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 Vol. 10, No. 2

to corolla except at apex, apex subentire to Soejarto et al 6456 (A, MO); Taytay Bay,
weakly crenate, glabrous inside, 3.8–5.0 times Apulit Island, Merrill 9426 (BO, BRI, L, P,
as long as alternipetalous lobes; alternipetalous SING); sine loc., Curran 4510 (L); sine loc.,
corona lobes simple or slightly bifid 0.7–1.7 Fenix 1183 (Z).
mm long; alternating corona lobes lacking. This species was included as a subspecies of
Stamens inserted at 0.7–0.8 of tube length from Wrightia pubescens by Ngan (1965), but it dif-
corolla base, exserted from tube; filaments fers substantially from that species in the shape
0.8–1.0 mm long; anthers 5.0–7.8 × 1.1–1.8 and form of the corolla and corona and in the
mm, pubescent on the outside or only at tips flower color. Therefore, it has been raised back
outside. Gynoecium of two connate carpels, to specific level.
ovary 1.5–2.6 mm high; style and style head
5–7 mm long. Fruit of a single follicle that 2. Wrightia dubia (Sims) Spreng., Syst. Veg. 1:
splits lengthwise at maturity as each of the two 638. 1825; King & Gamble, J. As. Soc. Beng.
component follicles dehisces, 20.5–22.0 cm 74, 2: 465. 1907; Ngan, Ann. Miss. Bot. Gard.
long, 1.4–2.2 cm wide, glabrous. Seeds not 52: 172. 1965; Whitmore, Tree Fl. Mal. 2: 24.
observed. 1973; Lý, Feddes Repert. 97: 622. 1986;
Distribution: Philippines. Turner, Gard. Bull. Sing. 47: 131. 1997;
Habitat: evergreen or mixed deciduous forest. Middleton, Fl. Thailand 7: 83. 1999. Fig. 2.
Additional material studied: PHILIP- Basionym: Cameraria dubia Sims in Curtis,
PINES. Sine loc.: Loher 6519 (M), 6530 (M), Bot. Mag. 39: t.1646. 1814. Scleranthera
6542 (M); Escritor 21296 (P); McGregor dubia (Sims) Pichon, Not. Syst. 14: 90.
18579 (P). BOHOL: Ramos 42712 (A). 1951. TYPE: Illustration in plate 1646 in
CAMOTES: Ramos 41607 (BO, L). CEBU: Curtis, Bot. Mag. 39 (1814).
Lopez & Reyes 27331 (BO, L, P). Synonyms: Strophanthus jackianus Wall. ex
GUIMARAS: Gammill 228 (NSW). LEYTE: G.Don, Gen. Syst. 4: 85. 1837; Hook. f.,
Hubasan, Baybay, Gaerlan et al PPI 10603 Fl. Ind. 3: 656. 1882; Ridley, Fl. Mal.
(GH). LUZON: Bataan: Alongapo, Bartlett Pen. 2: 355. 1923. TYPE: MALAYSIA.
14074 (A). Batangas: Bo. Biga, Lobo, Sulit Penang, Wallich 1643 (Lectotype: K;
PNH 7428 (A, PNH). Benguet: Twin Peaks, Isotype: K-W).
Elmer 6348 (NSW, P). Bulacan: Angat
Wrightia dubia var. membranifolia King
Watershed area, Norzagaray, Barbon et al PPI
& Gamble, J. As. Soc. Beng. 74, 2: 466.
22357 (PNH); near Kay Tianak and Ipo,
1907. TYPE: THAILAND, Phuket,
Bartlett 14720 (A). Cagayan: Brgy. San Juan,
Tongkah, February 1893, Curtis 2915
Sta. Praxedes, Garcia et al PPI 18505 (PNH).
(Lectotype: SING, designated here).
Nueva Ecija: Curran 8442 (SING).
Pangasinan: Alaminos, McGregor 41431 Wrightia cambodiensis Pierre ex Pitard
(KEP); Bolinao, Clemens 18171 (BO, C, S); in Lecomte, Fl. Gén. Indo-Chine 3: 1184.
sine loc., Ramos 8284 (L). Rizal: San Mateo, 1933; Kerr in Craib, Fl. Siam. En. 2: 456.
Ahern’s collector 1116 (BO, NSW); sine loc., 1939. Scleranthera cambodiensis (Pierre
Ahern’s collector 3155 (BO, P, SING); sine ex Pitard) Pichon, Not. Syst. 14: 89.
loc., Robinson 6757 (BRI, L). Zambales: Acoje 1951. TYPE: VIETNAM, Ninh Thuan,
Mine, Santa Cruz, Ridsdale 1418 (A, BO, KEP, Ca Na, Pierre 4402 p.p. (Lectotype: P;
L, MO); San Marcelino, Bgy. Sayasay, Mt. Isotype: A, P).
Mandakogdog, Majaducon & Fernando 8455 Wrightia rubriflora Pitard in Lecomte,
(L). – MASBATE: Elmer 3049 (P); Merrill Fl. Gén. Indo-Chine 3: 1185. 1933.
3082 (BO, L). MINDORO: occidental: Mt. TYPE: VIETNAM, Ninh Thuan, Ca Na,
Gonting, Lubang Island, Romero & Fuentes Poilane 5955 (Lectotype: P; Isotype:
PPI 37562 (A, PNH). Oriental: Mabaho Ridge, HM, NY, P).
Bo. Kabalwa, Mansalay, Sulit PNH 17044 (A, Wrightia kontumensis Lý, Feddes Repert.
PNH). NEGROS: Contreras 23404 (BO). 96: 173. 1985; Lý, Feddes Repert. 97:
Occidental: Danjugan Island, Bulata, Cauayan, 622. 1986. TYPE: Vietnam, Gialai-
Madulid & Majaducon PPI 36005 (L). Kontum, Loc P 2968 (Holotype: HNU;
PALAWAN: Puerto Princesa. Irawan River Isotype: HN).
Valley, Tatanarom, trail up Mt. Beaufort,
2005 MIDDLETON, A REVISION OF WRIGHTIA 165

FIGURE 2. Wrightia dubia (Sims) Spreng. (Sidek bin Kiah S.151, A).
166 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

