revision of wrightia

The genus Wrightia R. Br. was last revised byNgan (1965) who recognized 23 species andseveral subspecies. Since then a small number ofadditional species have been described (Bahadurand Bennet, 1978; Ly, 1985; Middleton andSantisuk, 2001) from India, Vietnam, andThailand. The genus is here revised for the forth-coming Flora Malesiana account of theApocynaceae, subfamilies Rauvolfioideae andApocynoideae (sensu Endress and Bruyns,2000). Eight species are recognized, one ofwhich has two subspecies. A new species fromrecently collected material in Palawan isdescribed, and the status of some of Ngan’s sub-species within Wrightia pubescens arereassessed. Ngan (1965) has given a very good

and thorough assessment of the morphologicalcharacters in Wrightia.

The genus shows some interesting biogeo-graphic patterns in Malesia. Most curious is thecomplete absence of any collections of the genusfrom Borneo, despite the widespread distribu-tion of taxa like Wrightia laevis and Wrightiapubescens subsp. lanitii in the surroundingregions. There are two species, Wrightia hanleyiand Wrightia palawanensis, endemic to Palawanin the Philippines, and another, Wrightia can-dollei, more widespread but still endemic to thePhilippines. Wrightia novobritannica is knownonly from one collection from the Papua NewGuinea Island of New Britain and would appearto have similarities to Wrightia candollei.

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 whichhas two subspecies. One new species, Wrightia palawanensis, is described, and one new combination, Wrightianovobritannica, is made. A key to the species is given, and the taxa are described. Lectotypes and neotypes aredesignated for a number of names.

Keywords: Apocynaceae, Apocynoideae, revision, Wrightia, Malesia.

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.

1Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Current address: RoyalBotanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland. E-mail: [email protected]

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

TAXONOMIC TREATMENT

Wrightia R. Br., Asclep. 62. 1810; R. Br., Prod.467. 1810; Benth. & Hook. f., Gen. Pl. 2: 712.1876; K. Schum. in Engler & Prantl, Nat.Pflanzenfam. 4, 2: 183. 1895; Pitard inLecomte, Fl. Gén. Indo-Chine 3: 1182. 1933;Pichon, Mém. Mus. Nat. Hist. Nat, sér. B, Bot.1: 73. 1950; Pichon, Not. Syst. 14: 77. 1951;

Ngan, Ann. Miss. Bot. Gard. 52: 114. 1965;Tsiang & Li, Fl. Reipubl. Popularis Sin. 63:119. 1977; Ly, Feddes Repert. 97: 619. 1986;Li et al., Fl. China 16: 174. 1995; Middleton,Fl. Thailand 7: 79. 1999. Wrightia sect.Bammatophyton A.DC., Prod. 8: 404. 1844.Type species: Wrightia pubescens R. Br.

MATERIALS AND METHODS

Approximately 350 herbarium specimensfrom Asia and Malesia have been studied. Theseare from the following herbaria: 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(Holmgren et al., 1990). All specimens cited

have been seen unless otherwise indicated.All dimensions given are for dried specimens

except for androecium and gynoecium charac-ters, which are from flowers reconstituted byboiling in water.

Author citation follows Brummitt and Powell(1992).

Synonyms: Annasser Blanco, Fl. Filipp. 112(1837). Type species: Anasser lanitiiBlanco (= Wrightia pubescens subsp.lanitii (Blanco) Ngan)

Scleranthera Pichon, Not. Syst. 14: 88.1951. Type species: Scleranthera cambo-diensis (Pierre ex Pitard) Pichon (=Wrightia dubia (Sims) Spreng.)

Wrightia sect. Walidda A. DC. in DC.,Prod. 8: 407. 1844. Walidda (A. DC.)Pichon, Not. Syst. 14: 87. 1951. Typespecies: Walidda antidysenterica (L.)Pichon (= Wrightia antidysenterica L.)

Shrubs or small trees. Leaves opposite; usu-ally with glands in the axils. Inflorescence a ter-minal cyme; few to many flowered. Sepals withlarge or small colleters in the sinuses inside, thelarger ones almost covering the inner surface ofthe sepals. Corolla lobes sinistrorse; matureflower rotate, salverform or infundibuliform;

corona of varying degrees of elaboration usu-ally present, rarely absent or completely adnateto the corolla. Stamens inserted at corolla mouthor in tube; strongly exserted from or completelyincluded in corolla tube; filaments usually quiteshort and wide; anthers narrowly triangular,apex acuminate, base sagittate, sterile at sidesand base with fertile locules on the inner upperhalf, adnate to the style head, often pubescenton the surface facing inward and/or the surfacefacing outward. Disk absent. Gynoecium of 2free carpels united into a common style or oftwo connate carpels, glabrous, rarely pubescent;style broadest near top; ovules numerous. Fruitof paired follicles or connate follicles; fusiform.Seeds linear or narrowly fusiform; with a comadirected toward the base of the follicle.

There are ca. 25 species in Africa, Asia, andAustralia, and 8 species in Malesia (one ofwhich may not be native, see note underWrightia religiosa).

