry forests.
Additional collections studied: EAST
TIMOR. Baucau, van Steenis 18029 (BO,
CANB, L). INDONESIA. BALI: Kuta, van
Balgooy 5691 (L); Mt Prapat Agung, NW Bali,
Kostermans et al KKSS 10 (A, BO, KEP, L, P);
Rapakagoeng, Beeking 45 (BO, L). JAVA: sine
loc.: de Vriese & Teysmann s.n. (L),
Leschenault 309 (P); Bangil, Backer 7571 (L);
Besuki, Backer 9529 (BO, L); Kupang near
Surabaya, Backer 26577 (BO, L); Res.
BanjoewangsAlas Rempis near Gentery,
Koorders 38905 ß (BO, L); Res. Besoeki,
Afd.Djember, Poeger, Koorders 199 ß (BO, L).
MALUKU: Sondo–sondo, Taylor 2731 B
(BO). Tidore, Teysmann HB 5181 (BO), de
Vriese s.n. (L). Kep. Tanimber: P.Jamdena,
Borssum Waalkes 3122 (BO, L); Olilit near
Saumlaki, Pleyte 27 (A, L); Leaut, Pereira s.n.
(SING). NUSA TENGGARA: Alor: Jaag 1084
(A, L, Z); Moroe-Gendok, Jaag 598 (Z).
Flores: Sine loc.: Verheijen 1005 (L), 2296 (L),
Wiriadinata s.n. (BO); (W), N part near Reo,
Kostermans & Wirawan 940 a (BO, L); (W), S
part, Mt Ndeki, Kostermans & Wirawan 185
(BO, L); P. Komodo, Verheijen 4902
(L);Ruteng, Schmutz 1022 (BO). Lombok:
Ampenan, Sun Hong-Fan Herb. 9064 (BO);
Sadjang, Elbert 742 (BO, L). Sumba, Jaag 21 a
(L, Z), Verheijen 4184 (L). Sumbawa: (W), east
of Sumbawa Besar, Kostermans 18001 A (A,
BO, CANB, L); Bima, Elbert 3832 (A, BO),
3898 (BO, L, SING), Warburg 17189 (E);
Kologebirge, Elbert 3650 (L). Timor: Sine loc.,
Riedlé s.n. (P), Teysmann HB 5121 (BO), 8754
(BO); Camplong, 46 km E of Kupang, Kuswata
& Kartawinata 1750 (L); Kp. Obe Naik,
Bloembergen 3346 (BO, L, SING); Kupang,
Teysmann 8744 (BO, L), Yoshida AN 62183
(BO). Wetar: Tarra, Bloembergen 3743 (BO, L,
NY, P, SING). SULAWESI: Kendari,
Batusanga, Neth. Ind. For. Serv. bb 24979 (L);
Lapakanrae, Noerkas 251 (BO, L); Muna,
Wasalangka, Neth. Ind. For. Serv. bb 21611 (L);
P. Saleyer, Teysmann 13602 (BO, L); Pulau
2005 MIDDLETON, A REVISION OF WRIGHTIA 175

FIGURE 6. Wrightia pubescens subsp. pubescens (Pleyte 27, A).

176 HARVARD PAPERS IN BOTANY
Lembeh, Alston 16098 (BO, L). WEST
PAPUA: near Achte Bivak, Merauke,
Branderhorst 284 (BO, L).
There are a number of specimens of this sub-
species where the fruit is very short and rather
misshapen. The dimensions are not included in
the description above as I suspect insect dam-
age in the formation of the fruit.
7b. Wrightia pubescens ssp. lanitii (Blanco)
Ngan, Ann. Miss. Bot. Gard. 52 (1965) 153;
Whitmore, Tree Fl. Mal 2 (1973) 23; Turner,
Gard. Bull. Sing. 47 (1997) 131. Fig. 7.
Basionym: Anasser lanitii Blanco, Fl. Filip.
(1837) 112. Wrightia lanitii (Blanco)
Merr., Gov. Lab. Publ. 27 (1905) 59.
TYPE: PHILIPPINES, Laguna, Makiling
National Park, Sulit PNH 22879
(Neotype: A, designated here; Isotype: L).
Synonyms: Wrightia javanica A.DC. in DC.,
Prod. 8: 405. 1844; King & Gamble, J.
As. Soc. Beng. 74, 2: 464. 1907; Ridley,
Fl. Mal. Pen. 2: 353. 1923; Kerr, Fl. Siam.