Shrub to 2 m tall. Branchlets glabrous or Perlis: Kaki Bukit, Burtt & Woods B 1729 (E),
sparsely puberulent, becoming glabrous and Kiah 35275 (A, BO, L), Kiah s.n. (SING).
lenticellate. Leaves: petiole 2–11 mm long; Terengganu: Gunong Tebu, Besut, Selvaraj FRI
blade papery to subcoriaceous, elliptic, oblong 13006 (KEP, L). SINGAPORE: Botanic
or obovate, 3.6–26.5 × 1.8–9.8 cm, 2.1–5.2 Gardens, Jumali 2866 (SINU), Kiah S 151 (E).
times as long as wide, apex acuminate to sub- This is the most distinctive species of
caudate, base cuneate to rounded, puberulent Wrightia in Malesia with its pink flowers and
abaxially, only on midrib and secondary veins without the anthers exserted.
or, rarely, glabrous and then both surfaces papil-
late, 6–14 pairs of secondary veins, ascending, 3. Wrightia hanleyi Elmer, Leafl. Philipp. Bot.
tertiary venation reticulate. Inflorescence 4: 1465. 1912; Ngan, Ann. Miss. Bot. Gard. 52:
2.5–4.5 cm long; sparsely puberulent or 168. 1965. TYPE: PHILIPPINES, Palawan,
glabrous; pedicels 4.0–5.5 mm long, robust. Puerto Princesa, Mt. Pulgar, Elmer 12873
Flowers inodorous. Sepals ovate, 1.3–3.0 × (Lectotype: GH, designated here; Isotype: A,
1.3–2.0 mm, 1.1–1.3 times as long as wide, apex BISH, BO, E, K, L, NSW, NY, US, Z). Fig. 3.
acute to obtuse; sparsely puberulent or glabrous, Shrub or treelet to 4 m tall. Branchlets
ciliate; colleters wide and large. Corolla orange, glabrous, sparsely lenticellate with age.
pink or reddish; infundibuliform; tube 6.0–15.5 Leaves: petiole 3 mm long, glabrous; blade nar-
mm long, 3.8–4.7 times as long as sepals, rowly ovate to elliptic, 3–11 × 1.3–4.6 cm,
0.5–0.6 times as long as lobes, glabrous or, 1.6–3.8 times as long as wide, apex acuminate
rarely, pubescent-papillose at top of tube out- with a sharp tip, base cuneate to rounded,
side, glabrous inside; lobes 10–28 × 6–9 mm, glabrous above and beneath, 6–15 pairs of sec-
1.9–3.2 times as long as wide, ovate, apex ondary veins, tertiary venation reticulate.
acuminate to obtuse, pubescent-papillose on Inflorescence terminal, 0.8–2.0 cm long, few-
lobes outside and slightly so inside; antepetalous flowered; peduncle 0.1–1.0 cm long, glabrous;
pedicels 1.5–4.0 mm long, glabrous. Sepals
corona lobes adnate to the corolla, 1.5–3.7 mm
ovate, 1.0–1.5 × 0.9–1.1 mm, 1.1–1.4 times as
long, 1.7–2.5 times as long as alternipetalous
long as wide, apex acute to obtuse, glabrous,
lobes, glabrous; alternipetalous corona lobes
ciliate, colleters small in the sinuses. Corolla
0.6–2.2 mm long; alternating corona lobes lack-
greenish-yellow to red; tube with a lower nar-
ing. Stamens inserted at 0.3–0.4 of tube length
row part and a campanulate upper part, with a
from corolla base, completely included in the raised lip inside between the lower and upper
tube; subsessile; anthers 4.9–5.6 × 1.0–1.2 mm, tube, 1.8–2.8 mm long, 1.5–1.9 times as long as
pubescent outside. Gynoecium of 2 free carpels sepals, 0.6–0.7 times as long as lobes, minutely
united into a common style, ovaries 1.6–2.5 mm puberulent at top of tube outside, glabrous
long, glabrous; style and style head 3.0–4.7 mm inside; lobes 3.0–3.9 × 1.5–3.0 mm, 1.3–2.0
long. Fruit of paired follicles, 13–30 cm long, times as long as wide, apex rounded, minutely
3.5–6.0 mm wide, glabrous, not lenticellate. puberulent inside and outside; antepetalous
Seeds linear, 12.5–25.4 × 1.2–2.5 mm; coma corona 1.5–2.9 mm long, entire, adnate to
2.6–5.5 cm long. corolla on medial line but with sides free, apex
Distribution: Cambodia, Laos, Vietnam, subentire or slightly crenate, glabrous inside,
Thailand, Peninsular Malaysia, Singapore. 4.1–7.5 times as long as alternipetalous lobes;
Habitat: growing in wide variety of forest alternipetalous corona lobes apparently absent
habitats at lower altitudes. or small and simple, 0.2–0.7 mm long; alternat-
Additional collections studied: MALAYSIA: ing corona lobes lacking. Stamens subsessile,
Kedah: Sine loc., Ridley 5521 (SING); Pulau inserted at 0.3–0.5 of tube length from corolla
Langkawi, Telok Apau, Haniff 7086 (SING); base, exserted from tube; anthers 2.6–5 ×
Pulau Langkawi, Stone et al 120 (KLU); Pulau 0.4–1.0 mm, pubescent only at the top outside.
Langkawi, Pulau Singa Besar, Zainudin et al Gynoecium of 2 free carpels united into a com-
4391 (L). Penang: Curtis 188 (SING); Curtis mon style, ovaries 0.7–1.1 mm high, glabrous;
s.n. (SING); below Waterfall gardens, Sinclair style and style head 1.8–4.0 mm long. Fruit
39023 (E, SING); valley of stream below paired, linear, 6.6–18.0 cm long, 2.5–4.0 mm
waterfall gardens, Sinclair [SING39023] 6627 wide, glabrous. Seeds 10.0 × 0.6 mm; coma 21
(E); Butterworth, O’Sullivan s.n. (SING). mm long.
2005 MIDDLETON, A REVISION OF WRIGHTIA 167

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

168 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

Distribution: Philippines (Palawan). 2118 (Lectotype: P; Isotype: P).

