A REVISION OF KOPSIA (APOCYNACEAE: RAUVOLFIOIDEAE)

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1 A REVISION OF KOPSIA (APOCYNACEAE: RAUVOLFIOIDEAE) DAVID J. MIDDLETON 1 Abstract. The Asian genus Kopsia (Apocynaceae: Rauvolfioideae) is revised. Calpicarpum and Kentrochrosia are included in synonymy. Approximately 2000 herbarium specimens from 35 major herbaria have been studied. The genus is found from southern China and Burma to northern Australia and Vanuatu and is most diverse in Peninsular Malaysia and Borneo. Twenty-three species are recognized, two of which have two varieties each. Three new species are described: Kopsia grandifolia, Kopsia rosea, and Kopsia sumatrana. A key to the species is given, all taxa are described with notes on distribution and habitat, distribution maps are provided for all taxa, and all names and synonyms are typified. Keywords: Kopsia, Apocynaceae, Rauvolfioideae, revision, Asia, Australia. This work on Kopsia is the latest in a series of revisions of the Asian genera of Apocynaceae, subfamilies Rauvolfioideae and Apocynoideae. The genus contains arguably the most attractive species of any of the Asian genera of Apocynaceae, and a few of them have become widely cultivated. It is possibly also the most frustrating genus in the family in Asia for reasons clarified later. Most of the work for this revision has been done through the study of herbarium specimens, but a number of the species have also been seen in the field or in cultivation. Kopsia was published by Blume (1823) in honor of the Dutch botanist J. Kops, ( ) with one species, K. arborea Blume. He added another, K. vincaeflora Blume, in his Bijdragen (Blume, 1826) and a third, K. flavida Blume, in Blume (1849). De Candolle (1844) transferred Cerbera fruticosa Roxb. to Kopsia before the publication of the third of Blume s species, and the number of species then grew slowly to the present day with significant publications of new taxa by Hooker (1882), King and Gamble (1907), Ridley (1923), Merrill (1926, 1929), Pitard (1933), Timmerman-Van der Sleesen (1960), Markgraf (1973), Allorge (1993), Allorge and Teo (1986), Middleton (2003), and others. The only modern work on the entire genus was a synopsis of Kopsia in a paper mostly on the chemotaxonomy of the genus (Sévenet et al., 1994). Twenty-six species were recognized in this paper, and many of the taxa were effectively typified (but with extensive application of Article 9.8 of the International Code of Botanical Nomenclature [Greuter et al., 2000]). However, no key was given and many of the taxa were not described. Other important works include Timmerman-Van der Sleesen (1959) on the Malesian species, Li et al. (1995) on the Chinese species, and Middleton (1999) on the Thai species. Kopsia Blume from 1823 is a conserved name against the Orobanchaceae genus Kopsia Dumort. from Calpicarpum was published by Don (1837) with two species, C. roxburghii and C. lamarkii. Unfortunately both species are illegitimate under Article 52.1 of the Code (Greuter et al., 2000) because Cerbera fruticosa Roxb. was cited in synonymy of Calpicarpum roxburghii, the epithet of which should have been taken up, and several species of Cerbera were cited in synonymy of Calpicarpum lamarkii, one of which should have been taken up. De Candolle (1844) effectively typified the genus with C. roxburghii by excluding the only other original species when he synonymized the genus under Kopsia. I would like to thank the Arnold Arboretum and the Harvard University Herbaria, the Nationaal Herbarium Nederland, Leiden Branch, and the Muséum National d Histoire Naturelle in Paris for their support of this revision. I would also like to thank the directors and curatorial staff of the herbaria that hosted my visits or loaned material. I am grateful to David Goyder for his help in locating specimens at Kew, the Harvard University Herbaria staff for their practical support, Kanchi Gandhi for fruitful discussions on nomenclatural problems, Holly Nixon for the illustrations, Martin Pullan for help with the Pandora database, and Carroll Wood and Peter Stevens for helpful comments on the manuscript. 1 Arnold Arboretum, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. dmiddlet@oeb.harvard.edu Harvard Papers in Botany, Vol. 9, No. 1, 2004, pp President and Fellows of Harvard College, 2004.

2 90 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Kentrochrosia was published by Schumann and Lauterbach (1900) with one species, K. monocarpa. Merrill and Perry (1941) suggested that all the species of Kopsia with spurred fruits should actually be in Kentrochrosia and those with unspurred fruits should remain in Kopsia. They acknowledged that the genera were indistinguishable on flower characters. If interpreted strictly this would leave only Kopsia arborea in Kopsia and all the rest would be moved to Kentrochrosia. Merrill & Perry (1941) made only two new combinations in Kentrochrosia to add to the original species. Markgraf (1973) later acknowledged that the single fruit character was not only insufficient to distinguish the two genera but also suggested that the character was only one of degree rather than of a qualitative nature. I agree with this interpretation. The genus Kopsia is most diverse in Peninsular Malaysia and in Sarawak. In both regions there are a number of species with very restricted distributions, and for some of the species the collections are few. It is interesting to note that the diversity is not as high in other parts of Sundaland such as Sumatra or Kalimantan, but it should also be recognized that these regions are not as intensively collected, and one could speculate that more taxa with equally restricted distributions, over and above the new species K. sumatrana described below, could be discovered when more extensive collections are made. This revision of Kopsia has taken much longer than it ought to have for a genus with only 23 species because of the ease with which I have been able to set the work aside and do something less frustrating. This frustration has arisen because a large percentage of the available herbarium material, generally speaking, has been very poorly collected. Many of the specimens have no corollas but rather only the inflorescences with the ovaries and calyx. The flowers are extremely delicate, and unless the plant is pressed in the field the corollas will fall off. Even plants pressed immediately often make rather poor specimens, as the corollas are very thin and delicate and tear and fall off easily. Markgraf (1973) hints at a similar frustration in the introduction to his work on the Malesian species, and he therefore provided a key for plants with incomplete inflorescences. Also, the majority of collections made have inadequate notes. Only a small percentage note such characters as tree height and diameter or bark color and texture, and few even note corolla color or whether the eye is colored in otherwise white flowers. This eye character would appear to be a good character for recognizing some species such as K. flavida and K. singapurensis, where the eye is red in an otherwise white corolla. However, it is difficult to know for sure whether it is consistent in other species with a yellow eye such as K. pauciflora, where the eye does not have as much of a contrast with the white of the corolla, and most labels mention the color only as white with no specific mention of the eye. In some species like K. arborea, the color is consistently white without an eye. It will be interesting to test this revision against concerted field observation and better collections in the future to see if it will stand the test of time. In this regard there are many areas where more collections are needed, particularly in Vietnam, Kalimantan, and Sumatra. In the most recent subclassification of the Apocynaceae (Endress and Bruyns, 2000) Kopsia is placed in the tribe Vinceae, subfamily Rauvofioideae. This tribe, however, is only weakly supported as monophyletic, after the addition of fruit and seed characters to the molecular data, in an analysis by Potgieter and Albert (2001). Vegetative Characters Habit. All species of Kopsia are shrubs or small trees as part of the understory vegetation in forests, at forest edges, or in the open. The largest of any species recorded is 14 m tall for K. arborea. However, this is one of the commonest species, and the heights given in the descriptions must be judged against the lack of data recorded on many specimens, the small number of collections made for several of the MORPHOLOGY species, and the fact that it is easier to collect smaller individuals. It is quite probable that many of the species have individuals taller than in the descriptions given. Bark. Bark characters are very rarely recorded on herbarium material so again the colors given in the descriptions should not be considered prescriptive. Branchlets. Most species of Kopsia have young branchlets that are somewhat angled.

3 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 91 This may be an artefact of drying in some, but not all, cases. In some species the angles become so pronounced as to become wing-like. Markgraf (1973) recognized a new species, K. lancifolia, based in part on this character. However, it is somewhat subjective as to where one draws the line between markedly angled and winged, and all stages in-between are manifest in some species, particularly in both varieties of K. pauciflora. Leaves. The leaves are always opposite. They may or may not be sessile, a character which is somewhat subjective as many species have very short petioles. However, in only two species, K. teoi and K. pauciflora, are there cases where the petioles do appear to be absent. In K. pauciflora this is of rare occurrence. The bases of the petiole weakly clasp the node but without the development of distinct ocrea. When young, the petiole bases envelop the terminal bud. Small colleters are present in the leaf axils. The blade is always entire and mostly elliptic. The venation is pinnate. The number of veins, the angle of the secondary veins with the midrib, and how distinct the secondary and tertiary venation are from each other are useful characters in some species. Reproductive Characters Inflorescence. Inflorescence structure has been discussed by Markgraf (1973). The basic structure is a terminal dichasium or sometimes a trichasium. When the inflorescence is young this is the predominant appearance in more or less all species. Some species keep this structure even as they develop and mature (e.g., K. griffithii, K. sleeseniana, K. tenuis, Fig. 6), often becoming somewhat thyrsoid (e.g., K. arborea, Fig. 2). In other species, the dichasial branches do not further branch and instead form cincinnate monochasia with flowers opening sequentially one at a time (e.g., K. grandifolia, K. macrophylla, K. pauciflora, K. rajangensis, Fig. 5). In several of the species with laxer, more dichasial and open inflorescences, the ultimate branches are short and the flowers congested (e.g., K. singapurensis, Fig. 7). Markgraf (1973) has suggested an evolutionary progression from the thyrsoid type in K. arborea to loose di- and trichasia, to di- or trichasia with cincinnate branches, to cincinnate branches only. There is a subtending bract at the base of each pedicel and most species have further bracts on the pedicels, but this is variable in some species, and only in K. teoi, with several bracts on the pedicel, is this a useful taxonomic character. Although the inflorescence is almost always terminal, very rarely are there also axillary inflorescences (e.g., in K. rajangensis). Sometimes the peduncle is very short before the inflorescence branches, giving the appearance of several inflorescences emerging from a single apex. In the taxon descriptions the term inflorescence axes is used to include the peduncle plus the branches of an inflorescence. Calyx. The sepals are mostly ovate, tending toward oblong. The apex can be acuminate to rounded, and this character is useful for recognizing species. All species have a small-tolarge gland just below the apex of the sepal. This gland is sometimes decayed or eaten, giving the sepal a secondarily retuse apex. There are no colleters in the sepal axils. Corolla. The basic salverform structure of the corolla is rather invariable in the genus. The tube is always slightly dilated around the stamens. The corolla lobes are dextrorsely contorted in bud, a character rather unusual in the subfamily Rauvolfioideae but invariable within the genus. They are held perpendicular to the tube in the open flower and can vary by as much as 25% in size within an individual flower. As has been discussed earlier, the corolla color is often not recorded on herbarium specimens, but in all known cases the flowers are showy and open during the day. The appearance would suggest butterfly pollination, although I am unaware of any studies having been conducted in the genus. Useful taxonomic characters include the overall dimensions of the corolla, the color and the shape of the corolla lobes, and the color of the area where the lobes join the tube, in this work referred to as the eye. Stamens. The stamens, as is the case for most members of subfamily Rauvolfioideae, are rather unspecialized. The filament is short and the anthers ovate and without lignified guide rails at the edges. They are attached to the inside of the corolla tube just above the height of the style head. The only taxonomically useful character in the stamens is their position in the corolla tube. Disk. There are two disk lobes alternating with the carpels. In almost all species they are more or less awl-shaped or simply oblong, with an acute or acuminate apex. In K. arborea they are more variable within the species and can be somewhat more complex in shape (see description below). Only in K. griffithii is the disk pubescent. Gynoecium. The gynoecium is of two ovaries, which are free from each other but are then

4 92 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 united into a common filiform style. The style head has a basal collar and a short cylindrical apex. The degree of pubescence, or otherwise, of the ovaries is a useful taxonomic character. Each carpel contains two ovules, although only one ever develops to maturity. The receptive stigmatic surface is on the sides of the style head. Fruit. The fruit of all species is a small drupe, less than 4.5 cm long, usually paired although sometimes with one aborting, especially in Kopsia arborea. In K. arborea the fruit is ellipsoid or slightly falcate. This is in contrast to all other species, where the falcate fruit has a spur, or at least a sharp angle, on the ventral side. Sévenet et al. (1994) suggest that there is an evolutionary progression from the fruit without an appendage to the large appendages of K. pauciflora and K. macrophylla. The largest appendages, however, are to be found in K. flavida, where it can be up to 13 mm long. No analyses have been done to examine these evolutionary theories more closely. Markgraf and Huber (1975) give a detailed description of the development of the spur in K. flavida. In a few species the fruit is not known, but in those where it is the shape of the fruit and spur is a useful taxonomic character. The color of the fruit is unknown in most species. Other Characters The paper by Sévenet at al. (1994), in which a synopsis of the genus Kopsia was given, was primarily a paper on the chemotaxonomy of the genus. They noted that alkaloids have long been known in the genus and went into some detail on the chemical structures and rearrangements possible. They then presented a table showing the alkaloid compounds that have been isolated from 15 species of the genus (only 12 recognized in this new revision) and suggested a number of relationships between taxa. I find only a couple of these suggestions likely. In particular they recognize K. arborea, K. pitardii, and K. jasminiflora as separate species, give no suggestions as to the chemical affinities of K. arborea but link each of the other two taxa to K. pauciflora and K. dasyrachis, respectively. It is clear to me that K. arborea, K. pitardii, and K. jasminiflora are synonymous. The suggestions that K. singapurensis and K. teoi are close, and K. dasyrachis and K. macrophylla are related, would, however, appear to be borne out by the morphology. These issues can be resolved only by a more thorough phylogenetic analysis, preferably also using molecular data. Chromosome numbers are known for K. arborea (n = 36; 2n = 72) and K. fruticosa (n = 18; 2n = 36) (see Van der Laan and Arends, 1985). In neither the Leeuwenberg classification system followed by Van der Laan and Arends (1985), nor by comparison to the Endress and Bruyns (2000) system, is there basic chromosome number consistency within the tribes in which Kopsia has been included. Approximately 2000 herbarium specimens from Asia and Malesia have been studied. These are from the following herbaria: A, AAU, ABD, AMES, B, BISH, BKF, BM, BO, BR, BRI, C, CAL, CANB, E, F, FU, G, GH, IBSC, K, L, M, MICH, MO, NY, P, PE, PNH, SING, TCD, U, UC, US, Z (Holmgren et al., 1990). All specimens have been seen unless otherwise indicated. All dimensions given are for dried specimens except for androecium and gynoecium characters, which are from flowers reconstituted by MATERIALS AND METHODS boiling in water. The number of flowers dissected is almost always fewer than the number of flowers measured for corolla tube and lobe dimensions. Therefore, it should be noted that dimensions given for characters like stamen insertion are based only on the rehydrated flowers that were dissected, leading to a few apparent discrepancies with the range of variation in the dried flowers. Author citation follows Brummitt and Powell (1992). Kopsia Blume, Catalogus , nom. cons.; G.Don, Gen. Hist. 4: ; A.DC. in DC., Prod. 8: ; Benth. & Hook.f., Gen. Pl. 2: ; Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2, 27: Type species: Kopsia arborea Blume TAXONOMIC TREATMENT Synonyms: Calpicarpum G.Don, Gen. Hist. 4: Type species: Calpicarpum roxburghii G.Don, nom. illeg. Kentrochrosia K.Schum. & Lauterb., Fl. Deutsch. Südsee Type species: Kentrochrosia monocarpa K.Schum. & Lauterb.