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

KEY TO THE SPECIES OF WRIGHTIA IN MALESIA

1a. Corolla infundibuliform; stamens completely included in the corolla tube . . . . . . . . . . . . . . . . . . . . . 2. W. dubia1b. Corolla rotate, salverform, or somewhat infundibuliform; stamens clearly exserted from the corolla tube. . . . . . 22a. Flowers pendulous; corona absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. W. religiosa2b. Flowers pendulous or not; corona present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33a. Corona with antepetalous lobes divided more than halfway to base, giving the whole corona a fimbriate

look. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. W. laevis3b. Corona with antepetalous lobes entire or divided less than halfway to base . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44a. Antepetalous corona lobes deeply divided or at least strongly dentate . . . . . . . . . . . . . . . . . . . . . 7. W. pubescens4b. Antepetalous corona lobes entire or at most weakly crenate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55a. Gynoecium apocarpus but united into a single style; fruits paired; Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . 65b. Gynoecium syncarpous; fruits solitary; Philippines and New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76a. Corolla tube 3.8–4.0 mm long; stamens inserted in the upper half of the corolla tube . . . . . . 6. W. palawanensis6b. Corolla tube 1.8–2.8 mm long; stamens inserted in the lower half or around the middle of the corolla

tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. W. hanleyi7a. Corolla tube 3.7–10.0 mm long, 1.7–5.5 times as long as sepals; Philippines . . . . . . . . . . . . . . . . 1. W. candollei7b. 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.Philipp. 186. 1885 (as cadollei). Fig. 1.Synonym: Wrightia pubescens subsp. candollei

(Vidal) Ngan, Ann. Miss. Bot. Gard. 52:153. 1965. TYPE: PHILIPPINES. Luzon,Manila, Cuming 1453 (Lectotype: P, des-ignated here; Isotype: BM, K).

Branchlets sparsely to densely puberulent.Leaves: petiole 4–8 mm long, glabrous tosparsely puberulent; blade elliptic, 4–20 ×3.0–6.5 cm, 1.3–3.7 times as long as wide, apexacuminate, base cuneate to rounded, sparselypuberulent to glabrous above, sparsely todensely pale puberulent all over or only on

midrib or glabrous beneath, 10–16 pairs of sec-ondary veins, tertiary venation reticulate.Inflorescence 2.5–3.5 cm long, often with leafybracts; peduncle 0.2–1.3 cm long, sparselypubescent, pedicels 4–7.5 mm long, pubescent.Sepals ovate, 1.1–4.0 × 1.8–4.5 mm, 0.3–1.1times as long as wide, apex rounded, sparselypubescent, ciliate, colleters large in sinuses.Corolla orange; tube 3.7–10.0 mm long,1.7–5.5 times as long as sepals, 0.4–0.5 timesas long as lobes, minutely puberulent at top oftube outside, glabrous inside; lobes 6.5–15.0 ×3.3–9.0 mm, 1.7–2.0 times as long as wide,minutely puberulent inside and outside; ante-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).

to corolla except at apex, apex subentire toweakly crenate, glabrous inside, 3.8–5.0 timesas long as alternipetalous lobes; alternipetalouscorona lobes simple or slightly bifid 0.7–1.7mm long; alternating corona lobes lacking.Stamens inserted at 0.7–0.8 of tube length fromcorolla base, exserted from tube; filaments0.8–1.0 mm long; anthers 5.0–7.8 × 1.1–1.8mm, pubescent on the outside or only at tipsoutside. Gynoecium of two connate carpels,ovary 1.5–2.6 mm high; style and style head5–7 mm long. Fruit of a single follicle thatsplits lengthwise at maturity as each of the twocomponent follicles dehisces, 20.5–22.0 cmlong, 1.4–2.2 cm wide, glabrous. Seeds notobserved.

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); McGregor18579 (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, Bartlett14074 (A). Batangas: Bo. Biga, Lobo, SulitPNH 7428 (A, PNH). Benguet: Twin Peaks,Elmer 6348 (NSW, P). Bulacan: AngatWatershed area, Norzagaray, Barbon et al PPI22357 (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); sineloc., Robinson 6757 (BRI, L). Zambales: AcojeMine, Santa Cruz, Ridsdale 1418 (A, BO, KEP,L, MO); San Marcelino, Bgy. Sayasay, Mt.Mandakogdog, Majaducon & Fernando 8455(L). – MASBATE: Elmer 3049 (P); Merrill3082 (BO, L). MINDORO: occidental: Mt.Gonting, Lubang Island, Romero & FuentesPPI 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 RiverValley, Tatanarom, trail up Mt. Beaufort,

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 ofWrightia pubescens by Ngan (1965), but it dif-fers substantially from that species in the shapeand form of the corolla and corona and in theflower color. Therefore, it has been raised backto 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; Ly, 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. Sclerantheradubia (Sims) Pichon, Not. Syst. 14: 90.1951. TYPE: Illustration in plate 1646 inCurtis, Bot. Mag. 39 (1814).