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

lobes, glabrous or minutely puberulent at top of Teminaboean, Versteegh BW 4971 (L). SUMA-
tube outside, glabrous inside; lobes 6.1–14.0 × TRA. Bt. Gajabuih, Ulu Gadut, 15 km E of
2.6–5.2 mm, 1.6–2.8 times as long as wide, Padang City, Hotta 25889 (BO); Palembang,
elliptic to slightly obovate, apex rounded, Unknown 1078 (L); Tanjong Ning, Palembang,
minutely puberulent outside and inside; corona Forbes 2766 (L, SING); near Sigirik mountain
of antepetalous, alternipetalous, and alternate within Sebelah Nature Reserve, Laumonier
lobes; antepetalous lobes adnate to the corolla TFB 4464 (L); Gunung Leuser NP, Ketambe,
at base, fimbriate, 2.8–9.4 mm long, 0.9–1.8 valley of Lau Alas, ca. 35 km NW of Kutatjane,
times as long as alternipetalous lobes, glabrous; de Wilde & de Wilde-Duyfjes 12369 (BO, KEP,
alternipetalous lobes free, simple, deeply bifid, L, US); Gunung Leuser NP, Middle Alas River,
or sometimes compound, 2.4–6.5 mm long; 15 km N of Galombang, de Wilde & de Wilde-
alternating lobes 0.6–5.8 mm long, simple, Duyfjes 20199 (BO, L, US, Z); vicinity of
shorter than the other two whorls. Stamens Loemban, Asahan, Rahmat Si Boeea 7533 (S,
inserted at 0.8–0.9 of tube length from corolla SING). MALAYSIA. Pulau Pomgoh, Curtis
base, completely exserted; filaments 0.2–1.7 2943 (SING); Johor: Kluang, Labis FR, Moton
mm long; anthers 3.9–5.6 × 0.7–1.6 mm, KEP 94083 (KEP, L, SING), Kochummen FRI
pubescent outside, sometimes only sparsely so 16446 (A, KEP, L, SING); Gunong Besar, Ulu
at base. Gynoecium of 2 free carpels united into Sungei Segamat, Samsuri & Ahmad SA 691 (C,
a common style, ovaries 0.7–1.8 mm long, G, KEP); Near Ulu Sungei Redong, Jumali 875
glabrous; style and style head 3.5–5.5 mm (SING); Ulu Sungei Redong, Kg. Tenang,
long. Fruit of paired follicles; 10–50 cm long, Labis, Jumali 4339 (SINU). Kelantan: Gua
5–17 mm wide, glabrous, lenticellate or not Panjong at Gua Ninik, Henderson 19599
(see note). Seeds linear; 18–22 x 1.7–2.5 mm; (SING); Kuala Mersing, Sg. Brok, Ng FRI
coma 3.3–4.0 cm long. 5392 (KEP); SE, Ulu S. Labir Kecil, Whitmore
Distribution: southern China and continental FRI 4405 (KEP, L, SING). Kuala Lumpur: Ulu
Southeast Asia through Malesia to Queensland. Gombak FR, Kochummen KEP 99607 (KEP).
Habitat: in a wide variety of forest habitats. Negri Sembilan: Johol, Ridley s.n. (SING); Ulu
Additional collections studied: Sine loc.: Pedas, Nur 11728 (SING). Pahang: Jalan
Forbes 2802 (L). EAST TIMOR. Bobonaro, Simpon, FR, Taman Negara, Ang FRI 23347
Balibo: Widjaja 1213 (L). INDONESIA: (KEP, L), Teo & P. 548 (L, SING); Krau Game
WEST PAPUA: Bernhard Camp, Idenburg reserve, Kuala Lompat, Saw FRI 36282 (KEP);
River, Brass 13893 (BRI, L); Kebar, Asiti, Vink Lanchang, Zainudin 3326 (L); Lesong Forest
BW 11420 (A, CANB, L, SING); Manokwari, Reserve, Rompin Dist., Maxwell s.n. (L,
Kostermans 256 (BO, L, SING); Ransiki, SINU); Nr Batu Balai, Burkill & Haniff 15826
Kostermans 100 (BO, L); Masni Plain, c.25 km (SING); S. Tembeling nr K. Keniyum S.
W of Manokwari, Koster BW 11046 (L); Redab, Whitmore FRI 8568 (KEP, L, SING);
Oemboei, near Andai, SW of Manokwari, Sungei Lenget, Rumping, Yeob 3205 (KEP);
Koster BW 11804 (L); Pulau Japen, Schram BW Taman Negara, 3 km from Kuala Tahan, van
10592 (BO, CANB, L), van Dijk 309 (BO); Balgooy 2436 (L, MO, Z), 2488 (L, Z), Chin &
Pulau Japen, Mariarotu, van Dijk 168 (BO), Mahmud 1289 (KLU). Perak: sine loc., King’s
196 (BO), 226 (BO); Pulau Japen, Wasabori Collector 8614 (BO, L), Scortechini 77 b
near Serui, Aet & Idjan 419 (BO, L); Pulau (SING), Ridley 14949 (SING); Padang Batang,
Japen, Seroei, Neth. Ind. For. Serv. bb 30530 Tapah Hills FR, Loh FRI 17384 (KEP, L,
(L, SING), bb 30391 (L, NY, SING), bb 30417 SING); Ulu Temango, Ridley 14605 (SING).
(L, SING), bb 30447 (SING); Plateau N of Selangor: Bukit Endong FR, Symington 57059
River Pami, 8 km NW of Manokwari, Koster (KEP), 57060 (KEP), 57061 (KEP); Bukit
BW 4349 (CANB, L), Koster BW 4355 Lagong FR, Kochummen FRI 2973 (KEP, L,
(CANB, L); Sausapor, Sorong, Versteegh BW SING), Sakat KEP 116122 (KEP), Chan FRI
3991 (CANB, L), BW 4628 (L); Sekoli Plain, 17654 (KEP, L), Sinclair SFN 40103 (BO, E,
near Jayapura, Iwanggin BW 9136 (BO, BRI, KEP, L, SING), Suppiah FRI 108885 (KEP, L),
L); Tiporra, Rijklof van Goensbaai, Stefels BW Pagi KEP 98711 (KEP, L); Dusun Tua, Ridley
5105 (L); Warsamson valley, E of Sorong, Moll s.n. (SING); Gaiting Bulai, Ridley 7566
BW 11560 (BO, CANB, L); Wersar, 6 km S of (SING);Gunung Nuang Ulu Langat, Symington
2005 MIDDLETON, A REVISION OF WRIGHTIA 171