5 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 93 Shrubs or small trees, buttresses absent. Branchlets terete to winged; glabrous or more rarely pubescent; lenticellate or not. Leaves opposite; mostly petiolate, rarely sessile, bases mostly clasping the terminal bud when young and the stem when older; blade mostly subcoriaceous to coriaceous, rarely papery, the opposite pair more or less equal in size, entire; colleters present in the axils; secondary venation pinnate, clearly distinguishable or not from the tertiary venation, often anastomosing into an intramarginal vein. Inflorescences terminal, very rarely also with some axillary; basically dichasial, more rarely trichasial, but often with elongated branches that do not further branch so as to appear somewhat cincinnate; pedunculate or not; glabrous to densely pubescent; subtending bracts and pedicel bracts small. Flowers 5-merous. Sepals erect, mostly ovate, sometimes somewhat oblong; without colletors in the axils; with a gland on the outside just below the apex. Corolla actinomorphic; lobes dextrorsely contorted in bud; salverform with a narrow tube, slightly wider around the stamens, and spreading lobes; tube generally pubescent inside around the anthers and more densely in the throat, sometimes glabrous in throat, mostly glabrous outside or rarely with a few hairs on the lobes and top of tube. Stamens inserted around the middle of the tube to near the tube throat, very rarely near base, not exserted from throat; filaments straight, short, thin; anthers ovate, fertile for most of length; free from style head. Disk of two lobes alternating with the two free carpels. Gynoecium 2-carpellate, apocarpous but apically united into a common style, glabrous or pubescent; style filiform, glabrous; style head with a collar at the base, otherwise short and cylindrical. Ovules 2 in each carpel although only 1 ever develops. Fruits of paired drupes, more rarely solitary, ellipsoid to falcate, more or less flattened (hardly at all in K. arborea), usually with spurlike appendages facing inward toward each other (except in K. arborea). Seed curved, broader at one end, other end acuminate. KEY TO THE SPECIES OF KOPSIA 1a. Stamens inserted very clearly around the middle or in lower half of corolla tube, 0.6 of corolla tube length b. Stamens inserted in upper half of corolla tube, > 0.6 of corolla tube length a. Sepals acute or acuminate, rarely obtuse but never rounded b. Sepals rounded a. Sepals acute or obtuse; secondary and tertiary veins clearly distinguishable; Peninsular Malaysia K. profunda 3b. Sepals acuminate; secondary and tertiary veins not clearly distinguishable; Vietnam K. harmandiana 4a. Inflorescences robust, glabrous to densely pubescent; branchlets glabrous to densely pubescent, terete to very weakly angled; Peninsular Malaysia K. macrophylla 4b. Inflorescences delicate, glabrous; branchlets glabrous, weakly angled to strongly winged; Borneo K. pauciflora var. mitrephora 5a. Sepals acuminate; inflorescence congested; corolla completely white; Thailand, Laos K. angustipetala 5b. Sepals emarginate to acute; inflorescence variable; corolla color variable; widespread a. Corolla tube 20 mm long b. Corolla tube > 20 mm long a. Branchlets sparsely to densely puberulent, glabrescent when older; petiole sparsely to densely pubescent; sepals densely puberulent b. Branchlets glabrous; petiole glabrous; sepals glabrous to sparsely puberulent a. Secondary veins pairs; corolla lobes times as long as wide K. grandifolia 8b. Secondary veins pairs; corolla lobes ca. 8.2 times as long as wide K. sumatrana 9a. Inflorescence lax with clear internodes, sparsely to densely puberulent; pedicels mm long b. Inflorescence with flowers clustered along short or elongated branches, glabrous or puberulent only in upper parts; pedicels 1 2 mm long a. Petiole 5 7 mm long; pedicels mm long; corolla tube throat pubescent, corolla lobes ca. 9.5 mm long; disk pubescent; ovaries densely pubescent K. griffithii var. pubescens 10b. Petiole 1 2 mm long; pedicels mm long; corolla tube throat glabrous, corolla lobes mm long; disk glabrous; ovaries glabrous K. tenuis 11a. Leaf apex caudate, secondary veins clearly distinguishable from tertiary venation, curved ascending; corolla white, lobes mm wide K. larutensis 11b. Leaf apex short acuminate, secondary veins not clearly distinguishable from tertiary venation, straight; corolla red, lobes mm wide K. lapidilecta 12a. Leaves sessile b.Leaves petiolate

6 94 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 KEY TO THE SPECIES OF KOPSIA CONT. 13a. Corolla white and tinged pink or pink with a red eye ; anther apex mm from corolla throat K. teoi 13b. Corolla completely white or white with a yellow eye, rarely pinkish-white; anther apex mm from corolla throat K. pauciflora 14a. Disk pubescent; inflorescence lax with distinct internodes K. griffithii 14b.Disk glabrous; inflorescence variable a. Petioles slender, 6 17 mm long, usually around 15% of total leaf length; inflorescence branches < 1 mm wide K. deverrei 15b. Petioles stout, 3 12( 15) mm long, less than 15% of total leaf length; inflorescence branches > 1 mm wide, rarely less but then corollas not white with a red eye a. Anther apex > 5 mm from corolla throat; inflorescences > 10 cm long, densely puberulent K. dasyrachis 16b. Anther apex 5 mm from corolla throat; inflorescences length variable but mostly < 10 cm long, puberulent or not a. Inflorescences with subtending bracts as large as the sepals and flowers clustered at the ends of the inflorescence branch; sepals mostly acute, more rarely obtuse; fruit without a spur or angle K. arborea 17b. Inflorescences with subtending bracts somewhat obscure or smaller than sepals, if as large as sepals then flowers not clustered at ends of the inflorescence branch; sepals obtuse to rounded; fruit with a spur or angle on one side a. Inflorescence axes 1 cm long; corolla tube < 23 mm long; Hainan K. hainanensis 18b.Inflorescence axes > 1 cm long, rarely less but then corolla tube > 25 mm long; corolla tube mostly > 23 mm long, rarely less but then axes > 1 cm long; not in Hainan a. Ovaries glabrous or with just a few isolated hairs b.Ovaries clearly pubescent a. Leaves with long caudate apex; secondary veins at to midrib; Borneo K. rajangensis 20b.Leaves acuminate; secondary veins at (45 )60 75 to midrib; not in Borneo a. Stamens inserted at < 0.8 of corolla tube length; Vietnam K. tonkinensis 21b.Stamens inserted at 0.8 of corolla tube length; southern Thailand and Malesia a. Stamen bulge slightly below top of corolla tube; flowers white or pinkish but not white with a strongly contrasted pink or red eye ; southern Thailand and peninsular Malaysia K. rosea 22b. Stamen bulge at top of corolla tube; flowers white with a strongly contrasted pink or red eye ; Philippines, eastern Malesia, western Pacific Islands K. flavida 23a. Inflorescence robust and lax with distinct internodes all through inflorescence K. sleeseniana 23b.Inflorescence robust or delicate with flowers congested, if lax then at least with ultimate branches congested a. Inflorescences without elongated branches; corolla pale to dark pink or white with a red or pink eye b. Inflorescences with elongated branches but with the flowers congested along these; corolla white or white with a yellow eye, very rarely with a pink tinge but then only ever one flower on an inflorescence branch open at a time a. Branchlets pubescent; native in Burma but widely cultivated K. fruticosa 25b.Branchlets glabrous; Malesia a. Peduncle to 1 cm long; fruit with a large deltoid or very slightly hooked projection on one side; Philippines, eastern Malesia, western Pacific Islands K. flavida 26b.Peduncle mostly > 1 cm long, very rarely under 1 cm; fruit with only a slight angle on one side; Peninsular Malaysia K. singapurensis 27a. Secondary veins at (55 )60 80 to midrib; anther apex mm from corolla throat K. pauciflora 27b. Secondary veins at to midrib; anther apex mm from corolla throat K. rajangensis 1. Kopsia angustipetala Kerr, Kew Bull. 1937: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: THAILAND. Nong Khai: Chaiyaburi, A.F.G. Kerr (Lectotype: K, designated by Sévenet et al. [1994]; Isolectotypes: BM, P, TCD). Map 1; Fig. 1. Tree to 5 m tall. Branchlets glabrous to densely puberulent, becoming glabrescent when older, sparsely lenticellate or not, weakly angled. Leaves: petiole 1 7 mm long, glabrous; blade cm, times as long as wide, papery, subcoriaceous to coriaceous, elliptic, oblong or obovate, apex caudate or long acuminate with a blunt tip, base cuneate, glabrous above and beneath, midrib shallowly sunken or raised above, secondary veins pairs with 2 8 mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable from tertiary venation or not above, clearly

7 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 95 MAP 1. Distribution of Kopsia angustipetala Kerr. distinguishable beneath, straight, tertiary venation prominent above or flat above and beneath, obscure or irregularly parallel to secondary veins, intramarginal vein straight or only weakly looped, inset from margin. Inflorescence dichasial, cm long with axes cm long and branches mm wide, glabrous to densely puberulent; peduncle cm long, mm wide, glabrous or puberulent; pedicels mm long, glabrous or densely puberulent, subtending bracts persistent, bracts present on pedicel. Sepals mm, times as long as wide, ovate or lanceolate, apex acuminate, ciliate or not, glabrous to densely puberulent outside, glabrous or puberulent on upper half inside. Corolla completely white; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent in upper part of tube above, around, and slightly below the stamens, throat glabrous, glabrous outside; lobes mm, times as long as wide, elliptic, apex acute to acuminate, not ciliate, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk 1 mm long, times as long as ovaries, glabrous, awlshaped or lanceolate, apex acute or acuminate. Ovaries mm high, sparsely pubescent on top; style 12.4 mm long; style head 0.6 mm long. Fruit falcate with a small blunt hooked spur, mm, spur 2 mm long, sparsely puberulent. Distribution: Thailand (Nong Khai Province), Laos (Vientiane province). Habitat: dry evergreen or deciduous forest at m altitude. Additonal collections studied: LAOS: Vientiane Province: Houei Hin Kanna Waterfall, 25 km N of Vientiane, Iwatsuki et al. IC (A); Nasaithong, Wongprasert et al. s.n. (BKF). THAILAND: Route 211 near Loei border, Murata et al. T (BKF); Phu Wua Wildlife Sanctuary, Pooma 1593 (A, BKF); Tham Phoon Falls, Wongprasert s.n. (BKF); Tadkham Falls, Wongprasert et al. s.n. (BKF). This is a very distinctive species with small white flowers, corolla lobes with acuminate to acute apices, and a congested inflorescence with acuminate sepals and bracts.

8 96 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 FIGURE 1. Kopsia angustipetala Kerr. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit. From Pooma 1593 (A, D) and Iwatsuki et al. IC 1594 (B, C). Scale bars = 1 cm (A, B, D); 1 mm (C).