Synonyms: Strophanthus jackianus Wall. exG.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 Pitardin Lecomte, Fl. Gén. Indo-Chine 3: 1184.1933; Kerr in Craib, Fl. Siam. En. 2: 456.1939. Scleranthera cambodiensis (Pierreex 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 Ly, Feddes Repert.96: 173. 1985; Ly, Feddes Repert. 97:622. 1986. TYPE: Vietnam, Gialai-Kontum, Loc P 2968 (Holotype: HNU;Isotype: HN).



FIGURE 2. Wrightia dubia (Sims) Spreng. (Sidek bin Kiah S.151, A).

Shrub to 2 m tall. Branchlets glabrous orsparsely puberulent, becoming glabrous andlenticellate. Leaves: petiole 2–11 mm long;blade papery to subcoriaceous, elliptic, oblongor obovate, 3.6–26.5 × 1.8–9.8 cm, 2.1–5.2times as long as wide, apex acuminate to sub-caudate, base cuneate to rounded, puberulentabaxially, only on midrib and secondary veinsor, rarely, glabrous and then both surfaces papil-late, 6–14 pairs of secondary veins, ascending,tertiary venation reticulate. Inflorescence2.5–4.5 cm long; sparsely puberulent orglabrous; 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, apexacute to obtuse; sparsely puberulent or glabrous,ciliate; colleters wide and large. Corolla orange,pink or reddish; infundibuliform; tube 6.0–15.5mm 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, apexacuminate to obtuse, pubescent-papillose onlobes outside and slightly so inside; antepetalouscorona lobes adnate to the corolla, 1.5–3.7 mmlong, 1.7–2.5 times as long as alternipetalouslobes, glabrous; alternipetalous corona lobes0.6–2.2 mm long; alternating corona lobes lack-ing. Stamens inserted at 0.3–0.4 of tube lengthfrom corolla base, completely included in thetube; subsessile; anthers 4.9–5.6 × 1.0–1.2 mm,pubescent outside. Gynoecium of 2 free carpelsunited into a common style, ovaries 1.6–2.5 mmlong, glabrous; style and style head 3.0–4.7 mmlong. 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; coma2.6–5.5 cm long.

Distribution: Cambodia, Laos, Vietnam,Thailand, Peninsular Malaysia, Singapore.

Habitat: growing in wide variety of foresthabitats at lower altitudes.

Additional collections studied: MALAYSIA:Kedah: Sine loc., Ridley 5521 (SING); PulauLangkawi, Telok Apau, Haniff 7086 (SING);Pulau Langkawi, Stone et al 120 (KLU); PulauLangkawi, Pulau Singa Besar, Zainudin et al4391 (L). Penang: Curtis 188 (SING); Curtiss.n. (SING); below Waterfall gardens, Sinclair39023 (E, SING); valley of stream belowwaterfall gardens, Sinclair [SING39023] 6627(E); Butterworth, O’Sullivan s.n. (SING).

Perlis: Kaki Bukit, Burtt & Woods B 1729 (E),Kiah 35275 (A, BO, L), Kiah s.n. (SING).Terengganu: Gunong Tebu, Besut, Selvaraj FRI13006 (KEP, L). SINGAPORE: BotanicGardens, Jumali 2866 (SINU), Kiah S 151 (E).

This is the most distinctive species ofWrightia in Malesia with its pink flowers andwithout 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. Branchletsglabrous, 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 acuminatewith 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. Sepalsovate, 1.0–1.5 × 0.9–1.1 mm, 1.1–1.4 times aslong as wide, apex acute to obtuse, glabrous,ciliate, colleters small in the sinuses. Corollagreenish-yellow to red; tube with a lower nar-row part and a campanulate upper part, with araised lip inside between the lower and uppertube, 1.8–2.8 mm long, 1.5–1.9 times as long assepals, 0.6–0.7 times as long as lobes, minutelypuberulent at top of tube outside, glabrousinside; lobes 3.0–3.9 × 1.5–3.0 mm, 1.3–2.0times as long as wide, apex rounded, minutelypuberulent inside and outside; antepetalouscorona 1.5–2.9 mm long, entire, adnate tocorolla on medial line but with sides free, apexsubentire or slightly crenate, glabrous inside,4.1–7.5 times as long as alternipetalous lobes;alternipetalous corona lobes apparently absentor 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 corollabase, 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. Fruitpaired, linear, 6.6–18.0 cm long, 2.5–4.0 mmwide, glabrous. Seeds 10.0 × 0.6 mm; coma 21mm long.



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

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 PPI24249 (PNH); Narra Municip., Bgy Calatigas,Purok Campsite, Soejarto et al 8602 (L); PuertoPrincesa (Mt. Pulgar); Puerto Princesa Munic.,Bacungan, Hills above Nagtabon beach,Soejarto et al 6479 (MO); Puerto PrincesaMunic., Takdua Zigzag, 35–40 km from PuertoPrincesa, Soejarto & Reynoso 6261 (A, L);Puerto Princesa, Tagburus, Stone et al PPI 262(PNH); Quezon, Tawa-tawa, Gaerlan et al PPI13448 (PNH); South of Narra, Bato-Bato river,Ridsdale 1080 (L); St Paul’s Bay, MtBloomfield, 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; Ly,Feddes Repert. 97: 623. 1986; Turner, Gard.Bull. Sing. 45: 37. 1993; Ba et al., PROSEA5.2: 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, “ChineseCollector” 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

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. variabilisTsiang, Sunyatsenia 4: 47. 1939. TYPE:Tsiang 2679 (Lectotype: A, designatedhere; Isotype: K, NY), CHINA,Guangdong, Sunyi District.