51751 (KEP); Semempih, Hume 8352 (SING); the subspecies. However, the specimens from
Ulu Gombak FR, Kuala Lumpur, Kochummen New Guinea in fruit show a shorter and wider
99607 (BO, L, SING); Ulu Gombak, Univ. fruit type than is found in the other parts of the
Malaya Field Studies Centre, Soepadmo & range, and the trees are reported as much taller.
Students FSC 832 (KLU, L). PHILIPPINES. In flower they are unremarkable except for the
Sine loc.: Mendoza 42092 (L, SING). LUZON: extremely long and fine corona lobes of the for-
Sorsogon: Irosin, Mt Bulusan. MINDANAO, mer subsp. novoguineensis. This variation does
Davao: de Mesa 27538 (BO, L, SING). not, however, correspond to the limits of any of
PALAWAN: Mt Cleopatra, Reynoso et al PPI the former subspecies and is not the basis for
24423 (L); Mt.Cleopatra Ranges, So. recognizing any formal ranks.
Calabayog, Brgy. Tanabag, Puerto Princesa,
Reynoso et al PPI 24244 (PNH); So. 5. Wrightia novobritannica (Ngan) D. J.
Calabayog, San Rafael, Puerto Princesa, Middleton, stat. nov.
Barbon et al PPI 18700 (PNH). SAMAR: Basionym: Wrightia pubescens subsp. novobri-
Loquilocon, Wright, Sulit 6055 (BO, L, SING); tannica Ngan, Ann. Miss. Bot. Gard. 52:
Mt Cansayao, Catarman, Sulit 14455 (BO, L). 153. 1965. TYPE: PAPUA NEW
PAPUA NEW GUINEA. Buna Hinterland, 7 GUINEA. New Britain, Nodup,
miles NW of Embi Lakes, Smith NGF 1272 Waterhouse 270 (Holotype: NY; Isotype:
(BRI, CANB, L). Central: Brown River forest A, F n.v., US).
station, Eddowes & Maru NGF 13112 (A, BO, Tree. Branchlets puberulent, glabrescent with
BRI, CANB, L, NSW, SING); near Maipa age. Leaves: petiole 4–6 mm long, puberulent;
airstrip, Kairuku subprov., Saunders 1108 (L). blade elliptic to oblong, 12.8–19.0 × 3.6–6.2
Gulf: near Putei, junction of Tauri & Kapau cm, 3.3–3.7 times as long as wide, apex acumi-
rivers, Craven & Schodde 964 (A, BRI, CANB, nate, base cuneate, very sparsely puberulent all
L). Madang: Brahman, Bundi subdist, Henty over above, at least when young, puberulent
NGF 49267 (BO, CANB), NGF 49267 (A, BRI, beneath, more densely so on midrib and vena-
E, L, M, NSW, SING); Gogol River, Katik NGF tion, 13–15 pairs of secondary veins, tertiary
46662 (A, BRI, L, NSW, SING); Utu village, venation reticulate. Inflorescence terminal,
Eddowes & Maru 13057 (A, BRI, E, NSW, many-flowered, 2.5–4.5 cm long; peduncle
SING, US); Ramu Valley, ca. 5 miles SE Faita 0.6–0.9 cm long, densely puberulent; pedicels
airstrip, Saunders 484 (BRI, CANB, L), 505 6–9 mm long, densely puberulent. Sepals
(BRI, L); Utu village, near Madang, Eddowes & ovate, 1.8–2.2 × 1.6–1.8 mm, 1.0–1.4 times as
Maru NGF 13057 (BO, CANB, L). Milne Bay: long as wide, apex obtuse to rounded, densely
E of Lake Lavu, Fergusson Island, Croft et al puberulent, ciliate. Corolla brick red; tube ca.
LAE 68795 (BRI, CANB, L), LAE 68795 (A, E, 3.5 mm long, ca. 1.6–1.9 times as long as
M, SING); near Ailuluai village, Esa’ala sub- sepals, ca. 0.3 times as long as lobes, glabrous
dist, Lelean & Katik NGF 49910 (A, BRI, E, L, or with a few hairs at top of tube outside,
US). Morobe: Bulolo, Havel & Kairo NGF glabrous inside; lobes ca.10.5 × 4.7 mm, 2.2
17009 (BO, BRI, CANB, CBG, SING, US); times as long as wide, elliptic, apex rounded,
Bulolo, Streiman & Kairo NGF 26050 A (A, minutely pubescent outside and inside; ante-
BO, BRI, CANB, L, M, NSW, SING, US); petalous corona adnate to the corolla lobe for
Oomsis near Lae, White NGF 10479 (A, BO, half of length, free at the margins, ca. 3.5 mm
BRI, CANB, L, SING). Northern: near Koreaf long, apex entire, glabrous inside, ca. 2.3 times
village, Tufi subdist., Hoogland 4840 (BO, BRI, as long as alternipetalous lobes; alternipetalous
CANB, L, MEL, NSW). Admiralty Islands, corona lobes simple, ca. 1.5 mm long; alternat-
Manus Island, Unknown NGF 543 (L). SINGA- ing corona lobes lacking. Stamens inserted at
PORE: Bukit Timah, Ridley s.n. (SING). ca. 0.8 of tube length from corolla base,
Wrightia laevis is an extremely widespread strongly exserted from tube; filament ca. 1 mm
and variable species. Ngan (1965) recognized long; anthers 6.5 × 1.0 mm, pubescent outside.
three subspecies across this range on the basis of Gynoecium of two connate carpels, ovary ca. 2
corona and inflorescence pubescence charac- mm high, glabrous; style and style head ca. 5
ters, for which I have been unable to find clear mm long. Fruit of connate follicles, splitting
demarcations. I am not, therefore, recognizing into two at dehiscence, ca. 17.5 cm long, 2.4
172 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

cm wide, sparsely puberulent, lenticellate. Stamens inserted at ca. 0.7 of tube length from
Seeds 13.3–14.8 × 1.5–2.3 mm; coma 3.4–3.8 corolla base, exserted from tube; filament ca. 1
cm long. mm long; anthers 5.0–7.1 × 1.1–1.7 mm, sagit-
Distribution: only from from the type col- tate, adnate to the style head, pubescent on the
lection from New Britain, Papua New Guinea. outside only at the very tip. Gynoecium of 2
Habitat: not known. free carpels united into a common style, ovaries
1.7–2.2 mm high, glabrous; style and style head
6. Wrightia palawanensis D. J. Middleton, sp. 3.0–5.5 mm long. Fruit unknown (see notes).
nov. TYPE: PHILIPPINES. Palawan: Irawan Distribution: Philippines (Palawan).
Valley head on the lower slopes of Mt. Habitat: reported from riverine and mixed
Beaufort, riverine forest, 150 m altitude, 29 forest and from ultramafic soils at 100–200 m
March 1984: Podzorski SMHI 698 (Holotype: altitude.
A; Isotypes: BO, BRI, KEP, L). Fig. 5. Etymology: the specific epithet refers to the
Frutex. Folia elliptica, apice acuminata, basi distribution of this species in Palawan.
cuneata, 2.6–11.1 cm longa, 1.0–3.3 cm lata. Additional collections studied: PHILIP-
Inflorescentia terminalis, 2–5-flora, 1–4 cm PINES. PALAWAN: Bataraza Munic., Bgy Rio
longa, glabra. Calyx lobis 1.9–2.1 mm longis, Tuba, Belanjao Range, Gamayon Settlement,
2.4–2.9 mm latis. Corolla tubus 3.8–4.0 mm Soejarto et al 8402 (L); Brookes Point munici-
longus; lobi 8.5–8.9 mm longi, 7.0–7.7 mm lati, pality, ca. 15 miles N, Infanta mining area,
alabastro sinistrosum obtegentes. Podzorski SMHI 2156 (A, L).
Shrub or small tree to 3 m tall. Branchlets
This new species would appear to have
glabrous, sparsely lenticellate. Leaves: petiole
affinities to Wrightia hanleyi Elmer, but it has
5–7 mm long, glabrous; blade elliptic, 2.6–11.1
larger flowers and the stamens are inserted
× 1.0–3.3 cm, 2.4–4.3 times as long as wide,
higher in the corolla tube. It differs from
apex acuminate, base cuneate, glabrous above
and beneath, 10–17 pairs of secondary veins, Wrightia candollei Vidal in the free carpels and
tertiary venation reticulate. Inflorescence termi- the lack of pubescence on the outside of the
nal, 2- to 5-flowered, 1–4 cm long; peduncle anthers. None of the specimens report the pres-
0.2–1.4 cm long, glabrous; pedicels 5.5–7.0 mm ence of white latex, but it would be surprising
long, glabrous. Sepals ovate, 1.9–2.1 × 2.4–2.9 if this were not present. I have not seen any
mm, 0.7–0.8 times as long as wide, apex obtuse fruit, but they are reported on one specimen to
to rounded, glabrous, ciliate. Corolla salmon be paired.
pink or yellow, infundibuliform; tube 3.8–4.0
mm long, widening at stamen insertion, 1.8–2.1 7. Wrightia pubescens R. Br., Prod. 467. 1810;
times as long as sepals, 0.4–0.5 times as long as R. Br., Asclep. 64. 1810; Miq., Fl. Ned. Ind. 2:
lobes, sparsely and minutely puberulent at top 432. 1857; Pichon, Not. Syst. 14: 83. 1951;
of tube outside, glabrous inside; lobes 8.5–8.9 × Ngan, Ann. Miss. Bot. Gard. 52: 150. 1965;
7.0–7.7 mm, 1.1–1.3 times as long as wide, Backer & Bakh.f., Fl. Java 2: 241. 1965; Tsiang
elliptic, apex rounded, minutely puberulent out- & P. T. Li, Fl. Reip. Pop. Sin. 63: 123. 1977;
side and inside; antepetalous corona lobes Lý, Fedd. Rep. 97: 623. 1986; Li et al., Fl.
adnate to the corolla lobe for entire length but China 16: 174. 1995; Ba et al., PROSEA 5.2:
free at margins, 4.3–4.5 mm long, apex suben- 513. 1995; Forster, Fl. Australia 28: 191. 1996;
tire to weakly crenate, glabrous inside, ca. 6 Middleton, Fl. Thailand 7: 86. 1999; Kessler et
times as long as alternipetalous lobes; al., Blumea suppl. 14: 15. 2002. TYPE:
alternipetalous corona lobes free but often so AUSTRALIA. North Coast (= Northern
small as to be difficult to see, simple, 0.7–0.8 Territories), Brown Iter Australiense 2861
mm long; alternating corona lobes lacking. (Lectotype: BM).