9 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) Kopsia arborea Blume, Catalogus ; G.Don, Gen. Syst. 4: ; A.DC., Prod. 8: ; Miq., Fl. Ind. Bat. 2: ; Markgr., Bot. Jahrb. 61: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Markgr., Blumea 20: ; Anderson, Checklist Trees Sarawak ; P.S.Ashton, Manual Nondipt. Trees Sarawak ; Goel & Sharma, Higher Pl. Ind. Subcont. 1: ; Gangop. & Chakr., J. Econ. Tax. Bot. 16: ; Sévenet et al., J. Ethnopharmacol. 41: ; Forster, Fl. Aust. 28: ; Beaman et al., Plants Mount Kinabalu 4: TYPE: INDONESIA. Java, Mt. Salak, C.L. Blume s.n. (Lectotype: L [ ], designated by Timmerman-Van der Sleesen [1959]; Isolectotype: L [ ]; possible Isolectotype: NY). Map 2; Fig. 2. Synonyms: Kopsia longiflora Merr., Philipp. Gov. Lab. Bur. Bull. 29: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: PHILIPPINES. Luzon, Bataan Province, Lamao River, T.E. Borden 611 (Holotype: PNH, destroyed; Lectotype: US, designated here; Isolectotypes: BM, K, NY; probable unnumbered Isolectotype: UC). Kopsia scortechinii King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Goel & Sharma, Higher Pl. Ind. Subcont. 1: ; Gangop. & Chakr., J. Econ. Tax. Bot. 16: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Perak sine loc., B. Scortechini 1878 (Lectotype: SING, designated by Timmerman-Van der Sleesen [1959]; Isolectotype: CAL). Kopsia laxinervia Merr., Phil. J. Sci., Bot. 13: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: PHILIPPINES. Luzon, Apayao, E. Fénix (Lectotype: A, designated here; Isolectotypes: BO, K, P, NY, US). Kopsia lancibracteolata Merr., Phil. J. Sci. 23: ; Tsiang, Sunyatsenia 2: ; Lý, Feddes Repert. 97: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: CHINA. Hainan: near Ka La, F.A. McClure 9183 (Lectotype: K, designated by Sévenet et al. [1994]; Isolectotypes: A, BM, ECON, MO, P, PNH, UC). MAP 2. Distribution of Kopsia arborea Blume.

10 98 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 FIGURE 2. Kopsia arborea Blume. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit. From Hu (A, B, C) and Kostermans 7673 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).

11 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 99 Kopsia jasminiflora Pitard, Fl. Gén. Indo-Chine 3: ; Kerr, Fl. Siam. Enum. 2: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: LAOS. Sine loc., Dussaud 114 (Holotype: P; Isotype: P). Kopsia pitardii Merr., Contr. Arnold Arbor. 8: ; Sévenet et al., J. Ethnopharmacol. 41: Kopsia cochinchinesis auct. non Kuntze: Pitard, Fl. Gén. Indo Chine 3: ; Lý, Feddes Repert. 97: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: VIETNAM. Dong Nai, Mt. Dinh, J.B.L. Pierre 32 (Lectotype: P, designated by Sévenet et al. [1994]; Isolectotypes: A, BR, F, HNU [n.v]., K, L, P, MICH, MO, NY, SING, US). See comment below. Kopsia pruniformis Reichb.f. & Zoll. ex Bakh.f., Blumea 6: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: INDONESIA. Java Timur, Rogodjampi, H. Zollinger 3832 (Lectotype: NY, designated by Sévenet et al. [1994]; Isolectotypes: K, L, P). Kopsia officinalis Tsiang & P.T.Li, Acta Phytotax. Sin. 11: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: CHINA. Yun-Ching-Hung, (=Junjinghong), Yunnan, Y.-M. Ting 54 (Holotype: CANT; Photographs: A, P). Tree to 14 m tall, to 30 cm dbh. Bark gray; inner bark pale brown. Branchlets glabrous or sparsely puberulent when young, sparsely lenticellate or not. Leaves: petiole 3 10 mm long, glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, elliptic, apex caudate to acuminate with a blunt tip, base acute or cuneate, glabrous above and beneath, midrib sunken to raised and with a central groove above, secondary veins 9 18 pairs with 4 13 mm spacing, from midrib, prominent above and beneath, clearly distinguishable from tertiary venation above and beneath, straight or ascending near margin, tertiary venation prominent or flat above and beneath, obscure or irregularly subperpendicular to midrib and oblique to secondary veins, intramarginal vein strongly looped. Inflorescence dichasial, cm long with axes cm long and branches mm wide, glabrous to sparsely puberulent; peduncle cm long, mm wide, glabrous or puberulent in upper parts; pedicels 0 5 mm long, glabrous to densely puberulent, subtending bracts persistent, bracts present on pedicel. Sepals mm, times as long as wide, ovate, lanceolate or oblong, apex obtuse to acute, ciliate, glabrous to sparsely puberulent outside, glabrous or puberulent on upper half inside. Corolla completely white; tube mm long, mm wide, times as long as lobes, times as long as calyx, glabrous to sparsely pubescent around stamens and slightly beneath inside, glabrous or sparsely puberulent at top of tube outside; lobes mm wide, times as long as wide, elliptic or oblong, apex rounded to obtuse, ciliate or ciliate only at lobe base, inside glabrous or sparsely pubescent in upper quarter, outside glabrous. Stamens inserted mm from corolla base which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long. Disk mm long, times as long as ovaries, glabrous, oblong, hourglass-shaped, or awl-shaped, apex shape variable and often quite complex, ranging from simply acuminate to rounded to horizontally V-shaped and flat on top or horizontally V-shaped and retuse on top. Ovaries mm high, glabrous to sparsely pubescent all over; style mm long; style head mm long. Fruit oblique ellipsoid or subglobose, blue-black, mm, only one carpel developing (see note), spur absent, glabrous. Distribution: Andaman and Nicobar Islands, southern China, Thailand, Vietnam, peninsular Malaysia, Borneo, Sumatra, Java, Lesser Sunda Islands, Sulawesi, Philippines, Queensland. Habitat: grows in a wide range of forest types as an understory tree and at forest margins on a wide range of soil types from sea level to 1500 m altitude. Geographical selection of the 185 collections studied: ANDAMAN AND NICOBAR ISLANDS: South Andaman, Prain s.n. (A, M, US); Great Nicobar Island: Dhar Bay, Hore 8750 (L). AUSTRALIA: Mossman Gorge, ca. 3 miles SW of Mossman, Schodde 4170 (A, BRI, CANB, L). CHINA: Guangdong: Guangzhou, Yip 153 (BKF). Hainan: Yaichow, How (A, K, NY, P, US). Hong Kong: Kadoorie Farm, Hu (A, K, PE). INDONESIA: Bali:

12 100 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Brambang, Sarip 14 (L). Java: Java Barat Province: Udjung Kulon Nature Reserve, Peutjang Island, Kostermans 47A (A, CANB, G, K, L, SAR). Kalimantan: Kalimantan Timur Province, Balikpapan, Kostermans 7673 (A, BO, K, L). Lesser Sunda Islands: Flores, Manggarai, Golo Sengang, Schmutz 2750 (L). Sulawesi: Sulawesi Utara Province, Sungai Ilanga, 220 km W of Manado, ca. 50 km inland from Pangi, Burley, Tukirin et al (A, BISH, E, K, KEP, L, MO). Sumatra: Lampung Province: Mt. Tanggamus, Jacobs 8043 (A, BISH, K, KEP, L, SING), Jacobs 8072 (A, BISH, K, L, SING). MALAYSIA: Penang: Waterfall Gardens, Haniff 3667 (BRI, SING, A). Sabah: Mt Kinabalu, Tenompok, Clemens & Clemens (A, BM, BO, L, K, NY, UC). Sarawak: Gunong Subis, Anderson S (K, L, SAN, SAR, SING). PHILIPPINES: Romblon, Elmer (A, BISH, BM, BO, E, G, L, MO, NY, US, Z). Mindoro Oriental: Subaan River, inland from San Teodoro, Ridsdale 1272 (A, BO, SAN). Negros Oriental: Lake Balinsasayao, Reynoso, Fuentes & Garcia 992 (GH). Leyte: Mt. Suiro, Sulit (L, SING). THAILAND: Chiang Mai Province: Ban Kong He, Kerr 3572 (ABD, BM, K). Nakhon Si Thammarat Province, Khao Luang, Beusekom & Phengkhlai 901 (BKF, K, L, P). VIETNAM: Binh Tri Thien, Hue, Mii Bach Ma Station, Poilane (P). Typification: King and Gamble (1907) typified Kopsia scortechinii with Scortechini 1878 collected in Perak. Timmerman-Van der Sleesen (1959) further lectotypified the name with the SING duplicate. Sévenet et al. (1994) gave the following typification for this name: Scortechini 57 b, 1878, holo-bm!, iso- SING! It would appear that they thought that the two collections were duplicates and that 1878 was a date, not a collecting number. However, it is clear from the style of the labels and the known collecting dates of Scortechini that 57b and 1878 are separate collections and that only 1878 could be a type. In addition, King and Gamble list only 1878 in the protologue and 57 is listed as a syntype of K. larutensis, a specimen of which it undoubtedly is. The type material of K. scortechinii from CAL is clearly synonymous with K. arborea. Some further confusion has occurred through Markgraf s (1973) distinction of K. scortechinii from K. arborea, partly based on the former s Ovary with long white hairs, even though his second key suggests it has a glabrous ovary, which would agree with the specimens and the original description. In most species of Kopsia, the two carpels both develop into paired fruits, with the hooks facing back toward each other. Sometimes only one of the carpels develops. In K. arborea, however, it would appear from the herbarium specimens that only one of the two carpels ever develops into a fruit. Although there is abundant herbarium material of this species, it should be borne in mind that the fruits are often fallen ones placed in a packet, so this statement should be treated with caution. The collection Kostermans 7673 has somehow become confused, and specimens with this number belong to both Alyxia reinwardtii and Kopsia arborea. The name Kopsia pitardii was applied by Merrill for Pitard s concept of K. cochinchinensis Kuntze, the type of which is a specimen of Tabernaemontana divaricata. Although Merrill referred to it as a nom. nov., it is actually a new species by Merrill, validated by reference back to Pitard s description of K. cochinchinensis. Sévenet et al. (1994) have effectively lectotypified the Paris element of the collection Pierre 32. In Paris there are three specimens of Pierre 32, but only one of them is labelled as K. cochinchinensis, the name taken up by Pitard (1933), on which K. pitardii was based. Therefore, it is unnecessary to make a second step lectotypification for this particular specimen. 3. Kopsia dasyrachis Ridl., Kew Bull. 1934: ; Masam., Enum. Phan. Born ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: MALAYSIA. Sabah, Lukan, Arsat 1211 (Holotype: K; Isotype: K; Photograph: A). Map 3; Fig. 3. Tree to 10 m tall, to 46 cm dbh. Bark gray or yellow, smooth; inner bark white. Branchlets glabrous, sparsely lenticellate, weakly winged. Leaves: petiole 5 7 mm long, glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, mostly drying reddish-brown, elliptic, apex long acuminate with a blunt tip, base acute to cuneate, glabrous above and beneath, midrib shallowly sunken or raised and with a central groove above, secondary veins 9 16 pairs with

13 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 101 MAP 3. Distribution of Kopsia dasyrachis Ridl mm spacing, from midrib, prominent to sunken above and prominent beneath, clearly distinguishable from tertiary venation above and beneath, straight, ascending near margin or curved ascending from midrib, tertiary venation prominent or flat above and beneath, obscure or irregularly subperpendicular to midrib and oblique to secondary veins, intramarginal vein straight, weakly or strongly looped, inset from margin. Inflorescence with dichasial branching followed by cincinnate branches, cm long with axes 7 16 cm long and branches mm wide, densely puberulent; peduncle cm long, mm wide, puberulent; pedicels ca. 1.2 mm long, densely puberulent, subtending bracts persistent. Sepals mm, times as long as wide, oblong, apex rounded to obtuse, ciliate, densely puberulent outside, glabrous inside. Corolla completely white; tube mm long, 1.4 mm wide, times as long as lobes, times as long as calyx, pubescent around and beneath stamens and in throat, glabrous or sparsely puberulent at top of tube outside; lobes mm, times as long as wide, elliptic, apex rounded, ciliate only at lobe base, glabrous outside and inside. Stamens inserted mm from corolla base, which is ca. 0.7 of corolla tube length in the rehydrated flowers measured; anthers mm wide, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, awlshaped, narrowly deltoid or lanceolate, apex acuminate. Ovaries mm high, densely pubescent on top or densely pubescent all over; style mm long; style head mm long. Fruit sparsely puberulent, with a blunt hooked spur, mm, spur mm long. Distribution: Borneo (Sabah). Habitat: in a variety of evergreen forest types, often on sandy soils from m altitude. Selection of the 83 collections studied: MALASIA. Sabah, Lahad Datu District, Ulu Segama, Argent et al (E, K, KEP, L, SAN, SAR); Danum Valley, Madani SAN (K, L, SAN). Possibly related to Kopsia pauciflora but differing from that species in the stamens being situated somewhat lower in the corolla tube and in the generally much more densely pubescent and often much longer inflorescences. In some areas in Sabah, such as in Danum Valley, it would appear to be the most common species.

14 102 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 FIGURE 3. Kopsia dasyrachis Ridl. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit. From Gibet (A, B, C) and Enggoh (D). Scale bars = 1 cm (A, B, D); 1 mm (C).

15 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) Kopsia deverrei L.Allorge, Phytologia 59: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Johor, 87 km milestone from Mersing to Johor Bahru, Deverre 25 (Holotype: P; Isotype: P). Map 4. Tree to 10 m tall. Branchlets glabrous, sparsely lenticellate or not, terete or weakly angled. Leaves: petiole 6 17 mm long, glabrous; blade cm, times as long as wide, papery or subcoriaceous, elliptic, apex caudate or short to long acuminate with a blunt tip, base acute or cuneate, glabrous above and beneath, midrib shallowly sunken above, secondary veins 7 11 pairs with 3 15 mm spacing, from midrib, prominent to sunken above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, ascending near margin or curved ascending from midrib, tertiary venation prominent above, prominent or flat beneath, subperpendicular to midrib and oblique to secondary veins and also somewhat reticulate, intramarginal vein straight to strongly looped, inset from margin. Inflorescence dichasial, cm long with axes cm long and branches mm wide, glabrous; peduncle cm long, mm wide, glabrous; pedicels mm long, glabrous or sparsely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate, apex rounded to obtuse, ciliate, glabrous outside and inside. Corolla white with a red eye ; tube mm long, 2 mm wide, times as long as lobes, times as long as calyx, pubescent in upper part of tube above, around, and slightly below the stamens, throat pubescent, glabrous outside; lobes mm, times as long as wide, elliptic or oblong, apex rounded, ciliate, pubescent at very base of lobes, glabrous outside. Stamens inserted mm from corolla base, which is ca. 0.9 of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, mm from corolla throat; filaments ca. 0.8 mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, awl-shaped, apex acute or acuminate. Ovaries ca. 0.9 mm high, densely pubescent on top; style mm long; style head mm long. Fruit unknown. Distribution: Peninsular Malaysia (Johor). Habitat: from 150 to 305 m altitude. MAP 4. Distribution of Kopsia deverrei L.Allorge.