Wrightia hainanensis var. chingii Tsiang,Sunyatsenia 4: 48. 1939. TYPE: Ching5539 (Holotype: IBSC n.v.; Isotype:NY), CHINA, Guangxi, Mung TungKon, N of Luchen.

Wrightia laevis subsp. novoguineensisNgan, Ann. Miss. Bot. Gard. 52: 138.1965. TYPE: Brass 5570 (Holotype: A;Isotype: BM, BO, BRI, K, NY, US),PAPUA NEW GUINEA, CentralProvince, Mt. Tafa.

Usage synonym: Wrightia tinctoria auct. nonR. 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 roughand flakey, gray, grayish-brown or cream-brown; inner bark white, cream, or pale brown.Branchlets glabrous, sparsely lenticellate withage. 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, apexacuminate, base cuneate to obtuse, glabrous,4–10 pairs of secondary veins, strongly ascend-ing, curved, tertiary venation mostly paler thanlamina in dried specimens, almost perpendicularto midrib and oblique to secondary veins andalso reticulate. Inflorescence 1.8–10.0 cm long;peduncle 1.0–1.6 cm long, glabrous to sparselypuberulent; 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, apexobtuse to rounded, glabrous to densely puberu-lent, ciliate; colleters wide and large. Corollawhite, pale yellowish-white to yellow-orange;subrotate; tube 1.1–3.0 mm long, 1.5–1.7 timesas long as sepals, 0.2–0.4 times as long as



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

lobes, glabrous or minutely puberulent at top oftube 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; coronaof antepetalous, alternipetalous, and alternatelobes; antepetalous lobes adnate to the corollaat base, fimbriate, 2.8–9.4 mm long, 0.9–1.8times 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. Stamensinserted at 0.8–0.9 of tube length from corollabase, completely exserted; filaments 0.2–1.7mm long; anthers 3.9–5.6 × 0.7–1.6 mm,pubescent outside, sometimes only sparsely soat base. Gynoecium of 2 free carpels united intoa common style, ovaries 0.7–1.8 mm long,glabrous; style and style head 3.5–5.5 mmlong. 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 continentalSoutheast 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, IdenburgRiver, Brass 13893 (BRI, L); Kebar, Asiti, VinkBW 11420 (A, CANB, L, SING); Manokwari,Kostermans 256 (BO, L, SING); Ransiki,Kostermans 100 (BO, L); Masni Plain, c.25 kmW of Manokwari, Koster BW 11046 (L);Oemboei, near Andai, SW of Manokwari,Koster BW 11804 (L); Pulau Japen, Schram BW10592 (BO, CANB, L), van Dijk 309 (BO);Pulau Japen, Mariarotu, van Dijk 168 (BO),196 (BO), 226 (BO); Pulau Japen, Wasaborinear Serui, Aet & Idjan 419 (BO, L); PulauJapen, Seroei, Neth. Ind. For. Serv. bb 30530(L, SING), bb 30391 (L, NY, SING), bb 30417(L, SING), bb 30447 (SING); Plateau N ofRiver Pami, 8 km NW of Manokwari, KosterBW 4349 (CANB, L), Koster BW 4355(CANB, L); Sausapor, Sorong, Versteegh BW3991 (CANB, L), BW 4628 (L); Sekoli Plain,near Jayapura, Iwanggin BW 9136 (BO, BRI,L); Tiporra, Rijklof van Goensbaai, Stefels BW5105 (L); Warsamson valley, E of Sorong, MollBW 11560 (BO, CANB, L); Wersar, 6 km S of

Teminaboean, Versteegh BW 4971 (L). SUMA-TRA. Bt. Gajabuih, Ulu Gadut, 15 km E ofPadang City, Hotta 25889 (BO); Palembang,Unknown 1078 (L); Tanjong Ning, Palembang,Forbes 2766 (L, SING); near Sigirik mountainwithin Sebelah Nature Reserve, LaumonierTFB 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 ofLoemban, Asahan, Rahmat Si Boeea 7533 (S,SING). MALAYSIA. Pulau Pomgoh, Curtis2943 (SING); Johor: Kluang, Labis FR, MotonKEP 94083 (KEP, L, SING), Kochummen FRI16446 (A, KEP, L, SING); Gunong Besar, UluSungei 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: GuaPanjong at Gua Ninik, Henderson 19599(SING); Kuala Mersing, Sg. Brok, Ng FRI5392 (KEP); SE, Ulu S. Labir Kecil, WhitmoreFRI 4405 (KEP, L, SING). Kuala Lumpur: UluGombak FR, Kochummen KEP 99607 (KEP).Negri Sembilan: Johol, Ridley s.n. (SING); UluPedas, Nur 11728 (SING). Pahang: JalanSimpon, FR, Taman Negara, Ang FRI 23347(KEP, L), Teo & P. 548 (L, SING); Krau Gamereserve, Kuala Lompat, Saw FRI 36282 (KEP);Lanchang, Zainudin 3326 (L); Lesong ForestReserve, 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, vanBalgooy 2436 (L, MO, Z), 2488 (L, Z), Chin &Mahmud 1289 (KLU). Perak: sine loc., King’sCollector 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); BukitLagong FR, Kochummen FRI 2973 (KEP, L,SING), Sakat KEP 116122 (KEP), Chan FRI17654 (KEP, L), Sinclair SFN 40103 (BO, E,KEP, L, SING), Suppiah FRI 108885 (KEP, L),Pagi KEP 98711 (KEP, L); Dusun Tua, Ridleys.n. (SING); Gaiting Bulai, Ridley 7566(SING);Gunung Nuang Ulu Langat, Symington