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
2005 MIDDLETON, A REVISION OF WRIGHTIA 173

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

174 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

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

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

Krawang, Tandjoengpoera, Koorders 13798 ß Mt. Natib, Reynoso et al PPI 17222 (GH);
(BO, L); Res. Pekalongan, N of Soebah, Alambra 23206 (A, SING); Alvarez 12939 (BO,
Hofmann 6539 (BO, GH); Res. Pekalongan, L); Gangan 22701 (A); Pascual 23062 (L);
near Soebah, Koorders 22549 ß (BO, L), 27294 Ranario 27780 (SING); Sulit 25679 (NSW);
ß (BO, L), 37066 ß (BO), 197 ß (BO, P), 201 ß Topacio 20009 (NSW). Batangas: Mt Lobo,
(BO), 13481 ß (BO, L), 13800 ß (BO, L), 13801 Sulit 15715 (A, L, PNH, SING), 15746 (A).
ß (BO, L), 13802 ß (BO, L), 14229 ß (BO, L), Bosoboso: Morong, Loher 4000 (M). Bulacan:
14232 ß (BO), 14244 ß (BO, L); Res. Preanger, Mt. Biak na Bato, San Miquel, Garcia et al PPI
Afd.Soemedang, Tomo, Koorders 195 (BO, L); 15071 (GH, L); Leuterio 24730 (BO, L, NSW).
Res.Preanger, Afd.Soekadoemi, Koorders Cagayan: near Penablanca, Adduru 197 (A, P);
34247 ß (BO); Res. Preanger, Afd.Soekaboemi, Alvarez 18541 (NSW); Bernardo 15138 (L, P).
Palaboeanratoe, Koorders 194 ß (BO, L), 200 ß Cavite: Ramos & Deroy 22555 (GH, SING).
(BO, L), 12232 ß (BO, L); Res. Rambang, Ilocos Norte: Adduru 22999 (A). Isabela:
Sedan, Ngandang, Koorders 36428 ß (BO); Barros 26288 (A); San Mariano, Ramos &
Res. Semarang, Karangassem, Koorders 198 ß Edaño 47008 (BO), Siriban 30396 (NSW,
(BO, L), 28200 ß (BO), 38874 ß (BO, L); Res. SING); Sierra Madre Mountains, San Mariano,
Semarang, Kedoengdjati, Koorders 272 ß (BO), Gutierrez 78044 (L). Laguna: Los Banos, Elmer
24516 ß (BO), 33704 ß (BO, L). Carita, 10 km 8069 (E); Mt. Makiling, Sulit 6875 (A),
N of Labuhan, Leeuwenberg 13217 (L); Whitford 19784 (L). Masbate: Bgy. Matipuron,
Cempaka putih, Backer s.n. (L); Simpang, Milagros, Barbon et al PPI 12598 (GH). Nueva
Bakh.v.d.Brink 4892 (L). SULAWESI: Bikerve Ecija: Curran 8448 (NSW), Tungol 24142 (A,
Balang, Warburg 16325 (A); central part, GH). Nueva Vizcaya: Dupax, McGregor 11298
Wayingan near Palu, Meijer 10226 (BO, L, (L), 11300 (L). Pangasinan: Mayor 24577
MO); Manado, Bralema, Bilatto, Neth.Ind.For. (NSW). Rizal: Ahern’s collector 90 (A), 2960
Serv. bb 15045 (L); North central, near Palu, 7 (BO, P, SING); Franco 17856 (GH); Merrill
km to Donggala, Meijer 9273 (L, MO); sp.Bl. 562 (A, BO, GH, L, MO, P); Ramos 2042
Sulawesi Tengah: W of Toboli, along road to (BO, BRI, L, P, NSW, SING); Robinson 6738
Palu, Leeuwenberg et al 14567 (A); Sulawesi (BRI, NSW). Zambales: Mt. Pinatubo, Balintay
Tengah: Wera Waterfall near Parampadene, 25 5626 (A). MARINDUQUE: So. Matalim, Bgy.
km SW of Palu, van Balgooy 3622 (A, BO, L). Dampulan, Romero & Chavez PPI 29153 (GH,
SUMATRA: sine loc., Unknown 230 (L). L). MINDANAO: Cotabato: Buayan, Ramos &
MALAYSIA. Jongkah, Curtis 3054 (SING). Edano 85136 (A). Zamboanga: Zamboanga,
Kedah: sine loc., Meh 9012 (KEP), Ridley Ramos & Edano 37465 (A). Zamboanga del
14950 (SING), Ridley 14951 (SING); Jitra, Sur: Bolong, Reynoso et al PPI 4486 (L, PNH).
Burkill 13349 (SING); Langkawi: Pulau Tuba, MINDORO: Pandarucan, Elmer 943 (GH),
Zainudin 5231 (L). Penang: Mainland, Tasoek Merrill 52932 (NSW). Occidental: Paluan,
Glugor, Curtis 3738 (SING). Perak: Lake Ramos 39677 (A). Oriental: East of Yagaw,
Gardens, Taiping, Corner s.n. (SING). Perlis: Conklin 37568 (L, SING); Puerto Galera,
Chupeny, Ridley 14952 (SING). Terengganu: Santos 5259 (L). NEGROS: Oriental:
Jalan Hubor, Kuala Terengganu, Sinclair 39986 Dumaguete (Cuernos Mts), Elmer 10291 (A,
(BO, L, SING); Jalan Kubor, Kuala Trengganu, BO, E, L, NSW). PALAWAN: Langen
Sinclair [SING39986] 7718 (E). PHILIP- (Malapakan) Isl., Podzorski SMHI 840 (A, BO,
PINES. sine loc., Unknown 460 (L). PANAY: KEP, L); Roxas, Langugan Zigzag, 85 km from
Antique: Brgy. Simirara Island, Caluya, Romero Puerto Princesa, Soejarto & Reynoso 6320 (A,
et al PPI 29509 (GH, PNH). BUSUANGA: L, SING); Taytay, Merrill 9200 (BO, GH, L, P,
Marche 320 (P). CULION: Elmer 676 (GH, SING). PANAY: Mt. Baloy, Musat,
SING); Herre 1049 (A). LUZON: Manila, Varadarajan et al 1537 (A, L, MO). SINGA-
Loher 3999 (M). Bataan: Lamao River, Mt PORE: Chinese Temple, Telok Belanga Drive,
Mariveles, Borden 770 (BO, P, NSW, SING); Ibrahim AI 66 (SING).
Lamao River, Mt Mariveles, Borden 3067 (BO, Wrightia pubescens ssp. lanitii would appear
SING); Mt. Natib, Bgy. Banawang, Fuentes & to be as different from ssp. pubescens as ssp.
de la Rosa PPI 38807 (L); Tawawa, slope of candollei, which has been resurrected as a
2005 MIDDLETON, A REVISION OF WRIGHTIA 179