16 104 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Additional collections studied: MALAYSIA: Johor: Sungai Kayu, Mawai Jemulang Road, Corner SFN28680 (K, L, SING); Bukit Tingau Laut, Corner SFN37067 (K, L, SING); Gunung Panti, Stone, Chew 6230 (L, MO). In this species, the corolla tube is particularly densely pubescent behind the anthers. Specimens of this species were included in Kopsia singapurensis by Timmerman-Van der Sleesen (1959) and by Markgraf (1973), although Timmerman-Van der Sleesen expressed her doubt that they belonged there. This species is close to K. singapurensis, differing from it in the remarkably long petioles as compared with the length of the leaf blade, the more delicate inflorescences, and the shorter, more obtuse sepals. However, there are also some specimens of K. singapurensis, most notably Hamid (KEP), with the shorter petioles, large leaves, and venation patterns of K. singapurensis but with the smaller flowers approaching those of K. deverrei, which somewhat bridge the gap between them. I have maintained K. deverrei here, as the material of this species is readily recognized by the long petioles and different leaf size and shape. However, more-detailed population studies of the two species may lead to a reevaluation of the distinctions between them. 5. Kopsia flavida Blume, Rumphia 4: ; Merr., Phil. J. Sc. 29: ; Miq., Fl. Ind. Bat. 2: ; Corner, Wayside Trees Malaya 1: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: NEW GUINEA. Sine loc. [but likely West Papua], Zippelius s.n. (Lectotype: L [ ], designated by Timmerman-Van der Sleesen [1959]; Isolectotypes: L [ , ]). Map 5; Fig. 4. Synonyms: Calpicarpum albiflorum Teijsm. & Binn., Tijdschr. Nederl. Ind. 25: TYPE: INDONESIA. Seram, J.E. Teijsmann HB5035 (Lectotype: BO, designated by Sévenet et al. [1994]; Isotype: L). Kopsia albiflora (Teijsm. & Binn.) Boerl., Handl. Fl. Ned. Ind. 2: ; Sévenet et al., J. Ethnopharmacol. 41: ; Kopsia fruticosa var. albiflora (Teijsm. & Binn.) King & Gamble, J. As. Soc. Beng. 4(2): Calpicarpum ornatum W.Bull, Retail List 199: ; Fraser & Hemsl., Johns Gard. Dict. new ed ; Mabberley, Taxon 34: No illustration or specimen has been found MAP 5. Distribution of Kopsia flavida Blume.

17 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 105 FIGURE 4. Kopsia flavida Blume. A, habit; B, flower dissection; C, ovaries and disk; D, fruit. From Brass 3110 (A) and Takeuchi 9028 (B, C, D). Scale bars = 1 cm (A, B, D); 1 mm (C).

18 106 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 that could be type material. However, from the description and its provenance it is clearly Kopsia flavida. Kentrochrosia monocarpa Laut. & K.Schum., Fl. Schutzgeb. Südsee ; Markgr., Bot. Jahrb. 61: ; Markgr., Nov. Guinea 16(2): ; Merr. & L.M.Perry, Phil. J. Sci. 76: TYPE: PAPUA NEW GUINEA. Oertzen Mountains, C.A.G. Lauterbach 2180 (Holotype: B, not found, presumably destroyed; Schumann & Lauterbach, Fl. Schutzgeb. Südsee t. XVIII, 1900, Lectoype, designated here). The original type specimen could not be found, but it is clear from the collecting locality, the description, and the illustration that it belongs to Kopsia flavida. Kopsia grandiflora Merr., Phil. J. Sci. 20: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: PHILIPPINES. Luzon, Camarines Sur Province, Paracale, M. Ramos, G.E. Edaño (Lectotype: US, designated here; Isolectotypes: A, K, P). Kopsia triangularis Quisumb. & Merr., Phil. J. Sci. 37: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: PHILIPPINES. Mindanao, Surigao, C.A. Wenzel 2648 (Holotype: UC; Isotypes: A, BO, BR, G, M, MO, Z). Kentrochrosia triangularis (Quisumb. & Merr.) Merr. & L.M.Perry, Phil. J. Sci. 76: Kopsia carolinensis Kaneh., Bot. Mag. Tokyo 45: ; Merr. & L.M.Perry, Phil. J. Sci. 76: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: PALAU ISLAND. Babeldayob, Aimiyou, R. Kanehira 502 (Holotype: FU; Isotype: NY); Kentrochrosia carolinensis (Kaneh.) Kaneh. & Hatus., Bot. Mag. Tokyo 53: ; Merr. & L.M.Perry, Phil. J. Sci. 76: Tree to 9.15 m tall, to 27.5 cm dbh. Bark ashy-brown, brown, or gray, smooth; inner bark straw colored. Branchlets glabrous, sparsely lenticellate or not. Leaves: petiole 3 10 mm long, glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, elliptic, apex acuminate with a blunt tip, base acute or cuneate, glabrous above and beneath, midrib deeply or shallowly sunken above, secondary veins 8 24 pairs with 3 15 mm spacing, from midrib, prominent above and beneath, clearly distinguishable from tertiary venation above and beneath, straight or ascending near margin, tertiary venation prominent above and beneath, reticulate, intramarginal vein obscure or more or less straight and inset from margin. Inflorescence dichasial or cincinnate, cm long with axes cm long and branches mm wide, glabrous or puberulent in upper parts; peduncle cm long, mm wide, glabrous; pedicels mm long, glabrous or sparsely puberulent, subtending bracts persistent, bracts present on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex rounded to acute, ciliate, glabrous outside and inside. Corolla white with a red eye or pale pink with a darker pink eye ; tube 26 38( 49) mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around and beneath stamens and in throat, rarely glabrous, glabrous outside; lobes mm, times as long as wide, elliptic or obovate, apex rounded to obtuse, ciliate or not, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, oblong, awl-shaped, narrowly deltoid or lanceolate, apex acute to acuminate. Ovaries mm high, glabrous or, rarely, very sparsely pubescent on top; style mm long; style head mm long. Fruit with a sharp deltoid spur, mm, spur 8 13 mm long, glabrous. Distribution: Philippines, Moluccas, New Guinea, Micronesia, Solomons, Vanuatu. Habitat: in a wide range of forest types from swamp forest to lowland evergreen forest, on limestone, ultrabasic, and poorly to welldrained soils from 0 to 270 m altitude. Geographical selection of the 86 collections studied: CAROLINE ISLANDS: Palau, Babeldaob, Aimiyou, Kanehira 502 (FU, NY). INDONESIA: West Papua: Merau River, South of Senajo, Royen 4674 (L). Moluccas: Buru, Vriese s.n. (L); Wae Toni, Eyma 3171 (A, SING, K, L); Kai Is, Jaheri 8 (BO, L); Seram, Vriese, Teijsmann s.n. (L). PAPUA NEW

19 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 107 GUINEA: Western Province: Wuroi, Brass 5789 (A, BM, BO, BRI); West New Britain Province: Gasmata, Kerenga, Obedi LAE62338 (L); Manus: Lorengau, Kerenga et al. LAE77481 (CANB, L). PHILIPPINES: Mani Kani Island, Quisumbing 2049 (A). Surigao: Lake Mainit, Ramos & Convocar (A). Eastern Samar: Balangiga, Madulid et al (L). SOLOMON ISLANDS: North Solomons: San Jorge, Talise Village, Hunt RSS2730 (A, K, L). Santa Cruz Islands: Vanikoro Island, Kajewski 581 (A, BRI, K, US). San Cristobal: Star Harbour, Brass 3110 (A, BO, BRI, L). Santa Isabel: Allardyce Harbour, Kinifu, Susui BSIP8279 (K, L, SING). VANUATU: Tisbel, Hallé RSNH 6374 (K, L, P). Efate, Burton 58 (A). Rutten 1895 from Seram has a corolla tube very much longer than all other known specimens of this species. Unfortunately there are no label notes on corolla color. In other characters it is similar to the other specimens from Seram. Additional collections may lead to a reappraisal of this decision. 6. Kopsia fruticosa (Roxb.) A.DC., Prod. 8: ; Hook.f., Fl. Brit. Ind. 3: ; Boerl., Handl. Fl. Nederl. Ind. 2: ; King & Gamble, J. As. Soc. Beng. 74(2): ; Merr., Enum. Born. Pl ; Ridl., Fl. Mal. Pen. 2: ; Tsiang, Sunyatsenia 2: ; Masam., Enum. Phan. Born ; Bakh.f., Blumea 6: ; Corner, Wayside Trees Malaya 1: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Markgr., Blumea 20: ; Lý, Feddes Repert. 97: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: ; Coode et al., Checklist Pl. Brunei Map 6. Basionym: Cerbera fruticosa Roxb. in Ker Gawler, Bot. Reg. t ; Roxb., Fl. Ind. 2: ; Roxb., Fl. Ind. 1: TYPE: Roxburgh Flora Indica illustration in Kew, number 2200 (Lectotype, designated here); Calpicarpum roxburghii G.Don, Gen. Syst. 4: ; Wight, Ic. Pl. 2: t ; Kopsia roxburghii (G.Don) Wehmer, Pflanzenst ; Sévenet et al., J. Ethnopharmacol. 41: Synonym: Kopsia vincaeflora Blume, Bijdr ; A.DC., Prod. 8: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: INDONESIA. Java sine loc., C.L. Blume s.n. (Lectotype: L [ ], designated by Sévenet et al. [1994]). MAP 6. Distribution of Kopsia fruticosa (Roxb.) A.DC.

20 108 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Tree to 6 m tall. Bark mostly grayish. Branchlets sparsely to densely puberulent, glabrescent when older, not lenticellate, weakly angled. Leaves: petiole 5 12 mm long, glabrous; blade cm, times as long as wide, subcoriaceous or coriaceous, elliptic, apex short to long acuminate with blunt tip, base obtuse to cuneate, glabrous above and beneath, midrib deeply sunken to flat above, secondary veins 8 17 pairs with 4 21 mm spacing, from midrib, slightly sunken to prominent above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, curved ascending from midrib, tertiary venation prominent or flat above and beneath, subperpendicular to midrib and oblique to secondary veins or obliquely scalariform, usually also somewhat reticulate, intramarginal vein obscure or straight to strongly looped and inset from margin. Inflorescence dichasial, cm long with axes cm long and branches mm wide, sparsely to densely puberulent; peduncle cm long, mm wide, puberulent at least in upper parts; pedicels mm long, sparsely to densely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex emarginate to obtuse, ciliate, glabrous to densely puberulent, glabrous or puberulent on upper half inside. Corolla mostly white or pale pink at the base of the tube and becoming pinker higher up, lobes whitish-pink to pink with a darker pink to almost red throat, more rarely whole corolla only pale pinkish; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside; lobes mm, times as long as wide, elliptic, apex rounded, ciliate or ciliate only at lobe base, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, oblong or awl-shaped, apex acute, acuminate or irregularly toothed and acuminate. Ovaries mm high, densely pubescent; style mm long; style head mm long. Fruit falcate with a blunt hooked spur, ca mm, spur ca. 3 mm long, densely puberulent. Distribution: Burma (Tenasserim), also widely cultivated. The original description suggests the plant comes from Pegu, but no collections have been seen from that far north. Habitat: no habitat information has been provided on the collections made in the wild. Noncultivated collections studied (77 collections, including cultivated specimens, studied in all): BURMA: Tavoy, Wallich 1583 (A, K), Wallich 1583a (BM, M), Wallich H.I.1583 (K); Mergui, Griffith s.n. (K), Parish 254 (K); Yanngwa Range, Parker 2740 (K, A); Mayinnge, Parkinson 1985 (K). Typification: The basionym of this species, Cerbera fruticosa, was originally published in Roxburgh (1814), but without a description, and later described in Ker Gawler s Botanical Register (1819). Roxburgh (1814) and Ker Gawler (1819) note that it originally came from Pegu, Burma, a site much farther north than any of the known collection localities. Ker Gawler further explains that it was introduced as a horticultural shrub to the Calcutta Botanic Garden and eventually to the United Kingdom. Ker Gawler gives credit to Roxburgh for the name and the description; the validating description itself is followed by Roxb. MSS and the wording is the same as that in Roxburgh s Flora Indica (1824). Curiously, Roxburgh has not been credited for the name in the bulk of the literature, with the authority invariably given as Ker Gawler (or variations on his name). It appears to me, however, that Ker Gawler has himself acknowledged that this species was named by Roxburgh and that the validating description comes from a Roxburgh manuscript, and therefore the authority should rather be Cerbera fruticosa Roxb. Roxburgh credits this name to himself but with Ker Gawler (1819) as the place of publication in his account of Cerbera fruticosa in Flora Indica (1824). This raises the issue of typification. Sévenet et al. (1994) typified this species with the illustration of a cultivated plant growing in Fulham in Southeast England in Ker Gawler s 1819 account. However, given that Roxburgh based his name on material cultivated in the Calcutta Botanic Garden, the material from Calcutta must be the type. Forman (1997) did not mention this species in his list of Roxburgh species, and I have been unable to find any