En. 2: 457. 1939; Back. & Bakh.f., Fl.
Java 2: 241. 1965. TYPE: INDONESIA.
Java: Kollmann s.n. (Holotype: G; scrap
from holotype in G-DC).
Wrightia ovata A. DC., Prod. 8: 405.
1844. TYPE: PHILIPPINES. Luzon:
Manila, Cuming 1279 (Lectotype: G,
designated here; Isotype: P).
Wrightia tomentosa var. cochinchinensis
Pierre ex Pitard in Lecomte, Fl. Gén.
Indo-Chine 3: 1186. 1933. TYPE:
VIETNAM. sine loc., Pierre 1047
(Lectotype: A; Isotype: BO, MICH,
MO). There is a possibility that Pierre
1047 and 1147 have become confused
and that they are in fact the same collec-
tion. There are additional specimens
labelled 1147 in NY and P.
Shrub or tree to 15 m tall, to 30 cm dbh, often
deciduous; wood pale. Branchlets sparsely
pubescent to tomentose, becoming glabrous;
sparsely lenticellate. Leaves: petiole 2–11 mm
long; blade ovate, elliptic or obovate, papery to
subcoriaceous, 1.5–15.0 × 1.0–7.5 cm, 1.2–4.0
times as long as wide, apex acuminate or apic-
ulate, base cuneate to rounded, densely tomen-
tose or puberulent above and beneath,
sometimes glabrate or only puberulent on vena-
tion, 7–15 pairs of secondary veins, tertiary
venation reticulate. Inflorescence terminal,
Vol. 10, No. 2
3.7–7.5 cm long; tomentose or puberulent;
peduncle 0.3–1.7 cm long, densely pubescent;
pedicels 4.3–11.0 mm long, densely pubescent.
Sepals ovate, 1–4 × 1.2–2.7 mm, 0.8–1.9 times
as long as wide, apex obtuse to acute, tomen-
tose or puberulent, colleters large and wide in
the sinuses. Corolla white and, sometimes,
pinkish; subrotate; tube 3.2–8.1 mm long,
2.0–3.9 times as long as sepals, 0.4–0.5 times
as long as lobes, minutely puberulent at top of
tube outside, glabrous inside; lobes 8.5–27.2 ×
4.8–9.6 mm, 1.6–4.0 times as long as wide,
oblong to slightly obovate, apex obtuse or
rounded, minutely puberulent outside and
inside; antepetalous corona 4.2–9.0 mm long,
crenate at apex, adnate to corolla lobes except
at apex and margins, sparsely pubescent inside,
0.9–1.4 times as long as alternipetalous lobes;
alternipetalous corona 2.5–7.5 mm long, nar-
row, bifid at apex, pubescent inside; alternating
lobes lacking. Stamens inserted at 0.7–0.9 of
tube length from corolla base, strongly exserted
from tube; filaments 0.7–2.0 mm long; anthers
5.9–8.5 × 1.3–2.1 mm, pubescent within and
without. Gynoecium of two connate carpels,
ovary 1.5–2.7 mm long, glabrous, carpels
medially connate; style and style head 8.0–10.5
mm long. Fruit of connate follicles, splitting
into two at dehiscence, 9.3–38.0 cm long,
0.9–2.2 cm wide, minutely puberulent or,
rarely, glabrous, minutely lenticellate or not.
Seeds linear; 10.7–15.7 × 1.0–2.6 mm; coma
1.2–4.5 cm long.
Distribution: China, Thailand, Cambodia,
Laos, Vietnam, Peninsular Malaysia, Sumatra,
western Java, Philippines, Sulawesi.
Habitat: in a wide variety of forest habitats.
Additional collections studied: INDONE-
SIA. JAVA: sine loc. Blume 1521 (L), Blume
1527 (L), Unknown 179 (L), Zippelius s.n. (L),
Zollinger 598 (A, G, L, P). Kedimdjali,
Koorders 24516 ß (BO); NW Moeriah, Docters
v.Leeuwen 997 (BO, L); Bantam, Afd.Menes,
Dist.Batoehideung, Koorders 196 ß (BO, L);
Res.Batavia, Backer 35432 (BO, L), van
Steenis 3055 (BO); Res. Batavia, G.Batoe,
Bakh.v.d.Brink 3131 (BO); Tandjoeng, Backer
35430 (BO, L); Res.Batavia, Tjimpaka poktih,
Backer 35431 (BO); Res.Batavia, Tjitajam,
Bakh.v.d.Brink 1662 (BO); Res.Batavia,
Wanajasa, Bakh.v.d.Brink 4851 (BO); Res.