species distinct from Wrightia pubescens. sparsely lenticellate. Leaves: petiole 1.5–4.0
Wrightia pubescens ssp. lanitii and ssp. pubes- mm long; blade papery, elliptic or oblong,
cens differ in the size and shape of the sepals 1.2–8.2 × 0.6–3.3 cm, 2.2–4.2 times as long as
and, for the most part, the pubescence of the wide, apex acuminate, rarely to rounded, base
antepetalous corona lobes. However, there are cuneate, puberulent on midrib beneath and
individuals that have some pubescence on the sometimes on midrib above, or completely
inside of the corona lobes, and specimens from glabrous, blade punctate beneath and some-
Sulawesi have the corona type and sepal shape times above, 3–8 pairs of secondary veins,
of ssp. pubescens but the sepal size of ssp. lan- ascending, often obscure, tertiary venation
itii. Because of these intermediates and the obscure. Inflorescence 1.7–4.2 cm long;
greater similarity in corolla shape and size glabrous or sparsely puberulent; pedicels 8–30
between these two subspecies compared to ssp. mm long, very delicate; flowers pendulous.
candollei, they are being maintained as sub- Sepals ovate or oblong, 1.0–2.5 × 0.7–1.0 mm,
species, whereas Wrightia candollei, which is 1.9–2.4 times as long as wide, apex obtuse to
also sympatric with W. pubescens ssp. lanitii, acute, glabrous, ciliate; colleters small and nar-
has been given specific status. row. Corolla white; subrotate; sometimes dou-
ble in cultivated plants; tube 2.1–4.2 mm long;
8. Wrightia religiosa (Teijsm. & Binn.) Benth. lobes 4.8–9.8 × 1.9–2.7 mm, 2.1–2.6 times as
ex Kurz, J. As. Soc. Beng. 46, 2: 258. 1877; long as wide, obovate, apex rounded; puberu-
Kurz, Fl. Burm. 2: 194. 1877; Hook. f., Fl. Brit. lent-papillose on lobes outside and inside,
Ind. 3: 653. 1882; King & Gamble, J. As. Soc. glabrous on tube outside and inside; corona
Beng. 74, 2: 464. 1907; Ridley, Fl. Mal. Pen. 2: absent. Stamens inserted at corolla mouth,
353. 1923; Pitard in Lecomte, Fl. Gén. Indo- strongly exserted from tube; filaments 1.5–2.5
Chine 3: 1183. 1933; Kerr, Fl. Siam. En. 2: 458. mm long; anthers 3.8–4.0 × 0.7–0.8 mm,
1939; Pichon, Not. Syst. 14: 85. 1951; Ngan, pubescent within and without. Gynoecium of 2
Ann. Miss. Bot. Gard. 52: 159. 1965; Backer & free carpels united into a common style, ovaries
Bakh. f., Fl. Java 2: 241. 1965; Whitmore, Tree 0.8–1.1 mm high, glabrous, of two separate
Fl. Mal. 2: 24. 1973; Lý, Feddes Repert. 97: carpels; style and style head 5.7–8.3 mm long.
622. 1986; Li et al., Fl. China 16: 175. 1995; Fruit of paired follicles; 8.5–17.5 cm long,
Turner, Gard. Bull. Sing. 47: 131. 1997; 3.3–4.5 mm wide; glabrous; not lenticellate.
Middleton, Fl. Thailand 7: 88. 1999. Fig. 8. Seeds linear; 7.0–8.6 × 1.1–1.3 mm; coma
Basionym: Echites religiosa Teijsm. & Binn., 3.0–3.5 cm long.
Tijdschr. Ned. Ind. 27: 34. 1864. - TYPE: Distribution: Thailand, Cambodia, possibly
Teijsmann s.n. (untraced). NEOTYPE: Peninsular Malaysia. The natural distribution
Collins 669 (Neotype: E, designated
of this species is now obscure because it is so
here; isoneotype: ABD, K, TCD),
widely cultivated, but is probably Thailand and
Thailand, Chon Buri, Sri Racha.
maybe Cambodia. It may also be native in
Shrub or small tree to 5 m tall. Branchlets Peninsular Malaysia, but is certainly widely
puberulent, soon becoming glabrous and cultivated in many parts of Malesia.