21 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 109 herbarium material that could unequivocally be considered original (although there is a specimen in BM labelled as Cerbera fruticosa in Roxburgh s handwriting but with no other information on the sheet). Sealy (1956) and Sanjappa et al. (1994) note that illustrations exist in the Roxburgh Flora Indica collections at Kew and Calcutta. Forman (1997) makes clear that these illustrations were made under supervision by Roxburgh and that there was a very close link between them and the descriptions in Flora Indica. Hence these illustrations can be considered part of the original material, and a lectotype can be made of the Kew illustration, number The illustration has also been reproduced by Wight (1841) as t. 431 under the name Calpicarpum roxburghii. The large majority of specimens known of this species are from cultivation throughout the tropics. Only the specimens that are likely to have come from the wild have been mapped. Only one fruiting specimen has been seen, which is quite curious for such a commonly collected plant, albeit mostly cultivated. There are a number of specimens labelled Kopsia vincaeflora in Blume s handwriting in the Leiden herbarium. It is not clear whether these are duplicates of a single collection or if they represent multiple collections, and therefore no isolectotypes can be distinguished from the syntypes with any degree of certainty. 7. Kopsia grandifolia D.J.Middleton, sp. nov. TYPE: MALAYSIA. Johor, Sungai Mupar, off 32nd mile Johor Bahru Mersing Road, H.M. Burkill 4230 (Holotype: A; Isotypes: K, L, SAR, SING). Map 7; Fig. 5. Frutex 2 3 m altus, ramulis puberulis. Folia elliptica, cm, nervis paribus. Corollae tubus mm, lobis mm. Tree to 3 m tall. Branchlets sparsely puberulent, sparsely lenticellate or not, terete or weakly angled. Leaves: petiole 7 11 mm long, sparsely to densely pubescent; blade cm, times as long as wide, papery or subcoriaceous, broadly elliptic or elliptic, apex caudate, rarely to short acuminate with a blunt tip, base obtuse to cuneate, glabrous or puberulent on midrib above, puberulent on midrib and sometimes on major veins beneath, at least when young, midrib shallowly sunken to flat above, secondary veins pairs with 3 15 mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable from tertiary MAP 7. Distribution of Kopsia grandifolia D.J.Middleton ( ) and Kopsia lapidilecta Sleesen ( ).

22 110 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 FIGURE 5. Kopsia grandifolia D.J.Middleton. A, habit; B, flower dissection; C, ovaries, disk, and base of style; D, fruit. From Burkill HMB.4230 (A, B, C) and David 265 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).

23 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 111 venation above and beneath, straight or ascending near margin, tertiary venation prominent or flat above, prominent beneath, parallel to secondary veins or subperpendicular to midrib and oblique to secondary veins, usually also somewhat reticulate, intramarginal vein straight or looped, inset from margin. Inflorescence dichasial, with or without longer unbranched branches, cm long with axes cm long and branches mm wide, sparsely to densely puberulent; peduncle cm long, puberulent, pedicels ca. 3 mm long, densely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate, apex rounded, ciliate, densely puberulent, glabrous inside. Corolla buds purple, yellow when open; tube mm long, ca. 1.8 times as long as lobes, ca. 9.7 times as long as calyx, densely pubescent only beneath stamens and becoming less so further down in upper half, throat glabrous, glabrous outside; lobes mm, times as long as wide, linear to oblong, apex rounded to acute, ciliate or not, glabrous outside and inside. Stamens inserted at ca. 15 mm from corolla base, which is ca. 0.8 of corolla tube length in the rehydrated flower measured, anthers mm, times as long as wide, apex mm from corolla throat. Disk mm long, ca. 0.8 times as long as ovaries, glabrous, awl-shaped or deltoid, apex acuminate. Ovaries mm high, glabrous; style mm long (see note); style head mm long. Fruit falcate with a blunt hooked spur in the lower part of the fruit, mm, hook 2 3 mm long, sparsely puberulent. Distribution: Peninsular Malaysia and probably the Anambas Islands (see comment below). Habitat: in forest to 60 m altitude. Additional collections studied: INDONE- SIA: Anambas Islands, North Jemaja, Gunong Datoh, Henderson (SING). MALAYSIA: Johor: Mersing, David 265 (P), Teo et al (P); Sungai Mupah, 53 km from Johor Baru towards Mersing, Teo & Pachiapu 1127 (K, L); Kluang Forest Reserve, Hutan Simpan Gunong Belumut, Teo & Pachiapu 1143 (K, L, SING). This new species is close to Kopsia larutensis, differing from it in its much larger flowers, leaves generally pubescent on the lower surface (at least when young), glabrous ovaries, and larger and longer fruits. Both species have very restricted distributions: K. larutensis in the Malaysian state of Perak and K. grandifolia in the Malaysian state of Johor and probably the nearby Indonesian Anambas Islands. Henderson from the Anambas Islands is a paratype of K. lapidilecta, the type of which is from the Natuna Islands. It is a rather poor specimen without a corolla. However, it does have leaves like K. grandifolia, pubescent when young, the remnants of the inflorescence are pubescent, and a comment on the label notes that the now-missing corolla lobes are narrow. Kopsia lapidilecta also has rather narrow corolla lobes but has glabrous leaves and a glabrous inflorescence. David 265 has an extremely short style, shorter than any other observed in the genus. The type collection of the species, Burkill HMB4230, has a style that places the style head at the same level as the stamens, as is the case in all the other species. Because there are so few specimens of this species, I am unsure whether the short style of David 265 is just an aberrant character or whether it has some pollination function. 8. Kopsia griffithii King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Malacca sine loc., W. Griffith s.n. (Lectotype: CAL, designated by Sévenet et al. [1994]; Isolectotypes BO, K, L, M). Map 8; Fig. 6. Synonym: Kopsia griffithii var. paucinervia King & Gamble, J. As. Soc. Beng. 74(2): TYPE: MALAYSIA. Perak sine loc., King s Collector (Lectotype: UC, designated here; Isolectotype: BO). KEY TO THE VARIETIES 1a. Pedicels glabrous; corolla tube > 30 mm long, lobes > 19 mm long var. griffithii 1b. Pedicels densely pubescent; corolla tube < 25 mm long, lobes < 15 mm long var. pubescens

24 112 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 MAP 8. Distribution of Kopsia griffithii King & Gamble.Var. griffithii ( ), and var. pubescens D.J.Middleton ( ). Kopsia griffithii King & Gamble var. griffithii Tree to 5 m tall; to 24 cm dbh. Bark ashybrown or gray. Branchlets glabrous, not lenticellate or sparsely lenticellate. Leaves: petiole 3 10 mm long, glabrous; blade cm, times as long as wide, papery to subcoriaceous, elliptic to oblong, apex caudate or long acuminate with a blunt tip, base acute or cuneate, glabrous above and beneath, midrib raised and with a central groove to sunken above, secondary veins 9 28 pairs with 1 10 mm spacing, from midrib, clearly distinguishable or not from tertiary venation above and beneath, prominent beneath, prominent or flat above, straight or ascending near margin, tertiary venation prominent above and beneath, reticulate or subperpendicular to midrib and oblique to secondary veins, intramarginal vein straight or only weakly looped, inset from or right at margin. Inflorescence dichasial, lax, 5 12 cm long of which the axes are cm long and branches mm wide, glabrous; peduncle cm long, mm wide, glabrous; pedicels mm long, glabrous, subtending bracts persistent, bracts present or absent on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex rounded to acute, ciliate, glabrous outside and inside. Corolla completely white or white with yellow eye ; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around and beneath stamens and in throat, glabrous outside; lobes mm, times as long as wide, elliptic, apex obtuse or acute, ciliate or not, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, pubescent, awl-shaped, apex acute. Ovaries mm high, densely pubescent on top or densely pubescent all over; style mm long; style head mm long. Fruit falcate with a small spur, mm, spur 3 4 mm long; sparsely puberulent. Distribution: Peninsular Malaysia. Habitat: in primary or secondary forest to 800 m altitude. Selection of the 35 collections studied: MALAYSIA: Selangor: Templer Park, Stone 5956 (MO, US), David 268 (P), Teo & Pachiapu 298 (CANB, K, L, SING), 1130 (K, L, SING); Ipoh Highway, 15 km NW of Kuala Lumpur, Worthington (L, MO, NY). The only material I was able to trace of

25 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 113 FIGURE 6. Kopsia griffithii King & Gamble. A, habit; B, flower dissection; C, ovaries, disk,and base of style; D, fruit. From Kasim 590 (A) and Wyatt-Smith (B, C, D). Scale bars = 1 cm (A, B, D); 1 mm (C).

26 114 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 King s Collector 10707, the type of Kopsia griffithii var. paucinervia King & Gamble, was from BO and UC. I have seen no material from CAL or K, where one would expect the type material to be. Therefore, the name has been lectotypified by the UC specimen. Kopsia griffithii King & Gamble var. pubescens D.J.Middleton, var. nov. TYPE: MALAYSIA. Johor, Sungai Hula Sembrong, 30 October 1892, Lake & Kelsall s.n. (Holotype: SING). A Kopsia griffithii var. griffithii pedicellis pubescentibus corollis parvioribus differt. Height unknown. Branchlets glabrous, not lenticellate, weakly angled. Leaves: petiole 5 7 mm long, glabrous; blade cm, times as long as wide, coriaceous, elliptic, apex long acuminate with a blunt tip, base acute, glabrous above and beneath, midrib shallowly sunken above, secondary veins pairs with 5 14 mm spacing, from midrib, ascending near margin, prominent and not clearly distinguishable from tertiary venation above, tertiary venation prominent and reticulate or subperpendicular to midrib and oblique to secondary veins above, intramarginal vein straight or only weakly looped, inset from margin. Inflorescence dichasial, ca. 11 cm long with axes ca. 9.3 cm long and branches ca. 1 mm wide, densely puberulent; peduncle ca. 5 cm long, ca. 2.5 mm wide, puberulent in upper parts; pedicels mm long, densely puberulent; subtending bracts persistent; bracts absent on pedicel. Sepals ca mm, ca. 1.7 times as long as wide, ovate, apex acute, ciliate, sparsely puberulent outside, glabrous inside. Corolla tube mm long, ca. 1.6 mm wide, ca. 1.9 times as long as lobes, ca. 9 times as long as calyx, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside; lobes ca mm, ca. 3.8 times as long as wide, elliptic, apex obtuse, not ciliate, glabrous outside and inside. Stamens inserted at ca. 18 mm from corolla base which is ca. 0.9 of corolla tube length in the rehydrated flower measured; anthers ca mm, ca. 3.5 times as long as wide, apex ca. 0.6 mm from corolla throat. Disk ca. 0.6 mm long, ca. 0.7 times as long as ovaries, pubescent, awlshaped, apex acute. Ovaries ca. 0.9 mm high, densely pubescent all over; style ca mm long; style head ca. 8 mm long. Fruit unknown. Distribution: Peninsular Malaysia (Johor). Habitat: not recorded. The single specimen known of this taxon is an old Lake & Kelsall s.n. specimen in SING collected on 30 October It is unfortunately not a very good specimen, and I have created this variety somewhat reluctantly. It is, however, quite different from typical Kopsia griffithii, and future collections of var. pubescens may determine that it deserves specific status. The main differences from var. griffithii are in the rather more lax and many-flowered inflorescence, the densely pubescent pedicels, and the smaller flowers. It shares with var. griffithii the lax inflorescence and a pubescent disk, and the two are the only taxa in the genus that have this latter character. 9. Kopsia hainanensis Tsiang, Sunyatsenia 2: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: CHINA. Hainan: Yaichow, F.C. How (Lectotype: IBSC, designated here; Isolectotypes: A, CANT, E, K, P). Syntype: N.K. Chun & C.L. Tso (A, NY). Map 9. Tree to 2 m tall. Bark gray. Branchlets glabrous, sparsely lenticellate or not. Leaves: petiole 4 8 mm long, glabrous; blade cm, times as long as wide, papery or subcoriaceous, elliptic, apex acuminate with a blunt tip, base cuneate, glabrous above and beneath, midrib raised and with a central groove, flattened or shallowly sunken above, secondary veins pairs with 2 10 mm spacing, from midrib, prominent above, prominent or not beneath, clearly distinguishable or not from tertiary venation above and beneath, tertiary venation prominent above, prominent or not beneath, reticulate, intramarginal vein straight or only weakly looped, slightly inset or right at margin. Inflorescence dichasial, cm long, of which the axes are cm long and branches mm wide, glabrous; peduncle cm long, mm wide, glabrous; pedicels mm long, glabrous, bracts present on pedicel. Sepals mm, times as long as wide, ovate, apex obtuse to acute, ciliate or not, glabrous outside, glabrous or puberulent at tips inside. Corolla completely white; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent in upper half of tube inside, glabrous outside; lobes mm, times as long as wide, elliptic or obovate, apex obtuse to