Charibon, Kalidjohopekik, Koorders 36598 ß
(BO, L); Res. Krawang, Purwakala near
Krawang, Koorders 13799 ß (BO); Res.
2005 MIDDLETON, A REVISION OF WRIGHTIA 177

FIGURE 7. Wrightia pubescens subsp. lanitii (Blanco) Ngan (Leeuwenberg et al. 14567, A).
178 HARVARD PAPERS IN BOTANY
Krawang, Tandjoengpoera, Koorders 13798 ß
(BO, L); Res. Pekalongan, N of Soebah,
Hofmann 6539 (BO, GH); Res. Pekalongan,
near Soebah, Koorders 22549 ß (BO, L), 27294
ß (BO, L), 37066 ß (BO), 197 ß (BO, P), 201 ß
(BO), 13481 ß (BO, L), 13800 ß (BO, L), 13801
ß (BO, L), 13802 ß (BO, L), 14229 ß (BO, L),
14232 ß (BO), 14244 ß (BO, L); Res. Preanger,
Afd.Soemedang, Tomo, Koorders 195 (BO, L);
Res.Preanger, Afd.Soekadoemi, Koorders
34247 ß (BO); Res. Preanger, Afd.Soekaboemi,
Palaboeanratoe, Koorders 194 ß (BO, L), 200 ß
(BO, L), 12232 ß (BO, L); Res. Rambang,
Sedan, Ngandang, Koorders 36428 ß (BO);
Res. Semarang, Karangassem, Koorders 198 ß
(BO, L), 28200 ß (BO), 38874 ß (BO, L); Res.
Semarang, Kedoengdjati, Koorders 272 ß (BO),
24516 ß (BO), 33704 ß (BO, L). Carita, 10 km
N of Labuhan, Leeuwenberg 13217 (L);
Cempaka putih, Backer s.n. (L); Simpang,
Bakh.v.d.Brink 4892 (L). SULAWESI: Bikerve
Balang, Warburg 16325 (A); central part,
Wayingan near Palu, Meijer 10226 (BO, L,
MO); Manado, Bralema, Bilatto, Neth.Ind.For.
Serv. bb 15045 (L); North central, near Palu, 7
km to Donggala, Meijer 9273 (L, MO);
Sulawesi Tengah: W of Toboli, along road to
Palu, Leeuwenberg et al 14567 (A); Sulawesi
Tengah: Wera Waterfall near Parampadene, 25
km SW of Palu, van Balgooy 3622 (A, BO, L).
SUMATRA: sine loc., Unknown 230 (L).
MALAYSIA. Jongkah, Curtis 3054 (SING).
Kedah: sine loc., Meh 9012 (KEP), Ridley
14950 (SING), Ridley 14951 (SING); Jitra,
Burkill 13349 (SING); Langkawi: Pulau Tuba,
Zainudin 5231 (L). Penang: Mainland, Tasoek
Glugor, Curtis 3738 (SING). Perak: Lake
Gardens, Taiping, Corner s.n. (SING). Perlis:
Chupeny, Ridley 14952 (SING). Terengganu:
Jalan Hubor, Kuala Terengganu, Sinclair 39986
(BO, L, SING); Jalan Kubor, Kuala Trengganu,
Sinclair [SING39986] 7718 (E). PHILIP-
PINES. sine loc., Unknown 460 (L). PANAY:
Antique: Brgy. Simirara Island, Caluya, Romero
et al PPI 29509 (GH, PNH). BUSUANGA:
Marche 320 (P). CULION: Elmer 676 (GH,
SING); Herre 1049 (A). LUZON: Manila,
Loher 3999 (M). Bataan: Lamao River, Mt
Mariveles, Borden 770 (BO, P, NSW, SING);
Lamao River, Mt Mariveles, Borden 3067 (BO,
SING); Mt. Natib, Bgy. Banawang, Fuentes &
de la Rosa PPI 38807 (L); Tawawa, slope of
Vol. 10, No. 2
Mt. Natib, Reynoso et al PPI 17222 (GH);
Alambra 23206 (A, SING); Alvarez 12939 (BO,
L); Gangan 22701 (A); Pascual 23062 (L);
Ranario 27780 (SING); Sulit 25679 (NSW);
Topacio 20009 (NSW). Batangas: Mt Lobo,
Sulit 15715 (A, L, PNH, SING), 15746 (A).