NOTE
Wrightia timorensis Miq., Fl. Ned. Ind. 2: 433. else a cultivated plant transported from India
1857. TYPE: TIMOR. Sine loc., Unknown s.n. earlier. There are no notes on the label to indi-
(Lectotype: L [898,112–484]; Isotype: U [frag- cate whether it was a cultivated plant. Until
ment]). such time as further material is collected in
Wrightia timorensis is a synonym of Wrightia Timor that matches this specimen and its status
tinctoria, a species otherwise known only from can be reassessed, I am not including this as a
India. Its appearance in Timor is a mystery and Malesian species to be discussed further.
is likely to be either a mistake on the label or
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 181

LITERATURE CITED
BAHADUR, K. N., AND S. S. R. BENNET. 1978. A new LÝ, T.
D. 1985. Zwei neue Arten der Familie Apocy-
species of Wrightia R. Br. (Apocynaceae) from naceae aus südwestlichen Gebieten Vietnams.
India. Indian Forester 104: 32–35. Feddes Repertorium 96: 173–178.
BRUMMIT, R. K., AND C. E. POWELL. 1992. Authors of MIDDLETON, D. J., AND T. SANTISUK. 2001. A new
Plant Names. Royal Botanic Garden, Kew. species of Wrightia (Apocynaceae: Apocynoideae)
ENDRESS, M.E., AND P. V. BRUYNS. 2000. A revised from Thailand. Thai Forest Bulletin 29: 1–10.
classification of the Apocynaceae s.l. Botanical NGAN, P. T. 1965. A revision of the genus Wrightia
Review 66(1): 1–56. (Apocynaceae). Annals of the Missouri Botanical
HOLMGREN, P. K., N. H. HOLMGREN, AND L.C.BARNETT. Garden 52: 114–175.
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 Ibrahim AI 66 (7b); Iwanggin BW 9136 (4).
419 (4); Ahern’s collector 90 (7b), 1116 (1), 2960 Jaag 21a (7a), 598 (7a), 1084 (7a); Jumali 875 (4),
(7b), 3155 (1); Alambra 23206 (7b); Alston 16098 2866 (2), 4339 (4).
(7a); Alvarez 12939 (7b), 18541 (7b); Ang FRI Katik NGF 46662 (4); Kiah S 151 (2), 35275 (2),
23347 (4). s.n. (2); King’s Collector 8614 (4); Kochummen
Backer 7571 (7a), 9529 (7a), 26577 (7a), 35430 FRI 2973 (4), FRI 16446 (4), KEP 99607 (4),
(7b), 35430 (7b), 35431 (7b), 35432 (7b), s.n. 99607 (4); Koorders 194 ß (7b), 195 (7b), 196 ß
(7b); Bakhuizen v.d. Brink 1662 (7b), 3131 (7b), (7b), 197 ß (7b), 198 ß (7b), 199 ß (7a), 200 ß (7b),
4851 (7b), 4892 (7b); Balintay 5626 (7b); Barbon 201 ß (7b), 272 ß (7b), 12232 ß (7b), 13481 ß (7b),
et al. PPI 12598 (7b), 18700 (4), 22357 (1); Barros 13798 ß (7b), 13799 ß (7b), 13800 ß (7b), 13801 ß
26288 (7b); Bartlett 14074 (1), 14720 (1); (7b), 13802 ß (7b), 14229 ß (7b), 14232 ß (7b),
Beeking 45 (7a); Bernardo 15138 (7b); 14244 ß (7b), 22549 ß (7b), 24516 ß (7b), 24516 ß
Bloembergen 3346 (7a), 3743 (7a); Blume 1521 (7b), 27294 ß (7b), 28200 ß (7b), 33704 ß (7b),
(7b), 1527 (7b); Borden 770 (7b), 3067 (7b); 34247 ß (7b), 36428 ß (7b), 36598 ß (7b), 37066 ß
Borssum Waalkes 3122 (7a); Branderhorst 144 (7b), 38874 ß (7b), 38905 ß (7a); Koster BW 4349
(7a), 284 (7a); Brass 5570 (4), 13893 (4); Burkill (4), BW 4355 (4), BW 11046 (4), BW 11804 (4);
13349 (7b); Burkill & Haniff 15826 (4); Burtt & Kostermans 256 (4), 18001 A (7a); Kostermans et
Woods B 1729 (2). al. KKSS 10 (7a), 100 (4); Kostermans &
Chan FRI 17654 (4); Chin & Mahmud 1289 (4); Wirawan 185 (7a), 940a (7a); Kuswata &
Clemens 18171 (1); Conklin 37568 (7b); Kartawinata 1750 (7a).
Contreras 23404 (1); Corner s.n. (7b); Craven & Laumonier TFB 4464 (4); Leeuwenberg 13217 (7b);
Schodde 964 (4); Croft et al. LAE 68795 (4), LAE Leeuwenberg et al. 14567 (7b); Lelean & Katik
68795 (4); Cuming 1279 (7b), 1453 (1); Curran NGF 49910 (4); Leschenault 309 (7a); Leuterio
4510 (1), 8442 (1), 8448 (7b); Curtis 188 (2), 2915 24730 (7b); Loh FRI 17384 (4); Loher 3999 (7b),
(2), 2943 (4), 3054 (7b), 3738 (7b), s.n. (2). 4000 (7b), 6519 (1), 6530 (1), 6542 (1); Lopez &
De Mesa 27538 (4); De Vriese & Teysmann s.n. Reyes 27331 (1).
(7a); De Vriese s.n. (7a); De Wilde & de Wilde- Madulid & Majaducon PPI 36005 (1); Majaducon &
Duyfjes 12369 (4), 20199 (4); Docters v.Leeuwen Fernando 8455 (1); Marche 320 (7b); Maxwell
997 (7b). s.n. (4); Mayor 24577 (7b); McGregor 11298 (7b),
Eddowes & Maru NGF 13057 (4), NGF 13112 (4); 11300 (7b), 18579 (1), 41431 (1); Meh 9012 (7b);
Elbert 742 (7a), 3650 (7a), 3832 (7a), 3898 (7a); Meijer 9273 (7b), 10226 (7b); Mendoza 42092
Elmer 676 (7b), 943 (7b), 3049 (1), 6348 (1), 8069 (4), 91195 (3); Merrill sp. Bl. 562 (7b), 3082 (1),
(7b), 10291 (7b), 12873 (3), 15595 (4); Escritor 9200 (7b), 9373 (3), 9426 (1), 52932 (7b); Moll
21296 (1). BW 11560 (4); Moton KEP 94083 (4).
Fenix 1183 (1); Forbes 2766 (4), 2802 (4); Franco Neth. Ind. For. Serv. bb 15045 (7b), bb 21611 (7a),
17856 (7b); Fuentes & de la Rosa PPI 38807 (7b). bb 24979 (7a), bb 30391 (4), bb 30417 (4), bb
Gaerlan et al. PPI 10603 (1), PPI 13448 (3); 30447 (4), bb 30530 (4); Ng FRI 5392 (4);
Gammill 228 (1); Gangan 22701 (7b); Garcia et al. Noerkas 251 (7a); Nur 11728 (4).
PPI 15071 (7b), PPI 18505 (1); Gutierrez 78044 O’Sullivan s.n. (2).
(7b). Pagi KEP 98711 (4); Pascual 23062 (7b); Pereira s.n.
Haniff 7086 (2); Havel & Kairo NGF 17009 (4); (7a); Pleyte 27 (7a); Podzorski SMHI 698 (6),
Henderson 19599 (4); Henty NGF 49267 (4), SMHI 840 (7b), SMHI 2156 (6).
NGF 49267 (4); Herre 1049 (7b); Hofmann 6539 Rahmat Si Boeea 7533 (4); Ramos 2042 (7b), 8284
(7b); Hoogland 4840 (4); Hotta 25889 (4); Hume (1) 39677 (7b), 41607 (1), 42712 (1); Ramos &
8352 (4). Deroy 22555 (7b); Ramos & Edaño 37465 (7b),
182 HARVARD PAPERS IN BOTANY Vol. 10, No. 2