27 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 115 MAP 9. Distribution of Kopsia harmandiana Pierre ex Pitard ( ) and Kopsia hainanensis ( ). acute, not ciliate, pubescent at very base of lobes inside, glabrous outside. Stamens inserted mm from corolla base, which is ca. 0.8 of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat. Filaments mm long. Disk mm long, times as long as ovaries, glabrous, awl-shaped, apex acute. Ovaries mm high, glabrous or very sparsely short pubescent on top; style mm long; style head mm long. Fruit oblique ellipsoid with an angle on one side, ca mm. Distribution: China (Hainan) Habitat: in forest or on river banks from 240 to 430 m altitude. Additional collections studied: China: Hainan: sin. loc. Katsumada s.n. (UC), Liang (K, NY), Wang (A, NY); Yaichow, How (E); Po-ting, How (A, BISH, BM, G, MO, P, PE), Fung (A, K, NY, UC, US); Bak Sa, Lau (A); Dung Ka, Chun & Tso (A, NY). Typification: Tsiang gives the location of the type specimens as in the herbarium of Sunyatsen University. These specimens are now to be found at IBSC. Therefore the lectotype has to be chosen from amongst the IBSC duplicates of How and Chun & Tso 43853, superceding Sévenet et al. s designation of the CANT duplicate of How (Sévenet et. al., 1994). E. D. Merrill has written the name Kopsia stenophylla Merr. n. sp. on Fung from Hainan, but there is no evidence that this name was ever published. 10. Kopsia harmandiana [Pierre in L.Planch., Prod. Apoc , nom. nud.] ex Pitard, Fl. Gén. Indo-Chine 3: ; Lý, Feddes Repert. 97: ; Sévenet et al., J. Ethnopharmacol. 41 (1994) 155. TYPE: VIET- NAM. Between the Mekong and Hué, Harmand in Pierre 5246 (Lectotype: P, designated by Sévenet et al. [1994]; Isolectotypes: BR, P). Syntypes: P.A. Eberhardt 2740 (P), P.A. Eberhardt 2698 (P). Map 9. Tree to 14 m tall. Branchlets glabrous, sparsely lenticellate. Leaves: petiole 2 8 mm long, glabrous; blade cm, times as long as wide, papery, subcoriaceous to coriaceous, elliptic or narrowly so, apex caudate to short acuminate with a blunt tip, base cuneate, glabrous above and beneath, midrib raised and with a central groove above, flattened or shallowly sunken above, secondary veins pairs with mm spacing,

28 116 HARVARD PAPERS IN BOTANY Vol. 9, No from midrib, slightly prominent but not clearly distinguishable from tertiary venation above and beneath, straight, tertiary venation prominent above and beneath, parallel to secondary veins and reticulate, intramarginal vein right at margin. Inflorescence dichasial, cm long, branches 0.7 mm wide, glabrous; peduncles cm long, 0.7 mm wide, glabrous; pedicels mm long, glabrous, bracts present on pedicel. Sepals mm, times as long as wide, lanceolate, apex acuminate, ciliate, glabrous outside, puberulent at tips inside. Corolla completely white; tube mm long, mm wide, ca. 1.4 times as long as lobes, times as long as calyx, pubescent in upper part of tube above, around, and slightly below the stamens and in throat, glabrous outside; lobes mm, times as long as wide, obovate, apex acute, not ciliate, glabrous or pubescent at very base of lobes inside, glabrous outside. Stamens inserted mm from corolla base, which is ca. 0.5 of corolla tube length in the rehydrated flowers measured; anthers ca mm, ca. 5.4 times as long as wide, apex mm from corolla throat; filaments 0.4 mm long, pubescent. Disk mm long, ca. 1 times as long as ovaries, glabrous, awl-shaped or lanceolate, apex acute to acuminate. Ovaries mm high, densely pubescent on top; style 8.5 mm long; style head 1.1 mm long. Fruit falcate with a small spur, ca mm, spur 2 mm long, sparsely puberulent. Distribution: Vietnam. Habitat: not recorded. Additional collections studied: Vietnam: Thua thien, Bo Giang, Eberhardt 2698 (P), 2740 (P); Nha Trang, Suôi Dâu, Vidal 4865 (L, P). The location of the type is given vaguely as between the Mekong and Hué. This locality has not been included on the map, but wherever it actually is would be rather to the west of the other location marked on the map. 11. Kopsia lapidilecta [Sleesen, Fl. Mal. Misc. Rec. 1: , no Latin description] ex Sleesen, Blumea 10: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: INDONESIA. Natuna Islands, Gunung Ranai, C.G.G.J. van Steenis 1384 (Holotype: L; Isotype: BO). Map 7. Habit unknown. Branchlets glabrous, densely lenticellate, terete. Leaves: petiole 3 8 mm long, glabrous; blade cm, times as long as wide, coriaceous, elliptic, apex short acuminate with a blunt tip, base obtuse, glabrous above and beneath, midrib flat to shallowly sunken or raised and with a central groove above, secondary veins pairs with 2 6 mm spacing, from midrib, prominent above and beneath, not clearly distinguishable from tertiary venation above, partly distinguishable or not from tertiary venation beneath, straight, tertiary venation prominent above and beneath, parallel to secondary veins or reticulate, intramarginal vein straight or only weakly looped, inset from margin. Inflorescence with few elongate branches, cm long with axes cm long and branches 2.3 mm wide, glabrous; peduncle cm long, 2 mm wide, glabrous; pedicels mm long, glabrous; subtending bracts persistent; bracts present or absent on pedicels. Sepals mm, times as long as wide, ovate, apex rounded, ciliate, otherwise glabrous outside and inside. Corolla bright red; tube mm long, mm wide, times as long as lobes, times as long as calyx, densely pubescent beneath stamens and becoming less so further down, throat glabrous, glabrous outside; lobes mm, times as long as wide, narrowly oblong, apex obtuse, not ciliate, glabrous, outside and inside. Stamens inserted mm from corolla base, which is ca. 0.8 of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex ca. 0.9 mm from corolla throat; filaments ca. 1 mm long, glabrous. Disk ca. 1.4 mm long, times as long as ovaries, glabrous, awl-shaped, apex acute. Ovaries mm high, glabrous; style ca. 16 mm long; style head ca. 0.7 mm long. Fruit not seen (see comment below). Distribution: Indonesia (Natuna Islands). Habitat: recorded from 200 m altitude. Additional collection studied: INDONE- SIA. Kalimantan Barat, Bunguran, Gunung Ranai, 1385 (BO,L,SING). Typification: Timmerman-Van der Sleesen is not entirely clear about the typification of this species, which under Art of the International Code of Botanical Nomenclature would only be validly published by reference to

29 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 117 a single type specimen. She has written Typus: Van Steenis 1384 (L); do 1385; Henderson SF Anambas & Natuna Islands. This could be interpreted as indicating that Van Steenis 1384 in Leiden is the holotype or that all three are syntypes. Only the former interpretation would make it a validly published species. However, Markgraf (1973) attempted to typify the species with Van Steenis 1385, which, if allowed, would actually invalidate the publication of the species. A type label has been placed on Van Steenis 1385 in Leiden, but the typed determination label by Timmerman-Van der Sleesen also has a handwritten note saying paratype, possibly in her hand. Sévenet et al. (1994) gave the holotype as Van Steenis 1384 (L), which, for the sake of convenience, I shall also accept as a holotype, this being the interpretation of the original publication that would effect no change. Henderson SF 20395, the only one of the paratypes from the Anambas Islands rather than the Natuna Islands, has now been moved to the new species Kopsia grandifolia. A fruit is illustrated in an accompanying illustration to the original description. However, none of the original material now extant has any sign of a fruit on it. The illustration shows a falcate fruit with only a small projection on one side. 12. Kopsia larutensis King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: Malaysia, Perak, Larut, L. Wray 2736 (Lectotype: CAL, designated by Markgraf [1973: 423], first step, and here, second step [see comment below]; Isolectotypes: CAL, SING). Syntypes: King s Collector 4269 (BM, K, L, SING), King s Collector 6165 (BO, L, SING), King s Collector 462 (K, P), King s Collector 2082 (CAL), L. Wray 3956 (not found), B. Scortechini 59b (BM, K), B. Scortechini 1704 (CAL), B. Scortechini 57b (BM). Map 10. Tree to 1.8 m tall. Branchlets glabrous, not lenticellate, weakly angled. Leaves: petiole 5 12 mm long, glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, elliptic or oblong, apex caudate or more rarely long acuminate, in both cases with a blunt tip, base acute to cuneate, glabrous above and beneath, midrib shallowly sunken to raised above, or raised and with a central groove, secondary veins MAP 10. Distribution of Kopsia larutensis King & Gamble.

30 118 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 pairs with 4 11 mm spacing, from midrib, prominent to shallowly sunken above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, curved ascending from midrib or only near margins, tertiary venation prominent above, prominent or not beneath, subperpendicular to midrib and oblique to secondary veins and then somewhat reticulate, intramarginal vein straight or only weakly looped, inset from margin. Inflorescence often many times branched near the base with short thick branches that appear very crowded, cm long with axes cm long and branches mm wide, glabrous or puberulent in upper parts; peduncle cm long, 2.2 mm wide, glabrous; pedicels mm long, glabrous or sparsely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate, apex rounded, ciliate, glabrous or sparsely puberulent outside, glabrous inside. Corolla completely white; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent in lower half of tube but not at base, throat glabrous, glabrous outside; lobes mm, times as long as wide, linear, apex acute, not ciliate, glabrous outside and inside. Stamens inserted ca. 6.5 mm from corolla base, which is ca. 0.8 of corolla tube length in the rehydrated flowers measured; anthers ca mm, ca. 3.5 times as long as wide, apex ca. 0.3 mm from corolla throat, filaments 0.5 mm long, glabrous. Disk mm long, times as long as ovaries, glabrous, awl-shaped, apex acute. Ovaries mm high, glabrous or very sparsely pubescent on top; style ca. 5 mm long; style head ca. 0.7 mm long. Fruit with a blunt hooked spur, mm, spur ca. 4 mm long, sparsely puberulent. Distribution: Peninsular Malaysia (Perak). Habitat: understory shrub in lowland forest to 200 m altitude. Selection of the 20 collections studied: Perak: Larut, Scortechini 57b (BM), 59b (BM, K), King s Collector 462 (P, K), 4269 (BM, K, L, SING), 6165 (BO, L, SING); Batu Gajah, David 188 (P); Kledang Saigong Forest Reserve, Ng FRI6024 (A, K, KEP, L, SING), FRI6063 (A, K, KEP, L, SING), Symington KEP (K, KEP, L); Batu Kurau, Haniff & Said (SING). Kedah: Kulim, Nagaswamy 19 (SING). Typification: King and Gamble listed several syntypes for this species. A first stage lectotypification in accordance with the St. Louis Code, article 9.14 (Greuter et al., 2000) was performed by Markgraf (1973). Sévenet et al. (1994) referred to a holotype in CAL but did not distinguish between the two specimens of this collection to be found there. Therefore, this was not an effective second step lectotypification. This is a very characteristic species with inflorescences that are branched near the base followed by crowded short cincinnate branches with small flowers that have narrow corolla lobes. The description given above will be expanded when further and better collections are made of this species. Although there are many collections, only Ng FRI 6024 had sufficiently mature and plentiful flowers to dissect, so it should be borne in mind that the androecium and gynoecium characters are based on measurements from this specimen only. 13. Kopsia macrophylla Hook.f., Fl. Brit. Ind. 3: ; King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: Singapore, T. Lobb s.n. (Holotype: K; Isotype: BM). Map 11. Synonym: Kopsia ridleyana King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: MALAYSIA. Negri Sembilan, Tinggi, H.N. Ridley (Holotype: K). Tree to 6.1 m tall. Branchlets glabrous to densely puberulent, sparsely lenticellate or not, terete to weakly angled. Leaves: petiole 3 8 mm long, glabrous to densely pubescent; blade cm, times as long as wide, papery or subcoriaceous, elliptic to obovate, apex caudate or short to long acuminate with a blunt tip, base acute or cuneate, glabrous to puberulent on midrib and major veins beneath, midrib shallowly sunken, flat or raised and with a central groove above, secondary veins 8 23 pairs with mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, straight or curved ascending from midrib or

31 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 119 MAP 11. Distribution of Kopsia macrophylla Hook.f. only near margin, tertiary venation prominent above and beneath, more or less parallel to secondary veins or subperpendicular to midrib and oblique to secondary veins and then also somewhat reticulate, intramarginal vein straight to strongly looped, inset from margin, glabrous or puberulent on midrib above. Inflorescence dichasial, mostly also with cincinnate branches, cm long with axes cm long and branches mm wide, glabrous to densely puberulent; peduncle cm long, mm wide, glabrous to puberulent; pedicels mm long, glabrous to densely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex rounded, margin ciliate, glabrous to densely puberulent, glabrous inside. Corolla completely white or white with a yellow eye ; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around stamens and slightly beneath inside or pubescent around and beneath stamens and in throat, glabrous inbetween, throat pubescent, sometimes so shortly so as to appear papillose, glabrous outside; lobes mm, times as long as wide, elliptic or broadly elliptic, apex rounded, ciliate or not, glabrous outside and inside. Stamens mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments 0.6 mm long. Disk mm long, times as long as ovaries, glabrous, awl-shaped, apex acute to obtuse. Ovaries mm high, sparsely to densely pubescent on top; style mm long; style head ca. 1 mm long. Fruit with a blunt hooked spur from about the middle of the fruit, mm, spur ca. 4 mm long, glabrous or sparsely puberulent. Distribution: Peninsular Malaysia. Habitat: Recorded from 100 to 152 m altitude. Selection of the 14 collections studied: MALAYSIA: Johor: Gunong Angsi, Ridley (K); Between Perhentian Tinggi and Gunong Angsi, Ernst 1074 (L). Melaka, Alvins 2178 (SING). Negri Sembilan: Seremban, Bukit Tangga, Napier s.n. (SING). Pahang: Dong, near Raub, Burkill & Haniff (SING); Bukit Serdam, Henderson (K, NY, SING); Rompin, Mahamud (KEP, SING), Watson CF3194 (K, KEP, SING); Lawang Ulu Lung, Mat Soh (K, KEP). Perak: sin. loc. King s Collector 4963 (K). SINGAPORE, T. Lobb s.n. (BM, K).