Bosoboso: Morong, Loher 4000 (M). Bulacan:
Mt. Biak na Bato, San Miquel, Garcia et al PPI
15071 (GH, L); Leuterio 24730 (BO, L, NSW).
Cagayan: near Penablanca, Adduru 197 (A, P);
Alvarez 18541 (NSW); Bernardo 15138 (L, P).
Cavite: Ramos & Deroy 22555 (GH, SING).
Ilocos Norte: Adduru 22999 (A). Isabela:
Barros 26288 (A); San Mariano, Ramos &
Edaño 47008 (BO), Siriban 30396 (NSW,
SING); Sierra Madre Mountains, San Mariano,
Gutierrez 78044 (L). Laguna: Los Banos, Elmer
8069 (E); Mt. Makiling, Sulit 6875 (A),
Whitford 19784 (L). Masbate: Bgy. Matipuron,
Milagros, Barbon et al PPI 12598 (GH). Nueva
Ecija: Curran 8448 (NSW), Tungol 24142 (A,
GH). Nueva Vizcaya: Dupax, McGregor 11298
(L), 11300 (L). Pangasinan: Mayor 24577
(NSW). Rizal: Ahern’s collector 90 (A), 2960
(BO, P, SING); Franco 17856 (GH); Merrill
sp.Bl. 562 (A, BO, GH, L, MO, P); Ramos 2042
(BO, BRI, L, P, NSW, SING); Robinson 6738
(BRI, NSW). Zambales: Mt. Pinatubo, Balintay
5626 (A). MARINDUQUE: So. Matalim, Bgy.
Dampulan, Romero & Chavez PPI 29153 (GH,
L). MINDANAO: Cotabato: Buayan, Ramos &
Edano 85136 (A). Zamboanga: Zamboanga,
Ramos & Edano 37465 (A). Zamboanga del
Sur: Bolong, Reynoso et al PPI 4486 (L, PNH).
MINDORO: Pandarucan, Elmer 943 (GH),
Merrill 52932 (NSW). Occidental: Paluan,
Ramos 39677 (A). Oriental: East of Yagaw,
Conklin 37568 (L, SING); Puerto Galera,
Santos 5259 (L). NEGROS: Oriental:
Dumaguete (Cuernos Mts), Elmer 10291 (A,
BO, E, L, NSW). PALAWAN: Langen
(Malapakan) Isl., Podzorski SMHI 840 (A, BO,
KEP, L); Roxas, Langugan Zigzag, 85 km from
Puerto Princesa, Soejarto & Reynoso 6320 (A,
L, SING); Taytay, Merrill 9200 (BO, GH, L, P,
SING). PANAY: Mt. Baloy, Musat,
Varadarajan et al 1537 (A, L, MO). SINGA-
PORE: Chinese Temple, Telok Belanga Drive,
Ibrahim AI 66 (SING).
Wrightia pubescens ssp. lanitii would appear
to be as different from ssp. pubescens as ssp.
candollei, which has been resurrected as a
2005 MIDDLETON, A REVISION OF WRIGHTIA
species distinct from Wrightia pubescens.
Wrightia pubescens ssp. lanitii and ssp. pubes-
cens differ in the size and shape of the sepals
and, for the most part, the pubescence of the
antepetalous corona lobes. However, there are
individuals that have some pubescence on the
inside of the corona lobes, and specimens from
Sulawesi have the corona type and sepal shape
of ssp. pubescens but the sepal size of ssp. lan-
itii. Because of these intermediates and the
greater similarity in corolla shape and size
between these two subspecies compared to ssp.
candollei, they are being maintained as sub-
species, whereas Wrightia candollei, which is
also sympatric with W. pubescens ssp. lanitii,
has been given specific status.
8. Wrightia religiosa (Teijsm. & Binn.) Benth.
ex Kurz, J. As. Soc. Beng. 46, 2: 258. 1877;
Kurz, Fl. Burm. 2: 194. 1877; Hook. f., Fl. Brit.
Ind. 3: 653. 1882; King & Gamble, J. As. Soc.
Beng. 74, 2: 464. 1907; Ridley, Fl. Mal. Pen. 2:
353. 1923; Pitard in Lecomte, Fl. Gén. Indo-
Chine 3: 1183. 1933; Kerr, Fl. Siam. En. 2: 458.