47008 (7b), 85136 (7b); Ranario 27780 (7b); 7428 (1), 14455 (4), 15715 (7b), 15746 (7b), PNH
Reynoso et al. PPI 4486 (7b), PPI 17222 (7b), PPI 17044 (1), 22879 (7b), 25679 (7b); Sun Hong-Fan
24244 (4), PPI 24249 (3), PPI 24423 (4); Ridley Herb. 9064 (7a); Suppiah FRI 108885 (4);
5521 (2), 7566 (4), 14605 (4), 14949 (4), 14950 Symington 51751 (4), 57059 (4), 57060 (4),
(7b), 14951 (7b), 14952 (7b); Ridsdale 1080 (3), 57061 (4).
1418 (1), SMHI 1620 (3); Riedlé s.n. (7a); Taylor 2731 B (7a); Teo & P. 548 (4); Teysmann HB
Robinson 6738 (7b), 6757 (1); Romero & Chavez 5121 (7a), HB 5181 (7a), 8744 (7a), 8754 (7a),
PPI 29153 (7b); Romero & Fuentes PPI 37562 (1); 13602 (7a); Topacio 20009 (7b); Tungol 24142
Romero et al. PPI 29509 (7b). (7b).
Sakat KEP 116122 (4); Samsuri & Ahmad SA 691 Van Balgooy 2436 (4), 2488 (4), 3622 (7b), 5691
(4); Santos 5259 (7b); Saunders 484 (4), 505 (4), (7a); Van Dijk 168 (4), 196 (4), 226 (4), 309 (4);
1108 (4); Saw FRI 36282 (4); Schmutz 1022 (7a); Van Steenis 3055 (7b), 18029 (7a); Varadarajan et
Schram BW 10592 (4); Scortechini 77 b (4); al. 1537 (7b); Verheijen 1005 (7a), 2296 (7a),
Selvaraj FRI 13006 (2); Sinclair [SING39023] 4184 (7a), 4902 (7a); Versteegh BW 3991 (4), BW
6627 (2), [SING39986] 7718 (7b), 39023 (2), 4628 (4), BW 4971 (4); Vink BW 11420 (4).
39986 (7b), SFN 40103 (4); Siriban 30396 (7b); Warburg 16325 (7b), 17189 (7a); Waterhouse 270
Smith NGF 1272 (4); Soejarto & Reynoso 6261 (5); White NGF 10479 (4); Whitford 19784 (7b);
(3), 6320 (7b); Soejarto et al. 6456 (1), 6479 (3), Whitmore FRI 4405 (4), FRI 8568 (4); Widjaja
8402 (6), 8602 (3); Soepadmo & Students FSC 1213 (4); Wiriadinata s.n. (7a).
832 (4); Spanoghe s.n. (7a); Stefels BW 5105 (4); Yeob 3205 (4); Yoshida AN 62183 (7a).
Stone et al. 120 (2), PPI 262 (3); Streiman & Kairo Zainudin 3326 (4), 5231 (7b); Zainudin et al. 4391
NGF 26050 A (4); Sulit 6055 (4), 6875 (7b), PNH (2); Zippelius s.n. (7b); Zollinger 598 (7b).

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) Wrightia laevis subsp. millgar (F.M.Bailey) Ngan (4)
Cameraria dubia Sims (2) Wrightia laevis subsp. novoguineensis Ngan (4)
Echites religiosa Teijsm. & Binn. (8) Wrightia lanitii (Blanco) Merr. (7b)
Nerium macrocarpum Span. (7a) Wrightia macrocarpa Pitard (4)
Scleranthera cambodiensis (Pierre ex Pitard) Wrightia millgar F.M.Bailey (4)
Pichon (2) Wrightia multiflora Zipp. ex Span. (7a)
Scleranthera dubia (Sims) Pichon (2) Wrightia novobritannica (Ngan) D.J.Middleton (5)
Strophanthus jackianus Wall. ex G.Don (2) Wrightia ovata A.DC. (7b)
Walidda antidysenterica (L.) Pichon (x) Wrightia palawanensis D.J.Middleton (6)
Wrightia antidysenterica L. (x) Wrightia pubescens R.Br. (7)
Wrightia balansae Pitard (4) Wrightia pubescens subsp. candollei (Vidal) Ngan (1)
Wrightia calycina A.DC. (7a) Wrightia pubescens subsp. lanitii (Blanco) Ngan (7b)
Wrightia cambodiensis Pierre ex Pitard (2) Wrightia pubescens subsp. novobritannica Ngan (5)
Wrightia candollei Vidal (1) Wrightia pubescens subsp. pubescens (7a)
Wrightia dubia var. membranifolia Wrightia religiosa (Teijsm. & Binn.) Benth. ex
King & Gamble (2) Kurz (8)
Wrightia dubia (Sims) Spreng. (2) Wrightia rubriflora Pitard (2)
Wrightia hainanensis Merr. (4) Wrightia sorsogonensis Elmer (4)
Wrightia hainanensis var. chingii Tsiang (4) Wrightia spanogheana Miq. (7a)
Wrightia hainanensis var. variabilis Tsiang (4) Wrightia timorensis Miq. (x)
Wrightia hanleyi Elmer (3) Wrightia tinctoria R.Br. (x)
Wrightia javanica A.DC. (7b) Wrightia tinctoria var. laevis (Hook.f.) Pichon (4)
Wrightia kontumensis Lý (2) Wrightia tomentosa var. cochinchinensis Pierre ex
Wrightia laevis Hook.f. (4) Pitard (7b)