51751 (KEP); Semempih, Hume 8352 (SING);Ulu Gombak FR, Kuala Lumpur, Kochummen99607 (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 PPI24423 (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, 7miles NW of Embi Lakes, Smith NGF 1272(BRI, CANB, L). Central: Brown River foreststation, Eddowes & Maru NGF 13112 (A, BO,BRI, CANB, L, NSW, SING); near Maipaairstrip, Kairuku subprov., Saunders 1108 (L).Gulf: near Putei, junction of Tauri & Kapaurivers, Craven & Schodde 964 (A, BRI, CANB,L). Madang: Brahman, Bundi subdist, HentyNGF 49267 (BO, CANB), NGF 49267 (A, BRI,E, L, M, NSW, SING); Gogol River, Katik NGF46662 (A, BRI, L, NSW, SING); Utu village,Eddowes & Maru 13057 (A, BRI, E, NSW,SING, US); Ramu Valley, ca. 5 miles SE Faitaairstrip, 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 alLAE 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 NGF17009 (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 Koreafvillage, 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 widespreadand variable species. Ngan (1965) recognizedthree subspecies across this range on the basis ofcorona and inflorescence pubescence charac-ters, for which I have been unable to find cleardemarcations. I am not, therefore, recognizing

the subspecies. However, the specimens fromNew Guinea in fruit show a shorter and widerfruit type than is found in the other parts of therange, and the trees are reported as much taller.In flower they are unremarkable except for theextremely long and fine corona lobes of the for-mer subsp. novoguineensis. This variation doesnot, however, correspond to the limits of any ofthe former subspecies and is not the basis forrecognizing 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 NEWGUINEA. New Britain, Nodup,Waterhouse 270 (Holotype: NY; Isotype:A, F n.v., US).

Tree. Branchlets puberulent, glabrescent withage. Leaves: petiole 4–6 mm long, puberulent;blade elliptic to oblong, 12.8–19.0 × 3.6–6.2cm, 3.3–3.7 times as long as wide, apex acumi-nate, base cuneate, very sparsely puberulent allover above, at least when young, puberulentbeneath, more densely so on midrib and vena-tion, 13–15 pairs of secondary veins, tertiaryvenation reticulate. Inflorescence terminal,many-flowered, 2.5–4.5 cm long; peduncle0.6–0.9 cm long, densely puberulent; pedicels6–9 mm long, densely puberulent. Sepalsovate, 1.8–2.2 × 1.6–1.8 mm, 1.0–1.4 times aslong as wide, apex obtuse to rounded, denselypuberulent, ciliate. Corolla brick red; tube ca.3.5 mm long, ca. 1.6–1.9 times as long assepals, ca. 0.3 times as long as lobes, glabrousor with a few hairs at top of tube outside,glabrous inside; lobes ca.10.5 × 4.7 mm, 2.2times as long as wide, elliptic, apex rounded,minutely pubescent outside and inside; ante-petalous corona adnate to the corolla lobe forhalf of length, free at the margins, ca. 3.5 mmlong, apex entire, glabrous inside, ca. 2.3 timesas long as alternipetalous lobes; alternipetalouscorona lobes simple, ca. 1.5 mm long; alternat-ing corona lobes lacking. Stamens inserted atca. 0.8 of tube length from corolla base,strongly exserted from tube; filament ca. 1 mmlong; anthers 6.5 × 1.0 mm, pubescent outside.Gynoecium of two connate carpels, ovary ca. 2mm high, glabrous; style and style head ca. 5mm long. Fruit of connate follicles, splittinginto two at dehiscence, ca. 17.5 cm long, 2.4


cm wide, sparsely puberulent, lenticellate.Seeds 13.3–14.8 × 1.5–2.3 mm; coma 3.4–3.8cm 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: IrawanValley head on the lower slopes of Mt.Beaufort, riverine forest, 150 m altitude, 29March 1984: Podzorski SMHI 698 (Holotype:A; Isotypes: BO, BRI, KEP, L). Fig. 5.