1939; Pichon, Not. Syst. 14: 85. 1951; Ngan,
Ann. Miss. Bot. Gard. 52: 159. 1965; Backer &
Bakh. f., Fl. Java 2: 241. 1965; Whitmore, Tree
Fl. Mal. 2: 24. 1973; Lý, Feddes Repert. 97:
622. 1986; Li et al., Fl. China 16: 175. 1995;
Turner, Gard. Bull. Sing. 47: 131. 1997;
Middleton, Fl. Thailand 7: 88. 1999. Fig. 8.
Basionym: Echites religiosa Teijsm. & Binn.,
Tijdschr. Ned. Ind. 27: 34. 1864. - TYPE:
Teijsmann s.n. (untraced). NEOTYPE:
Collins 669 (Neotype: E, designated
here; isoneotype: ABD, K, TCD),
Thailand, Chon Buri, Sri Racha.
Shrub or small tree to 5 m tall. Branchlets
puberulent, soon becoming glabrous and
NOTE
Wrightia timorensis Miq., Fl. Ned. Ind. 2: 433.
1857. TYPE: TIMOR. Sine loc., Unknown s.n.
(Lectotype: L [898,112–484]; Isotype: U [frag-
ment]).
Wrightia timorensis is a synonym of Wrightia
tinctoria, a species otherwise known only from
India. Its appearance in Timor is a mystery and
is likely to be either a mistake on the label or
179
sparsely lenticellate. Leaves: petiole 1.5–4.0
mm long; blade papery, elliptic or oblong,
1.2–8.2 × 0.6–3.3 cm, 2.2–4.2 times as long as
wide, apex acuminate, rarely to rounded, base
cuneate, puberulent on midrib beneath and
sometimes on midrib above, or completely
glabrous, blade punctate beneath and some-
times above, 3–8 pairs of secondary veins,
ascending, often obscure, tertiary venation
obscure. Inflorescence 1.7–4.2 cm long;
glabrous or sparsely puberulent; pedicels 8–30
mm long, very delicate; flowers pendulous.
Sepals ovate or oblong, 1.0–2.5 × 0.7–1.0 mm,
1.9–2.4 times as long as wide, apex obtuse to
acute, glabrous, ciliate; colleters small and nar-
row. Corolla white; subrotate; sometimes dou-
ble in cultivated plants; tube 2.1–4.2 mm long;
lobes 4.8–9.8 × 1.9–2.7 mm, 2.1–2.6 times as
long as wide, obovate, apex rounded; puberu-
lent-papillose on lobes outside and inside,
glabrous on tube outside and inside; corona
absent. Stamens inserted at corolla mouth,
strongly exserted from tube; filaments 1.5–2.5
mm long; anthers 3.8–4.0 × 0.7–0.8 mm,
pubescent within and without. Gynoecium of 2
free carpels united into a common style, ovaries
0.8–1.1 mm high, glabrous, of two separate
carpels; style and style head 5.7–8.3 mm long.
Fruit of paired follicles; 8.5–17.5 cm long,
3.3–4.5 mm wide; glabrous; not lenticellate.
Seeds linear; 7.0–8.6 × 1.1–1.3 mm; coma
3.0–3.5 cm long.
Distribution: Thailand, Cambodia, possibly
Peninsular Malaysia. The natural distribution
of this species is now obscure because it is so
widely cultivated, but is probably Thailand and
maybe Cambodia. It may also be native in
Peninsular Malaysia, but is certainly widely
cultivated in many parts of Malesia.
else a cultivated plant transported from India
earlier. There are no notes on the label to indi-
cate whether it was a cultivated plant. Until
such time as further material is collected in
Timor that matches this specimen and its status
can be reassessed, I am not including this as a
Malesian species to be discussed further.
180 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

FIGURE 8. Wrightia religiosa (Teijsm. & Binn.) Benth. ex Kurz (Forman 47, A).
2005 MIDDLETON, A REVISION OF WRIGHTIA
LITERATURE CITED
BAHADUR, K. N., AND S. S. R. BENNET. 1978. A new
species of Wrightia R. Br. (Apocynaceae) from
India. Indian Forester 104: 32–35.
BRUMMIT, R. K., AND C. E. POWELL. 1992. Authors of
Plant Names. Royal Botanic Garden, Kew.
ENDRESS, M.E., AND P. V. BRUYNS. 2000. A revised
classification of the Apocynaceae s.l. Botanical
Review 66(1): 1–56.