32 120 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Typification: Sévenet et al. (1994) typified Kopsia macrophylla with King 4963 from Perak, despite the fact that this collection was made in 1883, a year after the species was published. Hooker had noted that he had seen only one specimen collected by Lobb in Singapore. Lobb s.n. in Kew is, therefore, the holotype. Kopsia ridleyana was first included in synonymy of Kopsia macrophylla by Timmerman- Van der Sleesen (1959), and was followed by Markgraf (1973) and Sévenet et al. (1994). Timmerman-Van der Sleesen noted that the leaf characters, which, in part, were used by King and Gamble (1907) to distinguish the two species, break down with the larger number of specimens available since Kopsia ridleyana was described. However, there are differences between typical K. macrophylla and typical K. ridleyana. Kopsia macrophylla has a densely pubescent inflorescence and is sparsely to densely pubescent on the twigs and on the undersurface of the leaf blades. Kopsia ridleyana is glabrous and also tends to have smaller flowers. However, there are specimens that have a pubescent inflorescence but glabrous twigs and leaves, and specimens with small flowers but pubescent inflorescences, leaves, and twigs. These bridge the gap between the extremes and so prevent a demarcation of taxa. The variation in this species is rather akin to that found in K. pauciflora var. pauciflora. Of the form with densely pubescent inflorescences there appear to be two subforms: one with a distinctly pubescent corolla throat and another where the hairs in the throat appear to be so short as to appear more or less papillose. The first form is throughout the range of the species, the latter from Pahang in Peninsular Malaysia. There are otherwise no differences in other characters. No new material of either form has been collected since Kopsia pauciflora Hook.f., Fl. Brit. Ind. 3: ; King & Gamble, J. As. Soc. Beng. 74(2): ; Ridl., Fl. Mal. Pen. 2: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Anderson, Checklist Trees Sarawak ; P.S.Ashton, Manual Non-dipt. Trees Sarawak ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Malacca: Mount Ophir, A.C. Maingay KD 1056 (Holotype: K; Isotypes: K, L [scrap]; Photograph: A). Maps Synonyms: Kopsia parvifolia Merr., Phil. J. Sci. 29: ; Masam., Enum. Phan. Born ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Sévenet et al., J. Ethnopharmacol. 41: ; Beaman et al., Plants Mount Kinabalu 4: TYPE: MALAYSIA. Sabah: Banguey Island [=Pulau Banggi], A. Castro & F. Melegrito 1456 (Holotype: UC; Isotypes: A, BM, BO, K). Kopsia caudata Merr., Univ. Calif. Publ. Bot. 15: ; Masam., Enum. Phan. Born ; Sévenet et al., J. Ethnopharmacol. 41: ; Coode, et al., Checklist Pl. Brunei TYPE: MALAYSIA. Sabah, Sandakan, A.D.E. Elmer (Lectotype: UC, designated here; Isolectotypes: A, BISH, BM, BO, BR, F, G, K, L, M, MICH, NY, P, Z). Kopsia caudata var. glabra Merr., Univ. Calif. Publ. Bot. 15: ; Masam., Enum. Phan. Born ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: MALAYSIA. Sabah, Tawao, A.D.E. Elmer (Lectotype: UC, designated here; Isolectotypes: A, BISH, BM, BO, BR, BRI, F, K, G, GH, L, M, MICH, MO, NY, P, US, Z). Kopsia alba Ridl. ex M.R.Hend., Gard. Bull. Straits Settlem. 5: ; Kerr, Fl. Siam. Enum. 2: , pro parte; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Pahang, Pulau Tioman, Ayer Surin, M.R. Henderson (Lectotype: K, designated by Sévenet et al. [1994]; Isolectotype: UC). Kopsia lancifolia Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: MALAYSIA. Sabah, Mendulong, H.P. Nooteboom 1089 (Holotype: L; Isotypes: BISH, CANB, SAN, US). Usage synonym: Kopsia sp. Ridl., J. Straits Branch Roy. Asiatic. Soc. 59: Kopsia macrophylla auct. non Hook.f.: Beaman et al., Plants Mount Kinabalu 4:

33 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 121 MAP 12. Distribution of Kopsia pauciflora Hook.f. var. pauciflora. MAP 13. Distribution of Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton. KEY TO THE VARIETIES 1a. Corolla tube mm long; stamens inserted at mm from the base of corolla tube, anthers at mm from corolla throat var. mitrephora 1b. Corolla tube mm long; stamens inserted at mm from the base of corolla tube, anthers at mm from corolla throat var. pauciflora

34 122 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 Kopsia pauciflora Hook.f. var. pauciflora. Map 12. Tree to 10 m tall; to 15 cm dbh. Bark gray, olive-brown or white, smooth; inner bark pale brown, straw colored or white. Branchlets glabrous to sparsely puberulent, lenticellate or not, angled, weakly winged or markedly winged. Leaves: petiole 0 10 mm long, glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, elliptic, apex caudate or short to long acuminate with a blunt tip, base rounded to cuneate, glabrous or puberulent on midrib above, glabrous or puberulent beneath, midrib deeply to shallowly sunken above, or raised above, sometimes then with a central groove, secondary veins 7 24 pairs with 2 19 mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable or not from tertiary venation above and beneath, straight or curved ascending from midrib or only near margin, tertiary venation prominent or flat above and beneath, sometimes rather obscure, parallel or subperpendicular to midrib and oblique to secondary veins, then also somewhat reticulate, intramarginal vein obscure, straight or only weakly looped. Inflorescence dichasial or with cincinnate branches, cm long with axes cm long and branches mm wide, glabrous to densely puberulent; peduncle cm long, mm wide, glabrous to densely puberulent; pedicels mm long, glabrous to densely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex rounded, ciliate, glabrous to densely puberulent outside, glabrous or puberulent on upper half inside. Corolla completely white, white with yellow eye, white and green or, rarely, pinkishwhite; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous or sparsely puberulent at top of tube outside; lobes mm, times as long as wide, elliptic or obovate, apex rounded, ciliate or not, glabrous outside and inside. Stamens mm from corolla base which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, glabrous or pubescent. Disk mm long, times as long as ovaries, glabrous, awl-shaped or deltoid, apex acute, acuminate or irregularly toothed. Ovaries mm high, sparsely to densely pubescent, or mostly glabrous with a few hairs; style mm long; style head mm long. Fruit falcate with a blunt hooked spur from about the middle of the fruit, mm, hook 3 5 mm long, sparsely puberulent. Distribution: Southern Thailand, Peninsular Malaysia, Sumatra, Borneo, Java (but see note below). Habitat: in a wide variety of forest types, on a wide variety of soil types, from sea level to 1500 m altitude. Geographical selection of the 165 collections studied: BRUNEI DARUSSALAM: River Batu Apoi Sebatu watershed, Ashton BRUN310 (K, L). INDONESIA: Java: Cicurug, Jampango, Backer (K, L, SING). Natuna Islands: West of Kampong Ranai, Steenis 1097 (L). Sumatra: Riau, Bünnemeyer 5962 (L). MALAYSIA: Terengganu: Sungai Kerbat at Jeram Petang, Whitmore FRI20351 (SING, K, L, A, KEP). Perak: Piah Forest Reserve, Jaamat (SING). Sabah: Mt. Silam, 12 miles WSW of Lahad Datu, Beaman (GH, K, L, NY); Kota Merudu, Bukit Mandalong, Amin Gambating SAN (K, L, SAN). Sarawak: Maputi, Brooke (L), (BM, G, L, US), (BM, L). THAILAND: Krabi: Ao Luk, Charoenphol, Larsen & Warncke 3446 (L, P). Phangnga: Sanangmanora Forest Park, Middleton 570 (A, BKF). Narathiwat: Bacho National Park, Kiah (K). Typification: Sévenet et al. (1994) designated the BM duplicate of A. Castro & F. Melegrito 1456 the lectotype (but incorrectly used the term holotype) of Kopsia parvifolia. However, Merrill quite clearly states in the protologue that the still extant UC duplicate is the holotype. Similarly, Sévenet et al. (1994) designated the NY duplicates of A.D.E. Elmer and A.D.E. Elmer the holotypes (= lectotypes) of K. caudata and K. caudata var. glabra, respectively. Merrill, however, in an introduction to the paper in which these species were described, notes that he based his descriptions on his examination of the material in PNH and UC, and therefore the lectotypes must be chosen from amongst this material. Given that the PNH material has been destroyed, the UC material must be lectotypified for both taxa.

35 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 123 Only one collection, Backer 17199, is known from Java. Although it is not indicated on the label, this collection may have been taken from a cultivated plant and so the species may not be native to Java. The plant is described variously as a shrub or small tree on almost all specimens except for Madani et al. SAN142633, on which it is described as a 70 m tall tree with a 50-m clear bole and a girth of 60 cm. This is such an enormous tree (albeit with an unlikely girth) and so out of keeping with all the other material of this species, and of the genus as a whole, that I suspect a mistake has been made and so the dimensions are not included in the description above. This is an extremely variable species both in vegetative and floral morphology, although this variation is mostly manifest in the Bornean specimens rather than the continental ones. The fruit, however, is fairly uniform. The patterns of variation are reticulate. At the extremes, the specimens with thickly coriaceous leaves and anther apices only a few millimetres below the throat of the corolla tube appear very different from the specimens with thinner leaves and stamens inserted in the lower half of the corolla tube. However, around the Telupid area in Sabah specimens with particularly and characteristically thick leaves and a fairly uniform vegetative morphology show almost the entire range of stamen position within the tube. Beaman and Gardner 79, both from Tawai in Sabah, have leaves with a very short petiole but would otherwise appear to fall within the variation of Kopsia pauciflora. There are also specimens scattered elsewhere with stamens inserted high in the corolla tube and thin leaves or stamens inserted low in the corolla tube and thick leaves. There are, however, relatively few specimens that are really intermediate in stamen position, leading to intriguing and unanswered questions about the pollination biology of this species. Larsen et al from Ao Luk in Krabi in Thailand has a label describing the flower color as pinkish white. It is the only specimen not to have the flower color described as either white or white with a yellow eye but is otherwise unremarkable for Kopsia pauciflora. In the area where it was collected, the newly described and undoubtedly closely related K. rosea, which has either white or pink flowers, is also found, and the possibility arises that there has been some hybridization between the two taxa. Kopsia pauciflora var. mitrephora (Sleesen) D.J.Middleton, Gard. Bull. Sing. 55: Map 13. Basionym: Kopsia mitrephora Sleesen, Blumea 10: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: TYPE: MALAYSIA. Sabah: Lahad Datu District, Path between Sungei Sabahan and Sungei Dok, G.H.S. Wood SAN16118 (Holotype: L; Isotype: BRI). Tree to 3 m tall. Branchlets glabrous, sparsely lenticellate or not, weakly angled to markedly winged. Leaves: petiole 2 7 mm long, glabrous; blade cm, times as long as wide, subcoriaceous or coriaceous, elliptic or oblong, apex long acuminate with a blunt tip, base acute to cuneate, glabrous above and beneath, midrib shallowly sunken to raised above, secondary veins pairs with 3 14 mm spacing, from midrib, prominent above and beneath, clearly distinguishable from tertiary venation above and beneath, straight to curved ascending from midrib, tertiary venation prominent above and beneath, parallel to secondary veins or subperpendicular to midrib and oblique to secondary veins, usually also somewhat reticulate, intramarginal vein straight or looped, inset from margin. Inflorescence dichasial, cm long with axes cm long and branches mm wide, glabrous, peduncle cm long, mm wide, glabrous, pedicels mm long, glabrous or sparsely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm wide, times as long as wide, ovate or oblong, apex rounded, ciliate or not, glabrous outside and inside. Corolla completely white or white with a yellow eye, tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside; lobes mm, times as long as wide, elliptic, apex rounded to obtuse, ciliate or not, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat,

36 124 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, awl-shaped, apex acute to acuminate. Ovaries mm high, sparsely to densely pubescent; style mm long; style head mm long. Fruit falcate with a small blunt hooked spur on one side, mm, hook 3 5 mm long, sparsely puberulent. Distribution: Malaysia (Sabah). Habitat: in primary forest. Selection of the 29 collections studied: MALAYSIA: Sabah: Beluran, Sungai Ensuan, Aban SAN90031 (K, KEP, L, SAN, SAR); Labuk Sugut, Timimbang Labuk Sugut, Amin & Martin SAN69331 (K, SAN); Sandakan, Bukit Mangkop, SW of Buis Labuk, Meijer SAN51546 (K, L, SAN, SAR). 15. Kopsia profunda Markgr., Blumea 20: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Terengganu, Belara Forest Reserve, J. Sinclair (Holotype: L; Isotypes: E, K, US). Map 14. Synonym: Kopsia terengganensis L.Allorge & Wiart, Acta Bot. Gallica 142: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: MALAYSIA. Terengganu, Dungun, C. Wiart & L.E. Teo KL 4432 (Holotype: P; Isotypes: KEP, P). Tree to 3 m tall. Bark gray. Branchlets glabrous, sparsely to densely lenticellate or not. Leaves: petiole 2 12 mm long, glabrous; blade cm, times as long as wide, papery, elliptic or oblong, apex caudate to long acuminate with a blunt tip, base cuneate, glabrous above and beneath, midrib shallowly sunken, to raised and and then sometimes with a central groove above, secondary veins pairs with mm spacing, from midrib, prominent above and beneath, clearly distinguishable from tertiary venation above and beneath, ascending near margin or curved ascending from midrib, tertiary venation prominent above and beneath, parallel to secondary veins or irregularly subperpendicular to midrib and oblique to secondary veins, intramarginal vein straight or only weakly looped, inset from or right at margin. Inflorescence dichasial or cincinnate, cm long with axes cm long and branches mm wide, glabrous; peduncle cm long, mm wide, glabrous; pedicels mm long, glabrous or sparsely puberulent, subtending bracts MAP 14. Distribution of Kopsia profunda Markgr.