Frutex. Folia elliptica, apice acuminata, basicuneata, 2.6–11.1 cm longa, 1.0–3.3 cm lata.Inflorescentia terminalis, 2–5-flora, 1–4 cmlonga, glabra. Calyx lobis 1.9–2.1 mm longis,2.4–2.9 mm latis. Corolla tubus 3.8–4.0 mmlongus; lobi 8.5–8.9 mm longi, 7.0–7.7 mm lati,alabastro sinistrosum obtegentes.

Shrub or small tree to 3 m tall. Branchletsglabrous, sparsely lenticellate. Leaves: petiole5–7 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 aboveand beneath, 10–17 pairs of secondary veins,tertiary venation reticulate. Inflorescence termi-nal, 2- to 5-flowered, 1–4 cm long; peduncle0.2–1.4 cm long, glabrous; pedicels 5.5–7.0 mmlong, glabrous. Sepals ovate, 1.9–2.1 × 2.4–2.9mm, 0.7–0.8 times as long as wide, apex obtuseto rounded, glabrous, ciliate. Corolla salmonpink or yellow, infundibuliform; tube 3.8–4.0mm long, widening at stamen insertion, 1.8–2.1times as long as sepals, 0.4–0.5 times as long aslobes, sparsely and minutely puberulent at topof 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 lobesadnate to the corolla lobe for entire length butfree at margins, 4.3–4.5 mm long, apex suben-tire to weakly crenate, glabrous inside, ca. 6times as long as alternipetalous lobes;alternipetalous corona lobes free but often sosmall as to be difficult to see, simple, 0.7–0.8mm long; alternating corona lobes lacking.

Stamens inserted at ca. 0.7 of tube length fromcorolla base, exserted from tube; filament ca. 1mm long; anthers 5.0–7.1 × 1.1–1.7 mm, sagit-tate, adnate to the style head, pubescent on theoutside only at the very tip. Gynoecium of 2free carpels united into a common style, ovaries1.7–2.2 mm high, glabrous; style and style head3.0–5.5 mm long. Fruit unknown (see notes).

Distribution: Philippines (Palawan).Habitat: reported from riverine and mixed

forest and from ultramafic soils at 100–200 maltitude.

Etymology: the specific epithet refers to thedistribution of this species in Palawan.

Additional collections studied: PHILIP-PINES. PALAWAN: Bataraza Munic., Bgy RioTuba, 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 haveaffinities to Wrightia hanleyi Elmer, but it haslarger flowers and the stamens are insertedhigher in the corolla tube. It differs fromWrightia candollei Vidal in the free carpels andthe lack of pubescence on the outside of theanthers. None of the specimens report the pres-ence of white latex, but it would be surprisingif this were not present. I have not seen anyfruit, but they are reported on one specimen tobe 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;Ly, 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 etal., Blumea suppl. 14: 15. 2002. TYPE:AUSTRALIA. North Coast (= NorthernTerritories), Brown Iter Australiense 2861(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 slightlyreflexed; 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; antepetalouscorona lobes mostly slightly pubescent inside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7b. ssp. lanitii


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

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. Unknowncollector in Museum de Paris 1821(Holotype: G-DC).

Wrightia spanogheana Miq., Fl. Ned.Ind. 2: 434. 1857. Nerium macrocarpumSpan., 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. Basedon TIMOR. Zippelius s.n. (L [herb. no.898,112–449]).

Usage synonym: Wrightia tinctoria auct. nonR. 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 puberulenton midrib to sparsely pubescent all over above,sparsely puberulent on midrib and secondaryveins to densely rusty pubescent all overbeneath. Inflorescence terminal, 2.5–6.0 cmlong; peduncle 0.2–0.8 cm long, sparsely todensely 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 aswide, apex obtuse to rounded and slightlyreflexed or not, margins slightly reflexed,sparsely to densely pubescent, colleters largeand wide in sinuses. Corolla tube 3.6–6.0 mmlong, 1.2–1.5 times as long as sepals, 0.5–0.7times as long as lobes, minutely pubescent attop of tube outside, glabrous to sparselypuberulent inside; lobes 6.7–11.0 × 2.8–4.7mm, 1.8–2.8 times as long as wide, elliptic,apex obtuse; antepetalous corona lobes 3.5–5.2mm long, denate at apex, adnate to corolla lobefor most of length, glabrous inside, 1.3–2.6times 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 fromcorolla base, strongly exserted from tube; fila-ment 1 mm long; anthers 5.6–6.6 × 1.3–1.7mm, densely pubescent outside. Gynoecium oftwo connate carpels, ovary 1.3–2.0 mm high,glabrous; style and style head 6.7–8.2 mm

long. Fruit of connate follicles, splitting intotwo at dehiscence, 6.6–29.5 cm long, 1.2–1.6cm wide (see note), glabrous, lenticellate ornot. Seeds 12.3–17 × 1.3–2.3 mm; coma3.2–4.7 cm long.

Distribution: Australia (NorthernTerritories), Eastern Java, Lesser SundaIslands, Moluccas, New Guinea.