HOLMGREN, P. K., N. H. HOLMGREN, AND L.C.BARNETT.
1990. Index Herbariorum. IAPT, New York.
INDEX OF EXSICCATAE
The number in parentheses after each collection refers to the number given to each taxon above.
Adduru 197 (7b); Adduru 22999 (7b); Aet & Idjan
419 (4); Ahern’s collector 90 (7b), 1116 (1), 2960
(7b), 3155 (1); Alambra 23206 (7b); Alston 16098
(7a); Alvarez 12939 (7b), 18541 (7b); Ang FRI
23347 (4).
Backer 7571 (7a), 9529 (7a), 26577 (7a), 35430
(7b), 35430 (7b), 35431 (7b), 35432 (7b), s.n.
(7b); Bakhuizen v.d. Brink 1662 (7b), 3131 (7b),
4851 (7b), 4892 (7b); Balintay 5626 (7b); Barbon
et al. PPI 12598 (7b), 18700 (4), 22357 (1); Barros
26288 (7b); Bartlett 14074 (1), 14720 (1);
Beeking 45 (7a); Bernardo 15138 (7b);
Bloembergen 3346 (7a), 3743 (7a); Blume 1521
(7b), 1527 (7b); Borden 770 (7b), 3067 (7b);
Borssum Waalkes 3122 (7a); Branderhorst 144
(7a), 284 (7a); Brass 5570 (4), 13893 (4); Burkill
13349 (7b); Burkill & Haniff 15826 (4); Burtt &
Woods B 1729 (2).
Chan FRI 17654 (4); Chin & Mahmud 1289 (4);
Clemens 18171 (1); Conklin 37568 (7b);
Contreras 23404 (1); Corner s.n. (7b); Craven &
Schodde 964 (4); Croft et al. LAE 68795 (4), LAE
68795 (4); Cuming 1279 (7b), 1453 (1); Curran
4510 (1), 8442 (1), 8448 (7b); Curtis 188 (2), 2915
(2), 2943 (4), 3054 (7b), 3738 (7b), s.n. (2).
De Mesa 27538 (4); De Vriese & Teysmann s.n.
(7a); De Vriese s.n. (7a); De Wilde & de Wilde-
Duyfjes 12369 (4), 20199 (4); Docters v.Leeuwen
997 (7b).
Eddowes & Maru NGF 13057 (4), NGF 13112 (4);
Elbert 742 (7a), 3650 (7a), 3832 (7a), 3898 (7a);
Elmer 676 (7b), 943 (7b), 3049 (1), 6348 (1), 8069
(7b), 10291 (7b), 12873 (3), 15595 (4); Escritor
21296 (1).
Fenix 1183 (1); Forbes 2766 (4), 2802 (4); Franco
17856 (7b); Fuentes & de la Rosa PPI 38807 (7b).
Gaerlan et al. PPI 10603 (1), PPI 13448 (3);
Gammill 228 (1); Gangan 22701 (7b); Garcia et al.
PPI 15071 (7b), PPI 18505 (1); Gutierrez 78044
(7b).
Haniff 7086 (2); Havel & Kairo NGF 17009 (4);
Henderson 19599 (4); Henty NGF 49267 (4),
NGF 49267 (4); Herre 1049 (7b); Hofmann 6539
(7b); Hoogland 4840 (4); Hotta 25889 (4); Hume
8352 (4).
181
LÝ, T.
D. 1985. Zwei neue Arten der Familie Apocy-
naceae aus südwestlichen Gebieten Vietnams.
Feddes Repertorium 96: 173–178.
MIDDLETON, D. J., AND T. SANTISUK. 2001. A new
species of Wrightia (Apocynaceae: Apocynoideae)
from Thailand. Thai Forest Bulletin 29: 1–10.
NGAN, P. T. 1965. A revision of the genus Wrightia
(Apocynaceae). Annals of the Missouri Botanical
Garden 52: 114–175.
Ibrahim AI 66 (7b); Iwanggin BW 9136 (4).
Jaag 21a (7a), 598 (7a), 1084 (7a); Jumali 875 (4),
2866 (2), 4339 (4).