37 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 125 deciduous, bracts present on pedicel. Sepals mm, times as long as wide, ovate, apex obtuse to acute, ciliate, glabrous outside, glabrous or puberulent at tips inside. Corolla completely white or white with a yellow eye ; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent in upper part of tube above or around and slightly below the stamens, pubescent in throat, glabrous outside; lobes mm, times as long as wide, elliptic, apex obtuse to acute, ciliate or not, glabrous outside and inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm wide, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, awlshaped, apex acute. Ovaries mm high, densely long pubescent on top or densely pubescent all over; style mm long; style head mm long. Fruit falcate with a small or blunt hooked spur, ca mm, hook 4 mm long, sparsely puberulent. Distribution: Peninsular Malaysia. Habitat: in lowland evergreen forest from 230 to 330 m altitude. Additional collections studied: Terengganu: Bukit Kajang, Corner (K, SING); Ulu Brang, Moysey & Kiah SFN (K, L, SING); Dungun, Hutan Simpan Bukit Bauk, Anthony 604 (KEP), Teo & Pachiapu 1080 (K, L, SING), Sevenet & Deverre 242 (P), Deverre 135 (P), Teo, Pachiapu & Boichard 1186 (P); Dungun, David 217 (P), Hutan Simpan Bukit Kesing, Teo & Pachiapu 1136 (K, L, SING); km from Dungun to Kuala Berang, Teo & Din KL 5035 (P). The only differences between this species and Kopsia griffithii are in the stamens, which are borne lower in the corolla tube, and the lack of pubescence on the disk. They share the intramarginal vein, the yellow-eyed corolla, the acute sepals, and the lax inflorescence. Sévenet et al. (1994) distinguished Kopsia terengganensis from K. profunda on the position of the stamens in the corolla tube. They suggested that K. profunda had stamens situated at the base of the tube and K. terengganensis in the middle. In fact, the type specimens of K. profunda and of K. terengganensis have stamens situated at exactly a third of the length of the corolla tube from the base. They are indistinguishable in other characters. Sévenet et al. (1994) suggested that Kopsia terengganensis (= K. profunda) differed from K. macrophylla in being erect as opposed to prostrate. However, the only specimen of either species I have seen where the habit is described as prostrate was Sévenet & Deverre 242, which was incorrectly labelled as K. macrophylla but is in fact K. profunda. The leaves are also just as large in this specimen as in K. macrophylla. However, in K. profunda the inflorescence is delicate, lax, and few-flowered and the sepals acute, characters which do suggest that it is distinct from K. macrophylla. 16. Kopsia rajangensis D.J.Middleton, Gard. Bull. Sing. 55: TYPE: BORNEO. Sarawak, Kapit, Upper Rejang River, Clemens & Clemens (Holotype: MO; Isotypes: A, BM, BO, NY, SAR). Map 15. Usage synonyms: Kopsia sp. Sleesen, Fl. Mal. Misc. Rec. 1: Kopsia larutensis auct. non King & Gamble: Anderson, Checklist Trees Sarawak ; P.S.Ashton, Manual Non dipt. Trees Sarawak Kopsia arborea auct. non Blume: Markgr., Blumea 20: , pro parte. Small tree or shrub, to 4.6 m tall. Branchlets glabrous, not or sparsely lenticellate, terete. Leaves: petiole 4 9 mm long, glabrous; blade cm, times as long as wide, papery to subcoriaceous, elliptic or oblong, apex caudate, base obtuse to cuneate, glabrous above and beneath, midrib shallowly sunken or raised and with a central groove above, secondary veins 9 25 pairs with 6 25 mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, straight or slightly ascending near margin, tertiary venation prominent above, prominent or flat beneath, reticulate or sub-perpendicular to midrib and oblique to secondary veins, intramarginal vein looped and inset from margin. Inflorescences dichasial and then with cincinnate branches, 4 15 cm long with axes cm long and branches mm wide, glabrous to densely puberulent; peduncle cm long, mm wide; pedicels ca. 4 mm long; subtending bracts persistent. Sepals mm, times as

38 126 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 MAP 15. Distribution of Kopsia rajangensis D.J.Middleton ( ), Kopsia sleeseniana Markgr. ( ), and Kopsia tenuis Leenh. & Steenis ( ). long as wide, ovate, apex rounded, ciliate, otherwise glabrous outside and inside. Corolla white; tube mm long, ca. 2.3 mm wide, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside, times as long as lobes, times as long as calyx; lobes mm wide, times as long as wide, elliptic or oblong, apex obtuse or acute, not ciliate, glabrous outside and inside. Stamens mm from corolla base which is ca. 0.8 of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments ca. 0.7 mm long. Disk ca. 0.9 mm high, ca. 1 times as long as ovaries, glabrous, awl-shaped, apex acute. Ovaries ca. 0.9 mm high, glabrous to densely pubescent; style ca. 20 mm long; style head ca. 0.8 mm long. Fruits falcate with small blunt projection near the base, mm; projection 3 4 mm long. Distribution: Malaysia (Sarawak). Habitat: in primary or disturbed forest to 300 m altitude. Additional collections studied: MALAYSIA: Sarawak: Kapit, Upper Rajang River, Clemens & Clemens (K); Pelagus Rapids, Ashton S (L); Rajang, Haviland 3042 (BM, K, SAR, SING); Bukit Raya, Smith S (SAR). This recently described species is known from only a few specimens from Sarawak and, as can be seen from the citations, has long been confused with a number of other species. It is most immediately recognizable by its large leaves and the angle of the secondary veins with the midrib, and inflorescences that can become very long. 17. Kopsia rosea D.J.Middleton, sp. nov. TYPE: THAILAND. Krabi, Ao Luk, Thanbok Khorani National Park, D.D. Soejarto, T. Santisuk, K. Taylor & N. Nantasan 5945 (Holotype: A; Isotypes: F, L, MO, NY). Map 16. Frutex ad 3 m altus, ramulis glabris. Folia elliptica, cm, nervis 8 20 paribus. Corollae tubus mm longus, lobis mm. Usage synonyms: Kopsia alba auct. non Ridl. ex Henderson: Kerr, Fl. Siam. Enum. 2: , pro parte. Kopsia fruticosa auct. non (Roxb.) A.DC.: Middleton, Fl. Thailand 7: , pro parte. Kopsia macrophylla auct. non Hook.f.: Middleton, Fl. Thailand 7:

39 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 127 Tree to 10 m tall. Branchlets glabrous, lenticellate or not, terete or weakly angled. Leaves: petiole mm long, glabrous; blade cm, times as long as wide, coriaceous, elliptic, apex short acuminate with a blunt tip, base acute or cuneate, glabrous above and beneath, midrib shallowly sunken above, secondary veins 8 20 pairs with mm spacing, from midrib, prominent or flat above, prominent beneath, clearly distinguishable from tertiary venation above and beneath, curved ascending from midrib or only near margin, tertiary venation prominent or flat above and beneath, obscure or parallel or subperpendicular to midrib and oblique to secondary veins, then also somewhat reticulate, intramarginal vein straight or only weakly looped, inset from margin. Inflorescence dichasial, often with robust cincinnate branches, cm long with axes cm long and branches mm wide, glabrous to densely puberulent; peduncle cm long, mm wide, glabrous or puberulent; pedicels mm long, glabrous to densely puberulent; subtending bracts persistent; bracts present on pedicel. Sepals mm, times as long as wide, oblong, apex rounded, ciliate, glabrous to densely puberulent outside, glabrous inside. Corolla white, pale pink with darker pink eye, or white with a pink tinge; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent in upper part of tube above, around and slightly below the stamens, throat pubescent, glabrous or sparsely puberulent at top of tube outside; lobes mm, times as long as wide, obovate, apex rounded, not ciliate, glabrous outside and inside. Stamens mm from corolla base, which is ca. 0.8 of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, glabrous. Disk mm long, times as long as ovaries, glabrous, lanceolate, apex acuminate. Ovaries mm high, glabrous or with just one or two hairs; style ca mm long; style head ca. 1.2 mm long. Fruit falcate with a blunt hooked spur from around the middle of the fruit, mm, spur mm long, sparsely puberulent. Distribution: Southern Thailand, Peninsular Malaysia. Habitat: in forest to 300 m altitude. Recorded from limestone. Geographical selection of the 13 collections studied: MALAYSIA: Kelantan: Kuala Aring Forest Reserve, Kiew 42 (KEP). THAI- LAND: Surat Thani: Amphoe Phanom, Khao Song Pii Nong, Middleton 552 (A, BKF). Phangnga: Tap-put, Kerr (A, ABD, BM, E, K, L, P). Satun: Thung Wa, Kerr (A, E, BM). Krabi: Ao Luk, Larsen et al (BKF, L, P). This new species has similarities to Kopsia pauciflora, differing from it in the glabrous ovaries and in mostly having a pink or pinktinged corolla. The robust inflorescence structure is reminiscent of K. macrophylla, from which it differs in the stamens being inserted higher up the corolla tube. It differs from the pink-flowered K. fruticosa in the glabrous ovaries and in the glabrous twigs. Middleton (1999), working on the Thai species only, included the pink-flowered specimens of this species in a more variable K. fruticosa and the white-flowered specimens in a more variable K. macrophylla. With a broader knowledge of the ranges of variation in characters in the genus as a whole, it is now clear that K. fruticosa is native only in Burma and is morphologically less variable than previously thought, and that K. macrophylla is only in Peninsular Malaysia and contains only specimens with stamens inserted low down in the corolla tube. Kopsia fruticosa is, however, widely cultivated in Thailand and could easily be mistaken for K. rosea, except on closer examination. 18. Kopsia singapurensis Ridl., Fl. Mal. Pen. 2: ; Corner, Wayside Trees Malaya 1: ; Sleesen, Fl. Mal. Misc. Rec. 1: ; Whitmore, Tree Fl. Mal. 2: ; Markgr., Blumea 20: ; I.M.Turner, Gard. Bull. Sing. 45: ; Sévenet et al., J. Ethnopharmacol. 41: ; I.M.Turner, Gard. Bull. Sing. 47: TYPE: SIN- GAPORE. Chan Chu Kang, H.N. Ridley s.n. (Lectotype: K, designated here, see comment below). Syntypes: H.N. Ridley 59 (BM, SING), H.N. Ridley s.n. (L, Z), Lake & Kelsall s.n. (SING). Map 16; Fig. 7. Synonym: Kopsia fruticosa var. albiflora (Teijsm. & Binn.) King & Gamble, J. As. Soc. Beng. 4(2): , pro parte, not including the type (see comment below). Tree to 12 m tall, to 24 cm dbh. Branchlets glabrous, sparsely lenticellate or not, weakly angled. Leaves: petiole 5 12 mm long,

40 128 HARVARD PAPERS IN BOTANY Vol. 9, No. 1 MAP 16. Distribution of Kopsia rosea D.J.Middleton ( ) and Kopsia singapurensis Ridl. ( ). glabrous; blade cm, times as long as wide, subcoriaceous to coriaceous, elliptic, apex short acuminate with blunt or sharp tip, base acute or cuneate, glabrous above and beneath, midrib sunken above, secondary veins 7 15 pairs with 4 16 mm spacing, from midrib, prominent above, prominent or flat beneath, clearly distinguishable from tertiary venation above and beneath, curved ascending from midrib, tertiary venation prominent above, prominent or flat beneath, irregularly subperpendicular to midrib and oblique to secondary veins, intramarginal vein straight or only weakly looped, inset from or right at margin. Inflorescence dichasial, mostly many-flowered, cm long with axes (0.5 ) cm long and branches mm wide, glabrous or puberulent in upper parts; peduncle (0.2 ) cm long, mm wide, glabrous or puberulent; pedicels mm long, glabrous to densely puberulent, subtending bracts persistent, bracts present or absent on pedicel. Sepals mm, times as long as wide, ovate or oblong, apex rounded or obtuse, ciliate, glabrous to densely puberulent outside, glabrous inside. Corolla white with red eye ; tube mm long, mm wide, times as long as lobes, times as long as calyx, pubescent inside above, around and slightly below the stamens, throat pubescent, glabrous or rarely sparsely puberulent at top of tube outside; lobes mm, times as long as wide, elliptic or oblong, apex rounded to obtuse, ciliate, glabrous or, rarely, sparsely pubescent in upper quarter outside, glabrous or puberulent at base inside. Stamens inserted mm from corolla base, which is of corolla tube length in the rehydrated flowers measured; anthers mm, times as long as wide, apex mm from corolla throat; filaments mm long, pubescent. Disk mm long, times as long as ovaries, glabrous, oblong or awl-shaped, apex acute to acuminate. Ovaries mm high, sparsely to densely pubescent on top; style mm long; style head mm long. Fruit oblique ellipsoid with either a simple angle or a spur on one side, mm, angle/spur mm long, glabrous. Distribution: Peninsular Malaysia, Singapore. Habitat: in lowland evergreen forest, swamp forest or on river banks to 600 m.

41 2004 MIDDLETON, [REVISION OF] KOPSIA (APOCYNACEAE) 129 FIGURE 7. Kopsia singapurensis Ridl. A, habit; B, flower dissection; C, ovaries and disk; D, fruit. From Shah MS.1226 (A, B, C) and David 263 (D). Scale bars = 1 cm (A, B, D); 1 mm (C).