Habitat: in dry forests.Additional collections studied: EAST

TIMOR. Baucau, van Steenis 18029 (BO,CANB, L). INDONESIA. BALI: Kuta, vanBalgooy 5691 (L); Mt Prapat Agung, NW Bali,Kostermans et al KKSS 10 (A, BO, KEP, L, P);Rapakagoeng, Beeking 45 (BO, L). JAVA: sineloc.: de Vriese & Teysmann s.n. (L),Leschenault 309 (P); Bangil, Backer 7571 (L);Besuki, Backer 9529 (BO, L); Kupang nearSurabaya, 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), deVriese s.n. (L). Kep. Tanimber: P.Jamdena,Borssum Waalkes 3122 (BO, L); Olilit nearSaumlaki, 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), Spart, 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), eastof 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



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


Lembeh, Alston 16098 (BO, L). WESTPAPUA: 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 rathermisshapen. The dimensions are not included inthe 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, MakilingNational 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; scrapfrom 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. cochinchinensisPierre 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 Pierre1047 and 1147 have become confusedand that they are in fact the same collec-tion. There are additional specimenslabelled 1147 in NY and P.

Shrub or tree to 15 m tall, to 30 cm dbh, oftendeciduous; wood pale. Branchlets sparselypubescent to tomentose, becoming glabrous;sparsely lenticellate. Leaves: petiole 2–11 mmlong; blade ovate, elliptic or obovate, papery tosubcoriaceous, 1.5–15.0 × 1.0–7.5 cm, 1.2–4.0times 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, tertiaryvenation reticulate. Inflorescence terminal,

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 timesas long as wide, apex obtuse to acute, tomen-tose or puberulent, colleters large and wide inthe 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 timesas long as lobes, minutely puberulent at top oftube 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 orrounded, minutely puberulent outside andinside; antepetalous corona 4.2–9.0 mm long,crenate at apex, adnate to corolla lobes exceptat 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; alternatinglobes lacking. Stamens inserted at 0.7–0.9 oftube length from corolla base, strongly exsertedfrom tube; filaments 0.7–2.0 mm long; anthers5.9–8.5 × 1.3–2.1 mm, pubescent within andwithout. Gynoecium of two connate carpels,ovary 1.5–2.7 mm long, glabrous, carpelsmedially connate; style and style head 8.0–10.5mm long. Fruit of connate follicles, splittinginto 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; coma1.2–4.5 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), Blume1527 (L), Unknown 179 (L), Zippelius s.n. (L),Zollinger 598 (A, G, L, P). Kedimdjali,Koorders 24516 ß (BO); NW Moeriah, Doctersv.Leeuwen 997 (BO, L); Bantam, Afd.Menes,Dist.Batoehideung, Koorders 196 ß (BO, L);Res.Batavia, Backer 35432 (BO, L), vanSteenis 3055 (BO); Res. Batavia, G.Batoe,Bakh.v.d.Brink 3131 (BO); Tandjoeng, Backer35430 (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 nearKrawang, Koorders 13799 ß (BO); Res.


FIGURE 7. Wrightia pubescens subsp. lanitii (Blanco) Ngan (Leeuwenberg et al. 14567, A).

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), 27294ß (BO, L), 37066 ß (BO), 197 ß (BO, P), 201 ß(BO), 13481 ß (BO, L), 13800 ß (BO, L), 13801ß (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, Koorders34247 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 kmN of Labuhan, Leeuwenberg 13217 (L);Cempaka putih, Backer s.n. (L); Simpang,Bakh.v.d.Brink 4892 (L). SULAWESI: BikerveBalang, 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, 7km to Donggala, Meijer 9273 (L, MO);Sulawesi Tengah: W of Toboli, along road toPalu, Leeuwenberg et al 14567 (A); SulawesiTengah: Wera Waterfall near Parampadene, 25km 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), Ridley14950 (SING), Ridley 14951 (SING); Jitra,Burkill 13349 (SING); Langkawi: Pulau Tuba,Zainudin 5231 (L). Penang: Mainland, TasoekGlugor, Curtis 3738 (SING). Perak: LakeGardens, 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, Romeroet 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, MtMariveles, 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

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 PPI15071 (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, Elmer8069 (E); Mt. Makiling, Sulit 6875 (A),Whitford 19784 (L). Masbate: Bgy. Matipuron,Milagros, Barbon et al PPI 12598 (GH). NuevaEcija: 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); Merrillsp.Bl. 562 (A, BO, GH, L, MO, P); Ramos 2042(BO, BRI, L, P, NSW, SING); Robinson 6738(BRI, NSW). Zambales: Mt. Pinatubo, Balintay5626 (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 delSur: 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 fromPuerto 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 appearto be as different from ssp. pubescens as ssp.candollei, which has been resurrected as a


species distinct from Wrightia pubescens.Wrightia pubescens ssp. lanitii and ssp. pubes-cens differ in the size and shape of the sepalsand, for the most part, the pubescence of theantepetalous corona lobes. However, there areindividuals that have some pubescence on theinside of the corona lobes, and specimens fromSulawesi have the corona type and sepal shapeof ssp. pubescens but the sepal size of ssp. lan-itii. Because of these intermediates and thegreater similarity in corolla shape and sizebetween these two subspecies compared to ssp.candollei, they are being maintained as sub-species, whereas Wrightia candollei, which isalso 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, TreeFl. Mal. 2: 24. 1973; Ly, 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, designatedhere; isoneotype: ABD, K, TCD),Thailand, Chon Buri, Sri Racha.