Katik NGF 46662 (4); Kiah S 151 (2), 35275 (2),
s.n. (2); King’s Collector 8614 (4); Kochummen
FRI 2973 (4), FRI 16446 (4), KEP 99607 (4),
99607 (4); Koorders 194 ß (7b), 195 (7b), 196 ß
(7b), 197 ß (7b), 198 ß (7b), 199 ß (7a), 200 ß (7b),
201 ß (7b), 272 ß (7b), 12232 ß (7b), 13481 ß (7b),
13798 ß (7b), 13799 ß (7b), 13800 ß (7b), 13801 ß
(7b), 13802 ß (7b), 14229 ß (7b), 14232 ß (7b),
14244 ß (7b), 22549 ß (7b), 24516 ß (7b), 24516 ß
(7b), 27294 ß (7b), 28200 ß (7b), 33704 ß (7b),
34247 ß (7b), 36428 ß (7b), 36598 ß (7b), 37066 ß
(7b), 38874 ß (7b), 38905 ß (7a); Koster BW 4349
(4), BW 4355 (4), BW 11046 (4), BW 11804 (4);
Kostermans 256 (4), 18001 A (7a); Kostermans et
al. KKSS 10 (7a), 100 (4); Kostermans &
Wirawan 185 (7a), 940a (7a); Kuswata &
Kartawinata 1750 (7a).
Laumonier TFB 4464 (4); Leeuwenberg 13217 (7b);
Leeuwenberg et al. 14567 (7b); Lelean & Katik
NGF 49910 (4); Leschenault 309 (7a); Leuterio
24730 (7b); Loh FRI 17384 (4); Loher 3999 (7b),
4000 (7b), 6519 (1), 6530 (1), 6542 (1); Lopez &
Reyes 27331 (1).
Madulid & Majaducon PPI 36005 (1); Majaducon &
Fernando 8455 (1); Marche 320 (7b); Maxwell
s.n. (4); Mayor 24577 (7b); McGregor 11298 (7b),
11300 (7b), 18579 (1), 41431 (1); Meh 9012 (7b);
Meijer 9273 (7b), 10226 (7b); Mendoza 42092
(4), 91195 (3); Merrill sp. Bl. 562 (7b), 3082 (1),
9200 (7b), 9373 (3), 9426 (1), 52932 (7b); Moll
BW 11560 (4); Moton KEP 94083 (4).
Neth. Ind. For. Serv. bb 15045 (7b), bb 21611 (7a),
bb 24979 (7a), bb 30391 (4), bb 30417 (4), bb
30447 (4), bb 30530 (4); Ng FRI 5392 (4);
Noerkas 251 (7a); Nur 11728 (4).
O’Sullivan s.n. (2).
Pagi KEP 98711 (4); Pascual 23062 (7b); Pereira s.n.
(7a); Pleyte 27 (7a); Podzorski SMHI 698 (6),
SMHI 840 (7b), SMHI 2156 (6).
Rahmat Si Boeea 7533 (4); Ramos 2042 (7b), 8284
(1) 39677 (7b), 41607 (1), 42712 (1); Ramos &
Deroy 22555 (7b); Ramos & Edaño 37465 (7b),
182 HARVARD PAPERS IN BOTANY
47008 (7b), 85136 (7b); Ranario 27780 (7b);
Reynoso et al. PPI 4486 (7b), PPI 17222 (7b), PPI
24244 (4), PPI 24249 (3), PPI 24423 (4); Ridley
5521 (2), 7566 (4), 14605 (4), 14949 (4), 14950
(7b), 14951 (7b), 14952 (7b); Ridsdale 1080 (3),
1418 (1), SMHI 1620 (3); Riedlé s.n. (7a);
Robinson 6738 (7b), 6757 (1); Romero & Chavez
PPI 29153 (7b); Romero & Fuentes PPI 37562 (1);
Romero et al. PPI 29509 (7b).
Sakat KEP 116122 (4); Samsuri & Ahmad SA 691
(4); Santos 5259 (7b); Saunders 484 (4), 505 (4),
1108 (4); Saw FRI 36282 (4); Schmutz 1022 (7a);
Schram BW 10592 (4); Scortechini 77 b (4);
Selvaraj FRI 13006 (2); Sinclair [SING39023]
6627 (2), [SING39986] 7718 (7b), 39023 (2),
39986 (7b), SFN 40103 (4); Siriban 30396 (7b);
Smith NGF 1272 (4); Soejarto & Reynoso 6261
(3), 6320 (7b); Soejarto et al. 6456 (1), 6479 (3),
8402 (6), 8602 (3); Soepadmo & Students FSC
832 (4); Spanoghe s.n. (7a); Stefels BW 5105 (4);
Stone et al. 120 (2), PPI 262 (3); Streiman & Kairo
NGF 26050 A (4); Sulit 6055 (4), 6875 (7b), PNH
INDEX
Names in roman script are currently recognized species, those in bold are new taxa or combina-
tions, and those in italics are synonyms. The numbers in parentheses refer to the taxon numbers
above. (x) refers to taxa discussed elsewhere in the text.
Anasser laniti Blanco (7b)
Cameraria dubia Sims (2)
Echites religiosa Teijsm. & Binn. (8)
Nerium macrocarpum Span. (7a)
Scleranthera cambodiensis (Pierre ex Pitard)
Pichon (2)
Scleranthera dubia (Sims) Pichon (2)
Strophanthus jackianus Wall. ex G.Don (2)
Walidda antidysenterica (L.) Pichon (x)
Wrightia antidysenterica L. (x)
Wrightia balansae Pitard (4)
Wrightia calycina A.DC. (7a)
Wrightia cambodiensis Pierre ex Pitard (2)
Wrightia candollei Vidal (1)
Wrightia dubia var. membranifolia
King & Gamble (2)
Wrightia dubia (Sims) Spreng. (2)
Wrightia hainanensis Merr. (4)
Wrightia hainanensis var. chingii Tsiang (4)
Wrightia hainanensis var. variabilis Tsiang (4)
Wrightia hanleyi Elmer (3)
Wrightia javanica A.DC. (7b)
Wrightia kontumensis Lý (2)
Wrightia laevis Hook.f. (4)
Vol. 10, No. 2
7428 (1), 14455 (4), 15715 (7b), 15746 (7b), PNH
17044 (1), 22879 (7b), 25679 (7b); Sun Hong-Fan
Herb. 9064 (7a); Suppiah FRI 108885 (4);
Symington 51751 (4), 57059 (4), 57060 (4),
57061 (4).
Taylor 2731 B (7a); Teo & P. 548 (4); Teysmann HB
5121 (7a), HB 5181 (7a), 8744 (7a), 8754 (7a),
13602 (7a); Topacio 20009 (7b); Tungol 24142
(7b).
Van Balgooy 2436 (4), 2488 (4), 3622 (7b), 5691
(7a); Van Dijk 168 (4), 196 (4), 226 (4), 309 (4);
Van Steenis 3055 (7b), 18029 (7a); Varadarajan et
al. 1537 (7b); Verheijen 1005 (7a), 2296 (7a),
4184 (7a), 4902 (7a); Versteegh BW 3991 (4), BW
4628 (4), BW 4971 (4); Vink BW 11420 (4).
Warburg 16325 (7b), 17189 (7a); Waterhouse 270
(5); White NGF 10479 (4); Whitford 19784 (7b);
Whitmore FRI 4405 (4), FRI 8568 (4); Widjaja
1213 (4); Wiriadinata s.n. (7a).
Yeob 3205 (4); Yoshida AN 62183 (7a).
Zainudin 3326 (4), 5231 (7b); Zainudin et al. 4391
(2); Zippelius s.n. (7b); Zollinger 598 (7b).
Wrightia laevis subsp. millgar (F.M.Bailey) Ngan (4)
Wrightia laevis subsp. novoguineensis Ngan (4)
Wrightia lanitii (Blanco) Merr. (7b)
Wrightia macrocarpa Pitard (4)
Wrightia millgar F.M.Bailey (4)
Wrightia multiflora Zipp. ex Span. (7a)
Wrightia novobritannica (Ngan) D.J.Middleton (5)
Wrightia ovata A.DC. (7b)
Wrightia palawanensis D.J.Middleton (6)
Wrightia pubescens R.Br. (7)
Wrightia pubescens subsp. candollei (Vidal) Ngan (1)
Wrightia pubescens subsp. lanitii (Blanco) Ngan (7b)
Wrightia pubescens subsp. novobritannica Ngan (5)
Wrightia pubescens subsp. pubescens (7a)
Wrightia religiosa (Teijsm. & Binn.) Benth. ex
Kurz (8)
Wrightia rubriflora Pitard (2)
Wrightia sorsogonensis Elmer (4)
Wrightia spanogheana Miq. (7a)
Wrightia timorensis Miq. (x)
Wrightia tinctoria R.Br. (x)
Wrightia tinctoria var. laevis (Hook.f.) Pichon (4)
Wrightia tomentosa var. cochinchinensis Pierre ex
Pitard (7b)