Shrub or small tree to 5 m tall. Branchletspuberulent, soon becoming glabrous and

sparsely lenticellate. Leaves: petiole 1.5–4.0mm long; blade papery, elliptic or oblong,1.2–8.2 × 0.6–3.3 cm, 2.2–4.2 times as long aswide, apex acuminate, rarely to rounded, basecuneate, puberulent on midrib beneath andsometimes on midrib above, or completelyglabrous, blade punctate beneath and some-times above, 3–8 pairs of secondary veins,ascending, often obscure, tertiary venationobscure. Inflorescence 1.7–4.2 cm long;glabrous or sparsely puberulent; pedicels 8–30mm 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 toacute, 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 aslong as wide, obovate, apex rounded; puberu-lent-papillose on lobes outside and inside,glabrous on tube outside and inside; coronaabsent. Stamens inserted at corolla mouth,strongly exserted from tube; filaments 1.5–2.5mm long; anthers 3.8–4.0 × 0.7–0.8 mm,pubescent within and without. Gynoecium of 2free carpels united into a common style, ovaries0.8–1.1 mm high, glabrous, of two separatecarpels; 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; coma3.0–3.5 cm long.

Distribution: Thailand, Cambodia, possiblyPeninsular Malaysia. The natural distributionof this species is now obscure because it is sowidely cultivated, but is probably Thailand andmaybe Cambodia. It may also be native inPeninsular Malaysia, but is certainly widelycultivated in many parts of Malesia.


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 Wrightiatinctoria, a species otherwise known only fromIndia. Its appearance in Timor is a mystery andis likely to be either a mistake on the label or

else a cultivated plant transported from Indiaearlier. There are no notes on the label to indi-cate whether it was a cultivated plant. Untilsuch time as further material is collected inTimor that matches this specimen and its statuscan be reassessed, I am not including this as aMalesian species to be discussed further.


FIGURE 8. Wrightia religiosa (Teijsm. & Binn.) Benth. ex Kurz (Forman 47, A).


LITERATURE CITED

BAHADUR, K. N., AND S. S. R. BENNET. 1978. A newspecies of Wrightia R. Br. (Apocynaceae) fromIndia. Indian Forester 104: 32–35.

BRUMMIT, R. K., AND C. E. POWELL. 1992. Authors ofPlant Names. Royal Botanic Garden, Kew.

ENDRESS, M.E., AND P. V. BRUYNS. 2000. A revisedclassification of the Apocynaceae s.l. BotanicalReview 66(1): 1–56.

HOLMGREN, P. K., N. H. HOLMGREN, AND L.C.BARNETT.1990. Index Herbariorum. IAPT, New York.

LY, 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 newspecies 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 BotanicalGarden 52: 114–175.

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 & Idjan419 (4); Ahern’s collector 90 (7b), 1116 (1), 2960(7b), 3155 (1); Alambra 23206 (7b); Alston 16098(7a); Alvarez 12939 (7b), 18541 (7b); Ang FRI23347 (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); Barbonet al. PPI 12598 (7b), 18700 (4), 22357 (1); Barros26288 (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); Burkill13349 (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), LAE68795 (4); Cuming 1279 (7b), 1453 (1); Curran4510 (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.Leeuwen997 (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); Escritor21296 (1).

Fenix 1183 (1); Forbes 2766 (4), 2802 (4); Franco17856 (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); Hume8352 (4).

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); KochummenFRI 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 etal. 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 & KatikNGF 49910 (4); Leschenault 309 (7a); Leuterio24730 (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); Maxwells.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); MollBW 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), bb30447 (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),

47008 (7b), 85136 (7b); Ranario 27780 (7b);Reynoso et al. PPI 4486 (7b), PPI 17222 (7b), PPI24244 (4), PPI 24249 (3), PPI 24423 (4); Ridley5521 (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 & ChavezPPI 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 FSC832 (4); Spanoghe s.n. (7a); Stefels BW 5105 (4);Stone et al. 120 (2), PPI 262 (3); Streiman & KairoNGF 26050 A (4); Sulit 6055 (4), 6875 (7b), PNH

7428 (1), 14455 (4), 15715 (7b), 15746 (7b), PNH17044 (1), 22879 (7b), 25679 (7b); Sun Hong-FanHerb. 9064 (7a); Suppiah FRI 108885 (4);Symington 51751 (4), 57059 (4), 57060 (4),57061 (4).

Taylor 2731 B (7a); Teo & P. 548 (4); Teysmann HB5121 (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 etal. 1537 (7b); Verheijen 1005 (7a), 2296 (7a),4184 (7a), 4902 (7a); Versteegh BW 3991 (4), BW4628 (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); Widjaja1213 (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).


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 numbersabove. (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 Ly (2)Wrightia laevis Hook.f. (4)

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)

revision of wrightia

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